U. S. DEPARTMENT
OF HEALTH,
EDUCATION,
AND WELFARE
Public Health
Service
ALGAE AND
METROPOLITAN
WASTES
Transactions of the
1960 Seminar
-------
SEC TR W61-3
ALGAE AND
METROPOLITAN WASTES
Transactions of the Seminar on Algae and Metropolitan Wastes, held at
Cincinnati, Ohio, April 27-29, 1960, under the sponsorship of the Divi-
sion of Water Supply and Pollution Control and the Robert A. Taft
Sanitary Engineering Center.
U. S. DEPARTMENT OF HEALTH, EDUCATION, AND WELFARE
Public Health Service
Bureau of State Services
Division of Water Supply & Pollution Control
Robert A. Taft Sanitary Engineering Center
Cincinnati 26, Ohio
1961
-------
SEMINAR COMMITTEE
Dr. Alfred F. Bartsch
Chairman
Assistant Chief, Research Branch, Division of Water Supply and Pollution Control
Dr. William Marcus Ingram, In Charge
Biological Field Investigations, Technical Services Branch
Dr. C. Mervin Palmer, In Charge
Interference Organisms Studies
Research Branch, Division of Water Supply and Pollution Control
Dr. Herbert W. Jackson, Assistant Chief
Water Supply and Water Pollution Training, Training Program
CENTER PUBLICATIONS
The Robert A. Taft Sanitary Engineering Center is a national laboratory of the Public Health
Service for research, training, and technical consultation in problems of water and waste treat-
ment, milk and food safety, air pollution control, and radiological health. Its technical reports
and papers are available •without charge to professional users in government, education, and
industry. Lists of publications in selected fields may be obtained on request to the Director,
Robert A. Taft Sanitary Engineering Center, Public Health Service, Cincinnati 26, Ohio.
ii
-------
FOREWORD
The Public Health Service, particularly the Robert A. Taft Sanitary Engineering Center, has
a responsibility to promote free exchange of information on problems in the field of environ-
mental sanitation. This Seminar on Algae and Metropolitan Wastes was the first national meet-
ing devoted solely to this subject. It was part of a continuing program of seminars in the water
field at this Center.
The problem of algae is many faceted. Some species are a bane to men on earth, but others
are potentially the staff of life of men in space. It is important, however, to consider first
those problems most pressing and near at hand, such as the relationship of algae to this
Nation's spreading metropolitanism and to the disposal of urban wastes.
The Seminar on Algae and Metropolitan Wastes was limited to discussion of prevention and
control of objectionable blooms of algae resulting from enrichment by urban and other wastes.
This is a long standing problem which, as urbanization increases, will become more severe
if corrective measures are not developed and applied. We, therefore, regard this report as
particularly important and are pleased to present it under the sponsorship of this Center and
the Division of Water Supply and Pollution Control of the Public Health Service.
H. G. HANSON, Director
Robert A. Taft Sanitary Engineering Center
iii
-------
MEETING NOTE
The 1960 Seminar on Algae and Metropolitan Wastes was held in Cincinnati, Ohio, April
27-29, under the sponsorship of the Division of Water Supply and Pollution Control of the ]
Health Service Outstanding scientists and engineers from government, industry, and univei
sities were invited to participate. The seminar was attended by 139 registrants representing
27 states and the District of Columbia, and 4 foreign countries. The meeting consisted
discussions on (a) the problem, (b) growth characteristics of algae, (c) sources <
(d) methods of prevention and control, and (e) research needs. A tour of the Center s
was included, and an evening banquet at which Mr. R. G. Lynch of the Milwaukee Journal was
the principal speaker. Mr. Lynch's talk is presented as the Introduction to this volume.
Photo courtesy of Kenneth M. Mackenthun,
Wisconsin Committee on Water Pollution.
iv
-------
CONTENTS
INTRODUCTION
Page
Do They Dig You, Daddy-O, Or Are You Way Out? - R. G. Lynch 1
STATEMENT OF THE PROBLEM
Introduction
Induced Eutrophication - A Growing Water Resource Problem - A. F. Bartsch 6
Specific Problems in Lakes
Madison's Lakes: Must Urbanization Destroy Their Beauty and Productivity? - W. B. Sarles.... 10
Algae Blooms in Lake Zoar, Connecticut - R. J. Benoit and J. J. Curry 18
Klamath Lake, an Instance of Natural Enrichment - H. K. Phinney and C. A. Peek 22
Recent Changes in the Trophic Nature of Lake Washington - A Review -G. C. Anderson 27
Specific Problems in Rivers
Algae in Rivers of the United States - C. M, Palmer 34
GROWTH CHARACTERISTICS OF ALGAE
Nutrition
Fundamental Characteristics of Algal Physiology - R. W. Krauss 40
Micronutrients and Heterotrophy as Possible Factors in Bloom Production in Natural Waters -
L. Provasoli 48
Algal Density as Related to Nutritional Thresholds - J. B. Lackey 56
Productivity and How to Measure It
Methods of Measurement of Primary Production in Natural Waters - L. R. Pomeroy 61
Factors Which Regulate Primary Productivity and Heterotrophic Utilization in the Ecosystem -
E. P. Odum 65
SOURCES OF NUTRIENTS
Land Drainage
Land Drainage as a Source of Phosphorus in Illinois Surface Waters - R. S. Engelbrecht and
J. J. Morgan 74
Nutrient Content of Drainage Water from Forested, Urban and Agricultural Areas -
R. O. Sylvester 80
Wastes
Metropolitan Wastes and Algal Nutrition - W. J. Oswald 88
Limnological Relationships
The Role of Limnological Factors in the Availability of Algal Nutrients - G. H. Lauff 96
Nitrogen Fixation in Natural Waters under Laboratory Conditions - C. N. Sawyer and
A. F. Ferullo 100
Recent Observations on Nitrogen Fixation in Blue-Green Algae - R. C. Dugdale and J. C. Neess. 103
METHODS OF PREVENTION OR CONTROL
Limitation of Nutrients as A Step in Ecological Control
The Madison Lakes Before and After Diversion - G. W. Lawton 108
The Badfish River Before and After Diversion of Sewage Plant Effluent - T. F. Wisniewski 118
Spray Irrigation for the Removal of Nutrients in Sewage Treatment Plant Effluent as Practiced
at Detroit Lakes, Minnesota - W. C, Larson 125
-------
CONTENTS (Cont'd)
Chemical Methods for the Removal of Nitrogen and Phosphorus from Sewage Plant Effluents -
G. A. Rohlich 130
Stripping Effluents of Nutrients by Biological Means - G. P. Fitzgerald 136
The Use of Algae in Removing Nutrients from Domestic Sewage - R. H. Bogan ..... 140
Use of Algicides
The Practical Use of Present Algicides and Modern Trends Toward New Ones - K. M.
Mackenthun '. 148
RESEARCH NEEDS
Research Needs in Water Quality Conservation - B. B. Berger 156
PROGRAM 160
vi
-------
INTRODUCTION
This paper by Mr. Lynch was delivered as the dinner address of the Seminar. It is pre-
sented here to introduce the Transactions because it deals with the basic theme of achieve-
ment of better communication between scientist and layman and, indeed, between scientist and
scientist.
DO THEY DIG YOU, DADDY-O, OR ARE YOU WAY OUT?
R. G. LYNCH
The Milwaukee Journal, Milwaukee, Wisconsin
Mr. Toastmaster, scientists, technicans and
any others who may have wandered in:-
My editors sometimes complain that I am too
concerned with facts. They would like a lighter
touch, and I agree that it makes for better reading,
but I have observed that if I write that way and the
editor has to cut my copy to fit his space, it is the
facts that go out and the froth that remains.
The title of my talk, Do They Dig You, Daddy-o,
or Are You Way Out? is not intended to be funny.
Beatniks use that lingo as a barrier to outsiders—
to set themselves apart. Scientific and technical
people are guilty of the same thing. It is a form of
snobbishness and I think that you people cannot af-
ford the luxury.
As a fulltime reporter on natural resources,
the only one that I know of in the newspaper field,
I attend more or less technical meetings, work
from papers or summaries prepared by technicans
and scientists, and read extensively. And I have
observed a very serious lack of communication be-
tween scientific and technical people and the public.
It is particularly serious in the natural re-
sources field, where research and studies must
solve problems or develop approaches to them. The
men engrossed with these things meet to talk over
what they are doing, and they meet in something
like a vacuum—people who already know much a-
bout the subject telling each other more about it.
When they reach decisions or prepare plans,
they spring their ideas on the public and expect
them to be received gratefully, and implemented.
This seems to me to be a major reason why pro-
gress is so slow toward wise management of re-
sources.
Usually some changes are involved, some sup-
port must be generated, some money must be pro-
vided. People just do not accept changes unless
they are convinced of good reasons; they or their
representatives just will not dig up money for things
they don't understand. Exceptwhen they are fright-
ened, I should say. People of foreign countrieswho
want American dollars are shrewd enough to learn
the American language.
It is important that the people be taken along on
the whole trip, and that is not being done. Dr.
Carl O. Sauer, University of California geographer
and chairman of the 1955 international symposium
on "Man's Role in Changing the Face of the Earth",
said in summation:
"We are now come in 1955 to a revised version
of Aldous Huxley's 'bright new world' of the 1920's
— to a faceless, mindless, countless multitude,
managed from the cradle to the grave by a bril-
liant elite of madmen, obsessed with accelerating
technological progress".
Now, I don't want to be misunderstood. There
is exactness in scientific and technical terms.
Also, my friend, Joe Hickey, professor of wildlife
management at the University of Wisconsin, has
pointed out to me that scientific papers must be
written so that men of different languages may
understand them.
But technical and scientific people carry their
jargon outside and use it in addressing laymen,
sometimes unwittingly, sometimes to be impres-
sive. Even in their own meetings, they are some-
times less than comprehensible.
A year ago I attended a seminar on aquatic
biology here in Cincinnati. The first session was
devoted to radioactivity and the radiobiologists
talked a language that the others obviously had
difficulty in understanding.
"TLm" was one of the terms tossed around. A
speaker was kind enough to say that it was about
the same as LD-50, which I know is the dose lethal
to 50 per cent of an experiment population, so I
groped along with the speakers.
On the way out I asked a Ph.D. of my acquaint-
ance what TLm stood for. He said, "Oh, about the
same as LD-50." I said, "Yeah, I heard the
-------
ALGAE AND METROPOLITAN WASTES
man, too, but what do the letters stand for?" He
laughed and said he didn't know.
Another man turned around and said, "Toxicity
limit median, I think." A third man corrected him,
saying, "No, it's TOLERANCE limit median."
Some very interesting things were said at that
seminar and I think it is significant that the only
local newspaper coverage I saw was in the form of
interviews with Ernie Swift and Seth Gordon, a
couple of guest speakers from the wildlife conser-
vation field, and they were really just tails on the
kite. But newspaper men could understand what
they said.
Either at the seminar, or afterward by mail, I
got a notice of another seminar on benthic biology
out in California. I picked up my Webster's Col-
legiate dictionary and found benthos defined as
fresh water, bottom dwelling organisms. I wanted
more, and also had a hunch this was wrong, so I
lugged my big Merriam Webster to the dining room
table and found that benthos meant the bottom of
the sea and its flora and fauna.
Next I consulted encyclopedias at the office.
Americana said benthos was the fauna of ocean
depths under 100 fathoms; Britannica said it was
the sedentary bottom-living animal and plant life of
the sea.
And I said, "Why don't people say what they
mean!"
Incidentally, I replaced my 1943 Collegiate dic-
tionary with a new one, for benthos was in the new-
word section. As long as I can remember, I have
read with a dictionary at hand. I do not like to pass
over a word that I do not understand. But when I
must stop too often it destroys the continuity of my
reading and I may give up.
I gave up on Papini's Life of Christ and
Spengler's Drive to the East and Toynbee's History.
I have seven or eight volumes of Toynbee that one of
you can buy cheap. I have developed over the years
a deep resentment of men like Toynbee, who dis-
play their erudition with fancy language and lapses
into untranslated Greek, Latin, German and French.
I think that any man who has something of value
to say does not have the right to deny it to any mind
capable of understanding the thoughts or the facts,
simply because the possessor of the mind lacks
his own high level education.
Such men as Toynbee have forgotten that seman-
tics also is a science and, I think, an art as well.
There are others who realize this and try to com-
municate but fail because'they are too steeped in
the jargon of their disciplines.
Perhaps some of the Wisconsin men here are
familiar with Prof. John Curtis's new book on
Wisconsin vegetation. It is a wonderful book in its
field and I feel like a flea on a dog's back when I
mention it here, in this way. But it illustrates my
point.
Publication notice and wrapper said that the
book was written with a minimum of technical
terms, for the use of (among others) farmers,
conservationists and men in recreation and weed
control work. Well, Curtis really had me wearing
out that dictionary! I think I found 20 or 30 words
in the first couple of chapters that were not in my
Collegiate and some were not even in the big Mer-
riam. Mesic andxeric were terms constantly used;
neither was in the big dictionary. I called up two
foresters and a curator of botany who could not de-
fine the words. Finally I got what I wanted from a
university professor.
A man could as easily use "dry" and "drier" as
"xeric" and "more xeric" if he thought of it.
But Curtis made a sincere effort to get through.
He was on the right track when he used"podzoliza-
tion" with the explanation that it meant topsoil im-
poverishment by leaching and translocation. That
is something the scientific and technical man can
do. It is all right for him to employ words which
will be useful to his reader or listener, with an
explanation of the meaning. The type of reader who
wants information is apt to be receptive to new
words.
I must say that I stuck with Curtis and was well
rewarded, but I wonder how many farmers, weed
control men and conservationists went beyond such
expressions as "varietal endemism", "morpholo-
gical subspeciation", "altitudinal retardation of
phenoibgical events" or "Gaussian amplitude curve".
How many readers do you think will use a dic-
tionary, much less read with one at hand?
And when the offering is oral, there is no dic-
tionary and you can lose an audience quickly. Dr.
Luna Leopold of the geological survey addressed
a Milwaukee gathering of conservationists not long
ago and his paper was a polished masterpiece. I
admired his flow of language. But as we left the
hall an attorney said to me, "I should have brought
along a dictionary."
The lack of communication is being recognized
by more and more men in scientific and technolo-
gical fields as a serious thing. My presence here
is a manifestation of that.
Dr. Marston Bates, University of Michigan zool-
ogist, in a book published only last month, referred
to a new word, biocenosis. created as a substitute
-------
Introduction
for "community" in order to pinpoint a meaning.
"Biocenosis," he said, "leads easily to biomes
and biochores, to ecosystems and ecotones. These
are all lovely words but they don't really say any-
thing new. The trouble is that the word-coiner,
sinking blissfully into his additions, gradually loses
all communication with the outside world. He e-
merges from time to time to complain that the
world doesn't really understand or appreciate his
important thoughts—meaning his big words."
Prof. Hickey, whom I mentioned earlier, edited
the Journal of Wildlife Management for three years.
He ran across such gems as this:
"While piloting a Polar Cub 30 miles west of
Churchill, two foxes were seen in a highly abnor-
mal condition." Aside from indicating that two
foxes seem to have been piloting an airplane, this
is not a very informative statement.
And this one:
"A porcupine covered with Sarcoptes scabeii was
brought to the laboratory in a morbid condition."
That Journal, incidentally, recently had a fuss
in its letter column over editorial requests for
"polishing up" manuscripts, which led Justin
Leonard, a Ph.D. in zoology who heads up conser-
vation research in Michigan, to comment:
"The Ph.D. in science can make journal editors
quite happy with plain, unadorned, eighth grade-
level composition."
Dr. Watson Davis, director of Science Service,
in an address last December at the conference on
scientific communication of the American Associa-
tion for the Advancement of Science, said this:
"The great art in telling is to make the words
mean whatyou want them to, not in your own mind,
but in the minds of others. There is little need for
'writing down'. This is demonstrated by the daily
newspaper which serves children and old folks a-
like. Clearness and vividness are more needed
than vocabulary limitations based on word lists."
I cannot wholly agree with Dr. Davis. How can
there be clearness and vividness if the reader or
listener does not know the meaning of the words?
As for "writing down" (or talking down), it is dan-
gerous even to think in such terms.
Newspapers, too, have a responsibility in this
field and not all of them are making a real effort
to meet it. A few papers, like the Milwaukee Jour-
nal, try to give their readers a good ration of things
they should get, along with things they want, which
must be provided if papers are to be sold. After
all, newspaper publishing is a business. But The
Journal has lost no circulation because of its offer-
ing of substantial reading matter.
I think more and more papers in the next decade
or two will assign men to natural resources writing
and they will try to inform themselves, as I have
tried, in order to be interpreters of things scienti-
fic and technical.
But now the burden is heavily on you people and
your fellows. You must realize that competition
for newspaper space is very keen. The editor's
wastebasket is large and his reporters' assignment
sheet is crowded.
If your press releases do not get into print, if
reporters do not come to your meetings or, having
come, walk out and print nothing, examine your
efforts. Maybe your mimeographed papers are too
full of jargon, and also the summaries, which
usually are more appealing to the press. I think
authors of papers should not prepare the sum-
maries. That is a job for an interpreter, a public
relations man.
In my observation, a better job of communica-
tion results if a public relations man is a good
newspaper man with some scientific and technical
knowledge superimposed, rather than a scientific
or technical man drafted for a task which he may
not understand. Some ignorance of the jargon is an
asset; it guarantees a degree of translation.
I think that better meetings of this sort would
result if panel chairmen, in addition to summari-
zing at the end of a discussion, started it off with
an easily under stood explanation of its purpose and
probable content. Such prefaces might well be out
on the press table.
As for speaking and writing for popular con-
sumption, I think it is well to remind yourself
that, in the present state of biological evolution,
the people you approach probably have minds as
good as yours; they simply lack the tools to make
use of their minds as you use yours. Your job is to
supply the tools.
Words are the tools of the mind. I am sure that
you know that man could not think until he invented
words, or symbols, to think with. If you want peo-
ple to think about things that concern you and your
job, use words they know or supply new, useful
words. Do it casually, not in a way which seems
to say, "See how much more I know than you do!"
We had an incident in Wisconsin where a game
biologist told a group of farmers, "I won't attempt
to explain thatjyou wouldn't understand it anyhow."
The dropping of that gem into the water, you may
be sure, spread some ripples.
I think it is well to remind yourself, too, that
-------
ALGAE AND METROPOLITAN WASTES
people are not compelled to listen to you or read I used to become angry when I heard a speaker
you. Tou are in a buyer's market, competing with giving a very polished dissertation in the jargon of
murder, rape, scandal, politics and comics in the his profession and saw his audience looking at each
newspaper and sex, whodunits and westerns on other with raised eyebrows.
television and drugstore book counters.
But I have reached the point where I feel sorry
At least, you must make what you say under- for such a man and say to myself:
standable, and if you have the talent you had better
make it interesting. Regard it as a challenge, for "There he is, erudite as all get out. and
that it surely is. ignorant as hell."
-------
STATEMENT OF PROBLEM
-------
INTRODUCTION
INDUCED EUTROPfflCATTON - A GROWING WATER RESOURCE PROBLEM
A. F. Bartsch, Assistant Chief
Research Branch
Division of Water Supply & Pollution Control
Public Health Service
Cincinnati, Ohio
James Whitcomb Riley wrote sadly on "The
Passing of the Backhouse". Many others share his
nostalgic feelings and join Kim in quietly lament-
ing its passing from the American backyard scene.
Although there were sewers in ancient Rome and
Baghdad, the water carriage system of the modern
world really grew with the industrial revolution
that concentrated people in cities during the late
1800's. It is largely due to this passing of the back-
house and the train of events that followed that we
are faced now with problems involving algae and
metropolitan wastes. Streams and lakes, needed
as water supplies or used for other purposes, have
been receiving quantities erf sewage that serve as
nutrients for excessive growths of algae.
Algae of concern are mostly microscopic, free-
floating ones. Their spectrum of color includes
grass green, gold, brown, red, blue-green, and
various shades in between. In moderate quantities,
algae are extremely beneficial. Problems arise
only when they are too abundant. Although some
algae are more notorious than others for causing
undesirable blooms, almost all are objectionable
if present in sufficient quantity. We are made a-
ware of their presence by the color they impart to
the water, by the characteristic odors in the vi-
cinity, and by unsightly scums, drifting windrows
and drying drifts along the snore. In some urban
centers of the United States, the layman is quite
well acquainted with algae and what they do to the
utility of water.
Nutrients that support algal growth in lakes and
reservoirs originate in the surrounding drainage
area and enter with the runoff. If the soil is fertile,
nutrients for algae are likely to be abundant. Even
the time and pattern of applying fertilizer to agri-
cultural land may influence the-contribution of nu-
trients to the aquatic environment. In their simple
existence, algae utilize the mineral nutrients that
have come from the land as well as carbon dioxide
dissolved from the air or released in decay of or-
ganic matter. During warm seasons when growing
conditions are otherwise favorable, algal produc-
tion slows down and finally is stopped by depletion
of any one nutrient element. Because nitrogen and
phosphorus are not abundant in most surface wa-
ters, more commonly than other elements they
seem to act as a brake on further rapid growth as
the season progresses. Thus, although some nat-
urally fertile lakes habitually develop algal blooms
from year to year, as a rule, such blooms are less
frequent and objectionable than in lakes polluted
with sewage.
Sewage contains a multitude of different com-
ponents, of which many have not even been pre-
cisely identified. It is a rich source of nutrients
capable of growing dense populations of algae. Es-
pecially notable are nitrogen, phosphorus, potas-
sium and carbon, as shown in Table 1. There is
Table 1. SELECTED ALGAL NUTRIENTS IN
SEWAGE*
Nutrient
Concentration, ppm
Nitrogen
NH3
NO3
NO2
Org.
Carbon
Sol. phosphorus
Potassium
20- 50
7-40
0 -4
0-0.3
3-42
66 - 176
1-13
13 -44
*Taken from Fitzgerald and Rohlich, 1958. Based
on data for 14 sewages.
reason to suspect that sewage also contains organ-
ic substances that may possibly function as algal
stimulants at extremely low concentrations. The
algae-growing ability of sewage in undiluted form
is well known from experiences with laboratory,
pilot, and full-scale waste stabilization ponds.
Yields from sewage can be as high as 23 to 36 tons
per acre per year in ponds only 2 to 12 inches deep
(Gotaas, Oswald and Ludwig, 1954). Potential yields
obtainable in this way may be 10 to 20 times as great
as those from cultivated land. Sewage nutrients are.
also effective for growing algae, although some-
what restrained, after dilution in surface waters.
The objectives of conventional sewage treatment
are removal of suspended solids, microoganisms,
and BOD. In most cases, meeting these objectives
-------
Induced Eutrophication
makes treated sewage totally acceptable for dis-
charge to surface waters so far as health, esthetic,
and most water reuse considerations are con-
cerned. But even with so-called complete treat-
ment, removal of algal nutrients - nitrogen and
phosphorus in particular - is negligible. This de-
ficiency of waste treatment has been cited many
times. So far, little consideration has had to be
given to removal of these elements during or after
the conventional treatment process. In the final
analysis, conventional treatment will not prepare
sewage adequately for discharge to waters physi-
cally suitable for algal growth.
It is well known that lakes do not persist forever
as permanent marks on the landscape. During the
course of history, they change continually -
mature, grow old, and eventually change to dry
land. Examples of this development at all stages
can be found in numerous geographic areas. As
aging progresses, deep, infertile lakes accumulate
sediments and nutrients, become shallower, more
fertile and productive and thus qualify to be called
eutrophic lakes. Periodically they may have ex-
cessive quantities of algae as a sign that the wheels
of the aging process are fully in motion. Induced
eutrophication through sewage pollution acceler-
ates the aging process regardless of when in the
history of the lake such nutrients are applied
(Figure 1). At the algal bloom stage, people are
c **
il
o •»
K u
13
0 u.
e
g
a
EFFECT OF FERTILIZERS
ARTIFICIAL OR DOMESTIC
/
EXTINCTION
NATURAL
EOTROMIICATIM
ABE OF THE LAKE
Figure 1.
NATURAL AND INDUCED EUTROPHI-
CATION. (From Hasler, 1947).
concerned because of the immediate unpleasant-
ness of algae that interfere with water resource
enjoyment and not because this is a milestone in
the progressive extinction of the lake.
Throughout the world, there are many examples
of surface waters made fertile by sewage. Among
them are ponds and lakes intentionally enriched to
produce more fish or for other purposes. These
bodies of water are not the problem. Lakes un-
intentionally enriched with sewage, however, char-
acteristically have changed from sparkling, clear,
useful ones to unsightly and malodorous ones. The
nature and extent of the problem as it existed
worldwide in 1946 was well described by Hasler
(1947). A familiar thread of similarity and repe-
tition can be noted in each case. Sometimes it in-
cludes all but more commonly only several of the
following steps: (1) introduction of raw or treated
sewage, (2) replacement of prized deep water trout
orwhitefish by less desirable kinds, (3) increase in
plankton or free-floating plant and animal life, and
(4) explosive seasonal appearance of the blue-green
alga, Oscillatoria rubescens.
The Zurichsee, a lake in the foothills of the
Swiss Alps, is a celebrated example of induced
eutrophication. The lake is composed of two ba-
sins separated by a narrow passage. The upper
basin received no sewage and remained essentially
unchanged whereas the lower basin, receiving the
sewage from a group of small communities with
more than 100,000 people, underwent typical
changes as described above. Other Alpine lakes
suffered the same fate, as did also lakes in Sweden
and England. But, one need not go to Europe to
learn about the problems of induced eutrophica-
tions; there are excellent examples here in the
United States.
Many people know beautiful Madison, Wisconsin,
because of its chain of four lakes with the musical
names, described poetically by Longfellow. Others
know it also as a classical example of the conse-
quences of bringing together lake water, algae, and
sewage. The problem has a long history, going
back to 1920 or even earlier. Many technical people
have been fascinated by it and have studied one as-
pect or another. The voluminous historical record
of the problem - as it emerged and grew, how it
aroused the public, the technical, political and
legislative actions taken, and finally the remedy -
make instructive and fascinating reading.
The algal bloom problem has plagued the prop-
erty owners on the shores of Connecticut's Lake
Zoar also. This body of water is an impoundment
on the Housatonic River (Curry and Wilson, 1955;
Benoit, 1955) created in 1919. By 1947 production
of algae had increased to the nuisance level, stim-
ulating investigative action by the State government.
Again sewage and industrial wastes have been im-
plicated as sources of algal nutrients. A number
of potential remedies have been investigated.
Across the Nation, at Seattle, Lake Washington
is a newer entry to the algal problem field
(Sylvester, 1956; Edmondson, 1956; Bogan, 1956;
Edmondson, Anderson and Peterson, 1956). The
story begins about the turn of the century when
Seattle sewage was first discharged to the lake. As
the population grew and new lakeside communities
-------
ALGAE AND METROPOLITAN WASTES
were sewered, the quantity of sewage kept increas-
ing. By 1941 the sewage was finally diverted to a
treatment plant with the effluent going to a differ-
ent drainage. But the problem was not ended by
provision of the combined sewer system because
storm water and sewage frequently were bypassed
to the lake through numerous overflows. Since
1941, the growing communities in the surrounding
area provided additional sewage treatment facili-
ties draining their effluents to the lake. These
plants and a number of other sources upstream on
tributaries supply algal nutrients in quantity. There
is a history of parallel biological change also with
records going back to 1933. The quantity of algae
has increased significantly. Evidence of this is
shown both by increasing numbers of algae that
have been measured and recorded and by progres-
sively decreasing transparency of the water. By
1955, Oscillatoria rubescens. the rust-colored
form that often seems characteristic of induced
eutrophicatton, appeared on the scene and became
a prominent part of the algal population. Other
limnological changes, Including decrease in oxy-
gen resources in the deeper areas, have been stim-
ulated in turn by accumulation and settling out of
the increasing amounts of organic matter syn-
thesized in the lake. Intense efforts are being di-
rected to finding and executing a remedial and pre-
ventive program for this extremely valuable body
of water.
The Potomac River below Washington, D. C., is
another susceptible body of water. This tidal reach,
where the water rocks back and forth, is as much
like a lake as a river. The immediate tributary
land area has a concentrated sewered population of
1.7 million people. Records from a survey on the
river made by Purdy in 1913 show a moderate a-
mount of algal growth at that time. Many changes
have occurred since then in population size and
distribution pattern, in land uses upstream, in
sewerage and sewage treatment, and in silting of
the tidal flats and denuding them of aquatic plants.
Limited intermittent studies during the last few
years reveal that algal populations are much great-
er now than in the past. Blooms of blue-green algae
are common and become objectionable at least in
some of the bays and coves. Undoubtedly, the peak
of productivity has not yet been reached.
An interesting deviation from the usual pattern
just described occurred on Long Island in the area
famous for Long Island ducklings and Blue Point
oysters. In raising ducklings, it is customary to
have ponds accessible to the fowl for drinking and
exercise. Usually it is downslope from the feeding
troughs. Because of this arrangement, the pond
becomes a depository for manure from ducks pud-
dling in the water. Manure from others waddling
near the feed boxes reaches the pond asa result of
wet weather drainage down the slope. These fluid
wastes, rich in algal nutrients, drained or were
pumped to Moriches Bay. As they accumulated,
they stimulated formation of dense blooms of algae
of types unacceptable as the normal food of oysters
inhabiting the Bay. Oyster production declined un-
til the famous Blue Points were gone from the
market.
In spite of considerable study, the duck farm
waste problem has not been solved completely, but
in 1953 a passage from Moriches Bay to the Atlantic
Ocean was reopened by dredging and the action of
high seas. Now, the tides scour and flush the
wastes from the Bay, and harvesting oysters is
again a gainful occupation.
These are but a few examples of the conse-
quences of induced eutrophication. Many others
could be cited, but these are sufficient to show
that the trouble spots of the past are not isolated
and peculiar situations.
During the past year, more than $650,000,000
were spent for construction of sewerage facilities,
including waste treatment. Amounts spent during
other recent years are similarly impressive. The
citizens who pay this bill, like the people in
Madison or in communities on the shores of Lake
Washington or Lake Zoar who paid for secondary
treatment, have a right to expect safe, clean, at-
tractive, useable water in return. In some cases -
and fortunately so far they have been few - such
great expectations have not materialized. With the
mineralizing processes in sewage treatment, nitro-
gen and phosphorus are made more readily avail-
able as algal nutrients, and algal problems have
taken the place of the preceding, more typical pol-
lution problems.
It is obvious that the technique of waste treat-
ment has not yet gone far enough, that there are
communities that will have to do more than remove
suspended solids, microorganisms, and BOD from
sewage. It is necessary to think more seriously a-
bout additional- or tertiary - treatment to remove
algal nutrients where environmental factors re-
quire it. Lakes are more susceptible than streams
to algal troubles and must be considered with ex-
treme caution as potential basins to receive treated
sewage. Where flowing waters are available for
dilution, they are preferable. Some time ago, con-
sideration was given to discharging treated sewage
to Lake Tahoe. Apprehension of creating an irre-
versible algal problem in such valuable recreation-
al waters was an important factor in rejecting the
idea.
We stand now at a point where the need for more
knowledge is unquestioned. It is necessary to ex-
amine this growing algal problem, measure its
size, determine its exact nature, find how to rem-
edy existing situations, and learn to treat wastes
more effectively in order to prevent them. The
urgency of this need is emphasized by several
facts. Population increases continually. As a re-
-------
Induced Eutrophication
suit, there are ever more people in sewered com-
munities - not only to swell the quantity of sewage
with •which we are concerned, but also to compete
for recreational and other uses of available water.
As the quantity of sewage increases, its character
changes also in many ways. One is the per capita
increase in phosphorus content traceable to the
recent popularity of phosphorus-bearing deter-
gents. Most of it finds its way to the sewer. As
water resources development goes forward, we
find ourselves continually losing many miles of
free-flowing rivers and gaining many square miles
of quiescent standing water. With all of these
changes, we can expect to be faced more frequently
with problems of algae and metropolitan wastes.
Problems of the past have been costly ones. Prob-
lems of the future will be costly also unless we
spend the effort now to learn how to cope with them.
REFERENCES
Benoit, R. J. 1955. Relation of phosphorus content
to algae blooms. Sewage & Industrial Wastes 27:
1267-1269.
Bogan, R.H. 1956. A new critical phase of the Lake
Washington pollution problem: Treatment and con-
trol of nutritional elements. The Trend in Engi-
neering, April 1956, pp. 13-14.
Curry, J. J. and S. L. Wilson. 1955. Effect of
sewage-borne phosphorus on algae. Sewage & In-
dustrial Wastes 27: 1262-1266.
Edmondson, W. T. 1956. A new critical phase of
the Lake Washington pollution problem: Biological
aspects of the problem. The Trend in Engineering,
April 1956, pp. 11-12.
Edmondson, W. T., G. C. Anderson, and Dr. R.
Peterson. 1956. Artificial eutrophication of Lake
Washington. Limnology & Oceanography 1: 43-53.
Gotaas, H. B., W. J. Oswald, and H. F. Ludwig.
1954. Photosynthetic reclamation of organic
wastes. The Scientific Monthly 79: 368-378.
Hasler, A. D. 1947. Eutrophication of lakes by
.domestic drainage. Ecology 28: 383-395.
Sylvester, R. O. 1956. A new critical phase of the
Lake Washington pollution problem: A paradox of
water-pollution control. The Trend in Engineering,
April 1956, pp. 8-10.
-------
Specific Problems in Lakes
MADISON'S LAKES:
MUST URBANIZATION DESTROY THEIR BEAUTY AND PRODUCTIVITY?
WILLIAM B. SARLES
Department of Bacteriology, University of Wisconsin, Madison, Wise.
To the geologists, lakes such as those found
in and near Madison, Wisconsin, are, in geologi-
cal time, relatively short-lived widespreads in a
river that drains an extensive watershed. The four
principal lakes are on the Yahara River which flows
from the North to the Southeast where it empties
into the Rock River. These lakes are known by
Indian names given in 1858 by Dr. Lyman C.
Draper, founder of the Wisconsin State Histori-
cal Society, as Mendota, Monona, Waubesa, and
Kegonsa; a fifth, spring-fed, Lake Wingra, lies
South of Lake Mendota, and drains through Mur-
phy's creek into Lake Monona. Dr. Draper chose
the names from the language of the Chlppewa
Indians, who never dwelt in the vicinity of Madison,
but preferred to live In northern Wisconsin. His
choices were based on the beauty of the sound of
the names. However, the Winnebago Indians, who
lived in the vicinity of the Madison lakes, called
the entire chain "Taychoperah," which means "four
lakes." The Winnebago name, "Wingra" means,
literally, "dead lake", and it was aptly applied
because in the 19th century, that spring-fed, shal-
low, marl-bottomed body of water was nearly ex-
tinct. Extensive dredging operations, carried on
from 1912 to 1918, were needed to return Wingra to
the status of a lake.
We are concerned mainly with" the four lakes of
the Yahara drainage basin. They are relatively
young, according to the geologists, because it was
only 20,000 to 30,000 years ago that the ice of the
last glacier melted and left the water level of Lake
Mendota at approximately 8 50 feet above sea level.
The water level of Lake Monona stood originally
at about 844.5 feet, that of Lake Waubesa at 844.3
feet, and the water of Lake Kegonsa was originally
about 842.5 feet above sea level. Government sur-
veyors, who started their work in the southern part
of the State, and worked toward the North, desig-
nated the lakes by number in the order in which
they found them: Thus, Kegonsa was called First
Lake, Waubesa, Second Lake, Monona, Third
Lake, and Mendota, Fourth Lake. These numeri-
cal designations are gradually being dropped, but
"old-timers" still use them to show their sophis-
tication, and to confuse newcomers.
The area of land which drains to Lake Mendota
Is approximately 250 square miles, and the water
from it feeds four creeks, one small river (the
Yahara), and an unknown number of springs. Ac-
cording to the best estimates available, the three
lower lakes — Monona, Waubesa, and Kegonsa —
receive drainage from an additional 250 to 300
square miles of land. Hence, the total Yahara
drainage basin comprises 500 to 550 square miles
of fertile farm lands, wooded slopes, and city or
village properties. Lake Mendota is the largest
anddeepestof the four. Monona the second, Kegonsa
the third, and Waubesa the fourth in order of area
and depth.
The first white settlers in the Taychoperah area
were entranced by the beauty of the lakes and by
their productivity. They found that all of the lakes
contained an abundance of fish, wildlife and water-
fowl. Unfortunately, some of the early settlers
introduced carp into Lakes Mendota and Monona
sometime between 1850 and 1870.
No one knows exactly when nuisance conditions
caused by excessive growths of algae and rooted
plants first occurred in the four lakes, but the first
written report of such a phenomenon appeared in
1888 in the Transactions of the Wisconsin Academy
of Sciences, Arts, and Letters, VH (1883-1887) in
a paper written by Professor William Trelease of
the University of Wisconsin. This paper, which was
an essay, and which contained little factual infor-
mation, was entitled "The Working of the Madison
Lakes"; it describes nuisance conditions in Lakes
Mendota and Monona as early as June, 1882. Pro-
fessor Trelease also reported a bloom in Lake
Waubesa in 1886. Despite this report's inadequa-
cies as a scientific paper, it establishes the fact
that Madison's lakes could become overproductive
of algae and thus produce nuisance conditions be-
fore urbanization of the area became extensive.
Madison's first water works was completed in
1884. It pumped water from a highly productive
aquifer about 850 feet beneath the surface. This
water-bearing stratum continues today to supply
the needs of the city and its nearby communities
with good, but "hard" water. In 1884, the city,
built on an isthmus between lakes Mendota and
Monona, had a population of 12,000, but no sew-
erage. Privies, cesspools, and direct drains to
the lakes were used to dispose of sewage. Storm
water then, as now, flowed directly to the lakes.
10
-------
Madison's Lakes and Urbanization
11
In December of 1884, the Mayor and Common
Council of the City of Madison stated: "Considering
our finances, our sanitary desires, our presentand
prospective population, we want the cheapest, the
best, the safest, and the most flexible system —
(of sewage treatment and disposal) — available."
This was a prophetic statement; in it may be found
the needs, the limitations, and at least indications
of the problems that are with us today. The govern-
ment of Madison was faced then as now with the
problem of collecting, treating, and disposing of
the waste water of a rapidly growing community.
Also, then, as now, the wheels of government
turned slowly, and it was not until June of 1894,
ten years later, that Mr. John Nader reported to
the City Council that "the lakes are not properly to
be used as receptacles for sewage in the crude state"
— and that a sewerage system and treatment plant
should be constructed "at the earliest opportunity."
Immediately, arguments arose in the Council
over which system for sewage treatment and dis-
posal was the cheapest, the best, the safest, the
most flexible, and where the effluent was to be dis-
charged. Councilmen from wards with frontage on
Lake Mendota argued with those from wards with
frontage on Lake Monona. Finally, it was decided
to construct the treatment plant on the Yahara River
between Lakes Mendota and Monona with the hope
that during flow from the treatment plant to Lake
Monona, dilution of the effluent would be sufficient
to make the water of the river entering Lake Monona
as pure as that of the lake. Furthermore, this
location was chosen because it was on low ground
at the far northeast edge of the city, and thus ful-
filled the "out of sight, out of mind" criterion.
In August, 1894, the seventh of a succession of
engineers to study the Madison sewage treatment
and disposal problem submitted his report and, in
1895, construction of the sewerage system was
started. In October, 1897, construction was com-
menced of a sewage treatment plant which would
use chemical precipitation followed by filtration
through sand—a so-called "Inter national Process"
—with a predicted capacity of 200,000 gallons per
acre of sand filter per day. Unfortunately, the
treatment plant could not be made to operate pro-
perly, fell far short of the claim that it would pro-
duce an effluent "as pure as the water of Lake
Mendota", and was abandoned in January, 1901.
Mr. F. E. Turneaure, then City Engineer, later
Dean of the College of Engineering, was ordered
to construct a new treatment plant which would em-
ploy septic tanks followed by cinder filters which
would drain into the Yahara River near its outlet
into Lake Monona. This was done, and in 1902,
boatmen and residents along Lake Monona reported
that great improvement had occurred in the "puri-
ty of the Lake's water", and that "its odors and ex-
cessive weed growths had been decreased."
However, by 1906, the capacity of Turneaure's
treatment plant had been reached, by 1911 it was
overloaded, and it again became necessary to face
up to the problem of getting "the cheapest, the best,
the safest, and the most flexible system available."
Again, the "out of sight, out of mind" criterion was
employed, and on the basis of a study made by
engineer John M. Alvord of Chicago, E. E. Parker,
then City Engineer, built a new treatment plant in
the Town of Burke, two miles North of the Turneaure
plant. The Burke plant had a rated capacity of
5,000,000 gallons per day. It consisted of primary
settling tanks followed by sprinkling filters which
drained into a ditch leading to Starkweather Creek
at the North end of Lake Monona, about one mile
Northeast of the inlet of the Yahara River. The
plant was completed and put into operation in 1914.
Thus, from 1906 until 1914, Lake Monona was pol-
luted with poorly treated effluent from the old
Turneaure plant.
Nuisance growths of algae and offensive odors
occurred with increasing frequency in Lake
Monona; these undesirable conditions reached a
climax in the summer of 1918. Madison had grown
to a population of over 30,000, and the shores of
Lakes Monona, Waubesa.and Kegonsa were thickly
populated by cottagers. In addition, there were
several resorts on the shores of these lakes, and
the City of Stoughton, on the Yahara River, to the
Southeast of Lake Kegonsa, had grown to a popula-
tion of near 5,000. The lakes were beginning to
show the effects of urbanization, and people were
becoming aware of the problems created by use of
the lakes as "receptacles" for inadequately treated
sewage.
Much time has been spent in presentation of the
early history of sewage treatment and disposal in
Madison, and the development of public awareness
of the Madison lakes problem. This has been neces-
sary to gain an understanding of the studies which
have been made, and the steps which have been
taken since 1918 in an attempt to achieve desired
remedies. Time does not permit detailed descrip-
tion or discussion of investigations made, conclu-
sions reached, governmental agencies formed to
cope with the problem, legislation enacted, court
tests of new laws, and procedures followed to com-
ply with these laws. All that can be done is to
present these events in more or less chronological
sequence, and to comment on them briefly.
1919-1920. The Alvord and Burdick study and
"Report Upon the Cause of Offensive Odors from
Lake Monona, Wisconsin" blamed decomposition of
algae for the nuisance conditions encountered in
1918. In this work, the firm of Alvord and Burdick
had been aided by studies carried on under super-
vision of Dean H. L. Russell, a bacteriologist, and
Professor Chauncey Juday, a limnologist of the
University of Wisconsin. Dean Russell and Pro-
-------
12
ALGAE AND METROPOLITAN WASTES
fessor Juday agreed that "although the (treated)
sewage (from the Burke plant) may, to some degree,
stimulate the growth of the algae, we are of the
opinion that these growths would be sufficiently
luxuriant from natural causes to constitute a nui-
sance even if the sewage or effluent were not allowed
to reach the lake, or were subject to treatment to
exclude whatever nitrogenous matter it may con-
tain that would help support algae in the lake."
The report also stated "that there is no present
justification for attempting to dispose of the Madi-
son sewage plant effluent elsewhere than in Lake
Monona as is now done." The report recommended
that the Madison Board of Health, which had been
using copper sulphate to kill algae in Lake Monona,
should continue to do so only on the basis of con-
tinuing laboratory studies directed toward determi-
atton of the need for such treatment.
1920-1921. Studies were made on improvements,
recommended in 1920 by Alvord and Burdick, of
the Burke treatment plant, and the desirability of
building a new treatment plant near Nine Springs
Creek, which drains into the Tahara River just above
Lake Waubesa. The possibility of diverting Madi-
son's sewage to the Wisconsin River, approximately
18 miles West of the City, was considered, but
rejected as being too expensive. The Oscar Mayer
and Company packing plant commenced operations
in a location to the North and West of the Burke
treatment plant, and started to contribute its sew-
age to that of the City.
1923-24. Offensive odors from Lake Monona
were prevalent. The Lake Monona Improvement
Association was formed, and the statement was
made that "we are through with chemists and engi-
neers; let's get down to business." The Common
Council formed a Lakes and Rivers Commission to
supervise the sanitary conditions of the lakes and
streams. The City of Stoughton expressed opposi-
tion to establishment of the proposed Nine Springs
treatment plant, but the City of Madison, with ap-
proval from the State Board of Health, went ahead
with its plans for construction.
1928. The Nine Springs treatment plant, em-
ploying Imhoff tanks followed by trickling filters
and final clariflers was put into operation. Ef-
fluent from this plant flowed through the Nine
Springs Creek into the Yahara River, and thence
into Lake Waubesa. The Burke plantwas continued
in operation, but on a limited basis in line with its
rated capacity.
1929-30. The proposed sewage treatment plant
of the Village of Middleton, located at the northwest
end of Lake Mendota, and designed to discharge
effluent into Lake Mendota, was given preliminary
approval by the State Board of Health. These plans
were opposed by the City of Madison and by owners
of propertyon Lake Mendota. The controversy re-
sulted In creation of the Madison Metropolitan
Sewerage District under provisions of a law passed
by the 1927 State Legislature. Establishment of
the District made provision for collection and
transmission of Middle ton's sewage to the Nine
Springs treatment plant. After public hearings, the
Madison Metropolitan Sewerage District was up-
held by order of the Dane County Court on Febru-
ary 3, 1930. The decision of Judge George Kroncke
representated thorough study and was a carefully
worded statement of the Court's opinions. This
decision has stood the tests of time and of higher
courts, and may well serve as a model for future
use.
1931-33. Interceptor sewers, pumping stations,
and sewage treatment plants of the City of Madison
were transferred to the Madison Metropolitan
Sewerage District Commissioners of the District
stated in their plans for the future that "Inasmuch
as the waters of Lakes Mendota and Monona should
be maintained as free from contamination as pos-
sible, it is desirable to plan for the complete elim-
ination of the flow of effluent from sewage disposal
plants into the waters of these two lakes, and to
treat the sewage so that no unstable or improper
effluent will be permitted to enter the Yahara
River." This was a major statement of overall
policy that has been followed during the past 29
years.
A new "Clean Lakes Association" was formed in
1931 to protest against and to seek prevention of
pollution of Lakes Monona and Waubesa. This as-
sociation immediately requested that the Burke
plant be closed and that all of the District's sew-
age should be treated in the Nine Springs plant.
The summers of 1931 and 1932 were bad ones for
Lake Monona, and despite frequent and heavy
treatments with copper sulfate, the lake bloomed
and gave off bad odors. At the same time, in 1931,
Lake Mendota suffered an outbreak of disease in
its whitefish and perch. Despite the fact that there
had been no chemical treatment of Lake Mendota,
the general public blamed the Mendota fish deaths
on "sewage and chemicals." This is evidence of the
lack of confidence of the public in those responsible
for the conditions of the lakes and for the treatment
and disposal of sewage.
1933-35. The Dane County Board of Super-
visors established a County Parks Commission,
one duty of which was to control and improve the
condition of Lakes Waubesa and Kegonsa. The
Parks Commission was authorized to contract with
the Madison Board of Health for copper sulfate treat-
ment of Lakes Waubesa and Kegonsa. The Parks
Commission also appropriated a small sum to fi-
nance a study of the lower lakes. In 1935, the Lake
Kegonsa Protective Association was formed by
owners of property on the lake, with the expressed
purpose of "preventing the Madison Metropolitan
Sewerage District from converting Lakes Waubesa
and Kegonsa into their private privies." Members
-------
Madison's Lakes and Urbanization
13
of this Association then and later expressed them-
selves in earthy, powerful language; they were
action-minded, and were not interested in research
or explanations. They carried their problems to
the Governor and to the State Board of Health. One
member collected water from the Yahara River,
and from Lakes Monona, Waubesa and Kegonsa,
and mailed the samples to Dr. H. C. Bundeson, City
Health Commissioner of Chicago, for analysis.
This action exemplified the lack of confidence in
local authorities and in agencies of the State
government, and the belief that "someone from
outside" was needed to "get the answers and do the
job." Another very articulate citizen made a pro-
phetic statement: "There is only one way out —
pipe the stuff around the lakes." Oscar Mayer and
Company built and put into operation a plant for
treatment of its wastes that produced an effluent no
stronger than domestic sewage.
1936. The Dane County Board appropriated
funds to build locks designed to control the levels
of Lakes Waubesa and Kegonsa. It was believed
that maintenance of the "lower lakes" at the highest
possible levels would help remedy the algae nui-
sances. This same year, the report of an "outside"
laboratory that had been retained by the Lake Keg-
onsa Protective Association proved to be unaccept-
able because it showed that the Nine Springs sew-
age treatment plant was functioning efficiently.
The Nine Springs treatment plant was enlarged by
addition of an activated sludge treatment system
which was operated in parallel with the existing
facilities.
1937. The Burke sewage treatment plant was
closed and all sewage of the District was diverted
to the Nine Springs plants. The Dane County Board
established a County Sanitary District and a County
Conservation Committee. The latter was concerned
primarily with protection of fish and other wildlife
against the adverse effects of chemicals used to
control nuisance conditions in the lower lakes.
1938-40. The Southern Wisconsin Lakelands
Association, organized in 1939 to help promote
resort and recreational use of lakes in the southern
part of the State, added its support to anti-pollu-
tion and clean lakes efforts of the County Board.
The proposal was made by the County Board that
State and Federal aid should be obtained to build a
pipeline to carry effluent from the Nine Springs
plant to the Yahara River South of Stoughton.
1941-43. Through the efforts of the Southern
Wisconsin Lakelands Association, and representa-
tives from the Dane County Board, State Senator
H. A. Lewis of Boscoble, Grant County, introduced
a bill calling for revision of the Wisconsin anti-
pollution law. The Lewis bill would not permit the
effluent from the sewage treatment plant of a city
of 10,000 or more population to be introduced into
a lake of less than six square miles in area, or
into a stream within 15 miles of the city. In the
public hearings on this bill, strong opposition was
expressed by representatives of cities throughout
the State. Hence, Senator Lewis changed the bill
to apply only to cities of 45,000 persons or more;
to any lake of more than two square miles or less
than six square miles in area located within 10
miles of the treatment plant of the city or sewerage
district. Since the bill as changed applied only to
the Madison Metropolitan Sewerage District, oppo-
sition from other localities in the State was with-
drawn. In addition, the effective date of this leg-
islation was changed from "immediately" to Jan-
uary 1, 1943.
The Lewis bill, as amended, passed both houses
of the Legislature, but was vetoed by Governor Heil
in June of 1941. At that time the Governor said
that "I feel this matter should receive very care-
ful study and there should be a definite determina-
tion whether this particular disposal plant is the
cause of the trouble complained of in the lakes
near Madison". The Governor appointed a small
committee to study the problem and to report to
him immediately. The County Board stopped chem-
ical treatment of Lakes Waubesa and Kegonsa so as
to "shatter the public complacency" regarding the
lakes problem. Incidentally, copper sulfate was
then in short supply because of war needs.
The Governor appointed D. >V. Mead, Emeritus
Professor of Hydraulic and Sanitary Engineering,
as Chairman of his Committee, and included in its
membership Clifford Halverson of the County
Board, E. J. Tully of the State Board of Health,
J. L. Ferebee of the Milwaukee Sewerage Commis-
sion, and Dr. W. D. Stovall, Director of the Lab-
oratory of Hygiene of the State Board of Health.
This Committee recommended:
1. "That facilities be provided for the treat-
ment of the lakes with copper sulphate to abate the
nuisance odor caused by algae until a more satis-
factory solution can be arrived at. The city and
county should pay for this work.
2. "That immediate steps be taken to provide
the personnel necessary to begin an investigation
of the sources from which nitrogenous material,
phosphates, and other materials which act as food
substance for algae are contributed, and the amount
contributed from each source. This investigation
should last a year and be financed by the Metro-
politan Sewerage District.
3. "That the State provide sufficient funds for
the investigation of all the factors which are re-
sponsible for the growth of algae in the lakes in
the state, and that these funds be made available
to the State Board of Health,"
The County Board's reaction to this program,
conditioned by nearly 25 years of debate against
-------
14
ALGAE AND METROPOLITAN WASTES
State, Municipal and District agencies, was to rec-
ommend:
1. "That Dane County Officials ignore the Gov-
ernor's committee and go aheadwith a county pro-
gram to combat pollution of lakes Waubesa and
Kegonsa.
2. "That the Dane County Board appropriate
$3,500 to sponsor a chemical study of the pollu-
tion with the view of using the information gained
as a basis for court action to prevent the Madison
Metropolitan Sewerage District from continuing to
dump effluent in the lakes.
3. "That the County Board provide an appropri-
ation to spray the lakes with copper sulphate until
such time as conditions are corrected."
In June of 1942, the problems of the District,
the Governor's Committee, and the County Board
were further complicated by the decision of the
Federal Government to recondition and to use the
Burke plant to treat sewage of Truax Field, an
Army Air Corps base located to the North of Madi-
son.
At the end of June, 1942, Dr. Clair N. Sawyer
was engaged by the Governor's Committee to con-
duct the sanitary chemical studies needed to imple-
ment the Committee's program. The County Board
was unimpressed, and decided to conduct its own
survey.
The Governor's Committee made a preliminary
statement of the problem as follows:
"After considerable study and observation of the
local lake problem, the conclusion was reached that
the major part of the offensive conditions and sten-
ches produced are a direct result of the accumula-
tions of algal growths in advanced stages of
decomposition. It was further concluded that the
stimulation of such algal growths was not a direct
result of pollution of the lakes in an ordinary sense,
but rather was related to the fertilization of the
water by enrichment with minerals such as nitro-
gen, phosphorus, potassium, and possibly other
fertilizing elements and compounds. These con-
clusions were based on many considerations of
which the following are a few:
1. "Algal problems are mostly prevalent in
shallow lakes receiving drainage from rich agri-
cultural areas and/or urban communities.
2. "Chemical analysis of algae show nitrogen
to be one of the major component elements with
phosphorus a minor element.
3. "Stream pollution surveys where domestic
sewage is involved have shown zones of increased
biological productivity to occur in regions of the
streams below the sewer outfall.
4. "The science of hydroponics is based on the
principles of ordinary agricultural fertilization
applied to water.
5. "Fertilization of lakes is being practiced in
some areas with ordinary commercial fertilizers
to stimulate greater plankton growths which in turn
support fish populations.
6. "Some authorities place considerable impor-
tance on agricultural land drainage and thereby in-
fer that there is a cor relation between fertilization
and algal blooms.
7. "The growths causing nuisances in inland
lakes are largely members of the plant kingdom.
All ordinary plant growth is interdependent upon
a substrate containing nitrogen, phosphorus, potas-
sium, and other elements.
8. "Sewage of domestic origin although highly
purified by modern methods of treatment remains,
after treatment, abundantly rich in phosphorus,
nitrogen, potassium, and possibly other elements
of significance.
9. "High concentrations of certain fertilizing
elements which accumulate in lake waters during
the winter months are rapidly depleted in the sur-
face waters during the growing period of warm
weather."
SCOPE OF SURVEY
"Although the Committee was charged with
studying the pollution of Lakes Waubesa and Keg-
onsa, it was deemed advisable to include Monona
in the survey as it also is one of the lakes causing
considerable trouble. It was anticipated that sew-
age effluent from Truax Field would be added to
Lake Monona during the course of the survey, and
this was further reason for studying the lake as
diligently as the others mentioned above. Lakes
Wingra and Mendota were relegated to minor in-
vestigations because of their normally good behav-
ior so far as stench producing ability is con-
cerned."
METHODS OF SURVEY
1. "Measurement of fertilizing elements.
2. "Biological studies.
3. "Observations on other southeastern Wis-
consin lakes.
4. "Research in laboratories."
-------
Madison's Lakes and Urbanization
15
SIGNIFICANCE OF CHEMICAL OBSERVATIONS
"The soluble phosphorus, potassium, and inor-
ganic nitrogen content of tributary waters are con-
sidered more indicative of its fertilizing potency
since these represent readily available forms of
the elements. The organic nitrogen content is con-
sideredof secondary importance because the nitro-
gen is in a form which is made available incom-
pletely and slowly.
"The suspended organic matter contained in the
tributary waters is given consideration mainly for
the reason that certain groups have placed consid-
erable emphasis on the solids in the Madison sew-
age plant effluent as being the cause of the stenches
and unsightly conditions along the shores of Lake
Waubesa".
Early in 1943, Dr. James B. Lackey of the
United States Public Health Service studied the
problems faced by the Governor's Committee, con-
ducted investigations independently and in coop-
eration with Dr. Sawyer, and got the County Board
to suspend its proposed program of investigations
that would have duplicated the efforts of the Gover-
nor's Committee.
Senator Lewis reintroduced his anti-pollution
bill, but changed the date of enforcement to: "one
year after the termination of the present war as
proclaimed by the President or Congress." The
bill passed, and was signed by Governor Goodland
in April, 1943.
The first report of the Governor's Committee,
prepared by Dr. C. N. Sawyer, with consultation
and advice of Drs. J. B. Lackey and A. T. Lenz,
and covering investigations made from July 1942
to July 1943, was published in mimeographed form.
Excerpts from this report follow:
THE GOVERNOR'S COMMITTEE - SUMMARY
AND DISCUSSION
"This study shows that a large amount of nu-
tritive and pollutions! material is being added to
the lakes from various sources. It further shows
that the inorganic nitrogen contributed from these
sources and from deposits on the lake bottom is a
critical substance with relation to lake blooming.
It appears that a significant reduction in the amount
of the nutritive material reaching the lake would
reduce the frequency and density of lake blooms.
This should not be interpreted to mean that algae
growths in the lake will cease or that blooms will
be completely eliminated. While the Madison sew-
age effluent contributed by far the largest amount
of this substance to Lake Waubesa, 76.9%, it is
not the sole contributor. Lake Kegonsa receives
67.2% of its inorganic nitrogen and 88.6% of its
organic nitrogen from Lake Waubesa. In the case
of Lake Monona the storm and industrial sewers of
Madison are large contributors.
"As our studies have progressed, it has been
more and more apparent that the Madison sewage
disposal plant is functioning efficiently. The pop-
ular impression that this plant is not purifying the
sewage in a satisfactory manner is not correct.
There is a question as to the capacity of this plant
to handle the sewage and wastes from the metro-
politan area if all of that waste which is now going
into Lake Monona by various industrial and storm
sewers was passed through the plant.
". . The nuisance, as the results of this inves-
tigation show, is due to the excessive growth of
algae, and the odor created by it. Even though the
Madison sewage plant effluent is not directly the
cause of the odor that comes from the lakes in the
summer, the data clearly show that it is the larg-
est contributor of nutritive material upon which
algae feed."
SOME EXCERPTS REGARDING TECHNICAL AS-
PECTS OF THE SURVEY
"The question was, is there a critical relation
between these substances (nitrogen and phosphorus)
and the frequency of lake blooming and the density
of the bloom? The question bore directly upon the
quantitative relation between these substances, in-
organic nitrogen and phosphorus, and the sources
from which they are contributed to the lake. In
other words, if these nutritive materials or any one
of them could be shown to be critical, that is, nec-
essary in certain amounts to support algal growth,
then the amounts contributed from various sources
would be helpful in determining the significance of
the various sources to the blooming of the lakes."
"In June, before a heavy bloom, the inorganic
nitrogen in solution in the surface water of Lake
Waubesa amounted to almost 1.0 part per million.
After the algal growth had become so dense that it
was recognized as a scum on the lake surface, the
inorganic nitrogen in solution in the surface water
was found to have been reduced to 0.3 parts per
million. In other words, the algae in order to grow
to the density attained between these dates in June
used up more than half of the inorganic nitrogen in
solution in the surface waters. It is important to
note that this reduction of inorganic nitrogen took
place in water where the algal growth was most
dense.
"The organic nitrogen represents decomposable
matter whereas inorganic nitrogen does not. The
organic nitrogen is considered pollutional because
it undergoes decomposition in the lake. The inor-
ganic nitrogen is the result of completed decompo-
sition. It is stabilized nitrogen and is therefore
considered to be non-pollutional, nutritive materi-
al."
-------
16
ALGAE AND METROPOLITAN WASTES
"Raw sewage is by-passed from the city sewers
into Lake Monona during periods of mechanical or
prolonged electrical failure and during periods of
heavy rainfall. The main exit for the sewage to
Lake Monona is via the Burke Plant outfall sewer.
The sewage enters this sewer at the No. 1 pumping
station located at East Johnson and First streets."
"During the course of the survey no untreated
sewage was found in the effluent from the Madison
Sewage Plant. A total of 116 separate samples
were tested and the results showed the effluent to
be of better quality on an average than the effluents
produced by plants of similar design and facilities
operating elsewhere in the United States."
CONCLUSIONS BT OR. CLAIR N. SAWYER
1. "The biological productivity of the local lakes
is a function of the loading of inorganic nitrogen on
each lake.
2. "The soluble phosphorus content of water
maybe a factor in limiting the rate of biological
activity and in determining the nature of the
growths when its concentration drops below 0.01
part per million.
3. "Drainage from improved marsh land is ap-
proximately two to three times as rich in inorganic
nitrogen as drainage from ordinary farm lands.
4. "On the basis of agricultural drainage, Lake
Mendota should be from two to three times as pro-
ductive of biological life as Lake Monona and
Waubesa and Kegonsa should be some one to two
times as productive as Mendota. Lake Wingra on
the basis of lake volume should be the most pro-
ductive of alL
5. "High biological productivity and nuisance
conditions do not always occur simultaneously.
For example, Lake Wingra, a very shallow lake,
does not develop offensive conditions in spite of
its high productivity. The reason for this has not
been ascertained but may be related to its lower
phosphorus content."
THE FUTURE IN REGARD TO THE MADISON
LAKES PROBLEM AS SEEN BY THE GOVER-
NOR'S COMMITTEE AT THE TIME OF THE
FIRST "SAWYER" REPORT
"Any solution of the local problem must take
into consideration developments in the near and
perhaps distant future. The City of Madison and
its suburbs has increased its population by 9,430
persons during the past three years. The day is
not far distant when this center of government and
education will reach a population of 100,000, men
it will be 150,000 etc. It is reasonable to assume
that industrial growth will increase similarly.
Thus it would be logical to conclude that in the
course of another 50 or 100 years the contribution
of pollution and fertilization by the metropolitan
area will be doubled. During a similar period, it
is doubtful whether the contribution from unnatural
sources will increase materially. In the year 2000
A. D., the contribution of inorganic nitrogen from
the city may approximate 85% of the total contri-
bution to Lake Waubesa, and the soluble phospho-
rus 94% unless marked changes are made in
current practices."
DISCUSSION OF THE REPORT FINDINGS BY DR.
LACKEY, UNITED STATES PUBLIC HEALTH
SERVICE
"There can be no dispute as to the fact that
there is heavy fertilization of the lakes by the ma-
terials poured in. The use of sewage for fertilizing
lakes in Germany is too well known; the use of
sewage sludge as a fertilizer is widespread in this
country. These two facts constitute proof of the
fertilizing value of sewage, the first as to raw sew-
age, and the second as to solids and mineralized
portions of treated sewage. Snell gives calculated
monetary values of human excreta as fertilizer and
points out that 85% of the fertilizer value of excreta
is contained in the urine and is therefore lost in
the effluent of ordinary sewage treatment plants.
In other words, it passes on through the plants and
out with the effluent, not being incorporated into
the sludge. This fact should at once render unten-
able any argument that by-passed sewage solids,
if found, are a major cause of blooming in the
lakes.
"It would be well to remember that any removal
of nitrogen and phosphorus from effluent waters to
these lakes might not have an immediate remedial
effect of great magnitude. The layman might be
inclined to take such a view and be sharply critical
if obnoxious blooms did not stop at once following
removal or further treatment of the effluent. There
are, however, at least three reasons why the lakes
might continue to bloom for a considerable period:
"First - There may be other chemical factors
closely related to blooming in addition to nitrogen
and phosphorus contributed by sources enumerated
herein. These considerations do not obviate the
fact that present contributions of nitrogen and
phosphorus are readily available forms and exten-
sively used; probably a first choice1 relationship
exists. But should these sources be removed,
other possible sources should be remembered.
"Second - The bottom deposits which have been
building up over a long period of time should be
considered. One component part of these results
from sedimentation of dead plankton bodies; another
results from the annual death and decay of rooted
lake vegetation and attached algae (which at times
form very heavy growths both on the rooted plants
-------
Madison's Lakes and Urbanization
17
and on the bottom); and a third consists of sludge
from sewage treatment plants and sewers. The
depth of such deposits in the Madison lakes needs
more investigation, but in places at least,
according to Birge, Juday, and March, depths of
five meters have been encountered. This rich or-
ganic stratum is continually producing nitrates,
ammonia, and carbon dioxide; furthermore, any
of the gas bubbles, including methane and nitrogen,
rising from It afford excellent absorption of such
gases before ttie surface is reached, as well as
carrying up some bottom materials. One has but
to observe the black sludge brought up by gas bub-
bles at the Johnson Street sewer to realize this
last factor is an important one. Any mud or sludge
deposits, therefore, constitute reservoirs of nu-
tritive material from which the overlying water is
being continually enriched.
"Third - Another consideration has to do with
the standing crop of weeds in the lakes. In protec-
ted places the surface may be completely covered
with floating weeds, and they provide vast amounts
of food as they die and decay. That they tend to
cause heavy plankton production is well shown by
some of the other southeastern Wisconsin lakes."
1944. The second report of the Governor's
Committee, covering work done from July 1943
to July 1944, was prepared and published as before.
This report "strengthened the conviction that in-
organic forms of nitrogen and phosphorus are the
main factors in providing fertilizing elements for
algal blooms." The report summarized the lake
studies made as follows:
1. "Those lakes having large drainage areas
and/or receiving domestic sewage or sewage plant
effluents are most productive of plankton.
2. "Those lakes receiving sewage or sewage
plant effluents are most productive of nuisance
blooms, mainly due to blue-green algae.
3. "Inorganic nitrogen and phosphorus were
found to be critical factors in the productivity of
lakes, the .inorganic nitrogen appearing to be a
limiting factor in regard to the amount of growth
which could be produced and inorganic phosphorus
acting largely as a governor upon the rate at which
growths occurred.
4. "For Lake Waubesa, the most heavily fer-
tilized lake in the survey, during the year 1942-43
at least 65% of the inorganic nitrogen and 89% of
the inorganic phosphorus entering the lake was de-
rived from non-agricultural drainage. The contri-
bution of inorganic nitrogen and phosphorus in the
effluent and drainage reaching this lake is equiva-
lent to that derived from a population of 140,000."
The second report also summarized significant
laboratory and experimental studies, and made
proposals for future work. However, the program
of the Governor's Committee was terminated in
July of 1944 because the Legislature failed to pro-
vide funds for continuation, and the Madison Met-
ropolitan Sewerage District, having contributed
$20,000 to the two-year program, decided to end
its support.
In 1947, the late Thomas E. Brittingham, Jr.,
contributed $50,000 to the University of Wisconsin
to support a five-year study of lake pollution pro-
blems and of possible improvements in methods of
sewage treatment His contribution was supple-
mented by funds from the Wisconsin Alumni Re-
search Foundation, the Lake Mendota Association,
Oscar Mayer and Company, the National Insti-
tutes of Health, the Office of Naval Research, and
the Wisconsin Conservation Department The stud-
ies made since 1947 are too numerous and exten-
sive to be reported here in detail. They have been
aimed at finding explanations for nuisance condi-
tions, improvement of methods of sewage treat-
ment, thorough surveys of normal and of polluted
lakes and streams, and the development of manage-
ment practices which might be used to abate nui-
sance conditions in lakes and streams. Results of
some of these studies will be reported to this Sem-
inar. It is our firm belief that out of such basic
studies will come the facts needed for improved
practices and for practical management proce-
dures.
In 1947, Dr. A. F. Bartsch reported studies that
he had made during 1945-47, with the help of G. W.
Law ton, on Lake Mendota. These investigations
were made to get information needed by the State
Committee on Water Pollution to supplement the
1942-1944 studies of the Governor's Committee.
Dr. Bartsch's conclusions were that the "relative
infrequency of blooms in Lake Mendota, as com-
pared with the remaining three lakes, is directly
related to its lower rate of nutrient receipt. At
the same time, it is to be noted that Lake Mendota,
receiving non-urban drainage primarily, has suf-
ficient supplies of nutrients available to allow the
production of objectionable algal blooms when
other environmental factors are favorable." The
report also said that "algal conditions in Lake
Mendota are likely to become more severe unless
necessary steps are taken to decrease the rate at
which nutrients enter the lake."
Also, in 1947, the Burke treatment plant,
which had been shut down when Truax Field was
closed in 1946, was returned to operation by the
Metropolitan Sewerage District with the permis-
sion of the State Board of Health. This action was
taken because of the need to build a new pressure
sewer to carry sewage around the East end of Lake
Monona to the Nine Springs plant; the city's volume
of sewage exceeded the capacity of the existing in-
terceptors and pressure sewers.
-------
18
ALGAE AND METROPOLITAN WASTES
Return of the Burke plant to operation, and the
impatience of the increasing population living on
the shores of lakes Waubesa and Kegonsa, led in
1948 to the formation of the Southern Wisconsin
Anti-pollution Federation. The chief objectives of
this organization were: To combat pollution of
lakes and streams in southern Wisconsin; to en-
courage enforcement of existing laws on pollution;
and to secure additional anti-pollution legislation.
The thirteen organizations belonging to this Fed-
eration claimed a membership of 13,000. The Fed-
eration persuaded Governor Rennebohm to request
the State- Committee on Water Pollution to order
the Madison Metropolitan Sewerage Commission to
present preliminary plans for compliance with the
"Lewis" law when it was to become effective. This
order was issued jointly by the State Board of
Health and the State Water Pollution Committee in
July of 1948. By a 2 to 1 vote, the Commission
voted to petition the State agencies for a public
hearing on their joint order.
The public hearing, lasting four days, was held
by the State Committee on Water Pollution. The
joint order was upheld, and the Metropolitan Sew-
erage District was ordered to submit preliminary
plans by December 31, 1948 for diversion of the
Nine Springs treatment plant effluent to a river not
less than ten miles from the plant
The District appealed the affirmation of the joint
order, and the appeal was heard in Circuit Court,
Judge A. W. Kopp presiding, in January of 1949. In
March, Judge Kopp filed his decision which de-
clared void the order of the State agencies, based
on provisions of the Lewis act, because the Presi-
dent or Congress had not proclaimed that World
War n had ended.
The decision of the Dane County Circuit Court,
Judge Kopp presiding, was appealed to the State of
Wisconsin Supreme Court In August of 1951 the
Supreme Court upheld the order of the State Board
of Health and the State Committee on Water Pollu-
tion. Consequently, these agencies reaffirmed
their previous order, and the Madison Metropolitan
Sewerage District was thus forced to prepare plans
for diversion of its effluent to a river not less than
ten miles from its Nine Springs plant
Much more could be written concerning events
of the past nine years, but additional information
is believed to be unnecessary. Effluent from the
Nine Springs Sewage treatment plant is now diver-
ted via the Badf ish Creek to the Yahara River South
of Stoughton.
Will it'be possible to extend the limits of the
Madison Metropolitan Sewerage District, or to
create new sewerage districts to treat the sewage
of the growing populations surrounding Madison's
lakes? Can methods of sewage treatment be im-
proved to the degree necessary to produce an efflu-
ent free from plant nutrients? Can lake and
stream management practices be developed that
will control nuisance conditions, but will not harm
fish or aquatic life?
When we know the answers to these questions
we shall be able to answer the question raised in
this paper's title.
*************
Note: References to papers or reports cited are
not listed because these documents are not avail-
able in libraries. Much general information has
been taken from the Master's thesis, entitled "The
Madison Lakes Problem", by James J. Flannery,
University of Wisconsin, Department of Political
Science, 1949. Views or opinions expressed are
those of the author (W. B. Sarles) and do not carry
any official significance.
ALGAE BLOOMS IN LAKE ZOAR, CONNECTICUT
RICHARD J. BENOIT
Research and Development Dept, General Dynamics Corp., Groton, Conn.
and
JOHN J. CURRY
Water Resources Commission, Hartford, Conn.
The authors gratefully acknowledge the contributions of Dr. R. L. Holmes, Dr. S. L. Wilson, M.
Hupfer, J. Masselli, N. Masselli and B. Parker.
In 1919, when Lake Zoar was first formed by
Stevenson Dam on the Housatonic River, there was
no algae problem. As the years passed, algae in
the lake increased and by 1947 they were so plenti-
ful that a serious nuisance was created for lake-
side property owners. The problem was first re-
ferred to an ad hoc committee comprising the ag-
ricultural experiment station, the water commis-
-------
Algae Blooms in Lake Zoar
sion, the fish and game department, and the
health department, all of the State of Connecticut.
Although no funds were specifically appropriat-
ed, the Connecticut Agricultural Experiment Sta-
tion, being a research agency, conducted a general
study of the problem with the support of the other
interested agencies. In July 1954 a research study
was authorized.
PHYSICAL ASPECTS
Lake Zoar was formed behind a run-of-the-
river hydroelectric plant built by the Connecticut
Light and Power Company in 1919. The drainage
area of the Housatonic River at this point is about
1545 sq. mi. This company also operates a pump
storage facility at Lake Candlewood. Although
Lake Candlewood is not on the Housatonic River
proper, the water in the Lake is Housatonic River
water pumped to it for storage (Figure 2).
Figure 2. LAKE SYSTEM OF THE HOUSATONIC
RIVER AREA IN CONNECTICUT.
Before entering Connecticut, the Housatonic
River flows through the western end of Massachu-
setts where its waters are used extensively by in-
dustries in the Pittsfield region. The processes of
nature almost completely eliminate any pollution in
the water by the time it reaches the Connecticut
state line. As it flows through Connecticut, the
river receives little pollution before it reaches
New Milford. It is an esthetically satisfactory
stream and is used extensively for fishing and re-
creation. However, there are two aspects of the
water quality which are unusual for streams in this
area: (1) Flowing through limestone formations,
the water is hard (75 ppm.) compared with other
streams in the state; (2) total solids average about
170 ppm., whereas other esthetically acceptable
waters in the state average 65 ppm.
LIMNOLOGY
Below Boardman Bridge, the quality of the
Housatonic River changes materially. The major
factors causing this change were: (1) raw sewage
pollution from the Town of New Milford, totalling
approximately 200,000 gal. per day; (2) a quantity
of organic industrial wastes from the hat manufac-
turing industry in Danbury and Bethel, and the
trickling filter effluent from the Danbury sewage
treatment plant carried by the Still River. The
Shepaug, Pomperaug, and Pootatuck Rivers are
relatively large-sized tributaries carrying no
serious pollution. The Connecticut State Health
Department has eliminated practically all pollution
from development adjacent to Lake Zoar.
The lake volume is estimated to be 42,800 acre-
ft. and the average flow of the Housatonic River at
Lake Zoar is approximately 2,500 cfs, indicating
that the lake waters are completely replaced every
9 days. During the algae season, the low monthly
flow (550 cfs) of August and September results in
a replacement of the lake volume every 40 days.
However, power production causes great fluctua-
tions in the rate of flow.
Preliminary investigation showed that during
late July, as the seasonal reduction of river flow
becomes apparent, it might be possible for strati-
fication to develop. More detailed studies revealed
that the stratification was transient and did not
exist to an extent that would affect the problem.
TYPES OF ALGAL BLOOM
Although original complaints had specified root-
ed aquatic weeds as the cause of the disagreeable
condition, algal blooms were an equally objection-
able part of the problem. By mid-July Hydrodictyon
was found in great quantities in all the shallow,
quiet waters where it was intimately associated
with aquatic weeds such as Elodea, and Cerato-
phyllum. Large mats moved fro m place to place
during changes of flow. Early in the season large
quantities of Microcystis and Anabaena also ap-
peared (Curry and Wilson, 1955) usually in the up-
per four feet of lake water.
The most direct approach would have been the
addition of algicides such as copper sulfate. It was
calculated that a single treatment would cost
-------
20
ALGAE AND METROPOLITAN WASTES
$15,000. Laboratory tests with fungicides, bacteri-
ocides such as quarternary ammonium compounds,
and a wide variety of experimental compounds
showed some promise. Almost all of the materials
tested are included in the list reported by Palmer
(1956).
PHOSPHATE ASPECTS
Repeated tests indicated the level of nitrates
and phosphates in Lake Zoar was similar to that
noted in other lakes at the time of blooming. The
well-known ability of some blue-green algae to fix
atmospheric nitrogen indicates that the addition of
phosphate alone might result in intense blooms in a
lake like Zoar where all other aspects of the en-
vironment are favorable for bloom development.
Phosphorus concentrations varied from 12 to 41
parts per billion (ppb) and averaged about 25 ppb.
(Table 2).
Table 2. CONCENTRATIONS
IN LAKE ZOAR
OF PHOSPHORUS
Sample date,
1950-51
Sept. 2
Oct. 2
April 27
May 7
June 4
June 26
July3
July 13
July 18
July 23
July 30
Aug. 10
Flow,
c.f.s.
372
462
3,670
1,880
1,270
1,100
1,060
1,250
809
2,300
987
312
Phosphorus,
p.p.b. Ib./day
33
15
30
41
26
22
25
31
12
18
22
28
66
25
594
416
178
131
143
209
52
224
117
47
Average 1,289
25
183
The volume of phosphates entering the streams
in Connecticut from sewage outlets has reflected
the increased use of synthetic detergents. Most of
the sewage that is discharged to the Housatonic
River in this region is given a high degree of treat-
ment Such treatment oxidizes the organic phos-
phate to the mineral form making it available as
fertilizer to the aquatic plants.
The bulk of the phosphates fed into Lake Zoar
come from the Still River and the Pootatuck River.
The estimated amounts of phosphorus coming into
Lake Zoar from various sources are shown in Ta-
ble 3. The values from the analyses and computa-
tion check well in view of the fact that many more
individual samples were averaged for main stream
values than for tributary values. Contributions of
the Still River to the total phosphorus available in
Lake Zoar account for about 40 per cent of the to-
tal. If this source were eliminated, the phosphorus
concentration in Lake Zoar would be reduced to
15 ppb. The critical concentration of phosphorus
for blue-green algae growth is believed to be about
10 ppb. A reduction in phosphate should have an
important effect on the algae bloom.
Table 3- SOURCES OF PHOSPHORUS
BUTORY TO LAKE ZOAR
CONTRI-
Location
Housatonic River at Boardman
Bridge
Sewage from New Milford
Still River.
Housatonic River at Rt. 133
Shepaug River
Housatonic River above Pootatuck
Pootatuck River
Pomperaug River
Housatonic River at Lake Zoar
P.
Ib./day
98 a/
9
77
232 a/
2
167 a/
10 "
5
183 a/
Cumulative P,
Ib./day
184 /
186 /
201 /
a/ Based on analyses and flow rates at these points.
Lake Candlewood is filled with water taken from
the Housatonic River immediately below Boardman
Bridge. The soluble phosphorus content of the
Housatonic River at this point during periods of
high water is probably less than 30 ppb. However,
Lake Candlewood produces no algal bloom of any
consequence. Determinations made during the
algae season indicated phosphorus concentrations
of only 8 ppb. It is suggested that the phosphate is
utilized early in the season by diatoms which even-
tually sink to the bottom, removing phosphate from
surface waters. Since little additional water is
pumped into Lake Candlewood during the summer
the supply of nutrient is not renewed as it is in
Lake Zoar.
•
CHANGING CONDITIONS
During the past eight years pollution control
agencies have taken action which may affect the
problem in the future. The city of Danbury has en-
larged and renovated its sewage treatment plant
so the quantities of sewage by-passed to the Still
River are materially reduced. Six hat manufactur-
ing industries which discharge organic wastes di-
rectly to the Still River have constructed screens
and settling basins which keep much organic mat-
ter from entering the stream. The town of New
Milford and two state institutions in the watershed
have constructed sewage treatment facilities.
A dam has been constructed on the Housatonic
immediately below the confluence of the Shepaug,
creating another lake of similar characteristics to
Zoar. The new lake is in a position to intercept
the phosphorus load originating in the Still River.
Already there have been reports of a degree of
blooming but no lessening of bloom in Zoar has
been noted.
-------
Algae Blooms in Lake Zoar
21
PHOSPHATE REMOVAL
The physiographic features at the sewage treat-
ment plant of the State Hospital at Newtown on a
branch of the Pootatuck River offered an ideal op-
portunity to study the removal of phosphate by the
method of Lea, et al (1954) on a pilot plant scale.
This method uses alum and lime to precipitate
phosphate, with alum being largely recovered and
recycled. A pilot plant was constructed on the site
(see figure 3) with a special legislative appropria-
The ponds were 400 ft. long by 100 ft. wide with
depths from 4.5 to 8.0 ft. The pilot plant and pond
design enabled the simulation of conditions in Lake
Zoar and provided sufficient flexibility for a varie-
ty of controlled experiments. Both ponds were
filled with brook water and one of them was inocu-
lated with bloom algae from Lake Zoar. Both de-
veloped a good growth of algae, but no field ex-
periments were undertaken before the floods of
August and October 1955. The August flood des-
0AM
BROOK
Figure 3. SCHEMATIC DIAGRAM, ALGAE CONTROL PILOT PLANT.
Fairfield State Hospital, Newtown, Connecticut
tion. Two ponds were built for use in field experi-
ments to establish the critical levels of phosphate
for algae blooms. One pond was to receive sewage
or secondary sewage effluent and the other was to
serve as a control and receive only creek water or
secondary effluent from which phosphate had been
stripped.
Preliminary studies were made in 4000 gal.
batches in the pilot plant under conditions of 10-
nrinute mixing and 2-hour settling. Using 200 ppm
alum, a removal of better than 95 per cent from
effluents having a phosphorus content of 3.5 to 4.5
ppm was accomplished.
troyed the pond embankments and the piping from
the pilot plant and brook to the ponds.
Negotiations to rebuild the plant were carried
out with the United States Engineering Department
under their flood rehabilitation program and the
project was not back in operation until 1957. Dur-
ing this reconstruction, it was decided to reduce
the size of the lagoons to make them more amena-
ble to control. Before any substantial program
could be carried out with the restored facilities,
an unexplainable increase in the phosphate content
of the brook was indicated by analyses from an
outside laboratory. The reliability of these analy-
-------
22
ALGAE AND METROPOLITAN WASTES
sea had to be accepted in the absence of evidence
to the contrary, but no source of the phosphate
has ever been established. Nevertheless, the ad-
visability of carrying on the projected field pro-
gram was questionable since the authorized time
of the expenditure was running out It was there-
fore deemed prudent to conclude the project while
there was still time to make close down expendi-
tures. Such a decision was appropriate because
serious doubt had been cast on the feasibility of
keeping phosphate in waterways below the critical
level for blooms simply by removing phosphate
from specific sources with some suitable process.
NEW POSSIBILITIES FOR ALGIC1DE TREAT-
MENT • • . -.
Also relevant to the decision was the new oppor-
tunity for chemical treatment afforded by the con-
struction of the Shepaug Dam. For the volume of
water involved, chemical treatment no doubt will
always be very costly, but can be more easily jus-
tified for two large lakes with twice the recreation-
al development. Since Lake Zoar is downstream
from the new lake behind the Shepaug Dam, it will
probably only be necessary to apply algicides to
the upper lake to achieve a degree of control in
both lakes. Use of the Candlewood Lake discharge
as a mixing facility might provide economically
attractive chemical treatment for both lake devel-
opments.
REFERENCES
Curry, J. J. and S. L. Wilson, 1955. Effect of sew-
age-borne phosphorus on algae. Sewage and Indus-
trial Wastes 27(11): 1262-1266.
Palmer, C. M., 1956. Evaluation of new algi-
cides for water supply purposes. Jour. Amer.
Water Wks. Assoc. 48:1133-1137.
Lea, W. L., G. A. Rohlich and W. J. Katz, 1954.
Removal of phosphates from treated sewage. Sew-
age and Industrial Wastes 26(3): 261 -275.
KLAMATH LAKE, AN INSTANCE OF NATURAL ENRICHMENT
HARRY K. PHINNEY
Associate Professor of Botany, Oregon State College, CorvalUs
and
CHESTER A. PEEK
Research Assistant, Oregon State College
The City of Klamath Falls is located in south
central Oregon just north of the California State
line. The City is named for a turbulent, mile-long
stretch of-water, now known as the Link River,
that lies completely within the city limits. This
short river flows between Upper Klamath Lake
at the north of the City and Lake Ewauna at the
south. Lake Ewauna in turn drains into the Kla-
math River. In addition to the annual mean flow
of LI million acre-feet passing through the Link
River, there is a flow of more than 200 thousand
acre-feet that leaves Upper Klamath Lake by way
of the "A" canal of the Klamath Reclamation Pro-
ject's irrigation system (Stanley, 1954).
A flow of water of this magnitude would appear
to be adequate to supply dilution water for the dis-
posal of wastes from a population of 35 thousand.
Despite the fact thaLthere is neither urban devel-
opment nor heavy industry on the Upper Klamath
Lake drainage system, and that mere are far more
acres of wild than of agricultural land within the
basin, the Link and Klamath Rivers carry an or-
ganic load so heavy as to present serious problems
even before the discharge of the City's wastes.
A brief summary of geographical information
and a map (Fig. 4) are presented as an aid in under-
standing the scope of this problem. The total
area of the Upper Klamath Lake Basin that drains
into the Klamath River in Oregon is approximately
7450 square miles. The major source of water
flowing into the Klamath River is Upper Klamath
lake, which lies in the southwestern portion of the
Upper Klamath Basin. Upper Klamath Lake is
approximately 23 miles long and 5 miles wide and
at the northern end joins Agency Lake, which is
approximately 5 miles long by 3 miles wide. The
mean depth of Upper Klamath Lake is- eight feet
and the maximum depth approaches sixty feet.
The Lake is supplied by a major tributary, the
Williamson River, as mjell as a number of smaller
streams. The Williamson is fed by the Sprague
and the Sycan Rivers. Agency Lake receives flow
from Wood River and the Seven Mile Canal besides
numerous small streams. The Klamath Basin was
formed by block faulting followed by local volcanic
activity. To the north of Agency Lake, toward the
-------
Klamath Lake, Natural Enrichment
23
.. L
_i
Figure 4. MAP OF THE UPPER KLAMATH LAKE
BASIN IN OREGON.
Crater Lake-Mount Mazama site, are vast pumice
fields with much subsurface drainage into, or at
least in the direction of, the lakes. In the bottom
of both lakes and along both margins of Upper Kla-
math Lake are innumerable springs, some warm,
some mineral, some of artesian nature, that feed
into the lakes along multitudinous fault joints.
Outflow from Upper Klamath Lake is regulated
by a low dam on the Link River, constructed by the
Bureau of Reclamation in order to utilize the Lake
as a reservoir. Since 1917 the dam has been oper-
ated by the California-Oregon Power Company to
maintain the surface of the Lake between elevations
of 4143.3 and 4137.0 feet. This allows an active
storage capacity of 483 thousand acre-feet, which
is a little more than half the present total storage
capacity of the Lake basin.
The discharge of the Link River flows through
shallow Lake Ewauna and into the Klamath River.
The Klamath River flows through approximately 16
miles of flat land to Keno, Oregon, where it enters
a 236 mile long canyon that it has cut through both
the Cascade and the Coast Ranges, debouching into
the Pacific Ocean at Requa, California.
THE BIOLOGICAL PROBLEM
For at least sixty years the algal populations
of Upper Klamath Lake have been sufficiently large
to cause comment and speculation as to the cause
and effects of the growth. During the summer
months the Lake has been unsightly and has had an
offensive odor. The discharge from the Lake has
contained a heavy organic load, high in nitrogen.
This has caused portions of the upper reaches of
the Klamath River to become periodically anaerobic
despite the super saturation with oxygen that occurs
at the exit from the Lake during daylight hours.
Because of this situation a serious problem has
arisen in the disposal of metropolitan wastes.
From the correspondence files of the Bureau of
Reclamation comes evidence of early concern for
this situation. As early as January 1906, J. B.
Lippencot, Supervising Engineer for the then Re-
clamation Service (now Bureau of Reclamation)
wrote from his Los Angeles office in part: "...I
wish to call your attention to the fact that these
waters are filled with some sort of organic matter,
either animal or vegetable, so that they have a
decided green appearance. They are cutting up ice
now that has been formed from these waters, and
we will probably be asked to use this ice next sum-
mer. Last summer we were troubled a great deal
up there with stomach complaints. For that rea-
son I am somewhat interested in the sanitary
analysis of the water. This same material in the
water appears to have some fertilizing proper-
ties...."
Analyses were made at this time by the U. S.
Geological Survey office in Berkeley and interpre-
ted by them as follows in June 1906: "...the
amounts of nitrogen found as free and albuminoid
ammonia seem rather large, but as you say there
is no possibility of sewage contamination, this
would not necessarily condemn the water. The
manner in which these ammonias made their ap-
pearance and the comparatively small amount of
oxygen required point toward this organic matter
being of vegetable origin."
On the 25th of August 1928 the Oregon State
Board of Health reported to the Bureau.on samples
from the Link River as follows: "...You will note
that our results showed no colon bacillus and
therefore no sewage contamination in the water.
The dissolved oxygen in the water was between 72%
and 80% of saturation...
"...The murky condition of the water is princi-
pally due to an algae (sic) or watergrass growth.
This algae was found to be mostly spirogyra with a
few ulothrix and chroococcus present..."
The year 1933 saw a study of a problem arising
from enormous midge populations in the Lake and
an effort was made to associate the midge with the
plant problem. At this time the plankton algae
were identified by Dr. E. I. Sanborn of Oregon State
College as: Anabaena spiroides var. crass a,
Pediastrum sp., Melosira sp., Navicula sp.,
Microcystis sp. and Aphanizomenon sp.
The study that has provided most of the infor-
-------
24
ALGAE AND METROPOLITAN WASTES
mation reported here was initiated in 1955. It
represented an effort to define the problem and to
establish cause and effect relationships wherever
possible with the hope that ultimately information
might be forthcoming that would make possible
realistic control measures.
Although this survey has named 29 species of
algae as occurring in the plankton with some sea-
sonal regularity, four species of the Cyanophyta
(Aphanizomenon n. sp., Gloeotrichia echlnulata,
Anabaena circinalis, Anacystis (Microcystis or
Polycystis) aeruginosa), and two species of dia-
toms (AsterioneUa formosa and Melosira sp.)
have been most abundant at different seasons.
The prime offender in the summer bloom has
been the Aphanizomenon. Germinating spores and
single filaments were found in the plankton by the
end of March or by the first of April (Table 4). By
Table 4. PHYTOPLANKTON DURING THE
GROWING SEASONS OF 1956 and 1957 IN
COUNTS PER MILLILITER1
Densest population
Link River sampled
Date Aphanlzo- Total Aphanizo- Total
menon 2/ plankton menon 2/ plankton
1956
Apr.
May
June
July
Aug.
Sept.
Oct.
Nov.
1957
Mar.
Apr.
May
June
July
Aug.
Sept.
Oct.
Dec.
2.5
1,481
17,637
11,701
20,744
11,607
3,1173/
21.6
0.0
0.0
3.0
6,929
12,499
10,748
11,353
1,093
0.0
27.0
1,816
18,549
12,837
20,759
11,903
3,196
37.5
2 145^
' 42.747
28.0
6,935
12,570
10,805
12,249
1/124
41.54/
9.5
3,420
18,349
33,163
194,756
56,438
43,756
78.2
:
0.0
0.0
841
12,591
57
57
57
4,132
0.0
1,921
4,150
18,586
34,459
196,360
57,665
43,846
85.8
, 5/
64~.5
877
12,597
57
57
§/
4,222
I.IST4-/
1. The study reported herein was supported by the
Oregon Klamath River Commission, the California
Klamath River Commission, the Klamath County
Court, The City of Klamath Falls, The Klamath
County Chamber of Commerce, and the Oregon Agri-
cultural Experiment Station.
2. Counts of >ph»">«omenon are in filaments per mll-
lillter.
3. Mldlake Station, no sample taken at the Link River.
4. Primarily AsterioneUa formosa.
5. Also Link River.
the end of April the population had developed to the
point that several thousand filaments were con-
tained in each liter of water. By the end of May a
million and a half filaments per liter were present
and in June, July, August and September the counts
were in the tens of millions. Toward the end of
September the counts started to drop and through
the months of October and November the plants
died in great masses. In December, Aphanizome-
non was represented only by an occasional spore.
The overwintering spores appeared to be produced
in September and October. Spores formed earlier
than this could not be germinated in culture and
their viability in nature is suspect The density of
the population in various areas of the lake varied
from day to day as wind drift and currents caused
characteristic aggregations. In August 1956 (Table
5) the count at the head of Link River was 20 thou-
sand filaments per milliliter but the densest popu-
lation in the Lake, measured along Eagle Ridge,
was 195 thousand filaments per milliliter.
Table 5. STANDING CROP OF PHYTOPLANKTON
AT VARIOUS STATIONS IN UPPER KLAMATH
LAKE, AUGUST 22-24, 1956
Station
Counts
per ml
1 Link River 20,760
3 Wocus Bay 10,176
5 Mid Lake 28,387
GAlgomalnlet 2,475
7 Algoma Point 21,986
8 Eagle Ridge 196,361
10 North End 5,864
11 Mouth of Williamson River 76
12 Shoalwater Bay 36,179
14 Pelican Bay 2,077
17 Agency Lake-Westside 191
19 Agency Lake -Eastside 10,643
Throughout the growing season a rather minor
depletion of the population occurred through the
escape of water into the Link River and finally in-
to the Klamath River. The generally held belief
that the limnoplankton is rapidly destroyed or re-
moved in flowing water was not supported by the
evidence from an exploratory trip down the Kla-
math River. Aphanizomenon, considerably re-
duced in number it is true, survived the trip down
the turbulent river very well and was present in
relatively large amounts at the mouth of the river.
Informants at Requa stated that the characteristic
color of the water was distinguishable several
hundred yards out at sea.
Wherever along its length the river had been
impounded, whether behind a dam or in a back-
water or slough, the water had produced blooms
comparable with that in Upper Klamath Lake. It
can be predicted mat the construction of additional
impoundments on the Klamath River will greatly
increase the organic load of this already impossi-
bly burdened stream and will probably bring an
-------
Klamath Lake, Natural Enrichment
25
end to fish production in this stream.
The weight of organic material exported from
Upper Klamath via the "A" canal and the Link River
has been calculated from data taken for a very
different purpose. While known to be inaccurate,
it is also known to err on the conservative side by
a factor of between ten and one hundred. In the six
month growing period from May through October
of 1956, over seven thousand tons of naturally pro-
duced organic material were exported. In August
(the month of highest production) alone, 23 hundred
tons were exported.
Analysis of the freshly dried algal material de-
monstrated the reason for the surprise of the ear-
lier investigators concerning the high nitrogen
content of the water in the absence of contamina-
tion by sewage. Over sixty percent of the dry
weight of the algae was protein (Table 6). Every-
Table 6. PARTIAL ANALYSIS OF THE COMPOSI-
TION OF FRESHLY DRIED ALGAE FROM UPPER
. KLAMATH LAKE
Composition
Crude Protein
Ash
Phosphorus
Calcium
Potassium
61.10
5.73
0.60
0.54
1.08
one experienced in pollution problems can visua-
lize the septic conditions that occur from time to
time in this basin.
Two additional problems need to be mentioned.
The first was the occurrence of limberneck dis-
ease affecting thousands of the water fowl inhabi-
ting the lake. This paralytic condition results
from the presence of a population of the type C
strain of Cjostricuum botulinum on the bottom.
The development of this population is favored by
the anaerobic conditions caused by the decomposi-
tion of highly proteinaceous organic material. At
times the number of birds affected has been so
great that, in order to prevent decimation of the
water fowl population, it was necessary for the
public to join forces with employees of state and
national agencies to rescue the diseased birds. The
second problem concerns a very potent toxin pro-
duced by the bloom organisms. No concrete evi-
dence was obtained as to the effect of this toxin on
the biota of the Lake or River but experiments with
mice proved that ingestion of the algal material
was quickly lethal and intraperitoneal injections of
the aqueous extract almost instantaneous in causing
death. Since the toxin is water soluble and thermo-
stable it could well be the explanation for the
stomach complaints mentioned earlier. At the
present time all the water for domestic and indus-
trial use in the area of Klamath Falls is obtained
from deep wells. This situation minimizes the
danger of the toxin from the public health stand-
point but also indicates the probable difficulty of
using the Lake water for any purposes other than
irrigation or power generation.
CHEMICAL CONSIDERATIONS
The concentration of dissolved minerals in the
lake water declined somewhat during the summer
as these materials were incorporated into the
plankton organisms. After the decline and death of
the summer bloom population in the late fall, the
concentration of minerals increased in the winter.
There was a similar characteristic seasonal
shift in the pH of the Lake water. During the peri-
ods of great plankton development the alkalinity
generally increased, while during winter it de-
creased. This shift was attributed to three fac-
tors: 1) precipitation came mostly in late fall,
winter and early spring and the run-off had an al-
kalinity approaching neutrality, 2) heavy drainage
of humic waters from the marsh areas at the same
season increased the H-ion level, 3) in summer the
hydroxyl-ion exchange in photosynthesis of the
bloom population quickly depleted the H-ion sup-
ply and raised the alkalinity (Table 7).
Table 7. SEASONAL VARIATION IN THE CHEM-
ISTRY OF UPPER KLAMATH LAKE WATER
Measurement
Temperature in °C.
pH
Nitrate
Ammonia
Phosphate
Iron
Calcium
Potassium
Magnesium
Summer Winter
(June-July-Aug.) (Jan.-Feb.-March)
18.0°
8.2
- 28.0°
-9.9
Values in ppm
2.0°
7.6
-8.5°
-9.2
0.06 - 0.2 0.12 - 1.0
0.155 - 1.290 0.061 - 1.674
0.0 0.0
0.1 - 0.7 0.2 - 2.0
1.6 - 3.2 2.5 - 5.5
2.6 -"37.0 3.0 - 43.5
5.0 - 29.5 5.0 - 15.0
Repeated chemical analyses of the water from
many sample stations failed to establish the pre-
sence of chemical factors such as exceptional con-
centration or combination of nutrient mineral ele-
ments that might explain the exceedingly high pro-
ductivity. A general statement seems warranted
that there are no inorganic chemical constituents
that could be singled out as being responsible for
this condition. The Lake water was low in calcium,
extremely low in dissolved phosphorus, and quite
high in chloride. Other nutrients were present in
merely nutritionally adequate amounts but could
not be described as abundant. A review of the idio-
syncrasies of the various influent waters soon
concentrated attention upon the great influx of
humic waters into the Lake during late winter and
early spring.
-------
26
ALGAE AND METROPOLITAN WASTES
SOURCES OF HUMIC WATER ENTERING UPPER
KLAMATH LAKE
About the northern end of Upper Klamath and
Agency Lakes are a number of drainage systems
(some mentioned previously) whose basins include
approximately 136 thousand acres of marsh land
(Table 8). Some agricultural land has been included
in this estimate of natural marsh but this is land
irrigated by natural flooding and has retained most
of the characteristics of natural marsh. Water
from these lands drains directly into the local
drainage system. In addition to this unmodified
marsh there are an additional 92 thousand acres
of muck soil agricultural land (Table 8) that has
Table 8 - ACRES OF NATURAL MARSH AND OF
IRRIGATED AGRICULTURAL LANDS DRAINING
INTO UPPER KLAMATH AND AGENCY LAKES
Table 9. PARTIAL ANALYSIS OF THE SOLIDS IN
SEASONAL SAMPLES OF HUMC WATERS FROM
THE WILLIAMSON RIVER
Natural marsh
Wood River
Upper Klamath Marsh
(Seven Mile Canal)
Sycan Marsh
(Sycan River)
Klamath Marsh
(Williamson River)
24,300 acres
2,100
25,000
85,000
Irrigated agricultural land
Wood River
Upper Klamath Lake
Sycan Marsh
Sprague River
50,000
11,000
18,000
13,000
been drained and diked. This land lies flooded in
fall and winter and is pumped dry in late winter or
early spring. The drainage water, like the natu-
ral marsh drainage contains humic leachate from
the marsh soil. This marsh soil, or muck, con-
sists of a fiberous, Typha peat, several feet in
thickness lying on top of variously assorted layers
of tuff, pumice and diatomite.
Determination of the composition of the organic
leachate has proven quite difficult and only incom-
plete information is available. A comparison of the
analyses of samples showed the content of total
dissolved solids per unit volume in samples taken
in summer to be twice that in those taken in winter
(or early spring). The nitrogen content in summer
was more than double that of the winter, but the
phenolic content decreased somewhat in the sum-
mer (Table 9).
EFFECT OF HUMATES ON THE GROWTH OF
ALGAE IN CULTURE
Algal culture media were modified to include a
proportion of humic water and the final pH was ad-
Total solids
Kjeldahl nitrogen
Aromatic hydroxl groups
January
103 mg/1
0.7 mg/I
Spring
3.15 mg/1
August
247 mg/1
1.7 mg/1
Summer
2.5 mg/1
justed to a range in which our test organisms
grew well.. It was found that the production, meas-
ured as dry weight of algae, was 70% greater in
the humic-enriched cultures than in the unmodified
controls. The advantage gained ;by adding the hu-
mates could be overcome by quadrupling the min-
eral content of the control medium but the added
growth appeared in the controls only after three
weeks of growth. The vigorous growth stimulated
by the presence of the humates rapidly exhausted
the relatively lower mineral resources of the ex-
perimental medium while the four-fold fortification
of the controls allowed ample minerals for a longer
period of growth.
The mode of action of the humates remains to
be elucidated. Some workers (Burk, et al, 1932a,
1932b) have suggested that the high iron content of
humic waters provides considerable stimulation of
plant growth. We know, however, that there was
sufficient iron in the formulation of the control
medium to provide for very vigorous growth of the
test organisms for a month. The control medium
likewise contained a shotgun mixture of minor ele-
ments and it seems relatively unlikely that the
presence of one of these was the basis of the stim-
ulation by the humates.
It is our belief, as yet unsupported by concrete
evidence, that the action of the humates as chela-
ting and buffering agents was primarily responsi-
ble for the observed stimulation. It is also possi-
ble that colloidal absorption by the humates inacti-
vated metabolic end products that otherwise would
have exerted an anti-metabolic influence in older
cultures. Of these three possible explanations,
chelation appears the most reasonable. The bene-
fit derived from buffer action as well as action of
a protective colloid should appear as a prolongation
of the logarithmic phase of the growth curve of the
cultures. Contrarily, stimulation in our cultures
appeared in the earliest stages of growth and the
cultures tended to become exhausted or to stale
more quickly than the controls.
-------
Klamath Lake, Natural Enrichment
27
CONTROL OF THE ALGAL NUISANCE
Briefly stated, the problems involved in the
control of an algal population on such a large scale
are manifest The application of chemical control
agents would be expensive in material, manpower
and equipment. The dangers to the associated biota
would be great. Since such measures would give
only temporary relief, effects of the long term use
of chemical materials would require intensive
study before application. It seems necessary to
consider other approaches that lack at least some
of these objectionable features. The first sugges-
tion concerns the diversion of all influent humic
waters to reservoirs from which they would be
eventually distributed for the irrigation of agricul-
tural land. Besides the very probable major re-
duction in the algal population that would be effected
there would result a conservation of large amounts
of humates that are presently being lost.
A second possible approach would be the dis-
charge into the lake or its tributaries of a slurry
of some colloidal inorganic material, such as
Bentonite clay, in sufficient amount to reduce the
light intensity at the bottom over large areas of
the lake to below the compensation point. The eco-
nomic feasibility of these two suggestions must
be determined by a study of the engineering consi-
derations involved.
REFERENCES
Burk, D., H. Lineweaver, and C. K. Homer. 1932a.
Iron in relation to the stimulation of growth by
humic acid. Soil Science 33: 413-453.
Burk, D., H. Lineweaver, and C. K. Horner.
1932b. The physiological nature of humic acid
stimulation of Azotobacter growth. Soil Science 33:
455-487.
Stanley, Lewis A. 1954. Report of the Oregon
Klamath River Commission on Water Resources
and Requirements of the Upper Klamath Basin.
RECENT CHANGES IN THE TROPHIC NATURE OF LAKE WASHINGTON - A REVIEW*
G. C. ANDERSON
Department of Oceanography, University of Washington, Seattle
ABSTRACT
Until a few years ago Lake Washington was a
relatively clear, oligotrophic lake. Since the turn
of the century, Seattle and other communities bor-
dering the lake have been discharging sewage in
increasing amounts, and as of 1957 the lake was
receiving the discharge from 10 sewage treatment
plants plus septic tank drainage representing
76,300 people. Lake Washington has responded
with increased algal productivity and is now eu-
trophic.
Detailed limnological studies in 1933, 1950 and
1955-1958 illustrate the nature of the lake re-
sponse. Since 1950, prominent changes have oc-
curred, particularly in 1955 when, for the first
time, there appeared an increased growth of phy-
toplankton made up mainly by the blue-green alga
Oscillatoria rubescens, a notorious indicator of
pollution in many lakes. A regular annual increase
in crop of algae has occurred since this time. Due
to the increase in productivity, the transparency of
the water has decreased, and oxygen consumption
and nutrient release in the hypolimnion during
summer stratification have progressively in-
creased. The hypolimnetic oxygen deficit has
shown an almost threefold increase since 1933:
1.18 mg/cm2/month in 1933, 3.13 in 1955. In 1957,
for the first time, the deepest waters became
anaerobic for a short period.
If fertilization of the lake continues, it may be
expected that within a very few years nuisance
blooms of algae will be produced, unsightly scums
and decaying foul-smelling masses of algae will
accumulate in places along the shoreline, and well
developed stagnant conditions in the hypolimnion
will occur. The formation in 1958 of the Munici-
pality of Metropolitan Seattle, a corporation de-
signed to handle sewerage services on a metropo-
litan basis, was, in part, recognition of theurgency
of the Lake Washington situation. Long-range
plans include the ultimate diversion of all sewage
entering the lake.
INTRODUCTION
The purpose of this paper is to review some of
the limnological changes that have been observed
in a lake which is undergoing nutrient enrichment
by progressively increasing additions of sewage
Contribution No. 236 from the Department of Oceanography, University of Washington.
-------
28
ALGAE AND METROPOLITAN WASTES
discharge resulting from the growth of a large
metropolitan area. The situation of undesirable
lake response to the waste products of a growing
human population has been observed in many places
in this country and elsewhere; Hasler (1947) des-
cribed several such cases. Typically, the addi-
tion of nutrient rich material from domestic sew-
age will hasten the natural process of eutrophi-
cation of a lake. In a matter of a few years, de-
pending largely upon the amount of sewage effluent,
the size of the lake and the amount and nature of
inflow water from natural sources, the lake may
respond with increased growths of algae particu-
larly of the blue-green variety, larger populations
of zooplankton, depletion of dissolved oxygen in
the hypolimnion, and the replacement of game fish
with the scrap fish variety. The recreational uses
of the water are destroyed, nuisance blooms of
algae occur to the extent that undesirable odors
are produced from the accumulation of decaying
masses, and property surrounding the lake de-
clines in value. The community is faced with the
choice of losing the lake permanently as a recre-
ational and esthetic benefit or to correct the situa-
tion by the diversion of sewage inflow from the
lake. At the present time, Lake Washington is in
the process of deterioration from sewage enrich-
ment, but as yet algal blooms have not reached the
nuisance level.
The population of Seattle is estimated to rise
from 572,000 in 1957 to 1,250,000 in 1980. At
present, approximately 20 percent of the popula-
tion of the metropolitan area reside in the Lake
Washington drainage basin; ultimately 50 percent
will reside there. In 1957, the flow from ten sec-
ondary' sewage treatment plants, serving 64,300
people, was discharging into the lake. In addition,
the effluent from 4,000 septic tanks representing
12,000 persons reaches the lake. At the beginning
of the century, raw sewage entered the lake, but in
1926 steps were taken to remove this discharge and
the project reached completion in 1941. However,
during periods of appreciable rainfall, combined
sanitary and storm-water sewage systems over-
flow into the lake and contribute about 3.5 percent
of the total annual sewage generated in the area.
Lake Washington has received limnological at-
tention from time to time since 1913. Recently,
several studies have been conducted, particularly
since it was recognized that the lake was respond-
ing to ever increasing amounts of treated sewage.
The first published observation of the lake consist-
ed of a single series of chemical and plankton
samplings taken August 9, 1913 (Kemmerer,
Bovard and Boorman, 1923). A detailed investiga-
N| tion was first made in 1933 when an annual study
was conducted of the seasonal variation in temper-
ature, oxygen, nutrients and the net plankton
(Scheffer, 1936; Robinson, 1938; Scheffer and Rob-
inson, 1939). Unfortunately, phytoplankton esti-
mates were recorded as relative frequency and are
not quantitative. In 1950 a further study was made
of the seasonal fluctuations of the physical, chemi-
cal, and biological conditions which included
quantitative estimates of the phytoplankton and
zooplankton populations (Comita, 1953; Anderson,
1954;Comita and Anderson, 1959). In 1952, studies
by the Pollution Control Commission involved esti-
mates of bacterial pollution in selected areas of
the lake and in tributary rivers (Peterson, Jones
and Orlob, 1952). The same group investigated
surface chemical and chlorophyll conditions, in
1953, at 26 stations spread over the entire lake
(Peterson, 1955). The Department of Oceanography
at the University of Washington has obtained data
since 1950 on oxygen, temperature and salinity in
connection with a study on the intrusion of salt wa-
ter into the lake through the Lake Washington Ship
Canal (Rattray, Seckel and Barnes, 1954; Collias
and Seckel, 1954). During the summers of 1955
and 1956, members of the Department of Zoology
in cooperation with the Pollution Control Commis-
sion sampled the lake for temperature, oxygen,
phosphate and phytoplankton (Edmondson, Anderson
and Peterson, 1956; Sylvester, Edmondson and
Bogan, 1956).
Because of the marked increase in algal pro-
ductivity noted in the summers of 1955 and 1956,
a long-term study was initiated in September 1956
by Drs. W. T. Edmondson and Joseph Shapiro. The
immediate purpose of this study has been to obtain
a detailed description of the productivity and nutri-
ent condition of the lake and to compare it with
past and future conditions (Edmondson, 1960;
Shapiro, 1960). Also in 1956, it was realized that
a potentially serious civic situation existed, and an
engineering firm was appointed by city, county and
state officials to prepare a report on a long-range
program of sewerage and drainage improvements
for the metropolitan Seattle area (Brown and Cald-
well, 1958). A particularly useful feature of this
study to further limnological investigation of the
lake was the preparation of a nutrient budget giving
the increment of nitrogen and phosphorus from
analysis of measurements made in inlets, sewage
effluent and in the lake itself.
During the preparation of this paper, the writer
has drawn extensively from those reports cited in
the references. Especial thanks are due to Dr. W.
T. Edmondson, University of Washington, and Dr.
Joseph Shapiro, The Johns Hopkins University,
for allowing the writer to examine papers in press
and unpublished data from recent years and to
utilize some of these data in this review.
PHYSICAL CONDITIONS
Lake Washington lies in an elongate glacial
trough sculptured by the Vashon ice sheet. The
surface area is 8762 ha, maximum depth 65.2m,
mean depth 32.9 m, and volume 2,884.2 x 106m3
(Comita and Anderson, 1959). In cross section,
-------
Trophic Changes in Lake Washington
29
the lake is W-shaped rather than the more con-
ventional U-shaped. After the withdrawal of the
Vashon glacier, the lake underwent a transitory
marine phase, evidence for which was found in
deep sediment cores (Gould and Budinger, 1960).
Typically, the lake has an epilimnion of 10 m,
thermocline 10 m, and hypolimnion 40 m. Stra-
tification usually takes place in late March or
early April and remains so until November or
December. After stratification breaks down in
late autumn, full circulation takes place during
the winter at a temperature of 5 to 8°C. Trans-
parency of the water fluctuates with the season and
is highest in winter and lowest during periods of
summer algal blooms. The arithmetic mean of
Secchi disc measurements made in 1950 was 3.82
m (Comita and Anderson, 1959).
LMNOLOGICAL CONDITIONS TO 1950
Generally speaking, there were few variations
noted in most conditions between the studies made
in 1933 and 1950 apart from those which might be
expected from normal annual variations due to
differences in climatic conditions from year to
year. For this reason and because the 1933 study
did not include quantitative estimates of the phyto-
plankton, 1950 has been chosen as representing
typical lake conditions prior to its response to
added nutrient income. Some of the deteriorative
changes between 1933 and 1950 will be mentioned
in a later section.
During 1950, the pH was near neutrality except
hi midsummer when it rose to above 8 due to re-
moval of CO£ by photosynthesis. Bicarbonate
fluctuated between 16-32 mg/L HCO3. Dissolved
oxygen was abundant at all depths, although a min-
imum of 5.6 mg/L was measured at 55 m during
summer stratification. The phosphate concentra-
tion remained near 15 ppb PO4-P during winter
mixing but decreased to 0-1 ppb in the epilimnion
during summer. An accumulation of phosphate oc-
curred in the hypolimnion as summer stratification
progressed, and a maximum of 23 ppb was reached.
Data on nitrogen concentrations during this year
are scanty, but the maximum observed during the
winter was 560 ppb N(>j.N which decreased to less
than measurable quantity in the epilimnion at times
in the summer. The fluctuations of chemical con-
ditions described above compare favorably with
those described in 1933 with the exception that the
nutrient concentration was lower in the earlier
year, i.e., approximately 8 ppb PO^.P and 150 ppb
N03.N during winter mixing.
The annual fluctuation in quantity of phy top lank-
ton from measurements of cell volume and chloro-
phyll followed a typical bicyclic pattern. The
maximum bloom occurred in spring, a midsummer
minimum followed, and a second bloom of lesser
.extent was observed in late August. Diatoms made
up the greatest part of the spring bloom. Common
among these were Stephanodiscus niagarae, Melo-
siravarians, M. italica, M. italica var. tenuissima
Fragilaria crotonensis, Rhizosolenia gracilis, As-
terionella formosa, Synedra acus and Cyclotella
bodanica. The late August bloom was composed
mainly of dinoflagellates, especially Peridinium
divergens and to a lesser extent Ceratium hirun-
dinella. Among the blue-green algae, Oscillatoria
agardhi and Phormidium sp. were common but did
not contribute significantly to the size of the major
blooms. The Chlorophyta, or green algae, were
common at times but quantitatively unimportant.
In contrast, Anabaena lemmermanni and a number
of diatoms were of greatest relative importance in
the summer phytoplankton during 1933.
LIMNOLOGICAL CONDITIONS SINCE 1950
Some of the significant changes observed in the
lake since 1933 are summarized in Figure 5. It is
immediately obvious that the most drastic changes
have occurred since 1950.
The standing crop of phytoplankton has in-
creased significantly in recent years as indicated
by measurements of phytoplankton volume and
chlorophyll concentration (Fig. 5, Panels A and B).
During the summer months of July, August, and
September, the phytoplankton volume in the epilim-
nion increased from 0.6 mm3/L in 1950 to 1.6
mm3/L in 1955 and further to 4.2 mm3/L in 1956,
the last year for which data on phytoplankton vol-
ume are available at the present time. Judging
from the measurements of chlorophyll, a further
large increase in phytoplankton crop occurred in
1958.
As mentioned previously, the major components
of the phytoplankton in 1933 were diatoms and
Anabaena lemmermanni. In 1950, diatoms made
up the spring bloom and dinoflagellates the late
summer bloom. Phormidium sp. and Oscillatoria
agardhi were conspicuous components in 1950, but
the largest crops attained by the blue-green algae
were 0.33 mm3/ Lin February and again in Septem-
ber. The maximum standing crop of all phyto-
plankton observed during the period of sampling
in 1955 was 2.89 mm3/L in July of which 2.78
mm3/L or 96 percent was composed of the blue-
green alga Oscillatoria rubescens. O. rubescens
remained abundant throughout the samplings and
Aphanizomenon flosaquae became dominant in
September.
O. rubescens is a well-known algal species,
especially in the Swiss lakes, as an indicator of
pollution. Late in the nineteenth century, Zurich-
see, Switzerland, changed in a relatively short
period of time from an oligotrophic to an eutrophic
lake. The phytoplankton increased rapidly and was
composed largely of O_. rubescens. About 10 years
later, the cladoceran Bosmina coregoni was re-
-------
30
ALGAE AND METROPOLITAN WASTES
- A
JULY AUG. SEPT.
A. VOLUME OF ALGAE
10
\
- 1.8
2.1
8.4
6.1 6.1
1950 1955 1956 1957 1958
B. CHLOROPHYLL IN EPILIMNION
8'
10
»
25
^ '*2 (1.6) 1.5
Sccehi disc
C. MINIMUM TRANSPARENCY
\ 5-
8.0
_
8,|
6.9
4.6
5.1
1933 1950 1955 1957 1958
D. OXYGEN IN HYPOLIMNION
I
S
I
BELOW
10 m.
--jT^rr-—AJ
••* 4/*k_
20m.
1933 1950 1958
E.RATE OF OXYGEN CONSUMPTION
s?
30
\
I
F
.20
26.4
19.1
_ 9.6
6.1
16.6
1933 1950 1955 1957 1958
PHOSPHATE PHOSPHORUS
IN HYPOLIMNION
Figure 5. CHANGES THAT HAVE OCCURRED IN SOME PROPERTIES IN LAKE WASHINGTON IN RECENT
YEARS.
Panel A - Phytoplarikton volumes in the epilimnion.
B -Average chlorophyll concentration in the epilimnion during July, August and September of each year.
C - Minimum observed transparency during each year.
D - Mean dissolved oxygen concentration below 10 m at the end of summer stratification.
E -Oxygen deficit between June 20 and August 20 below 10 m and below 20 m.
F - Mean concentration of phosphate below 10 m at the end of summer stratification.
(Modified from Edmondson, 1960).
-------
Trophic Changes in Lake Washington
31
placed by B. longirostris. A somewhat similar
situation has been observed in Lake Washington
although the change of cladocera species occurred
prior to the appearance of O. rubescens. As early
as 1940, B. longirostris replaced B. longispina
Leydig (= B. coregoni longispina). By 1955, phy-
toplankton volume had increased significantly and
0. rubescens became dominant. Deevey (1942) has
shown from fossil evidence that the same change in
Bosmina species occurred during the transition
of Linsley Pond from oligotrophic to eutrophic.
The transparency of a lake is related in part to
the quantity of plankton material present. Due to
increased algal growth in Lake Washington, Secchi
disc measurements are becoming less (Fig. 5,
Panel C). The minimum transparency in 1950 was
2.5m whereas in 1956 it was 1.2 m. One measure-
ment of 4.1 m was made in 1913. Data are not
available for 1933.
Seasonal changes in the hypolimnetic waters of
lakes are in part related to epilimnetic processes.
Organic material, originally produced or derived
from the epilimnion, settles into the deeper waters.
This material consists of living and dead phyto-
planktonand zooplankton, fecal material from zoo-
plankton, and other debris entering the lake. As
the dead organic matter decomposes by bacterial
action and living material respires, oxygen is con-
~sumed and nutrients other than those absorbed by
bacteria and living algal cells are released to the
waters. Some fraction of the material is sedi-
mented to the bottom. It is therefore not surpris-
ing that some workers have found the consumption
of oxygen in the hypolimnion in a series of lakes to
be related roughly to the mean quantity of seston
(Hutchinson, 1938) or standing crop of net plankton
(Rawson, 1942) in the epilimnion although the true
relationship is probably with the total production
in the epilimnion rather than with the standing crop.
In LakeiWashington after 1950, it can readily be
seen that during summer the average concentration
of dissolved oxygen in hypolimnetic waters has be-
come much less, and in 1957, the deepest waters
became anaerobic (Fig. 5, Panel D). The rate of
removal of dissolved oxygen from the hypolimnion,
termed the oxygen deficit, has been calculated be-
low 10 m and below 20 m for all years since 1933
that sufficient data are available and expressed on
an areal basis (Edmondson, Anderson and Peter-
son,. 1956; Edmondson, 1960). In earlier years,
when the lake was relatively transparent, it is
likely that some photosynthesis took place in the
upper part of the hypolimnion so that values cal-
culated below 10 m in those years are probably
minimal and the 20 m estimate is more reliable.
However, as the lake has become less transparent,
especially since 1955, and because an interesting
oxygen minimum between 10-15 m has increased
in magnitude particularly since 1956, the values
below 10 m may be better estimates in more re-
cent years. In any case it is apparent that a sig-
nificant increase in the deficit has taken place
since 1933 (Fig. 5, Panel E). Although the larg-
est rate (3.1 mg/cm^/month 02 consumed below
20 m) was recorded in 1955, the general trend is
that of progressive increase.
One of the features of the distribution of dis-
solved oxygen in the lake is the previously referred
to appearance of a metalimnetic oxygen minimum
beginning usually early in June and growing pro-
gressively stronger until its disappearance at the
end of summer stratification. Shapiro (1960) has
reported an increase in the magnitude of this mini-
mum correlated with conditions of increasing eu-
trophication, the minimum recorded level reaching
2.2 mg/L at 15 m in 1957. Shapiro has concluded
that the agent responsible for this minimum is the
respiration of a metalimnetic population of non-
migrating copepods and that the recent increase in
magnitude of the minimum is due to an increase in
the numbers of copepods.
The concentrations of all measured nutrients
have been increasing greatly in recent years, and
phosphate, in particular, has shown a marked in-
crease. This is adequately illustrated by noting
that phosphate accumulation in the hypolimnion
during summer stratification has steadily in-
creased since 1950 (Fig. 5, Panel F). The cause
for this has, of course, been attributed to greater
amounts of organic material produced in the epi-
limnion due to an increase in the nutrient income
to the lake with a resultant rise in the amount of
material settling into the hypolimnion, subse-
quently decomposing and releasing nutrients to
these waters. Epilimnetic waters, in all years
studied, contained low, sometimes unde tec table,
quantities of phosphate, indicating near maximum
utilization of nutrients in the trophogenic zone
during the summers. As a result, the major dif-
ferences between the years are reflected in chan-
ges in the hypolimnion. For example, the maxi-
mum concentration of phosphate reached in the
deepest waters was 23 ppb in 1950, 89 ppb in 1957,
and 74 ppb in 1958. Although the maximum record-
ed measurement in 1958 was somewhat less than
that in 1957, greater concentrations were found at
intermediate depths in 1958 so that the total phos-
phate present was greater than before.
Although the principal source of nitrogen and
phosphorus to Lake Washington at the present time
is from the major tributary streams, the amount
contributed by sewage discharge would in the near
future greatly exceed that from natural sources.
Figure 6 shows the enrichment of the lake by phos-
phorus from natural sources and from sewage
sources, and the projected trend from sewage in
future years if effluent continues to enter the lake
(Brown and Caldwell, 1958). Table 10 gives the in-
crement of N and P to the lake from natural and
sewage sources (Brown and Caldwell, 1958). The
-------
32
ALGAE AND METROPOLITAN WASTES
estimated amount of nitrogen entering the lake
from sewage sources in the past 40 years has more
than doubled, and the phosphorus income has al-
most tripled. The contribution of nitrogen from
sewage is expected to increase from the 195 7 level
of 6.5 percent of the total input from all sources to
an ultimate value of 35.2percent; phosphorus from
43 percent to 92 percent.
NATURAL SOURCES
O O O
• O M
O O
YEARS
Figure 6. THE TREND OF PHOSPHORUS EN-
RICHMENT IN LAKE WASHINGTON FROM NAT-
URAL AND SEWAGE SOURCES. In 1916, the
Cedar River was diverted to the lake which ac-
counts for the rise in phosphorus from natural
sources since that time (from Brown and Caldwell,
1958).
Table 10 - CONTRIBUTION OF TOTAL DIS-
SOLVED NITROGEN AND PHOSPHORUS TO
LAKE WASHINGTON.
Category
Natural sources
Sewage sources
Total
Mean content In lane
Date
1957
1916-1930
1931-1940
1941-1950
1957
1957
1933
1957
Nitrogen*
5,850
181
240
226
409
6,259
1,970
4,180
Phosphorus*
151
42
54
74
114
265
114
184
* Nutrient values expressed as 1000 Ibs/year. Values taken from Brown
and Caldwell (1958).
In summary, Lake Washington is exhibiting an
increase in productivity, resulting from fertiliza-
tion with treated sewage. The mean quantity of
phytoplankton and nutrient concentration within the
lake has progressively increased, especially with-
in the past decade. Also, significant annual in-
creases have occurred in oxygen consumption and
phosphorus accumulation in hypolimnetic waters
during summer stratification. If fertilization con-
tinues, the series of changes already observed
can be expected to increase in magnitude until se-
rious conditions of algal nuisance blooms, odors,
scums, bottom stagnation, etc. develop.
The only practicable manner at present of con-
trolling fertilization in a lake of this size is to di-
vert the sewage elsewhere. The Pollution Control
Commission, in 1956, established a policy requir-
ing that all sewage treatment plants eventually di-
vert their effluent from the lake into Puget Sound.
However, a project on this scale requires the co-
operation of several responsible communities
bordering the lake and for this reason it has long
been felt by many interested parties in Seattle
that a central sewerage authority is necessary to
deal adequately with the problem. In 1957, an act
providing for the formation of Metropolitan Muni-
cipal Corporations was passed by the Washington
State Legislature. By a vote of the residents of the
greater Seattle area on September 9, 1958, the
Municipality of Metropolitan Seattle was established
and empowered to plan, finance, and administer
sewage services on a metropolitan basis. If the
present schedule of "Metro" is maintained, more
than 80 percent of the sewage discharges presently
entering Lake Washington will be diverted by 1962.
Limnologists will be keenly interested in the bio-
logical response of the lake as the nutrient income
declines during and after sewage diversion.
REFERENCES
Anderson, G. C. 1954. A limnological study of the
seasonal variations of phytoplankton populations.
Ph.D. Thesis. Univ. Washington. 268 pp.
Brown and CaldwelL 1958. (Civil and Chemical
Engineers). Sewage disposal in Lake Washington.
Chapter 10 in Metropolitan Seattle sewerage and
drainage survey. A report for the city of Seattle,
King County and the State of Washington on the col-
lection, treatment and disposal of sewage and the
collection and disposal of storm water in the met-
ropolitan Seattle area. Seattle, pp. 219-247.
Collias, E. E. and G. R. SeckeL 1954. Lake Wash-
ington Ship Canal data. Univ. Washington, Dept.
Oceanogr. Spec. Rept. No. 2. pp. 1-27.
Comita, G. W. 1953. A limnological study of plank-
tonic copepod population. PhuD. Thesis. Univ.
Washington. 195 pp.
Comita, G. W. and G. C. Anderson. 1959. The sea-
sonal development of a population of Diaptomus
ashlandi Marsh, and related phytoplankton cycles
in Lake Washington. Limnol. Oceanogr., £(1):
37-52.
-------
Trophic Changes in Lake Washington
33
REFERENCES (Cont'd.)
E. S. 1942. Studies on Connecticut lake
sediments, ffl. The biostratonomy of Linsley
Pond. Amer. J. Sci., 240: 233-264, 313-338.
Edmondson, W. T. 1960. Changes in Lake Wash-
ington following an increase in the nutrient income.
Proc. Int. Assoc. Theor. Appl. LimnoL (In press).
Edmondson, W. T., G. C. Anderson and D. R. Pe-
terson. 1956. Artificial eutrophication of Lake
Washington. Limnol. Oceanogr., 1^(1): 47-53.
Gould, H. R. and T. F. Budinger. 1960. Control of
sedimentation and bottom configuration by convec-
tion currents, Lake Washington, Washington. J.
Mar. Res. (In press).
Easier, A. D. 1947. Eutrophication of lakes by
domestic drainage. Ecology. 28: 383-395.
Hutchinson, G. E. 1938. On the relation between
the oxygen deficit and the productivity and typology
of lakes. Int. Rev. Hydrobiol., 36: 336-355.
Kemmerer, G., J. F. Bovard and W. R. Boorman.
1923. Northwestern lakes of the United States:
Biological and chemical studies with reference to
possibilities in production of fish. Bull. U. S. Bur.
Fish., 39: 51-140.
Peterson, D.R. 1955. An investigation of pollution -
al effects in Lake Washington. Washington Pollu-
tion Control Commission. Tech. BulL No. 18. 18
pp.
Peterson, D. R., K. R. Jones and G. T. Orlob. 1952.
An investigation of pollution in Lake Washington.
Washington Pollution Control Commission. Tech.
BulL No. 14. 29 pp.
Rattray, M., Jr., G. R. Seckel and C. A. Barnes.
1954. Salt budget in the Lake Washington Ship Can-
al system. J. Mar. Res., 1JJ: 263-275.
Rawson, D. S. 1942. A comparison of some large
alpine lakes in western Canada. Ecology 23:143-
161.
Robinson, R. J. 1938. Chemical data for Lake
Washington. Typewritten report. Univ. Washington
Library.
Scheffer, V. B. 1936. The plankton of Lake Wash-
ington. Ph.D. Thesis. Univ. Washington.
Scheffer, V. B. and R. J. Robinson, 1939. A limn-
ological study of Lake Washington. Ecol. Monogr.,
9: 95-143.
Shapiro, J. 1960. The cause of a metalimnetic
minimum of dissolved oxygen. Limnol. Oceanogr.
(In press).
Sylvester, R. O., W. T. Edmondson and R. H.
Bogan. 1956. A new critical phase of the Lake
Washington pollution problem. The Trend in En-
gineering, J5 (2): 8-14.
-------
Specific Problems in Rivers
ALGAE IN RIVERS OF THE UNITED STATES
C. MERVIN PALMER, In Charge
Interference Organisms Studies
Research Branch, Division of Water Supply and Pollution Control
Concern for the quality of river waters in the
United States increases as the many uses for these
waters are intensified. For example, the algae,
which are microscopic organisms common in riv-
ers, have a significance that is being emphasized
in relation to self purification, radioactivity, wa-
ter treatment, fish and other aquatic animal life,
industrial and sewage pollution, taste and odor
production, and recreational uses of the stream
water.
Unlike studies of lakes, there have been few de-
tailed or long continued investigations of algae in
American rivers. The general impression regar-
ding planktonic algae in rivers seems to be that
they are relatively few in number, that many of
them are transients multiplying in lakes and mere-
ly existing passively and temporarily in streams,
or that they are primarily benthic organisms that
have broken away from their moorings at the bot-
tom and sides of the stream.
The unattached algae in rivers have been con-
sidered to be so few that often they have been re-
corded in numbers per liter or cubic meter, rather
than per ml. In such data converted to numbers
per ml, the algae are generally fewer than 100.
A real difficulty in evaluation, however, has been
the dearth of recorded information on the abun-
dance of algae in rivers.
It is necessary to know the algal population of
rivers quantitatively and qualitatively, if we are to
be concerned with assessing their value or their
significance-as stream purifiers, pollution indica-
tors, or as excessive growths; their role in water
treatment problems; and their function as the pri-
mary food for fish. It can be important to know the
algal population of a river before any major change
is made in the use of the stream. Also, we need to
know the algal population of rivers throughout the
year and not merely for the warmer months.
This type of basic information has been almost
completely lacking in the United States. It should
be obtained over a long period of time, for all
months of the year, and for several representa-
tive locations on all important rivers of the coun-
try. The same procedure for enumeration must be
used at all stations if the various records are to be
easily compared. While records of all species
would be helpful, it can hardly be expected that a
comprehensive plan could be carried beyond gen-
eric level since much more time would be required
for identification of species.
Fortunately some standardized information on
stream algae is now becoming available. In 1957
the National Water Quality Network program was
inaugurated by the Public Health Service. Samp-
ling stations on 16 rivers and the Great Lakes
were chosen. where water could be obtained at
regular intervals for various examinations, one of
these being for plankton organisms (Palange and
Megregian, 1958).
During 1958, the Network's first year, plank-
ton analyses were made once per month of sam-
ples from 47 stations in the United States (Figure
7). Rivers included in the survey during the first
year were the Arkansas, Colorado, Columbia,
Delaware, Detroit, Hudson, Merrimack, Missis-
sippi, Missouri, Ohio,Potomac, Red, Rio Grande,
Savannah, Snake, and Tennessee. The numbers of
stations and rivers sampled are being increased
gradually so that the accumulated data may be re-
presentative of a larger part of the country. More-
over, samples are now being collected semi-
monthly rather than monthly.
The clump count procedure is used in recording
the plankton organisms as numbers per ml. In this
method of enumeration, isolated cells and colonies
are recognized as the unit. Analysis is commonly
made on unconcentrated samples at a magnification
of 200X. Two longitudinal strips (representing a-
bout 200 fields) of the Sedgwick-Rafter slide are
examined, rather than only 10 fields as often rec-
ommended.
The first annual compilation of data has been
published (Anon., 1958) together with a statistical
summary of selected data (Anon., 1959). The re-
sults of the second year's work are completed and
ready for publication. From these records some
information as to the numbers and kinds of algae in
the country as a whole has been obtained for pre-
sentation at this time.
Over a period of two years the average count
per monthly sample was 3625. algae per ml, the
first year averaging 3460 and the second year 3850.
The monthly average ranged from 1376 to 6745.
November, December, and March had the lowest
average counts for the two year period while A-
pril, September, and October had the highest. The
34
-------
Algae in U. S. Rivers
Figure 7. PHS WATER QUALITY BASIC DATA PROGRAM NETWORK OF SAMPLING STATIONS
indications are that the plankton algae in rivers
average much larger numbers than has generally
been assumed. During the early winter the count
may be lower but seasonal fluctuation is less than
had been suspected.
Occasional very high counts may raise the av-
erages significantly. For example, the median for
all rivers during the first year was only 1460 per
ml, in contrast to a mean'of 3460 per mL Some
individual counts exceeded 20,000, but only one
was above 50,000.
The five rivers with the highest average counts
for the first year were the Mississippi, Arkansas,
Merrimack, Missouri, and Columbia, while the
five with the lowest were the Red, Detroit, Colo-
rado, Savannah, and Tennessee rivers. High
plankton counts are to be expected in rivers en-
riched by land drainage from productive soils, or
by treated or untreated sewage from cities and
towns, particularly where toxic industrial wastes
are not abundant. This relationship of plankton
count to enrichment will undoubtedly become more
evident when data are available from more fre-
quent sampling and more sampling stations over a
period of several years. Plankton fluctuations due
to sporadic variations in weather or other environ-
mental factors will then have less influence on data
representing average plankton populations.
The same genera of algae tend to be dominant
in all of the rivers of the country, and only a lim-
ited number of genera comprise the list of the
dominant forms that are generally encountered.
Leading the list are several diatoms, the first six
in order of decreasing frequency being Cyclotella,
Synedra, Melosira, Navicula, Asterionella, and
Stephanodiscus. Other algae high in the list are
Anacystis, Chlorella, Chlamydomonas, and An-
kistrodesmus.
Various comparative studies of the plankton
records are being made at the Taft Center. One of
these deals with the planktonic green algae of the
Mississippi and the Ohio rivers. Another is con-
cerned with the number of genera of diatoms in
several river systems.
An indication of what the plankton counts may
-------
36
ALGAE AND METROPOLITAN WASTES
reveal is found when four-month averages for the
year 1959 of the planktonic green algae at 8 sta-
tions on the Mississippi River are compared. The
average count decreases from the uppermost sta-
tion at RedWing, Minnesota, to the lowermost sta-
tion located at New Orleans, La. (Figure 8). The
average count per ml for the whole year was 2087
at Red Wing, and 151 at New Orleans, the count at
each intermediate station being less than the one
above. In the upper portion of the river, represen-
ted by the first 3 stations, the number of genera
and the count were considerably lower during the
Nov.-Feb. period than for the remainder of the
year. However, from East St. Louis to New Or-
leans for each third of the year, both the number
of genera and the count were fairly constant at all
stations.
NOV.-FEB.
MAR.-JUNE
JULY- OCT.
NOV.-FEB.
MAM.-JUNE
JULY-OCT.
RED WING,
MINNESOTA
OUBUQUE.
IOWA
BURLINGTON.
IOWA
E. ST. LOUIS.
ILLINOIS
CAPE GIRARDEAl
MISSOURI
WEST MEMPHIS,
ARKANSAS
VICKSBUR6.
MISSISSIPPI
NEW ORLEANS.
LOUISIANA
LEGEND:
AVERAGE NO. GER€RA/«I
AVERAGE COUNT/•!
AVERAGE COUNT/ »l OVERLAP
Figure 8. PLANKTONIC GREEN ALGAE — MISSISSIPPI RIVER ~ 1959
-------
Algae in U. S. Rivers
37
For the Ohio River the planktonic green algae,
averaged by four-month periods for a total of 2
years, indicate a very distinct difference of popu-
lation in 1958 and 1959 (Figure 9). During the
March-June and the July-Oct. periods of 1959, the
average numbers of genera and the average counts
were much higher than in the previous year. It is
believed that either an increase in algal nutrients
or a decrease in toxic materials, such as certain
industrial wastes in the rivers, was responsible
for the large growth of green algae during 1959.
A report from Pittsburgh, Pa., (Anon., 1960)
for September 1959 states that bass, bluegills, and
minnows were returning to the Ohio, Monongahela,
and Allegheny rivers. This change was ascribed
to the temporary absence of acid wastes in the
streams while the steel mills were idled by a pro-
longed strike in the steel industry.
Determination of the effect of particular factors
on the biota of rivers will require detailed studies
that are planned for that purpose. The basic data
will provide background information giving, for ex-
ample, an idea of the range in numbers and kinds
of algae in different seasons, in different years,
and at different locations for all of the larger riv-
ers of the United States.
Appreciation is expressed to Dr. Louis G. Wil-
liams, Aquatic Biologist, Sanitary Engineering
Center, who is in direct charge of the plankton
analyses, for the planning and preparation of Fig-
ures 8 and 9.
NOV.-FEB.
1958
MAR.-JUNE
JULY- OCT
NOV.-FEB.
1999
MAIt-JUNC
JULY-OCT.
EAST LIVERPOOL
OHIO
HUNTINGTON
WEST VIRGINIA
CINCINNATI
OHIO
CAIRO
ILLINOIS
LEGEND: §§§§s AVERAGE NUMBER GENERA /mi
AVERAGE COUNT/ml
Figure 9. PLANKTONIC GREEN ALGAE — OHIO RIVER
-------
38 ALGAE AND METROPOLITAN WASTES
REFERENCES
Anon., 1958. National water quality network, an- Anon., 1960. Percolation and runoff. Jour. Amer.
nual compilation of data, October 1, 1957-Septem- Water Wks. Assn. 52(1):38,40 P&R.
ber 30, 1958. U. S. Dept Health, Educ., and Wel-
fare, Public Health Service Publication 663. Palange, R. C. and S. Megregian. 1958. Moni-
toring of stream water quality — USPHS program.
Anon., 1959. National water quality network, Sup- Jour. Amer. Water Wks. Assn. 50:1214-1219.
plement 1, Statistical summary of selected data,
October 1, 1957-September 30, 1958. U. S. Dept.
Health, Educ., and Welfare, Public Health Ser-
vice Pub. 663, Suppl. 1. .
-------
GROWTH CHARACTERISTICS OF ALGAE
-------
Nutrition
FUNDAMENTAL CHARACTERISTICS OF ALGAL PHYSIOLOGY
ROBERT W. KRAUSS
Department of Botany, University of Maryland, College Park, Md.
Control of biological communities as complex
as those encountered in natural waters is at best
a difficult and at worst an impossible occupation.
Yet increasing requirements for water have placed
a growing burden of performance on biologists re-
sponsible for maintaining water sources in the best
possible condition for human consumption and en-
joyment. The dual role of predicting and control-
ling the biological balance in natural waters serv-
ing as sources to metropolitan areas requires a
thorough appreciation of the subtleties of ecologi-
cal interaction. The consequences of changing a
rapidly moving stream into a relatively stagnant
lake, by what may appear to be the praiseworthy
conservation practice of damming, are often as
difficult to predict as the results of the obviously
deleterious practice of using a river as a drain for
organic wastes. In spite of what can be determined
about the physical and chemical characteristics of
a given water source, the degree to which it can
initiate the rapid growth of a bacterium or the
even more spectacular "bloom" of an alga depends
on the specific environmental requirements of
these microorganisms. Such requirements, con-
trary to common belief, are not only nutritional.
Accumulating evidence suggests other avenues
through which the environment can affect the
growth of microorganisms. Sound judgements of
cause and effect will be reliable in direct propor-
tion to the completeness of knowledge concerning
the nature of the individuals comprising the com-
munity. For this reason it is appropriate to focus
attention on the physiology of the organisms, es-
pecially the algae, which play a major role in de-
termining the aquatic community, and reciprocally
affect the quality of the water that supports them.
It would be most desirable if an approach to the
fundamental principles of algal physiology could be
made with the ambition and hope for intellectual
satisfaction that can be enjoyed by physicists in
developing the unified field theory. No equivalent
advance in biological theory is available for de-
veloping such insight into the physiology and bio-
chemistry of any group of organisms, much less
the algae. Except for the evolutionary concept of
Darwin, and the modern consequences of Mendel's
experimentation, biology in general, and physiolo-
gy more especially, has not been graced with the
intermeshing laws and certainties with which the
physical sciences have made such striking advan-.
ces in the control of nature. However, from the
masses of data that are recorded in the thousands
of biological publications now issued yearly, cer-
tain principles seem to be reliable enough for
science to accept with some measure of security.
In making certain principles worth stating the
student of algal physiology is faced with a bewil-
dering array of differences in the organisms he
studies. Chlorella pyrenoidosa was one alga a few
years ago, now it is many. Chlorophyll a, the sine
qua non of photosynthesis until recently, is now at
least 3 chlorophylls a and probably more. One
pathway for hexose metabolism is now ancient
history, and a given organism may have three or
more alternatives in the initial steps, not to men-
tion new modifications in the reactions of the
Krebs cycle. Much of this information is recorded
and discussed in detailed reviews and books dealing
with algal physiology (Fogg, 1953; Krauss, 1958;
Myers, 1951; and Smith, 1951). It is not the pur-
pose of this paper to dwell on these details, but
rather to discuss some of the generalities, which
might be dignified by the term principles, that can
be useful in interpreting the performance of algae,
and to mention those areas where our understand-
ing is weak, but where the rewards of knowledge
should be great in enhancing our present abilities
to predict and control the algae in natural popula-
tions.
TAXONOMY
Although it may at first seem out of place in a
discussion of algal physiology, a few comments
should be made about the classification of the or-
ganisms which we call algae. There are 7 divi-
sions of algae, each distinct enough in its own
characteristics to be worthy of independence as
major segments of the plant kingdom ranking with
the ferns, gymnosperms, and seed plants. Mor-
phology plays a role in this system of classifica-
tion, but unlike the criteria for the separation of
the groups of higher plants the more critical dif-
ferences between the divisions of algae have been
the peculiarities of their physiology. The blue-
green algae (Cyanophyta) were very early recog-
nized as different from the green (Chlorophyta),
red (Rhodophyta), or brown (Phaeophyta) algae be-
cause of their pigmentation. Less obvious, but
clearly separable because of color, were the yel-
low-green (Chrysophyta) and the golden-brown
(Pyrophyta) groups. Only the Euglenas (Eugleno-
phyta) are not primarily thought of as being inde-
pendent because of a peculiarity of pigmentation.
More careful study over the years has revealed
more and more discomforting exceptions to the
40
-------
Algal Physiology
41
initially obvious pigmentation. The blue-green al-
gae possess phycobilins essentially the same as
those in red algae, and the red algae contain the
blue bilins of the blue-greens. Even certain green
algae have been shown to contain the blue-green
pigments and the cryptomonads are occasionally
blue or red (Allen, 1959). Such modifications in
the pigmentation of individuals have not resulted in
changes in classifications but rather in an aware-
ness of the breadth of biochemical capacity of the
members of widely different groups. The nature
of the reserve substances of the algae believed to
be taxonomically characteristic has not received
the attention given the pigments, but it seems cer-
tain that further investigations in this area also
will show that the organisms have as much disre-
spect for classific ation systems in this area as in
the case of the pigments. X-ray powder diagrams
of the reserve material paramylon, long consi-
dered the characteristic of the Euglenophyta, have
demonstrated its identity with the hydroglucan of
yeast (Kreger and Meeuse, 1952) and leucosin the
Chrysophycean reserve is essentially the same as
the laminar in of the brown algae (von Stosch,
1951). Other taxonomic criteria based on physio-
logical characters too are showing weakness.
This does not mean that all individuality is ne-
cessarily lost by the breaking of man-made taxo-
nomic fences. Evidence of remarkable individual-
ity is also available. The complete absence of
brown algae from fresh waters is a characteristic
that some might devoutly wish could be shared by
other groups. The unique coupling of CO2 fixation
in either light or dark with the normal illuminated
photosynthetic intermediates by Euglena is another
case in point (Lynch and Calvin, 1953).
The situation concerning the overlapping of
physiological characteristics between the major
algal divisions is, however, of less interest here
than the problem of the identity of individual spe-
cies. It is becoming increasingly obvious that the
former classification of species particularly
among the unicellular algae, is only a general
guide to the requirements and capabilities of the
organisms which fall within the available morpho-
logical descriptions. The list of algae at the cul-
ture collection at the University of Indiana (Starr,
1960) gives some indication of the situation. Num-
erous strains, many of them different in their
growth characteristics, are listed for Anabaena,
Nostoc, Lyngbya. Chlorella, Euglena, Chlamy-
domonas, Pandorina, and others. Few of the de-
tailsof the strain differences have been thoroughly
investigated, but the degree to which strains of the
same species can differ is seen in the tabulation of
data for two strains of Chlorella in Table 11 (Soro-
kin, I960).
At present we are attempting a revision of the
classification of the entire genus Chlorella based
on physiological characters alone. It may well be
that the only satisfactory measure of individuality
in any genus of algae will be the growth perfor-
mance under rigidly controlled environmental con-
ditions in specific media.
Table 11 - CHARACTERISTICS OF GROWTH,
PHOTOSYNTHESIS, AND RESPIRATION FOR
HIGH AND LOW TEMPERATURE STRAINS OF
CHLORELLA PYRENOIDOSA
Strain of Chlorella
Characteristic
Emerson
7-11-05
Temperature optimum for:
Growth
Photosynthesis
Endogenous respiration
Glucose respiration
Growth rate at light
saturation:
at 25° C
at 39° C
Rate of apparent photosyn-
thesis at light saturation:
at 25° C
at 39° C
Rate of glucose
respiration:
at 25° C
. at 39° C
Saturating light inten-
sity for growth:
at 25° C
at 39° C
Degrees C
25-26
32-35
30
30
38-39
40-42
40-42
40-42
Number of doublings
per day
3.1
3.0
9.2
i3 cells/hour
43
rapidly
declining
47
170
« 3
mm 02/mm cells/hour
4.5
1.6
8
18
Foot candles
500
500
1400
What fundamental principle of algal physiology
does this reveal to the student of algal growth in
natural waters? It is simply that the performance
of a given organism cannot be predicted on the ba-
sis of morphological description alone. Neither
can it be predicted by references to published
physiological characteristics of that species or
even of the larger groups. Strain differences pre-
clude certain identification without thorough phys-
iological study of the organisms. A "bloom" of
Anabaena flos-aquae in two bodies of water may,
or may not, be the result of matching environmen-
tal conditions. This will depend on the physiologi-
cal identity of the strains growing in the separate
locations.
Studies of artificially induced mutants have
shown that large numbers survive in enriched me-
dia and may be peculiarly adapted to survival
there (Wetherell and Krauss, 1957). The high
number of mutations which can be expected in a
large microorganismal population should in time
permit a selection of strains peculiarly adapted to
the conditions in which they were generated. En-
vironments can be expected to select and perpe-
-------
42
ALGAE AND METROPOLITAN WASTES
tuate strains as well as to determine which spe-
cies will survive.
GROWTH RATES
Regardless of the identity of the organisms the
phenomenon most familiar to those concerned with
water supplies is the periodic rapid growth of al-
gae. The capacity for generating the population
explosion known as the "bloom" apparently is a
quiescent characteristic of the organism which
manifests itself only when conditions are ideal. In
the laboratory, in suitable media and in appropri-
ate apparatus, the maximum potential growth rate
of a species can be achieved and maintained inde-
finitely (Myers and Clark, 1944; and Myers and
Graham, 1959). While it might be unwise to give
the term normal to a perpetual exponential growth
rate, at least the cells from such cultures give
uniquely standard populations which have been use-
ful for physiological study.
In nature or, in a self contained laboratory cul-
ture, the period of maximum growth rate endures
for only a restricted period during the growth of
the alga. Figure 10 gives a typical plot of the
growth curve of an alga growing in a finite medium
in which, in time, one or more of the requirements
for growth have become limiting. This sigmoid
curve may be distorted somewhat in nature by
Figure 10. THE RELATION BETWEEN THE
PLOTS OF A TYPICAL SIGMOID GROWTH CURVE
OF AN ALGAL CULTURE AND THE GROWTH
RATE, K, CALCULATED FOR THAT SAME CUL-
TURE USING THE FORMULA, logo -£f- « K
(t2-tl). C1
special conditions, but it is essentially a model for
what one might expect during a bloom. Several
features of the curve are worth mentioning. First,
it should be recognized that the period of maximum
growth rate is given by the portion of the curve be-
tween A and B. After this time, although the cul-
ture continues to grow, the rate is actually falling.
The second curve plotted on the ordinates gives a
measure of the rate of change in the growth rate
from the optimum, during the exponential phase,
to the falling rate during the arithmetic phase.
However during this period of falling growth rate
the yield of the culture, in terms of increase in
weight or volume per unit time, is still increasing
because of the larger population base. This means
that an algal bloom may be increasing most rapid-
ly long after the conditions that initiated it have
ceased to be optimum. It is easy to see how an
analysis of waters at the time of maximum bloom
may overlook the most important factors causing
it
The fact that a plot of the growth curve of a
population of algae is smooth tends to obscure an-
other facet of the multiplication process which may
affect the course of growth. Single-celled algae
cultured in the laboratory in so called "steady
state" cultures are in all stages of maturity. Newly
released autospores and mature cells as well as
. all possible intermediates are mixed and growing
together. The rates of respiration, photosynthe-
sis and growth of the cells of different ages are not
the same. There is reason to believe that the re-
quirements of growth of cells at different ages may
also be different (Sorokin and Krauss, 1959). The
coincidence of conditions suitable for the maximum
rates of progression through all stages may be
rare in natural waters. Conversely the lack of a
suitable environment for the passage of the cells
through one stage of development may halt a bloom
when otherwise conditions are ideal.
The laboratory technique for studying the cells
at various stages is known as synchronization. By
alternating periods of light and darkness the cul-
ture can be made to adjust itself to the point where
all of the cell divisions take place at the same
time, and, consequently, all of the cells at any
given time are the same age. The success of this
technique depends on the fact that cell division is
inhibited in the light and all cells capable of divi-
sion divide rapidly in the dark (Sorokin and Krauss,
1959). The useful cycles of light and dark are of
the order of 12 hours of light and 12 hours of dark,
but the ideal time varies somewhat with the species
and with the conditions of culture. This time inter-
val is of the order of natural day/night cycles,
and it is quite possible that algae growing at their
optimum rate in nature may become synchronized
to a considerable degree. Although harder to de-
tect, synchronization can exist in the filamentous
blue-greens such as Anabaena or Rivularia, as
well as in the essentially unicellular forms such as
Microcystis.
The degree of natural synchronization and the
opportunities this facet of algal physiology offers
to the biologist interested in control are unknown
at this time. How much the suddenness of a bloom
may be due to the precipitate division of cells that
-------
Algal Physiology
43
have been accumulating the necessary metabolites
for explosive growth and division can be deter-
mined only by investigating this aspect of algal
physiology in natural populations.
AUTOTROPHIC AND HETEROTROPHIC GROWTH
A feature of the algae that is basic to an analy-
sis of their physiology is the general capacity for
both autotrophic and heterotrophic growth. The
capacity for photosynthesis sets the algae apart
from most other microorganisms, especially the
fungi. Though colorless species are found in all of
the major divisions, except the brown algae, the
use of light as an energy source and water as the
hydrogen donor (photolithotrophy) is the dominant
mode of life. Most species, if the medium is sat-
isfactory, act as facultative chemoorganotrophs
using sugars or organic acids as both the energy
source and as a source of reduced carbon. These
two basic forms of existence have variations in
numerous species in which the ability to synthe-
size certain organic moieties such as amino acids
and vitamins is limited or blocked completely.
The overall control of metabolic reaction rates
in the algae seems linked to the photosynthetic
mechanism even when the organism does not em-
ploy the light reaction in growth. Beginning with
the studies of Bristol-Roach (Bristol-Roach and
Muriel, 1928) to the present (Krauss, 1958; and
Myers, 1957) the evidence indicates a rate regula-
ting function of the light reaction. The growth rate
of algae, with the exception of such photosyntheti-
cally crippled forms as Ochromonas (Myers and
Graham, 1956), can not be accelerated above that
during photosynthesis at saturating light intensi-
ties by the addition of a source of reduced carbon.
The regulatory mechanism takes the extreme form
in those cases of obligate phototrophy where growth
of the algae is not possible regardless of the rich-
ness of the medium in the absence of light (Weth-
erell, 1958). Whether this effect is strictly a mat-
ter of production of some photosynthetic intermedi-
ate or is due to some other regulatory effect of
light cannot yet be determined.
Of similar nature is the effect of the photosyn-
thetic process on the fate of the carbon which is
fixed during the reaction. Calvin (Bassham and
Calvin, 1957) has shown that during illumination
the normal path of reduced carbon is into storage
products presumably starch and related com-
pounds. In the dark the labeled Cl4 from C14O2
quickly appears in the compounds of the Krebs
Cycle. Whether there is a gate to dark respira-
tion which is closed by light is not yet certain. The
gate is open wide enough to permit the incorpora-
tion of both exogenous reduced carbon and photo-
synthetically reduced carbon simultaneously.
Myers has demonstrated that when photosynthesis
is exogenously subsidized by glucose 50% of the
carbon incorporated by the organism comes from
glucose and 50% from CO2 (Myers, 1957).
The dependence of algae on light, and the dom-
inance and even control of the photosynthetic me-
chanism on the other aspects of the metabolism of
the algae, has often led to the assumption that the
more light the better. In fact, for even the more
hardy autotrophic species like Chlorella and Ana-
cystis the saturation level for the photosynthetic
process and for growth is well below 1000 foot can-
dles , or less than 1/10 of full sunlight. The point
at which damage to growth takes place is at, or
just above, 1000 foot candles for the more suscep-
tible species and only 2000 foot candles for those
which are celebrated as resistant to high light in-
tensities. Figure 11 gives the light intensity curves
for four algae. Although the light intensities given
B 2
o
too
400
600
FOOT
•OO 2000
CANDLES
60OO
IOOOO
Figure 11. THE GROWTH RATES OF FOUR SPE-
CIES OF ALGAL AT 25° C MEASURED AT LIM-
ITING, SATURATING, AND INHIBITING LIGHT
INTENSITIES. The symbols are: Chlorella pyre-
noidosa (van Niel), circles; Chlorella vulgar is,
crosses; Scenedesmus obliquus, triangles; and
Chlamydomonas reinhardti, squares. Open sym-
bols show growth under fluorescent light. Closed
symbols show growth under incandescent light.
will vary somewhat with the geometry of the appa-
ratus in which the measurements are taken, it is
clear that most of the radiant energy of full sun-
light is either not used or is actually damaging to
cell growth.
Such data, which should be supplemented by that
for other species, coupled with measurements of
light intensities at various depths could be useful
in computing the theoretical total production capa-
city of a given body of water during a period of
bloom. It would be instructive to know how great a
contribution to the total weight of the algae in a
given situation was produced by the heterotrophic
vs. the autotrophic form of metabolism. Such in-
formation could serve to identify the fundamental
cause of an algal bloom.
-------
44
ALGAE AND METROPOLITAN WASTES
Not only is the source of the carbon of the algae
a fact of particular importance, but the reaction
sequence by which the algae convert the photo-
synthate or absorbed organic carbon to energy
yielding processes is perhaps of even greater con-
cern. Biochemical studies of algal metabolism
have demonstrated alternative pathways for the
utilization of organic substrates. The classical
Meyerhoff-Embden system for phosphorylation and
degradation of hexoses now has two alternative
routes established for this process — the glucu-
ronic or gluconic acid shunt (Table 12) and the
galactose-6-phosphate route (Figure 12) (Galloway
and Krauss, 1959b). Each of these routes for the
metabolism of sugar has an entirely different bat-
Table 12. REACTIONS OF THE GLUCONIC ACID
SHUNT
H. -Ol«l»
C—-j
HCOM I
CM.OM
ctvcosr -i- pws
«t o«
MCOM I
HOCH 0 I
MCO» I
HC '
CH.OH
tiucase
'?"—I
HCOH I
HOC" 0
HCOM !
HC 1
HC-
CM,OI>O,-
fLUCOiC-t-
CM,O«
MC^
HOCHN,
-«- o •
•*?"J
MC - 1
C-.OM
f -i- mwsMurr
M. .OM
MCOM |
• MOCM 0
HOCH
HC '
CM, OH
UHCTOSl
I.
HCOM I
MOCH O ••-
MOCH
CH.OPO,-
C4lACTOSf-S- PMOStMAlt
HC -
CM.OPO,'
mvcnsc-t-rmsfiarc
OCH I
HC '
CH.C
-t-f
!
CH,OK
• MOCH O
HC—J
CM.OK),'
6C6H120
60] -
6H20
4C$H1005 2C7Hl4<>7 «• 2CjH6<>3 2C6H1206 + 2C4H804
peotoae heptose trlose bexose tetrose
3C6H1206
tetrose bexose
CH.OPO,-
CM.OFO,*
"?v^ "9-«.
MOCM^ i^ *. MOCHAS
MCOM O ~- HOCH 9
HC 1
rrvcftfl T.t &r*osf»trr
• MOCM
MC
nttroit-i.i-
tery of enzymes. It seems clear that certain algae
utilize one or another of these pathways preferen-
tially, and may, indeed, be obliged to use only one.
Similar alternatives exist in the systems involved
in the synthesis and degradation of nitrogen com-
pounds within the algae.
It is likely that the variability of these enzyme
systems is largely responsible for the differential
selectivity of the chemical agents that have been
utilized to control algae. Studies have shown that
the susceptibility of an alga to a given toxic chemi-
cal may be due to the destruction of certain en-
zymes. Susceptible algae are unable to bypass
such blocks, but algae which do have the enzymes
for different or alternative pathways can survive at
high concentrations of the toxic agent. Evidence
for just such an explanation for differential toxi-
city has been obtained in our laboratory in studies
of the susceptibility of phosphoglucose isomerase
(Galloway and.Krauss, 1959a; Galloway and Krauss,
1959b). The enzymatic reactions of the photosyn-
thetic cycle proposed by Calvin (Bassham and Cal-
vin, 1957) might also be examined in a similar
fashion. The characteristics of the unique enzyme,
carboxydismutase, in the initial step of carbon
dioxide incorporation into the pentose cycle of
photosynthesis should be better known. This one
key biochemical reaction is characteristic of pho-
tosynthesizing algae and is a pregnant subject for
study by those with control as an ultimate aim.
It is not possible to cover all the aspects of the
photosynthetic process. Reviews, some especially
related to the algae, may be found elsewhere
(Bassham and Calvin, 1957; Krauss, 1956; and
CH,0«>,- CM.OM
«trcr»M- tHxienoir
ffurtf-l tccnxr
Figure 12. THE SEQUENCE OF HEXOSE INTER-
MEDIATES OF AN ALTERNATE PATHWAY IN
RELATION TO THE CLASSICAL GLYCOLYTIC
SCHEME.
Anon., 1958). In fact most of the current investi-
gation in this area is moving into the realms of
solid state physics. However, one aspect of the
process which has been uncovered in recent years
and is especially pertinent here is the identification
of the process of cyclic photosynthetic phosphory-
lation. The intricacies of this process are in-
volved, but can be studied in detail in the papers of
Arnon (Arnon, 1959), Jagendorf (Jagendorf, 1959)
and others. The outstanding feature of this system
is that ATP can be generated in the chloroplast
through a cyclic process by which water is first
split into H4* and OH~, and then, after passage
through certain enzymes, is recombined to yield
water again. This process, quite different from
the Hill reaction which provides the reducing power
for CO£ reduction and liberates oxygen, results in
no oxygen liberation at alL Nevertheless, although
no gas exchange can be measured, electromagnetic
energy is converted to chemical energy for use by
the plant, How extensive this system may be and
what portions of the ATP requirement are satisfied
in this fashion are not established. Nevertheless it
is certain to be of major significance in the energy
economy of growing populations, and must be taken
into account in the calculations of efficiency. Gas
measure of the energy exchange can no longer be con-
sidered the exclusive trapping activity of the cell.
-------
Algal Physiology
45
NUTRITION
Studies of algae concerned with their growth in
nature have been strongly oriented toward inorgan-
ic nutrition. The availability of good data has not,
however, always made the picture clearer. At-
tempting to decipher the producing capacity of a
body of water based on its nutrient composition is
as difficult as determining what the algae require
in terms of their internal composition to insure
optimum growth. Apparent contradictions in the
data stem from three problems related to algae
growth. The first deals with the selection of the
most useful criterion for measuring growth; the
second relates to the availability of nutrients in
waters; and the third involves the estimation of re-
quirements when more than one of the factors af-
fecting growth are in rate-limiting supply.
With regard to the first and third of these it is
important to recall that there are various ways of
evaluating growth in microorganisms. An alga,
blocked from further protein synthesis by lack of
nitrogen, may continue to synthesize carbohydrate
reserves. The capabilities of the cells in this re-
gard have been well tabulated (Spoehr and Milner,
1949; and Thomas and Krauss, 1955). In a real
sense this cell is still growing. In fact, as it shifts
from protein synthesis to carbohydrate or even fat
synthesis, the increase in cell weight may be re-
duced very little. The amount of light reaching the
cell may actually be more influential in determin-
ing the amount of cell growth than the nitrogen
limitation. This becomes increasingly involved
when the factor limiting growth is not limiting
growth absolutely, but only reducing the rate of
growth. Then as the cell grows in an environment
shifting from darkness to damaging light intensi-
ties, as it shifts to heterotrophic or autotrophic
growth, and as its synthetic directions change, the
degree to which each of the factors can be said to
be the "limiting" one becomes almost impossible
to determine. A sophisticated system of integra-
tion, perhaps employing computers, would be ne-
cessary for an effective solution. Nor is it even
safe to say what the minimum levels of a given es-
sential metabolite may be for the different forms
of nutrition. An excellent example is the low re-
quirement for manganese shown by algae when
growingheterotrophically in contrast to a require-
ment, at least two times higher, for the same or-
ganism obtaining its reduced carbon by photosyn-
thesis (Reisner and Thompson, 1956). Such consi-
derations are borne out by the difficulty encountered
by Gerloff and Skoog (Gerloff and Skoog, 1954; and
Gerloff and Skoog, 1957) in deciding the degree to
which nitrogen limits growth in the notorious Wis-
consin lakes.
The second problem has been of interest for a
number of years. It concerns the way in which the
algae absorb their inorganic components from the
medium. The elemental requirements for the algae
are fairly well worked out and can be shown in
comparison to those of other microorganisms in
Table 13. These requirements have been described
in detail in several reviews (Ketchum, 1954; and
Pirson, 1955). However the presence of a given
Table 13. CHEMICAL REQUIREMENTS OF MI-
CRORGANISMS
Element
Well established
and general:
C,H,0
N, P, S
Mg
Ca
Co
Cu
Fe
Me
K
Zn
Algae Fungi Bacteria Protozoa
7
?
7
7
7
?
?
?
Poorly established
or less general:
B
Ga
Mo
Si
Na
Va
Sr
Rb
* Signifies demonstrated essentiality
*? Signifies essentiality in doubt although a require-
ment has been demonstrated in some species.
element in the medium does not in itself assure
that the algae can utilize this element. Though ex-
ogenous proteolysis and amino acid sources for
nitrogen are common in the algae, even nitrogen
in some organic combinations is unavailable. This
can be often overlooked when waters show a high
nitrogen analysis. In a similar fashion numerous
phosphate compounds, even the polyphosphates,
are not suitable for algal growth.
The micronutrients which are just as important
to algae as nitrogen in spite of their quantitatively
inconspicuous characters are probably most sub-
ject to conditions which make them unavailable.
The shift from soluble molecules to colloidal ag-
gregates may be too subtle to be detected by ordi-
nary analytical means, but is widely accepted as
the cause of failure of discrete media to support
optimum growth in laboratory cultures. Routine
dictates the use of some type of chela ting agent in
the preparation of all media now used in studies of
algal physiology. Nevertheless relatively little is
known about the levels and effectiveness of chela-
ting agents found in nature, and these may deter-
mine the availability or lack of availability of an
element much more than elementary analyses in-
dicated.
Of recent concern has been the mechanism of
exchange by which chelated trace metals are ab-
sorbed by algae. It appears that chelates may re-
lease the metals to the cell by weak though pro-
gressive dissociation; that the chelates may be
destroyed by such agents as light thereby liberat-
-------
46
ALGAE AND METROPOLITAN WASTES
ing their metal complement; or the entire chelate
complex may be absorbed and metabolized by the
algae (Krauss and Specht, 1958). Further investi-
gations of the mechanism of chelate action should
prove useful in understanding their role in support-
ing algal growth in nature as well as in the labora-
tory.
ALGAL DECOMPOSITION AND ALGAPHAGE
So far we have directed attention to the factors
in the environment responsible for algal growth and
to the ways in which the algae utilize these factors
in their metabolism. The final question is what
happens to the algae after they have grown and ex-
hausted the supply of whatever has generated
them? In a pure laboratory culture a mature popu-
lation may live for months intact in a condition
characterized by extremely low endogenous respi-
ration. Even if the culture is exposed to outside
bacteria it may continue to exist indefinitely. This
is not the case in nature. Algal blooms do disap-
pear and this very fact is some cause for hope in
their control. Of course in moving waters the
wash-out rate will account for the rapid dissolution
of a bloom once growth ceases, but they disappear
almost as rapidly in relatively stagnant bodies.
One should expect that if the algae decomposed that
they would liberate into the water those same pro-
ducts that gave them life and the cycle would regen-
erate. In general this is not the case and it seems
likely that we must look to the activity of other or-
ganisms for the answer.
Grazing by larger animals undoubtedly accounts
for the consumption of some of the bloom, but most
of the algae are disposed of by other microorgan-
isms. Little is known of algal pathology. With the
exception of the Chytrids no parasite has been iden-
tified on them. A great deal of laboratory experi-
ence does show that pure cultures of algae con-
taminated with bacteria or fungi grow poorly, but
in fact many grow very well on inorganic media
with bacteria and some even grow better when con-
taminated. Anaerobically the algae decompose
rapidly (Golueke, Oswald and Gotaas, 1957) and
this is the fate of some of the bloom which has
settled into the bottom mud. However in the light
of what we know about microorganismal infection
of other higher plants, animals, and microorgan-
isms it seems that a very large vacuum exists with
regard to our knowledge of this phase of algal
growth. Especially pertinent is the question of
whether phages play a role in the degeneration of
algae in nature.
It is not the purpose of this paper to make more
than a general comment in this regard. However it
seems almost impossible to avoid the conclusion
that phages or related viruses must infect algae.
About this subject which could be of the greatest
practical value we know absolutely nothing. Some
experiments are being performed in our laboratory
on a limited scale exploring the techniques of phage
identification and culture, but results are too new
to warrent further statements. From the point of
view of the physiologist few things could be more
exciting than being able to study the nucleic acid-
protein relationships of algae and infecting phages.
Much of what we know of inheritance and DNA-RNA
mechanisms in other organisms have come from
experiments with viruses. It will be a distinct
service to algal physiology and to practical aquatic
biology when algaphages are finally identified and
examined.
CONCLUSION
The general impression that these remarks were
intended to convey is that there are sound begin-
nings to the understanding of the ways of algal life.
Already the exploitation of what is known should be
a help to the practical biologist. Even greater re-
wards must come from a study of the algae for the
sake of our own intellectual curiousity.
REFERENCES
Allen, M. B. 1959. Studies with Cyanidium cal-
darium, an anomalously pigmented chlorophyte.
Archiv fur Mikrobiologie 32: 270-277.
Anon., 1958. The photochemical apparatus its
structure and function. Report of Symposium held
June 16-18, 1958. Biology Department, Brook-
haven National Laboratory, Upton, N. Y.
Arnon, D. L 1959. Chloroplasts and photosynthe-
sis. In: The Photochemical Apparatus Its Struc-
ture and Function, p. 181-235. Brookhaven Nation-
al Laboratory, Upton, N. Y.
Bassham, J. A. and Melvin Calvin. 1957. The path
of carbon in photosynthesis. Prentice Hall Inc.,
Englewood Cliffs, N. J. 104 pp.
Bristol-Roach and B. Muriel. 1928. On the influ-
ence of light and of glucose on the growth of a soil
alga. Ann. Bot. 42: 317-345.
Fogg, G. E. 1953. The metabolism of algae. Me-
thuen and Co. Ltd., London. 149 pp.
Galloway, R. A. and Robert W. Krauss. 1959a.
The differential action of chemical agents, espe-
cially Polymyxin B, on certain algae, bacteria,
and fungi. Am. Jour. Bot 46: 40-49.
Galloway, R. A. and R. W. Krauss. 1959b. Mechan-
ism of action of Polymyxin B on Chlorella and
Scenedesmus. Plant Physiol. 34: 380-389.
-------
Algal Physiology
47
REFERENCES (ContU)
Gerloff, Gerald C. and Folke Skoog. 1954. Cell
contents of nitrogen and phosphorous as a measure
of their availability for growth of Microcystis
aeruginosa. Ecology 35: 348-353.
Gerloff, Gerald C. and Folke Skoog. 1957. Nitro-
gen as a limiting factor for the growth of Micro-
cystis aeruginosa in southern Wisconsin Lakes.
Ecology 38: 556-562.
Golueke, C. G., W. J. Oswald, and A. B. Gotaas.
1957. Anaerobic digestion of algae. Applied Mi-
crobioL j>: 47-55.
Jagendorf, A. T. 1959. The Relationship between
electron transport and phosphorylation in spinach
chloroplasts. In: The Photochemical Apparatus
Its Structure and Function, p. 236-258. Brook-
haven National Laboratory, Upton, N. Y.
Ketchum, B. H. 1954. Mineral nutrition of phy-
toplankton. Ann. Rev. Plant Physiol. J5: 55-74.
Krauss, R. W. 1956. Photosynthesis in the algae.
Ind. and Eng. Chem. 48: 1449-1458.
Krauss, R. W. 1958. Physiology of the fresh wa-
ter algae. Ann. Rev. Plant Physiol. £: 207-244.
Krauss, R. W. and Alston Specht 1958. Nutrition-
al requirements and yields of algae in mass cul-
ture. In: Transactions of the Conference on the
Use of Solar Energy ~ The Scientific Basis, p.
12-26. Edwin F. Carpenter, Editor. Vol. IV.
Photochemical Processes. University of Arizona
Press, Tucson, Arizona.
Kreger, D. R. and B. J. D. Meeuse. 1952. X-ray
diagrams of Euglena paramylon, of the acid-insol-
uble glucan of yeast cell walls, and of Laminarin.
Biochim. et Biophys. Acta 9: 699-700.
Lynch, V. H. and M. Calvin. 1953. CO2 fixation by
Euglena. Ann. N. Y. Acad. Sci. 56: 890-900.
Myers, J. 1951. Physiology of the algae. Ann.
Rev. Microbiol. 5_: 151-180.
Myers, J. 1957. Suppression of carbon dioxide
assimilation by glucose in Chlorella. Plant Phy-
sioL 32_: Suppl. xvi-xvii.
Myers, J. and L. B. Clark. 1944. Culture condi-
tions and the development of the photosynthetic me-
chanism. II. An apparatus for the continuous cul-
ture of Chlorella. J. Gen. Physiol. 28: 103-112.
Myers, J. and J. R. Graham. 1956. The role of
photosynthesis in the physiology of Ochromonas.
J. Cell and Comp. Physiol. 47: 397-4131
Myers, J. and J. R. Graham. 1959. On the mass
culture of algae. II. Yield as a function of cell con-
centration under continuous sunlight. irradiance.
Plant Physiol. 34: 345-352.
Pirson, A. 1955. Functional aspects in mineral
nutrition of green plants. Ann. Rev. Plant Physiol.
6: 71-114.
Reisner, G. A. and J. F. Thompson. 1956. Man-
ganese deficiency in Chlorella under heterotrophic
carbon nutrition. Nature 178; 1473-1474.
Smith, G. M. (Editor) 1951. Manual of phycology.
375pp. Chronica Botanica Co., Waltham, Mass.
Sorokin, C. 1960. Tabular comparative data for
the low and high temperature strains of Chlorella.
Nature 184: 613-614.
Sorokin, C. and R. W. Krauss. 1959. Maximum
growth rates of Chlorella in steady state and in
synchronized cultures. Proc. NatL Acad. of Sci.
(U. S.) 45: 1740-1744.
Spoehr/H. A. and H. W. Milner. 1949. The chemi-
cal composition of Chlorella; effect of environmen-
tal conditions. Plant Physiol. 24: 120-149.
Starr, Richard C. 1960. The culture collection of
algae at Indiana University. Am. Jour. Bot. 47:
67-86. ~
Thomas, W. A. and R. W. Krauss. 1955. Nitrogen
metabolism in Scenedesmus as affected by environ-
mental changes. Plant Physiol. 30: 113-122.
von Stosch, H. A. 1951. Uber das Leukosin, den
Reservestoff der Chrysophyten. Die Naturwissen-
shft 38: 192-193.
Wetherell, D. F. 1958. Obligate phototrophy in
Chlamydomonas eugametos. Physiologia Plantar-
urn 11: 260-274.
Wetherell, D. F. and R. W. Krauss. 1957. X-ray
induced mutations in Chlamydomonas eugametos.
Am. Jour. Bot 44: 609-619.
-------
48 ,
ALGAE AND METROPOLITAN WASTES
MICRONUTRIENTS AND HETEROTROPHY AS POSSIBLE FACTORS
IN BLOOM PRODUCTION IN NATURAL WATERS
LUIGI PROVASOLI
Haskins Laboratories, 305 E. 43rd Street, New York, N. Y.
INTRODUCTION
The heterotrophic abilities and heeds of the
algae are receiving increased attention. Since the
algae are at the origin of the vegeto-animal clea-
vage' they offer a variety of nutritional potencies,
from obligate photoautotrophy to facultative heter-
otrophy and to phagotrophy (consult Lwoff, 1943;
and Provasoli and Pintner, 1953b, for evolution-
ary trends). In our discussion we will consider
only the transition phototrophy-osmotrophic het-
erotrophy without discussing phagotrophy. The
word heterotrophy, of nebulous etymology (het-
ero - other), is now generally accepted to signify
a need for preformed organic molecules. The
early distinction between need of organic carbon
sources and "building blocks" has become unten-
able because of the complexity of the web of path-
ways of synthesis. Though the need for growth
factors is clearly heterotrophy, a new name, auxo-
trophy, has gained recognition. It seems oppor-
tune to accept it, especially in the algae, because
this need is often the only heterotrophic need, is
independent of the source of energy utilized, and
is quantitatively extremely small in contrast to the
large heterotrophic needs for carbon sources and
building blocks. For clarity we will consider auxo-
trophy and heterotrophy separately.
ORGANIC MICRONUTRIENTS
For many years the algae were considered to
require only inorganic salts for growth. We know
now mat many of them require vitamins.. Though
photoautotrophic algae requiring growth factors de-
pend quantitatively almost solely on inorganic nu-
trients, the micro-requirement for growth factors
is as important because of the absolute nature of
this requirement, hence the necessity of consider-
ing them as ecological factors.
Most of the ecologically important species, both
marine and freshwater, have not been cultured and
we do not know if they are auxotrophic or not.
Some guiding principles can, however, be postulat-
ed because the phototrophic algae are quite stereo-
typed in their vitamin requirements. It is remark-
able that the trends perceived several years ago on
about two-dozen auxotrophs (Provasoli and Pintner,
1953b) have remained the same for a much larger
number of species. The sampling is still very
small, but the apparent constancy of trends lends
weight to them. The available data were tabulated
in a recent review (Provasoli, 1958). Since then
J. and R. Lewin (1959) have studied the require-
ments of 25 species of marine diatoms, Droop
(1959) of Oxyrrhis marina, McLaughlin and Zahl
(1959) of two symbiotic dinoflagellates, Fries
(1959) of a red alga, and E. G. and O. Pringsheim
(1959) of 23 species of Volvocales. Table 14 sum-
marizes the old and new data. All the phototrophic
species and their related counterparts (Chilomon-
as, Polytoma, Astasia, etc.) require only vitamin
Bi2» thiamine and bio tin, alone or in various com-
binations. This restriction to only three vitamins
is unexpected because the other groups of micro-
organisms (bacteria and molds) have widely diverse
vitamin requirements. The specificity of the algae
can hardly be fortuitous because they colonize en-
vironments rich in all vitamins. In contrast, pha-
gotrophic co lor less species, such as Peranemaand
Oxyrrhis, need additional vitamins and building
blocks. This narrow selection appears, then, as
a peculiarity of the numerous algae and flagellates
of the transition "chlorophyte-leucophyte" (sensu
Lwoff, 1943). Whatever the reason, the task of
theecologist and the water engineer is then limited
to the charting of these three vitamins in the wa-
ters.
The general order of incidence (Table 14, total
of single vitamins) appears to be cyanocobalamin
(vit. BIZ), thiamine, and biotin. Even though most
Table 14. VITAMIN REQUIREMENTS OF ALGAE
Algal group
Chlorophyeeae
• Bugteninae
Cryptophyceae
Dtooophyceae
Carysophyceae
BaciUariophyeeae
Cyanopnycese
Rhodopbyceae
Totals
Totals for stack
TttamiBS
Handier
of
species
58
9
9
17
13
37
10
1
154
No
Titamin
requirement
25
0
0
1
1
20
9
p
56
Require
vitamins
33
9
9
16
12
17
1
_!_
98
B«
3
2
2
11
3
10
1
1
33
83
TWainlnc
6
1
2
1
3
"Is"
63
Bl2 + Biotin +
Biotin ^Mnfflifrg thi^BfilfHf
24
6
5
1 1
5 1
4
45 2
7
BJ2 * biotin
+ thiamine
3
2
_^_
5
-------
Nutrients in Bloom Production
49
algae require Bj2> it is dangerous to attribute
more ecological importance to this vitamin as a
possible bloom factor, than to thiamine and biotin.
The role and relative importance of the vitamins
can only be assessed when we know in detail their
seasonal and spacial fluctuations, (i.e. the balance
between producers and consumers) and we corre-
late them with the growth of the different algae in
the waters.
The general trends are:
1. No correlation seems to exist between need
of vitamins and the ability of algae to employ vari-
ous sources of energy. Strictly photoautotrophic
species, such as several marine dinoflagellates,
the fresh-water Woloszynskia limnetica, Synura
petersenii, S_. caroliniana, and several marine
Crypto monads need vitamins just as do photosyn-
thetic species endowed with developed heterotro-
phic abilities, such as several freshwater Euglena,
Phacus, Trachelomonas, and Ochromonas. The
permanent loss of chlorophyll (colorless species)
is not necessarily accompanied by loss of the
power of synthesizing vitamins. Polytoma uvella
and P. obtusum do not need vitamins, while other
colorless Polytoma, Polytomella coeca and Chilo-
monas paramoecium are auxotrophic. Thus auxo-
trophy represents a loss of function which occurs
independently.
2. No correlation can be found between environ-
ment and need for vitamins. Synura needs vitamins
as well as Euglena, Volvox, Trachelomonas, etc.,
which colonize polluted waters. Asterionella and
Tabellaria do not need vitamins, but neither do the
mesosaprobe Fragillaria and Chlorella which can
live in the oxidation ponds. Furthermore, the
marine, brackish, and freshwater species belong-
ing to the same algal group have apparently the
same order of incidence of auxotrophic species.
3. On the contrary, there is a definite homo-
geneous trend in each algal group. The Cyanophy-
ceae, the Bacillariophyceae, and the Chlorophy-
ceae are the algal groups in which photoautotrophy
probably predominates and the need for vitamins
is restricted to fewer species. In our table we have
included only the species of Chlorophyceae and
Cyanophyceae for which the vitamin requirements
have been specifically studied. This requires a
series of precautions for chemical asepsis which
is not normally employed in the maintenance of
cultures. We have excluded a few hundred Chloro-
phyceae and 20 or more species of blue-green al-
gae which are maintained in various culture col-
lections on mineral media in cotton plugged tubes.
Cotton during sterilization releases vitamins and
Robbins et a_L (1951) have shown that microorgan-
isms grow and produce appreciable vitamin Bj2 in
distilled water allowed to stand for a few days in
the laboratory. Nonetheless, judging from the gen-
eral trend of the well-studied species, it is most
probable that the majority of the Chlorophyceae
and Cynanophyceae not included in our table does
not require vitamins. The blue-green algae, Ana-
baena cylindrica, A. variabilis, Nostoc muscorum
(1013J Wisconsin), Anacystis nidulans, Phormidi-
um autumnale do not require vitamins; only Phor-
midium persicinum requires cyanocobalamin.
Robbins et al. (1951) found that Plectonema nosto-
corum, Aphanizonemon flos -aquae, Diplocystis
aeruginosa, and Calothrix parientina produce
(the uninoculated mineral medium was assayed and
had no
Conversely, in all the other algal groups, nearly
the totality of the species so far studied needs vita-
mins. It seems safe to expect that most Eugleninae,
Dinophyceae, Chrysophyceae, and Cryptophyceae
are auxotrophic. We believe that the present samp-
ling, though small, is sufficiently representative
because the newly studied species were not pre-
selected by the choice of media. They were ob-
tained bacteria -free either directly from nature
or from bacterized cultures, and grown initially
in media enriched by a mixture of the known B
vitamins.
The data indicate that auxotrophic species pre-
dominate in algal groups with strong animal ten-
dencies (high incidence of colorless, holozoic
forms, predominance of the flagellated or ame-
boid species) like the euglenids, dinoflagellates,
chry so monads, and cryptomonads and that few
species require vitamins in the algal groups which
have well developed vegetal tendencies. This
might explain why the need for vitamins is corre-
lated with algal groups and not with environments.
4. Minor trends that might not be at all signi-
ficant with a larger sampling are that: a) the thia-
mine requirement is more evident in the Chloro-
phyceae, while the need for vitamin Bj2 predom-
inates in the other algal groups; _b) both 6^2 and
thiamine are required by the majority of the Eug-
leninae, Cryptophyceae and Chrysophyceae; c)
most Dinophyceae require only B}2-
ECOLOGY OF VITAMINS
The above data clearly indicate that Bi2, thia-
mine, biotin, and perhaps other unknown growth
factors, are ecological factors which we cannot
afford to neglect. The main problem, then, is how
to measure them. No difficulty stands in our way
for the measurement of vitamins in freshwaters.
Vitamin 612 is routinely measured in blood, urine,
and other organic fluids and organs, and many
hospitals now have an algological section with tem-
perature-controlled rooms and fluorescent light -
banks, because cyanocobalamin is titrated with the
freshwater algae Euglena gracilis and Ochromonas
malhamensis. Freshwaters are easier to analyze
than blood and other organic fluids yet no ecologi -
cal laboratory is equipped with a bioassay section.
-------
50
ALGAE AND ME TROPOLJTAN WASTES
Curiously, the little advance achieved has been for
the marine environment where salinity makes
things difficult. The assay of vitamin Bj2 in sea-
water has been done by using the above-mentioned
freshwater Euglena and reducing interference from
inorganic salts either by dialysis (Provasoli and
Pintner 1953, Kashiwada et al. 1957), by extract-
ing the 612 (Cowey 1956), by diluting the sea water
(Daisley, 1958), or by employing marine organisms
for the assay(Adair and Vishniac 1958). This pro-
blem does not exist for freshwaters; Euglena gra-
cilis and Ochromonas malhamensis can be em-
ployed reliably for measuring the various cobala-
mins present in the waters and many suitable
bacteria for bioassay are available for thiamine
and biotin. In fact, long ago Hutchinson (1943) and
Hutchinson and Setlow (1946) measured thiamine,
biotin, and nicotinic acid in Lindsley Pond and Ban-
tam Lake. A recent paper by Hutner et aL (1958)
covers adequately the methods, selection of bio-
assay organism, and known pitfalls.
We will discuss now what is Known of the cycle of
vitamins in waters. The situation being on the
whole similar, data will be taken from work done
both on freshwaters and the sea. Bacteria and
other microorganisms are the main producers of
vitamins in nature.
A great part of the vitamins in freshwaters and
in the littoral zone of the sea can be assumed to
come from any soil run-off especially during the
spring floods. The Lochhead school (Burton and
Lochhead, 1951; and Lochhead and Thexton, 1951)
found that 65% of the actinomycetes and 70 to 84%
of the bacteria in the soil produce vitamin Bj2 and
that 14% required soil extract for growth (for 50%
of these bacteria the soil extract requirement could
be met by Bj^). The predominance of Bj^ produ-
cers over 612 consumers in the soil should result
in an accumulation of Bj2- This is actually so:
Robbins etaL (1950) found 1-8 ug. of Bj2 per liter
of cold water soil extract; soil extract has been a
requirement for all sorts of algae and its effect can
often be duplicated by vitamins plus trace metals.
Muds are another source of vitamins. Starr
(1956) and Burkholder and Burkholder (1956) fol-
lowed the fate of vitamin B12 from a Spartina
marsh into a tidal river, a sound, and then the
open sea on the coast of Georgia. The vitamin is
concentrated by adsorption on the suspended parti-
cles which flow toward the sea; it originates in the
marsh where the content of Bj2 in the mud is 0.7
ug/g dry matter. Burkholder and Burkholder
(1956) found that suspended solids in brown waters
from several Georgia rivers had an extremely
high content of 812 in October (1-6 ug Bi2/gdry
'weight). Depending upon the amount of particles
suspended in the river waters, the B}£ content per
liter varies from 2-40 mpg. The same authors
(1958) found 0.4-3 ug. of vitamin Bi2 per gram of
suspended particle in the phosphorescent bay of La
Parguera, Puerto Rico. The sediments of the same
bay had an average content of 280 mpg. of Bj2> ?
mug. of biotin and 73 mpg. of thiamine per gram of
dried mud. Starr et al. (1957) found that 70% of
34 isolates of marine bacteria produced various
cobalamins. Burkholder (1959) studied the produc-
tion of B vitamins by 344 bacteria isolated from
waters and muds of Long Island Sound. They found
that when cultured with appropriate enrichments,
27% of these bacteria give off vitamin 612, 50%
biotin, 60% thiamine, and 11% nicotinic acid. The
same picture holds for freshwaters: Robbins et aL
(1950) report that fungi and many bacteria, isolated
from the water and mud of a pond in which Euglena
blooms, produce Bj2; they also demonstrated that
these bacteria, grown with Euglena on agar plates
of a medium deprived of Bi2> diffused enough vita-
min to support growth of Euglena.
The exchange of nutrients between microorgan-
isms and algae can be quite direct as in the case of
filamentous algae and seaweeds which produce a
mucilaginous slime inhabited by bacteria and yeast
Most bacteria epiphytic on seaweeds produce B}2
(Ericson and Lewis 1953).
A third source is the vitamins present as so-
lutes in waters. For 812, the ratio between the
quantity absorbed on suspended particles and the
quantity dissolved in seawaterwas about 2.5 (Burk-
holder and Burkholder, 1958). The quantity of dis-
solved vitamin Bj2 in sea water varies greatly.
Coastal waters are the richest: Lewin (1954) and
Droop (1955) found 5-10 mug Bj^/liter; Kashiwada
et al. (1957b) found up to 55 mpg/liter in the fer-
tile Kagoshima Bay. In the open waters the content
is far less: Cowey (1956) found in the North Sea
and Norwegian deeps values from 0.1-2 mpg/liter
and Kashiwada et aL (1957) in the North Pacific,
values fluctuating from 0 to 1 mpg/liter, for sur-
face waters.
The data on vitamin content in freshwater are
even scantier. Thiamine varied seasonally in
Lindsley Pond from 0.1-0.2 pg/liter; lower values
(0.03-0.04 pg/liter) were found in Waramung and
Bantam Lakes, (Hutchinson, 1943); biotin in Lind-
sley Pond varied from 0.3-4 mpg/liter, (Hutchin-
son and Setlow, (1946); Benoit (1957) found 80 mpg/
liter of Bj2in the same pond, and Kashiwade et aL
(1957b) found variations between 0-10 mpg Bi2/li-
ter in the first ten meter depth of Lake Ikedo.
The seasonal variations in vitamins found by
Hutchinson were also found by Cowey in sea water:
the dip from 2 mpg/liter in March to 0.3 mpg/liter
in May-June coincides with the diatom blooms.
Similarly Vishniac and Riley (1959) record for
Long Island Sound in the early spring a fall in Bj2
which parallels the consumption of NO3- However,
vitamin 612 even at the low value (4.5 mpg/liter)
appears not to be limiting. On the contrary, thia-
mine is barely detectable in Long Island Sound
-------
Nutrients in Bloom Production
51
(0-20 mug/liter) and they suggest that thiamine is
derived from land drainage (63 mug/liter at the
breakwater of Indian River) and is diluted as the
river mixes with sound waters, or that thiamine
is destroyed in alkaline seawater more rapidly
than it is produced. Even before the data of Vish-
niac and Riley, which tend to support Droop's
opinion, Droop (1957) concluded that vitamin Bi2
is not limiting in the sea because it is always pre-
sent in quantities far exceeding those needed by
Skeletonema costatum. In fact many species of
algae requiring B^2 are very sensitive and require
in vitro quantities between 0.1-5 mug/liter. Very
sensitive organisms could then still grow in the de-
pleted summer waters of the North Sea which con-
tain 0.1 mug/liter (Cowey, 1956) and bloom in the
lowest values (4.5 mug/ liter) found in Long Island
Sound.
However, Daisley (1957) pointed out that in na-
ture the rate of cell division controlled by the vari-
ous levels of Bi2was more important than the pos-
sible total yield. Another important consideration
is that data in vitro of cell yield per given concen-
tration of 612 are determined in a system lacking
the competition of other organisms requiring vi-
tamin 612; in nature, on the contrary, we have
such a competition. The competition for vitamin
Bj2 is further complicated by two other peculiari-
ties.
1. Holm-Hansen et al. (1954) have shown that
blue-green algae not requiring Bi2 employ 612
readily as a cobalt source; as cobalt is generally
scarce in water, even organisms not requiring Bj2
may compete for it
2. Several forms of cobalamins are produced by
bacteria; Burkholder and Burkholder (1958) have
shown that both in Puerto Rico and Long Island
muds Bi2 analogs predominate over cyanocobala-
min ( ="true"Bi2 = antipernicious anemia factor).
Droop et al. (1959) show that most auxotrophic al-
gae utilize only cyanocobalamin, while several dia-
toms utilize all cobalamins; the ratio of cyanoco-
balamin total cobalamins therefore becomes ecolo-
gically important.
These intricacies call for far more detailed
studies. Since the assay organisms which permit
differential assay for cyanocobalamin and the other
cobalamins are freshwater organisms, a fresh-
water location would be preferred. A further ad-
vantage is that we can select oligotrophic and eu-
trophic locations where the runoff, river contri-
bution, flow, and circulation can be followed more
easily. Introduction of radioactive Co may help
to trace the production of cobalamins by micro-
organisms and their consumption byphytoplankton,
zooplankton, and other microorganisms. The sit-
uation for biotin and thiamine is even more vague
because of the scarcity of data: thiamine seems,
however, to be a limiting factor.
The water environment favors the exchange of
"external metabolites" postulated by Lucas (1955),
creating a very complex situation. The cycle of
vitamins may be far more intricate than that of
phosphorus and the other scarce and needed metab-
olites. Not only do the algae compete for vitamins
between themselves but against other microorgan-
isms which, like the bacteria, reporduce and me-
tabolize more rapidly. Burkholder (1959)(Table 15)
shows the amazing interrelations of vitamin ex-
change between some marine bacteria. Similar
interrelations were found by the Lochhead school
between Bi2~producers and Bj^-users in soil.
Here the nutritional interdependency is dramati-
cally illustrated by vitamin producers themselves
being dependent on other vitamins. This interde-
pendency is a chemical symbiosis mediated and en-
hanced by the continuum which is the water envi-
ronment.
HETEROTROPHIC ABILITIES OF THE ALGAE
The subject has been recently reviewed by
Table 15. PRODUCTION OF B VITAMINS BY VITAMIN-REQUIRING
MARINE BACTERIA (BURKHOLDER 1959)
No. of
cultures
29
7
2
3
3
5
1
1
1
Pattern of requirements
Biotin Thiamine Niacin Bj2
+
+
+
+
-t- +
+ t
+ -e
+ t t
+ t +
No.
Biotin
6
1
3
1
of vitamin producers
Thiamine Niacin Bj2
3 29 1
7
1 3
3
5
1
-------
52
ALGAE AND METROPOLITAN WASTES
Krauss (1958) and Pringsheim (1959). Since algae
are at the origin of plants and animals, and the
evolution still proceeds, the energy requirements
vary greatly. The commonest carbon source for
the photosynthetic algae is probably CO2« Many
such algae are obligate autotrophs; they cannot be
grown in darkness on exogenous carbon sources
and do not utilize carbon sources even in light.
Among them are several Chlamydomonas; many of
the marine flagellates (Dunaliella, Rhodomonas,
Amphldinium, Gonyaulax, Gymnodinium, Peridin-
ium, Isochrysis, Syracosphaera); the freshwater
Synura caroliniana, S_. petersenii, Asterionella
formosa, Fragilaria capucina, Tabellaria floccu-
losa, and Woloszynskia limnetica (Provasoli and
Pintner, unpublished); Anabaena cylindrica (Fogg
1953); Anabaena variabilis, Anacystis nidulans and
Nostoc muscorum (Kratz and Myers, 1955). Con-
versely, all the colorless species (Polytoma, Poly-
tomella, Chilomonas, Astasia, etc.) are dependent
completely on heterotrophy, and, according to E.
and O. Pringsheim (1959) so are several colonial
Volvocales. For them a carbon source is needed
even in light: glucose is indispensable for Gonium
sacculiferum, and acetate for Chlamydobotrys,
Astrophoneme gubernaculifera, Gonium octonari-
um, G. quadra turn, Stephanosphaera pluvialis, and
Volvulina steinii. These species grow even better
in peptone and yeast extract, and reproduce in
darkness.
These two extremes, obligate phototrophy and
obligate heterotrophy, are bridged by species which
are bipotent; however they have different degrees
of phototrophic and heterotrophic abilities. Some
are predominantly photoautotrophic and the addition
of organics is only stimulatory: Eudorina elegans,
Gonium sociale, (E. &O. Pringsheim, 1959) Tra-
chelomonas pertyi, T. abrubta, Phacus pyrum,
Cryptomonas ovata, var. palustris (Provasoli and
Pintner, unpublished), and several species of
Chlorella.
Euglena gracilis and Ochromonas danica grow
exceedingly well phototrophically and also hetero-
trophically.
Several marine pennate diatoms are able to
grow in darkness on glucose, lactate, or acetate
(J. C. & R. A. Lewin, 1959). Pleodorina californi-
ca, P. illinoiensis and Volvox globator (Pringsheim
1959) grow poorly phototrophically; good growth is
obtained with dilute peptone, yeast extract, and
acetate.
Other species are predominantly heterotrophic
like Ochromonas malhamensis which grows poorly
phototrophically (Myers and Graham 1956). Species
with very high heterotrophic abilities like Euglena
gracilis and the two Ochromonas in opportune con-
ditions are able to utilize a variety of organic com-
pounds such as fatty acids, alcohols, sugars, amino
acids, etc. Other species are apparently more
exacting: Chlorella pyrenoidosa grows with high
efficiency in darkness but only on glucose, galac-
tose and acetate (Samejima & Myers, 1958); the
"azetatflagellaten" utilizes acetate and not glucose
and the "zuckerflagellaten" utilizes glucose and
not acetate (Pringsheim 1935, 1954, 1958). How-
ever, this apparently specialized utilization may
depend in part on preferential penetrabilities of the
other metabolites which were not met by the ex-
perimental or natural pH tried (Hutner and Prova-
soli, 1951, 1955). In some cases special triggers
may be needed: E. gracilis (an "azetatflagellaten")
can utilize glucose only in high CO2 or in the pre-
sence of small amounts of organic acids (Cramer
and Myers 1952) or amino acids (Hutner et al.
1956). In general the species that utilize very well
the exogenous carbon sources grow well in dark-
ness (Euglena, Ochromonas, some Gonium, Chlo-
rella, Chlororogonium, Astrophoneme, Volvulina,
Stephanosphaera). Less efficient heterotrophs do
not grow in darkness (Pleodorina), or grow poorly
. (two Oscillatoria, Lyngbya sp., Phormidium foveo-
larum. Plectonema notatum and Nostoc muscorum)
(M. G. Allen 1952).
CONSIDERATIONS ON HETEROTROPHY AS A
POSSIBLE ECOLOGICAL FACTOR
The effect of organics in natural waters varies
greatly with concentration. Therefore we will con-
sider separately three-types of environment differ-
ing essentially in organic content.
Richest are the polysaprobic (i.e. septic, sapro-
bic) waters. The flora of these waters consists
predominantly of heterotrophic and phagotrophic
algae. These environments can be colonized only
by algae which can both withstand and utilize high
concentrations of organic matter, products of
bacterial decomposition (including NH4), high CC>2
concentrations, reducing condition and, often,
gradients of H2S. Colorless flagellates (Distigma,
Astasia, Menoidium, Peranema, Monasj etc.),
euglenids, and chrysomonads are the usual algal
members of these communities. Oxidation ponds
are an artificial environment of this type, especial-
ly the lagoon receiving the organic influent. The
succeeding lagoons have intermediate conditions
leading, in the best cases, to a p-mesosaprobic
(i.e. subcontaminated) water.
The flora of the first two lagoons differs greatly
depending upon the type of waste being processed.
The richest in organics are those receiving meat
and milk products. The very high concentrations
of organics at high load favor active microbial
fermentations and anaerobiosis — conditions fa-
vorable only to a few tough obligate heterotrophic
algal species which do not contribute oxygen. It
is questionable whether the pigmented algae which
can live there (especially at low organic load) con-
tribute net oxygen. Depending on their heterotro-
phic powers, they may prefer heterotrophy to
-------
Nutrients in Bloom Production
53
photosynthesis. It will be interesting to determine
the CO2/O balance of the facultative and obligate
heterotrophic pigmented algae in the presence of
graded concentrations of highly utilizable carbon
sources and building blocks. An extreme case is
Ochromonas malhamensis (Hutner et aL 1953).
When the medium is extremely rich, the organisms
grow at first almost chalk-white in light (they pre-
fer to live heterotrophically and the chloroplast is
reduced to 1/10 - 1/20 or more of the normal
area); when the medium has become depleted the
organisms require a normal chloroplast but even
then, as shown by Myers and Graham (1956), they
photosynthesis. It will be interesting to determine
the CO2/O balance of the facultative and obligate
heterotrophic pigmented algae in the presence of
graded concentrations of highly utilizable carbon
sources and building blocks. An extreme case is
Ochromonas malhamensis (Hutner et al. 1953).
When the medium is extremely rich, the organisms
grow at first almost chalk-white in light (they pre-
fer to live heterotrophically and the chloroplast is
reduced to 1/10 - 1/20 or more of the normal
area); when the medium has become depleted the
organisms require a normal chloroplast but even
then, as shown by Myers and Graham (1956), they
are very poor photosynthesizers.
In oxidation ponds fed by domestic sewage, Chlo-
rella, Scenedesmus, Chlamydomonas and Anki-
strodesmus seem to predominate. Allen (1955)
found that these species utilize neither sewage nor
its bacterial oxidation products efficiently enough
to grow in darkness at the pH of the ponds. Other
evidence, both laboratory and field, supports her
conclusions that they are photoautotrophic under
these conditions. Oxidizable organic matter did
not decrease when this Chlorella was grown on
sterilized sewage —i.e. bacteria and other hetero-
trophs are responsible for the oxidation of organic
matter. Chlorella grows better in sewage when the
bacteria are present, but the favorable action of
bacteria depends on the amount of oxidizable ma-
terial present in the sewage. A similar increase
in growth of Chlorella can be elicited in low BOD
sewage when CO2 is supplied artificially.
Furthermore, the growth of Chlorella in the
presence of adequate amounts of CO£ is neither
stimulated nor inhibited by the presence of sewage
bacteria — i.e. the bacteria supply CO«j to Chlo-
rella. The situation is highly favorable for the
normal operation of the oxidation ponds. Chlorella
— and the shallowness of the ponds —provides the
oxygen necessary for the oxidative processes of
the heterotrophs which in turn create a high CC>2
content for Chlorella. Though the content in organ-
ics may not be high, presumably utilizable C sour-
ces and building blocks should be present in these
oxidation ponds favoring the establishment of bi-
potential organisms (i.e. photosynthesizers with
facultative heterotrophy). In the case analyzed by
Allen the lack of predominance of these organisms
may be due to inhibitions; Ankistrodesmus, Oocy-
stis, and Euglena which are also a part of this
flora grew at a low rate in sewage of relatively low
BOD and did not grow at all in high BOD sewage.
This emphasizes the need to consider in any en-
vironment not only the substances which may en-
hance growth but also inhibitory substances and
metabolite concentrations which may be unfavor-
able for more sensitive organisms. It is well
known that high concentrations of mineral ferti-
lizers (as normally found in fish ponds) favor the
growth of green algae. Pollution by duck excreta
in Great South Bay (Long Island) favored growth of
Nannochloris and Stichococcus and eliminated the
normal flora of diatoms. Ryther (1954) found that
Nannochloris and Stichococcus utilize NH4, urea,
and uric acid, and that a Nitzschia isolated from
the same bay grew very poorly in NH4 and organic
N. The change of flora, in my opinion, is not due
to inability to utilize NH4, because all algae utilize
NH4 at a pH below neutrality, but probably to in-
hibition. Many marine algae are inhibited or
killed, depending upon the concentration, by am-
monia at alkaline pH. Production of antimetabo-
lites by the green algae is also probable.
As the quantity of organics becomes less — be-
tween 10-100 ppm. — the action of organics be-
comes subtler indeed to analyse, yet its effect is
ecologically very important These are the condi-
tions prevailing in freshwater and bays affected by
civilization. The creation of blooms and often the
change in flora cannot be considered as depending
principally on the heterotrophic effect. There is
undoubtedly an advantage for the algae (facultative
heterotrophs) which can utilize organic compounds
as building blocks or carbon sources. A small body
of evidence from tracer data indicates that minor
syntheses maybe bypassed when the synthetic pro-
ducts are present in the environment. Thus adenine
is taken up by Ochromonas malhamensis (Hamilton,
1953), and uptake of aspartic acid suppresses en-
dogenous synthesis of lysine in E. gracilis (Vogel,
1959). While such by-passings of synthesis maybe
individually minor, it is conceivable that collective-
ly they permit a noticeable, even substantial, in-
crease in growth. We have been concerned in he-
terotrophy for the most part with great stimulations
and all-or-nothing effects; these smaller biosyn-
thetic stimulations are just beginning to be ex-
plored.
A similar effect has been noted by many when a
few milligrams percent of various peptones, soil
extract, other infusions of natural products, and
even purines and pyrimidines, are added to bac-
teria-free cultures of algae grown in synthetic
media enriched with trace metals and vitamins.
However, the stimulation of these additions is of-
ten eliminated when the mineral part and the trace
metals in the medium are better adjusted to fit the
particular organism. This implies a metal buffer-
-------
54
ALGAE AND METROPOLITAN WASTES
ing action, or supplying of metals by the organics
(peptones and infusions are rich in metals), rather
than a micro-heterotrophic advantage. Other ef-
fects may also be due to pH buffering, rH poising,
organic sulphur compounds — all known to bene-
fit the growth of some algae. In short, it is diffi-
cult to dissect the remarkable effects of minor
pollution and eutrophication.
Fogg (1959) is inclined to think the heterotro-
phic effect of the dissolved organics is minor and
he considers that the yellow acids and algal pro-
ducts which are a part of the organics may play an
important role. Shapiro (1957) found that the or-
ganic acids of a yellow pigmented fraction present
in freshwaters are stable to biological decompo-
sition for a few months and that they chelate metal
ions effectively.
The nitrogenous extracellular products given off
by blue-green algae are capable of binding copper
and other trace metals very strongly. Death of
Anabaena cylindrica occurs at 2 mg/liter of copper
in the absence of these substances while in their
presence copper kills at 16 mg/liter (Fogg and
Westlake 1955). "Humic acids" brought in by soil
runoff act similarly. All these substances bind
and solubilize trace metals making available to the
algae a non-toxic continuous supply. The vitamins
may play an important role because whether limit-
ing or in ample supply they will tend to modify the
flora. When limiting, the absence of auxotrophs
will leave the non-auxotrophic algae free from
competition for the same nutrients; when plentiful
the auxotrophs have a chance to predominate. The
effluent of sewage disposal plants is likely to be
rich in vitamins; sludge contains high quantities of
612 (Hoover et aL 1951; Sathyanarayama et aL
1959). Nothing is known of the vitamin content of
the effluent of sewage oxidation ponds, but it would
be surprising if they lacked vitamins; many bac-
teria and algae are high vitamin producers.
Organic enrichments whether derived from al-
gae growing in lakes or. directly from civilization
have, then,a complex action:
(a) they may be a direct source of nutrients;
(b) they may solubilize P and trace metals;
(c) they are vitamin sources;
(d) as supporters of microbial growth they are
likely to influence algal growth through products of
microbial degradation including NIL}, NO3> and
through (b), and (c); and
(e) they may create favorable pH and rH conditions.
In seawater and freshwater unknown factors other
than the usual nutrients enhance algal growth. Per-
haps some of them are similar to plant harmones.
The algae in oligotrophic waters are either pho-
toautotrophs or photoauxotrophs. Blooming condi-
tions will then mainly depend upon inorganic sub-
stances, metal solubilizers, and vitamins. Soil
run-off and rivers are the most likely suppliers.
REFERENCES
Adair, E. J. and H. S. Vishniac 1958. Marine fun-
gus requiring vitamin Bj2. Science 127: 147-48.
Allen, M. B. 1952. The cultivation of Myxophyceae.
Arch. f. Mikrobiol. 17: 34-53.
Allen, M. B. 1955. General features of algai 'growth
in sewage oxidation ponds. State Water Pollution
Control Board. Publ. No. 13, Sacramento, Calif.
Benoit, R. J. 1957. Preliminary observations on
cobalt and vitamin 8^2 in freshwater. Limnol. &
Oceanogr. 2: 233-40.
Burkholder, P. R. 1959. Vitamin-producing bac-
teria in the sea. Inter. Oceanog. Congress. Pre-
prints, A.A.A.S., Washington p. 912-13.
Burkholder, P. R. and L. M. Burkholder 1956. Vi-
tamin Bj2 in suspended solids and marsh muds
collected along the coast of Georgia. LimnoL &
Oceanogr. 1: 202-08.
1958. Studies on B vitamins in re-
lation to productivity of the Bahia Fosforescente,
Puerto Rico. Bull. Mar. Sci. of the Gulf & Carri-
beanS: 201-23.
Burton, M. O., and A. G. Lochhead 1951. Studies
on the production of vitamin Bj2~active substances
by microorganisms. Canad. J. Bot. 29: 352-59.
Cowey, C. B. 1956. A preliminary investigation
of the variation of vitamins Bj2 in oceanic and
coastal waters. J. mar. biol. Ass. U.K. 35: 609-20.
Cramer, M. and J. Myers 1952. Growth and photo-
synthetic characteristics of Euglena gracilis.
Arch. f. MikrobioL 17: 384-402.
Daisley, K. W. 1957. Comment on article by M.
R. Droop. Nature 180: 1042-43.
Daisley, K. W. 1958. A method for the measure-
ment of vitamin Bj2 concentration in sea water.
J. mar. biol. Ass. U.K. 37: 673-81.
Droop, M. R. 1955. A suggested method for the
assay of vitamin Bj^ in sea water. J. mar. biol.
Ass. U.K. 34_: 435-440.
Droop, M. R. 1957. Vitamin Bl2 in marine eco-
logy. - Nature 180: 1041-42.
-------
Nutrients in Bloom Production
55
REFERENCES (Cont'd.)
Droop, M. R. 1959. Water-soluble factors in the
nutrition of Oxyrrhis marina. J. mar. biol. Ass.
U. K. 38: 605-620.
Droop, M. R., McLaughlin, J. J. A., Pinter, I. J.,
and L. Provasoli 1959. Specificity of some proto-
phytes toward vitamin Bl2~like compounds. Inter.
Oceanogr. Congress, Preprints, A.A.A.S. Washing-
ton, p. 916-18.
Ericson, L. E. and L. Lewis. 1953. On the occur-
rence of vitamin Bi2~factors in marine algae. Ark.
f. Kemi. 6: 427-42.
Fogg, G. E. 1953. The metabolism of algae.
Methuen & Co. Ltd. London 149 pp.
Fogg, G. E. 1959. Dissolved organic matter in
ocean and lakes. In New Biology, vol. 29, Penguin
Books, p. 31-48.
Fogg, G. E. and D. F. Westlake 1955. The impor-
tance of extracellular products of algae in fresh-
water. Verh. int. Ver. Limnol. 12_: 219-32.
Fries, L, 1959.' Goniotrichum elegans: a marine
red alga requiring vitamin Bi2. Nature 183: 558-
59.
Hamilton, L. 1953. Utilization of purines for
nucleic acid synthesis in chrysomonads and other
organisms. Ann. N. Y. Acad. Sci. 56: 961-8.
Holm-Hansen, O., Gerloff, G. C., and F. Skoog
1954. Cobalt as an essential element for blue-
green algae. Physiol. Plantarum _7: 665-75.
Hoover, S.R., Yaseweiez, L., Pepinsky, J.B. and
N. Porges. 1951. Vitamin Bi2 in activated sewage
sludge. Science 114: 213.
Hutchinson, E. G. 1943. Thiamine in lake waters
and aquatic organisms. Arch. Biochem. 25: 143-50.
Hutchinson, E. G. and J. K. Setlow. 1946. Limno-
logical studies in Connecticut. Vm. The niacin
cycle in a small inland lake. Ecol. 27: 13-22.
Hutner, S. H., Bach, M. K. and G. L. M. Ross.
1956. A sugar-containing basal medium for vitamin
Bl2-assay with Euglena; application to body fluids.
J. Protozool. 3: 101-12.
Hutner, S. H., Cury, A. and H. Baker 1958. Mic-
robiological assays. Anal. Chem. 30_: 849-67.
Hutner, S. H. and L. Provasoli 1951. The Phyto-
flagellates. In "Biochemistry and Physiology of
Protozoa" A. Lwoff edit. Vol. I Academic Press
N. Y.
1955. Comparative biochemistry of
flagellates. In: "Biochemistry and Physiology of
Protozoa", Hutner and Lwoff edit. Vol. n Academ-
ic Press N. Y.
Hutner, S. H., Provasoli, L. and J. Filfus 1953.
Nutrition of some phagotrophic freshwater chry-
somonads. Ann. N.Y. Acad. 56: 852-62.
Kashiwada, K., Kakimoto, D., Morita, T., Kana-
zaqa, A. and K. Kawagoe 1957. Studies on vita-
min B12 in sea water, n. On the assay method
and the distribution of this vitamin Bl2 to the ocean.
Bull. Jap. Soc. Sci. Fisheries. 22: 637-40.
Kashiwada, K., Kakimoto, D. and K. Kawagoe 1957b.
Studies on vitamin Bi2 in sea water, in On the
diurnal fluctuation of vitamin Bi2 in the sea and
vertical distribution in the lake. Bull. Jap. Soc.
Sci. Fisheries. 23: 420-53.
Kratz, W.A. and J. Myers. 1955. Photosynthesis
and respiration of the blue-green algae. Plant
Physiol. 30: 275-80.
Krauss, R. W. 1958. Physiology of the fresh water
algae. Ann. Rev. Plant Physiol. 9: 207-44.
Lewin, R. A. 1954. A marine Stichococcys sp.
which requires vitamin Bi2- J. Gen. MicrobioT.
10: 93-96.
Lewin, J. C. and R. A. Lewin 1959. Auxotrophy
and heterotrophy in marine littoral diatoms. Inter.
Oceanograph. Congress, Preprints. A. A. A. S.,
Washington p. 928-29.
Lochhead, A. G. and R. H. Thexton 1951. Vitamin
612 as a growth factor for soil bacteria. Nature
167: 1034.
Lwoff, A. 1943. L'evolution physiologique, Her-
manm et Cie. Paris.
Lucas, C. E. 1955. External methabolities in the
sea. Papers in marine Biology and Oceanography
Suppl. to Vo. 3 of Deep Sea Research p. 139-148.
McLaughlin, J. J. A. and P. A. Zahl 1959. Vitamin
requirements in symbiotic algae. Inter. Oceano-
graph. Congress, Preprints. A.A.A.S., Washing-
ton p. 903-31.
Myers, J. and J.Graham 1956. The role of photo-
synthesis in the physiology of Ochromonas. J.Cell.
comp. Physiol. 47: 397-413.
Pringsheim, E.G. 1935. Ueber Azetatflagellaten.
Naturwiss. 23: 110-114.
1954. Ueber Zuckerflagellaten. Natur-
wiss. 41: 1-3.
1958. Ueber mixotrophie bei flagella-
tion Planta 52: 405-30.
1959. Heterotrophie bei Algen und
Flagellaten. in Handb Pflanzenphysiol. -Springer -
Verlag. j): 303-26.
Pringsheim, E. G. and O. Pringsheim 1959. Die
Ernahrung Koloniebildender Volvocales. Biol.
Zentralbl. _78: 937-71.
Provasoli, L. 1958. Nutrition and ecology of Pro-
tozoaand Algae. Ann. Rev. Microbiol. 12: 279-308.
-------
56
ALGAE AND METROPOLITAN WASTES
REFERENCES (Cont'd.)
Provasoli, L. and I. J. Pintner 1953, Assay of vi-
tamin Bi2 in sea water. Proc. Soc. Protozool.
Provasoli, L. and I. J. Pintner 1953b. Ecological
Implications of in vitro Nutritional Requirements of
Algal Flagellates. Ann.N. Y.Acad.Sci. 56: 839-51.
Robbins, W. J., Hervey, A. and M. E. Stebbins
1950. Studies on Euglena and vitamin Bi2- Bull.
Torrey Bot. Club, 77: 423-41.
Robbins, W. J., Hervey, A. and M. E. Stebbins
1951. Further observations on Euglena and B}2-
BulL Torrey Bot. Club. 78: 363-375.
Ryther, J. 1954. The ecology of phytoplankton
blooms in Moriches Bay and Great South Bay,
L. L, N. Y. Biol. BulL 106: 198-209.
Samejima, H. and J. Myers 1958. On the hetero-
trophic growth of Chlorella pyrenoidosa. J. gen.
microbioL 18: 107-17.
Shapiro, J. 1957. Chemical and biological studies
on the yellow organic acids of lake water. Limnol.
and Oceanogr. 2: 161-79.
Sathyanarayana, R., Jastry, C. A., Bhagavan, H.
N. and B. R. Balija 1959. Vitamin Bj^ in sewage
sludges. Science 129: 276.
Starr, T. J. 1956. Relative amounts of Vitamin
Bj2 in detritus from oceanic and estuarine envi-
ronments near Sapelo Island, Georgia. Ecol. 37:
658-64.
Starr, T. J., Jones, M. E., and D. Martinez. 1957.
The production of vitamin B^active substances
by marine bacteria. Limnol. and Oceanogr. 2:
114-19.
Vishniac, H. S., and G. Riley, 1959. Bi2 and thia-
mine in Long Island Sound: patterns of distribution
and ecological significance. Inter. Oceanogr. Con.
gress, Preprints, A.A.A.S. Washington, p. 942-43.
Vogel, H. J. 1959. Lysine biosynthesis in Chlo-
rella and Euglena: phylogenetic significance. Bio-
chim. et Biophys. Acta 34: 282-3.
ALGAL DENSITY AS RELATED TO NUTRITIONAL THRESHOLDS*
JAMES B. LACKEY
Dept. of Civil Engineering, University of Florida, Gainesville
Definition of oligotrophic and eutrophic as poor-
ly productive or highly productive presumes know-
ledge of how many organisms or how great a bio-
mass a given body of water contains or, literally,
produces in a given interval of time. Algal den-
sity is a term demanding clarification in regard to
the location of the algae and whether we are talking
of the numbers of algal cells, or their bio mass.
Algal density in the top three inches of a lake may
bear little relationship to the original concept if
the lake is thirty feet deep. In a moving stream
the density may be uniform thruout the depth. Pre-
sumably the topic should include total numbers or
biomass (of algae) in a body of water under con-
sideration, as well as whether there is a relation-
ship between nutrition and local aggregation of
algae. Furthermore, the consideration should in-
clude aggregates, top three inches, or patches, for
example, and continuity of the density in time.
Algae may be benthic or planktonic. For pur-
poses of this seminar benthic algae may be unim-
portant Unless they are great masses such as
Spirogyra sometimes exhibits, and are anchored
to the bottom, they tend to be confined to a thin
layer, up to 5mm thick. These bottom dwelling
forms neither clog filters nor produce tastes and
odors as a general thing. But we can learn much
from them. In 1937, slides were hung in the Ohio
River below the Stream Pollution Investigations
Station, at Third and Kilgour Streets, in Cincinnati
Ohio; within a few days they were colonized by
various organisms. Most of these were not the
organisms present as plankton in the river. This
work has been repeated in Santa Fe Lake in Flori-
da since October 1959. The same results have
been obtained, but the organisms are different,
Santa Fe being an acid lake.
Slides left in the lake four and five months have
developed thick and dense coatings of algae and
other organisms, but almost the same species oc-
cur now as were present a few days after setting
out. Given a constant exposure to such nutrients
as are brought to them by wind-circulated water,
they do not die out, but increase, albeit slowly.
Water temperatures have fluctuated between 11°
and 16° C. The nutrients have not been identified
or measured. But the significant thing is that spe-
cies of Frustulia for example have not died out.
Dominance has changed, but not the species list.
The nutritional threshold has been sufficient to
maintain a continuous but slow growth.
*This work is incident to and partly done under a grant-in-aid from the Environmental Sciences Branch,
Division of Biology and Medicine,Atomic Energy Commission but was not financially supported thereby.
-------
Nutritional Thresholds
57
We do not have to postulate a very sharp in-
crease in the division rate of an algal cell to show
a great increase in numbers within a short time.
Birge and Juday (1922) decided that algae in the
Madison (Wis.) lakes divided once or twice in 24
hours. On Aug. 31, 1956, there was a population
of one Chlorella per ml. in a settling basin at Oak
Ridge, Tennessee; three days later this pond was
green with a population of about 20,000 per ml.
This represented 15 divisions in three days rather
than three —a five-fold increase. But the biomass
had increased many thousandfold. Particular at-
tention was paid to the size of the cells. They were
about the same average size throughout the three
days. Since this bloom maintained itself for about
two weeks, despite a heavy loss downstream every
24 hours, the necessary nutrients were evidently
present during these 2 weeks in sufficient abun-
dance to support this growth. Actually the amount
could have been quite small, even for a 20,000, OOO/
liter population.
The nutrition of Chlorella pyrenoidosa has been
carefully worked out (Burlew, 1953) and it is com-
mon knowledge that the organism requires so little
mineral salts as to be a pest in laboratories by in-
vading distilled water dispensers. Birge and Juday
(1922) give the composition of the total plankton
crops in the Madison lakes, a type of study which
has been repeated many times for individual spe-
cies of algae. Generally the dry weight is about
one-tenth the wet weight, and the ash from 3.5
to 10.0 percent of the dry weight, except in diatoms
where the silica makes the ash content much high-
er. In some prepilot-plant experiments on cultur-
ing Chlorella, Mituya et al (Burlew, 1953) set up
about 1200 liters of culture medium containing
roughly 8000 grams of nutrients, 3000 grams of
this being urea. After ten days operation, algae
had taken up .0065 per cent of the available nutrients
as ash content. These are rough computations
from their figures, but are mass figures rather
than small scale laboratory cultures. The results
are a confirmation of the idea that contributed nu-
trients in very small amounts account for enor-
mous growths.
It becomes of interest, then, to determine the
quality of nutrients necessary for maintenance of
an algal species, and the quantity necessary to pro-
duce a sharp increase in the population of that spe-
cies. The first statement is that the matter is sub-
ject to very little generalization — each species
has its own optimum.
We are quite familiar with this. Thus we say
that it takes one amount of copper as CuSO4 to kill
species A and a different amount for species B.
All attempts to grow Spirogyra in our laboratory
on spring water and distilled water were unsuc-
cessful until glass distilled Cu-free water was
used, after which the cultures grew well. The
copper in the spring water is 0.01 ppm. In a long
series of tests on chlorine-and iodine-resistant
algae such as Anabaena, Navicula, Gomphosphae-
ria, and some others of interest to swimming pool
owners, it was found that after an initial inocula-
tion with perhaps 50 species of microorganisms
most were killed by extremely small amounts of
these two chemicals, but it was very difficult to
kill the above three algae. These, and a consider-
able number of other species grew well in the con-
trols. The questions at once arose as to whether
this was a direct toxicity, or whether the ratios of
nutrients were not so adjusted that other algae
could live, or even if other species died because of
antibiotic production. Since several nutrient sub-
strates were used in the experiments, it seems
that direct toxicity was the killing agent. But se-
lecting a single factor is not easy.
DENSITY AS A MULTI-FACTOR RESULT
Yount (1955) concluded that "there is, in any habi-
tat, no one factor which determines the species
density of an area, but always a combination of
factors." One may question how such amazing
blooms as are often encountered come into being
if they are dependent on a combination of factors.
Certainly proper nutrients, and adequate nutrients
are two of the several factors. But figures can be
presented to show that these are not always effec-
tive. Thus phosphorus, supposedly contributed by
the rich phosphate mining areas of central Florida
has been repeatedly suggested as a cause of the
blooms of Gymnodinium breve in the Gulf of Mexi-
co. Lackey and Hynes (1955) studied the nitrogen-
phosphorus ratios in sea water where G. breve was
present in varying numbers and could find no cor-
relation either as to total NO3 or PO^ present, or
as to N/P ratios. Samples from the vicinity of
Englewood, Florida, on June 18, 1956, were cen-
trifuged. The results of some of the 18 analyses
are shown in the following tabulation:
Organisms
per liter
4,814,000
9,500
212,000
3,235,000
211,000
NOg , ppm
0.507
0.149
0.188
0.541
0.160
PO4 ppm
0.
0.021
0.011
0.
0.
The decanted water contained virtually no par-
ticulate matter, and since there had been a marked
kill of fish and other animals, with considerable
decomposition, the failure to find more PO4 was
unexpected. It is unfortunate that total P was not de-
ter mined. However, these figures indicate that
certainly not all of it was tied up in the dinof lagel-
late bodies. The nitrogen figures are likewise in-
explicable. The essential point here is that it is
not possible to say too much or too little of either,
or an improper ratio, is limiting. We have here a
verification of Yount's point, mentioned above.
-------
58
ALGAE AND METROTROPOIJTAN WASTES
NEED FOR CHEMICALLY DEFINED MEDIA
Even with bacteria free cultures, the medium
must be chemically defined if we are to arrive at
the nutritive threshold for unlimited production.
Ketchum and Redfield (1938) used sea water, en-
riched by the addition of 20 atoms of nitrogen to
1 of phosphorus, to secure quantities of Nitzschla
closterium. The cultures were bacteria free, and
by following the cell count they could determine
when to add more nutrients. They found that 2 x
10® cells/liter completely exhausted the nutrients.
At the end of 15 days the division rate had dropped
to 0.07 per day. The only unknowns here are the
substances in sea water which might have had an
adverse effect on the division rate, or possibly
metabolites produced by the diatoms themselves.
Manifestly the division rates they obtained are far
lower than those which occur in some cultures.
We cannot say the same thing for populations in na-
ture, unless we can rule out the possibility that
dense blooms are aggregates, drawn together from
a large volume of water, by some unknown mechan-
ism. However, it has been shown that Entosiphon
maintains a division rate of between 1 and 3 per
day (Lackey, 1929; Bennett, In Press) for many
generations as long as daily isolations are made,
whereas in mass cultures the division rate soon
shows and after ten days dead organisms as well
as stunted ones are found. Where flow-through
culture conditions are maintained, or under such
conditions as provided by Ketchum and Redfield
(1938) it is at least possible to determine the nu-
tritive levels necessary to maintain or step up the
division rate. In their cultures it was evident that
nitrate and phosphate were rapidly used up, and
had to be replaced.
This is a rather general assumption for algae.
In a stream where there is less chance of deple-
tion than in a lake, absorption is probably at a
steady rate by users. Rice (1953) has shown this .
for PO4 in the case of bacteria free strains of
Nitzschia. Sawyer and Lackey (Sawyer and Lackey,
1945; Lackey, 1945) after a study of 17 lakes in
southeastern Wisconsin concluded that when NOs
and PC-4 in a lake reached values of 0.3 and 0.015
ppm, respectively, a bloom appeared, sometimes
one species, sometimes another, sometimes sev-
eral. In Lake Santa Fe, Florida, the lowest popu-
lation recorded in over a year was following a
heavy rain which considerably diluted the lake,
but did not bring in appreciable nutrients since
there is exceedingly little run-off into the lake.
The rain occurred March 31-April 1, and on April
6 the NOs in the lake was 0.16 ppm and the PO4
0.005 ppm. Almost the entire population, aside
from bacteria, consisted of Anabaena filaments,
182 per mL Considering Anabaena as potentially
able to fix its own nitrogen and therefore probably
limited principally by available PO4, its lone posi-
tion is understandable.
INABILITY TO DO MORE THAN GENERALIZE
Among the most frequent of the bloom forming
organisms are Euglena species and their green and
colorless relatives. It would seem that the nutrient
thresholds for such organisms would be well
known. Unfortunately, most of our knowledge per-
tains to their environments, rather than the nutri-
ent substrates necessary for dense blooms. Thus
we know that Euglena mutabilis favors H2SO4 aci-
dity, that Entosiphon and Peranema form dense
cultures in a medium containing autoclaved wheat,
and that many genera and species of the group oc-
cur abundantly in barnyard pools. But almost all
of these are either not grown in the laboratory or,
if so, are grown along with other organisms so that
defined substrates are not known. Astasia klebsii
(von Dach, 1940) is easily grown in pure culture on
an acetate medium; Euglena gracilis has been
grown by many workers in bacteria-free media
and its status in this respect has been reviewed by
Pringsheim (1956) in a paper which indicates the
need for pure culture work with this group. Cul-
tures are easily developed and the Indiana Culture
Collection (Starr, 1960) lists 55 species or strains
of Euglenophyceae which they maintain either in
unialgal, or bacteria-free condition. Most of these,
however, are grown in media containing beef ex-
tract, yeast extract, and tryptone, viz., not a
chemically defined medium. Many of their other
algae are maintained in media not possible to
chemically define.
When we note the conditions of blooms in the
field, it is at once apparent that there is much
variation of the conditions under which they appear,
and that at best our knowledge of what nutrient
substances and amounts are related to algal densi-
ty is merely a guess. Johnson, working on raising
shrimp in controlled pools at Marineland, Florida,
added commercial fertilizer. The pools developed
heavy concentrations of Coccolithophora, not al-
ways the same species, and occasionally another
genus such as Platymonas bloomed. Nevertheless,
there seems little doubt, from records of too many
workers to cite, that amounts of NOsand PO4 can
readily be limiting for many organisms, probably
including most blooms of Euglenophyceae. How-
ever, for these last it is highly probable that un-
defined extractives and dissolved organic substan-
ces (often from the excreta and dung of animals)
are also necessary. The use of soil extract also
indicates such is the case.
We recognize the role of NOsand PO4in a prac-
tical way. Pringsheim (1949) varies the ratio of
NOs to PO4 between 4 and 8 to 1, in his Chu 10
formulae. He gives other formulae as Knop's so-
lution, 11 parts NOs to 2 Parts PO4; Pringsheim's
own medium is 10 parts NOs to * PO4- The *or~
mulae of Ketchum and Redfield (1938) are by
weight, 9 to 1 and 6.8 to 1; by atoms, 20 and 15 to 1.
-------
Nutritional Thresholds
59
BLOOMS DURING AND FOLLOWING WASTE
TREATMENT
Anyone familiar with the accepted methods of
biological treatment of sewage and trade wastes
knows that eventually certain blooms are inevita-
ble; in trickling filters, only on the top of the
medium; in lagoons and oxidation ponds, on the
surface; and after treatment, almost invariably in
the receiving water. These blooms may be blue
greens or filamentous greens on trickling filter:
small Chloroeoccales such as Chlorella, Schene-
desmus, and Golenkinia, or Volvocales such as
Chlorogonium or Chlamydomonas on oxidation
ponds. Sometimes Euglena is a heavy contributor
to oxidation pond blooms. Trade waste lagoons
may vary a great deal in bloom composition, de-
pending apparently on specific substances in the
waste. Thus the lagoons at!tPlymouth, Florida,
which handled citrus wastes bloomed heavily with
Euglena gracilis, 12. pisciformis and Chlorogonium
euchlora. Receiving water blooms are frequently
mixed, but with numerous Euglenophyceae. Some-
times such waters fail to bloom, as is often the
case with the polishing ponds at the University of
Florida campus treatment plant.
Assuming that the above methods of treatment
have produced a satisfactory reduction in BOD and
in bacterial numbers, what can we say regarding
the algal densities produced, either coincident, or
subsequently? What do they — at least the subse-
quent densities — indicate regarding eutrophication
of receiving waters ?
It is well known that lakes in the vicinity of
cities, if they receive urban drainage, may become
eutrophic. There seems little doubt that this is due
to an increased addition of phosphate and nitrate,
and, what we are prone to forget, other necessary
growth promoting substances. This is not a case
of secondary BOD contributions which, if they oc-
cur at all, lag far behind the algal blooms. There
has been considerable discussion of the possibility
of removing PO^ from the effluents of conventional
treatment plants. It is suggested here, that this
may change only the composition of the bloom.
There are algae whose PO4 requirements are ex-
tremely low, below the amounts we might remove.
In short, it would seem that conventional treatment
has solved the major waste treatment problems.
While algal blooms are often a nuisance, and
sometimes highly objectionable, they are not nec-
essarily a menace to health. Perhaps we had bet-
ter concentrate investigation on means of lowering
algal densities in receiving waters, by biologic
means, such as predator-prey relationships or by
harvesting the algae.
SUMMARY
In summary, we can say that for some species
of algae, density is directly related to the amounts
of nitrate and phosphate present, and that minima
of these for producing blooms are 0.3 and 0.015
ppm. However, it should also be determined
whether the algal bloom is merely a local aggregate
due to some probably unknown factors, or whether
it is present in a great volume of water or is wide-
spread.
We are not ready to set absolute values on nu-
trients, because we still know too little about the
requirements of individual species. This is espe-
cially true of those species which are not obligate
autotrophs, as seems to be the case for many Eu-
glenids and Coccolithophora or Chlorophyceae. We
also know too little about the production of anti-
biotic metabolites by other species in the environ-
ment. But we have reached the stage where it is
possible to generalize, just as the agriculturist
can generalize about the particular fertilizer
needs, if we realize that sometimes he, too, is not
successful despite following a pattern.
REFERENCES
Bennett, Carrie F. In Press.^ Sublethal effects of
Co60 Gamma radiation on Entosiphon sulcatum.
Paper delivered before the 21st annual meeting
of the Assn. of Southern Biologists, New Orleans,
April 22, 1960.
Birge, Edward A. and Chancey Juday. 1922. The
inland lakes of Wisconsin. The plankton. I. Its
quantity and chemical composition. Bull. 64,
Scientific Series 13. Wisconsin Geological and
Natural History Survey.
Burlew, John S., Editor. 1953. Algal culture from
laboratory to pilot plant. Publication 600, Carnegie
lost, of Washington.
Ketchum, Bostwick H. and Alfred C. Redfield,
1938. A method for maintaining a continuous sup-
ply of marine diatoms by culture. Biol. Bull. 75,
1: 165-169.
Lackey, James B. and Jacqueline A. Hynes. 1955.
The Florida Gulf Coast Red Tide. Bull. Series No.
70, Florida Eng. and Ind. Exp. Station. 1-24.
Lackey, James B. 1945. Plankton productivity of
certain southeastern Wisconsin lakes as related to
fertilization. H. Productivity. Sew. Wks. J.,
17(4): 795-802.
-------
60
ALGAE AND METROPOLITAN WASTES
Lackey, James B. 1929. Studies in the life his-
tories of Euglenida. n. The life cycles of Entosi-
phon sulcatum and Peranema trichophorum.
Archiv. fur. Protistenkimde. 67(1): 127-156.
Pringsheim, Ernst Georg. 1956. Contributions
toward a monograph of the genus Euglena. Nova
Acta Leopoldina. 18(125): 1-168.
Pringsheim, E. G. 1949. Pure cultures of algae.
Cambridge University Press. Xn - 119.
Rice, Theodore R. 1953. Phosphorus exchange in
marine phytoplankton. Fishery Bull. 80, U.S. Fish
and Wildlife Service, 54: 1-89.
Sawyer, C. N. and J. B. Lackey. 1945. Investiga-
tion of the odor nuisance occurring in the Madison
Lakes, particularly Monona, Waubesa, Kegonsa,
from July 1943 to July 1944. Report to the Gover-
nor's Committee, Madison,Wisconsin. 1-92.
Starr, Richard C. 1960. The culture collection of
algae at Indiana University. Am. J. Bot. 47(1):
67-86.
von Dach, H. 1940. Factors which affect the
growth of a colorless flagellate, Astasia klebsii,
in pure culture. Ohio J. Sci. 40: 37-48.
Yount, James L. 1955. Factors that control spe-
cies numbers in Silver Springs. Productivity of
Florida Springs. NR 163-106 (NONR 580°2).
Third Annual Report to Biology Branch, Office
of Naval Research. Jan. 1 to Dec. 31, 1955. Un-
paged.
ZoBell, Claude E. 1946. Marine Microbiology.
Chronica Botanica Co., Waltham, Mass. XV - 239.
-------
Productivity and How to Measure It
METHODS OF MEASUREMENT OF PRIMARY PRODUCTION IN NATURAL WATERS*
LAWRENCE R. POMEROY
University of Georgia Marine Institute, Sapelo Island, Georgia
ABSTRACT
To estimate primary production of natural plant
populations it is necessary to measure the rate of
photosynthesis of the populations, or portions of
them, under natural environmental conditions.
Measurements may be carried out in the field or
under simulated natural conditions in the labora-
tory. Some of the methods by which this has been
done are the following:
1. Small samples of the natural plant communi-
ty are confined in transparent vessels, and the rate
of change of some product or raw material of photo-
synthesis is measured. Changes in dissolved oxy-
gen, pH, or uptake of C -labeled carbon may be
followed.
2. Changes in the concentration of a product or
raw material of photosynthesis are measured in
unconfined natural waters.
3. By measuring the chlorophyll content of a
system and the radiant energy reaching the plant
populations the rate of photosynthesis may be de-
duced empirically.
4. Plant material may be harvested periodi-
cally to deduce the photosynthetic rate from chan-
ges in plant biomass.
The accuracy of most of these methods is limi-
ted by the methods of sampling the plant popula-
tions or parameters of photosynthesis. Improved
sampling accuracy can be achieved by the use of
proper statistical methods and by making many
more observations. To take a significant number
of observations, techniques of automation can be
used. One possibility is to continuously record the
oxygen tension, either in vessels or in nature, with
polarographic electrodes. Another is to use auto-
matic sample-collecting devices.
In principle the measurement of photosynthesis
is the same in the laboratory and in the field. In
practice complexities and uncertainties creep in
when we attempt to estimate the photosynthetic
rate of natural populations under natural condi-
tions. While laboratory studies of photosynthesis
are likely to be concerned with defined conditions,
field studies seek to find the rate of photosynthesis
under existing conditions in nature. These condi-
tions may be difficult to define and are variable
with time.
Several recent reviews (Lund and Tailing,
1957; Steele, 1959; Strickland, 1960) and sym-
posia (Cons. Int. Explor. Mer, Rapp. et Proc.
Verb., Vol. 144; Second all-Union Conference on
Photosynthesis: translation by Office of Technical
Services, 123 pp, 1957) have dealt with problems
of measuring primary production.
I shall briefly review the more promising meth-
ods in the light of these critiques, and point out
what seem to me to be the important shortcomings
of our present methods and the most likely means
of making improvements. Four general approaches
have been made in estimating primary production,
and I shall review each briefly.
1. Small samples of the natural community are
confined in transparent vessels where the rate of
change of concentration of some product or raw
material of photosynthesis can be measured. Ex-
amples of this approach are the light-and-dark-
bottle method (Gaarder and Gran, 1927) and the
C14 method (Steeman-Nielsen, 1952). Similar
methods have been used with benthic populations as
well as phytoplankton(Odum, 1957; Pomeroy, 1959;
1960a).
This general approach has some important limi-
tations.
a) The samples are small and may not be re-
presentative. This difficulty may be overcome in
part by the use of integrating sampling methods
(Griffith, 1957) or by abundant replication of ob-
servations and statistical treatment.
b) Natural eddy diffusion is lacking in small
*Contribution No. 19 from the University of Georgia Marine Institute, Sapelo Island, Georgia.
61
-------
62
ALGAE AND METROPOLITAN WASTES
vessels. This may result in poor exchange of ma-
terials. It certainly prevents vertical movement
of the cells. Experiments on the effect of stirring
on photosynthesis in closed vessels are conflicting
(Tailing, 1960), and further experimental work is
needed to clarify this point.
c) Small closed vessels may become depleted
of essential materials. Especially in the study of
blooms or macrophytes (cf. Emerson and Green,
1934) the investigator is pressed by possible la-
tent periods on one side and by depletion of CC«2
or nutrients on the other. The exposure period
must be chosen with care.
Closed-vessel methods usually involve the ex-
posure of the vessels in nature at the depths from
which the samples were collected. This is labori-
ous and expensive. It has become increasingly
popular to place the bottles instead in some sort
of chamber, in the laboratory or aboard ship, that
duplicates the natural physical environment. The
chambers vary in sophistication from washtubs
to illuminated constant-temperature baths with rol-
ling or shaking devices. The principal difficulty in
using such chambers is the duplication of the radi-
ant energy spectrum and energy flux found in na-
ture (Strickland, 1958; 1960). Daylight flourescent
lamps may have energy spectra that depart widely
from that of sunlight (MacAdam, 1958). The use of
simple light meters to estimate radiation may lead
to both qualitative and quantitative errors. Fur-
thermore, there is some evidence that flourescent
lights inhibit the growth of some algae (Kain and
Fogg, 1960).
In spite of all these limitations and others,
closed-vessel methods are useful. The C meth-
od potentially offers much greater accuracy than
the light-dark-bottle method, but this accuracy, is
largely potential rather than actual at present.
Interpretation of the results of C14 uptake studies
is in a state of controversy (cf. Steeman-Neilsen,
1952; 1959; Ryther, 1956). Corrections for iso-
tope exchange and for respiration vary. Ryther
(1956) found that newly-respired carbon is used
preferentially for photosynthesis in cultures of
Dunaliella (see also Myers and Johnston, 1949).
If this is true under all natural conditions, it will
greatly improve the accuracy of the C14 method.
However, more experimental work under more
varied conditions is needed to see if this is true
under all natural conditions. At present the meth-
od offers little advantage except in very oligotro-
phic conditions where no other method is suffi-
ciently sensitive.
The light-dark-bottle method, used with proper
precautions (cf. Lund and Tailing, 1957; Steeman-
Nielsen, 1957; Strickland, 1960), is satisfactory
in situations of moderate to high productivity,
where observations need not be longer than 12
hours. This method is not fashionable at present,
but it requires a minimum of equipment and can be
made to yield satisfactory results in many situa-
tions.
2. Changes in the concentration of some product
or raw material of photosynthesis may be measured
in unconfined natural waters, to estimate net pho-
tosynthesis of all plant populations in the system.
The difficulties of confined samples are avoided,
and if the system is well mixed, the sampling pro-
blem is less. However, we must contend with the
estimation of import and export of the substance
measured from an open system.
A useful application of this type of approach is
the measurement of diurnal changes in dissolved
oxygen (Sargent and Austin, 1949; Odum, 1956;
1957). The method works well in highly produc-
tive, flowing systems. The limit on accuracy would
seem to be the estimation of the oxygen exchange
coefficient and, in unproductive or incompletely-
mixed systems, this is a serious limitation.
It is not certain whether the diurnal curve meth-
od is comparable with the light-dark-bottle meth-
od (cf. Jackson and McFadden, 1954; Tailing, 1957;
Odum and Hoskins, 1959; Pomeroy 1960a). Tailing
suggested that both methods were rather crude and
that close agreement should not be expected. Pro-
bably results within an order of magnitude should
be looked upon as in agreement.
Changes in pH can be used in much the same
way as changes in oxygen (Verduin, 1956a; 1956b).
The gas exchange difficulties are less. The meth-
od is less sensitive in sea water, and in any case
requires a very good pH meter.
Long-term changes in phosphorus have been
used to estimate production in the sea and coastal
waters (Riley, 1951; 1956; Steele, 1956). In its
present form it is only suitable for relatively sim-
ple systems, such as the sea, where interchange
of phosphorus with the land and the bottom is min-
imal. Short-term changes in phosphate concentra-
tion cannot be correlated with photosynthetic rates,
because the turnover of phosphate is rapid and
partly independent of photosynthesis (Pomeroy,
1960b).
.3. The measurement of chlorophyll content and
available energy offers a rapid, inexpensive proce-
dure (Ryther and Yentsch, 1957). It would not ap-
pear to offer great precision, regardless of re-
finements in technique. The accuracy of the meth-
od, aside from problems of sampling and pigment
analysis, depends on the estimation of the assimi-
lation number of the plant populations. The assi-
milation number may vary widely (Steeman-
Nielsen, 1959; Strickland, 1960), probably even
within a given system.
4. Harvest methods may be used to estimate
production where much of the plant material pro-
-------
Measurement of Production
duced accumulates in situ over a considerable
period. Such methods have been used to estimate
production of seaweed (Blinks, 1955), sea grasses
(Grntved, 1958), salt-marsh grasses (Smalley,
1959; Odum and Smalley, 1959) and some other
aquatic plants (Penfound, 1956). The scientist in-
evitably is harvesting side by side with the fishes,
snails or grasshoppers. To estimate net produc-
tion, it is necessary to know the food consumed by
the herbivores. This is possible (Teal, 1958) but
laborious.
In selecting a method for estimating primary
production, it is necessary to consider the kind of
-system to be studied and the questions to be asked
about it. The use of the C method is very pro-
mising, although its present widespread use does
not seem to be justified by our present knowledge
of the path of newly-respired carbon under diverse
natural conditions. With more complete knowledge
of these details the C method may permit quite
accurate work with rather simple field equipment.
Probably the most serious problems in estima-
ting production are the interpretation of the physio-
logical requirements and responses of the plants,
and the proper sampling of natural populations of
aquatic plants. Much attention has been given to
the physiological problems, and such as remain
for the ecologist may soon be solved. The samp-
ling problem has received considerably less at-
tention and, in my opinion, it is the greatest
limitation on accuracy of most current methods of
estimating production.
Natural plant populations typically are not ran-
domly distributed, and methods of statistical anal-
ysis suited for clustered, over-dispersed distri-
butions must be used. Such methods exist, and
they have been used occasionally in plankton stu-
dies (Barnes and Marshall, 1951; Barnes and
Hasle, 1957; Bliss and Calhoun, 1954) but seldom
if ever in production studies. The chief impedi-
ment to their application to production studies is
the need for many replications of the observations.
This becomes impossibly laborious and expensive.
The best way to surmount this difficulty would
seem to be by the use of techniques of automation.
The necessary equipment need not be more ex-
pensive than that needed for some of the methods
currently popular.
The use of polarographic electrodes makes it
possible to obtain a continuous record of changes
in oxygen tension in nature or in closed vessels
(Carritt and Kanwisher, 1959; Kanwisher, 1959).
Much more detail can be obtained in this way, and
it should be possible to incorporate oxygen recor-
ders in buoys, so a number of samples or stations
can be studied concurrently with a single ship or
field party. By the use of time switches the input
from several pairs of electrodes can be fed into
one recorder, or a multi-channel recorder can
make continuous records of all inputs. Incident
radiation and water tempe rature can be recorded
simultaneously, often on the same recorder.
A different automation technique is employed by
H. T. Odum, who has in operation an automatic
sampling buoy that collects a 24-hour series of
samples for Winkler determinations. While this
may lack some of the advantages of a continuous
record, it permits sampling of a wide spectrum of
chemical information. Similar installations along
the shoreline are now fairly common.
These, then, are some suggestions for improved
estimates of primary production: randomized or
otherwise statistically planned sampling programs,
data collected automatically over broad segments
of space and time, and statistical treatment of the
observations to give both an estimate of production
and a parameter of reliability. While it is not al-
ways possible to set up a program such as this, it
is a goal we might try to attain.
REFERENCES
Barnes, H., and G. R. Hasle. 1957. A statistical
examination of the distribution of some species of
dinoflagellates in the polluted inner Oslo Fjord.
NyttMag. Bot., !5: 113-124.
Barnes, H., and S. M. Marshall. 1951. On the var-
iability of replicate plankton samples and some ap-
plications of 'contagious' series to the statistical
distribution of catches over restricted periods. J.
Mar. Biol. Assoc. U.K., 30: 233-263.
Blinks, L. R. 1955. Photosynthesis and productiv-
ity of littoral marine algae. J. Mar. Res., 14j 363-
373.
Bliss, C. I., and D. W. Calhoun. 1954. An outline
of Biometry. New Haven, Yale Co-operative Corp.
272 pp.
Carritt, D. E., and J. W. Kanwisher. 1959. An
electrode system for measuring dissolved oxygen.
Anal. Chem., 31: 5-9.
Emerson, R., and L. Green. 1934. Manometric
measurements of photosynthesis in the marine al-
ga Gigartina. J. Gen. Physiol., 1/7: 817-843.
Gaarder, T., and H. H. Gran. 1927. Investigations
of the production of plankton in the Oslo Fjord.
Cons. Int. Explor. Mer, Rapp. et Proc.-Verb., 42j
3-48.
-------
64
ALGAE AND METROPOLITAN WASTES
Griffith, R. E. 1957. A portable apparatus for
collecting horizontal plankton samples. Ecology,
38: 538-540.
Grntved, J. 1958. Underwater macrovegetation in
shallow coastal waters. J. du Cons., 24: 32-42.
Pomeroy, L. R. 1959. Algal productivity in salt
marshes of Georgia. Limnol. Oceanogr., 4: 386-
397.
Pomeroy, L. R. 1960a. Primary productivity of
Boca Ciega Bay, Florida. Bull. Mar. Sci. Gulf and
Carib. In press.
Jackson, D. F. and J. McFadden. 1954. Phyto-
plankton photosynthesis in Sanctuary Lake, Pyma-
tuning Reservoir. Ecology, 35: 1-4.
Pomeroy, L. R. 1960b. Residence time of dis-
solved phosphate in natural waters. Science. In
press.
Kain, J. M., and G. E. Fogg. 1960. Studies on the
growth of marine phytoplankton. HI. Prorocen-
trum micans Ehrenberg. J. Mar. Biol.Assoc.U.K.,
39: 33-50.
Riley, G. A. 1951. Oxygen, phosphate, and nitrate
in the Atlantic Ocean. Bull. Bingham Oceanogr.
Coll., 13: 1-126.
Kanwisher, J. W. 1959. Polarographic oxygen
electrode. Limnol. Oceanogr., 4: 210-217.
Lund, J. W. G., and J. F. Tailing. 1957. Botanical
limnological methods with special reference to the
algae. Bot. Rev., 23: 489-583.
Mac Adam, D. L. 1958. Continuously recording
spectroradiometer. J. Opt. Soc. Amer., 48: 832-
840.
Myers, J., and J. A. Johnston. 1949. Carbon and
nitrogen balance of Chlorella during growth. Plant
Physiol., 24: 111-119.
Odum, E. P., and A. E.Smalley. 1959. Comparison
of population energy flow of a herbivorous and a
deposit-feeding invertebrate in a salt marsh eco-
system. Proc. Nat. Acad. Sci., 45: 617-622.
Odum, H. T. 1956. Primary production in flowing
waters. Limnol. Oceanogr., 1: 102-117.
Odum, H. T. 1957. Trophic structure and produc-
tivity of Silver Springs, Florida. Ecol. Monogr.,
27: 291-320.
Riley, G. A. 1956. Oceanography of Long Island
Sound. IX. Production and utilization of organic
matter. Bull. Bingham Oceanogr. Coll., 1J5: 324-
344.
Ryther, J. H. 1956. Interrelation between photo-
synthesis and respiration in the marine flagellate
Dunaliella euchlora. Nature, 178: 861-862.
Ryther, J. H., and C. S. Yentsch. 1957. The es-
timation of phytoplankton production in the ocean
from chlorophyll and light data. Limnol. Ocean-
.ogr., 2: 281-286.
Sargent, M. C., and T. S. Austin. 1949. Organic
productivity of an atoll. Trans. Amer. Geophys.
Union, 30: 245-249.
Smalley, A. E. 1959. Comparison of energy flow
in two invertebrate populations in a salt marsh
ecosystem. Doctoral thesis, University of Georgia.
Steele, J. H. 1956. Plant production on the Fladen
Ground. J. Mar. Biol. Assoc. U. K., 35: 1-33.
Steele, J. H. 1959. The quantitative ecology of ma-
rine phytoplankton. Biol. Rev., 34: 129-158.
Odum, H. T., and C. M. Hoskins. 1959. Compara-
tive studies on the metabolism of marine waters.
Pub. Mar. Sci. (Univ. Texas). In press.
Steeman-Nielsen, E. 1952. The use of radio-active
carbon (C^) for measuring organic production in
the sea. J. du Cons., 18: 117-140.
Penfound, W. T. 1956. Primary production of
vascular aquatic plants. Limnol. Oceanogr., 1:
92-101.
Steeman-Nielsen, E. 1957. Experimental methods
for measuring organic production in the sea. Cons.
Int. Explor.Mer, Rapp.et Proc.-Verb., 144: 38-46.
-------
Measurement of Production
65
Steeman-Nielsen, E. 1959. Untersuchungen uber
die Primarproduktion des Planktons in EinigenAl-
penseen. Osterreichs. Oikos, 10: 26-37.
Strickland, J. D. H. 1958. Solar radiation pene-
trating the ocean. A review of requirements, data
and methods of measurement, with particular re-
ference to photosynthetic productivity. J. Fish.
Res. Bd. Canada, 15: 453-493.
Strickland, J. D. H. 1960. Measuring the produc-
tion of marine phytoplankton. Bull. Fish. Res. Bd.
Canada. In press.
Tailing, J. F. 1957. Diurnal changes of stratifi-
cation and photosynthesis in some tropical African
waters. Proc. Roy. Soc. London, B147: 57-83.
Tailing, J. F. 1960. Comparative laboratory and
field studies of photosynthesis by a marine plank-
tonic diatom. Limnol. Oceanogr., _5_: 62-77.
Teal, J. M. 1958. Energy flow in the salt marsh
ecosystem. Proc. Conf. Salt Marshes, pp. 101-
107.
Verduin, J. 1956a. Primary production in lakes.
Limnol. Oceanogr., ^:85-91.
Verduin, J. 1956b. Energy fixation and utilization
by natural communities in Western Lake Erie.
Ecology, J57: 40-50.
FACTORS WHICH REGULATE PRIMARY PRODUCTIVITY AND
HETEROTROPfflC UTILIZATION IN THE ECOSYSTEM*
EUGENE P. ODUM
Department of Zoology,
University of Georgia, Athens, Gegrgia
In this paper algae are considered as compon-
ents of the ecosystem and the algal bloom is viewed
as an unbalance between production and consump-
tion of organic matter in the ecosystem. We might
even go so far as to think of the noxious bloom as
a sort of cancer in the metabolism of Nature. Ab-
normal growths, whether at the cellular or at the
ecosystem level, result in the final analysis from
the failure of, or at least the overtaxing of, regu-
latory processes. In the case of the metropolitan
waste problem the excess growth results, of
course, from the overloading of the system with
growth-promoting inorganic and/or organic sub-
stances. If control is to be more than a trial and
error proposition we must study and understand
not only the growth itself (i.e. the "bloom")but also
the system in which the growth occurs (i.e. the
body of water).
A couple of other general points may well be
emphasized before we get down to cases. First,
descriptive information alone rarely solves prac-
tical problems; functional information is also usu-
ally needed. Dr. Pomeroy in the previous paper
has discussed some of the methods of measuring
one important rate function, that is, primary pro-
duction. It was evident from his paper that pro-
cedures such as the diurnal oxygen curve method,
which measures function of the whole ecosystem,
are potentially of greater practical value than
methods which are restricted to small samples in
bottles. Much remains to be done to establish the
reliability of total system measurements, but un-
til we do have such measurements we will be greatly
handicapped in attempts to understand any specific
component within the system. As will be pointed
out later there are important functions other than
primary production which must be considered.
A second point is that in applied research we
observe again and again that no single method of
control can be counted on to work or to be feasible
under all of the varied conditions of Nature. The
control of algal blooms will certainly prove to be
no exception to this rule. Consequently, we should
not restrict our thinking to one or a few approaches
just because a measure of success has been a-
chieved with them. We need to have other possi-
bilities ready in case the recommended method
Contribution No. 20 from the University of Georgia Marine Institute, Sapelo Island, Georgia.
-------
66
ALGAE AND METROPOLITAN WASTES
does not work. We must also be prepared for un-
foreseen changes in the nature of wastes resulting
from social and economic development beyond the
control of the mere scientist. Sooner or later, at
some time or place, any theoretically sound ap-
proach may prove practical. There should be no
limit to the arsenal of the mind 1
At this point it would be well to. review briefly
the basic theory of ecosystem function. Figure 13
illustrates in a simplified manner what I like to
call "the two basic principles of functional ecolo-
gy", namely, the one-way flow of energy and the
cycling of materials (nutrients, etc.). The "boxes"
in the diagram represent the standing crops of
functional autotrophs and heterotrophs in terms of
weight or calories per unit area. The "pipes" re-
present energy flow in terms of calories per unit
time. The "stippled circle" represents movement
of nutrients and other materials from one biolog-
ical unit to another, or to and from a pool within
the system. A point to emphasize is that while ma-
terials circulate and inorganic nutrients may be
used over and over (if not lost from the system)
the flow of energy is one-way; once utilized in
respiration the energy is dispersed as heat and
lost from the system. Imports and exports of both
materials and fixed energy are also shown on the
diagram; such imports and exports, of course, are
of primary concern in waste disposal situations.
HETEROTROPHS
figure 13. SIMPLIFIED FLOW DIAGRAM OF AN
ECOSYSTEM SHOWING THE ONE-WAY FLOW OF
ENERGY AND THE RECYCLING OF MATERIALS.
The "boxes" represent the standing crops of auto-
trophic and heterotrophic components, the open
"pipes" the flow of energy and the stippled circuit
the flow of materials (nutrients, etc). PG = gross
production; PN = net primary production; P =
heterotrophic production which may be consumed
within the system (by secondary consumers, i.e.
carnivores, etc., which are not shown on the dia-
gram). It maybe stored or exported from the sys-
tem.
Another point to emphasize is that there are
two kinds of primary production but only one kind
of secondary production. In the diagram, total
phytosynthesis is designated as gross production
while that portion of the fixed energy not used in
plant respiration and ending up as "growth" is
designated as net production. At the heterotro-
phic level the only thing "produced" is "growth".
In this broad sense "growth" asaphenomenon
includes not only additions to the biomass but
production of stored food and soluble organic ma-
terials which may leak out or be excreted from
the organisms. A source of much confusion is that
the various methods of measuring primary produc-
tion, as described in the preceding paper, do not
measure the same thing; some methods measure
gross primary, some net primary, others appar-
ently estimate a quantity in between, while still
other methods measure net community production
(i.e. gross production minus community respira-
tion). Thus, it is often difficult to compare differ-
ent situations where different methods have been
employed. Finally, as clearly shown in Figure 13,
energy flow at any level, or for any specific spe-
cies or population, is the sum of production and
respiration. If the population is growing rapidly
production may be a major part of energy flow,
but if the population is not growing then all of the
energy flow may be respiration. In our preoccu-
pation with production and its measurement we
must not forget that consideration of respiration
is equally important. None of these important func-
tions can be determined by merely describing the
composition of the standing crop or by making sin-
gle estimates of its size.
Diagrams such as that of Figure 13 can be made
quantitative to represent a particular situation.
The area of the "boxes" can be made proportional
to the standing crops, and the diameter of the ener-
gy flow "pipes" and the nutrient "pipes" can be
made proportional to the rates of flow. Above all,
however, such graphic representations of energy
flow enable us to visualize ways in which unbalance
between production and consumption may come
about, and, by the same token, we may visualize
various ways in which the unbalance might be cor-
rected. For example, if the inflow of nutrients is
increased gross production will increase unless
the inflow is counteracted by (1) loss from the sys-
tem or into an unavailable pool or (2) a decrease
in the available light energy (as might result from
increased turbulence, increased turbidity or shad-
ing). Increase in gross production will result in
an increase in standing crop of autotrophs and per-
haps the development of a bloom situation unless
there is a compensatory energy flow through the
community respiration "pipes" (i.e. plant respi-
ration plus heterotrophic respiration) or unless
there is immediate and efficient conversion of net
production into herbivore biomass. Thus, we
quickly visualize that there are several quite dif-
-------
Factors Regulating Productivity and Utilization
67
ferent ways by which a bloom resulting from ex-
cess inorganic nutrients-might be controlled, as
for example: (1) reducing inflow of nutrients (i.e.
"stripping"). (2) diverting nutrients into a pool (as
by converting into unsolubleor unavailable forms),
(3) reducing effective light, (4) increasing respira-
tion of the algae themselves (i.e. "self destruc-
tion"), (5) increasing direct consumption of net pro-
duction by herbivores (i.e. increasing grazing),
and (6) exporting (i.e. removing) organic matter
(and with it some of the nutrients) either as pri-
mary (algae) or as secondary production (for ex-
ample, fish). How effective any of these approaches
would be depends, of course, entirely on quantita-
tive considerations, that is, how much or how fast
materials or energy can be moved along the routes
in question.
Increasing imports of organic matter result in
a somewhat different situation as can be visualized
in the diagram. Nature develops several mechan-
isms to deal with the situation but, of course, these
mechanisms are commonly overtaxed by the mag-
nitude of metropolitan wastes. Increased consump-
tion by microorganisms and other heterotrophs,
and increased export downstream or storage with-
in the system are the usual direct responses of the
ecosystem. The increased community respiration,
of course, puts critical demands on the oxygen
supply which becomes the major limiting factor.
As organic matter piles up between the auto tro-
phic and heterotrophic "boxes" (see diagram) some
of it may flow backwards, as it were, and the auto-
trophs may become heterotrophs. From what little
we know about this at present it would appear that
this route cannot be very effective as a means of
reducing the organic matter. From the standpoint
of the ecosystem as a whole it would appear that
extensive shifts from autotrophic to heterotrophic
nutrition (in those species of algae which are capa-
ble of doing so) is a sort of last-ditch regulatory
mechanism. Certainly, organisms adapted to pho-
tosynthetic metabolism can hardly be expected to
be as efficient consumers of organic matter as ob-
ligatory heterotrophs, or can they?
One of the reasons metropolitan wastes present
such difficult problems is that they consist of com-
plex mixtures of inorganic nutrients and energy-
containing organic materials (plus poisons of vari-
ous kinds!). It is a striking but little understood
phenomenon that blooms resulting from organic nu-
trients seem to be more resistant to consumption
by the community than are those resulting from in-
organic fertilization. The kinds of algae (blue-
greens or filamentous types, for example) which
respond to organic nutrients often seem to be un-
palatable to consumers. Or perhaps antibiotics
are produced which inhibit consumers. Either pos-
sibility produces the same thing; something about
the chemical composition of the bloom favors de-
layed consumption often resulting in a big pile up of
algae followed by decomposition of dead matter and
depletion of oxygen. The well documented situation
at Moriches Bay in Long Island Sound (Ryther,
1954) illustrates what often seems to happen. In
this example when the nutrient flow shifted from
inorganic to organic form (as a result of duck
farming along the shore) previously rare and un-
known species of algae, which were able to use
organic phosphorous and nitrogen, produced large
blooms. Unlike the diatoms and other phytoplankton
previously present in the bay the new forms were
not readily eaten by zooplankton and the shellfish.
As a result both the human bathers and the oysters
suffered. We certainly need to know a great deal
more about why certain types of blooms are more
resistant to natural consumption than are others.
And of course we would like to know if by regula-
ting the type of inflow we can avoid encouraging the
resistant -type of bloom.
In discussing the functional processes in the
ecosystem it would be well to keep in mind some
orders of magnitude. The world distribution of
primary production has been summarized by E. P.
Odum (1959) and Ryther (1959). The quantitative
relationships between production and consumption
(i.e. Community P/R ratios) have been considered
from the theoretical standpoint by H. T. Odum
(1956, 1960), and with reference to lakes by Verduin
(1956), Ohle (1956) and others, and with reference
to estuaries by Odum and Hoskins (1958). The most
naturally fertile parts of the biosphere are the
shallow waters, areas such as marshes, shallow
(eutrophic) lakes and ponds, estuaries and shallow
coastal seas (especially with "upwhelling") as well
as the alluvial plains, deltas and coastal terraces.
Gross primary production in such areas may aver-
age 3-10 grams of dry matter per square meter
per day for extended periods (to visualize these
figures in approximate pounds per acre, multiply
by 10) with even higher rates occurring during the
most favorable seasons. In these fertile ecosys-
tems the autotrophic and heterotrophic components
(the "boxes" of Figure 13) are close together physi-
cally (thus promoting rapid regeneration of nutri-
ents) while light, water and other conditions of
existence tend to be favorable. In lakes, produc-
tivity is inversely related to depth (Rawson 1952)
or to put it another way, is inversely related to
distance between major autotrophic and hetero-
trophic components. Production rates may rise as
high as 60 gms/M2/day in bloom situations. Some
of the highest sustained production rates so far
"officially" recorded have been found in sewage
ponds (see Bartsch and Allum, 1957). At the other
extreme, deep lakes, the deep oceans and land de-
serts may average less than 0.5 gm/M2/day. A
large portion of the earth is in the low production
-------
68
ALGAE AND METROPOLITAN WASTES
category where even temporary blooms are rare,
unless, of course, production Is augmented by man.
In most naturally fertile aquatic ecosystems
production tends to get ahead of respiration, at
least seasonally, with the result that some organic
matter is either stored or exported. However,
there are many fertile aquatic ecosystems which
rarely develop blooms in the sense of a large ex-
cess of standing crop accumulating at anyone time.
Such systems need to be carefully studied for clues
as to how man might handle situations where he
inadvertently boosts fertility (i.e. "artificial eu-
trophication"). In our program at the Marine In-
stitute of the University of Georgia we are intense-
ly interested in one such fertile system which com-
prises the vast salt marshes and estuaries of the
Georgia coast. At present we are promoting a
multi-disciplined study in which ecologists, bio-
geochemists, microbiologists and geologists are
taking part Primary production in the marsh-
estuary system may reach 5-10 gms/M2/day for
a good portion of the year, yet except for an oc-
casional outburst of dinoflagellates in limited areas
(Pomeroy, Haskins and Ragotzkie, 1956) blooms
are rare. Strong tidal mixing and export to the
sea, of course, are important factors, but both
macro-consumers (fiddler crabs, mussels, snails
and fish) and micro-consumers (bacteria) are very
active. Algae living on or near the surface of the
vast intertidal sediments have a surprisingly high
productivity which is maintained throughout the
year (Pomeroy, 1959), yet the standing crop re-
mains small. What portion of the net production
is removed by currents'and what portion is utilized
by the abundant heterotrophs associated with the
sediments has not yet been determined.
The coral reef is another, very fertile natural
system where community respiration is, well bal-
anced against community production; In our work
on Eniwetok reefs (Odum and Odum, 1955) we were
impressed not only with the importance of water
movement but also with the following: (1) the inti-
mate association of algae with corals and many
other animals, (2) the highly developed symbiosis
between autotrophs and heterotrophs in general,
and (3) the large standing crop of consumers in-
cluding large numbers of herbivorous fish which
were capable of consuming algae as fast as pro-
duced. The important point I would like to make is
that there is nothing in nature to indicate that high
fertility must necessarily generate noxious blooms.
Nor is there any reason to assume that poisons or
other chemical controls are the only possible ap-
proaches to artificial eutrophication. It ought to be
possible to establish reasonable balance by physi-
cal manipulations of water movement and by stim-
ulating heterotrophic utilization. It may be neces-
sary, however, to get man into the food chain. We
seem to have a paradox here. In many parts of the
world people are crying for more production, while
at this seminar we seem to be crying equally loudly
for less production! Perhaps it would be well to
begin thinking in terms of making use of the in-
creased productivity created by cities rather than
attempting to destroy it.
The diagram in Figure 13 is over simplified in
that only one broad energy flow is shown. Actually,
between the first and second trophic levels in
most ecosystems the energy flow of the community
is divided into two broad streams resulting in two
types of primary consumption: (1) direct and im-
mediate utilization of living plant tissues by her-
bivores and plant parasites, and (2) delayed uti-
lization of dead tissues and stored food by other
consumers; In our work on the Georgia estuaries
we have been impressed with the fact that the ratio
of detrital consumers to direct herbivores appar-
ently is an important factor in determining the pat-
tern of heterotrophic utilization (Odum and Smal-
ley, 1959). If detrital consumers (bottom fauna
and microorganisms, for example) predominate
then a pulsating energy flow may be expected with
the likelihood of periodic pile-ups of primary pro-
duction assuming little or no export If herbivo-
res (i.e. "grazers" such as zooplankton) predomin-
ate then a more nearly steady-state may be main-
tained. In many natural waters, such as Long
Island Sound, grazing by zooplankton provides an
important check on spring blooms (Riley, 1956).
As already indicated, waste disposal situations do
not favor grazing by microcrustacea. The rela-
tive size of autotrophs and heterotrophs and the
"size-metabolism law" (i.e. higher rates of metab-
olism per grams are small as compared to large
organisms) are important considerations. To be
effective, herbivores must be large enough indi-
vidually to consume the algae (or else capable of
breaking them down to "bite size") yet have a high
enough rate of metabolism per gram to bring about
a high rate of consumption. Also the question of
antibiotic effects comes up again. There is no
doubt that algae often produce antibiotics which af-
fect their growth (Proctor, 1957). What is not
clear is whether algae may produce in significant
quantities metabolites which inhibit herbivores.
Harvey's (1955) "exclusion theory" (to explain
patchy distribution of phyto - and zoo-plankton in
the sea) is based on such an assumption, but there
is yet no proof. While we are thinking along these
lines it would be well to point out that Nature rare-
ly works only one side of the street Thus, there
may be substances which stimulate the herbivores
to graze. Perhaps in the future we can look for-
ward to the time when we may be able to apply an
"environmental hormone" to a body of water to
bring about increased heterotrophic consumption
of the growth in the same manner as hormone ther-
-------
Factors Regulating Productivity and Utilization
69
apy might be employed to correct unbalance in the
human body. While we are letting our imagination
run wild, it might also be possible some day to
shift the enzyme systems in the algal blooms so
that plant respiration is increased at the expense
of photosynthesis thus bringing about "self destruc-
tion" of the excess growth. Although sufficient
data are not yet available, it would seem that the
autotrophic component in many subtropical and
tropical ecosystems exhibits a rather low net pro-
duction because a high rate of respiration counter-
acts high gross production.
To return to the more immediately practical
mere conversion of net production into herbivores
may not provide the answer if the rate of importa-
tion of fertilizer continues high. Some sort of har-
vest (i.e. "export") must be worked out so that some
of the nutrients as well as the organic matter can
be removed from the system or that part of the
system which is causing trouble. Harvest of large
organisms which turn over slowly and accumulate
energy over periods of time (i.e. long periods of
individual growth) will likely be most practical.
Harvesting algae or small herbivores on a large
scale requires frequent or continuous operations,
whereas harvest of larger units can be periodic.
Even if filters or screens can be developed, main-
tenance problems would be great in "gunky" envi-
ronments. Therefore, it appears that truly herbi-
vorous species of fish which consume algae directly
and are adapted to tolerate organically fertile wa-
ters provide very definite candidates for a "con-
version-harvest" solution. We know very little
about such fish in this country. Fish management
or fish culture in fresh water in the United States
is almost exclusively concerned with sport fishes
which are grown on a long food-chain (usually
through bottom fauna) involving delayed utilization
of primary production and consequent low yields
per acre. Sports fish would be completely ineffec-
tive, and, in fact, could not survive in many waste
disposal situations. As I understand it there are
species of fishes in India, in the Orient and in the
Philippines which eat algae, even blue-greens and
filamentous types (which as already indicated are
resistant to direct consumption). Yields of 1000
pounds or greater, per acre per year, are common
in oriental ponds stocked with herbivores (Hickling,
1948). Such yields would amount to a secondary
production energy flow of 100 gms/M^/year or
about 0.3 gms/M^/day. If the assimilation efficien-
cy (ratio production to total assimilation)of the fish
population were no more than one-third, which is
to say that the respiratory energy flow would be at
least twice the production, then the fish would con-
sume at three times their production rate or about
one gram dry weight of algae per square meter per
day. Referring to the primary production levels
previously discussed it can be seen that such a
herbivorous fish population could be effective ex-
cept in the more explosive bloom situations. I
shall not attempt to estimate the "biological strip-
ping" effect which an annual harvest of 1000 pounds
per acre would have on the nutrients, but it would
seem that the effect could be important, at least
with reference to micronutrients and perhaps to
phosphorus.
The big ecological point that I am making is this:
When large amounts of organic wastes, etc., are
introduced into natural waters essentially a new
ecosystem is produced which may lack some bio-
logical components simply because there are no
adapted species available. A definite "open niche"
is certainly present in sewage ponds which contain
no fish or other grazers. Under such conditions
the possibilities of filling the niche are worth in-
vestigating. We might find what we need in some
other part of the world where organic pollution
has been in effect for a longer period of time; or
we might breed a strain especially adapted for the
situation. Such a procedure would amount to "bio-
logical engineering" in that we seek to adapt a
population to a function. The game bird situation
in the Middle West will illustrate the point. When
natural grasslands and forests were converted by
man into corn fields, the productivity was not
changed, but the habitat was. No native species
of gallanaceous bird could adapt and take advan-
tage of the high productivity of such lands. The
ring-necked pheasant, which had a long history of
adaptation in the intensive grain farming regions
in Europe and Asia, filled the niche very well when
it was introduced. We are all leary, of course, of
introducing exotics because they sometimes be-
come pests. In my opinion the "open niche" situa-
tion is the only situation where we should consider
introductions, and we should make very certain
that our introduction is very specialized for the
intended niche. If specialized for the job it will not
only be more efficient but it will be less likely to
spread into waters where it is not wanted.
One other factor which may affect heterotrophic
utilization should be mentioned. Species diversity
may be very important in determining the pattern
of energy flow. Our studies on organic production
and turnover in old-field succession (Odum, 1960)
as well as the work of Margalef (1958) on phyto-
plankton succession indicate that the process of
diversification which usually occurs during eco-
logical succession favors stability of the ecosystem
often at the expense of total production at primary
level.
Typically, early stages of succession are char-
acterized by a small number of species, which,
however, have rapid growth rates. As the number
of species increases, production is divided among
-------
70
ALGAE AND METROPOLITAN WASTES
more kinds and is thus spread out more uniformly
in both time and space. Since many of the later
invaders maybe low-yielding types, the total com-
munity production may be reduced, but from the
functional standpoint the ecosystem is more stable
and is better able to regulate itself and to adapt to
changing climate or other conditions imposed from
without. Seasonal succession, as well as the slower
annual changes, seems to conform to this pattern.
Anything which can be done to increase diversity at
both autotrophic and heterotrophic levels should
help stabilize productivity. Tailing (1951) has
pointed out that the element of chance may play a
large part in determining what species become
dominant in new ponds. It would be interesting to
experiment with seeding new waste stabilization
ponds with a variety of species of algae to see if
the composition of the community could be regu-
lated and the pattern of primary production stabi-
lized.
To summarize, the basic theory of ecosystem
function, as graphically illustrated by the energy
flow diagram, points to a number of approaches
to control of the unbalance between production and
consumption which is inherent in the noxious bloom
situation. Some possibilities are: (1) reducing in-
flow of nutrients, (2) converting nutrients to un-
available form, (3) poisoning production machin-
ery, (4) reducing effective light, (5) increasing
respiratory energy flow in autotrophs, (6) in-
creasing "grazing" by herbivores, and (7) har-
vesting net production (and with it some of the nu-
trients) either as primary or secondary produc-
tion. The first three possibilities were not
discussed in this paper, since they are considered
in detail in other papers in the symposium. In-
creased depth and increased vertical turbulence
do reduce productivity in natural waters. While
the first three or four approaches are the ones
being currently investigated, they are essentially
"negative" in that attempts are made to prevent or
destroy natural productivity. The viewpoint of this
paper has been that accumulation of undesirable
organic materials following algal blooms may be
as much a problem of delayed or incomplete utili-
zation as it is a problem of increased primary
production. Consequently, attention was focused
on the utilization side of the picture, with the sug-
gestion that we should at least be thinking about
using the fertility of cities rather than doing away
with it. A consumer-harvest approach involving
herbivorous fish was suggested as deserving prac-
tical consideration. Estimation of population en-
ergy flow based on recorded yields of herbivorous
fish in the Orient indicates that such an approach
could be effective.
REFERENCES
Bartsch, A. F. and M. O. Allum, 1957. Biological
factors in the treatment of raw sewage in artificial
ponds. LimnoL andOceanogr., 2: 77-84.
Harvey, H. W. 1955. The chemistry and fertility
of sea water. Cambridge Univ. Press, Cambridge,
England.
Hickling, C. F. 1948. Fish farming in the Middle
and Far East Nature, 161; 748-751.
Odum, H. T. 1956. Efficiencies, size of organ-
isms, and community structure. Ecol, 37: 592 -
597.
1960. Ecological potential and
analogue circuits for the ecosystem. Amer. ScL,
48: 1-8.
Odum, EL T. and C. M. Hoskins. 1958. Compara-
tive studies on the metabolism of marine waters.
PubL Inst Mar. ScL, Univ. Texas, 5: 16-46.
Odum, H. T. and E. P. Odum. 1955. Trophic struc-
ture and productivity of a windward coral reef
community on Eniwetok AtolL Ecol. Monogr., 25:
291-320.
Odum, E. P. 1959. Fundamental of Ecology, Se-
cond Edition. W. B. Saunders Co., Philadelphia.
pp 1-546.
1960. Organic production and
turnover in old-field succession. Ecol, 41; 34-49.
Odum, E. P. and A. E. Smalley, 1959. Comparison
of population energy flow of a herbivorous and a
deposit-feeding invertebrate in a salt marsh eco-
system. Proc. Nat Acad. ScL, 45; 617-622.
Ohle, W. 1956. Bioactivity, production and energy
utilization of lakes. Limnol. and Oceanogr., 1:
139-149.
-------
Factors Regulating Productivity and Utilization
71
REFERENCES (Cont'd.)
Margalef, Ramon. 1958. Temporal succession and
spatial heterogeneity in phytoplankton. Perspec-
tives in Mar. Biol. Univ. California Press, pp 323-
349.
Pomeroy, L. R., H. H. Haskins, and R. A. Ragotz-
kie. 1956. Observations on dinoflagellate blooms.
LimnoL and Oceanogr., 1: 54-60.
Pomeroy, L. R. 1959. Algal productivity in salt
marshes of Georgia. Limnol. and Oceanogr., 4:
386-397.
Proctor, Vernon W. 1957. Studies on algal anti-
biosis using Hemato coccus and Chlamydomonas.
LimnoL and Oceanogr., 2: 125-139.
Rawson, D. S. 1952. Mean depth and fish produc-
tion in large lakes. EcoL, 33: 513-521.
Riley, Gordon A. 1956. Production and utiliza-
tion of organic matter. Bull. Bingham Oceanogr.
ColL, 15: 324-344.
Ryther, John H. 1954. The ecology of phytoplank-
ton blooms in Moriches Bay and Great South Bay,
Long Island, New York. Biol. Bull., 106: 198-209.
1959. Potential productivity of
the sea. Science, 130: 602-608.
Tailing, J. R. 1951. The element of chance in pond
populations. The Naturalist (London); Oct. - Dec.:
157-170.
Verduin, Jacob. 1956. Energy fixation and utili-
zation by natural communities in western Lake
Erie. Ecol, 37: 40-50.
-------
SOURCES OF NUTRIENTS
-------
Land Drainage
LAND DRAINAGE AS A SOURCE OF PHOSPHORUS IN ILLINOIS SURFACE WATERS*
R. S. ENGELBRECHT and J. J. MORGAN
Respectively, Professor of Sanitary Engineering and Instructor in Sanitary Engineering,
Department of Civil Engineering, University of Illinois, Urbana, Illinois
INTRODUCTION
Phosphorus in surface waters may stem from a
variety of sources. Major sources of phosphate
substances are human wastes, chemicals em-
ployed in water conditioning, synthetic detergents,
certain industrial wastes, and drainage of agricul-
tural lands. The purpose of this paper is to dis-
cuss some of the factors which bring about various
concentrations of phosphate substances in the
drainage water from agricultural lands, and to
present the results obtained in stream surveys of
several Illinois surface waters, particularly the
River. These studies indicate the im-
portance of land drainage as a source of ortho-
phosphate, hydrolyzable phosphate, and organic
phosphate in streams which drain highly cultivated
lands.
PHOSPHORUS IN AGRICULTURAL LAND
Bennett (1939) reported that analyses of 389
samples of surface soils throughout the United
States showed an average phosphoric acid con-
tent of 0.15 per cent, as P2Os. Waggaman (1952)
stated that phosphoric acid is the main fertilizer
element used in American agriculture, and the pre-
dominant ingredient of nearly all mixed fertilizers.
Waggaman observed a phosphoric acid content of
0.07 to 0.25 per cent for the first nine inches of
good productive soils, or 1750 to 6250 pounds per
acre, as PgOs. Only a small fraction of this total
is available as soluble phosphoric acid. Melsted
(1960) has indicated that the available PaOs of farm
lands in the Kaskaskia River basin of Illinois var-
ied from 43 to 50 pounds per acre. It is a well
known fact that the readily soluble phosphates of
commercial fertilizers soon revert in the soil to
less soluble phosphoric acid compounds.
TRANSPORT OF PHOSPHORUS TO SURFACE
WATERS
Phosphorus carried to surf ace waters maybe in
the simple orthophosphate form or as soluble hy-
drolyzable phosphate, or it may be adsorbed on
clay particles. It is known that much of the soil
phosphate exists in an adsorbed form on soils and
clays. According to Kurtz (1945), as these adsorbed
forms of phosphate increase in amount their solu-
bility in water increases rapidly. Silvey (1953) re-
ports that clays act as ion exchange compounds and
that certain phosphates in calcium bearing soils
would be converted to calcium phosphate and re-
main in the humus or the sub-soil until the pH
closely approaches 7.0, at which time the soluble
phosphate would be eluted by runoff, percolation,
and subsurface seepage.
Kohnke and Bertrand (1959) suggest that typical
surface runoff water is high in solid soil particles,
especially clay and organic matter, high in ad-
sorbed phosphorus, and low in soluble salts. If
runoff occurs quickly after the start of a rain, sol-
uble salts, accumulated in the surface soil during
a dry period, may be washed off before infiltration
carries them into the body of the soil. Percolation
water contains a relatively high concentration of
soluble salts. Thus, for an area which is drained
by tile, surface waters might be expected to re-
ceive significant amounts of both adsorbed and
soluble phosphorus in times of high rainfall and
runoff.
Concentrations of available phosphate in seepage
waters from agricultural soils were determined in
lysimeter studies conducted by the Department of
Agronomy at the University of Illinois. Melsted
(1960) reported P£O5 concentrations in percola-
tion water of from 0.2 to 0.7 mg/1, depending upon
the rate of percolation. A concentration of approxi-
mately 0.5 mg/1 PoOs per acre-inch of water was
typical of the results obtained.
The contribution of phosphate to surface waters
by land drainage in the Madison, Wisconsin, area
was investigated by Sawyer (1944). He reported to-
tal phosphorus from land drainage of approximately
1.6 Ib PjjOs/day/sq. mile, of which about 75 per
cent was in the organic form.
PHOSPHATES IN ILLINOIS SURFACE WATERS
The Sanitary Engineering Laboratory of the
University of Illinois conducted surface water sur-
veys in 1956 and 1957 to determine concentrations
of phosphates in Illinois surface waters. The meth-
*This investigation represents part of a research project supported by the Association of American Soap and
Glycerine Producers, Inc.
74
-------
Land Drainage as a Source of Phosphorus
od used in the analyses of samples for orthophos-
phateand hydrolyzable phosphate was that reported
by the Association of American Soap and Glycerine
Producers Subcommittee on Phosphates (Anon,
1958).
Figure 14 is a map of Illinois, showing the loca-
tions of lakes and reservoirs, streams, and sew-
age treatment effluents sampled during the surveys.
Table 16. TYPICAL LEVELS OF DIFFERENT
SURFACE WATER SOURCES, ILLINOIS, 1956
LAKE OR RESERVOIR
SOURCE
STREAM SOURCE
SEWAGE TREATMENT
EFFLUENT
Figure 14. SAMPLING LOCATIONS IN ILLINOIS
The overall results of the various surveys and the
details of sampling and analysis have been reported
by Engelbrecht and Morgan (1959). Table 16 pre-
sents typical results for P2Os concentrations found
in different surface waters. It is seen that phos-
phate concentrations in surface waters vary consi-
derably.
One of the samples from a relatively pollution-
free reservoir, Lake Bloomington, showed an
ortho plus hydrolyzable P2Os concentration of 0.23
mg/1, of which only 10 per cent was orthophos-
phate. This reservoir has a drainage area of 60
square miles. In contrast to the low phosphate
concentrations found in reservoirs were those ob-
served in a small stream receiving the industrial
Source
Lakes and reservoirs,
8 samples.
Major drainage basins,
moderate domestic pollution
25 samples.
Streams with high
domestic pollution,
33 samples.
Stream with high
industrial pollution,
7 samples.
Ortho P2°5
mg/1
0.04
0.37
6-15
77
Ortho t Hydrolyzable
P205, mg/1
0.09
0.63
7-16
81
waste from a metal finishing plant, where levels of
80 mg/1 P2Os were noted.
KASKASKIA RIVER BASIN SURVEY
In order to assess the relative amounts of phos-
phate from land drainage and domestic sources,
one major drainage basin was selected for an in-
tensive survey. Phosphorus and stream flow data
were then analyzed according to the scheme out-
lined in Table 17. The Kaskaskia River was chosen
Table 17. STREAM PHOSPHATE BALANCE
(1) S = L + W + 1
or
(2) Q-c = A-a + P-p -t- 1
S = Q-c = Pounds in stream
Q = Runoff
c = Concentration
L = A-a = Pounds from land drainage
A = Area drained
a = Pounds per unit area
W = P-p = Pounds from domestic sources
P = Total population
p = Per capita quantity
1 =. Industrial quantity
for this survey because it was convenient to the
laboratory, because of the existence of a system
of rain gages and USGS stream gaging stations
along its length, and because of the availability of
hydrological and agricultural data for the water-
shed area.
The location of the Kaskaskia River is shown in
Figure 14, while the locations of the various stream
sampling points and sewage treatment effluents
sampled are shown in Figure 15. Six stream sam-
pling points are shown. Descriptive data for each
stream sampling point are given in Table 18. Farm
lands in the Kaskaskia basin are drained mainly by
tile drain placed at a depth of 24 inches.
-------
76
ALGAE AND METROPOLITAN WASTES
A Stream sampling station
Sewage treatment plant
effluent
Figure 15. KASKASKIA RIVER BASIN. SHOWING
SEWAGE TREATMENT EFFLUENTS AND
STREAM SAMPLING STATIONS
Table 18. DESCRIPTIVE
RIVER BASIN
DATA, KASKASKIA
Stream
1
1
3
4
5
Mile
273
245
170
120
65
0
Tributary
population
0
820
6.670
12.850
18.320
90,240
Total Drainage
area,sq. mi.
12
125
1030
1980
2680
5220
Per cent
cultivated
86
86
82
76
69
77
Phosphate from Domestic Sources
There were no known phosphates discharged
separately as an industrial waste along the Kas-
kaskia River. Therefore it was only necessary to
account for phosphate contributions from sewage
treatment plant effluents in order to estimate the
magnitude of the phosphate contributions from land
drainage above each of the stream sampling points.
Figure 16 shows the variation in ortho and ortho
plus hydrolyzable P2Ogfor three sewage treatment
plant effluents in the Kaskaskia basin. Per capita
values of ortho plus hydrolyzable P2Osin the three
effluents ranged from 0.003 to 0.045pounds per day.
Id
a
ui
h
•z.
u
UI
o:
I
u
H
25
20
15
10
5
Plant A
Population 600
Ortho plus
Hydrolyxabls
0
.-
•
!*
•gs
s:
5
t
• • i
Ortt
io
25
20
15
10
5
1 —
Plant B
Population 1600
0
e
0^1
W
e
v/y
rdf(
.«
-
to
yfa
0
p
•c.
IS8"
' •
7\
Ort
us
»/«]
ho
•
O
•
o
i
o
5 10 30 50 70 9O 95
PER CENT EQUAL TO OR LESS THAN
Figure 16. DOMESTIC P2Os IN 3 KASKASKIA BA-
SIN SEWAGE TREATMENT EFFLUENTS, 1956
Variation of Runoff and Stream Phosphates
Stream samples were collected approximately
once each week at three points in the upper Kas-
kaskia. from April through September, 1956. Sam-
ples were collected at the stations in the lower ba-
sin in April and May of the same year. Figure 17
describes the variations in stream runoff, calcu-
lated pounds per day of P2&5 in the stream runoff,
and concentration at Station 1. This station re-
ceived no domestic P2Os, and received runoff from
a cultivated drainage area of 11 square miles.
Thus, all phosphate found in the stream at this sta-
tion is attributed to land drainage. Concentration
of ortho plus hydrolyzable P2C>5 varied from 0.03
to 0.96 mg/1 in the 27 samples collected. Pounds
per day of ortho plus hydrolyzable P2Os ranged
from 0.1 to 7.5, with an average value of 1.1 pounds
per day. This corresponds to a value of 0.1 pound
of P2C>5 per day per square mile of drainage area.
-------
Land Drainage as a Source of Phosphorus
M
_
u
c
e
I
n
Variation in
weight of PZ09
9
E
0
0.6
0.2
Variation in
concentration of PtOB
APRIL MAY
JUNE JULY AUG.
MONTHS, 1956
SEPT
Figure 17. VARIATION IN RUNOFF AND ORTHO
AND ORTHO PLUS HYDROLYZABLE P2Os. Sta-
tion 1, Kaskaskia River. 11 Sq. miles cultivated.
No domestic
Total phosphate values may be 20 to 30 per cent
higher, because only a portion of the organic phos-
phate is accounted for by the method of analysis
used.
Figures 18, 19 and 20 describe variations in
phosphate and stream runoff for the remainder of
the Kaskaskia sampling stations. Both domestic
and agricultural phosphates are expected to be pre-
sent at these locations. A line corresponding to the
maximum 90 per cent value of domestic phosphate
has been shown on the plots describing variation
in weight of P2^ at each of the stations. It is
apparent that the quantity of stream phosphate at-
tributable to land drainage varies with the stream
runoff. Similar correlations with rainfall have been
observed, as would be expected.
Overall Results of Kaskaskia Survey
For each of the sampling locations on the Kas-
kaskia River, Table 19 presents mean values of
phosphate concentrations and the approximate per
cent of the ortho plus hydrolyzable P£Os in the
samples collected which could be attributed to
land drainage. These per cents are based on as-
sumed average contributions of P2&5 from domes-
tic sources. Orthophosphate concentrations are
seen to average from 0.11 to 0.39 mg/1 P2C>5,
while ortho plus hydrolyzable phosphate averages
range from 0.16 to 0.68 mg/1
it
c
B
C
&
a
I
200
IOO
240
160
D
M
0.
Ortho plus
—Hydro/viable
Variation in
concentration of P209
APRIL MAY
JUNE JULY
MONTHS, 1956
AUG. SEPT
Figure 18. VARIATION IN RUNOFF AND ORTHO
AND ORTHO PLUS HYDROLYZABLE P2Os. Sta-
tion 2, Kaskaskis River. 109 sq. miles cultivated.
No domestic P2O5-
2400
APRIL MAY
JUNE JULY
MONTHS, 1956
AUG. SEPT
Figure 19. VARIATION IN RUNOFF AND ORTHO
AND ORTHO PLUS HYDROLYZABLE P2O5. Sta-
tion 3, Kaskaskia River. 842 Sq. miles cultivated.
Maximum domestic P2Os, 260 Ib/day.
-------
78
ALGAE AND METROPOLITAN WASTES
DISCUSSION
The stream survey results obtained for the Kas-
kaskia River, though subject to considerable varia-
tion, do indicate that land drainage is a significant
source of phosphates in streams which drain agri-
cultural lands. The Kaskaskia basin, as shown in
Table 18, is extensively cultivated. The farm lands
contain in the order of 40 to 50 pounds of available
P2Og per acre, and are drained by tile drains.
High rainfalls and resultant high rates of percola-
tion and runoff might be expected to carry some
portion of both soluble and adsorbed phosphorus
forms to the stream. This expectation is borne
out by the presence of measurable phosphate con-
centration at a stream location receiving no dom-
estic phosphate, and by the presence at other
stream locations of phosphates in excess of those
expected from domestic sources alone.
The amount of agricultural phosphate trans-
ported to streams undoubtedly depends upon a num-
ber of factors: nature and amount of phosphates in
the soil, mode of drainage, topography, intensity
and distribution of rainfall, rates of infiltration
and percolation, and probably others. For the 100
samples reported here, the calculated pounds of
¥2*0$ per day per square mile varied from 0 to 58
for ortho plus hydrolyzable. The mean pounds of
P2O§ per square mile for all samples was 1.4,
which is on the same order of magnitude as that
reported by Sawyer (1944).
Table 19. RESULTS OF KASKASKIA STREAM SURVEY, 1956
Stream
station
1
2
3
4
5
6
Drainage
area,
sq. mi.
12
125
1030
1980
2680
5220
No. of
samples
27
24
25
10
7
7
Mean Ortho
P205, mg/1
0.11
0.29
0.39
0.14
0.13
0.20
Mean Ortho +
hydrolyzable
P205> mg/1
0.16
0.40
0.68
0.34
0.24
0.43
Mean per cent
Ortho + hydrolyzable
P20g, land drainage
100
55
45
25
55
35
REFERENCES
Anon. 1958. Determination of orthophosphate, hy-
drolyzable phosphate, and total phosphate in sur-
face waters. AASGP Committee Report. Jour.
Amer. Water Works Assoc., 50: 1563-1574.
Bennett, H. H. 1939. Soil Conservation. 1st Ed.
McGraw-Hill Book Co., N. Y., pp. 9-10.
Engelbrecht, R. S. and J. J. Morgan. 1959. Stu-
dies on the occurrence and degradation - of con-
densed phosphates in surf ace waters. Sew. and Ind.
Wastes, 31: 458-478.
Kohnke, H. and A. R. Bertrand. 1959. Soil Con-
servation. McGraw-Hill Pub. Co., N. Y., pp. 91-
92.
Kurtz, L.T. 1945. Adsorption and release of phos-
phate ions by soils and clays. Unpublished Ph.D.
Thesis, University of Illinois, Urbana, Illinois.
Melsted, S. W. April, 1960. Personal communi-
cation. Agronomy Department, University of Illi-
nois, Urbana, Illinois.
Sawyer, C.N. 1944. Fertilization of lakes by agri-
cultural and urban drainage. Jour. New England
Wat. Works Assoc., 61: 109-127.
Silvey, J. K. G. 1953. Relation of irrigation to
taste and odor. Jour. Amer. Water Works Assoc.,
45: 1179-1186.
Waggaman, W. H. 1952. Phosphoric acid, phos-
phate, and phosphatic fertilizers. 2nd Ed., Rein-
hold Pub. Corp., N. Y., pp. 20-24.
-------
Land Drainage as a Source of Phosphorus
STATION 4
1500 SQ. MILES CULT.
DOM. P205, 500 LB/DAY
APRIL
MAY
STATION 5
1900 SQ. MILES CULT.
DOM. P205, 700 LB/DAY
Ortho +Hydrolyzabl9
APRIL MAY
MONTHS, 1956
STATION 6
4000 SQ. MILES CULT.
DOM. P2 09,3500 LB/DAY
Domestic P9O*-
-^=±^1-
APRIL
MAY
Figure 20. VARIATION IN RUNOFF AND ORTHO AND ORTHO PLUS HYDROLYZABLE P0Oc, KASKASKIA
RIVER.
-------
80
ALGAE AMD METROPOLITAN WASTES
NUTRIENT CONTENT OF DRAINAGE WATER FROM FORESTED, URBAN AND AGRICULTURAL AREAS
ROBERT O. SYLVESTER, Professor of Sanitary Engineering
Department of Civil Engineering, University of Washington, Seattle
SYNOPSIS
Nutrient concentrations are presented from field
observations on drainage waters originating in
timbered areas of little habitation and land use;
from urban street drainage; sub-surface and sur-
face drains carrying irrigation return flows; urban
streams; a eutrophic lake, and a comparison is
made of the nutrient change in a stream of multi-
purpose usage. The effect of land and water use on
nutrient concentrations is clearly shown. Nutrient
loss through sedimentation and incorporation in the
bottom sediments and loss to littoral and other
attached vegetation is indicated.
INTRODUCTION
Nutrient data in the form of soluble and total
phosphorus, nitrates and total Kjeldahl nitrogen,
were obtained during field studies of a eutrophic
lake and its environment (Sylvester and Anderson,
1960) and from a study1 now in progress on the
character and significance of irrigation return
flows in a large river basin. Nitrite values were
found to be very low and are not included for the
sake of brevity. Ammonia is included in the total
Kjeldahl nitrogen.
Natural drainage or runoff waters contain vary-
ing quantities of nutrients (fertilizers) depending
upon their source. These natural nutrient concen-
trations become altered through man's multi-
purpose land andwater usage. This alteration nor-
mally consists of an increase in the phosphorus'
and nitrogen content of the drainage water. An
increase in the nutrient content of a water, along
with a sufficiency of other elements, results in an
increased plant productivity in the water that may
reach such proportions that normal water uses are
harmed or jeopardized. This plant productivity
may be in the form of algae; emergent or non-
emergent sessile plants, such as willows, cattails,
water weeds; or it might consist of slime growths,
such as the extensive Sphaerotilus beds of the lower
Columbia River (Anon.) that have been attributed to
unnatural increases in several nutrients, among
which are wood sugars. An increase in the pri-
mary producers, or phytoplankton, will usually re-
sult in an increase in the zooplankton and thus a
better habitat is provided for fish production.
When this algae production exceeds the fish food
needs, the water may become undesirable as a
source of water supply or as a medium for recrea-
1. "Character and Significance of Irrigation Return Flows"
of Washington, study in progress.
tion. The stimulated growth of water weeds and
other vegetation increases the cost or difficulty of
ditch maintenance, shoreline maintenance and of
insect control.
Nutrient concentrations necessary to produce
nuisance water growths will vary, depending upon
the concentrations of other substances, the water
temperature, light penetration in the water, and
the rate at which nutrients can be supplied, once
growth commences. The temperature of and the
light penetration in the aquatic environment are
normally subject to change only from natural phe-
nomena (with a few exceptions), leaving nutrients
as that requirement for plant growth that has been
altered by man. Many waters have become highly
productive of luxuriant plant growths in the ab-
sence of man's machinations and many others
would naturally reach this stage in the course of
time (witness the many eutrophic lakes). Man's
water and land uses have usually served to hasten
this process of natural maturing by his addition of
nutrients to the water through land cultivation and
fertilization and through the discharge of his spent
or waste waters into the natural water bodies of his
environment. Sawyer, in his study of Wisconsin
lakes (1947) reports that large phytoplankton
blooms maybe supported when phosphorus concen-
trations are in or above the range of 10 ppb PO4 P
and nitrogen concentrations are above 300 ppb
NO, N. Hutchinson (1957) reports mean nitrate
nitrogen values of 50 to 86 ppb in three Wisconsin
lakes that produce regular nuisance blooms of algae.
Some lakes have high nitrogen values but no regu-
lar nuisance blooms because of a deficiency in
some other factor. Nuisance algal blooms were
observed (Sylvester and Anderson, 1960) to com-
mence in Seattle's Green Lake (a very soft-water
lake) when nitrate nitrogen levels were generally
above 200 ppb and soluble phosphorus was greater
than 10 ppb. These blooms usually stopped when
the soluble phosphorus dropped to zero. On one
occasion, the bloom was attenuated when the ni-
trate concentration was depleted and the soluble
phosphorus was still above 10 ppb.
DATA PRESENTED
The information presented herein consists of
nutrient data (summarized) that were obtained in-
cidental to comprehensive water quality studies.
Data are generally presented by showing the range
of values observed together with a weighted-aver-
N. I. H. Research Grant (RG6412C1) to University
-------
Nutrient Content of Drainage
81
age value. Whenever possible, other factors are
given, such as nutrient ratios, rate of water flow,
soil types and concentration of nutrients discharged
per acre. Weighted-average values (to account
partially for flow variations) are obtained by multi-
plying each individual determination by the flow
rate at the time of the determination, summing
these individual products and then dividing by the
summation of the flow rates. Table 26 gives mean
rather than weighted average values since flow
rates were not available. In most cases, the mean
value was reasonably close to the median value.
Phosphorus concentrations are reported as the
element phosphorus, and nitrogen as the element
nitrogen. Soluble phosphorus was determined by
the stannous chloride method (Anon, 1955) and to-
tal phosphorus by evaporating and igniting a sam-
ple containing magnesium chloride followed by the
stannous chloride method. Nitrate nitrogen was
determined by the phenoldisulfuric acid method
(Anon, 1955). Samples were collected at frequen-
cies of from once a week to about once a month.
Urban Street Drainage
Table 20 shows the nutrient content in street
gutter drainage water collected from major high-
ways, arterial and residential streets, under
varying conditions of antecedent rainfall. Samples
were dipped from the gutters anywhere from 30
Table 20. NUTRIENTS IN URBAN STREET
DRAINAGE1, MAY - NOVEMBER, 19592
Type of
street
Arterial St.
Arterial st.
Arterial St.
Residential St.
Major highway
Major highway
Major highway
Residential st.
Residential st.
Residential st.
Residential st.
Arterial St.
Arterial st.
Arterial st.
Arterial st.
Residential St.
Major highway
Major highway
Major highway
Major highway
Arterial St.
Arterial st.
Arterial st.
Arterial st.
Arterial St.
Arterial st.
Arterial St.
Arterial st.
Major highway
Major highway
Major highway
Residential st.
Residential St.
Arterial st.
Arterial st.
Median
Mean
Antecedent
rainfall,3
inches
0.21
0.21
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.78
0.78
0.78
0.78
0.78
0.78
0.0
0.0
0.0
0.0
0.0
0.0
0.22
0.22
0.22
0.22
0.0
0.0
0.55
0.55
0.55
0.55
0.55
0.55
0.07
0.07
Nitrogen - rog/1 N
Total
Kjeldahl
—
—
0.44
1.01
1.4
0.4
0.45
0.37
0.32
2.78
1.31
1.34
0.57
1.43
1.84
6.68
9.06
7.45
5.36
7.21
8.01
0.38
0.60
0.41
0.88
1.67
1.54
0.22
0.62
0.37
0.91
0.39
0.39
0.17
0.24
0.41
2.01
Nitrates
0.185
0.10
0.56
0.16
0.86
0.52
0.22
0.17
0.17
1.10
0.52
0.29
0.23
0.29
0.23
0.65
2.24
2.80
0.96
0.53
0.52
0.65
1.00
1.00
0.80
0.48
0.36
0.12
0.03
0.03
0.03
0.02
0.10
0.42
0.11
0.42
0.53
Phosphorus as P, ppb
Soluble
—
—
—
—
—
—
—
—
—
Tr
16
Tr
2
20
Tr
14
54
72
30
28
14
22
32
156
784
280
154
20
10
10
8
70
78
58
48
22
76
Total
10
49
104
328
—
228
—
228
440
154
308
144
90
108
126
166
352
404
102
162
81
146
198
300
1400
280
182
21
13
14
21
98
78
212
280
154
208
minutes to several hours after a rainstorm had
commenced and they were dipped so as not to in-
clude particulate matter that would be retained by
a catch basin. Since most of the area sampled was
served by combined sewers, the gutter water sam-
ples did not generally include roof drainage, but
did include all types of surface drainage. As one
would expect, low antecedent rainfall conditions
gave the highest nutrient concentrations. Since
these samples were not all collected at the very
beginning of rainstorms nor at the end of prolonged
rainstorms, they can be considered quite repre-
sentative of urban surface runoff under varying
conditions of street sweeping and flushing, ante-
cedent rainfall, and the presence and absence of
deciduous vegetation. The major highway (not a
limited access freeway) had the greatest nitrogen
values; the arterial streets contained the most
soluble phosphorus; and the residential streets the
highest total phosphorus concentrations.
Forested Areas
Nutrient values in three streams as they emerge
from forested areas are given in Table 21. Each
Table 21. NUTRIENTS IN STREAMS FROM
FORESTED AREAS1
Characteristics
Mean annual flow, els
Mean flow for data period
Drainage area, sq. mi.
Phosphorus as P, ppb
Total
Range of values
Weighted average
Soluble
Range of values
Weighted average
Nitrates as N, mg/ 1
Range of values
Weighted average
Total Kjeldahl N, mg, 1
Range of values
Weighted average
Total nitrogen as N. mg/ 1
Total phosphorus, Ibs.. acre/ year
Total nitrogen, Ibs./ acre/ year
Total N
R»UO Total P
Yakima River
at Easton
(12 months)
587
630
182
19-140
10
0-23
9
0.05-0.50
0.20
0-0.22
0.08
0.28
0.74
2.96
4
Tleton River
(7 months)
559
520
239
32-200
115
0-23
8
0.03-0.18
0.126
0-0.13
0.068
0.194
0.77
1.30
1.7
Cedar River2
at Landsberg
(12 months)
690
587
125
15-85
22
2-7
4
0.018-0.154
0.065
__
--
0.32
-
-
1. Samples collected from gutters on Seattle streets.
t. From Sylvester and Anerson (1960).
3. Antecedent rainfall in inches during week prior to sample collection.
1. Areas subject only to logging and road construction - large reservoirs
present on all headwaters.
2. Seattle Engineering Department data.
watershed contains large reservoirs, roads and
some logging but no human habitation that would
contribute any significant amount of waste water to
the streams. The Cedar River originates on the
western slope of the Cascade mountains, is the
source of Seattle's water supptyand it is generally
a less turbid river than the other two. The Yakima
River at Easton is the point of diversion for the
70,000 acre Kittitas irrigation project while the
Tieton River at the point of sampling is just below
the diversion structure for the 28,000 acre Tieton
project (see figure 21). Discharge of phosphorus
and nitrogen from the eastern slope streams is
over twice as great as that from the western slope
streams.
-------
82
ALGAE AND METROPOLITAN WASTES
•KITTITAS
PROJECT
Volume Count
Kile* if Of Counfy
SUNNYSIDE PROJECT
I
KENNEMCK PROJECT-
ScoH in Miles
s o s 10 ts
Figure 21. YAKIMA RIVER BASIN. 1. Yakima River at Easton; 2. Yakima River at Parker; 3. Yakima
River at Kiona; 4. Tieton River; 5. Wilson Creek; 6. Granger Drain; 7. Main Reservation Drain; 8. Sunny-
side 35.4 Drain; 9. D-2 Sub-surface Drain; 10. D-4 Sub-surface Drain; 11. D-5 Sub-surface Drain; 12. Sub-
surface Drain; 13. D-14 Sub-surface Drain.
Irrigation Return Flow Drains
About 450,000 acres of land are irrigated in the
arid Yakima River Basin. Water application
ranges from 2.5 to 11 acre-feet per acre per year,
averaging around 3.7. From 20 to 75 percent of
this irrigation water is returned to the main river
each year as return flow. Irrigated land must be
well drained to prevent an accumulation of salts in
the surface portion of the soil. A favorable salt
balance is attained when the drainage water con-
tains a higher salt load than the applied water.
Return flow drains may consist of ditches (sur-
face) or buried pipe that is porous or has open
joints (sub-surface). The water in these return
flow drains is highly mineralized. For example,
the Yakima River in receiving the return flow from
-------
Nutrient Content of Drainage
83
some 450,000 irrigated acres between Easton and
Kiona (see figure 21) has its specific conductance
increased from 30 to 240 micromhos during the
height of the irrigation season.
Tables 22 and 23 give the nutrient variations in
typical surface and sub-surface drains and table
24 shows the difference in the soil characteristics.
The study from which tables 22 and 23 were pre-
pared A/ is still progressing and it is too early
to definitely speculate on what causes or does not
cause the differences in the nutrient content of the
drain water. Variables herein involved include the
soil types (surface and sub-surface); quality and
quantity of water applied to the land; presence of
irrigation waste (excess) water; nature of crops
being grown; fertilizers applied; and the depth of
the drain and water table. The principal difference
between the nutrient content in the surface and sub-
surface drains is the higher mineralized nutrient
content of the sub-surface drains. Nitrates are
twice as high in the sub-surface drains and the N/P
ratio of the sub-surface drains averages 2.5 times
as high as the surface drains.
In the surface drains, the total phosphorus con-
tained in the drainage water varied from 0.9 to 3.9
Ibs./acre/year while the total nitrogen varied from
2.5 to 24 Ibs./acre/year. Sub-surface drains re-
moved total phosphorus at the rate of 2.5 to 8.9
Ibs./acre/year and total nitrogen at the rate of 38
to 166 Ibs./acre/year. Sawyer has reported (1947)
Wisconsin agricultural lands as discharging 0.398
Ibs. of phosphorus and 7.03 Ibs. of nitrogen per
acre per year. With the limited data now available
on fertilizers applied to the land (Blake, 1960), it
would appear that a considerable portion of them
are being carried off in the drainage water.
Multi-Purpose Water Usage
The Yakima River below Easton receives the
treated sewage discharge from some 80,000 per-
sons, irrigation return drainage from 450,000
acres and industrial waste effluents having a pop-
ulation equivalent of over 100,000 (Sylvester,
Weston, Suzuki and Dailey, 1951; and Anon., 1952).
The drainage area below the mountains is quite
arid (average rainfall is 7 inches) so that most of
the runoff in the area comes from the irrigated
land. Changes in nutrient concentrations in the
river between Parker and Kiona (see figure 21),
where there are no significant year-around tri-
butaries other than spent waters, are given in
Table 25. Between Parker and Kiona, a distance of
72 river miles, the tributary irrigated acreage is
250,000 acres and the sewered population plus
industrial equivalent is about 93,000 persons. The
Table 22. NUTRIENTS IN SURFACE IRRIGATION RETURN FLOW DRAINS1 YAKIMA VALLEY, MARCH
1959 - MARCH 1960
Characteristics
Approx. drainage area, sq. mi.
Approx. mean flow, cfs
Predominate soil type
Total phosphorus, P ppb
Range of values
Weighted average
Soluble phosphorus, P ppb
Range of values
Weighted average
Nitrates as N, tag/I
Range of values
Weighted average
Total KJeldahl N, mg/1
Range of values
Weighted average
Total Nitrogen as N, mg/1
Total phosphorus, Ibs./acre/year
Total Nitrogen, Ibs./acre/year
Ratio N/P
Main drain
Wapato Project
150
379
Naches soils
105-240
165
75-255
127
0.03-4.2
1.19
0.05-0.47
0.15
1.34
1.28
10.45
8.1
Granger drain
Sunnyside Project
13
47
Sagemoor
sandy loam
120-380
260
88-300
210
0-5.70
1.54
0-0.34
0.13
1.67
2.88
•18.5
6.4
Drain 35.4
Sunnyside Project
2
7
Esquatzel fine
sandy loam
130-650
360
115-295
180
0-5.0
1.90
0.08-1.30
0.33
2.23
3.88
24.0
6.2
Wilson Creek
Kittitas Project
252
340
Naches fine
sandy loam
135-300
220
72-192
130
0-1.40
0.38
0.02-0.48
0.21
0.59
0.92
2.45
2.7
1. All drain areas of diversified farming except for Wilson Creek which drains areas of hay, corn and grains.
-------
ALGAE AND METROPOLITAN WASTES
Table 23, NUTRIENTS IN SUB-SURFACE IRRIGATION RETURN FLOW DRAINS YAKIMA VALLEY, MARCH
1959 - MARCH 1960
Characteristics
Soil type
Avg. discharge, gpm
Crops growu
Fertilizers used
Approx- water applied-- Ac. Ft./ Yr.
Type irrigation
Approx. area drained, acres
Drain
Length - Feet
DUmeter - inches
Depth -Ft.
Age
Total Phosphorus, P, ppb
Range of values
Weighted average
Soluble Phosphorus
Range of values
Weighted average
Nitrate* as N, mg/1
Range of values
Weighted average
Total Kjeidahl N, mg/1
Range of values
Weighted average
Total Nitrogen, N, mg/1
Total Phosphorus, Ibs./acre/year
Total Nitrogen, tos./acre/year
Ratio N/P
D-2
Klttitas Project
Nacbes loam
67
Sweet corn
Commercial
4.1
Furrow
15
960
8
3-7
1.5
73-276
173
29-156
89
0.40-5.0
2.73
0-0.58
0.15
2.88
3.4
56
16.7
D-4
Kittttas Project
Esquatzel and
Selah loam
157
Wheat
Barnyard t
commercial
5
Furrow
27
2400
8-10
5-8
2
195-460
320
120-460
320
0.42-4.1
8.03
0-0.32
0.09
2.12
8.1
54
6.7
D-5
Tieton Project
Ahtanum loam
275
Barley b pasture
Gypsum & commer,
3
Furrow
35
2500
10-15
4-10
1
175-465
260
175-350
260
0.20-6.5
2.55
0-0.32
0.13
2.68
8.9
92
10.3
D-12
Sunnyside Project
Sagemoor loam
830
Hops, grapes,
wheat, hay, mint
All types
4.3
Furrow
100
8000
10
4-10
43
75-405
195
49-170
150
0.10-9.0
4.37
0-1.86
0.18
4.55
7.1
166
23
D-14
Tieton Project
(8 months)
Varied
42
Apples
—
—
Sprinkler
10
400
8
4
2
82-173
133
82-127
103
0.01-5.9
1.77
0.08-0.83
0.31
2.08
2.5
38
15.6
Table 24. SOIL CHARACTERISTICS (from Anon., 1958; and Smith, Dwyer and Schafer, 1945)
Soil type
Ahtanuin loam
Esquatzel silt loam
Naches loam
Sagemoor loam
Selah loam
Surface soil
Pale-brown; strongly
alkaline; calcareous
Brownish -gray, neutral
to mildly alkaline; non-
calcareous
Brown; neutral; non-
calcareous
Light brownish-gray,
netnral to mildly
alkaline; non-calcareous
Light brown; neutral
to mildly alkaline;
non-calcareous
Subsoil
Greyish-brown loam
over time hardpan;
strongly alkaline;
calcareous
Pale-brown; mildly
alkaline; calcareous
in lower part
Brownish gravelly
clay; neutral; non-
. calcareous
Light brown to gray
sandy loam; alkaline;
strongly calcareous
Brown, compact, sandy
clay; lime-silica
hardpan in lower part
alkaline; calcareous
Inherent fertility
Very low
Moderate to high
Moderate
Moderate
Moderate
Internal drainage
Medium to very stow
Medium
Medium to high
Medium
Very slow
-------
Nutrient Content of Drainage
85
Table 25. CHANGE IN RIVER NUTRIENTS FROM
VARIOUS SPENT WATERS YAKIMA RIVER,
WASHINGTON1 — MARCH 1959 - MARCH 1960
Characteristics
Mean annual flow - cfs
Mean flow for data period - cf a
Drainage area - sq. ml.
Runoff drainage area - sq. ml.
Irrigated land tributary - acres
Sewered population tributary - persons
Industrial waste tributary - pop. equlv. '
Industrial waste flow -mgd3
River miles from headwaters
Total Phosphorus as P, ppb
Range of values
Weighted average
Soluble Phosphorus as P, ppb
Range of values
Weighted average
Nitrates as N, mg/1
Range of values
Weighted average
Total Kjeldahl Nitrogen as N, mg/1
Range of values
Weighted average
Total Nitrogen as N, mg/1
Phosphorus - Ibs/acre/year*
Nitrogen - Ibs./acre/year*
Phosphorus - tons discharged/year
Nitrogen - tons discharged/year
Ratio N/p
Yakima
River at
Parker
2,529
3,180
3,650
2,290
200,000
59,260
39,160
4.1
110
49-210
70
29-75
43
0.04-0.45
0.25
0-0.23
0.12
0.37
0.30
1.575
219
1,150
5.3
Yakima
River at
Kiona
3,916
4,550
5,600
2,930
450,000
80, 340
111,510
6.0
182
55-240
135
23-94
51
0-0.78
0.32
0-0.73
0.175
0.495
0.64
2.35
602
2,210
3.7
1. 85% of drainage basin is relatively uninhabited—1,064,100 acre-
feet of storage on headwaters.
2, Below the mountains, the land Is arid and runoff comes
principally from the irrigated areas.
3. Food processing wastes are mostly seasonal.
4. Using "runoff drainage area".
river flow at Parker frequently becomes very low
during the irrigation season (less than 150 cfs)
while at downstream Kiona, the flow will be over
1700 cfs, this additional flow coming from irriga-
tion drains. Because of these spent waters, one
would expect the river to show a very large in-
crease in nutrients between Parker and Kiona with
nutrient values that would approach the levels of
those observed in the drains. Although all nutrient
values did increase in this river stretch, only the
Kjeldahl nitrogen values approached the concentra-
tion found in the drains. The readily assimilable
phosphates and nitrates showed little increase indi-
cating their uptake in the stream plant life. This
plant life is abundant and is sufficient to frequently
cause dissolved oxygen saturation values of 120
percent in the daytime and 60 percent just before
sunrise (Sylvester, 1958). River turbidity has a
median value of about 9 units. Much of this plant
growth is flushed from the stream bed during the
autumn and spring freshets. The nitrogen-phos-
phorus ratio dropped in this stream stretch indi-
cating a proportionately greater uptake of nitrogen.
Eutrophic Lakes
Table 26 gives some mean nutrient values for
surface waters in the City of Seattle. Green Lake
has regular nuisance blooms of algae in the spring,
summer and early autumn. This lake had reached
a trophic state long before it was affected by man.
The Lower Woodland and Densmore drains shown
in table 26 are typical of those feeding Green Lake.
Nutrient concentrations in the lake water are much
less than in the source water. Calculation of a nu-
trient budget (Sylvester and Anderson, 1960) indi-
cated that 55 percent of the phosphorus was lost to
the sediment (permanently), through deposition of
algae and particulate matter, and through the thou-
sands of fish taken from the lake by fishermen. 2/
Phosphate or nitrate values will reach zero during
periods of heavy algal blooms. The algae will then
die and promptly settle to the lake bottom. The in-
flow of nutrient-rich water and regeneration of nu-
trients from the mud permit another algal pulse
within a short period.
Thornton Creek drains into Lake Washington.
This nutrient-rich stream is typical of many tri-
butaries to the Lake that are, along with sewage
inflows, cause for concern as to the future trophic
state of the Lake.
SUMMARY
Nutrient concentrations from various water
sources have been presented. A summary of these
values is given in Table 27 with a visual compari-
son presented in Figure 22. Although "clean"
streams from forested areas contain the lowest
assimilable nutrients, their concentrations are
close to the mean concentrations in a eutrophic
lake and it would appear from Table 27 that an in-
crease in phosphate (soluble phosphorus) is all
that would be needed to create a productive stream
out of the "clean" stream.
The greatest concentrations of total phosphorus
were found in surface irrigation return flow drains;
of soluble phosphorus and nitrates, in sub-surface
irrigation return flow drains; and of total Kjeldahl
nitrogen, in urban street drainage.
2/lt is estimated that approximately 100,000 fish, averaging about 0.3 Ib. each, were taken from Green Lake
in 1959 by fishermen. According to Professor Lauren Donaldson, these fish would contain about 292 mg of
phosphorus per 100 grams of fish. The phosphorus thus removed from the lake in the form of fish flesh might
amount to 88 Ibs. per year which is but 14 percent of the mean phosphorus content of the lake water.
-------
86
ALGAE AND METROPOLITAN WASTES
ACKNOWLEDGMENTS
The study of Green Lake was supported by the
City of Seattle and the irrigation water study is be-
ing supported by research grants from the National
Institutes of Health. Grateful acknowledgment is
given George C. Anderson, Research Assistant Pro-
fessor of Oceanography, and Robert W. Seabloom,
Assistant Professor of General Engineering, both
of the University of Washington, for their assistance
in data collection and analysis and for their review
of this manuscript.
Table 26.
1960)
NUTRIENTS IN MISCELLANEOUS WATERS, SEATTLE URBAN (from Sylvester and Anderson,
Location
Green Lake
Lower Woodland Drain
Densmore Storm Drain3
Thornton Creek4
Nitrates - mg/ 1 N
Range of
values
0-0.47
0.20-0.85
0.53-2.02
1.26-1.70
Mean
0.084
0.46
1.24
1.48
Total Kjeldahl N
mg/1 N
Range of
values
0.02-1.0
0.01-0.88
0.10-1.34
Mean
0.34
0.32
0.57
Soluble phosphorus
ppb
Range of
values
0-58
23-112
58-128
45-85
Mean
16
75
85
66
Total phosphorus
PPb
Range of
values
38-178
49-292
69-300
74-213
Mean
76
103
136
110
1. 256 acre lake in central Seattle receiving nutrient rich drainage - samples April to January, 1959.
2. Subsurface drain in park; discharging to Green Lake - some surface water - samples April to Jan., 1959.
3. Small creek in urban area with duck ponds.
4. Creek in urban Seattle area with mean annual flow of about 23 cfs draining 13.5 sq. mi. (Data from
Seattle Engineering Department).
Figure 22. SOURCE COMPARISON OF MEAN NUTRIENT CONCENTRATIONS
It
I).1«.M II
-------
Nutrient Content of Drainage
87
Table 27. SOURCE COMPARISON OF MEAN NUTRIENT CONCENTRATIONS (See Fig. 22)
Source
Streams from forested areas -
little habitation or land use.
Sub-surface irrigation
retrun flow drains
Surface irrigation return
flow drains
Urban street drainage
Urban street drainage
Multiple - use river*
2
Urban streams
Eutrophic lake3
Total
phosphorus
ppbP
69
216
251
208
154*
135
123
76
Soluble
phosphorus
ppbP
7
184
162
76
22*
51
76
16
Nitrates
ppbN
130
2690
1250
527
420*
320
1360
84
Total Kjeldahl
Nitrogen
ppbN
74
172
205
2010
410*
175
570
340
1. Yakima River at Kiona
2. From table 26
3. Green Lake, Seattle
* Median values
REFERENCES
Sylvester, R. O. and G. C. Anderson. 1960. An
engineering and ecological study of Green Lake .
Seattle Park Department (In press), February.
Anon. Columbia River study, 1956-1958. Progress
Report by the Washington Pollution Control Com-
mission and Crown Zellerbach Corp.
Sawyer, C. N. 1947. Fertilization of lakes by
agricultural and urban drainage. J. N. E.W.W. A.
61: 2, June.
Hutchinson, G. E. 1957. A Treatise on Limnology.
Vol. I, New York Wiley. 1015 pp.
Anon., 1955. Standard methods for the examina-
tion of water, sewage and industrial wastes. A.P.
H.A., 10th Edition.
Anon., 1958. Soil survey, Yakima County, Wash-
intion, U.S.D.A., Soil Conservation Service Series
1942, No. 15, Issued April 1958.
Smith, L. H., C. H. Dwyer, and G. Schafer, 1945.
Soil survey, Kittitas County, Washington, U.S.G.
P.O., Series 1937, No. 13, Issued January.
Blake, Larry J. 1960. Water quality studies of
irrigation sub-surface return flow sand influencing
factors. M.S. Thesis, University of Washington.
Sylvester, R. O., M. J. Westin, J. T. Suzuki, and
M. G. Dailey, 1951. An investigation of pollution
in the Yakima River Basin. Washington Pollution
Commission Technical Bulletin No. 9.
Anon., 1952. A comprehensive pollution control
program for the Yakima River Basin, Washington
Pollution Control Commission Bulletin, March.
Sylvester, R. O. 1958. Water quality studies in
the Columbia River Basin. U.S.F. & W. S. Special
Scientific Report - Fisheries No. 239.
-------
Wastes
METROPOLITAN WASTES AND ALGAL NUTRITION
WILLIAM J. OSWALD, Associate Professor
Sanitary Engineering and Public Health, University of California, Berkeley
INTRODUCTION
Organic wastes rank high among the many sour-
ces of nutrients which increase the fertility of na-
tural waters. Computations indicate that in the
United States alone approximately 20 million tons
of agricultural, food processing, domestic, and
industrial wastes must annually be deposited di-
rectly or indirectly into natural waters. The quan-
tity of these wastes is increasing not only in direct
proportion to the population increase of our coun-
try, but also in proportion to the current tendency
for people to spend a greater percentage of their
recreational time near, on, or in the water. Algal
growths in natural waters maybe either heterotro-
phic, autotrophic, or a combination of the two.
Heterotrophic algal growth is not considered in
this paper except to note that such growth in com-
petition with bacteria must be small in magnitude
when comparedwith autotrophic algal growth which
is benefited by bacterial activity. As wastes added
to water undergo bacterial decomposition, carbon
dioxide, ammonia, phosphate, sulfate, nitrate and
other elementary substances are liberated and be-
come available to the autotrophic algae, which in
the presence of adequate light and suitable tempera-
tures, tend to grow in direct proportion to concen-
tration of added decomposable nutrients.
Autotrophic algal growth, which occurs as a
result of addition of organic wastes to natural
waters, is of interest not only because it may lead
directly or indirectly to gross nuisances in lakes,
streams, or reservoirs, but also because it may
be.employed in controlled processes to produce
oxygen and eliminate odors in stabilization ponds,
to remove the bulk of nutrients from wastes prior
to dilution in receiving waters thus controlling un-
wanted algal growth, and under special circum-
stances to accomplish both nutrient recovery and
water reclamation (Oswald, Golueke and Gee,
1959). While each of these applications is of suffi-
cient importance to be considered individually,
they involve the same basic questions, namely; to
what extent will a given organic waste support
autotrophic algal growth, and what factors govern
the magnitude of algal growth that will occur in a
given waste? An approach to the answers to these
basic questions will be the subject matter of this
paper.
Autotrophic growth in a given waste and body of
water is a function of temperature, light and nutri-
ent. Although taste and odor nuisance may occur
at low temperatures, algal blooms reach nuisance
growth proportions mainly when nutrients have
accumulated and water temperature is between 20
and 30° C. Light has an important influence on
autotrophic algal growth because it is the energy
source for such growth. During periods of optimum
temperature the amount of light per unit of area is
rarely ample so that nutrient seldom becomes a
predominantly important factor in determining the
instantaneous magnitude of algal growths in natural
waters. However, if light is abundant on a sus-
tained basis, repetitive crops of algae may occur
until some nutrient becomes exhausted. Thus it
is important to know the nutrient limiting magni-
tude of algal growth in a given water.
Control of nutrients entering natural waters may
have considerable impact on the problem of nui-
sance blooms. Such control may stem from ex-
cluding wastes from particular waters, from spe-
cialized processing of wastes to remove bloom-
producing nutrients before they enter natural
waters, or from the use of compounds which se-
quester critical nutrients in an unavailable form.
Thus to accomplish control we become concerned
with the nature of bloom-producing wastes and
their nutrient composition. One such bloom-
producing waste is domestic sewage. From the
standpoint of its quality, domestic sewage has been
known for 60 years or more to encourage algal
growth, since it was incorporated into media used
by many of the pioneers in algal culture. The ele-
mentary chemical composition of an average do-
mestic sewage is shown in Table 28. The elemen-
tary analysis of an artificially constituted algal
nutrient is also shown. This artificial nutrient was
used by Fisher (1953) for pilot plant experiments
on Chlorella cultures also shown in Table 28. It
is evident from a comparison of the data for the
two types of nutrient that although domestic sewage
contains all of the essential macro nutrients for
Chlorella growth, these are much more dilute than
in the pilot plant nutrient, and are present in dif-
ferent forms and ratios. On the basis of macroele-
88
-------
Metropolitan Wastes and Algal Nutrition
89
Table 28 - COMPARISON OF SEWAGE AND AN
INORGANIC MEDIUM FOR ALGAL GROWTH
Concentration, ppm
Item
Sewage
Medium
Tbtal Solids
Volatile
Ash
Total N
Organic N
Ammonia N
Nitrate N
Nitrite N
Phosphorus
Sulphur
Potassium
Magnesium
Calcium
Sodium
Iron
Alkalinity
BOD
PH
574.0
247.0
327.0
61.3
26.6
33.6
1.4
0.0
10.7
9.5
13.0
18.0
6.0
72.0
0.4
240.0
168.0
9.3
__
—
—
350.00
Nil
0.05
350.00
—
296.00
322.00
1330.00
246.00
15.00
5.50
0.15
—
—
6.00
mentary composition in comparison with pilot plant
media, one might conclude that domestic sewage
is a poor nutrient for Chlorella. However, out-of-
doors and due mainly to light limitations, domestic
sewage will produce a standing crop of Chlorella
practically equal to that in pilot plant media.
In the work reported here there has been no
effort to evaluate the nutrient value of the various
wastes on the basis of elementary composition.
Rather, we have sought to establish quantitative
relationships between waste concentration as mea-
sured by total solids and B.O.D., and algal growth
employing the waste, diluting water, bacteria and
algae in simple bioassays in which no factor other
than the nutrient species relationship was limiting
to algal growth.
In the assays we have sought to measure the po-
tential of wastes to produce algal growth in excess
of that which occurs in dilution waters without
wastes added. We have also sought to determine
the nutrient-limiting magnitude of such growth.
On a short-time practical basis the magnitude of
a bloom may not be the most significant factor
since nuisance blooms are not always large, but
on a long-time basis eutrophication is a function of
bloom magnitude. Another feature of the assays
reported here is interpretation of the data in terms
of B.O.D. and oxygen production. Theoretically,
a waste which will support a growth of algae of
such magnitude that the oxygen produced during
algal growth exceeds the B.O.D. of the waste, can
be considered to be "treatable"; that is, it should
be possible to remove the fertility compounds from
the waste by means of intensive algal culture.
MATERIALS AND METHODS
Selection of Wastes
The wastes used in the assays were selected on
the basis of convenience as well as their signifi-
cance to receiving streams. The list of wastes
assayed is certainly far from complete and in-
cluded milk, abattoir, reduction plant, packing
house, cattle-holding pen, chicken pen, ground
garbage, vegetable cannery, winery, tomato can-
nery, masonite, monosodium glutamate, glutamic
acid, domestic sewage, and high rate domestic
sewage pond supernatant. Glutamic acid, domestic
sewage supernatant, domestic sewage and ground
garbage were included in the assays because these
have been subjected to analyses by ourselves and
other workers.
Dilution Water
The concentration of certain essentials to algal
growth, such as phosphorus, potassium, calcium,
magnesium and trace elements is influenced or
sometimes determined by the composition of dilu-
tion waters. Thus, for a valid assay of a waste it
was considered important that the potential receiv-
ing water be incubated with the waste in determin-
ing maximum probable algal growth.
Normally the concentrated waste was diluted
with the type of water used for dilution in the re-
ceiving stream and endogenous bacteria comprised
the bacterial seed. At times, sub-dilutions were
made with distilled water or with water of special
interest and sewage seed was introduced sepa-
rately.
Selection of Algal Species
Since our procedure called for determination
of the maximum algal crop which could be grown
in a given waste and at a given dilution, it was im-
portant to minimize the loss of algal nutrients,
such as carbon dioxide and ammonia which some-
times escape during the early stages of decomposi-
tion. This was done by selecting algae which would
grow well in actively decomposing wastes. E. gra-
cilis was used in the early experiments, but a
comparison of growth of 12. gracilis with that of C^.
pyrenoidosa showed that the latter attained a higher
growth rate than the former. Since it will grow so
rapidly, the extent of the growth of Chlorella in a
given waste indicates more accurately than does
the growth of Euglena the maximum standing crop
to be attained in a water receiving the waste. Con-
sequently, Chlorella was the alga used in most of
the succeeding studies.
Methods of Evaluating Growth
Algal growth was evaluated through cell counts
used as a base for estimates of culture weight as
a function of dilution of wastes. Because of the
large volume of material normally required, de-
terminations of culture weight, volume and light
penetration, though commonly used to determine
growth, were not used for these small cultures.
-------
90
ALGAE AND METROPOLITAN WASTES
la these studies, cell counts were taken daily and
the weight of Euglena or of Cfalorella cells was es-
timated from earlier studies correlating cell count
with cell weight for culture of various ages.
Figure 23 shows the culture weight in micro-
4 68
CULTURE AGE, DAYS
Figure 23. WEIGHTS OF ONE MILLION CHLO-
RELLAOR EUGLENA CELLS ASA FUNCTION OF
CELL AGE; WEIGHT OF OXYGEN EVOLVED BY
GROWING ONE MILLION CELLS OF CHLORELLA
OR EUGLENA.
grams per million Euglena and Chlorella cells as
a function of cell age as determined from large
continuous culture in which data on cell weight and
cell age were accurately obtained (Ludwig, Oswald,
Gotaas and Lynch, 1950). Also shown is the equi-
valent weight of oxygen determined by multiplying
the weight of algae by 1.6, the ratio of oxygen pro-
duced to algal cell material synthesized, a factor
also determined in continuous cultures (Oswald,
Ludwig, Gotaas and Lynch, 1953). The cell count
in millions of cells per liter was multiplied by the
cell weight in milligrams per million cells obtained
from Figure 23. This gave cell weight in milli-
grams per liter. This weight was then multiplied
by 1.6 to estimate oxygen production.
Cell counts are quickly made, require little ex-
perimental material, and are not subject to inter-
ference by the presence of extraneous materials
which do interfere seriously with gravimetric,
volumetric, and spectrophotometric determina-
tions. A disadvantage of the cell count method is
the relatively low reproducibility of algal counts
and the uncertainty regarding the weight of algal
cells in various cultures. Enumeration with a hae-
mocytometer of a large number of replicate algal
samples gave information on the accuracy of our
algal counts. An analysis of variance made on the
repetitive algal counts showed that 70% of the Eu-
glena counts in excess of 10s cells per milliliter
were within 20 per cent of the stated values, and
that 70% of the Chlorella counts in excess of 10"
cells per milliliter were within 10 per cent of the
stated value. The same order of reproducibility is
to be expected then for the dry weight estimations.
With regard to the accuracy of the assumed weights
of algal cells in confirmatory experiments, a large
number of replicate algal cultures were grown on
filtered wastes and their cell weights determined
volumetrically by determining centrifuged packed
volume and the relationship between packed volume
and cell weight. Good correlation was obtained
between the cell count and volumetric methods for
growth evaluation.
Incubation Apparatus
A further advantage of the cell count method
of growth evaluation is the simplicity of culture
apparatus. In Figure 24 a cross-section of the
simple incubator used for the assays is shown. It
consists of several wooden shelves (A) and is il-
luminated by a 30-watt daylight fluorescent lamp
(B) mounted parallel to each shelf. A window glass
shield (C) reduces the amount of heating within the
sample culture (D), and also decreases the magni-
tude of the ultraviolet component of the fluorescent
spectrum to a point where it has little effect on the
cultures. Their radiance within a 200 milliliter
Erlenmeyer flask in place in the incubator is about
10 calories per liter per minute, and under con-
tinuous illumination, the total energy available to
a 75 milliliter culture within a flask is about 14,000
calories per liter of culture per day. Because of
their high surface area to volume ratio, the flasks
dissipate waste heat to the surrounding air during
incubation and therefore maintain a uniform tem-
perature of about 25°C.
Performance of Tests
In the actual test procedure, 75 milliliter sam-
ples of serial dilutions made of the wastes under
study were inoculated with unialgal cultures of
Chlorella and of Euglena and, if required, of sew-
age bacteria. The cultures were then incubated
under the standard conditions previously described
until the daily algal growth increment became
negligible. Daily cell counts were made with a
haemocytometerand total growth computed as pre-
viously described. Growth rate constants were
also computed from some of the cell counts. The
average growth rate kg, was computed from the
expression kg « r log S2- in which C0
O I \_,J ^
-------
Metropolitan Wastes and Algal Nutrition
91
is the population at the end of the time period and
Cj, the population at the beginning of the time
period. These rates were determined merely as
a matter of interest. The wastes were analyzed
initially for total solids, pH, alkalinity, and for
B.O.D. as determined either according to Standard
S'-K/
'" ""'^^^f^s^s
•^z^^^*z^^
8"
"Of
'•*)•"•" '"-fj^f^^fj
-'•'•'•'•'•'-' *r*f-*
YENT
I BALLAST
VENT
Figure 24. INCUBATION APPARATUS FOR BATCH
CULTURES OF ALGAE IN ORGANIC WASTES.
A - shelf; B - lamp; C - window glass; D - Sam-
ple flask.
Methods or with a Warburg respirometer. In the
confirmatory experiments, packed solids were
analyzed for initial and final B.O.D., pH, alkalinity
and daily algal counts. The B.O.D. of the final ef-
fluent was determined by centrifuging the culture
at 500 times gravity for 10 minutes and determin-
ing the B.O.D. of the decanted supernatant.
Values of the deoxygenation constant k were
computed on the basis of B.O.D. data from the ox-
ygen versus time curves using the simplified meth-
od for analysis of B.O.D. data as described by
Moore, Thomas, and Snow (1950). Avalue for algal
reoxygenation termed ka was also computed in cer-
tain cases using the Moore, Thomas, and Snow
method for analyzing data on oxygen produced.
RESULTS
The Influence of Time and Algal Species
Figure 25 shows the growth curves of Euglena
500
ALGAL OXYGEN
PRODUCTON.CHLORELLA
ALGAL OXYGEN
PRODUCTION,
EUGLENA
66
TIME, DAYS
Figure 25. COMPARABLE GROWTH CURVES
FOR EUGLENA AND CHLORELLA ON FRESH
CHICKEN-PEN WASTE.
and Chlorella cultured on fresh chicken pen waste
which had been diluted first with tap water to 1,000
ppm, and then with distilled water to obtain a
B.O.D. of about 275 milligrams per liter. The
growth curves are given in terms of total oxygen
produced, i.e., 1.6 times the dry weight as deter-
mined from Figure 23, in milligrams per liter.
According to the figure, Chlorella had a growth
rate sufficient to result in the production of oxygen
-------
92
ALGAE AND METROPOLITAN WASTES
at a rate greater than that required to meet the
B.O.D. of the waste. Given sufficient time, how-
ever, the maximum dry weight of algal material
produced in a given culture was approximately the
same for both Chlorella and Euglena. Neverthe-
less, it was decided that although Euglena are
characteristically polysaprobic, their generation
time is too long to permit their use as valid indica-
tors of maximum potential algal production. It is
considered likely that, had it been measurable,
the exertion of B.O.D. by bacteria in some of the
cultures with Euglena would have been retarded by
a lack of oxygen. Chlorella produced more total
oxygen than Euglena. In fact, within five days
Chlorella produced more than 1.6 times as much
oxygen as required to meet the five-day B.O.D. of
the waste, although, over a period of ten days
Euglena produced more than 1.4 times as much
oxygen as was required.
The Influence of Waste Dilution
In Figure 26 is shown the influence of time and
solids dilution upon the B.O.D. of chicken pen
wastes and upon oxygen production by Chlorella.
According to Figure 26(A) algal oxygen production
exceeded the B.O.D. of the wastes at the beginning
of the experiment and was the equivalent of twice
the B.O.D. after five days. At a solids concentra-
tion of 250 ppm (Figure 26(B), algal oxygen pro-
duction did not meet oxygen demand until 3.5 days
had elapsed after initiating the experiment. How-
ever, after six days, oxygen production was 1.7
times the B.O.D. As shown in Figure 26(C), at a
waste concentration of 500 ppm, almost five days
elapsed before oxygen production by Chlorella was
equal to the B.O.D. of the waste. The curve in
Figure 26(D) shows that when the concentration of
the waste was 1,000 ppm the B.O.D. of the waste
was exerted at a rate much more rapid than that of
oxygen production by Chlorella.
The Influence of Dilution Water
In Figure 27 is shown the five-day Standard
Methods B.O.D. of reduction plant wastes as a func-
tion of waste concentration. Dilution of the wastes
for B.O.D. determination was made with dilution
ToM MMd* odstd- 2SO Mm
CHCXEM MANURE
CMLOBELLA
CMOCN MANURE
CHUWELLA
TcM Mi* odwd- SOOmm
Tott Mid* oddtd-'OOO
- CMCKEN MANURE/
CMUDRELLA '
(D)
9OO
800
TOO
O
O
O
6OO
O 500
O
O
tc
"-400
O 300
•X
I
ZOO -
CHLORELLA o
REDUCTION PLANT WASTE /
(Variable dilution water) /
' . 5 DAY BOO
"Standard Method*"
67
4OO 8OO BOO
SOLIDS CONCENTRATION, MG/L
Figure 26. THE INFLUENCE OF SOLIDS DILU-
TION UPON THE B.O.D. OF CHICKEN-PEN
WASTES AND UPON OXYGEN PRODUCTION BY
CHLORELLA.
Figure 27. THE INFLUENCE OF WASTE DILU-
TION AND OF DILUTION WATER TYPE ON
GROWTH OF CHLORELLA IN REDUCTION PLANT
WASTE.
-------
Metropolitan Wastes and Algal Nutrition
93
water prepared according to Standard Methods.
Oxygen production by Chlorella grown in reduction
plant wastes diluted with three types of dilution
water to the solids concentration indicated are
shown in curves identified according to the type of
water used. The curves show that with the excep-
tion of river water, the oxygen produced by Chlo-
rella increased with waste concentration between
0 and 400 or 500 milligrams per liter, and then
decreased. In the case of river water, no algal
growth occurred at a waste concentration of 200
milligrams per liter, and growth was barely de-
tectable at a waste concentration of 400 milligrams
per liter. Growth was enhanced when the concen-
tration of the waste was increased to 816 milli-
grams per liter. When either well water or Rich-
mond tap water was used, the oxygen produced
during five days of growth exceeded the five-day
B.O.D. up to a waste solids concentration of about
650 ppm. Since growth was plentiful when either
well water or Richmond tap water was used as a
diluent, it is evident that some factor in the river
water inhibited the growth of algae and conse-
quently, the production of oxygen.
Influence of Type of Waste
In Table 29, the various wastes subjected to
Table 29. ALGAL GROWTH AND OXYGEN PRO-
DUCTION
Wute
ifilk
Abattoir
Reduction plant
Packing bouse
Cattle holding pen
Chicken pen
Ground garbage
Vegetable cannery
Winer?
Tomatoe cannery
Uuonlte
KoooKxHum glatamate
Glntamlc acid
Domeatlc sewage
Domestic aevage •upenalant
Relative (1)
Growth
++
+*+
*++
»t
*
***
++
+*
+•»
*•*
«**
Equivalent (1)
cone. ppm.
Don*
ISO
HO
900
150(2)
500
350
150(1)
None (2)
100
None
None
400
370
SdayBC
l^i^i
_
in
400
wo
—
300
50
100
None
BO
—
—
200
300
35
1C ppm.
Final
_
—
10
10
— -
50
-
— -
40
SO
10
(1) Dry weight concentration of the wait* >l which BOD 1> equal to algal oxygen
production.
(J) t Detectible (rowth.
** Growth leaa than required to meet BOD.
*•• Good growth oxygen produced equal to BOD at high dilution..
**•« Excellent growth oxygen production greater than BOD at low dtumons.
assay are listed, together with data on the equiva-
lent concentration of the waste at which the B.O.D.
of a particular dilution equalled the oxygen pro-
duced by the algae grown at that dilution, and data
on the initial and final B.O.D. It is evident that, in
most cases, at a concentration of waste less than
the listed equivalent concentration, algal oxygen
production usually exceeded the oxygen demand of
the waste. A waste was labeled "none", when the
extent of algal growth was insufficient to produce
the oxygen necessary to meet the estimated B.O.D.
at any of the several dilutions tried.
The fourth column in Table 29 lists the five-
day 20°C B.O.D. of the wastes at their equivalent
concentration. A waste for which no B.O.D. is
listed either was not tested or gave highly erratic
results. Determinations were made of the final
B.O.D. of reduction plant, packing house, chicken
pen wastes, and of domestic sewage and glutamic
acid. In all cases, the final B.O.D. was low, an
indication of the exhaustion of the readily available
organic matter in the wastes. The supernatant
from domestic sewage was reinoculated with Chlo-
rella and incubated under standard conditions for
20 days. No detectable growth of algae was ob-
served during this period, an indication that only
one crop of algae could be grown in this dilute
waste. It is evident from the table that algal growth
was favored by the more nitrogenous wastes, since
relatively poor growths occurred in vegetable
wastes. Although pigmented and acid wastes only
permitted poor growths, the extent of the algal
growth observed at the higher dilutions of these
wastes indicates their potential for producing long-
time nuisance blooms.
In Table 30 are presented the results of experi-
ments referred to in the introduction as confirma-
tory, conducted primarily to verify the results
listed in Table 29 which were obtained on the basis
of cell counts. As noted previously, at the time a
given sample was incubated in the light, after being
inoculated with algae, a replicate was placed in
the Warburg to determine its Warburg B.O.D. The
Table 30. ALGAL OXYGEN PRODUCTION IN
COMPARISON TO WASTE BOD
Non-fat milk
Reduction plant*
Chicken manure*
Fresh sewage*
Monasodlum glutamate*
BOD
180
220
100
160
190
Algal 02
production
60
350
130
375
Nil
U)
*a
.25
.30
.09
.075
(2)
k
.21
.30
.10
.15
(»)
^
.45
.60
.90
.50
(1) ka * Oxygen production.
(2) k - Oxygen utilization.
(3) kg = Average population growth rate.
• Filtered.
Warburg B.O.D. and the packed volume of the algal
cell material were determined daily. Inasmuch as
the wastes had been filtered or were highly dilute,
most of the extraneous material was excluded, and
consequently the centrifuged packed volume gave a
relatively valid approximation of the dry weight of
cells produced. It is seen from the table that
filtered reduction plant waste, chicken manure,
and fresh sewage sustained an algal growth suffi-
cient to produce oxygen in excess of the equivalent
Warburg B.O.D. whereas the simulated milk wastes
supported little algal growth even when highly di-
lute. Few algae could grow in wastes containing
monosodium glutamate end-liquor, probably be-
cause light could not penetrate the dark liquid. Its
high solids content may have been a contributing
inhibitory factor. Values of k are presented in
Table 30 to give an indication of the relative rates
-------
94
ALGAE AND METROPOLITAN WASTES
of 'algal oxygen production and bacterial oxygen
utilization in the waste study. Comparable to the
k values normally used in B.O.D. analyses, the lar-
ger the value of k and kg, the more rapid the rate
of oxygen utilization or oxygen production respec-
tively. It is interesting to note that algal oxygen
production was rapid in milk wastes but that the
total amount was so small it never exceeded B.O.D.
Hence, although algal growth occurred, the con-
centration was never such that, oxygen production
equalled oxygen utilization, as was the case with
other wastes studied.
DISCUSSION
The criterion that the nutrient quality of a waste
for algal growth is indicated by the algal produc-
tion of oxygen in excess of the B.O.D. of the waste,
is open to some discussion. Its validity certainly
may be considered to be proved with respect to do-
mestic sewage but the amount of research done in
culturing algae on other types of wastes has been
so limited as to exclude all but the most prelimin-
ary of conclusions. It seems reasonable to believe,
however, that a waste having sufficient nutrients
in balanced form to produce a strong surge of algal
growth in a laboratory growth unit is also likely to
promote a comparable growth in any body of natural
water, providing that light and temperature are
within permissible limits as to magnitude. It is
unlikely that a more concentrated growth of algae
would occur outdoors than under the laboratory
condition used, because the total light energy avail-
able in the laboratory incubator exceeded several-
fold that available under usual outdoor conditions.
In some of our earlier studies, attempts were
made to culture algae in sterile organic wastes.
Algae did grow well in many such wastes but the
growth was much less intense under sterile con-
ditions than when the waste inadvertently became
contaminated. This led to the adoption of the prac-
tice of intentional bacterial inoculation with endo-
genous organisms which was followed in this study.
It now appears certain that bacterial decomposition
of the waste just prior to or simultaneously with
the onset of algal growth liberates essential nu-
trients and leads to a more extensive algal growth
than would otherwise occur. In natural waters
many of the bacterial sludges which form during
decomposition find their way into benthal deposits
where decomposition to methane decreases the
amount of carbon available for photosynthesis.
Direct utilization of compounds by algae has
been a subject of great interest, and there is much
documentary evidence that certain soluble sub-
stances are assimilated by the cells in the dark.
Nevertheless, there is little question that during
times when significant increases in algal concen-
tration do occur, photosynthesis is the major me-
chanism for increase. Under such conditions am-
monium is the principal energy containing com-
pound incorporated into cell material. Such rapid
incorporation of ammonia into algal cells prevents
its oxidation to nitrate, and inasmuch as algae
eventually settle, this portion of the nitrogen may
be removed from soluble form.
Each of the wastes in which strong algal growth
occurred, also showed an optimum concentration
for maximum algal growth. Such concentrations
probably were much higher than those which would
be encountered in natural waters and therefore al-
gal growth occurring naturally would be more com-
parable to that obtained at the higher dilutions
studied. The decrease observed in the algal con-
centration obtained at waste concentrations above
a certain maximum, probably was due to a decline
in the total available light. Some of the stronger
wastes were extremely turbid. This turbidity ab-
sorbed light and inhibited algal growth to the ex-
tent that less oxygen was produced than was re-
quired to satisfy the B.O.D. Several of the strong
wastes became foul smelling and acid conditions
persisted, with an apparently detrimental effect on
Chlorella. After long periods of incubation these
foul cultures also became fresh smelling as a re-
sult of algal-bacterial stabilization.
Perhaps the most interesting, if not the most
expected finding in this study, is the strong influ-
ence of diluting water upon the growth of algae at-
tained in reduction plant wastes. For example,
wastes diluted with water from a river in the dis-
posal area strongly inhibited algal growth. This
inhibition could be attributed to some nutrient pre-
sent in well water and in Richmond tap water, but
missing in the river water, or it could be attributed
to some toxic substance in the river water. A more
extensive study of the phenomenon is now under
way.
Highly pigmented wastes supported algal growth
only at highest dilutions. Because some of the
wastes, such as masonite waste, also contained
substances toxic to bacteria, such as phenols, their
nutrient content was released only at the higher
dilutions. In natural lakes or streams the slow re-
lease of nutrient from such wastes, despite their
toxicity, could cause nuisance blooms over a long
period of time in spite of the fact that there was
little evidence of such tendency in the laboratory.
Milk wastes, because of their acidity, do not
favor algal growth, but under conditions where oxi-
dation is favored, algae do grow in their presence.
For example, a small rapidly flowing stream at
Novato, California, receives the waste from a
cheese factory and courses over sandstone for
several hundred yards. A one millimeter thick
layer of Chlorella grows upon the submerged sand-
stone surface, where it is protected from the
strong sunlight and perhaps from whey acids by a
thin layer of aerobic bacteria growing upon its
surface. The little stream is highly aerobic in
-------
Metropolitan Wastes and Algal Nutrition
95
spite of a B.O.D. load which, on the basis of the
atmospheric reaeration theory, should render it
anaerobic.
SUMMARY
A series of bioassays was performed for the
purpose of determining the nutritional value of the
more common organic metropolitan wastes for the
algae, Euglena gracilia, and Chlorella pyrenpido-
sa. The algae were seeded into a series of flasks
containing the whole and diluted wastes of known
B.O.D., which previously had been seeded with bac-
teria and were undergoing active bacterial decom-
position. In each experiment a standard sewage
was used as a reference and control. Inasmuch as
the cultures were incubated at a temperature of
about 20°C, in fluorescent light of optimum wave
length, moderate intensity and long duration, the
magnitude of algal growth obtained was mainly a
function of the quality and availability of nutrients
in the waste and its dilution water undergoing
assay.
Any algal growth in excess of that which oc-
curred in the dilution water was assumed to indi-
cate the potentiality of long term problems. It
was assumed that wastes in which oxygen produc-
tion by algae exceeded the B.O.D. of the waste could
be expected to produce heavy algal blooms under
natural conditions, whenever dilution, light and
temperature permitted, and when bacterial oxi-
dation of the waste was in progress.
The metropolitan wastes studied were: milk,
abattoir, reduction plant, packing house, cattle
holding-pen and chicken pen wastes, ground gar-
bage, mixed vegetable cannery wastes, tomato
cannery wastes, masonite manufacturing wastes,
monosodium glutamate end liquor, glutamic acid,
winery waste and domestic sewage. Analytical
work was carried out to determine: (1) the influ-
ence of waste type on algal growth; (2) the influence
of waste concentration on algal growth; (3) and the
influence of algal growth on stabilization of the
wastes as indicated by the relationships between
algal growth, oxygen production and B.O.D.
It was found that Chlorella because of its high
growth rate was of greatest value in waste assays.
Each of the wastes would support some algal
growth. However, milk, monosodium glutamate,
winery waste and masonite did not support a suffi-
cient growth of algae to produce the oxygen needed
to meet the B.O.D. requirements of the diluted
wastes and therefore it is possible that these
wastes would cause long term nuisance problems.
Algal growth in each of the other wastes was suffi-
cient to satisfy the B.O.D. at some specific dilution.
These wastes could, therefore, be expected to sup-
port heavy algal growths under natural conditions
with odorless natural removal of nutrients occur-
ring, providing the waste is properly diluted, and
the waste pond correctly designed.
ACKNOWLEDGMENTS
I am indebted to Dr. C. G. Golueke for review-
ing this manuscript and to Mr. Henry Gee for pre-
paration of the figures. This research was sup-
ported in part by a Research Grant from the Na-
tional Institutes of Health of the United States
Public Health Service.
REFERENCES
Fisher, A. E. 1953. Pilot plant studies in the pro-
duction of Chlorella. Chapter 17, page 248 in Algal
Culture from Laboratory to Pilot Plant. Carnegie
Institution of Washington Publication 600.
Ludwig, H. F., W. F. Oswald, H. B. Gotaas and
V. H. Lynch. 1950. Algae symbiosis sewage oxi-
dation ponds. University of California IER Series
44, !_!.
Moore E. W., H. A. Thomas and W. B. Snow. 1950.
Simplified method for analysis of B.O.D. data.
Sewage and Ind. Wastes 10_: 1343. (1959)
Oswald W. J., C. G. Golueke and H. K. Gee. 1959.
Waste water reclamation through production of al-
gae. Contribution 22, Water resources center,
University of California, Berkeley.
Oswald W. J., H. F. Ludwig, H. B. Gotaas, V. H.
Lynch. 1953. Photosynthetic oxygenation. Sewage
and Industrial Wastes 25, 6, 692.
-------
Limnological Relationships
THE ROLE OF LIMNOLOGICAL FACTORS IN THE AVAILABILITY OF ALGAL NUTRIENTS *
GEORGE H. LAUFF
Department of Zoology, the University of Michigan and
University of Georgia Marine Institute
ABSTRACT
The availability and utilization of algal nutrients
in lentic habitats are influenced by a complex of
factors that vary from one body of water to another.
In a broad consideration, these factors are related
to the morphometry of the basin and the nature of
the surrounding drainage area. Climatic conditions
such as wind and solar radiation are also involved
inasmuch as they affect the distribution and utili-
zation of nutrients once they enter the basin.
The relative availability of nutrient materials is
closely linked with the shape of the basin and its
mean depth. The cyclic relations involved in plant
growth, decomposition and nutrient regeneration
are more efficient when the zone of active plant
growth is in close proximity to the zone of most
active decomposition of organic matter and regen-
eration of nutrients. The extent of water movement
and the influence of water temperature on the me-
tabolism of the biota are also important considera-
tions in these cyclic phenomena.
Biotic factors are chiefly involved with utiliza-
tion of nutrients and their regeneration from or-
ganic matter produced within the habitat or derived
from its drainage area. The rate and thoroughness
of the decomposition of organic matter in the water
and the release of inorganic and organic materials
from the bottom sediments are important factors
in the cycle of some nutrients. Dissolved organic
matter may function as a nutrient or accessory
growth factor for planktonic algae; it can also act
as a toxin, or in the chelation of trace metals.
In reviewing some of the major limnological
factors that influence the availability and utiliza-
tion of algal nutrients, it is apparent that the his-
tory of limnology and knowledge of the factors
associated with the growth of phytoplankton have
been so completely interwoven that the two can not
be conveniently separated. Much of the early work
in limnology involved attempts at lake classifica-
tion based on their ability to produce phytoplankton
-and other organic matter. Naumann (1919) estab-
lished the nutrient-poor "oligotrophic" and the
nutrient-rich "eutrophic" lake types. He also added
a third type, the "dystrophic", in which humic
substances were regarded as important limiting
factors in the availability and utilization of nutri-
ents. Welch (1941; 1952) and Findenegg(1955) have
provided extensive reviews of the general subject
of lake classification. Despite the many attempts,
it remains to be demonstrated that a generally sat-
isfactory system of lake classification based on
nutrients or other indices of productivity can be
derived owing to the milieu of interrelated factors.
In attempting to evaluate the physical and chem-
ical parameters that influence the dynamic proces-
ses in lake metabolism, Rawson (1939; 1955; 1958)
has considered three primary groups:
a. Morphometric factors which relate to the
size and shape of the basin, and which may affect
the availability of nutrients.
b. Edaphic factors which affect the supply of
dissolved nutrients in the water.
c. Climatic factors which affect the utilization
of nutrients and, indirectly, their availability.
Depending on conditions, human influence may be
considered a fourth group. These primary groups
consist of many related factors which interact to
determine the lake's ability to produce and circu-
late organic matter; depending on specific condi-
tions, each of these may have a large or small ef-
fect on the lake.
As a background to the more pertinent limno-
logical factors, it may be well to review some of
the physical and chemical conditions which occur
in lakes and reservoirs, as they relate to this dis-
cussion.
During the spring when the temperature is uni-
form throughout the lake, the effect of increased
solar radiation is to warm the water at all depths,
but decreasingly so at increasing depths. The den-
*Contribution No. 22 from the University of Georgia Marine Institute, Sapelo Island, Georgia.
96
-------
Limnological Factors of Algal Nutrients
97
sity differences thus produced eventually lead to
the formation of three relatively distinct layers of
water: the upper layer or epilimnion and the lower
layer or hypolimnion, separated by the thermo-
cline or discontinuity layer. Under most condi-
tions, the temperature is approximately the same
at all depths in the epilimnion; the temperature
within the discontinuity layer falls rapidly with in-
creased depth and the resultant density gradient
functions in separating the epilimnion from the
hypolimnion. In the hypolimnion, the temperature
is lower than the overlying water, and it does not
change appreciably with increasing depth. When
meteorological conditions result in reduced air
temperatures for prolonged periods, the strati-
fication is destroyed by alteration of the density
structure and by action of strong winds on the lake
surface. The lake is then considered to enter the
period of overturn and the entire water mass is
freely circulated. An inverse stratification may be
effected by the occurrence of an ice cover, and is
followed by another overturn during the spring.
It is generally considered that the primary syn-
thesis of organic matter takes place in the epilim-
nion (trophogenic zone) and these materials ulti-
mately find their way into the hypolimnion and to
the bottom (tropholytic zone). Here the processes
of decay and mineralization result in the release
of various inorganic and organic compounds, and
the utilization of dissolved oxygen. The latter may
result in oxygen depletion during periods of pro-
longed stratification.
It is known that the chemical composition of
plankton is usually higher in protein, and fat con-
tent than organic matter derived from terrestrial
vegetation. Its incomplete mineralization is evi-
denced by the chemical composition of upper lake-
bottom sediments of eutrophic lakes which invari-
ably contain considerable amounts of nutrient ele-
ments in the form of resistant organic compounds.
It has been supposed that these materials are
mineralized in bottom sediments to a considerable
extent, with the nutrients thus released becoming
available in the trophogenic zone during periods of
overturn. The existence of extensive mineraliza-
tion has not been supported by chemical analyses
of the bottom sediments, however (Kleerekoper,
1953). Mortimer (1942) described the existence of
an oxidized microzone at the mud surface which he
considered to be maintained by molecular diffusion
of oxygen into the mud. It was demonstrated to
exert a profound influence upon the exchange of
substances across the mud-water interface. Re-
cently, Gorham (1958) has presented evidence
indicating that the thickness of the oxidized micro-
zone may depend upon the turbulent displacement
of the uppermost sediments into the overlying aer-
ated water, as well as upon the reducing power of
the .sediments themselves.
In general, the redox potentials in the organic
matter determine the extent to which organic sub-
stances can be hydrolyzed by bacterial enzymes
or decomposed by abiotic chemical processes. In
eutrophic lakes, these processes are limited by
the temporary or permanent lack of oxygen in the
tropholytic zone. Under anerobic condition, the
course of decomposition is different and less well-
known; decomposition does not proceed to complete
mineralization, but stops short at intermediate or-
ganic stages such as methane (Ruttner, 1953).
Studies by Kleerekoper (1953) on the mineralization
of plankton suggest that most of the decomposition
of the sinking planktonic detritus may take place in
the epilimnion with the resultant liberation of much
of the nitrogen; the liberation of phosphorus ap-
peared to be slower, and both phosphorus and
silica accumulated in the surface bottom sedi-
ments.
The availability of certain nutrients such as
iron, manganese, nitrogen and phosphorus is
closely associated with the cyclic chemical con-
ditions which exist during periods of stratification
and overturn. Under certain conditions, bicarbon-
ate salts of iron and manganese are precipitated in
the presence of oxygen; phosphorus may behave
similarly, though it is also made available in a dis-
solved form through bacterial putrifaction or by
activities of protozoa (Hooper and Elliott, 1953).
These materials may thus be relatively abundant
in the dissolved state in the oxygen depleted hypo-
limnion, along with silicic acid which occurs in a
dissolved or colloidal form. Nitrogen may be pre-
sent as nitrate in the epilimnion, but it appears
chiefly as ammonia, derived from breakdown of
protein, in the oxygen-free hypolimnion.
It has already been pointed out that increased
solar radiation in the spring eventually may result
in the production of a stratified condition. Of the
sunlight which strikes the lake surface, a certain
fraction is reflected and never enters the water.
The remainder penetrates to varying depths, de-
pending upon the concentration of dissolved colored
substances and the absorbing and scattering ma-
terials such as plankton and inorganic and organic
particulate matter. All of the light is eventually
absorbed by the water and phytoplankton, except
for that reflected at the surface or lost by back-
scatter.
Thus, the amount of light which penetrates in
different bodies of water is variable, depending
upon prevailing conditions; it is also altered in
spectral composition by differential absorption of
certain wave lengths. In stained water, for exam-
ple, the short wave lengths of the violet and blue
portion of the spectrum are absorbed more readily
than the orange and red. In distilled water, essen-
tially the reverse is true. Since the rate of photo-
synthesis of algae is controlled by both the inten-
sity and spectral composition of light, and by tem-
perature, the extent to which solar radiation
-------
98
ALGAE AND METROPOLITAN WASTES
penetrates the water influences the utilization of
nutrient materials. Specific relationships between
photosynthesis and various components of the visi-
ble spectrum are not well recognized. It is known,
however, that the light and temperature relations
are not the same for all algae but the same general
limiting factor relation applies; namely, at any
given temperature, photosynthesis is directly pro-
portional to the light intensities at low light, but at
high intensities, it is independent of light intensity
and can be increased only by a rise in temperature
(Lund, 1958). The seasonal variation in light and
temperature has been used to explain the shift in
composition of phytoplankton from spring to fall;
some investigators attribute the observed changes
to the availability of nutrients and/or the produc-
tion and effect of metabolites.
The cyclic relations involved in the utilization,
regeneration, and circulation of nutrients is most
efficient when the zone of active plant growth is in
close proximity to the zone of most active decom-
position of organic matter and regeneration of nu-
trients. From this viewpoint, one would anticipate
the most efficient system to be that in which there
was a continual circulation, with light and dissolved
oxygen being present to the bottom. In this situa-
tion, there would be relatively little lag between
regeneration of nutrient material and its utili-
zation in plant synthesis. These are essentially
the conditions that exist in the western end of Lake
Erie and other shallow lakes and reservoirs where
there is little or no stratification owing to a re-
duced mean depth and a favorable exposure to pre-
vailing winds. The mechanisms involved are also
those which function in the sewage stabilization
ponds employed in some regions of the United
States.
From the' physical point of view, the absorption
of the ma]°r portion of the incident solar radiation
in the surfacewater, owing to suspended materials
or water color, may result in the establishment of
an appreciable density difference that can not be
destroyed by prevailing wind conditions. In such
instances, a relatively thin epilimnion could deve-
lop. The nature and extent of the thermocline may
vary greatly, but in general the depth tends to in-
crease as the summer advances and the ratio of the
volume of epilimnion to volume of hypolimnion
therefore increases.
The availability of nutrients in bodies of water
tint have become stratified is dependent on the
water circulation patterns and basin morphometry.
The density structure that is established by in-
creased solar radiation and by wind energy can be
visualized as a series of circulating cells, with the
surface wind activity providing the driving force.
y flow exists ai""g the boundaries of the
circulating cells only when relative current move-
ments along the density interface are below a cer-
tain velocity. Depending on the existing conditions
of viscosity and density, a velocity may be reached
at which the laminar flow becomes turbulent:
wave-like disturbances develop on the interface
and grow in amplitude, and finally break into vor-
tices, resulting in a great increase in friction and
mixing. The production of waves and vortices in-
creases the area along the density interface many
times, and the mixing is completed by microtur-
bulence and molecular motion. It is this phenom-
enon which results in the erosion of the thermo-
cline and the increase in the volume of the epilim-
nion as the summer progresses.
The mixing caused by turbulence and eddy for-
mation not only functions in the transfer of heat,
but also results in the movement of nutrients into
the trophogenic zone where they may again take
part in organic matter synthesis. The extent to
which the waters of the hypolimnion are involved
in such circulation is not well known. The work
of Gorham (1958) on bottom sediments and that of
Mortimer (1951) on water movements suggests that
hypolimnetic circulation may be appreciable and
may provide an explanation for the development of
oxygen depletion near the bottom sediments at a
rate more rapid than would be expected on the
basis of molecular diffusion alone.
In shallow lakes where the volume of the epilim-
nion is large in comparison to the volume of the
hypolimnion, one could expect a more rapid and
efficient circulation of regenerated nutrients. In
shallow lakes, particularly those with irregular
shore lines, the intimate contact of the shore and
bottom sediments with the productive volume of the
lake should not be discounted in considerations of
nutrient availability. When the volume of the hypo-
limnion is appreciably greater than that of the
epilimnion, as is often the case in deep lakes, cir-
culation of nutrients would be expected to be much
slower and less efficient. The general relation-
ships between mean depth and organic matter pro-
duction of lakes have been studied by Raw son (1955).
His findings show an excellent correlation between
average kike depth and the unit area production of
plankton, benthic organisms, and fish, and provide
evidence for the efficient circulation and utilization
of nutrients in shallow bodies of water.
Only a few of the many factors associated with
the availability of nutrients have been briefly dis-
cussed. It should be pointed out that the products
of mineralization are not the only important com-
ponents in algal nutrition. Dissolved organic mat-
ter is known to exert its effects on phytoplankton in
four general ways (Saunders, 1957). Investigations
have demonstrated that certain algae are capable
of utilizing organic substances as an energy source
or as a nutrient. In some instances, dissolved or-
ganic matter may supply accessory growth factors
which are required for or stimulate growth of phy-
toplankton, whereas toxic substances are known to
be produced and inhibit growth. Organic complexes
-------
Limnological Factors of Algal Nutrients
99
may be formed with various trace metals and, de-
pending on circumstances, may function to remove
a particular nutrient from the trophogenic zone or
cause it to be retained. Recent investigations con-
ducted in some of Michigan's hard-water lakes
(Schelske, 1960) demonstrated that the addition of
certain organic compounds can enhance algal pro-
duction by apparently chelating iron and perhaps
other nutrients so as to prevent their chemical
precipitation as insoluble carbonates.
In summary, it may be stated that the availa-
bility and utilization of algal nutrients are associ-
ated with many physical, chemical and biological
factors that are related to the basin's morphome-
try, as well as the edaphic and climatic conditions
that prevail. Since these factors vary in impor-
tance from one body of water to another, and from
time to another, anything more than simple gen-
eralizations is difficult; however, information is
available which will permit detailed evaluation of
particular situations if desired.
REFERENCES
Findenegg, V. I. 1955. Trophiezustand und See-
typen. Schweizarische Zeitschrift Fur Hydrologie.
17: 87-97.
Gorham, E. 1958. Observations on the formation
and breakdown of the oxidized microzone at the
mud surface in lakes. Limnol. and Oceanogr. 3:
291-298.
Hooper, F. F., and A. M. Elliott. 1953. Release of
inorganic phosphorus from extracts of lake mud by
Protozoa. Trans. Am. Micro. Soc. 72j 276-281.
Kleerekoper, H. 1953. The mineralization of
plankton. J. Fish. Res. Bd. Can. 10j 283-291.
Lund, J. W. G. 1958. Primary production in in-
land waters. In the Biological Productivity of Bri-
tain, Publ. Inst. of Biol., London: 1-6.
Mortimer, C. H. 1942. The exchange of dissolved
substances between mud and water in lakes. J.
Ecol. 30: 147-201.
1951. Water movements in stra-
tified lakes, deduced from observations in Winder-
mere and model experiments. Union Geodes et
Geophys. int., Assoc. int. d'Hydrol. scientif., As-
semblee gen. de Bruxelles. ^: 336-349.
Naumann, E. 1919. Nagra sympunkter angaende
planktons okologi. Svenske. Bot. Tidskrift 13.
Rawson, D. S. 1939. Some physical and chemical
factors in the metabolism of lakes. In Problems of
Lake Biology, Publ. Am. Assoc. Adv. Sci., No. 10:
9-26.
1955. Morphometry as a dominant
factor in the productivity of large lakes. Proc.Int.
Assoc. Theor. and Appl. Limn. 12: 164-175.
1958. Indices of lake productivity
and their significance in predicting conditions in
reservoirs and lakes with disturbed water levels.
In Invest, of Fish-Power Problems. H. R. MacMil-
lan Lectures in Fisheries. Univ, of British Colum-
bia: 27-42.
Ruttner, F. 1953. Fundamentals of limnology.
Univ. of Toronto Press. 242 pp.
Saunders, G. W. 1957. Interrelations of dissolved
organic matter and phytoplankton. Bot. Rev. 23:
389-410.
Schelske, C. L. 1960. Preliminary report of the
influence of iron, organic matter and other factors
upon primary productivity of a marl lake. Rept.
No. 1586, Inst. for Fish. Res., Mich. Dept. Cons.,
Ann Arbor.
Welch, P. S. 1941. Dissolved oxygen in relation
to lake types. In a Symposium on Hydrobiology.
Univ. of Wise. Press: 60-70.
Col, New York.
1952. Limnology.
538 pp.
McGraw-Hill
-------
100
ALGAE AND METROPOLITAN WASTES
NITROGEN FIXATION IN NATURAL WATERS UNDER CONTROLLED LABORATORY CONDITIONS
CLAIR N. SAWYER and ALFRED F. FERULLO
Metcalf & Eddy, Engineers, Boston, Massachusetts
The subject of nitrogen fixation in aquatic areas
and its relationship to the productivity of lakes and
impoundments appears to be a sorely neglected to-
pic of investigation, particularly in view of the fact
that the productivity of so many aquatic areas
seems to be unrelated to the gross contribution of
fertilizing matter in tributary waters. Clarke
(1954) in his book on "Elements of Ecology" states
—"the supply of nitrogen in the soil and in the wa-
ter may be further augmented as the result of ni-
trogen fixation by specialized bacteria belonging to
the genera Azotobacter and Clostridium which are
free living..." He also goes on to say — "Free ni-
trogen can also be fixed by certain photosynthetic
bacteria, sulfate reducing bacteria, and blue-
green algae." The significance of this phenomenon
appears to be left to the reader's imagination,
however.
Hutchinson (1957) discusses nitrogen fixation in
somewhat greater detail and points out that a con-
siderable number of bacterial species including
Azotobacter. Clostridia. Azotomonas. Aerobacter.
Methanomonas. and Pseudomonas are capable of
fixing nitrogen. He lists a number of blue-green
algae that have been reported to fix nitrogen.
These include such well known forms as Anabaena.
Gloeocapsa, and Nostoc.
Allen (1956) points out that the nitrogen fixing
ability of blue-green algae has been established
since 1928 and discusses the macro and micro nu-
trient requirements in considerable detail. Moly-
bdenum, cobalt, and sodium have been found to
be necessary items in the nutrition of blue-green
algae, and the significance of domestic sewage as
a source of these elements is well correlated with
the stimulation of blue-green algae blooms that
occur in aquatic areas fertilized by such waste wa-
ters.
STUDIES AT MADISON, WISCONSIN
The senior author's interest in nitrogen fixation
in aquatic areas stems from personal observations
made during the conduct of the Madison Lakes Sur-
vey, 1942-44 (Sawyer, 1944; and Sawyer, 1945).
A part of the survey method consisted of obtaining
visual evidence to demonstrate to a committee
composed of laymen as well as scientists that the
productivity of the aquatic areas under considera-
tion was related to the degree of fertilization. For
this demonstration, samples of waters collected
from the various lakes and streams were kept in
loosely stoppered, partially filled, glass containers
under natural light conditions in a green house.
The visual evidence obtained was quite dramatic
and perhaps, had more weight in convincing some
members of the committee of the significance of
fertilizing matters than all the analytical data ob-
tained.
During the course of the above-mentioned stu-
dies it was noted that many of the lightly fertilized
waters showed a productivity greatly in excess of
what would have been predicted, particularly when
the exposure period was continued for several
weeks. In addition, it was noted that blue-green
algae often came into dominance. This led to a
series of studies in which the nitrogen content of
the specimens was placed under close surveillance.
The results of these studies are presented in Table
31 and show that nitrogen fixation occurred in all
waters with the exception of the specimen from
Lake Wingra.
Table 31. NITROGEN FIXATION IN NATURAL
WATERS UNDER LABORATORY CONDITIONS
(NATURAL LIGHT)
Lake Waters
Mendota
Mendota
Monona
Monona
Waubesa
Kegonsa
Wingra
Streams
Six Mile Creek
Crawfish River
Yahara River
Days
58
167
41
61
115
116
45
151
167
116
Total
Start
0.48
0.51
1.03
0.58
1.72
0.82
1.88
1.97
_
2.00
nitrogen,
Finish
0.79
2.61
2.03
1.17
5.35
5.80
1.58
7.26
5.45
7.83
mg/L
A
0.31
2.10
1.00
0.59
3.63
4.98
-0.30
5.29
?
5.83
Nitrogen fixation appeared to be most prolific in
those lake waters receiving significant amounts of
fertilization from domestic sewage or drainage
from farm lands. An experiment was designed,
therefore, in an attempt to show the significance
of available nitrogen and phosphorus as factors in
nitrogen fixation. These studies were not made in
replicate and interpretations drawn from them
were considered tentative. The experiment con-
sisted of adding normal sewage treatment plant
effluent, effluent minus its nitrogen and phospho-
rus, and effluent minus its nitrogen to Lake Men-
dota water. The results of the study are shown in
-------
Nitrogen Fixation in Natural Waters
101
Table 32 and indicate that phosphorus is a key ele-
ment in the stimulation of nitrogen fixation.
Table 32. NITROGEN FIXATION IN LAKE MEN-
DOTA WATER FERTILIZED WITH SEWAGE EF-
FLUENTS
Mixture
Lake water, control
Plus 10% effluent
Plus 10% effluent
minus its N ft P
Phis 10% effluent
minus its N
Days
167
181
170
188
Total nitrogen, mg/L
Start Finish A
0.51
2.67
0.67
0.67
2.61 2.10
4.39 1.72
2.17 1.50
11.85 11.18
STUDIES AT BOSTON, MASSACHUSETTS
Taste and odor problems related to
algal
growths are becoming more and more common and,
oftentimes, a logical explanation of why the pro-
blem occurs is not always evident. It is always
embarrassing to a consulting engineer to have to
resort to some of the stock answers that have been
used in the past for, to him, there is a cause for
every effect.
In an attempt to explore more fully the signifi-
cance of nitrogen fixation in the productivity of
aquatic areas, our laboratory undertook a series
of experiments. These studies were designed to
evaluate the significance of nitrogen, phosphorus
and increased alkalinity, all items of considerable
concern insofar as fertilization by human wastes
is concerned.
The experiments were performed in the follow-
ing manner: Pond waters were brought to the la-
boratory, fortified with ammonium and/or phos-
phate salts to increase the level significantly and
the alkalinity was adjusted as desired by adding
sodium bicarbonate. The use of calcium salts was
avoided because of the known stimulatory effect of
calcium ion (Allen, 1956).
Two pond waters were selected for the initial
studies and plans were made to make repetitive
runs around the calendar in order to evaluate sea-
sonal changes. The samples were kept under flu-
orescent lighting at an intensity of 500 foot-candles
for 12 hours of each day.
Spy Pond. This pond has an area of 129 acres
and is located in Arlington, Mass. It is not used
as a public water supply since it is used for re-
creational purposes and is fringed by homes. It is
subjected, therefore, to some street drainage and
possibly some human wastes. Algae blooms of
moderate intensity do occur on occasion, in spite
of the fact that nutrient levels of nitrogen and
phosphorus are quite low.
Samples of Spy Pond water were fortified with
0.2 mg/L of phosphorus and/or 0.5 mg/L of am-
monia nitrogen. The aklalinity was increased
when desired by adding 200 mg/L of NaHCOs, equi-
valent to 120 mg/L of CaCOs- Three series of
studies have been completed and the results are
given in Tables 33, 34, and 35. The data for the
Table 33. NITROGEN FIXATION IN SPY POND
WATER (FLUORESCENT LIGHTING - 12 HR/
DAY) JULY 21 - AUGUST 25, 1959
Control
Control + Alk.»
Plus N
Plus N + Alk.
PlusP
Plus P + Alk.
Plus N t P
Plus N + P + Alk.
Total P,
mg/L
Tr.
Tr.
Tr.
Tr.
0.2
0.2
0.2
0.2
Total nitrogen.
Start Finish
0.57
0.57
1.07
1.07
0.57
0.57
1.07
1.07
0.78
077
1.38
1.04
1.69
1.60
1.86
1.61
mg/L
A (35 days)
0.21
0.20
0.31
-0.03
1.12
1.03
0.79
0.54
•200 mg/L NaHCOj
Table 34. NITROGEN FIXATION IN SPY POND
WATER (FLUORESCENT LIGHTING - 12 HR/
DAY) AUGUST 27 - NOVEMBER 25, 1959
Control
Control + Alk.»
PlusN
Plus N » Alk.
PlusP
Plus P «• Alk.
Plus N t P
Plus N «• P <• Alk.
Total P,
mg/L
Tr.
Tr.
Tr.
Tr.
0.2
0.2
0.2
0.2
Total nitrogen,
Start Finish
0.92
0.92
1.42
1.42
0.92
0.92
1.42
1.42
1.23
1.30
2.26
1.94
2.43
2.58
2.81
3.00
mg/L
A(90 days)
0.31
0.38
0.84
0.52
1.51
1.66
1.39
1.58
•200 mg/L NaHCOj
Table 35. NITROGEN FDCATION IN SPY POND
WATER (FLUORESCENT LIGHTING - 12 HR/
DAY) DECEMBER 3, 1959 — FEBRUARY 17,
1960
Control
Control + Alk.»
PlusN
Phis N + Alk.
Phis P
Plus P + Alk.
Phis N 4 P
Plus N + P + Alk.
Total P,
mg/L
Tr.
Tr.
Tr.
Tr.
0.2
0.2
0.2
0.2
Total nitrogen, mg/L
Start Finish A (70 days)
0.98
0.98
1.48
1.48
0.98
0.98
1.48
1.48
1.55 0.57
1.32 0.34
1.55 0.07
1.72 0.24
2.02 1.04
1.95 0.97
1.70 0.22
1.79 0.31
•200 mg/L NaHCOj
first two periods are summarized graphically in
Figures 28 and 29. From these data we conclude
that phosphorus plays a key role in nitrogen fixa-
tion and that the bicarbonate ion is relatively un-
important, since Spy Pond water has a .natural
alkalinity of only 30 mg/L.
Hagar's Pond. Hagar's Pond is a natural im-
poundment of about 27 acres in Hagar's Brook and
the water is heavily fertilized by the sewage plant
effluent from the city of Marlboro, Mass. The pond
water, as brought to the laboratory, contained over
-------
102
ALGAE AND METROPOLITAN WASTES
ui
o
o
n
IS,
CONTROLS
m— +P
POND WATER
•map
Figure 28. NITROGEN FIXATION IN SPY POND
WATER. (35 days, July-Aug.). Shading indicates
alkalinity added.
c
-I
9
-
u I
0
o
fc
z
0
_
—
_^_
—
n
ni
CONTROLS 4N
1
1
I
M
1
?v:
*S
»:
S
I
S
1
1
9
|
1
1
1
1
1
1
1
1
—
—
+p -map
POND WATER
Figure 29. NITROGEN FIXATION IN SPY POND
WATER. (90 days, Aug.-Nov.). Shading indicates
alkalinity added.
3 mg/L of total phosphorus and in excess of 5 mg/L
of total nitrogen. It was considered useless to at-
tempt studies by fortifying this water with phos-
phorus and nitrogen. Instead a study was made to
determine the nitrogen fixing ability of the water
when diluted with a high quality water. Boston tap
water was used for this purpose. A series of dilu-
tions were made as shown in Table 36. The sam-
ples were kept under laboratory conditions for 72
days and then analyzed. The data are given in Ta-
ble 36 and Figure 30. Only the mixture containing
100 percent and 20 percent of pond water showed
significant increases in nitrogen fixed. The ex-
periment was not repeated so no explanation except
biological variation can be offered at this time.
Table 36. NITROGEN FIXATION IN HAGAR'S
POND - BOSTON TAP WATER MIXTURES - JULY
29 - October 9, 1959
Hagar's Pond
water. %
100
50
20
10
5
2
1
0
Total P,
mg/L
3.3
1.65
0.66
0.33
0.17
<0.10
<0.10
<0.10
Total nitrogen, mg/L
Start Finish A (72 days)
5.38
2.92
1.45
0.95
0.71
0.56
0.50
0.46
6.74 1.36
3.46 0.54
4.55 3.10
1.61 0.66
1.03 0.32
1.09 0.53
0.97 0.47
0.81 0.35
-J 2
»*
o
UI
o
o
n
IOO
50 2O K> 5 2 I
t» POND WATER IN MIXTURE
Figure 30. NITROGEN FIXATION IN HAGAR'S
POND. Boston tap water mixtures. (72 days, Aug.
-Oct.).
DISCUSSION
In the laboratory studies conducted to date, no
attempt has been made to ascertain whether nitro-
gen fixation was due to algae or bacterial action.
It seems important to note, however, that when-
ever unusual amounts of nitrogen were fixed blue-
green algae were found in the test specimens. The
dominant genus noted was Aphanizomenon.
The studies conducted in the relatively soft wa-
ters of Massachusetts have confirmed the impor-
tance of phosphorus as a key element in nitrogen
fixation. This points up another important facet
-------
Nitrogen Fixation in Natural Waters
103
of the effect of human wastes in determining the
productivity of receiving bodies of water. Domes-
tic wastes, particularly since the advent of syn-
thetic detergents, are extremely rich in phospho-
rus in relation to nitrogen. Biological utilization
of the available nitrogen leaves a water that is
abundantly rich in phosphorus and ripe for exploi-
tation by nitrogen fixing forms. It is quite likely
that this condition accounts for the explosive
blooms of blue-green algae which occur in many of
our lakes and reservoirs.
CONCLUSIONS
1. Phosphorus is a key element in nitrogen fixa-
tion.
2. Fertilization of aquatic areas by domestic
wastes stimulates biological productivity.
3. Sewage plant effluents contain phosphorus in
excessive amounts.
4. The excess phosphorus can stimulate exten-
sive blooms of nitrogen fixing blue-green algae.
5. The productivity of most aquatic areas
probably related to their phosphorus budgets.
is
REFERENCES
Allen, M. B. 1956. Photosynthetic nitrogen fixa-
tion by blue-green algae. Scientific Monthly, p.
100, August.
Clarke, G. L. 1954. Elements of ecology. John
Wiley & Sons, N.Y.C.
Hutchinson, G. E. 1957. A treatise on limnology.
John Wiley & Sons, N.Y.C.
Sawyer, C. N. 1944. Investigation of the odor nui-
sance from Madison Lakes. July 1942-July 1943.
Sawyer, C. N. 1945. Investigation of the odor nui-
sance from Madison Lakes. July 1943-July 1944.
RECENT OBSERVATIONS ON NITROGEN FIXATION IN BLUE-GREEN ALGAE
RICHARD C. DUGDALE, Department of Biological Sciences,
University of Pittsburgh, and
JOHN C. NEESS, Department of Zoology, University of Wisconsin
INTRODUCTION
The fixation of N£ by organisms occupies one
facet of an extremely complex and interesting bio-
geochemical cycle. An excellent review of the
steps through which unequivocal proof was obtained
of the ability of certain of the blue-green algae to
fix nitrogen has been made by Fogg and Wolfe
(1954). In that article, a list showing the known
distribution of the ability to fix nitrogen in the My-
xophyceae is given; although several species have
been added since then, the early conclusion that the
family Nostocaceae is most important is still sup-
ported.
With the ability of blue-green algae to fix nitro-
gen definitely established, speculation has natural-
ly arisen regarding the role of the process under
natural conditions. Hutchinson (1941) noted a very
considerable increase in the combined nitrogen
content in Linsley Pond during a bloom of blue-
greens dominated by Anabaena circinalis and sug-
gested that biological fixation by the algae was pro-
bably responsible. Early recognition of the possi-
ble importance of this process was made by Saw-
yer, et al (1945) whose work on the Madison lakes
has already been reported to this conference. In a
recent appraisal, Hutchinson (1957) concludes that
this source of combined nitrogen can probably be
expected to be important when Anabaena is found in
heavy concentrations. Allen (1956) also concluded
from laboratory culture studies that fixation of ni-
trogen by blue-green algae could, under optimum
conditions, amount to about 480 pounds per acre
per month, more than double the quantity which
would be fixed by a legume-Rhizobium combination
for an entire crop.
A serious hindrance to an understanding of the
role of biological fixation in the aquatic environ-
ment has been the lack of a suitable method for
direct measurement of this contribution. Although
a balance-sheet can be constructed, as e.g., Roh-
lich and Lea (1949) have done, the possibility of
concomitant denitrif ication makes the magnitude of
the fixed component uncertain even if it were pos-
sible to determine accurately the loss to the sedi-
ments over the study period.
-------
104
ALGAE AND METROPOLITAN WASTES
The N*^ Method for Measuring Fixation Rates
Investigations in progress at the University of
Wisconsin and at the University of Pittsburgh em-
ploy the heavy isotope of nitrogen, N*5, as a tra-
cer, the method having been adapted from that used
by Burris, et al (1943). The procedure involves
primarily the removal of atmospheric dissolved
N2 from the water sample, held in specially de-
signed 1-liter vessels, and the subsequent replace-
ment with a quantity of nitrogen gas heavily en-
riched in N (95 atom-percent in most of these
experiments). The sample is then incubated by
suspending it from a buoy in the lake or in a tank
in the laboratory. Following a suitable incubation
period, usually 24 hours or less, the sample is
boiled down, Kjeldahl-digested, distilled, and ti-
trated with acid. After converting the ammonia to
nitrogen gas by the method of Rittenberg, et al
(1939) the final isotope ratio is determined with a
mass spectrometer. The total amount of N2 incor-
porated during the experiment is given by the sim-
plified expression:
N fixed = atom-percent excess N x total N in
atom-percent N supplied sample
if the increase in total N is reasonably small. The
entire procedure has been carefully calibrated; the
least significant enrichment in N* * has been found
to be 0.03 ug of N£- Ammonium sulfate is used as
a standard; excess N^"* is computed by subtracting
the atom-percent N^ of untreated lake water,
drawn from the same lotas that used in the experi-
ment, from that of the incubated sample.
Results Obtained from the
Method
One full year of work on Sanctuary Lake, the
uppermost portion of Pymatuning Reservoir in
northwestern Pennsylvania, has been completed.
Sanctuary Lake has a surface area of about 2, 000
acres, a maximum depth of 3 meters and a mean
depth of 2 meters. The lake has been shown by
Tryon and Jackson (1952) and by Hartman (in press)
to exhibit extremely dense blooms of blue-green
algae. Essentially no light penetrates to the lake-
bottom.
In Figure 31, the rate of nitrogen fixation, ex-
pressed as a percentage fixed per day of the exist-
ing total combined nitrogen, concentration of Ana-
baena spp., nitrate, nitrite, and ammonia, is
plotted. The data refer to fixation at the lake sur-
face and represent a composite of samples incubat-
ed in the lake and in the laboratory. Significant
fixation rates appeared concurrently with the pres-
ence of Anabaena in numbers. Following a decline
which coincided with a short-lived decrease in the
Anabaena population, the fixation rate rose and re-
mained at about 1 percent newly fixed nitrogen per
day until late August when a very considerable in-
crease in both Anabaena and fixation rate took
place. Two features are especially noteworthy.
ISO
160
140-
120-
S MOO
« x
3,5.00-
40-
20
•6
•
C
4*
• 2
Figure 31. RELATIONS AMONG RATE OF NI-
TROGEN FIXATION, CONCENTRATIONS OF
AMMONIA AND NITRATE AND NUMBER OF
CELLS OF ANABAENA AND DATE IN SANCTU-
ARY LAKE.
Percent fixation rate
Anabaena
NO--N
NH»-N
The period of rapid increase in the Anabaena popu-
lation occurred when the levels of inorganic com-
bined nitrogen had fallen to nearly undetectable
quantities. The concurrent rise in fixation rate
suggests that the expected advantage nitrogen fix-
ers would have over non-fixers under these condi-
tions is indeed realized, eventually resulting in
what Professor Hutchinson calls "Anabaena soup".
The second noteworthy event is the startling de-
cline and final disappearance of fixation at a time
when the Anabaena population was still nearly at
peak abundance. Although we have no experimental
evidence, this decline can probably be ascribed to
the simultaneous increase to very high values of
ammonia concentration in view of the well-docu-
mented depressing effect of available combined
nitrogen on nitrogen fixation (Fogg, 1954). The
subsequent rise of nitrate and the first appearance
of nitrite indicate the beginning of active nitrifica-
tion.
Phosphorus concentrations (not shown) fell with
the onset of the spring blue-green bloom and re-
mained low for the rest of the summer, indicating
a rapid rate of turnover or a high rate of exogen-
ous supply. The total combined nitrogen in the lake
increased several fold in late August as a result of
the very high rates of fixation prevailing at that
time. The process of accumulation of fixed nitro-
gen is self-accelerating wherever percentage rates
of fixation remain consistently high.
-------
Nitrogen Fixation in Blue-Green Algae
105
It is, of course, possible that the observed fixa-
tion is attributable to organisms other than the
algae. We have, however, shown a clear depen-
dence of the process of fixation in lake water upon
light by incubating samples in ordinary transparent
flasks together with similar samples in darkened
flasks, finding that fixation is reduced or obliterat-
ed in the latter. The process, as it occurs in lake
water, seems clearly to be energized by photosyn-
thesis directly or indirectly.
Some observations made in Lake Mendota (at
Madison, Wisconsin) during the summer of 1959
suggest that a very similar series of events takes
place there. In mid-September, daily percentage
fixation rates were measured above the thermocline
as follows: 0.019 (at 1 meter below the surface),
0.023(2 meters) and 0.014(4 meters). At this time
the phytoplankton was not dense and among the
blue-greens only Gleotrichia echinulata was present.
Nitrate was at undetectably low concentrations in
the entire epilimnion(0-12 meters), although there
was a small maximum in the thermocline; ammo-
nia concentrations were not measured. Nitrate be-
gan to appear in the upper water after October 13,
immediately after complete turnover of the lake,
and at the end of that month fixation rates had
dropped to zero. By themselves, these data are
not particularly useful, but they would be expected
if one assumed an inverse relationship between ni-
trate concentration and fixation rate such as was
found in Sanctuary Lake.
Fixation rates so low that they should probably
be termed non-existent have been measured under
the completely different conditions found in two
lakes on Afognak Island, Alaska.
Conditions for Intense Nitrogen Fixation
A working hypothesis maybe constructed at this
point as a guide for further experimentation and
observation. The following conditions are likely to
be necessary for the support of intense fixation in
a body of water:
1. The general physical and nutritional charac-
teristics of the body of water must be such as to
encourage the growth of blue-green algae.
2. Some factor(s) must operate to reduce the
concentrations of the various forms of combined
nitrogen to very low levels. This may be accom-
plished by the phytoplankton population itself, or by
some other agency such as denitrification.
3. The work of Sawyer, et al. (1945) indicates
that phosphorus is important. Although prevailing
concentrations apparently need not be strikingly
high, an adequate supply would appear to be a cri-
tical factor. The experiment of Buljan (1957) pro-
vides dramatic support for this point of view. In
it, a clear, blue, unproductive lagoon of the Adri-
atic Sea, whose water contained very low concen-
trations of phosphorus and combined nitrogen, was
fertilized with phosphorus alone, the assumption
being that, with an adequate supply of phosphorus
available, the nitrogen requisite for increased pro-
ductivity could be supplied by fixation. The re-
sults of the experiment amply proved this to be
the case, the blue water of the lagoon changing to
an unappealing green. These results are similar
to those obtained very much earlier by investiga-
tors studying fertilization of central European carp
ponds, showing that increased yields of fish from
ponds fertilized with combined nitrogen, phospho-
rus and the two in combination depended upon the
quantities of phosphorus provided, whether this
was used alone or with nitrogen, and not upon
amounts of added nitrogen. The latter was pre-
sumably supplied by local fixation.
4. Certain elements in trace amounts are now
known to be specifically necessary to permit nitro-
gen fixation by particular species of blue-green
algae; e.g., Eyster (1959) has shown that Ca, B,
and Mo are necessary for fixation by Nostoc mus-
corum, and has determined amounts necessary for
optimum growth in the absence of an external
source of combined nitrogen. A comparison of
these amounts with those ordinarily available under
natural conditions suggests that Ca is usually pre-
sent in sufficient concentration in fresh waters,
but that B and Mo are likely to be in short supply.
Little is known about the distribution of the latter
in fresh waters; the few analyses available indi-
cate that it will seldom be found in concentrations
greater than about 1 ug/1, about one-hundredth the
concentration required for optimum fixation by
algae in culture (Fogg, 1956). The success of Bul-
jan's experiment may well be due to the more-than-
adequate supply of B and the nearly-adequate sup-
ply of Mo (about 10 ug/1) ordinarily present in
sea-water. Billiard (personal communication) has
found in a series of brackish lagoons near Cape
Thompson, Alaska, the same three species of Ana-
baena (flos-aquae, spiroides and circinalis) that
have apparently been responsible for fixation in
Sanctuary Lake. These findings are not inconsis-
tent with the results of Allen (1956), who found that
Anabaena cylindrica could tolerate NaCl in con-
centrations as great as 1.5 percent. Allen (1959)
has also isolated nitrogen-fixing species of Calo-
thrix from coastal waters of California. All of
these observations seem to indicate that dilute
sea-water is a reasonably good medium for nitro-
gen-fixing blue-green algae, perhaps owing more
than anything else to favorable quantities of the
trace elements named above. It seems possible
that some of these elements are concentrated in
sewage, resulting under certain circumstances in
the stimulation of nitrogen fixation by this material.
We wish to acknowledge the support of the Na-
tional Institutes of Health for our studies on nitro-
gen fixation being carried out jointly at the Uni-
versities of Wisconsin and Pittsburgh. A very
considerable portion of the work reported for
-------
106
ALGAE AND METROPOLITAN WASTES
Sanctuary Lake and for lakes in Alaska has been
done by Vera Dugdale; John Goering has likewise
carried out a share of that reported for the Madi-
son lakes.
REFERENCES
Allen, M. B. 1956. Photosynthetic nitrogen fixa-
tion by blue-green algae. Sci. Month., 83: 100-106.
Allen, M. B. 1959. Photosynthetic Nitrogen Fixa-
tion in the Marine Environment. International
Oceanographic Preprints: 194.
Buljan, M. 1957. Report on the results obtained
by a new method of fertilization experimented in
the marine by"Mljetska Jezera", Acta Adriatica
6:6 1-54.
Burris, R. H., F. J. Eppling, H. B. Wahlin, and
P. W. Wilson. 1943. Detection of nitrogen fixation
with isotopic nitrogen. Jr. Biol. Chem., 148: 349-
357.
Eyster, C. 1959. Mineral requirements of Nostoc
muscorum for nitrogen fixation. Proc. IX Int. Bot.
Congr., 2: 109.
Fogg, G. E. 1956. Nitrogen fixation by photosyn-
thetic organisms. Ann. Rev. Plant Phys., 7; 51-
70.
Fogg, G. E., and M. Wolfe. 1954. The nitrogen
metabolism of the blue-green algae. Symp. Soc.
Gen. Microbiol., 4: 99-125.
Hartman, R. T. and J. Herbert Graffius. 1960.
Quantitative seasonal changes in the phytoplankton
communities of Pymatuning reservoir. Ecology
44: 333-340.
Hutchinson, G. E. 1941. Limnological studies in
Connecticut. IV. The mechanism of intermediary
metabolism in stratified lakes. Ecology Mon., 11:
21-60.
Hutchinson, G. E. 1957. A Treatise on Limnology,
Vol. I. New York, Wiley. 1015pp.
Rittenberg, D., A. S. Keston, F. Rosebury, and R.
Schoenhoomer. 1939. Studies in protein metabo-
lism, n. The determination of nitrogen isotopes
in organic compounds. Jr. Biol. Chem., 127: 291-
299.
Rohlich, G. A., and W. L. Lea. 1949. The Origin
of Plant Nutrients in Lake Mendota. Rept to Univ.
of Wis. Lake Invest. Comm. (mimeo.) 7 pp.
Sawyer, C. N., J.B. Lackey, and R. T. Lenz. 1945.
An Investigation of the Odor Nuisance Occurring in
the Madison Lakes, Particularly Monona, Waubesa
and Kegonsa, from July 1943 j-July 1944. ReptT
Gov. Comm., Madison, Wis. 92 pp., 25 fig., 27-
tables.
Tryon, C. A. Jr., and D. F. Jackson. 1952. Sum-
mer plankton productivity of Pymatuning Lake,
Pennsylvania. Ecol. 33: 342-350.
-------
METHODS OF PREVENTION OR CONTROL
-------
Limitation of Nutrients as a Step in Ecological Control
THE MADISON LAKES BEFORE AND AFTER DIVERSION
GERALD W. LAWTON
Hydraulic & Sanitary Laboratory
University of Wisconsin, Madison
The Madison lakes are among the most inten-
sively studied lakes in this part of the world. The
early work of Birge and Juday is well known
among limnologists. Domogalla (1926, 1935 and
1941) reported on studies of the lakes. Sawyer
(1943 and 1944), acting as director of the Gover-
nor's Committee, published his investigations of
the odor nuisances in the Madison lakes. Sawyer
and Lackey (1945) reported on plankton produc-
tivity in Wisconsin lakes which included the Madi-
son chain. Lawton and Bartsch (1949) reported on
an investigation of Lake Mendota and its tribu-
taries. Following 1947, twenty-five thesis studies
were made by the Hydraulic and Sanitary Engi-
neering Laboratories on the Madison lakes. Twelve
of these theses (Darrow and Jackson, 1949; Hag-
gerty, 1950; Levihn, 1951; and Teletzke, 1953)
covered chemical and biological characteristics of
the lakes and thirteen covered the hydrological
characteristics. Flannery (1949) summarized the
Madison lakes problem from a political science
point of view in a Master of Science thesis. Other
reports of investigations have appeared from time
to time, including that of Bryson, et al. (1955).
Presently a chemical and biological study of the
lakes is in progress at the Sanitary Laboratory of
the University of Wisconsin. Data for this dis-
cussion have been drawn from the above sources.
A map of the Madison lakes and their tribu-
taries is shown in Figure 32. Pertinent physical
characteristics of these lakes are given in Table
37.
Figure 32. MAP OF MADISON LAKES
The algae nuisance and odor problem has
plagued Madison for many years as indicated by
Professor Sarles earlier in this program. The
odor nuisance over the years has been variously
attributed to sewage, to industrial wastes, and to
algae. The first really comprehensive study of
the problem was made by the engineering firm of
Alvord and Burdick (1920). They traced the foul
odors to the decomposition of algae, and showed
Table 37. PHYSICAL CHARACTERISTICS OF THE MADISON LAKES*
Lake
Mendota
Monona
Waubesa
V •*
Wingra
Length,
miles
5.90
4.16
4.20
300
1.00
Width,
miles
4.60
2.40
1.40
2 2*)
0.37
Area,
sq. miles
15.20
5.44
3.18
4 91
Max.
depth,
ft.
84.0
74.0
36.6
31 4
14.1
Mean
depth,
ft.
39.7
27.6
16.1
15.1
8.9
Volume,
M.G.
126,385.35
31,409.95
10,634.58
15 603.63
1,497.52
Ave. drop
between
lakes, in.
36.0
3.4
18.0
••••••
* From Domogalla (1926)
108
-------
Madison Lakes and Diversion
109
that the nitrogen and phosphorus contained in do-
mestic and industrial wastes were important con-
tributing factors in the production of algal blooms.
A resume of the progress of waste treatment
in Madison is given in Table 38.
The Governor's Committee, headed by Sawyer,
sought to determine the quantity and quality of all
fertilizing matter entering Lakes Monona, Waubesa
and Kegonsa. A thorough two year study deter-
mined the sources of the nutrients, phosphorus
and nitrogen, as shown in Figures 33 and 34. The
data show clearly that the bulk of the nutrients in
the lower lakes (Waubesa and Kegonsa) is derived
from the liquid wastes of the Madison metropolitan
area even though these wastes are subjected to
complete treatment.
In 1943 the Lewis bill requiring effluent diver-
sion was passed by the legislature and signed by
the Governor. Following various legal battles the
diversion sewers and channels were finally com-
pleted near the end of 1958 and diversion started
in December of that year. The route of the diver-
sion via Badfish Creek bypasses Lakes Waubesa
and Kegonsa as shown in Figure 35 (Woodburn,
1959).
This route involves pumping the effluent over
a divide 80 feet higher than the treatment works.
The pumping is accomplished by means of four
24-inch single stage double volute centrifugal
pumps, each with a rated capacity of 25,000 gpm
at 110 feet of head. The present flow can be han-
dled by one pump, but two will be necessary for
peaks. The excess pumpage capacity was provided
for emergency use and expansion. The effluent is
pumped from two equalizing basins, each 130 feet
in diameter, and discharged after chlorination
through a 54-inch pipeline consisting of 5.1 miles
of 100 psi reinforced concrete pipe. The effluent
then flows by gravity through 3. 8 miles of new
open channel and into the Badfish Creek. To re-
store oxygen to the effluent two cascade aerators
are provided in the new channel. One is located
at the end of the pipeline, and the other 0.6 miles
above the point of discharge into the creek. The
creek channel was widened and deepened for 10. 3
miles to the Dane-Rock County line. The remain-
der of the creek channel was considered to be
adequate to carry the increased flow.
The average flow in the Yahara River, as com-
puted from data contained in Sawyer's reports, is
approximately 100 mgd. Domogalla's calculations
of volumes for Lakes Waubesa and Kegonsa are
Table 38. WASTE TREATMENT AND DISPOSAL IN MADISON
Date
1886
1899-1902
1902-1914
1914-1926
1926
1930
1936
1942
1950
1952
1958 (Dec.)
Development
Sanitary sewer construction started
Chemical treatment
Septic tank and filter bed
Burke plant-contact bed and trickling filters
First unit of Nine Springs plant-trickling filters
Metropolitan district formed - completely covered
Lake Mendota and Lake Monona shoreline
Burke plant closed
U. S. Army leased Burke plant and reactivated it
East Side interceptor placed in operation (all
waste to Nine Springs plant)
Brewery ceased waste discharge to Lake Monona
Diversion started
Effluent disposal to
Lake Monona
Lake Mendota
Lake Monona
Lake Monona
Lake Monona
Lake Waubesa
Lake Monona
Lake Waubesa
Lake Waubesa.
Lake Monona
Lake Waubesa
Yahara River
(below Madison Lakes)
-------
ALGAE AND METROPOLITAN WASTES
MMUAL
TOTALS
§
<
8
H 125
K
ItJ
£ "o
E
m
o
| n
z
(O
I so
I
u
35
29,601 LBS.
i
c
o
3
— K
5 •
M °
i v
|!
i :
f~ « X
[ i *
fil 1 J jl
129, 366 LBS
&
U-J
-t
o^
1«
X
•
1
I
n°
n
r"
|
3
0
#
MF-1
118.874 LBS.
i
>*
E-"
J*-
!i
o
K
*
—
U ^ .
I
; —
w
M
O —
K
O
g
£ #
nri
ANNUAL
TOTALS
L MONONA
L WAUBESA
L KE60NSA
Figure 33. ANNUAL CONTRIBUTIONS OF INOR-
GANIC PHOSPHORUS - 1943-1944. (From Sawyer
and Lackey, 1944)
10,634 and 15,603 million gallons, respectively;
thus the theoretical flow through times are about
3.5 and 5.2 months. Estimates of this type led
many of the proponents of diversion to believe that
the removal of the effluent from the lower lakes
(Waubesa and Kegonsa) would quickly reduce the
nitrogen and phosphorus to levels where algal
blooms would occur only occasionally and of mod-
erate intensity.
Sawyer's studies pointed out the fact that the
unconsolidated bottom muds of these lakes re-
lease appreciable quantities of nitrogen and phos-
phorus to the overlying water. Reductions in the
concentration of these elements in the lower lakes
thus may be considerably less pronounced than
anticipated.
In an attempt to evaluate the effects of diver-
sion the present chemical and biological study was
started in June, 1959. Data are being obtained at
twelve stations located on lakes Mendota, Monona.
Wingra, Waubesa, and Kegonsa; on Badfish Creek
just above the junction with the Yahara River; and
on the Yahara River above and below the junction
to
a
§
a
6OO
K
X
u-
2
-
8
O
400
200
--
|
313,573 LBS
6
- S
3
—
O
a
0
S 3 ;
* 1
' £ '
nn
911, 065 LBS.
tit
C ° ~"z
«
•
>• u,
«i Z
o z
S
i
z
1
nn"
49O, 524 LBS.
w
_l
3
—
1 — »
^
U (
o
1
o
fin
L, MONONA
L. WAUBESA
L. KEGONSA
Figure 34. ANNUAL CONTRIBUTIONS OF INOR-
GANIC NITROGEN - 1943-1944. (From Sawyer
and Lackey, 1944)
*
Figure 35. NEW OUTFALL ROUTE via Badfish
Creek bypasses Lakes Waubesa and Kegonsa.
(From Woodburn, 1959)
-------
Madison Lakes and Diversion
111
with the Badfish Creek. Weekly samples are taken
during the summer and bi-weekly samples during
the winter. Determinations include temperature,
pH, alkalinity, ammonia, nitrate and organic ni-
trogen, B.O.D., C.O.D., soluble phosphorus,
volatile suspended solids, and estimates of
amounts of predominant algal species.
Before attempting to draw conclusions from
comparisons of data taken before and after diver-
sion, it is important to consider other factors that
enter into the production of algal blooms. Cli-
matological factors that are of importance include
variations in air temperature, water temperature,
sunshine, rainfall, and velocity and direction of
wind. Copper sulfate treatment of the Madison
lakes, to control the growth of algae, was practiced
with some regularity from 1925 to 1954. The
effect of this treatment on the early data cannot be
completely determined. A consideration of these
factors emphasizes the fact that one must be ex-
tremely cautious when interpreting the available
data.
CHEMICAL DATA
The data considered in this discussion include
concentration of inorganic nitrogen, organic nitro-
gen, soluble phosphorus and volatile suspended
1.2
z
bJ
O|X>-
O
K
I a
o
I6
o
5.4
solids, all expressed in mg/1. Average values for
each of the lakes are presented graphically for
both the growing (June through October) and non-
growing (November through May) seasons. These
values are based on samples taken at the outlet of
each lake.
Summer average concentrations for Lake Men-
dota are shown in Figure 36. A considerable
variation in concentration of inorganic and organic
nitrogen over the years is apparent, but there
appears no general trend. Soluble phosphorus is
slowly but steadily increasing as might be expected
with the increased use of land fertilizer.
Figure 37 presents average summer concentra-
tions for Lake Monona. This lake received all of
Madison's sewage treatment plant effluent until
1926 and smaller amounts until 1936. A decrease
in both inorganic nitrogen and soluble phosphorus
is noted following 1925. Burke plant effluent was
again discharged to Monona between 1943 and 1950
and is reflected in the higher values of nitrogen
and phosphorus in 1950, followed by decreased
values in 1959. Organic nitrogen and volatile
suspended solids content show no pronounced
trends; however, they are appreciably higher than
in Mendota, indicating a generally higher level of
algae during the frowing season.
•
ul.6
o
01.0-
z
< .8-
o
a. ..
O *
.5
.2
1922 1925 1940 (942 1943 I95O 1959
1922 1925 I94O 1942 1943 1950 1959
c
8
o.
4
3-
.1
E 14
in
o 12
a.
in
10
8
6-
2-
1922 1925 1940 1942 1943 I95O 1959 «22 1925 1940 1942 1943 1950 »59
Figure 36. SUMMER AVERAGES, LAKE MENDOTA
-------
112
ALGAE AND METROPOLITAN WASTES
gun
o
-------
Madison Lakes and Diversion
.113.
C. 12
*** 10
o
a.
t 8
o 6
z
«s
S .4
.2
1922 1925 1940 1942 1943 I960 1952 I9S9
il.8-
t 1-2-
ZIO
Me
S.6H
5.4-
2
^ 16-
-------
114
ALGAE
METROPOLITAN WASTES
12
uo
§
„ JB
*
Illlll
1942 1943 1948 1949 1950 1959
<2.0-
,5 1.8-
z 1.6-
ui
8 '
t **\
z 1.0
1 8
o .6"
0 .4
.2
1942 1943 1948 1949 1950 1959
~ J
v>
K 6
O
sU
™ 3
m
32
1942 1943 1948 1949 1950 1959
Figure 40. WINTER AVERAGES, LAKE MENDOTA
i 12
10
o -6
S -*
o
^ 22
5 2O-
z 16
UJ
§ .4
t '2
Z liH
u
z 8
tD 6
o:
O 4
S5O 1959
1942 1943 1948 1949 1950 1959
o
I
a
ui
at
1942 1943 1948 1949 1950 1959
Figure 41. WINTER AVERAGES, LAKE MONONA
-------
Madison Lakes and Diversion
115
201
~ .2
1942 1943 1948 1949 1950 1952 1959
1942 1943 1948 1949 I95O 1952 1959
1942 1943 1948 1949 1950 1952 1959
Figure 42. WINTER AVERAGES, LAKE WAUBESA
o
cc
o .o
2
O 4
CC
o
5 2
^- I 8-
5 16-
O 14
cc
t I2H
2 .OH
'i 8
<* 6
O
CC 4
O *
1942 1943 1948 1949 1950 1952 1959
*"^
r-i
Va-
.7
i
.4
.3
.2
1942 1943 1948 1949 1950 1952 1959
Figure 43. WINTER AVERAGES, LAKE KEGONSA
-------
ns.
ALGAE AND METROPOLITAN WASTES
ditions found in these lakes; hence it is likely that
a considerably longer period will be required to
thoroughly flush them out.
PLANKTON DATA
Over the years of study of these lakes, the con-
centration of algae has been expressed in various
terms—cell counts, volumetric standard units,
mg/1, and per cent of various species. The
methods of concentrating the algae have included
the following: filtration through plankton nets,
filter paper, sand, and membrane filters; cen-
trifuging; settling; and flotation. Samples have
been taken at various depths with various types of
sampling equipment during the many investiga-
tions. Methods of preserving the samples until
examined also have varied widely. Any correla-
tion of the plankton data obtained by these various
methods was considered to be of such a low order
as to be of little value. Consequently, only the
plankton data obtained during the growing seasons
of 1955, 1957, and 1959 are considered in this dis-
cussion. All samples during these years were
examined by Or. George Fitzgerald and are thus
directly comparable.
In 1955 and 1957, the net plankton from 5 to 10
quarts of sample were further concentrated and
preserved with the aid of formaldehyde and wet-
ting agent. In 1959, the total plankton were con-
centrated 20-fold using Rohde's modification of
Lugol's iodine-wetting agent method of concentra-
tion and preservation. The examination of the
plankton consisted of identification of the predomi-
nant species and an estimation of the relative
amounts of each species.
Table 39 summarizes the plankton data. Dur-
ing the period before diversion (1955 and 1957)
there were many species of algae in Lakes Monona
Table 39. ALGAL SPECIES IN MADISON LAKES
IMt n/M to 1WS)
t-n
MdMlrm, CMtoufcuriui. wd MI? othtn
tM1(T/» tat/4)
>rU.
IHt (VIS to Wl)
U-M
in. OKUtuwlm.
and Kegonsa, with the dominant species changing
from sample to sample. During the same period
the algae of Lake Waubesa always consisted of
99 per cent or more Microcystis.
Samples taken after diversion indicated that
again there were many species of algae in Lakes
Monona and Kegonsa, but the importance of
Microcystis in the plankton of Lake Waubesa de-
creased sharply to a point where it made up only
25 to 90 per cent of the total. Other algae of
significance in Lake Waubesa during 1959 were
Melosira. Oscillatoria. and Ceratium. plus others
in lesser amounts.
It appears that, in general, the number of
species and changes in dominant species in Lakes
Monona and Kegonsa have not changed during the
sampling periods, whereas the number of species
in Lake Waubesa has been markedly increased
since diversion.
SUMMARY
1. Variations in the amount of sewage treat-
ment plant effluent added to Lake Monona over the
years are reflected in the levels of nutrient con-
centration in that lake.
2. Diversion has completely stopped the ad-
dition of sewage treatment plant effluent to Lakes
Waubesa and Kegonsa.
3. The first growing season following diver-
sion showed no appreciable trend in the nutrient
content of Lakes Waubesa and Kegonsa.
4. The first full winter season following di-
version showed appreciable reductions in both the
inorganic nitrogen and soluble phosphorus content
of Lake Waubesa. Lake Kegonsa exhibited little
change from previous years.
5. The number of predominant species of
algae in Lake Waubesa has sharply increased
since diversion. The number of predominant
species in Lakes Monona and Kegonsa has shown
no appreciable change during the same period.
6. ft is realized that, due to the many factors
affecting the growth of algae, one cannot arbitrar-
ily conclude that the observed trends and changes
are entirely due to diversion.
7. Data obtained over a longer time period
will be necessary before the full impact of diver-
sion can be determined.
-------
Madison Lakes and Diversion
117
REFERENCES
Alvord and Burdick Engineers. 1920. Report upon
the cause of offensive odors from Lake Monona.
Bryson, R.A., G. P. Fitzgerald, H. M. Kaneshige,
and G.A. Rohlich. 1955. The Madison Lakes
Problem, Part I. Report on investigation of a
project sponsored by Oscar Mayer & Co.
Darrow, R.A.,andH.H. Jackson. 1949. A sani-
tary survey of the Yahara River. B. S. Thesis in
Civil Engineering, University of Wisconsin.
Domogalla, B. 1926. Treatment of algae and
weeds in lakes at Madison, Wis. Engineering
New-Record 97: 950-54.
Domogalla, B. 1935. Eleven years of chemical
treatment of the Madison lakes: Its effect on fish
and fish foods. Transactions of the American
Fisheries Society 65: 115.
Lawton, G.W. and A.F. Bartsch. 1949. Report
on Lake Mendota studies. Wisconsin Committee
on Water Pollution.
Levihn, Paul. 1951. A sanitary survey of the
Yahara River and bottom muds of Lake Mendota.
M.S. Thesis in Civil Engineering, University of
Wisconsin.
Sawyer, C.N., and J.B. Lackey. 1943. Inves-
tigation of the odor nuisance occurring in the
Madison lakes particularly Monona, Waubesa,
Kegonsa from July, 1942 to July, 1943. Report of
the Governor's Committee.
Sawyer, C. N., and J. B. Lackey. 1944. Investi-
gation of the odor nuisance occurring in the Madison
lakes particularly Monona, Waubesa, Kegonsa from
July, 1943 to July 1944. Report of the Governor's
Committee.
Domogalla, B. 1941. Scientific studies and
chemical treatment of the Madison lakes. A Sym-
posium of Hydrobiology, University of Wisconsin
Press: 303-09.
Flannery, J. J. 1949. The Madison lakes problem.
M. A. Thesis in Political Science, University of
Wisconsin.
Haggerty, J.R. 1950. A sanitary survey of the
Yahara River. M. S. Thesis in Civil Engineering,
University of Wisconsin.
Sawyer, C.N., and J.B. Lackey. 1945. Plank-
ton productivity of certain southern Wisconsin
lakes as related to fertilization. Sewage Works
Journal 17: 573-85.
Teletzke, G.H. 1953. A sanitary survey of the
Yahara River and Badfish Creek. M. S. Thesis in
Civil Engineering, University of Wisconsin.
Woodburn, J. G. 1959. Outfall around the Madi-
son lakes. Water and Sewage Works 106: 497-500.
-------
118
ALGAE AND METROPOLITAN WASTES
THE BADFISH RIVER
BEFORE AND AFTER DIVERSION OF SEWAGE PLANT EFFLUENT
THEODORE F. WISNIEWSKI
Director, Wisconsin Committee on Water Pollution
Madison, Wisconsin
INTRODUCTION
After Supreme Court decisions validated orders
of the Committee on Water Pollution issued pur-
suant to a statute prohibiting discharge of effluent
from treatment plants serving municipalities of
specified size to lakes of specified area, it be-
came necessary for the Madison Metropolitan
Sewerage District to provide for diversion of its
effluent in accordance with statutory provisions.
The Madison Metropolitan Sewerage District
operates a complete treatment plant serving a
population of about 135,000. The flow through the
plant normally averages about 20 million gallons
per day. Primary treatment consists of screen-
ing, grit collection, and sedimentation. About
one-fourth of the sewage receives secondary
treatment in a trickling filter, and the remainder
receives secondary treatment in an activated
sludge process. The combined final effluent from
the secondary processes is chlorinated before dis-
charge.
Engineers were retained to make preliminary
studies of possible diversion routes and to make
estimates of costs for diversion of effluent from
the Madison lakes. Fourteen separate routes were
considered and it was concluded that a route uti-
lizing Badfish River was the most practicable for
'several reasons, including the probable use of the
effluent for irrigation of lands adjoining a pro-
posed ditch leading to the river.
Badfish River was a small meandering stream
which flows through typical agricultrual lands in
Dane and Rock Counties. Many years ago, the
Dane County portion was straightened and widened
to serve as a drainage ditch to lower the water
table under adjoining lands. From the Dane-Rock
County Line to its junction with the Tahara River,
however, It has a substantially larger natural flow
and a correspondingly greater channel capacity.
Badfish River has an average slope of six feet per
mile, and portions of the stream have been con-
sidered as marginal trout water. Effluent from
the trickling filter treatment plant of the Village of
Oregon, Wisconsin, amounts to 65,000 gallons per
day and is discharged to Badfish River near its
headwaters.
The route chosen for the diversion necessitated
construction of five miles of 54-inch pipeline,
straightening and reconstruction of about four
miles of the drainage ditch, and straightening and
improvement of 10 miles of Badfish River to the
Rock County line. The improved ditch is 16 feet
wide and the improved river channel is 16 to 20
feet wide. From the Dane-Rock County line to its
junction with the Yahara River, the natural channel
has adequate capacity for handling the added
effluent flow without serious change in level, and
no reconstruction was necessary. The Yahara
River, after six miles, discharges into the Rock
River.
To restore dissolved oxygen to the effluent
after its passage through the pipeline, two cascade
aerators are provided in the new channel. One,
located at the discharge end of the pipe, is 35.5
feet wide and consists of a seven-step cascade
with a total fall of 6.5 feet; the second is 3,240
feet above the point of discharge into Badfish
River and is 30 feet wide with a five-step cascade
having a total fall of 5.5 feet. The new ditch
channel has a slope of 2.6 feet per mile while the
improved river channel has an average grade of
4.75 feet per mile for the first four miles and an
average grade of 8.05 feet per mile for the re-
maining six miles.
ORGANIZATION OF STUDY
The Madison Metropolitan Sewerage Commis-
sion and the Wisconsin Committee on Water Pol-
lution concluded that here an opportunity would be
afforded for study of the effect of sewage plant
effluent on a stream and proposed a joint study to
determine conditions before and after diversion of
effluent. Surveys were conducted at one-week
intervals during 1956 to determine stream charac-
teristics before diversion; and after diversion of
effluent to the Badfish River in December, 1958,
the surveys were repeated during 1959 at the
identical stations. Arrangements were also made
by the Madison Metropolitan Sewerage District for
installation of a U. S. Geological Survey stream
gauging station on Badfish Creek at a point 9.2
miles above its confluence with the Yahara River,
-------
Badfish River and Diversion
this point being about nine miles down from the
point of discharge of the 54-inch pipeline. Thus, a
continuous record of stream flow at this point on
the river is available.
The data reviewed in this report are based only
on the bi-weekly samples collected by the repre-
sentatives of the Committee on Water Pollution
and analyzed by them and by the State Laboratory
of Hygiene and represent the results of deter-
minations for a 26-sample period prior to diver-
sion.and a similar period subsequent to diversion.
Badfish River is approximately 16-1/2 miles
long, and, for the purpose of this study, three
sampling stations were selected. Station 1 is ap-
proximately one mile below the confluence of the
effluent ditch with the river; Station 4 is approxi-
mately four miles downstream from Station 1 and
in the immediate vicinity of the gauging station;
Station 8 is the farthest downstream station on
Badfish River and is eight miles below Station 4
(Figure 44).
MADISON MCTKOPOI.ITM
SCWCC
BADFISH CREEK
DIVERSION PROJECT
WISCONSIN
Figure 44. BADFISH CREEK DIVERSION PROJ-
ECT, WISCONSIN.
Three stations were chosen on the Yahara
River, one above the confluence with Badfish
Creek (Station 10) and two below this confluence
(Stations 9 and 14). The distance of the Yahara
River which is now affected by the effluent is 6.4
miles. On the Rock River, Station 15 is approxi-
mately two miles above the confluence of the Ya-
hara and Rock Rivers, and Stations 16 and 17 are
located six and ten miles, respectively, from this
confluence.
Chemical determinations were made in accord-
ance with procedures detailed in the Tenth Edition
of Standard Methods for the Examination of Water,
Sewage, and Industrial Wastes. Enumeratioa of
phytoplankton was carried out according to the
procedures outlined by Prescott.
The important chemical data and the phyto-
plankton were analyzed statistically (Table 40).
The mean and the 95% confidence interval were
calculated for 26 paired dates prior and subse-
quent to diversion for nitrogen, phosphorus, 5-day
B. O. D. and phytoplankton. The coefficient of
variation on these data normally do not appear
excessive, especially when the climatic and sea-
sonal influence is taken into consideration.
RESULTS OF STUDY
Physical Observations
As the effluent leaves the 54-inch pipeline, it
passes over a seven-step reaeration cascade and
enters a rather straight ditch with steep banks.
The first half mile of this ditch often carries a
blanket of detergent foam. For the next mile, the
banks are less steep, and within a year after di-
version was commenced, there was evidence of
some vegetation encroachment, principally round-
stemmed bulrush. Badfish River was dredged to
a bottom width of 16 feet for about four miles
and a width of 20 feet for six miles. This is a
great change from the original meandering stream
bed. Along with the changes created by physical
disturbance, there is a change in flow produced by
the introduction of slightly over 20 million gallons
per day of effluent. Prior to diversion, Badfish
River, at about its midpoint (Station 4) between its
source and confluence with the Yahara River, had
an average flow of 9.6 c.f.s. for the 2-1/2 years
in which records were kept. Following the diver-
sion, the flow averaged 43 c.f.s. for the summer
portion of the study with 32.8 c.f.s. being con-
tributed as effluent from the Madison Metropolitan
Sewerage District plant. In the unimproved stretch
of Badfish River, there was little gross physical
change noted.
Badfish River originally contained many riffle
areas with a bottom composed principally of small
rock and gravel. The bottom was, of course,
altered in the improved section, but remains a
coarse gravel over much of the area.
The effluent normally contains about 25 p.p.m.
of suspended solids and a substantial amount of
these solids has settled to build a sludge deposit
over most of the upstream portion of Badfish
River. In some areas, especially in small pock-
ets along the side of the stream, this deposit ap-
proaches six to ten inches in depth. In most of
the upstream region, as well as the ditch itself,
the sludge is of sufficient thickness to provide a
suitable habitat for a bountiful population of midge
larvae.
-------
120
ALGAE AND METROPOLITAN WASTES
Chemical and Bacterial Determinations
The organic nitrogen (Table 40) shows a size-
able increase at all stations in Badfish River fol-
lowing diversion. At Station 4, where stream
flow information was available, calculations show
a mean of 30 pounds per day of organic nitrogen
prior to diversion, and a mean of 286 pounds per
day following diversion.
la the Yahara River, there appears to be no
statistical difference between the samples col-
lected before or after diversion. There is an
indication, however, as shown at Station 10 for
the 1959 samples, that the organic nitrogen is less
in the Yahara River above the entrance of Badfish
River than it is below this confluence. The lower
value at this point can be attributed to diversion,
as the effluent formerly discharged through Lakes
Waubesa and Kegonsa and down the Yahara River.
There was no significant change in the organic ni-
trogen concentrations in the Rock River. Calcu-
lations show that the effluent contributed 732
pounds per day of organic nitrogen to the stream,
and that this was reduced to 286 pounds per day in
the Badfish by the time the flow reached Station 4.
The reduction was most likely due to settling of
the suspended solids to create sludge deposits in
the upstream region.
The inorganic nitrogen, as shown in Table 40,
Includes the total of ammonia, nitrite and nitrate
nitrogen. The 1959 samples indicate a 5-fold in-
crease in concentration. Again at Station 4, de-
terminations showed a mean of 110 pounds per day
before diversion and 3,153 pounds per day after
diversion, a more than 30-fold increase. The
effluent contributed 3,192 pounds per day or in-
organic nitrogen during the test period. Most of
the increase in the stream is attributed to the am-
monia nitrogen contributed in the effluent although
there was a slight increase in nitrite and nitrate
nitrogen. The inorganic nitrogen persists in so-
lution in the Badfish River, and the effect of its
presence is demonstrated in the Yahara River and
in the Rock River by the increase in concentration
in these rivers after diversion.
Soluble phosphorus is characteristically high
in sewage effluent as compared to natural drain-
age. Following diversion, Badfish River dis-
played a great increase in the soluble phosphorus
content. Consideration of soluble phosphorus pas-
sing Station 4 reveals 9 pounds per day in 1956
and 1,351 pounds per day in 1959. The effluent
discharged to Badfish River contributed 1,458
pounds of soluble phosphorus per day. The Yahara
River samples were all quite high in 1956 reflect-
ing the discharge from Lakes Waubesa and Kegonsa
which received the effluent that year. In 1956,
there was no significant difference between the
determinations at any of the three stations on the
Yahara River. In 1959, after diversion, stations
below the confluence of the Badfish River showed
a 100% increase in soluble phosphorus as com-
pared to the station on the Yahara River above the
confluence. The Rock River samples indicated a
similar differential between the station above the
confluence with the Yahara and the stations below.
High B. O. D. (biochemical oxygen demand)
values and low D. O. (dissolved oxygen) values
were encountered in Badfish River during the
post-diversion survey. Considering the summer
period (June 1 to October 1), there were 75 pounds
of B.O. D. per day and 475 pounds of D. O. per
day prior to diversion at Station 4. After diver-
sion for this same period in 1959, the water at
Station 4 was carrying 1,602 pounds per day of
B. O. D. and 900 pounds of D. O. per day, definitely
a D. O. deficit. The effluent during this period
contributed 3,724 pounds of B. O. D. per day. It
should be noted that in the four-mile stretch be-
tween Station 1 and 4, a substantial amount of the
contributed B. O. O. was removed. The Yahara
and Rock Rivers did not appear to be appreciably
affected by this B. O. D. -D.O. relationship.
The most probable number of coliform organ-
isms per 100 ml. was quite variable throughout
the course of the surveys (Table 40). The effluent
from the Oregon sewage treatment plant was not
chlorinated and did have an effect on Badfish River
prior to diversion, with readings of above normal
concentrations of colif orm organisms. Following
diversion, the MPN determinations for Badfish
River were higher than those recorded for 1956.
The influence of the effluent was also noticeable
in the Yahara River. On the Rock River, MPN
determinations were higher at all stations than the
determinations made in 1956 and can be attributed
to factors other than diversion.
Biological Determinations
The phytoplankton volume throughout the course
of the study displayed considerable variability as
would be expected (Table 40). However, the mean
volume, although showing an increase for the
larger rivers, showed no statistical difference
either between the three stations on a given river
or between the two periods of study for the same
station. It thus appears that a sizeable increase
in nutrients in a flow water situation has no sub-
stantial effect upon a volumetric production of
phytoplankton.
The phytoplankton volume of Badfish River was
generally lower than that of the Yahara River or
Rock River. Although there was little change hi
phytoplankton volume evidenced as a result of di-
version, the change was most pronounced at Sta-
tion 1 on Badfish River. There was an indication
of a volumetric reduction following diversion
which suggested inhibited growth. The blooms of
Euglena which were present before diversion at
-------
Badfish River and Diversion
121
Table 40. SUMMARY OF BIOLOGICAL AND CHEMICAL DATA BEFORE AND AFTER DIVERSION ON
BADFISH CREEK, YAHARA RIVER, AND ROCK RIVER - BASED UPON 26 BI-WEEKLY PAIRED DATES
EXTENDING FROM JUNE 6, 1956 TO MAY 22, 1957, AND MARCH 4, 1959 TO FEBRUARY 17, 1960.
Range
Pfaytoplankton2
Organic N.2
Inorganic N.2
Soluble P.2
B.O.D.*
D.O.2
PH
M.P.N. If 103)
_3t
1
4
8
10
9
14
IS
16
17
1
4
8
10
9
14
15
16
IT
1
4
8
10
9
14
IS
16
17
1
1
8
10
9
14
15
16
17
1
4
8
10
9
14
IS
16
17
1
4
8
10
9
14
IS
16
17
1
4
8
10
«
14
IS
16
17
1
4
8
10
9
14
15
16
17
1956
0.12-40.93
1.09-15.4}
1.35-16.74
0.04 -37.49
O.S3-24.S8
0.16-37.38
0.58-82.19
0.55-56.51
1.06-62.24
0.13-2.20
0.27-2.00
0.21-2.30
0.60-4.34
0.64-1.32
0.60-2.70
0.90-3.71
0.88-3.51
0.94-3.31
1.98-5.53
1.73-3.64
1.38-4.49
0.09-1.24
0.10-2.18
0.10-1.51
0.09-0.68
0.09-0.91
0.05-1.13
0.30-1.56
0.10-0.30
0.01-0.12
0.46-1.30
0.16-1.30
0.40-1.28
0.01-0.19
0.01-0.41
0.02-O.44
1.8-7.7
0.8-5.4
0.6-8.8
1.3-14.5
1.8-15.7
1.5-14.3
3.5-1T.1
2.7-14.3
3.0-15.4
S.1-1S.4
7.8-15.9
6.6-16.7
4.2-21.6
8.9-19.3
5.6-1S.4
4.4-20.9
6.6-18.J
5.3-18.1
7.5-8.2
7.7-«.e
7.7-S.8
8.0-8.9
8.2-9.8
8.1-9.6
8.3-9.2
7.8-9.7
7.9-9.4
7-540
0.5-160
0.4-240
0.02-18
0.08-35
0.05-54
0.2-35
0.2-17
0.2-54
1959
0.37-10.16
0.55-12.45
0.22-13.99
0.11-56.87
0.30-30.71
0.15-52.53
0.53-57.55
1.03-62.20
0.23-53.67
1.33-11.74
0.83-9.50
1.03-7.10
0.00-2.44
0.94-3.34
0.60-2.65
0.94-3.54
0.74-3.34
0.84-3.24
13.4-21.1
10.0-18.7
7.2-17.6
0.09-2.82
1.09 til. 31
0.80-4.64
0.10-3.32
0.15-3.10
0.19-3.09
5.5-12.0
4.4-7.3
3.0-8.4
0.23-1.5
0.56-6.4
0.58-2.6
0.01-0.68
0.15-0.94
0.13-1.40
4.1-39.4
3.1-55.8
3.3-38.4
1.5-7.70
2.5-19.7
2.4-15.4
2.2-15.3
2.1-12.9
2.4-10.2
0.1-8.9
1.7-10.7
2.2-11.1
5.9-17.5
3.9-16.7
2.9-15.7
3.0-25.8
6.4-22.0
6.1-20.5
7.4-4.1
7.S-3.2
7.7-8.1
7.7-9.2
7.0-8.9
7.7-8.9
7.7-9.1
7.6-9.3
7.8-9.2
3.3-790
4.9-350
0.8-1,200
0.2-49
0.8-130
1.3-430
0.5-210
0.3-170
0.3-160
Heu
±95% Confidence Interval
1556 1555
6.53*4.04
3.98*2.58
6.35*1.54
10.62*4.19
7.59*2.55
9.56*3.79
25.59*8.59
25.43*7.07
24.64*7.61
0.73*16
0.6U.17
O.S4*.18
1.6U.41
1.30*.19
1.54*.27
1.99*. 25
1.86*. 29
1.86*.2S
3.73*.35
2.65*.08
2.34*.31
0.471.14
0.66*. 21
0.55*. 16
0.27*.07
0.29*.08
0.30*. 10
1.07*.12
0.19*.02
0.08*.01
0.94*.10
0.83*.12
0.8U.10
0.05*.02
O.lSt.04
0.17*.04
3.63*.60
2.11*2.30
2. 12*. 64
6.14*1.75
4.90*1.24
5.90*1.50
7.61*1.38
7.18*1.26
7.00*1.42
2.74±.95
3.24*1.78
4.86*1.70
12.70*6.34
9.09*3.10
11.63*5.49
18.37*6.15
21.36*6.96
19.50*6. 5«
4.13*1.14
J.22..89
2. 59*. 66
1.29*.25
1.49*.23
1.58*. 23
1.77*. 27
1.78*. 29
2.14*.56
17.98*2.07
15.17*.91
ll.42t.97
0.62*. 25
3.53*1.11
2.00t.37
1.34*.47
1.42..41
1.43*. 39
8.224.64
S.96*.37
5.22*.64
0.8g*.12
1.86*. 52
1.22*.49
0.22*.01
0.46*0
0.48*. 02
21.01*8.13
17.25*5.59
13.99*4.44
3.89*.89
5.86*1.44
5.08*1.13
5.23*1.17
5.46*1.19
4.95*.8»
Standard
Deviation
1956
10.03
3.20
3.85
10.17
6.73
9.41
20.38
17.15
18.08
0.40
0.41
0.42
0.96
0.48
0.66
0.60
0.71
0.60
0.87
0.21
0.77
0.34
0.52
0.40
0.17
0.21
0.26
0.33
0.05
0.03
0.26
0.33
0.28
0.05
0.11
0.10
1.46
5.29
1.56
4.23
3.08
3.70
3.46
3.09
3.51
1959
2.28
2.70
4.06
14.35
7.51
13.32
14.91
16,89
15.93
2.69
2.17
1.59
0.63
0.56
0.54
0.65
0.69
1.42
2.03
2.20
2.35
0.61
2.73
0.93
1.18
1.01
0.97
1.51
0.90
1.56
0.33
1.28
0.41
0.02
0.01
0.03
18.80
13.56
10.75
2.17
3.55
2.79
2.91
2.94
2.21
Coefficient
of Pounds Per Day
Variation (%) Range Mean
1956
153
80
60
95
83
98
82
67
73
55
67
77
60
37
43
30
38
32
23
8
33
72
79
73
63
72
86
30
25
37
28
40
35
95
61
63
40
250
74
69
63
63
45
43
50
1959 1956
83
83 56-791
83
112
82
114
81
79
81
65
67 13-71
61
49
38
34
37
39
66
11
IS 89-143
21
98
77
46
88
71
68
18
15 7-12
30
38
69
34
8
3
6
89
79 39-113
77
56
61
55
56
52
45
368-636
1959 1956 1959
247-1,435 259 622
208-147 30 286
2,171-4,246 110 3,153
996-1,701 9 1,351
755-2,333 75 1,602
413-1,749 475 904
1. Pounds of material per day on 9 bi-weekly paired dates (June 1 - October 1) for Station 4.
Flow in c.t.s. In 1956 ranged from 8.0-10.0 with a mean of 8.7; in 1959 the flow In c.f.s.
ranged from 40.0-48.0 with a mean of 43.0.
2. Parts per million.
-------
122.
ALGAE AND METROPOLITAN WASTES
Station 1 did not appear in the 1959 samples. Os-
cillatoria sp. did appear in the samples following
diversion and quite possibly came from the rather
extensive growth of this genus over the bottom
deposits. The principal diatoms occurring in the
1956 samples consisted of Navicula. Nitzschia.
Gomphonema. and Synedra. In the 1959 samples,
populations were dominated by species of Navicula
and Nitzschia with other genera appearing only
occasionally and in very small numbers. On the
occasions when green algae appeared, these con-
sisted of Chlamydomonas and Closterium. both in
the 1956 and 1959 samples. In general, the 1959
samples, especially at Station 1, appeared more
heterogeneous to class and more homogeneous to
genera than those collected in 1956.
The tendency toward inhibited growths was ap-
parent although much reduced at Station 4, follow-
ing diversion. In 1956, the greatest diatom volume
appeared in late summer and consisted principally
of Navicula with several other genera represented
in varying numbers. The 1959 samples did not re-
veal as great a volume nor as great a variety of
species, but did indicate a more equal represen-
tation between the diatoms, blue-green algae, and
green algae in the phytoplankton.
At Station 8, the principal constituents of the
diatom population prior to diversion were Navicula
and Nitzschia with Synedra. Cyclotella. Gom-
phonema. and Cocconeis contributing to the total
volume regularly. In 1959, Navicula and Nitzschia
were the principal constituents of the diatom popu-
lation, with the other genera appearing only
occasionally and contributing less to the total
volume. Green algae and blue-green algae ap-
peared occasionally in the 1959 samples and not in
the 1956 samples, although the total plankton
volume was rarely affected by these occurrences.
The Eugtena. group appeared more often in the 1959
samples', but they, too, seldom affected the total
phytoplankton volume.
The phytoplankton volumes on the Yahara River
stations reveal little change subsequent to diver-
sion. The diatom population at Station 10 above
the confluence with Badfish River prior to diver-
sion was dominated by Melosira with species of
Navicula. Nitzschia and Cyclotella appearing regu-
larly but in lesser numbers. After diversion, the
same genera of «M«,tnmp were encountered, but
the volume became more equally proportioned
among those present and no particular genus pre-
dominated. Blue-green algae appeared in both the
1956 and 1959 samples with Anacystis and Aphani-
zomenon predominating. The most commonly
recorded genera of green algae were Scenedesmus
PIMJ Anirifftrodes***"^, c^attiTdomonag j. and Coe-
lastrum in both the 1956 and 1959 samples. On
only two occasions in the spring and early summer
of 1959 did the green algae volume exceed the
diatom volume.
Stations on the Yahara River below its conflu-
ence with Badfish River generally revealed a
greater proportionate volume of green algae fol-
lowing diversion. Blue-green algae at these sta-
tions were noted only occasionally and contributed
little to the total volume. The diatom population,
especially in the summer months, appeared simi-
lar both before and after diversion. In mid-winter
of 1959, a bloom of Cyclotella approached a popu-
lation of 7,000 organisms per ml. and extended
over a period of six weeks. The species were
very small and contributed little to the total
volume.
The phytoplankton in the Rock River revealed
no detectable difference between stations in a
given year. The prominent genera in both 1956
and 1959 were Stephanodiscus. Melosira. and
Cyclotella. Navicula and Nitzschia appeared con-
sistently scattered but rarely exceeded one p. p.m.
in volume. Cyclotella was a major constituent of
the population during the entire year. During
December, 1958 and January, 1959, it was the
principal genus found, and populations at this time
approached 30,000 organisms per ml.
All stations on the Rock River revealed a sub-
stantial volume of blue-green algae during all ex-
cept the winter months. This consisted principally
of Anacystis and Aphanizomenon. Green algae
appeared more prominent in the 1959 samples,
particularly in the spring and summer months.
Volumes of green algae exceeded 10 p. p. m. only
rarely. The principal constituents were Clos-
terium and Coelastrum in 1956 and Coelastrum in
1959. Scenedesmus appeared regularly but the
volume seldom exceeded 2 p. p. m. in any particu-
lar sample.
The organisms which dwell upon and within the
bottom, deposits were studied at seven separate
stations on four different dates in Badfish River.
Pre-diverslon surveys were conducted on August 1,
1956 and March 1, 1957, whereas post-diversion
surveys were conducted on September 17, 1959
and December 1, 1959. Prior to diversion at Sta-
tion 1, the stream was 3 to 6 feet wide and ap-
proximately 6 inches deep. It gradually increased
in width downstream until a width of around 30
feet was attained before the confluence with the
Yahara River. The depth at this point, however,
was still relatively shallow, varying between 6 and
18 inches. The bottom material at the sampling
stations consisted principally of rock and coarse
gravel and, at some points, gravel mixed with
sand. Submerged aquatic vegetation was abundant
prior to diversion and, at some points, streamers
-------
Badfish River and Diversion
123
of filamentous algae were attached to the sub-
merged vegetation. In September, 1959, following
diversion, the improved portion of Badfish River
still maintained a coarse gravel bottom and, in
the Station 1 area, the stream was already choked
with submerged vegetation. In the downstream
areas, this vegetation appeared to be less dense
than in 1956. Long streamers of filamentous
green algae (Stigeoclonium and Rhizoclonium).
some of which were estimated to be 50 feet in
length, were attached to bottom materials at
numerous locations. In the upper areas of the
stream, there was a green blanket of Osclllatoria
covering the bottom. Sludge had deposited along
the edge of the stream and covered portions of the
vegetation. A definite sewage odor was present in
the Station 1 area in September, and this odor ex-
tended the full length of Badfish River in Decem-
ber, 1959. Much of the stream bottom was covered
with a slimy mat of the blue-green algae, Oscilla-
toria. and, especially in the December survey,
much of the vegetation was covered with a prolific
growth of a stalked protozoa belonging to the
family, Epistylidae. These formed a gray mass
not unlike a dense growth of fungus.
The degradation of the stream following diver-
sion is apparent when one examines the community
of biological life living upon and within the bottom
materials. Prior to diversion, between 10 and 14
different invertebrate species were recovered
from each of the samples collected. Following
diversion, the number of species was reduced to
about five.
Prior to diversion, also, a balanced community
of intolerant and tolerant organisms was observed.
At nearly every station, caddis fly larvae (Cheu-
matopsyche and Hydropsche), mayfly nymphs
(Baetis and Caenis). and riffle beetle larvae were
found in association with cranefly larvae, horsefly
larvae, scuds, and miscellaneous midges. Very
tolerant forms such as sludge worms (Tubificidae)
were also found, but occurred in very low num-
bers. In some locations, the intolerant caddis
fly larvae formed the bulk of the total population.
Following diversion, all stations in the ditch
and in the improved portion of Badfish River sup-
ported a bottom-dwelling population comprised of
sludge worms (Tubificidae) and at least three
species of very tolerant midge larvae (Tendipes
plumosus, T. tendipediformis, and T. decorus).
These are all considered to be very tolerant or-
ganisms and were found to be living in the sludge
deposits on the bottom and along the sides of the
stream. Near the lower end of Badfish River in
the unimproved portion, tolerant and very tolerant
bottom-dwelling organisms predominated. Occa-
sionally, an intolerant form was observed, but
this was only one among many of the more tolerant
forms.
SUMMARY
1. Studies have been conducted on the bio-
logical and chemical effects resulting from the
diversion of approximately 20 million gallons a
day of effluent from the Madison, Wisconsin,
Metropolitan Sewerage Commission Treatment
Plant to a small stream which originally had a
flow of 9.6 cubic feet per second. This stream,
Badfish River, discharges into the Yahara River,
and the Yahara River into the Rock River. The
effects upon all three river systems were investi-
gated.
2. In addition to physical and biological ob-
servations, and bottom fauna studies made at in-
tervals, 26 bi-weekly samples were collected and
analyzed from selected stations before and after
diversion for chemical and phytoplankton deter-
minations.
3. Considering that 10 of the 14.5 miles of
stream were improved to a bottom width of 16 and
20 feet, that the flow was increased nearly five-
fold, and that a deposition of solid materials
created substantial sludge deposits in some areas,
a tremendous physical change, especially in the
upper regions, was exerted upon Badfish River
as a result of diversion.
4. The water chemistry of Badfish River es-
pecially responded to diversion with substantial
increases in organic nitrogen, inorganic nitrogen
(influenced principally by ammonia nitrogen),
phosphorus, and B. O. D. The dissolved oxygen
was reduced to a critical level many times
throughout the summer, and a D. O. deficit of 700
pounds per day existed at Station 4 during this
period.
5. Phytoplankton populations were of substan-
tially the same concentration between the three
stations on a given stream and between the two
periods of study for similar stations on the same
stream, but were greater in the Yahara River
than in Badfish River, and greater in the Rock
River than in the Yahara River. There was an
indication of a population depression following di-
version at the upper stations on Badfish River and
a difference in genera encountered between the
pre-and post-diversion samples.
6. Submerged aquatic vegetation was abundant
prior to diversion and, already in 1959, had be-
come abundant in the dredged portion of the River.
Perhaps it is yet too early to judge, but the sub-
merged plants do not now present a problem.
Long streamers of filamentous algae were attached
to plants and bottom materials at numerous loca-
-------
124 ALGAE AND METROPOLITAN WASTES
tions. A blanket of Oscillatoria covered much of containing few species and only very tolerant
the bottom of the upper Badfish River. sludge worms and midges following diversion.
7. A study of bottom organisms indicated 8. The benthos in Badfish River exhibited a
severe stream degradation following diversion. much greater response than the phytoplankton to
Stream biota changed from a balanced population the addition of nutrients, suspended solids, and
containing several species and many intolerant B. O. D. contained in the effluent of the Madison
organisms, prior to diversion, to a population Metropolitan Sewerage District Treatment Plant.
REFERENCES
Mackenthun, K.M., L. A. Lueschow, and C. D. Standard methods for the examination of water,
McNabb. 1960. A study of the effects of diverting sewage and industrial wastes (1955). Amer. Publ.
the effluent from sewage treatment upon the re- Health Assoc., Inc., 1790 Broadway, New York 19,
ceiving stream. To be published in Volume 49 of N.Y., 10th Ed.
the Trans. Wis. Acad. of Sci., Arts and Letters. Woodburn> James G. 1959. Outfall around the
Madison lakes. Water and Sewage Works 106 (11):
497-500.
-------
Spray Removal of Nutrients in Detroit Lakes
125
SPRAY IRRIGATION FOR THE REMOVAL OF NUTRIENTS IN SEWAGE TREATMENT
PLANT EFFLUENT AS PRACTICED AT DETROIT LAKES, MINNESOTA
WINSTON C. LARSON
Consulting Engineer,
Detroit Lakes, Minnesota
Detroit Lakes is a resort city located in a
recreational area of 412 lakes within a radius of
25 miles.
The first sewage treatment plant at Detroit
Lakes was erected in 1929, though as early as
1909 there was in existence a septic tank for the
treatment of waste from a limited sewage system.
The 1909 septic tank drained to the same water
course as the 1929 plant addition. The 1929 plant,
by reason of the growth of the city, was enlarged
and improved by additional construction in 1941.
The 1950 census gives Detroit Lakes a population
of 5, 734. The plant now treats on an average of
750,000 gallons of waste dally. The present
sewage treatment plant consists of a coarse
screen, a fine screen, a primary high-rate trick-
ling filter, an intermediate settling tank, a sec-
ondary low-rate trickling filter, a final settling
tank, and a chlorination tank. Sludge is digested
and removed to sludge drying beds and sludge
lagoons.
The effluent from the plant was originally dis-
charged into a ditch which runs through what was
formerly the easterly arm of St. Clair Lake, then
into Lake St. Clair proper, and from there into
St. Clair Ditch, which empties into the Pelican
River. The Pelican River, after its confluence
with St. Clair Ditch, empties into Muskrat Lake
which empties into Lake Sally. Lake Sally is a
navigable lake and, as the water flows, lies be-
tween 4 and 5 miles from the sewage treatment
plant. It is one of the recreational lakes in the
vicinity of Detroit Lakes in the so-called Pelican
River chain. The recreational uses of the lake
are swimming, boating, and fishing; and, with its
wooded shores, it has attracted many residents
for summer living. It appears that until the year
1947, there was no particular complaint about the
algal bloom or weed growth in Lake Sally, though
there were statements made that there had been a
gradual increase in the growth of lake vegetation
since 1940. The bloom is due to a large increase
in the number of algae in the upper part of the
water.
The summer of 1947 was unseasonably warm
with a low amount of precipitation, and the amount
of algae in the lake was particularly extensive, and
the odor, at times, extremely offensive. On
August 18, 1947, there was a fish kill in the lake.
A large number of dead fish were lodged on the
shores, and that, together with the great amount
of algae in the lake, some of which were washed on
the shore, caused a great deal of disturbance and
alarm among the summer residents. Bathing in
the lake was unpleasant on account of the algae
and recreational use of the lake except for fishing
was greatly reduced.
Complaints were made by the residents to the
City of Detroit Lakes, alleging that the source of
the trouble was the effluent from the sewage treat-
ment plant;, while there was no dissatisfaction with
the effluent from a sanitary viewpoint, it was al-
leged to be rich in nutrients, particularly phos-
phorus and nitrogen, which were conducive to the
excessive growth of algae and weeds in Lake Sally.
STUDY BY THE DEPARTMENT OF HEALTH AND
CONSERVATION
As this problem was an entirely new one for the
city officials, it was referred to the Minnesota
Department of Health and Conservation for inves-
tigation and study. Sampling stations were set up
at various points between the waste treatment plant
and Lake Sally and at other points atwater courses
in the area which were not affected by sewage
treatment plant effluent. Reports issued in 1948
and again in 1951 indicated that some artificial
fertilization was due to the effluent from the
sewage treatment plant, although test data indi-
cated a reduction of 90 per cent in phosphorus
between the sewage treatment plant and Lake
Sally. In the summary and conclusion of the 1951
report, the following was stated:
"To obtain a proportionate evaluation of the
effect of the sewage treatment plant effluent on the
present rate of enrichment of Lake Sally, the fol-
lowing program is offered:
"(a) A continuous study should be carried out,
consisting of at least weekly sampling and flow
estimation at all significant stations for at least
a year.
-------
126
ALGAE AND METROPOLITAN WASTES
"(b) A sanitary survey of all lakeshore and
watershed watershed property should be made.
"(c) The nutrient content of ground waters and
their effect on Lake Sally should be adequately
studied.
"(d) -The relative algal productivities of Lakes
Sally, Monson, Mellissa, and Detroit Lakes should
be scientifically determined over at least one
year's time.
"(e) The kinds and amount of nutrients from
all sources should be evaluated according to the
nutrient requirements of plant life in general and
of blue-green algae in particular."
OTHER STUDIES
During the period from 1948 to 1951, Investi-
gations were made of studies and reports on simi-
lar conditions throughout the nation. Information
and data were very limited. We found that
Madison, Wisconsin, had a similar condition and
that the University of Wisconsin was carrying on
various studies on the nutrition of algae, methods
of control, removal of nutrients from sewage plant
effluent, etc. At that time, no practical solution
was available.
EXPERIMENT
coNsm
AT THE UNIVERSITY OF WIS-
On February 13, 1950, W. L. Lea and G. A.
Rohlich Issued a paper on "The Removal of Phos-
phates from Sewage Treatment Plant Effluent."
Their data showed that approximately 96 per cent
of the soluble phosphate phosphorus was removed
from effluent employing alum as a coagulant hi the
amount of 200 parts per million. Sixty-two per
cent of the nitrogen containing organic compounds
was also removed from the effluent. The data
showed that addition to the treatment plant effluent
of 200 parts per million of alum produced no sig-
nificant change in the concentration of free am-
monia, nitrites or nitrates.
The data presented In that study showed that,
in order to remove 95-99 per cent of the soluble
phosphates from a sewage plant effluent containing
5.0 parts per million of phosphates, approximately
6-10 times as much coagulant is required as is
ordinarily used in the clarification of surface
water supplies employed by a community as a
source of potable water. Obviously then, the cost
of chemicals required to remove the phosphates
from, one million gallons of effluent would be 6-10
times the chemical cost ordinarily incurred in
treating one million gallons of a surface water
supply.
REPORT OF THE MINNESOTA DEPARTMENT
OF HEALTH
The Minnesota Department of Health, Division
of Water Pollution Control, issued a "Report on
the Experimental Removal of Phosphorus from
Sewage Plant Effluents with Lime" in April, 1950.
Experiments were conducted at the Detroit Lakes
Treatment Plant.
The summary and conclusions stated that:
1. Approximately 700 ppm slaked lime or
530 ppm of unslaked lime were required to remove
80 per cent of the total phosphorus from the ef-
fluent of the low rate filter at Detroit Lakes.
2. The existing facilities did not lend them-
selves readily to full scale trial of the efficiency
of the lime method, and It Is estimated that the
removal of phosphorus could be increased and the
amount of chemical used decreased with proper
mixing and efficient flocculation and settling of the
treated waste.
3. The volume of sludge produced would be
approximately three times the volume without the
use of lime.
4. Under the conditions existing during the
survey, the chemical costs only for the year-
around treatment with unslaked lime would be ap-
proximately $7, 600. Assuming a sewage flow of
0.5 mgd, changes and additions would have to be
made to the treatment plant units and equipment in
order to provide chemical treatment and additional
operating personnel would be needed.
5. The use of this method of phosphorus re-
moval would not be recommended for this plant on
the basis of this study.
SUGGESTED SOLUTIONS
Another suggested solution to the problem was
to pond the effluent in large gravel pits to the
north of the city. This possibility was ruled out
as it is believed that the outwash gravel in the pits
was directly connected with aquifers that supplied
wells within the city.
A non-recreational lake to the north of the city
was also studied with the possibilities of using it
as a ponding reservoir. This lake had no outlet
and its level is evidently maintained by surface
run-off. However, studies indicated that the in-
troduction of effluent would raise the water level
approximately 6 inches per year. This would
cause flooding of additional land and consequently
lead to trouble and possible litigation.
-------
Spray Removal of Nutrients In Detroit Lakes
127
LITIGATION
All actions of the city council and studies and
reports by public agencies were open for public
review and comments. What originally was the
theory of a few, under proper influence, became
the conviction of many. It was finally concluded
by certain of the cottagers and their representa-
tives that fantastic sums of money could be col-
lected. Civil action was served on the City of
Detroit Lakes on or about August 22, 1951, and
additional ones shortly thereafter. In these actions
the plaintiffs asked for a permanent injunction
against the City of Detroit Lakes, together with
money damages. A total of 26 claims were even-
tually filed totaling $258, 500.
In the decision handed down by Gunnar H. Nord-
bye of the United States District Court, District of
Minnesota, Sixth Division, he stated that he could
not assume to speculate or conjecture the causes
of any excessive growth of algae or weeds in Lake
Sally, when the weight of the evidence establishes
that the present data were insufficient to form any
definite conclusions thereon. The evidence did not
reflect any other practical method which would
reduce the nutrients in the effluent more effec-
tively than the present system.
The case was dismissed, without prejudice to
the plaintiffs' right to apply again for equitable
relief and damages In the event sufficient data and
evidence exist to justify the granting of such re-
lief.
IRRIGATION FOR THE DISPOSAL OF EFFLUENT
As the citizens of Detroit Lakes were interested
in protecting the recreational lakes in their area
from possible damage, the City Council Instructed
our firm to Investigate further possible means of
preventing the effluent from reaching these waters.
An Investigation was made of a 55-acre wooded
knoll lying directly west of the treatment plant.
Borings Indicated that gravel lay to a depth of 12
feet. A contour map was prepared of the area.
On the west and east side of this knoll, there exist
drainage ditches which converge to the south of
the knoll and enter St. Clair Lake. We were quite
confident that the porosity of the soils and trans-
piration by the trees would dispose of an enormous
amount of water. Laboratory tests indicated that
phosphorus would precipitate In filtering through
the soils. Growth of vegetation would aid in the
removal of nitrates. Oxidation of organic wastes
would take place.
Factors which were unknown and had to be found
out by experimentation were:
1. Soil permeability and irrigation rates
2. Effect on vegetation
3. Odors
4. Operating conditions during the winter
months
5. Effect on ground water
As there were a number of factors which were
unknown and could cause failure of the project, we
proposed to proceed slowly. The initial installa-
tion included a turbine pump with a capacity of
600 gallons per minute against a total pumping
head of 110 feet and driven by a 20 horsepower
motor. This unit was installed in the chlorination
tank and made automatic with a float switch. A
weir was installed in the outlet to the chlorination
tank and set at an elevation approximately six
inches above the high water level of the float
switch. This would permit an automatic by-pass
of effluent in the event of a surge greater than the
pump capacity or in the event of pump failure and
still provide for practically 100 per cent of the
effluent going to the irrigation system during nor-
mal operation. The pump was equipped with a
check valve, gate valve and a propeller type main
line meter.
The initial piping system consisted of galvan-
ized steel irrigation pipe with 29 sprinkler heads.
Sizes and lengths of piping were as follows: 1312'
of 8", 288' of 4", and 2304' of 3".
Sprinkler heads were those manufactured by
the Skinner Irrigation Company: 1" utility with
7/32" main nozzle and 3/16" secondary nozzle.
These were to have a sprinkling radius of 45 feet
at 30 pounds of pressure and discharge 12. 3 gal-
lons per minute each.
Pipes were laid on the surface of the ground
and at a gradient so as to be self-draining. The
eight-inch main was provided with a two-inch
valved tap at the low point which would be left
open as desired during cold weather so as to pro-
vide drainage when the pump stopped. The 1, 300
feet of eight-inch pipe were covered with hay and
weighed down to help retain the water tempera-
ture. End caps of the laterals were tapped to pro-
vide drainage and prevent freeze-up during periods
of pump stoppage. Pipes were set on blocks where
necessary to provide the gradient. Trees were
removed only where necessary to provide for
reasonable pipe alignment.
Equipment and materials for the initial instal-
lation, in place, cost approximately $6, 500. Cost
of land was $3,000.
-------
128
ALGAE AND METROPOLITAN WASTES
A test well was installed in the irrigation area
and a sample of the ground water was taken before
operation began. The static level from the top of
the test well was measured and found to be 11' -
10-1/2". Operations began on October 14th, 1955.
Ground water analysis showed the following in
ppm:
Total hardness 300
Alkalinity (M.C.) 310
Chlorides 9
Nitrate nitrogen 1
Nitrite nitrogen NF
Ammonia nitrogen 0.19
Organic nitrogen 1.4
Total phosphorus 0.6
Total nitrogen 2.6
The average of samples of sewage treatment
plant effluent showed a pH of 7.4 and the following
analysis in ppm:
Suspended solids 34
B.O. D. (5 day) 33
Soluble phosphorus 7.8
Total phosphorus 9.1
Nitrate nitrogen 2.9
During the month of March, 1956, effluent was
applied to a 4-acre tract at the following rates:
Effluent pumped.
Pumping time in 31 days
Gals/Acre/Day
Gallons per acre foot . .
Total gallons per foot,
4 acres
Total feet of Irrigation .
Total inches of-
irrigation
Sprinkling rate
Sprinkling period daily .
21,547,500 gals.
(695, 080 G. P. D.)
651 hours
173,470
325,829
1,303,400
16.5 per month
198 per month
0.3 inches per hour
20 hours
The flow was 360 gallons per minute through
the sprinkler heads and 190 gallons per minute
through drains during this period of the year. At
this high rate of application, we found that during
the summer months, vegetation would not grow in
the sprinkled areas and a number of trees died.
Some washing and ponding occurred but no objec-
tionable odors were present.
In the fall of 1956, the irrigation area was
doubled in size and a 6-inch main installed to feed
the 4-inch and 3-inch laterals. Valves were in-
stalled on each lateral so that one branch could be
shut off "at a time for maintenance of nozzles and
providing rest periods for each area served by the
lateral.
In a four year period, 994,426,300 gallons of
effluent have been pumped through the irrigation
system during a total pumping time of 26,871
hours for an average rate of 617 gallons per
minute. The pump operates approximately 20
hours per day.
The static water level in the test well on No-
vember 21, 1958, was 8' - 9-1/2" having raised
3'-l" during the four year period. Analysis of the
sample of ground water taken November 21, 1958,
was as follows in ppm:
Total hardness ................ 420
Alkalinity (M.O. ) .............. 320
Chlorides ................... 130
Nitrate nitrogen ............... 31.0
Nitrite nitrogen ............... NF
J ammonia nitrogen , ,
*'2
w
N2l organic nitrogen
Total phosphorus .............. 2.9
Total nitrogen ................ 33.2
COST OF OPERATION
Cost of operation of the system is largely power
costs. At its designed condition, the pump motor
will require 0.47 kilowatt per 1,000 gallons
pumped. This would be an increase in power con-
sumption of 11,000 to 12,000 K. W. per month for
an average of 24,000,000 gallons. Power meter
readings during the years 1954 through 1959 con-
firm this figure. At two cents per kilowatt, the
cost of power for irrigation by this method is
slightly less than one cent per 1,000 gallons.
CONCLUSIONS
Limited personnel, equipment, and finances do
not permit the gathering of sufficient data to
thoroughly weigh the effects of the system insofar
as nutrient removal is concerned. It is our opin-
ion that results are being obtained chiefly through
visual observance of the water course and limited
test data. No doubt a large amount of the organic
wastes are being oxidized in the soil.
The system has definite value as an additional
unit to the treatment works.
If normal vegetation were to be grown, five feet
of irrigation water a year would perhaps be the
maximum allowable, considering the limited grow-
ing season of our area. If this rate were used to
dispose of 287,000,000 gallons of effluent per
year, approximately 200 acres would be required.
This would require frequent moving of irrigation
pipe and, consequently, additional personnel. Such
an extensive system would be very difficult to
operate during the winter months. H the higher
rates can be maintained in a smaller space without
-------
Spray Removal of Nutrients in Detroit Lakes
129
ill effects and without regard to the growth of
vegetation, forced irrigation appears to be the
most economical method to waste effluent in this
area, where rainfall is quite adequate.
Experience has shown that with proper pro-
visions, an irrigation system can be operated
during the winter months without difficult main-
tenance problems, if the object is chiefly to dis-
pose of water.
If the topography and soil characteristics are
satisfactory, we believe that a "ridge and furrow"
system could be used quite well as a method for
inducing the effluent into the soils. The operating
costs could be reduced considerably over spray
irrigation as it would not be necessary to pump
against a high head; however, the benefits gained
by aeration and evaporation in the overhead spray
irrigation system would be lost. Again, more
research and investigation is necessary.
-------
130
ALGAE AND METROPOLITAN WASTES
CHEMICAL, METHODS FOR THE REMOVAL OF NITROGEN AND
PHOSPHORUS FROM SEWAGE PLANT EFFLUENTS
GERARD A. ROHLICH
Hydraulic and Sanitary Laboratories
University of Wisconsin, Madison, Wisconsin
As part of the program of lakes and streams
investigations at the University of Wisconsin,
studies have been carried out on the removal of
plant nutrients from sewage and sewage effluents.
In particular, attention has been given to methods
for the removal of nitrogen and phosphorus, es-
sential elements which are usually considered of
principal concern in supporting algal blooms. In-
asmuch as other contributors to this seminar will
discuss removal of nutrients by biological growths,
this paper will be confined to a discussion of
chemical methods for the removal of nitrogen and
phosphorus from sewage and sewage effluents.
Rudolfs (1947) has reported on the quantities of
phosphates in sewage and sewage effluents. Ob-
viously the amount of phosphate will vary depend-
ing on the quantity and composition of industrial
wastes present. In a report on the removal of
phosphorus from sewage plant effluent, Owen's
(1953) analyses of samples of domestic sewage
from communities in Minnesota, with populations
varying from 1200 to 940,000, showed that the raw
sewage of those communities contained 1.5 to 3.7
grams of phosphorus per capita per day, with a
median of 2.3 grams. His study showed also that
the amount of phosphorus removed in conventional
sewage treatment processes varied from 2 to 46
per cent. The higher removals generally were
found in plants with secondary treatment and in
those which showed the highest 5-day B.O. D. re-
ductions.
Following laboratory studies, Owen (1953)
carried out an investigation at a municipal plant
consisting of fine screens, high-rate trickling
filter, secondary clarifier, low-rate trickling
filter, and final clarifier. Slaked lime was added
to the influent of the final settling tank in a con-
centration of 545 ppm CaO, and the results indi-
cate that the total phosphorus was reduced from a
concentration of 7.4 ppm to 1.7 ppm indicating a
removal efficiency of 77 per cent. He estimated
that the chemical costs for treatment with un-
slaked lime would be approximately $7,600 when
treating a sewage flow of 0.5 mgd. His laboratory
studies indicate that with the same dosage, a pH of
10.9 resulted, and the phosphorus concentration
was reduced from 6 ppm to 0.3 ppm, a reduction
of 95 per cent. In the laboratory studies, a set-
tling period of one hour was used. He further
showed that when the settling period was increased
to 18 hours, the phosphorus was reduced to .015
ppm, a reduction in excess of 99 per cent. Owen
concluded that the plant scale tests would have
more closely approached the laboratory results if
adequate mixing and efficient flocculation could
have been obtained. He found that the sludge pro-
duced was approximately three times the* volume
ordinarily handled at the plant.
Graphite Anode.
C"
Sea Water-,
C^>
Development of .- ,,
Chlorine Gas I/
1
.-^»
^
F-
naj
-i-
£-•
/
v^Scum
...Diaphram
-/Iron Cathode
.Development of
< -sodium hydroxide,
_' magnesium hydroxide
and hydrogen gas.
r- Sewage + Sea Water
Figure 45. PRINCIPLE OF DR. E.F0YN'S DIA-
PHRAGM CELL.
Dr. Ernst Ffyn at Oslo, Norway, developed a
process referred to as the Electrolitic Sewage
Purification Method. The principle involved in
Dr. Fjrfyn's method is shown in Figure 45. A di-
vided container is equipped with electrodes con-
nected to the negative and positive poles of a
battery. One portion of the chamber contains sea
water and the other sewage mixed with approxi-
mately 10 to 15 per cent sea water. Chlorine is
developed at the graphite anode and hydrogen and
the alkali in the chamber containing the iron cath-
ode. In this way, the chemical conditions neces-
sary for precipitation of the phosphorus are
established. The phosphate is adsorbed on the
magnesium hydroxide floe and is floated to the
surface by the hydrogen bubbles that are formed.
-------
Chemical Removal of Nitrogen and Phosphorus
131
Following F^yn's laboratory study, a pilot plant
investigation was carried out under the direction
of Mr. L. R. Hougen of the Institute for Industrial
and Technical Research at the Technological Uni-
versity in Trondheim. In Hougen's scheme, the
diaphragm in the electrolitic cell is eliminated.
The electrodes are arranged horizontally and the
difference in the density between the sea water
and sewage is assumedly sufficient to separate the
two liquids. Using a retention time of about 1/2
hour, the results indicated a removal of phosphate
of between 90 and 95 per cent. The pilot plant test
showed power requirements of 1 kilowatt hour per
cubic meter of sewage flow. The patents to Dr.
F^yn's invention have been assigned to Elektro-
kemisk. In addition to the phosphate reduction, it
is reported that the Kjeldahl nitrogen was reduced
from 18.8 to 5.4 mg/1.
In our laboratories, and in a pilot plant con-
structed adjacent to the outfall sewer of the Madi-
son Nine Springs Sewage Treatment Plant, studies
were conducted on phosphate removal using fer-
rous sulfate, ferric sulfate, cupric sulfate, dia-
tomaceous earth, and aluminum sulfate. The most
intensive study of phosphate removal was made
using aluminum sulfate in the form of filtered
alum as the coagulant. Initial studies were made
to determine the effects of concentration of coagu-
lant and pH in the coagulating system. In addition,
studies were made on the influence of mixing time.
Details of these studies have been presented else-
where (Lea, et al., 1954).
In summary, the data showed that an alum
dosage of about 200 mg/1 was required to effec-
tively remove from 95 to 99 per cent of the soluble
phosphates from a sewage treatment plant effluent.
Approximately 6 to 10 times as much coagulant
is required as is ordinarily used in the clarifica-
tion of surface water supplies employed by a com-
munity as a source of potable water. Obviously,
the cost of the chemicals required to remove the
phosphates from 1 mg of effluent will be 6 to 10
times the chemical cost ordinarily incurred in
treating 1 mg of a surface water supply. As an
approach to the problem of reducing the costs for
the removal of soluble phosphates from sewage
treatment plant effluents, a study was made of
coagulant recovery and purification.
After preliminary studies of alum recovery at
low pH values involving acidification and the use
of an ion exchange resin, this approach was aban-
doned in favor of an alkaline or high pH recovery
process. The chemical reactions involved in the
alkaline recovery and purification of the aluminum
hydroxide floe are quite simple, as shown by the
following:
A1(OH)3 ' [P04] + 4NaOH —•+* NaAlO2 +
Na3PO4 + 2H2O + 3OH - NaAlO2 + 2Na3PO4 +
3CaCl2 —*> NaAlO2 + Ca3(PO4)2 + 6NaCl
The mechanics of the process also are simple.
The aluminum hydroxide floe with its adsorbed
phosphate is pumped from the bottom of a sedi-
mentation basin to a recovery tank. The concen-
tration of aluminum hydroxide in the settled floe
normally is equivalent to a 10,000 ppm solution of
aluminum sulfate. Sodium hydroxide is added to
the floe suspension until the pH of the solution is
raised to approximately 11.9. At pH 11.9, the
insoluble aluminum hydroxide is converted to
soluble sodium aluminate and the phosphate to a
soluble sodium phosphate. Addition of calcium
chloride to the solution at this point results in the
formation of insoluble tricalcium phosphate. The
calcium phosphate is readily separated from the
sodium aluminate solution by sedimentation and is
a by-product of the process. The comparatively
phosphate-free sodium aluminate solution is then
adjusted in strength and pumped back to the floc-
culation process to be re-used as a coagulant for
removing more phosphate from sewage treat-
ment plant effluent.
The chemical aspects of alum recovery and re-
use are shown by the following equations and ex-
planatory remarks:
A12(S04)3 ' 14H20 + H20 —>• 2A1(OH)3 +
3H2S04 (1)
200 ppm A12(SO4)3 ' 14H2O yields 52.5 ppm
A1(OH)3 and 99 ppm H2SO4
A1(OH)3 + NaOH —>- NaAlO2 + 2H2O (2)
52.5 ppm A1(OH)3 =55.2 ppm NaAlO2
NaAlO2 + H2CO3 + H2O —*• A1(OH)3 +
NaHCO3 (3)
55.2 ppm NaAlO2 requires 29.6 ppm of CO2
(Neutralization)
The requirement of 29.6 ppm of CO2 is close
to the average concentration of carbon dioxide
present in the Nine Springs effluent. Consequently,
the addition of excess sodium hydroxide over the
amount required to form sodium aluminate must
be avoided. Excess sodium hydroxide will neu-
tralize or consume a part of the carbon dioxide
naturally present in the effluent and thereby re-
sult in incomplete hydrolysis of the recovered
coagulant, sodium aluminate. If the coagulant
loss in the over-all process is 10 per cent, or
20 ppm of alum, and this loss is made up through
-------
132
ALGAE AND METROPOLITAN WASTES
the addition of 20 ppm of new alum just previous
to the addition of the recovered sodium alumi-
nate, advantage may be taken of the acidity of the
added alum. The amount of acidity, as carbon
dioxide, made available in this manner is shown
by the following: 200 ppm of alum upon hydrolysis
produce 99 ppm of sulfuric acid (see Eq. 1).
H2SO4 •
98
2H2O + 2CO2
2 x 44 =88
-CaSO4 + 2H2CC>3
(4)
Therefore, hydrolysis of 200 ppm of alum will
produce, through reaction with the bicarbonates
present in the effluent, 88 ppm of carbon dioxide.
Ten per cent of this amount of alum will produce a
proportionate amount of carbon dioxide. This
proportionate amount, 8.8 ppm, is 29.6 per cent
of the amount of carbon dioxide required to neu-
tralize an amount of sodium aluminate equivalent
to 200 ppm of alum.
PILOT-PLANT STUDIES
With the laboratory studies as a background, a
pilot plant for continuous operation was construc-
ted adjacent to the outfall sewer of the Madison
Nine Springs sewage treatment plant.
With the cooperation of the Madison Metropoli-
tan Sewerage District, a concrete blister was con-
structed on the effluent outfall structure. Installed
in the concrete blister was a 4-inch gate valve.
This construction permitted the withdrawal of
sewage effluent from the outfall structure.
As the effluent was pumped from the blister into
a constant-head tank, an automatic sampler was
used to collect a composite sample of the effluent.
From the constant-head tank, the effluent flowed
into a baffled mixing tank, to which the alum so-
lution was added. Mixing was accomplished by
the use of air.
The effluent flowed over the mixing tank weir,
through a trough into the settling tank, in which
the theoretical detention time was about 2 hours at
a flow rate of 10 gpm. Another sampler was used
at the effluent end of the settling tank, from which
the treated supernatant flowed into a ditch.
A manifold was provided for drawing off the
settled floe. At the drawoff points, valves were
provided to control the flow of sludge, which was
drawn off by gravity into a thickening tank, where
the supernatant was removed and from which a
sump pomp lifted the sludge into the reaction tank.
The reaction tank was 4 ft. in diameter and 5 ft.
high, and was constructed with a conical bottom to
facilitate removal of the calcium phosphate pre-
cipitate. Sodium hydroxide was added to the re-
action tank for pH adjustment and calcium chloride
to precipitate the calcium phosphate. From the
reaction tank, the precipitate was drawn off to a
drying bed. The supernatant of the reaction tank
was then discharged to a storage tank, from which
it was pumped to the alum feed tank. With the
exception of the conical bottom, the storage tank
was of similar dimensions to the reaction tank.
Both tanks were designed to hold about 400 gallons.
As a result of the studies, the following con-
clusions may be made:
1. Laboratory studies show it is possible to
remove approximately 96 to 99 per cent of the
soluble phosphates from the effluent of a sewage
treatment plant. This removal can be accom-
plished in a coagulation process employing any of
the following coagulants: (a) aluminum sulfate,
(b) ferrous sulfate, (c) ferric sulfate, or (d) cop-
per sulfate.
2. The use of copper sulfate as a coagulant is
not advisable because the floe formed by hydroly-
sis of this salt in the sewage treatment plant ef-
fluent has very poor settling characteristics.
3. Filter alum appears to be the most suitable
coagulant for removal of soluble phosphates from
sewage treatment plant effluent because: (a) The
residual phosphate concentration of the effluent
following coagulation with 200 ppm of alum is, on
the average, 0.06 ppm, expressed as P. (b) The
optimum pH range for the removal of phosphates
through coagulation with alum is 7.1 to 7.7. The
average pH value of Nine Springs Treatment Plant
effluent lies hi this range; therefore, no adjust-
ment of pH would be required at this plant, (c) The
concentration of aluminum hydroxide in the effluent
of the coagulation process is only approximately
1.0 to 1.5 ppm and represents a loss of only 0.75
per cent of the coagulant. It is assumed that 1.0
to 1.5 ppm of aluminum hydroxide in the effluent
is not objectionable, (d) The aluminum hydroxide
floe resulting from the hydrolysis of alum may be
recovered, purified by removing the adsorbed
phosphates in the form of tricalcium phosphate,
and re-used for further phosphorus removal in the
form of sodium aluminate. This recovery and
purification reduces by 80 per cent the cost of
chemicals required to remove phosphates from
sewage treatment plant effluent.
4. A study was made of the mechanism by
which phosphates are removed through coagulation
with alum, cupric sulfate, ferric sulfate, or fer-
rous sulfate. This study revealed that for the
first three coagulants the phosphates are removed
through adsorption upon the hydroxide floes
formed. The data obtained on phosphate removal
through coagulation with ferrous sulfate indicated
-------
Chemical Removal of Nitrogen and Phosphorus
133
that adsorption is not the sole means of removal,
but that precipitation may also enter into the
reaction during the transition from ferrous to
ferric hydroxide.
5. The results of this study showed that ap-
proximately 85 per cent of the biologically oxi-
dizable organic compounds and 68 per cent of the
organic nitrogen compounds are removed along
with the soluble phosphates in the coagulation
process employing 200 ppm of alum.
6. The study showed that there was no re-
moval of inorganic nitrogen compounds by the use
of alum coagulation.
7. Pilot-plant studies show that with the use
of the alum recovery process, from 77 to 89 per
cent of the soluble phosphates can be removed.
Filtering of the effluent showed that from 93 to 97
per cent of the soluble phosphate can be removed.
Improved settling facilities should give phosphorus
removals that lie between the unfiltered and fil-
tered values obtained in the pilot-plant study.
NITROGEN REMOVAL
Much less attention has been given to the re-
moval of nitrogen from sewage and sewage effluent
by chemical means obviously because soluble nitro-
gen compounds are least affected by precipitation
processes.
It is of interest, however, to note that Gleason
and Loonam (1933), in reporting their work on the
development of the Guggenheim process, used the
removal of nitrogen as a criterion for the effec-
tiveness of the process.
The Guggenheim process, as described by
Gleason and Loonam (1934), consisted of the fol-
lowing steps:
1. Removal of suspended solids by coagula-
tion with iron compounds and lime, and settling of
the coagulated solids.
2. Disposal of the sludge by filtration and
circulation and regeneration of the iron as ferric
sulphate from the incinerated ash.
Lime
Sewoqe
Effluent
Recloimed
Linw
Solid*
Contact
Process
Basin
Lime
Kiln
Reclaimed Lime
Worts
Re (eneront
Condition in
Tank
A Heot
CaC03
Steam
or Air'
Dso ninoniot( '
or
Forced
Draft
Aerator
Reclaimed
Salt
Solution
Figure 46. PROCESS DESIGN NITROGEN AND PHOSPHORUS REMOVAL FROM SEWAGE EFFLUENT.
Ammonia by Cation Exchange.
-------
134
ALGAE AND METROPOLITAN WASTES
3. Removal of the basic nitrogen compounds
by an exchange reaction using a zeolite.
4. Regeneration of the zeolite and concentra-
tion of the basic nitrogen compounds in a salt so-
lution, and subsequent recovery of ammonia from
this solution. Results reported by these workers
indicated that the total nitrogen in the raw sewage
was 24 - 28 ppm and in the effluent from the pro-
cess, 2-3 ppm. The ammonia nitrogen was re-
duced from a concentration of 12 - 14 ppm in the
raw to 0.5 to 1.0 ppm in the effluent.
Subsequent studies by these workers (Gleason
and Loonam, 1934) carried out at the North-side
Sewage Treatment Works at Chicago over a seven-
month period showed organic nitrogen removals of
about 79.3% and ammonia nitrogen removals of
about 67.4%.
From studies in our laboratories carried out
by Nesselson (1954) using ion exchange, the fol-
lowing conclusions were made:
1. Strong base anion exchangers, regenerated
with common salt, perform satisfactorily for the
removal of nitrate. A number of evaluations es-
tablished that Amberlite IRA-410, in the treatment
of trickling filter effluent, has an exchange capac-
ity of 8.7 to 11.7 kilograms/cu. ft. as CaCO3. It
operates under an efficiency of 4.0 to 6.8 Ibs. of
NaCl/Mlograin of anions removed. Nalcite SAR
had respective values of 6.5 to 7.7 kgrs/cu. ft.
as CaCOs and 6.0 to 9.9 Ibs. NaCl/kgr of anions
removed. Both media were regenerated to an end
point of 1 ppm of nitrogen in the waste regenerant.
A minimum volume of about 7 per cent of the in-
fluent feed was required for this operation.
2. The removal of ammonia nitrogen by nu-
clear sulfonic cation exchangers was investigated.
Nalcite HCR has an exchange capacity of 16.0 to
22.0 kgrs/cu. ft. as Ca(X>3 in the treatment of
activated sludge effluent. It operates with an ef-
ficiency of 1.4 to 2.5 Ibs. NaCl/kgr cations re-
moved. Amberlite IR-120 has respective values
of 13.0 to 17.0 kgrs/cu. ft. as CaCX>3 and 1.3 to
2.6 Ibs. NaCl/kgr cations removed. A minimum
volume of about 6 pejr cent of the influent feed was
required to perform regeneration.
Sowogo
Effluont
Limo
&
RKtounod
J Solids L
I Contact |
I PTOMM I
X **" /
CoCOj
Slurry
Vacuum
RWor
H2S04
V
Forced
Draft
Aorator
Goto
1
"^^nrfMMMiti
m
1
\
Contact
Tank
1
URM
Kiln
Hoot
Figure 47. PROCESS DESIGN NITROGEN AND PHOSPHORUS REMOVAL FROM SEWAGE EFFLUENT.
Ammonia by Forced Draft Aeration.
-------
Chemical Removal of Nitrogen and Phosphorus
135
Kuhn (1956) in further work at Wisconsin
carried out studies on air stripping in packed
towers to remove ammonia nitrogen. As a result
of these studies the following conclusions were
made.
It is possible to remove ammonium nitrogen
from sewage effluent by stripping with air.
Desorption may be accomplished in a tower
packed with Raschig rings.
In the studies conducted, the optimum pH for
stripping was 11.0.* The optimum pH value was
determined from a study of ammonia nitrogen re-
movals through the range pH 8.0 to pH 12.0. In-
creased removals were obtained by increasing pH
values. No significant difference existed between
removals at pH 11.0 and pH 12.0.
The effect of air/liquid loading, expressed as
cfm of air per gpm of liquid, was studied at ratios
of: 40, 59, 85, 230, and 447 cfm/gpm. Respec-
tive removals obtained at these ratios were: 15.1,
28.5, 37.8, 67.0, and 78.7 per cent, respec-
tively. These results are consistent with theory
which predicts best removals at an air/liquid ratio
of 453 cfm/gpm.
Ammonium nitrogen removals were studied at
depths of 1/2" Raschig rings of: 2.5, 4.0, 5.0,
5.5, and 7.0 feet in an 8" diameter column. At
the optimum pH value, pH 11.0, removals of am-
monia nitrogen from sewage effluent of 53.9, 71.1,
78.5, 82.2, and 92.3 per cent, respectively, were
obtained at the depths above. Loadings for these
removals were 52 to 55 cfm of air and 0.10 gpm
of effluent. These results are consistent with the
theory of design of desorption columns.
The studies on phosphorus and nitrogen re-
moval have suggested again two possible process
designs originally proposed by Nesselson (1954).
The units described next are suggested pri-
marily for removal of nutrients from the effluent
of an activated sludge or high-rate filtration plant.
The extent of nitrification in these two types of
plants can be minimized by proper operational
control.
The essential features of the treatment scheme
shown in Figure 46 are as follows: Sewage efflu-
ent enters a softening unit for hardness reduction.
The effluent stream from this unit is then recar-
bonated to convert NH4OH to NH4+. The liquid
then passes through the cation exchanger and is
discharged to the watercourse. Reclamation of
waste regenerant is effected by precipitating the
calcium and magnesium and diverting the slurry
to the lime recovery units. Supernatant liquid
containing NaCl and NILjOH is passed through
either a forced draft aerator or a deammoniator.
The liquid effluent from this unit is recirculated
to the brine tank and the stripped ammonia is re-
covered in a sulfuric acid solution.
Treatment of sewage effluent in the process
design shown in Figure 47 is as follows: Effluent
from the pH adjustment tank flows to a forced
draft aerator. Since air requirements for com-
plete ammonia removal are quite high, only suf-
ficient air is supplied to reduce the ammonia con-
centration to a lower level amenable to economi-
cal treatment by chlorine. Following recarbona-
tion and chlorination, it may be necessary to de-
chlorinate.
REFERENCES
Rudolfs, W. 1947. Phosphates in sewage and
sludge treatment. I. quantities of phosphates.
Sewage Works Journal 19: 1, 43.
Owen, R. 1953. Removal of phosphorus from
sewage effluent with lime. Sewage and Industrial
Wastes Journal 25; 5, 548.
Lea, W. L., G. A. Rohlich, and W. J. Katz. 1954.
Removal of phosphates from treated sewage. Sew-
age and Industrial Wastes Journal 26: 3, 261.
Gleason, G.H. and A.C. Loonam. 1933. The de-
velopment of a chemical process for treatment of
sewage. Sewage Works Journal 5: 1, 61.
Gleason, G.H. and A.C. Loonam. 1934. Results
of six months operation of a chemical sewage puri-
fication plant. Sewage Works Journal 6_: 3, 450.
Nesselson, E. J. 1954. Removal of inorganic ni-
trogen from sewage effluent. Unpublished Ph. D.
Thesis on File, University of Wisconsin Library.
Kuhn, P. A. 1956. Removal of ammonia nitrogen
from sewage effluent. Unpublished M.S. Thesis
on File, University of Wisconsin Library.
-------
136
ALGAE AND METROPOLITAN WASTES
STRIPPING EFFLUENTS OF NUTRIENTS BY BIOLOGICAL MEANS
GEORGE P. FITZGERALD
Sanitary Laboratory
University of Wisconsin, Madison, Wisconsin
Studies have been made over a two year period
on the factors influencing the absorption of algal
nutrients from treated and untreated domestic
sewages by algae, both in controlled laboratory
experiments and in field experiments with a half-
acre pond. This work was supported by the Oscar
Mayer Company and the Rockefeller Foundation
and was carried out by the author and Dr. Harry
Kaneshige.
Laboratory experiments were carried out to
clarify or substantiate some of the data obtained
in the pond studies. These studies have shown that
(1) there is little influence of different lake waters
TIME IN DAYS
as a source of inocula (1 ml/100 ml) on the sub-
sequent growth of algae in secondary sewage ef-
fluent, (2) the amount of growth of algae in
treated effluents is influenced by temperature and
the level of nutrients available for the algae,
(3) growth and nutrient utilization by Chlorella
pyrenoidosa is similar in both primary and secon-
dary effluents (see Figures 48 and 49), and (4) the
CO2 content of effluents will influence the pH,
solubility of nutrients, and the growth and nutrient
utilization of Chlorella (see Figure 50).
12 K
TIME IN DAYS
Figure 48. GROWTH AND NUTRIENT UTILIZA-
TION BY CHLORELLA IN SEWAGE EFFLUENT.
Figure 49. ALGAL GROWTH AND NUTRIENT
UTILIZATION IN PRIMARY AND FINAL EF-
FLUENT.
-------
Biological Stripping of Nutrients
137
TROGCM IAIN * T90 MO C/MTI
ITHOGCN 0.1) » 400 ua C/MV1
TKO6CN (Alt ONLTI
48 72
TIME IN DAYS
Figure 50. GROWTH AND NITROGEN UTILIZA-
TION BY CHLORELLA IN SEWAGE EFFLUENT
WITH DIFFERENTIAL LEVELS OF CO2-
In general, laboratory experiments have indi-
cated that the growth of an alga, such as Chlor-
ella. in domestic sewage effluents, should effi-
ciently remove algal nutrients from solution during
that portion of the year when the temperature is
conducive to the growth of algae.
In contrast to the results of controlled labora-
tory experiments, data from the operation of the
half-acre pond have indicated that there were con-
siderable periods of time, during what might be
considered the optimum algae growing period of
the year, when there was very little algal growth
or changes in nutrient levels.
The data obtained in 1956 with secondary efflu-
ent as feed are summarized in Figure 51 as the
amount of suspended solids in the outlet of the
pond, the percentage removal of the inorganic ni-
trogen as the waters passed through the pond, and
the water detention time in the pond.
An early bloom of Euglena in April had little
effect on nitrogen removal. The peak and valley
of nitrogen removal at this time is due to a slug of
ammonia from a fertilizer plant and its subsequent
passage through the pond.
During the latter part of May, a bloom of
Chlamydomonas brought about a removal of 50% of
the inorganic nitrogen.
A period in the latter part of July, with 10 days
or more detention time, brought about nitrogen
removals in excess of 80%, despite only minor
amounts of algae (Closteridium and Eudorina).
The feed to the pond was changed from secon-
dary effluent about the first of September. The
early nitrogen removals recorded during the first
part of September are due to dilution of the sewage
with the pond waters and the fact that the flow to
the pond was not continuous due to mechanical
failures.
Despite heavy blooms of Euglena and Chlamy-
domonas during October and November, the nitro-
gen removal was only about 20% with detention
times of about 5 days.
During 1957, the feed to pond was secondary
effluent. Since the data of 1956 Indicated that 9
or 10 days detention time appeared best, this flow
was used for most of 1957. The data obtained are
summarized in Figure 52.
The peaks in inorganic nitrogen removed ap-
pear to coincide with the peaks in algal content of
the pond.
The data do not indicate the reason for the low
algal content of the pond waters during June, de-
spite periods of very low flow. During this time,
the pond was inhabited by a quite dense bloom of
Daphnla. However, observations did not indicate
whether the Daphnia brought about the decline in
the algal population or if the Daphnla came into
dominance after the algae had decreased.
There was a period of high inorganic nitrogen
removal during July and August brought about by a
dense bloom of Euglena. This was followed by
another peak in nitrogen removal due to the growth
of a mixture of Chlorella, Chlamydomonas. and
Anklstrodesmus.
The algal population declined during the middle
of September and was replaced with Daphnia.
The pond operation is summarized as follows:.
1. The impoundment of final effluent without
artificial aeration causes the D. O. content to de-
crease considerably below the 6-8/mg/l D. O. of
the influent. Lower levels are obtained during the
summer than in winter. With the growth of algae
in the water, the D. O. rises to levels consider-
ably higher than that of the influent.
2. Variations in the depth of the pond (2 feet
vs. 3.5 feet) had little effect on D. O. during
periods 6f low algal activity.
3. Vertical mixing by means of an air com-
pressor and 1-1/2 inch plastic pipe with 1/8 inch
holes every two feet was found to be very effec-
-------
138
ALGAE AND METROPOLITAN WASTES
I M I I I I I I M I I I I i I I I I I I I I I I I I I I I I i I I I I I I I I I I i I i I
APR. I MAY ' JUN.
SECONDARY EFFLUENT
SEPT. > OCT. ' NOV. DEC.
PRIMARY EFFLUENT
Figure 51. NUTRIENT UTILIZATION BY ALGAE IN A HALF-ACRE POND RECEIVING SEWAGE EF-
FLUENT, 1956.
tive (as measured by suspended solids (algae) and
D. O. concentrations). No apparent effect on B. O.
D. removals was noted.
4. Recirculation inside each cell or from the
effluent end to influent end of the pond had no effect
on B. O. D. removals.
5. Detention time variations between 2 to 5
days during the winter did not cause B. O. D. re-
moval to vary. During summer periods, 10 days
or more detention time appears to be required for
substantial Inorganic nitrogen removal.
6. Suspended solids removal and B. O. D, re-
duction during periods of low algal activity took
place in Cell 1 of the pond, with little change
taking place in the other cells. During two such
periods in 1957, the suspended solids and B. O. D.
were decreased 70-80% in Cell 1.
7. The pond appears capable of removing
of the influent B. O. D. when either primary or
secondary effluents are the feed.
8. The average nitrogen removal throughout
the year is about 30%, with removals in the sum-
mer reaching about 70%. It should be pointed out,
however, that there were only 33 days in 1956 and
76 days in 1957 when the inorganic nitrogen re-
moval exceeded 50% with secondary effluent as
feed.
9. Phosphorus removal during periods of high
algal activity coincides with high pH values in the
pond and is probably due to precipitation. This is
borne out by the fact that during winter periods
the effluent phosphate concentrations frequently
surpassed influent levels—probably due to the
dissolution of phosphorus previously precipitated
(similar results were obtained in laboratory ex-
periments when the pH of solutions were adjusted
to 9.5 and then back to 8.0).
-------
Biological Stripping of Nutrients
139
9
CO
-i
130
120 j
no
100
90
80
70
60
50
40
30
20
10
0
90
80
70
60
50
40
30
2O
10
0
5
0
i it i i n i i i i i n >i> i iilI Iiii I I i
01-
IS 251 5 IS 251 5 IS 2S I 5 15 ZS 5 15 29 I 5 IS 2SI 5 IS Z5 I S I! SS S 15 25 5 15 Zi
JAN. I FEB. ' MAR. '- APR. MAY ' JUN. ' JUL. ' AUG. SEPT. OCT NOV. DEC.
SECONDARY EFFLUENT
Figure 52. NUTRIENT UTILIZATION BY ALGAE IN A
FLUENT, 1957.
HALF-ACRE POND RECEIVING SEWAGE EF-
-------
140
ALGAE AND METROPOLITAN WASTES
THE USE OF ALGAE IN REMOVING NUTRIENTS FROM DOMESTIC SEWAGE
R. H. BOGAN
Associate Professor
Civil Engineering Department
University of Washington, Seattle, Washington
INTRODUCTION
Excessive enrichment of receiving waters by
nutrient rich wastes appears to be emerging as a
major water pollution problem in many areas.
Limnologlcal investigations indicate that, where
domestic sewage serves as the primary source of
eutrophication, treatment for the removal of phos-
phorus would provide a practical and effective
means of control (Sawyer, 1953). Treatment, un-
like the use of algicides, would be permanent and,
as in the case of diversion, would not suffer the
possible disadvantage of degrading other drainage
basins. In spite of these obvious advantages, a
suitable method of treatment has yet to be placed
in practice.
Phosphorus may be removed from sewage by
biological and by chemical means. Either ap-
proach Is aimed at converting soluble inorganic
phosphorus into recoverable insoluble matter. Of
the two, chemical coagulation has received the
greatest attention. Several promising but costly
chemical treatment methods have been proposed.
Little has been done to date with biological treat-
ment.
The purpose of this paper is to explore the con-
cept of employing algae as a means of removing
nutrients from domestic sewage. The concept of
employing algae for this purpose does not appear
to have received much attention, hence much of
what follows is based on research carried out at
the University of Washington during the past three
years. Wherever possible, pertinent data reported
by others has been drawn upon.
THEORETICAL CONCEPTS
Phosphorus Metabolism
'The concept of removing nutrients by biological
means can hardly be considered as new or unique.
In any actively growing biological system, nutrient
i^ntTlpi" are constantly being extracted from the
environment through conversion to cell tissue, the
rate and degree of removal being dependent upon
the biological system employed and upon the en-
vironmental conditions provided.
Rate of removal, other things being equal, Is a
function of the rate of cell tissue synthesis. Growth
rate varies considerably with type of organism, as
shown In Table 41. The mixed mlcroblal culture
Table 41. COMPARISON OF
PORTED GROWTH RATES.
COMMONLY RE-
Type of organism
Algae
Protozoa
Bacteria
ka (day -1)
0.20 to 2.0
1.0 to 4.0
2.0 to 60
a Based on Nj = NQe
kt
provided by the activated sludge process would
appear to be the most effective biological system
in terms of removal rate. Theoretical dally phos-
phate removal rates are shown In Figure 53 for
several concentrations of cell tissue having growth
rates comparable to those commonly reported for
algae.
Cell tissue composition together with the min-
eral content of a sewage will determine the amount
of phorphorus which can be extracted. Examination
of the data presented in Table 42 indicates that
assimilation of 1 mg/1 of phosphorus.
Table 42. COMPOSITION OF ACTIVATED
SLUDGE AND COMMON FRESH WATER ALGAE.
Biological system
Activated stodge*' b>
Algae c
C
%dry wt.
41 -53
49-60
N
%dry wt.
8-12
1.4 - 11
P
%dry wt.
0.7 - 2.2
0.9 - 2.0
a. Hoover. S. R.. and N. Forges. 1952.
b. Helmers, E. N. et al.. (1952).
c. Burlew. 1953. See R. W. Krauss, Chapter 8.
-------
Use of Algae in Nutrient Removal
141
would be accompanied by the metabolism of 25 to
50 mg/1 or more of carbon and 2 to 12 mg/1 of
nitrogen. It is interesting to note the similarity
in the C, N, and P content of algae and of acti-
vated sludge.
20.0
CALCULATED FROM
6"
P-0.02(Nt-N0)
0 0.2 0.4 0.6 08 1.0
GROWTH RATE V, (DAYS'1 )
Figure 53. THEORETICAL RELATIONSHIP BE-
TWEEN CELL TISSUE CONCENTRATION,
GROWTH RATE AND METABOLIC CONVERSION
OF PHOSPHORUS BY ALGAE. Light intensity
will determine cell tissue concentration. Tem-
perature, diet and species will regulate "k". In
the Seattle area, algal concentration ranged from
25 to 50 mg -1 in 4 ft. deep lagoons. The mean Vk"
was 0. 30 days.
Nutritional Limitations
Ordinary domestic sewage does not provide a
balanced diet for activated sludge, being deficient
in carbon and nitrogen with respect to the amount
of phosphorus normally present. Organic carbon
is usually limiting. Algae do not suffer from the
same dietary restrictions. Adequate amounts of
carbon are normally available in the form of alka-
linity. The inorganic nitrogen content of domes-
tic sewage, however, still may be in short supply.
There is evidence that atmospheric nitrogen fixa-
tion might be made to serve as a significant
source of nitrogen (Sawyer, 1953). Agricultural
grade NH4-N could also be employed as a rela-
tively inexpensive source of nitrogen.
Viewed in terms of nutritional requirements,
algae appear to offer the most effective biological
system for extracting phosphorus from domestic
sewage.
Observed Phosphorus Reductions
Based on a carbon to phosphorus ratio of 100
to 1 (Helmers, et al., 1953) and a settled raw
sewage B.O.D. lying in the range of 100 to 200
mg/1, it becomes obvious that phosphorus reduc-
tions during the course of biological sewage treat-
ment would on the average be limited to about 1 to
2 mg/1. Owen (1953), in an investigation of sewage
treatment plant performance in Minnesota, found
phosphorus removals ranged from an average 2 per
cent for promary treatment plants to an average
23 percent for plants employing biological treat-
ment; this was equivalent to approximately 1 to 2
mg/1 of P. Analysis of biological sewage treatment
plant effluents in the Seattle area disclosed a re-
duction ranging from 15 to 40 per cent which was
equivalent to 0.80 to 2.0 mg/1 of P.
A number of excellent research reports have
appeared in the literature dealing with algal be-
havior in oxidation ponds and in raw sewage la-
goons (Burlew, 1953; Oswald, et al., 1953; Os-
wald and Gotaas, 1955; and Anon., 1957). Atten-
tion has been centered mainly upon the use of
photosynthetic oxygen production. Very little data
has been reported regarding nutrient reductions
realized in such processes. Available information
indicates phosphorus reductions ranging from 10
to 90 per cent or better. Performance appears to
be erratic and unpredictable. Considerable diffi-
culty has been experienced in harvesting algal cell
tissue; this coupled with slow growth rate would
account for some of the wide fluctuations noted in
the mineral composition of lagoon effluents.
The Use of Algae
The most direct method of employing algae as
a means of recovering nutrients would appear to
be an oxidation pond followed by a separation
operation for harvesting algae. With such an ar-
rangement sewage would be continually mixed
with actively growing algae and the nutrients
gradually converted to cell tissue; cell tissue
would be recovered for reuse or wasted according
to need. Experience indicates that the principal
problem would be one of harvesting the algae.
Several unit operations have been investigated
as a means of harvesting algal cell tissue (Burlew,
-------
142
ALGAE AND METROPOLITAN WASTES
1953). Those most frequently referred to In the
literature are screening, settling, centrifuging
and chemical coagulation. All have been found
wanting in some aspect, generally in terms of ef-
ficiency and often In terms of cost. From the
standpoint of performance and economy, some
type of screening device appears to be the most
promising.
Screen performance obviously would be related
to the nature of the algal culture. It was reasoned,
however, that if some readily recovered alga could
be established, then through the simple mechan-
ism of recovery and reuse, It could In turn be
made to predominate. Other things being equal,
the process would naturally tend to contain the
most readily utilized population. The ideal or-
ganism would appear to be a large, rapidly grow-
ing, filamentous alga.
There is little evidence that filamentous species
of the type desired normally grow in oxidation
ponds or sewage lagoons. This does not preclude
the possibility of their being cultivated under such
conditions. Failure of filamentous species to con-
stitute a significant part of the population of most
oxidation ponds may be due to the fact that they
are relatively slow growers and simply tend to be
overgrown by Chlorella and Scenedesmus.
EXPERIMENTAL RESULTS
The use of algae In removing phosphorus from
sewage was studied in both the laboratory and the
field. Several fundamental concepts relative to
using algae for such a purpose came under Inves-
tigation. Space does not permit a detailed ac-
counting of all major aspects of this research.
There follows a brief summary of some of the
more significant findings dealing with the topics of
available algae, mechanism of phosphorus re-
moval, physical chemical behavior of orthophos-
phate, photosynthetic pH adjustment, and cell tis-
sue recovery.
Available Algae
Several common fresh water algae were grown
in the laboratory on mixtures of lake water and
raw and treated sewages. However, except for
Chlorella and Scenedesmus, most species gradu-
ally died out after a brief period of active growth.
A large filamentous alga, subsequently identified
as Stigleoclonium stagnatile, was recovered from
the rock of a biological filter in the area and suc-
cessfully cultured. This alga, when grown under
aeration, developed into settleable floe particles
resembling activated sludge. Photomicrographs
of Stigleoclonium stagnatile are shown in Figure
54. Its growth characteristics and nitrogen and
phosphorus content are shown in Tables 43 and 44.
'•
>
(a)
Figure 54. PHOTOMICROGRAPHS OF STIGLEOCLONIUM STAGNATILE. (a) Floe-like colonies which
developed In aerated cultures (220x). (b) View of Individual organism (520x).
-------
Use of Algae in Nutrient Removal
143
Table 43. EFFECT OF TEMPERATURE AND
CULTURE MEDIA ON GROWTH RATE, STIGLEO-
CLON1UM STAGNATILE.
pH variation 8.3 - 9.5. Illumination 400 ft. candles
Temp. °C
10
15
20
Synthetic
sewage
employing
NO3 - N
0.165
0.188
0.252
k dajrs "X*
Synthetic
sewage
employing
NH3 - N
0.140
0.179
0.131
Secondary STP
effluent
0.170
0.215
0.131"
kt
* k computed from Nj = NQe
*• Scenedesmus appeared and began to predominate
after approximately 15 days.
phosphate reductions comparable to those ob-
served In laboratory batch and pilot plant studies.
PO| Behavior
Under suitable conditions, orthophosphate may
combine with a number of substances commonly
present in sewage to form relatively Insoluble
complexes. Calcium Ion concentration and pH
were found to be the principal controlling factors
in determining PO| solubility; their relationship
is shown in Figure 57. Ammonia, Iron and mag-
nesium in the amounts generally encountered in
domestic sewage were not found to exercise any
discernible effect on phosphate solubility.
Constituent
N
P
N/P
Synthetic sewage
NO3 - N NH3 - N
5.71 6.59
2.16 1.81
2.64 3.63
Secondary STP
effluent
6.52
1.89
3.44
1/2 STP Eff
1/2 N03 - N Syn Sew.
6.00
2.07
2.89
a Expressed as percent dry cell weight. Values reported represent an average of several
determinations.
Table 44. PHOSPHORUS AND NITROGEN CONTENT OF STIGLEOCLONIUM STAGNATILE HARVESTED
FROM VARIOUS CULTURE MEDIAa.
Mechanism of PO| Removal
It was expected that phosphorus removal would
follow a pattern predicted by cell tissue composi-
tion and growth rate (see Figure 53). Contrary to
expectations, orthophosphate residuals in batch
fed cultures were found to decrease at a rate con-
siderably in excess of that predicted by biological
uptake. Response to repeated heavy phosphate
doses was most interesting; in general, some 80
to 90 per cent of the phosphate added was removed
from solution within two hours as shown in Figure
55. Obviously, more than metabolic uptake was
involved. Examination of culture characteristics
disclosed that phosphorus removal was related to
pH. It thus appeared that coagulation and adsorp-
tion may have played a significant role.
ADDITIONS
225 MG/L POj AS NA MjPO,
SOMS/U CA*»AS CACL,
SUSPENDED SOLIDS
0 HR - 240 MCA.
12 • -412 •
2JS H«-4«4 •
9 $.
12 1C
TIME-HOURS
At moderately high pH levels, generally in the
range of 9.5 to 10.0, large amounts of phospho-
rus were extracted rapidly from solution without
the use or need of auxiliary chemicals. At lower
pH levels, the rate of phosphorus removal should,
of course, be determined largely by algal growth
rate. This hypothesis was subsequently tested in
the laboratory and in the field. Laboratory pilot
plant performance is shown in Figure 56. At pH
levels of 9. 5 and above, field cultures exhibited
Figure 55.
CULTURES
RESPONSE OF BATCH FED ALGAL
TO REPEATED DOSES OF
P0|.
Aerated Stigeoclonium cultures exhibited a re-
markable capacity for coagulating and adsorbing
orthophosphate. Photosynthetic response was not
visibly impaired by repeated use of cell tissue.
-------
144
ALGAE AND METROPOLITAN WASTES
M
^
i
u 10
0
~L_
1
H
1_
INfLUINI
MOML/HR
HO«L/H
n
— t
i
LMOCN
CONTACT p,^
(.*_
» SLUMC t
-
Sr
——ACTUAL POJ CONC .
TMCOMT POJ CONC
STM K«*0e P0( .90 HO/L
TIMP >I9*C
f— 1 _1 •
LJ
— r^
»
« • KI n
TIME -OATS
PMOSPMOKOUS HtMOVALS IN LAKWATMr PILOT PLANT
9 MO
si
0
w
I
D
»— ^>
*>~*
LAOOON
^
ETFLU
«««|
f
1
(
1
V
1000 f
^x
^
x
h"^
/ ;
TIUC-OATS
» L»idl«T(XIY PILOT Ft«NT OPW»TW« CONP1T1<)«$
Figure 56. TYPICAL LABORATORY PILOT
PLANT PERFORMANCE.
Photosynthethetic pH Adjustment
A study was made of the role of algae in adjust-
ing pH. Such factors as temperature, cell tissue
concentration, composition of culture media and
alkalinity naturally would be expected to influence
the rate of photosynthetic pH adjustment. In-
terestingly, light Intensity was found to be the
principal rate controlling factor. Photosynthetic
pH response under carefully controlled laboratory
conditions and under field conditions is shown in
Figure 58 and Table 45. Where light was not
limiting, pH response followed a pattern such as
that shown in Figure 58. Under field conditions
light intensity became limiting and the rate of
change of pH was consequently markedly reduced.
Judged solely on the basis of rate of pH ad-
justment, minimum light intensity requirements
appear to lie in the vicinity of 100 to 200 f. c.
Where such light intensities are possible, the pH
of raw and treated sewages may be photosyn-
thetically increased to 9. 0 and above within 4 to
12 hours.
r
_i
a
i
UJ
I
a.
i
Q-
o
tc
o
30
20
in
8
7
6
5
4
9
a
1.0
0.8
0.7
as
as
04
as
az
ai
A
; &
E V-
: \
E
— i —
*v
\X
Vs
\ \
vV
\\
\ \
\ \
\ \ *
\\ \
-
.
.
.
.
-
•
-
5-
\\
\
\
\
3
N^
X ^'
v
\
s SN
3
v
\
\
V x
V s
S~
x
* —
s
^ "**•
S
^
N
NXN
X.
>s
"V
^\
'v
X
X
POJ CONC. INFILTRATE, ^s.
USING:
SYNTHETIC SEWAGE
SECONDARY PLANT EFF
— 1 —
-
•
».
pH-9 -
"Xy^,^
-
-
pH "10 "
-
s
^x.-
•
pH-ll .
uUENT
0 20 40 60 100
CALCIUM CONCENTRATION-MG/L CA**
Figure 57 INFLUENCE OF pH AND CALCIUM
CONCENTRATION ON ORTHOPHOS PHATE
SOLUBILITY. Calcium ion concentration and pH
adjusted by means of CaCl2, Ca(OH)2 and NaOH.
Samples of synthetic sewage and sewage treatment
based on filtered supernatant.
SCEHEDE5MUS IN STP EFFLUENT
CELL TISSUE CONCENTRATION APPHOX Z5O MOA
TEMP.'w-20'C LIGHT. IOOO FC
600 BOO
TIME-MINUTES
Figure 58. INFLUENCE OF HCO3- CONCEN-
TRATION ON PHOTOSYNTHETIC pH SHIFT. At
light intensities above 200 f. c., Scenedesmus,
Chlorella and Stigleoclonium caused a rapid In-
crease in the pH of raw and treated sewages.
Alkalinity in amounts generally encountered In
most sewages had little effect on pH response.
-------
Use of Algae in Nutrient Removal
145
Table 45. PHOTOSYNTHETIC pH CHANGES IN LAGOON CELLS - Autumn, 1958.
Lagoon cell
1
2
3
4
Time of sampling
Weather
Temperature °C
Susp. solids mg/1
Day
1
7.7
8.0
8.1
8.1
1:30 PM
Cloudy
2
7.6
7.7
8.0
7.5
11: AM
Cldy
28
3
8.4
8.4
8.5
8.2
11: AM
Pt. Cldy.
14
33
5
8.7
8.8
9.1
10.4
11:30 AM
Rain
36
7
8.5
8.9
9.3
10:30 AM
Sunny
12
36
10
9.4
9.2
9.7
3 PM
Pt Cldy.
11
62
12
9.5
9.2
9.7
11:30 AM
Sunny
10
62
15
10.1
9.6
10.4
12N
Sunny
11
70
Remarks
Intermittent aeration
plus artificial illumi-
nation (400 F.C.) PM
Int. aeration: no
artificial ilium.
No. aeration; no
artificial ilium.
Int. aeration. Air
off after day 6
All lagoon cells
filled with secondary
treatment plant ef-
fluent and seeded
with equal volume of
algal culture.
Cell Tissue Recovery
When it was found that Stigleoclontum stagnatile
could be effectively recovered by sedimentation,
the concept of screening as a recovery device was
abandoned. Repeated use of a Stigleoclontum cul-
ture, such as during successive batch culture
feedings or during cell tissue recycling in pilot
plant operations, resulted In a marked improve-
ment in settling characteristics. It thus appeared
that adsorption of insoluble calcium phosphate
salts by the algae caused coagulation of cell tis-
sue, thereby increasing its rate of subsidence.
During pilot plant studies, the Stigleoclonlum
culture became contaminated with Chlorella and
Scenedesmus. At temperatures above 20°C,
Chlorella and Scenedesmus tended to predominate;
Stigleoclonlum was simply overgrown. This gave
rise to some serious misgivings until it was ob-
served that culture settling characteristics varied
little with species owing to the coagulation effect
of the insoluble phosphate salts produced at high
pH levels. This is most interesting for it implies
that the algae in most oxidation ponds might be
effectively recovered by sedimentation If they are
first permitted to Increase pH levels above ap-
proximately 9. 5 before recovery Is attempted.
DISCUSSION
Algae are capable of removing phosphorus
from solution both by metabolic uptake and by
chemical coagulation and adsorption. Adsorption
and coagulation appear to play the major role
where rapid removal of large concentrations of
phosphorus is involved. The relative significance
of biological uptake depends, of course, on algal
growth rate, environmental conditions, and upon
time available for growth. In either case, it is
the photosynthetic activity of the algae which
governs the rate of removal.
High Rate Process
Laboratory pilot plant studies employing an il-
luminated contact unit followed by sedimentation
have demonstrated that a high rate continuous flow
process is functionally feasible when light is not
limiting. Orthophosphate concentrations can be
reduced to less than 1 mg/1 within 6 to 12 hours.
This is equivalent to a 90 to 95 per cent reduction
in the phosphorus content of most sewages. Re-
sidual phosphate concentrations of about 3 to 5
mg/1 were realized with contact times as brief as
2 to 4 hours.
While a high rate algal treatment process ap-
pears to be functionally sound, economic consid-
erations restrict the use of algae to the more
leisurely conditions prevailing in oxidation ponds
and sewage lagoons. Process costs are related
to the cost of pH adjustment, which in turn is a
function of light requirements and holdup time.
Where artificial illumination is employed, power
requirements and hence, cost can be computed
from a knowledge of lamp performance and the
rate of light attenuation hi the algal culture. Cost
of pH adjustment employing artificial and natural
illumination is compared in Figure 59 together
with the cost of pH adjustment by lime alone.
Lime requirements were computed from titration
curves of a number of treated sewages in the
Seattle area. Economical considerations appear
to preclude the use of artificial illumination.
Lighting Limitations
Owing to a high rate of light attenuation, it is
exceedingly difficult to maintain adequate Illumi-
nation in large scale cultures. Theoretical con-
siderations based on the Beer-Lambert Law indi-
cate that it ordinarily would not be possible to
maintain light intensities above 100 f. c. by natural
or artificial means in depths much greater than
-------
146
ALGAE AND METROPOLITAN WASTES
1000
UJ
Q
<
IOO
10
ARTIFICIAL ILLUMINATION
15 WATTS/FT3 „*
OXIDATION PONDS
AT tO-OS/FT8
9
PH
10
Figure 59. ECONOMIC COMPARISON OF VARI-
OUS METHODS OF ADJUSTING pH. Cost of elec-
trical energy was taken at $.01/kw-hr. Power
requirements were based on high voltage fluor-
escent elements and a culture extinction coeffi-
cient of 2 x 10-3 cm2/mg. Oxidation pond costs
were based on a construction cost of $0.05/ft3, a
useful life of 20 years, and a liquid depth of 4 feet.
Lime requirements were based on titration cur-
ves for sewages in the Seattle area; mechanical
equipment costs were neglected.
one foot. Current practice is to construct oxida-
tion ponds and lagoons with depths ranging from
3 to 5 feet.
As a practical matter where thorough mixing
occurs, the effect of employing depths greater
than that involved in photosynthesis is roughly
equivalent to illuminating the entire culture at a
proportionately lesser intensity. The net photo-
synthetic response is thus reduced and a longer
contact time must be provided to achieve a given
degree of pH adjustment. For example, it was
found that 12 hours contact time at 100 f. c. was
approximately equivalent to 10 to 12 days lagoon
retention time under field conditions prevailing
in the Seattle area during the fall of the year.
Detention Time Requirements
Theoretically, an oxidation pond should be ca-
pable of very high efficiencies of phosphorus
removal. In order to realize this potential, active
photosynthesis and/or high pH conditions must
prevail for some time prior to discharge. Where
biological fixation is the principal mode of phos-
phorus removal, efficiency will be a function of
detention time, growth rate, and cell tissue con-
centration.
During field pilot plant studies, algal cell tis-
sue concentrations in lagoon units having a depth
of 3 to 4 feet varied from 25 to 50 mg/1. Average
growth rate was equivalent to a k of 0.30 day-1.
Thus, in order to biologically extract 5 mg/1 of P
(equivalent to 80 to 90 per cent reduction for most
sewages), it appears that lagoon retention times
on the order of 14 to 28 days would be required.
Theoretical retention time requirements for any
other set of circumstances can be calculated from
the data presented in Figure 53. These consider-
ations suggest that pond volume in excess of that
employed in most oxidation ponds may be neces-
sary where a high degree of phosphorus removal
is a treatment objective.
SUMMARY AND CONCLUSIONS
The concept of employing algae as a means of
removing phosphorus from sewage was studied
in the laboratory and in the field on a pilot plant
scale. A high rate process was developed in the
laboratory whereby soluble phosphate reductions
equivalent to 90 per cent or better were achieved
with contact times as brief as 6 to 12 hours. The
process subsequently was studied in the field
along with the behavior of conventional sewage
lagoons. Certain aspects of this work were dis-
cussed in this paper and are briefly summarized
below.
1. In the presence of adequate amounts of
light, it is possible to realize rapid biological ex-
traction of phosphate. Minimum light require-
ments appear to be in the vicinity of 100 to 200 f. c.
2. Under normal field conditions, adequate
light intensities seldom prevail in algal cultures
at depths much in excess of one foot. The use of
deeper ponds, in common practice today, serves
in effect to decrease net illumination roughly in
proportion to the ratio of light and dark volumes.
Photosynthetic reaction times are markedly in-
creased.
3. Adsorption and coagulation appear to play
the major role where rapid removal of large
amounts of phosphate is involved. Metabolic con-
version is the principal removal mechanism under
the more leisurely conditions prevailing in oxida-
tion ponds and sewage lagoons. In the latter case,
efficiency of phosphate removal is proportional to
detention time, other things being equal.
4. Three algae, Chlorella, Scenedesmus,
and Stigleoclonium, were grown in raw and treated
-------
Use of Algae in Nutrient Removal
147
sewages. Repeated reuse of these algae generally
improved subsidence properties without noticeably
impairing photosynthetic response. Settleability
appeared to be influenced by the formation of in-
soluble phosphates at elevated pH. It appears
that the photosynthetic pH shift may be employed
as a means of enhancing the recovery of algae by
sedimentation.
ACKNOWLEDGMENTS
A number of the writer's former students par-
ticipated in this work. Messrs. Orris E. Albert-
son and James C. Pluntze were responsible for
process development and study.
Initial phases of this work, including the labor-
atory pilot plant studies, were supported by the
University of Washington Engineering Experiment
Station.
The city of Seattle provided funds for construc-
tion of the field scale pilot plant.
REFERENCES
Anon. 1957. Development of design criteria for
waste stabilization ponds. University of Texas,
Civil Engineering Department, Final Report to the
AEC, March 1, 1957.
Burlew, John S., Editor. 1953. Algal culture
from laboratory to pilot plant. Carnegie List, of
Washington, Publication 600, Washington, D. C.
Helmers, E. N., et at. 1952. Nutritional require-
ments in the biological stabilization of industrial
wastes. Sewage and Industrial Wastes 24: 496.
Hoover, S. R., and N. Forges. 1952. Assimila-
tion of dairy wastes by activated sludge - n. Sew-
age and Industrial Wastes 24: 306.
Oswald, W. J., et al. 1953. Algal symbiosis in
oxidation ponds - n Growth characteristics of
Chlprella pyrenoidosa cultured in sewage. Sewage
and Industrial Wastes 25: 26.
Oswald, W.J., and H.B. Gotaas. 1955. Photo-
synthesis in sewage treatment. ASCE Proceed-
ings, 81, Sep. No. 636.
Owen, R. 1953. Removal of phosphorus from
sewage plant effluent with lime. Sewage and In-
dustrial Wastes 25: 548.
Sawyer, C. N. 1953. Some new aspects of phos-
phates in relation to lake fertilization. Sewage
and Industrial Wastes 24: 708.
-------
Use of Algicides
THE PRACTICAL USE OF PRESENT ALGICIDES AND MODERN TRENDS TOWARD NEW ONES
KENNETH M. MACKENTHUN
Public Health Biologist
Wisconsin Committee on Water Pollution, Madison, Wisconsin
Prolific and extensive algal growths give rise
to many damaging effects and hinder man in his
full utilization of this great natural resource,
water. These effects have been well documented
(Bartsch, 1946; Bartsch, 1954; Mackenthun, 1952;
and Mackenthun, 1958). Algae can cause tastes
and odors in water supplies, clog filters in indus-
trial and municipal treatment operations (Mathe-
son, 1952) and Interfere with the manufacture of a
product in industry, such as paper. They may de-
crease the flshablllty of water, destroy a fish
population through the elimination of the dissolved
oxygen resulting from decomposition (Mackenthun,
Herman and Bartsch, 1948), and decrease such
recreational uses as swimming, skiing, speed
boating, skin diving, etc. Some species have been
found to produce toxic blooms which upon acciden-
tal or purposeful ingestion have been known to
cause abdominal discomfort to man, and kill fish,
pets, all types of wildlife, and domestic animals
(Davidson, 1959; Ingram and Prescott, 1954;
Mackenthun, Herman and Bartsch, 1948; Prescott,
1948; Rose, 1954; and Senior, 1960). Some wild-
life kills, especially, have been most extensive.
The decomposition of a large algal mass may
liberate sufficient hydrogen sulfide to react with
the lead compounds used as paint pigments, and
discolor paint on lake shore residences. The stain-
ing is especially noticeable on weathered white
paint, forming brown specks or blotches. The
odors arising from the decomposition of this mass
of algae and dead fish are so vile that cottages
remain unrented, vacations are cut short, busi-
ness is hampered, and property values are
lowered.
Today our waters are subjected to the onrush
of civilization, population pressures, year-around
shoreline dwellings, Increased pollution, and the
concept of multiple use. Because of these in-
creasing demands placed upon our recreational
waters, adequate control of an algal nuisance is
of utmost importance, especially to those who live
in close association with highly productive lakes.
An ideal alglcide for adequate control should
possess certain attributes (Mackenthun, 1959). It
must kill the specific nuisance plant or plants, be
non-toxic to fish and most fish-food organisms at
the plant-killing concentration, not prove seriously
harmful to the ecology of the general aquatic area,
and be of reasonable cost. Since 1904 (Moore and
Kellerman, 1904) and up to the present time, the
chemical which has most nearly met these speci-
fications for the control of algae has been copper
sulphate (blue vitriol). Despite this extensive
usage, copper sulphate does have shortcomings in
that it may poison fish and other aquatic life In
excessive concentrations, it may accumulate in
bottom muds as an insoluble compound following
extensive usage, and it is corrosive to paint and
equipment (Bartsch, 1954).
Initial attempts at prescribing definite dosage
rates for the control of various types of algae with
copper, sulphate were first undertaken by Moore
and Kellerman in 1905 (Moore and Kellerman,
1905) and have been extensively reprinted in tabu-
lar form giving specific dosages for some 70
organisms (Hale, 1954). The practical application
of such a table is difficult, however, because of
the many variables which one encounters in nature.
Because solubility of copper in water is influenced
by pH and alkalinity as well as temperature, the
dosage required for control depends not only upon
the chemistry of the water itself, but also upon
the species or genus variation of the organisms
and their resistance to copper sulphate. Thus,
rather arbitrary dosage rates have been success-
fully used, especially in the midwestern states,
for a number of years (Mackenthun, 1958; and
Bartsch, 1954). Since a total alkalinity of 40.0 ppm
seems to be a natural separation point between
soft and hard waters (Moyle, 1946), those lakes
which have a total methyl orange alkalinity of
40 ppm or greater are treated at a rate of 1 ppm
blue vitriol for the upper 2 feet of water regard-
less of actual depth. On an acreage basis, this
concentration would amount to 5.4 pounds of com-
mercial copper sulphate per surface acre. The
2-foot depth has been determined to be about the
maximum effective range of a surface application
of copper sulphate in such water since algae will
be killed with increasing depth only if the rate of
downward diffusion exceeds the rate of copper
precipitation. The algae killed by such a treat-
148
-------
Use of Algicides
149
ment are those which are suspended near the
water surface and which commonly occur with
blooms in calm weather. In lakes with a total
methyl orange alkalinity below 40 ppm, a concen-
tration of 0.3 ppm commercial copper sulphate
for the total volume of water has been selected.
This is roughly comparable to 0.9 pounds of cop-
per sulphate per acre foot of water. It is obvious
that if a low alkalinity lake has an average depth
of about 6 feet, the dosage would be about the
same as in a high alkalinity lake of the same area.
When lesser average depths are involved, a
greater concentration of chemical results in the
high alkalinity lakes. This apparent paradox would
be even more striking were it not for the fact that
in low alkalinity lakes, the algae frequently are of
the filamentous types that may lie at the bottom
and, therefore, the entire volume of water must
be calculated to insure that a sufficient concentra-
tion of the chemical will reach the algae to effect
a kill. In high alkalinity lakes, algae frequently
are planktonlc and tend to concentrate near the
surface which is the only stratum at which appre-
ciable concentrations of soluble copper can be
expected. Certainly, when copper sulphate is used
as the algiclde, the best and most lasting control
will result if the lake water has a total alkalinity
around 50 ppm or less.
Algal control treatments may be marginal or
complete and the type applied to a given body of
water must be determined by the size, shape, and
relative fertility of the water, and the estimated
cost of the project. Complete treatment in which
the calculated amount of copper sulphate is sys-
tematically applied over the entire surface area is
the most satisfactory. It insures that a major
portion of the total algal population is eliminated
at one time so that a long algal recovery period is
required before a bloom condition may again oc-
cur. The interval of time between necessary
treatments will be directly correlated with clima-
talogical conditions and the available nutrients
utilizable by the remaining algal cells which are
not killed as a result of chemical application. One
to three complete treatments per season may be
sufficient to give reasonable control. Marginal
treatment, on the other hand, is a method designed
to obtain temporary relief in a restricted area
where more extensive activity Is not practicable
or financially possible. It is a procedure in which
a lane 200 to 400 feet wide, lying parallel with the
shore, and all protected bays are sprayed in the
same manner as for complete treatment. All
other parts of the area are not affected even though
much algae may be present.
As a result of treatment, the algal population
and the intensity of odors along the periphery of
the lake are reduced. The duration of freedom
from the algal nuisance following this type of
treatment is dependent upon the density of the
algal population in the center of the lake and its
ability to Infiltrate the treated area through the
action of wind, waves, and currents, in addition
to those repopulation factors presented for a com-
plete treatment operation. Any marginal control
operation should definitely be considered on a
periodic repeat basis. Providing fertility is not
excessive, large bodies of water mijht gain enough
relief from marginal treatment to warrant this
type of control. However, It would probably be a
waste of money to consider this type of control for
a large, fertile lake which is long and narrow and
subject to considerable wind and wave action.
Complete treatment In this case might be the only
present answer, and the cost of complete treat-
ment might be prohibitive.
Copper sulphate may be applied in a variety of
ways: bag-dragging, dry feeding (Monie, 1956),
liquid spray (Mackenthun, 1958), and airplane ap-
plication of either dry or wet material (Anon.,
1959). Regardless of the method, rapid and uni-
form distribution of the algiclde Is essential.
Therefore, the size and scope of the problem
would determine to some extent the method em-
ployed. In general, liquid spraying systems oper-
ated from a boat or barge have thus far been most
widely used.
Copper sulphate Is a highly corrosive chemical.
Therefore, materials which are used in the con-
struction of spraying equipment should be resis-
tant to its corrosive nature. These include red
brass, containing 85% copper and 15% zinc; com-
mercial bronze, containing 90% copper and 10%
zinc; stainless steel; and a good grade of copper.
Cast iron is unsuitable. A more complete list of
materials may be found in the Chemical Engineer1 s
Handbook (Anon., 1950).
The life of chemical spraying equipment de-
pends to a great extent upon the care given the
equipment. Equipment which is used in spraying
a corrosive chemical should be thoroughly cleaned
in clear water following the spraying operation.
This may be accomplished by spraying clear water
at the end of treatment before the equipment is
taken apart. It is most Important that a residue
of copper sulphate not be left In the equipment at
the end of the operation, thus producing corrosion
over a long period of time.
Essential equipment for the distribution of a
solution of copper sulphate is illustrated in Figure
60. The equipment, as shown, can be easily fab-
ricated for about $200. The copper sulphate,
crystals or powder, is placed in the chemical so-
lution tank to which water Is constantly added as
the saturated copper solution is withdrawn. The
saturated copper solution is bled into the suction
line containing clear lake water producing a 2% to
5% solution. This is drawn through the pump and
discharged through a smooth fire hose nozzle
which is continuously swept back and forth over
-------
150
ALGAE AND METROPOLITAN WASTES
HOSE RETURN-
TO WASH CRYSTALS
3? PRESSURE HOSE
« AMD NOZZLE WITH
DISCHARGE
37 PRESSURE HOSE
STWLESS STEEL
STRAINER WITH
£ HOLES
POWER
SOURCE
WITH
VALVE
SCREEN
Figure 60. BASIC EQUIPMENT DESIGN
the area receiving treatment so that good distri-
bution Is obtained. A small portion of the dis-
charge solution is returned to the chemical solu-
tion tank to furnish dilution water far the crystals.
This will naturally reduce the nozzle pressure and
decrease the diameter of the spray pattern. To
compensate for this, the equipment may Include a
small auxiliary pump whose sole function is to
maintain a constant level of dilution water in the
chemical solution tank. Other modifications may
Include the substitution of a California redwood
baffled wooden box for the chemical solution drum,
the substitution of a burlap bag of chemical for the
stainless steel strainer in a chemical solution
tank, and the addition of mechanically oscillating
spray nozzles. A wooden barrel may be used as a
chemical solution tank since it will not corrode
and lasts well; however, an ordinary steel drum
will usually suffice for one season providing the
treatment Is not too extensive. A good quality, of
pressure hose or steam hose, with similar fittings
and clamps, should be used for all hose connec-
tions in assembling the equipment.
The selection of the pump which will exert the
right pressure on a particular size of nozzle to
produce the desired volume of discharge and spray
combination should be given consideration. Early
experiments (Fleming, 1918) on fire hose and
small nozzles supplied basic information on the
distance of throw when the nozzles are operated
under varying pressures (Table 46). It Is indi-
cated that the tip of the nozzle should be reamed
out for a distance of at least one-half Inch in order
to obtain a good fire stream.
The boat, barge, or launch which Is used to
transport the spraying equipment Is equally as
Important as the equipment itself—and generally,
more costly. It must be adequate to carry the
equipment, required personnel, and additional
chemical. Speed and ease of operation are essen-
tail for good chemical distribution.
The amount of commercial copper sulphate to
apply to a given area can be readily calculated
either by arithmetic or through the use of a dosage
Table 46. DISCHARGE FROM SMALL NOZZLES AT VARYING PRESSURES.
Results of Experiments at University of Illinois with Small-size Nozzles attached to 1-1/2-inch hose.
9/16" Nmtsle
1/2" Nozale
fttbue
aCnozsle
(PAL)
Good;
tori- Extreme
until tort-
flre zontal
Discharge stream drops
UM>.IB.) let (ft) (feet)
10
SO
40
50
60
TO
«0
90
100
12
16
17
1*
01
23
24
16
as
IS
18
21
23
20
28
29
30
31
53
63
71
78
84
90
96
102
107
Discharge
(g-p-m.)
33
40
46
52
57
61
65
69
73
Good
hori-
zontal
fire
stream
jet (ft.)
15
20
25
30
33
37
40
43
46
Extreme
hori-
zontal
drops
(feet)
63
79
91
102
111
120
127
134
140
-------
Use of Algicides
151
zoo
19
18
9280
9000
Figure 81. COMMERCIAL COPPER SULPHATE
DOSAGE CHART FOR LAKES WITH A METH-
YL ORANGE ALKALINITY GREATER THAN 40
PPM. Treating a mile of shoreline for a width of
200 feet requires 130 pounds of blue vitriol.
chart (Figure 61). The area of the spray pattern
and the concentration of chemical solution as it
leaves the nozzle determine the concentration in
the receiving water. It thus remains for the
operator to effect an even distribution of the cal-
culated amount of chemical within a given specified
area. This may be quite easily accomplished by
adjusting the percentage of the solution sprayed
with the diameter of the spray pattern and the
speed of the boat. Acidified copper sulphate
standards containing a 1%, 2%, 3%, 5%, and 8%
solution of commercial copper sulphate should
always be available for testing purposes. The test
is made by collecting a small sample from the
pump discharge, adding sufficient acid (powdered
citric acid has been found easiest to handle in the
field) to clear up the precipitate and matching the
test color with the standard colors to determine
the percentage of solution being sprayed. Other
methods include the use of the known pump ca-
pacity to determine that a given percentage solu-
tion will involve the distribution of a given number
of pounds of copper sulphate within a given number
of minutes. In this method, the speed of the boat
must be regulated so that the entire area is
covered in the specified time. Calculations can
also be made using the average speed of the boat
or barge coupled with the diameter of the spray
pattern to determine the pounds of chemical which
must be used in a given amount of time.
The relative cost for the control of algae
through the use of copper sulphate on high alka-
linity lakes is in the neighborhood of $1 per acre
for chemicals, plus the cost of distribution for
each treatment undertaken. The cost of distribu-
tion may range anywhere from 25? to $1 per acre,
depending upon the methods used in control and
the circumstances surrounding the particular con-
trol project. In a liquid spray operation from a
boat or barge, it is possible to distribute about
300 to 350 pounds of commercial copper sulphate
per hour of spraying time, thus covering about 60
acres per hour.
The effect of copper sulphate treatment on an
algal population can be noted soon after the treat-
ment is applied. Within a few minutes, the color
of the water will change from dark green to
grayish-white. Although at no time are all the
algae in the lake entirely eliminated (Domogalla,
1926), the water should be visibly free of cells two
or three days following a complete application.
Treatment has proven beneficial, both when ap-
plied to large bodies of water (Domogalla, 1935;
and Domogalla, 1941) and also to small fish-
rearing ponds (O'Donnell, 1945). Both in Wiscon-
sin and Minnesota (Moyle, 1949), there is an indi-
cation that certain species of algae, particularly
Aphanizomenon. seem to have acquired an in-
creased tolerance to copper as a result of many
years of treatment. Two to five times as much
copper sulphate must now be used as was neces-
sary some 26 years ago to achieve similar control.
Much has been recorded on the toxicity of cop-
per sulphate to various forms of aquatic life
(Doudoroff and Katz, 1953; Ellis, 1937; Marsh and
Robinson, 1910; Prescott, 1948; Rushton, 1924-
Schaut, 1939; and Anon., 1939). Extensive field
and laboratory studies have clearly shown that
fish are not killed by copper sulphate at concen-
trations normally used for algal control and that
fishing and fish yields of lakes which have been
treated over a long period of time have not de-
teriorated (Moyle, 1949). It is well known that
copper will accumulate upon the lake bottom fol-
lowing repeated treatments and that the greatest
amount of accumulation is found in the profundal
region (Nichols, Henkel and McNall, 1946). At-
tention has also been directed to the possible de-
leterious effect of this accumulation on lake
ecology (Hasler, 1947; Schoenfeld, 1947; and
Schoenfeld, 1950). However, it has been shown
-------
152
ALGAE AND METROPOLITAN WASTES
experimentally that the accumulation of copper in
bottom muds resulting from the use of nearly two
million pounds of copper sulphate to control algae
in a hard-water lake over a period of 26 years was
considerably lower In concentration than the
amounts determined to have a deleterious effect
on the profundal bottom-dwelling organisms (Mac-
kentnun and Cooley, 1952). More recent work
(Antonie and Osness, 1959) on this.same lake
(Lake Monona, Madison, Wisconsin) has shown
that the copper content is significantly less hi the
upper 1" of bottom mud than in the stratum 6"
lower. Since this lake has received reduced
treatment with copper sulphate since 1947 and
practically no control since 1953, this would sug-
gest that the constant seepage of silt into the
deeper areas of the lake will in time cover up
the heavy concentrations of copper that were de-
posited during the years of continuous and heavy
treatment.
In recent years, there has been a constant
search to find a chemical replacement for copper
sulphate which would be more effective, non-
accumulative, and perhaps even less costly over
an extended period of time (Maloney and Palmer,
1956; and Palmer and Maloney, 1955). Work has
been done on Phygon XL (Fitzgerald, Gerloff, and
Skoog, 1952; and Fitzgerald and Skoog, 1954),
which is rather selective for blue-green algae,
and more recently on related naphthoqulnone com-
pounds which are believed to be less toxic to fish.
Delrad (dehydroabletylamine acetate) has also
proven successful under specific situations (John-
son, 1955; and Lawrence, 1954), but care must be
exercised hi its use because of its toxicity to fish
hi some waters. Some newer alglcides are just
now entering the field-testing phases following
preliminary work. These will require many tests
and observations under the varying conditions
found hi nature before their future potential can be
determined. In all cases, where a newer algicide
is involved, the cost of treatment of a large area
has thus far exceeded the cost based upon the use
of copper sulphate (Hooper and Fukano, 1955) and
none have made up the difference in the amount of
control offered. As a result, present day usage
of some of the newer alglcides has been confined
to the smaller operations where a more costly
chemical Is feasible. Copper sulphate remains as
the most effective, least harmful, and least ex-
pensive algicide for large-scale control at the
present time!
Occasionally the 'need arises for an algicide to
control a specific type of growth. By and large,
however, the problem is one of a general algicide
to control a general algal nuisance under the ma-
jority of circumstances encountered hi field oper-
ations. This, then, is the basic challenge which
must be met by a striving "new" chemical—it
must control a general algal nuisance at a con-
centration not seriously harmful to lake ecology
and It must achieve control at a competitive cost
to existing standard procedures. If the cost is
greater, it must have more desirable qualities to
offer, and, generally speaking, It must either do a
better or a more lasting job of control.
Any algal control measures should be under-
taken at the right tune in the development of the
algal bloom and this is especially true in treating
confined areas. If, for some reason, treatment
is not applied to a given area until the algal cells
have become dense, one must use good judgment
in determining the extent of the area which should
receive treatment at one time. Usually, it is a
good practive to sub-divide the area Into sections
hi an attempt to control the nuisance hi one section
at a given time. Other sections may be treated
later to insure that sufficient dissolved oxygen is
present to satisfy the demand of the decomposing
algae.
Recently, a questionnaire was directed to each
state to obtain information on aquatic nuisance
control activities. The problems of algal nui-
sances vary considerably from state to state.
Many states do receive requests and many do
carry out some type of control program. Some
states report that no problem exists and some re-
port a much reduced problem because of high
water turbidity. In many states, It is necessary
to receive authorization prior to the treatment of
public waters for algal control. About 40 per cent
of the states regulate the introduction of chemicals
for the control of aquatic nuisances by statute or
executive order and require a permit for the In-
troduction of chemicals into public waters. About
57 per cent of the states report complete super-
vision of field application of chemicals and an ad-
ditional 21 per cent spot check on field applica-
tions.
' Wisconsin is the only state with a long-standing
special statute pertaining specifically to aquatic
nuisance control and the only state which assesses
an established fee for permits and a definite rate
of charge for supervisory service. To set up a
project, an application for permit and a $10 appli-
cation fee Is filed with the Wisconsin Committee
on Water Pollution. The Committee determines,
through the law enforcement division of the Con-
servation Department and the appropriate county
clerk, if there are any objections on the part of
citizens to the proposed chemical application. If
objections are forthcoming within a specified
period of time, a meeting or possibly a hearing is
held and as a result the permit is granted, re-
jected, or modified. Upon receipt of a permit,
the sponsoring party may hire one of the commer-
cial operators who will carry out treatment hi ac-
cordance with a specified plan which the operator
submits to and is approved by the Committee, or
the sponsoring party may provide suitable equip-
ment and all necessary materials and labor to
complete the project. Under either method, the
-------
Use of Algicides
153
distribution of chemicals into public waters for
purposes of aquatic nuisance control must be
supervised by a representative of the Committee
on Water Pollution. There is a charge of $8 per
hour or fraction thereof for this supervisory ser-
vice, or a minimum seasonal charge of $20 if the
seasonal supervision requirements do not exceed
two hours of supervisory service.
Although 21 states report the use of topper
sulphate for the control of algae, the greatest
large-scale use is apparently confined to the mid-
western lake states. Minnesota, in 1959, reports
the use of 221, 745 pounds of copper sulphate in
the treatment of 17, 574 acres of water. In Wis-
consin, 6, 270 acres on 29 lakes received 54, 765
pounds of copper sulphate for algal control. Other
states report the use of lesser amounts. The ma-
jority of this chemical, at the present time, is
distributed through the use of pumps powered by
gasoline engines and transported by boats or
barges.
REFERENCES
Anonymous. 1939. Chemical treatment of lakes
and streams. Wisconsin Committee on Water Pol-
lution, Wis. State Board of Health.
Anonymous. 1950. Chemical Engineers' Hand-
book. Textbook Edition, McGraw Hill Book Com-
pany, p. 1426.
Anonymous. 1959. Great Lakes Newsletter. Great
Lakes Commission, Rackham Bldg., Ann Arbor,
Michigan ^(6): 1-10, July 1959.
Antonie, C.J. and W. H. Osness. 1959. Toxicity
and accumulation of copper as used in the copper
sulfate treatment of lakes. West High School,
Madison, Wis. (Mimeo)
Bartsch, Alfred F. 1946. Aquatic nuisance con-
trol in Wisconsin. Wisconsin Committee on Water
Pollution, Madison, Wis. 35 pp.
Bartsch, Alfred F. 1954. Practical methods for
control of algae and water weeds. Public Health
Reports 69(8): 749-757.
Davidson, Floyd F. 1959. Poisoning of wild and
domestic animals by a toxic waterbloom of Nostoc
rivulare Kuetz. Jour. Amer. Water Works Assoc.
51(10): 1277-1287.
Domogalla, Bernard. 1926. Treatment of algae
and weeds in lakes at Madison, Wisconsin. Engi-
neering News-Record, pp. 3-7, Dec. 9, 1926.
Domogalla, Bernard. 1935. Eleven years of
chemical treatment of the Madison lakes - its
effects on fish and fish foods. Trans. Amer. Fish.
Soc. 65: 115-120.
Domogalla, Bernard. 1941. Scientific studies and
chemical treatment of the Madison lakes. A Sym-
posium on Hydrobiology, University of Wisconsin
Press, Madison, Wis., pp. 303-310.
Doudoroff, Peter and Max Katz. 1953. Critical
review of literature on the toxicity of industrial
wastes and their components to fish. n. The
metals, as salts. Sewage and Ind. Wastes 25(7):
802-839. ~
Ellis, M. M. 1937. Detection and measurement of
stream pollution. Bulletin No. 22, U. S. Bur. of
Fisheries; Bull. Bur. Fisheries 48: 365-437.
Fitzgerald, George P., Gerald C. Gerloff, and
Folke Skoog. 1952. Studies on chemicals with se-
lective toxicity to blue-green algae. Sewage and
Ind. Wastes 24(7): 888-896.
Fitzgerald, George P. and Folke Skoog. 1954.
Control of blue-green algae blooms with 2, 3-Di-
chloronaphthoquinone. Sewage and Ind. Wastes,
26(9): 1136-1140.
Fleming, Virgil R. 1918. Fire streams from
small hose and nozzles. University of Illinois,
Bulletin No. 105 XV(37): 43-60.
Hale, F. E. 1954. Use of copper sulphate In con-
trol of microscopic organisms. Phelps Dodge Re-
fining Corp., New York, 30 pp., 6 plates.
Hasler, Arthur D. 1947. Antibiotic aspects of
copper treatment of lakes. Wis. Acad. Sci., Arts
and Lett., 39j 97-103.
Hooper, F. F. and K. G. Fukano. 1955. Labora-
tory and field experiments with new algaecides,
1944-1955. Report No. 1461 of the Institute for
Fisheries Research, Michigan Dept. of Cons.,
Ann Arbor, Mich.
Ingram, W. M. and G. W. Prescott. 1954. Toxic
fresh-water algae: Summary article of literature.
Amer. Mid. Nat. j>2(l): 75-87.
Johnson, Leon D. 1955. Control of Ulothrix zonata
in circular ponds. Prog. Fish Culturist r?(3):126-
128.
-------
154_
ALGAE AND METROPOLITAN WASTES
Lawrence, J.M. 1954. Control of branded alga,
Plthophora, in farm fish ponds. Fish Culturist 14
(2): 83-88.
Mackenthun, Kenneth M. 1952. Cleaner lakes can
be a reality. Wis. Cons. Bulletin 17(1): 1-4.
Maffcpnth"", Kenneth M. 1958. The chemical con-
trol of aquatic nuisances. Wis. Committee on Water
Pollution, Madison, Wis. , 64 pp.
Mackenthun, Kenneth M. 1959. Summary of aquatic
weed and algae control research and related activi-
ties in the United States. Wis. Committee on Water
Pollution, Madison, Wis., pp. 1-14. (Mimeo)
Elmer F. Herman, and
A heavy mortality of
Kenneth M. ,
Alfred F. Bartsch. 1948.
fishes resulting from the decomposition of algae in
the Yahara River, Wisconsin. Trans. Amer. Fish.
Soc. , 75: 175-180, (1945).
Mackenthun, Kenneth M. and Harold L. Cooley.
1952. The biological effect of copper sulphate treat-
ment on lake ecology. Trans. Wis. Acad. Scl.,
Arts and Lett. 41: 177-187.
Maloney, Thomas E. and C. Mervln Palmer. 1956.
Tozlcity of six chemical compounds to thirty cul-
tures of algae. Water and Sewage Works 103(10):
509-513.
Marsh, M.C., and R.K. Robinson. 1910. The
treatment of fish-cultural waters for the removal of
algae. Bull. Bur. Fisheries 28 (Part 2), 871-890,
(1908).
Matheson, D.H. 1952. The effects of algae In water
supplies. General report prepared for the 1952
Congress for the International Water Supply Assoc. ,
Parts, 82 pp.
Manic, W.D. 1956. Algae control with copper sul-
fate. Water and Sewage Works 103(9); 392-397.
Moore, George T. and Karl F. Kellerman. 1904.
A method of .destroying or preventing the growth of
algae and certain pathogenic bacteria in water sup-
plies. Bulletin 64, Bureau of Plant Industry, U. S.
Dept. of Agriculture.
Moore, George T. and Karl F. Kellerman. 1905.
Copper as an algiclde and disinfectant in water
supplies. Boll. Bureau of Plant Industry, U. S.
D. A. 76: 19-55.
Moyle, John B. 1946. Some indices 01 iase pro-
ductivity. Trans. Amer. Fisheries Soc. 76: 322-
334.
Moyle, John B. 1949. The use of copper sulphate
for algae control and its biological implications.
Limnological Aspects of Water Supply and Waste
Disposal. Pub. of the Amer. Assoc. for the Adv.
ofSci., Washington, D. C., 79-87.
Nichols, M. Starr, Theresa Henkel and Dorothy
McNall. 1946. Copper in lake muds from lakes of
the Madison area. Trans. Wis. Acad. Sci. 38:
333-350.
O'Donnell, D. John. 1945. Control of Hydro-
dictyon reticulatum in small ponds. Trans. Amer.
Fisheries Soc., 73: 59-62, 1943.
Palmer, C. Mervln and Thomas E. Maloney.
1955. Preliminary screening for potential algi-
cides. Ohio Jour, of Sci. 55(1): 1-8.
Prescott, G.W. 1948. Objectionable algae with
reference to the killing of fish and other animals.
Hydrobiological, 1-13.
Rose, Earl T. 1954. Blue-green algae control at
Storm Lake. Proceedings of Iowa Acad. of Sci.,
61: 604-614.
Rushton, W. 1924. Biological notes. Salmon and
Trout Magazine, No. 37, 112-125.
Sawyer, C. N., J. B. Lackey, and A. T. Lenz.
1945. An investigation of the odor nuisance oc-
curring In the Madison lakes, particularly Monona,
Waubesa, and Kegonsa from July, 1943 - July,
1944. Kept. Governor's Committee, 92 pp., 25
fig., 27 tables.
Schaut, G. G. 1939. Fish catastrophes during
droughts. Jour. Amer. Water Works Assoc.,
31(5): 771-882.
Shoenfeld, C. 1947. Don't let 'em spray. Field
and Stream, No. 4: 46, 79, 80, 81, August 1947.
Shoenfeld, C. 1950. The case against
Hunting and Fishing, 11, 12, July 1950.
copper.
Senior, V. E. 1960. Algal poisoning in Saskatche-
wan. Canadian J. of Comparative Medicine and
Veterinary Science 24(1): 26-30, Jan. 1960.
-------
RESEARCH NEEDS
-------
RESEARCH NEEDS IN WATER QUALITY CONSERVATION
BERNARD B. BERGER, Chief
Research Branch, Division of Water Supply and Pollution Control
I should like to thank the Program Committee
for approving the subject of this talk. Such as-
signments force us to contemplate our position
with respect to the adequacy of the tools and tech-
niques of the profession. This is a hard and
sometimes thankless task—but it is an essential
function of the Robert A. Taft Sanitary Engineering
Center.
The SEC is in an advantageous position for se-
curing such information because of the many re-
quests we receive for assistance on difficult and
unusual problems. Although our relatively small
resource permits us to respond to only a few of
these requests, knowledge of important new prob-
lems is literally forced on us. This is desirable
from our point of view. Not only do we learn of
emerging problems, we obtain guidance as to the
direction our research should take. I propose to
base my discussion largely on the information so
obtained.
We find that our uncertainty touches on every
aspect of a waste's origin, behavior and ultimate
fate. Proceeding in logical fashion from the ori-
gins of waste to ultimate water reuse, our broad
areas of ignorance are encompassed by the follow-
ing questions:
Do we account for all pollution reaching
streams?
Having a new waste in hand, can we predict its
potential polluttonal effects?
Assuming we can determine tolerance levels
for this waste, can we really treat it effectively?
In disposal of waste effluents, do we make ef-
fective use of the receiving water?
Having disposed of the waste effluent, can we
monitor effectively the quality of waters which
have received it?
Can we effectively treat polluted water for re-
use?
Each of these questions offers its major chal-
lenge to research.
ORIGIN OF WASTES
Do we account for all pollution reaching
streams?
When we think of pollution, we normally con-
sider sewage and industrial wastes. This is not
surprising. One expects an industrial plant to
produce an industrial waste just as a community
produces sewage. But only recently have we begun
to consider the growing pollutional effect of land
drainage, particularly that from agricultural
lands. Even more recently have we become aware
of pollution resulting from chemical poisons de-
liberately applied to the water course to kill
weeds, insect larvae, fish, and other forms of
aquatic life. We thus have at least four major
avenues through which man-made pollution may
reach streams: sewage, industrial wastes, land
drainage, and direct chemical applications. Each
warrants close study by the sanitary engineer.
1. Sewage which is assuming new, important
characteristics. For example, synthetic organic
chemicals are often undetected and unsuspected
components of sewage. It is probable that the
presence of alkyl benzene sulfonate, the basic in-
gredient of household synthetic detergent, would
be unsuspected if it were not associated with foam.
One could speculate on the presence of other sub-
stances, equally resistant to treatment and equally
persistent in the stream, which give no physical or
chemical clues to their identity.
2. It is not necessary to belabor the point that
the "traditional" industrial wastes, the dinosaurs
of our experience, are still major problems. How-
ever, new and perhaps even more troublesome
wastes are being encountered. Increasing interest
is being directed to radioactive wastes from uran-
ium ore refining plants and nuclear power reac-
tors, thermal pollution from power plants and in-
dustrial cooling operations, and the vast complex
of waste substances discharged by the chemical
industries. Synthetic organics are a particular
problem because of the very rapid growth of the
petrochemical industries. Many of these organic
compounds resist destruction by the communities
of microorganisms on which we depend for or-
156
-------
Research Needs
157
ganic waste stabilization in the treatment plant
and in the stream. They may persist almost in-
definitely in the stream and reach locations of
water use many miles below the point of waste
discharge. An interesting example is a benzene
derivative we followed 1000 miles down the Mis-
sissippi River.
3. The increasing use of agricultural poisons,
particularly insecticides, is reflected in the ap-
pearance of these chemicals in bodies of water
receiving farm land drainage. DDT has frequently
been recovered from streams, and even from
treated water supplies. One of the new chlorinated
hydrocarbon insecticides is the most toxic ma-
terial yet encountered for fish. It has been used
in Pennsylvania to destroy rodents in orchards.
Like other such substances, it is persistent in
water solution and is not easily removed by con-
ventional water treatment processes. As materi-
als of increased potency come on the market,
public concern with their impact on water use
must inevitably grow. One aspect of this problem
is to determine how to control a waste whose
source is general land drainage rather than a spe-
cific outfall sewer.
4. The growing use of synthetic organic poi-
sons to destroy nuisance water vegetation, mos-
quito larvae, and undesirable fish has introduced
new water quality control problems. Dickinson,
North Dakota, provided us with a typical case.
To destroy worthless coarse fish prior to restock-
ing with game fish, it was proposed that a fish
poison be applied to the municipal raw water im-
poundment in use. As one would expect, the health
authorities were concerned with the ability of the
water treatment plant to prevent this material
from reaching the treated water. With the in-
creasing recreational appeal of stored waters, it
is clear that problems of this kind will be experi-
enced widely and frequently.
Is our concern with these synthetic organics
exaggerated? Of the many thousands of com-
pounds of industrial origin reaching waterways
daily, we have information on only a few. How-
ever, we can't help being impressed by the low
concentrations in which these materials may exert
their pollutional effects. For example, I have
mentioned a certain chlorinated hydrocarbon in-
secticide which will kill fish in a concentration of
less than 1 ppb. Phenols in concentration of just
a few ppb have been implicated in taste and odor
problems in drinking water. Alkyl benzene sul-
fonate has been associated with frothing in con-
centrations of about 1 ppm. Ignorance of the ef-
fects of other compounds does not certify their
harmlessness.
The possibility of toxicity for humans is a
dominating concern. However, we know of no
case of human illness in which synthetic organic
substances in a public water supply have been in-
volved. Yet, we cannot escape the fact that as the
number and size of plants engaged in synthetic
organic chemical manufacture increase, their
wastes will increase in flow and complexity. No
matter how slowly the concentrations build up in
receiving waters, this situation may have ultimate
public health implications. Here I should like to
emphasize that it is not the acute episode we fear
—this would be readily identified. It is the in-
sidious effects that could be associated with re-
peated exposure over long periods of time to low
concentrations of the waste.
POLLUTIONAL CHARACTERIZATION OF A NEW
WASTE
Having a new waste in hand, can we predict its
potential pollutional effects?
Can we confidently predict the effect of the
waste on the dissolved oxygen of the stream, its
toxic effect on fish and other aquatic life, its
tainting effect on fish flesh, its possible damage
to water treatment processes, its effect on water
palatability, and its harmful effect on persons who
drink that water?
I think you will agree that few agencies are
presently equipped to do more than estimate the
waste's probable effect on the dissolved oxygen of
the stream,and to determine, within rough limits,
its toxic effect on fish in the receiving water.
Several years ago, the Center undertook to
evaluate its ability to make such predictions. We
selected a group of six organic cyanides as the
wastes of challenge. Chemists, engineers, bi-
ologists, and toxicologists were assigned to de-
termine the pollutional character of these com-
pounds. They were instructed to exercise
ingenuity wherever standard methods and equip-
ment could not be used. Improvisation was
necessary. For example, it was found that con-
ventional organic analysis could not detect nor
measure the low concentrations of compounds
found to be significant. Quantitative analysis
based on fish bioassay solved this particular
problem.
Our experience indicates that a well-equipped
and well-staffed laboratory can obtain the infor-
mation needed to predict with confidence a new
waste's probable impact on certain important
downstream water uses. However, it is an ex-
pensive, time-consuming project not one to be
undertaken as a routine task. The toxicological
phase of the study is perhaps its most perplexing
aspect. The specialized services and cost neces-
sary for determining the effect of repeated expo-
sure to low concentrations of the waste for long
periods of time would inevitably place this job out
of reach of most public agencies. Equally dis-
couraging, perhaps, is the probability that the
toxicological study may take as long as two years.
-------
158
ALGAE AND METROPOLITAN WASTES
WASTE TREATMENT
Assuming we can determine tolerance levels
for this waste, can we really treat it effectively?
Many yardsticks of waste strength and charac-
ter are in use: Biochemical Oxygen Demand
(BOD) is unquestionably the most common. With
the best treatment processes available, a BOD
reduction of approximately 95% may be obtained.
A reduction in BOD does not mean that the objec-
tionable mineral salts—nitrates and phosphates
—have been removed. Just the opposite effect is
produced in sewage stabilization by secondary
treatment. Likewise, the soluble inorganic con-
stituents of wastes are not removed by conven-
tional secondary waste treatment processes. And
it has already been noted that many persistent or-
ganics discharged in sewage and in chemical
industry wastes do not have a BOD. They are
either non-oxidizable or they exert their oxygen
demand so slowly as to be considered non-
oxidizable. Obviously, a reduction in a waste's
BOD gives no assurance that such compounds have
been removed or destroyed.
The possibility that so-called complete treat-
ment of wastes may not always be sufficient, was
pointed out by G. E. McCallum, Chief, Division
of Water Supply and Pollution Control, Public
Health Service, at the 1959 Purdue Industrial
Wastes Conference. A new interest has been
stimulated in the future's need for tertiary treat-
ment processes and, beyond that, for ultracleans-
ing of wastes. You may be interested in a few of
the investigations under consideration at the
Center.
1. Fundamental studies on microbial enzyme
systems important in waste treatment. The po-
5 important
of biologic
tential of biological communities for breaking
down organic compounds should be fully realized.
This naturalistic treatment must, in our opinion,
be exploited to the maximum because it costs less
than other methods, and because it is a common!
sense preliminary to any necessary treatment of
organic substances by non-biological methods.
Research may yield valuable information on how
to break down certain kinds of chemical structures
now invulnerable to microbial attack and how to
stimulate development of more adaptable microbial
enzyme systems.
2. Fundamental studies on particle separation
by filtration. The ultimate potential of filters for
particle removal should be established. The re-
lationship of filter design to the size of particles
to be removed has not hitherto been adequately in-
vestigated. Because of the obvious usefulness of
filters at points of waste production,and their very
wide employment at points of water use, we should
know the "innate value of this unit process. As
part of this project, methods could be explored for
producing chemical floes of controlled size, den-
sity, and toughness. Such floes deposited on
filters of various kinds will permit the realization
of maximum filter potential.
3. Application of adsorption principle to waste
treatment. Effective treatment of complex organic
wastes may require, in many cases, the intelligent
use of the principle of adsorption. For a number
of years, the Center has studied intensively the
adsorption of soluble organics from sewage and
industrial waste streams and receiving waters.
Many materials have been evaluated for their ad-
sorption properties, with special attention directed
to activated carbon which has been found, so far,
to be the adsorbant of choice. In the summer of
1958, a special evaluation was made of the factors
influencing adsorption of organic contaminants.
The problem of regeneration of adsorbants is of
paramount importance.
4. Destruction of organics by strong chemical
oxidants. The feasibility of destroying the organic
content of wastes by strong chemical oxidants
such as ozone and hydrogen peroxide should be
investigated. In this way, complex organic com-
pounds may be converted to relatively simple in-
organic structures which might be removed by
some appropriate procedure.
Other physical-chemical phenomena may be
useful in waste treatment. Advanced methods of
freezing, ion exchange, solvent extractions, and
foam fractionation should also offer interesting
research opportunities in water cleansing.
WASTE DISPOSAL
In disposal of waste effluents, do we make ef-
fective use of the receiving water?
Presumably, available knowledge and experi-
ence on lateral mixing and dispersion of wastes in
streams should permit full use of the stream's
dilution ability. However, streaks of waste may
often be traced many miles along a water course.
Obviously, the full dilution effect is not being used.
This often poses stream sampling problems be-
cause of variation in waste concentration in a
cross-section of a stream. It also raises perti-
nent questions regarding the adequacy of outfall
design techniques.
Many of you have personal knowledge of the
relatively poor state of the art of waste disposal
into tidal waters. A profound question is, "What
is the fate of a particle of waste discharged into a
mass of tidal water?" The intelligent use of
coastal waters and estuaries for waste disposal
was the subject of an International Conference
held last year at the University of California in
Berkeley. Every one of the 24 coastal and Gulf
States is seeking, or will inevitably seek, effec-
tive techniques to answer this question. The use
of models to supplement field studies is a recent
interesting development.
-------
Research Needs
159
WATER QUALITY MONITORING
Having disposed of the waste effluent, can we
monitor effectively the quality of waters which
have received it?
Until recently, few programs of water quality
intelligence could present a realistic picture of
changing quality conditions in a stream, and those
were usually developed at great expense in special
problem situations. You may recall that in De-
cember, 1955, the unsatisfactory condition re-
garding knowledge of the quality of our water
impelled the President's Advisory Committee on
Water Resources to note that improvement in
basic data on water quality was urgent.
The monitoring problem has three important
phases: to define what constitutes meaningful data
for a given stream; to apply appropriate sampling
and analytical procedures; and to fit these data
into a scheme of known water uses, waste dis-
charges, and stream flow to permit correct inter-
pretation of data. Obviously, mere collection of
data is not helpful if they do not truly reflect
changes in water quality.
We are not badly off with respect to detection
of certain types of pollution. Recent improvements
in sampling and analytical techniques for coliform
organisms and enterococci permit us to discern
fecal contamination more readily than before. The
successful development of equipment for continu-
ous recording of dissolved oxygen represents a
significant advance in the monitoring of streams
for oxygen demanding wastes. The current em-
phasis on radioactivity monitoring is providing
steadily improved instrumentation for this pro-
gram. However, many important shortcomings
remain.
Perhaps the most significant defect in data
collection programs is the inadequacy of proced-
ures for detecting pollution by highly stable or-
ganic compounds of industrial origin. A good
example is the compound described earlier which
could be recovered 1000 miles below its point of
introduction into the stream. Its presence was
suggested by an unusual peak in the infrared ab-
sorption spectrum. Only by using the most ad-
vanced techniques of sampling and analysis may
we hope to detect such substances. An additional
shortcoming is the failure to take full advantage of
stream biota as useful indicators of pollution.
Although biologists have repeatedly stressed the
sensitivity of aquatic life to water quality changes,
our approach remains relatively crude and expen-
sive.
It must be re-emphasized that collection of data
is not important if they do not relate to water
quality. The time is ripe for close, critical
scrutiny of the kind of information now being re-
corded in water quality basic data programs.
WATER TREATMENT
Can we effectively treat polluted water for re-
use?
The main function of the modern water treat-
ment plant is to remove particles suspended in the
water and to destroy microorganisms that manage
to survive flocculation and filtration procedures.
Except for removal of hardness, little attempt is
made to remove soluble chemicals in drinking
water. Where taste and odor are highly objection-
able, carbon is usually applied. But carbon is
selective in its action and sometimes it is ineffec-
tive. We actually know little about the principle
of adsorption as it applies to the use of carbon.
Moreover, there is no assurance that harmful
substances in solution will signify their presence
by producing taste and odor. Here, as in waste
treatment, interesting research opportunities
exist.
CONCLUSION
My assignment has been to consider major
needs of water quality research. I have addressed
my remarks to the following questions:
1. Do we account for all pollution reaching
streams?
2. Having a new waste in hand, can we pre-
dict its potential pollutional effects?
3. Assuming we can determine tolerance
levels for this waste, can we really treat it effec-
tively?
4. In disposal of waste effluents, do we make
effective use of the receiving water ?
5. Having disposed of the waste effluent, can
we monitor effectively the quality of waters which
have received it?
6. Can we effectively treat polluted water for
reuse?
These, I believe, encompass the important
areas of concern to agencies responsible for water
quality conservation.
Perhaps in stressing areas of ignorance, I have
been guilty of painting an unduly gloomy picture of
the state of our art. We need not apologize. The
success of sanitary engineers is controlling filth
in water and, by so doing, eliminating the epi-
demic diseases associated with filth — one of the
great triumphs of public health. We were able to
solve the problems of the past. We may look for-
ward with confidence to solving the problems now
facing us.
-------
PROGRAM
Wednesday, April 27, 1960
8:15 - 8:55 Registration and Assembly
8:55 - 9:00 Opening Remarks:
A. F. Bartsch, Assistant Chief, Research Branch,
Division of Water Supply and Pollution Control
9:00 - 9:10 Welcome:
H. G. Hanson, Director,
Robert A. Taft Sanitary Engineering Center
9:10 - 12:30 STATEMENT OF THE PROBLEM:
Chairman: A. F. Bartsch
Introduction
Induced Eutropnication — A Growing Water Resource Problem:
A. F. Bartsch
Specific Problems In Lakes
Madison's Lakes: Must Urbanization Destroy Their Beauty and Productivity?
William B. Sarles, Department of Bacteriology,
University of Wisconsin, Madison, Wisconsin
Phosphate from Treated Sewage and Algae Blooms in Lake Zoar, Connecticut:
Richard J. Benoit, Research & Development,
General Dynamics Corporation, Groton, Connecticut and
John J. Curry, Water Resources Commission,
Hartford, Connecticut
Klamath Lake, an Instance of Natural Enrichment:
Harry K. Phinney, Department of Botany & Plant Pathology,
Oregon State College, Corvallis, Oregon
Recent Changes in the Trophic Nature of Lake Washington:
G. C. Anderson, Research Department of Oceanography,
University of Washington, Seattle, Washington
Specific Problems In Rivers
Algae in Rivers of the United States:
C. M. Palmer, In Charge, Interference Organisms Studies,
Research Branch, Division of Water Supply
and Pollution Control
1:30 - 4:30 GROWTH CHARACTERISTICS OF ALGAE:
Chairman: Jack Myers, Department of Zoology,
University of Texas, Austin, Texas
Nutrition
Fundamental Characteristics of Algal Physiology:
Robert W. Krauss, Department of Botany,
University of Maryland, College Park, Maryland
Micro-Nutrients and Heterotrophic Requirements of Algae as Possible Factors
in Bloom Production:
Luigi Provasoli, Raskins Laboratory, 305 E. 43rd St.
New York, New York
Algal Density as Related to Nutritional Thresholds:
James B. Lackey, Department of Civil Engineering,
University of Florida, Gainesville, Florida
Productivity And How To Measure It
Methods of Measurement of Primary Production in Natural Waters: -
Lawrence R. Pomeroy, Marine Institute,
University of Georgia, Athens, Georgia
Factors Which Regulate Primary Productivity and Heterotrophic Utilization
in the Ecosystem:
Eugene P. Odum, Department of Zoology,
University of Georgia, Athens, Georgia
160
-------
Program 161
Thursday, April 28, 1960
9:00 - 12:30 SOURCES OF NUTRIENTS:
Chairman: Harold B. Gotaas, Northwestern
Technological Institute, Northwestern University,
Evanston, Illinois
Land Drainage
Land Drainage as a Source of Phosphorus in Illinois Surface Waters:
R. S. Engelbrecht & James J. Morgan, Department of Civil
Engineering, University of Illinois, Urbana, Illinois
Urban, Agricultural and Forested Areas:
R. O. Sylvester, Department of Civil Engineering,
University of Washington, Seattle, Washington
Wastes
Metropolitan Wastes and Algal Nutrition:
William J. Oswald, School of Public Health, University
of California, Berkeley, California
Limnological Relationships
The Role of Limnological Factors in the Availability of Algal Nutrients:
George H. Lauff, Department of Zoology,
University of Michigan, Ann Arbor, Michigan
Nitrogen Fixation in Natural Waters under Controlled Laboratory Conditions:
C. N. Sawyer & Alfred F. Ferullo, Metcalf & Eddy,
Engineers, Statler Building, Boston, Massachusette
Recent Observations on Nitrogen Fixation in Blue-Green Algae:
Richard C. Dugdale, Department of Biological Sciences,
University of Pittsburgh, Pittsburgh, Pennsylvania
1:30 - 4:00 METHODS OF PREVENTION OR CONTROL:
Chairman: M. A. Churchill, Stream Pollution
Control Section, Division of Health & Safety,
Tennessee Valley Authority, Chattanooga, Tennessee
Limitation of Nutrients As A Step In Ecological Control
The Madison Lakes Before and After Diversion:
Gerald W. Lawton, Hydraulic & Sanitary Laboratory,
University of Wisconsin, Madison, Wisconsin
The Badfish River Before and After Diversion of Sewage Plant Effluent:
Theodore F. Wisniewski, Committee on Water Pollution,
Madison, Wisconsin
Spray Irrigation for the Removal of Nutrients in Sewage Treatment Plant
Effluent as Practiced at Detroit Lakes, Minnesota:
Winston C. Larson, Winston C. Larson and Associates,
Detroit Lakes, Minnesota
Chemical Methods for Removal of Phosphorus and Nitrogen from Sewage Plant
Effluents:
Gerard A. Rohlich, Hydraulic & Sanitary Laboratory,
University of Wisconsin, Madison, Wisconsin
Stripping Effluents of Nutrients by Biological Means:
George P. Fitzgerald, College of Engineering,
University of Wisconsin, Madison, Wisconsin
The Use of Algae in Removing Nutrients from Domestic Sewage:
R. H. Bogan, Department of Civil Engineering,
University of Washington, Seattle, Washington
5:30 - 6:30 Social Hour, Hotel Sinton
6:30 - 8:30 Banquet
Do They Dig You, Daddy-O, Or Are You Way Out?
R. G. Lynch, The Milwaukee Journal, Milwaukee, Wisconsin
-------
162 ALGAE AND METROPOLITAN WASTES
Friday, April 29, 1960
Chairman: Frank Woodward, Division of Environmental
Sanitation, State Department of Health,
Minneapolis, Minnesota
9:30 - 10:30
Use of Alglcides
The Practical Use of Available Algicides and Modern Trends Toward New Ones:
Kenneth M. Mackenthun, Committee on Water Pollution,
Madison, Wisconsin
10:30 - 11:00 RESEARCH NEEDS
Research Needs in Water Quality Conservation:
Bernard B. Berger, Chief, Water Supply and Water Pollution
Research, SEC
11:00 - 11:05 Concluding Remarks:
A. F. Bartsch
11:15 - 12:30 Tour of SEC Facilities:
H. W. Jackson, Biologist, Water Supply and Water
Pollution Training Training Program, SEC
-------
|