United States
                   Environmental Protection
                   Agency
Environmental Research
Laboratory
Duluth MN 55804
                   Research and Development
EPA-600/S3-83-075 Nov. 1983
x°/ERA         Project  Summary
                   Effects of  Phosphorus  Loading
                   on  Phytoplankton  Distribution
                   and  Certain  Aspects  of
                   Cytology  in  Saginaw Bay,  Lake
                   Huron
                   E.F. Stoermer, L Sicko-Goad, and LC. Frey
                     Saginaw Bay has always been one of
                   the more productive regions of the Great
                   Lakes system. At the present time, it is
                   also one of the most modified. Excessive
                   nutrient  and conservative element
                   loadings are factors which have led to
                   severe perturbation of primary producer
                   communities in the region. Because of
                   the physical dynamics of the bay region,
                   idealized dilution gradients are grossly
                   modified by transport of water masses
                   and their entrained chemical consti-
                   tuents, fauna and flora into, as well as
                   away from, the Bay. However, there
                   appears to  be considerable selection
                   among population components of the
                   assemblages transported. For example,
                   blue-green algae appear to be conserved
                   in the Bay while diatoms are subjected
                   to great losses.
                     The major effort in this investigation
                   was to provide data on phytoplankton
                   biovolume that would support a model
                   of processes occurring in Saginaw
                   Bay. A method of estimating the actual
                   viable fraction of the cell volumes of
                   representatives of the various physio-
                   logical groups of phytoplankton found
                   in Saginaw Bay was developed,  and
                   polyphosphate  body formation was
                   studied. Results showed that substan-
                   tial phytoplankton  populations were
                   exported from the Bay to Lake Huron.
                   Under average wind conditions, most
                   export occurred along the southern
                   coast. These populations were then
                   entrained in the general Lake  Huron
                   circulation and were spread down the
Michigan coast southward from the
Bay. Under certain advective conditions,
however, phytoplankton  were dis-
charged from the Bay either to the north
or directly offshore.
  Cytological analysis showed that
many species sequestered phosphorus
in excess of their immediate  physiologi-
cal needs, in the form of polyphosphate
bodies. Populations exported from the
Bay also contained these polyphosphate
bodies. Analysis of the polyphosphate
bodies showed that significant quanti-
ties of certain toxic  metals,  notably
lead, were  incorporated  into these
inclusions.
  Analysis of the relationship of total
phytoplankton cell  volume to proto-
plasmic constituent volume  showed
that crude cell volume measurements
furnished a poor estimate of actual
living biomass in many populations. It
was concluded that more  refined
techniques are required to correctly
convert  estimates of cell  number to
estimates of biomass.
  This Project Summary was developed
by EPA's Environmental Research
Laboratory, Duluth. MN. to announce
key findings of the research project that
is fully documented in a separate report
of the same title (see Project Report
ordering information at back).

Introduction
  Saginaw Bay has always been one of
the more productive regions within the
Great Lakes system. The productivity of

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the fishery resource was undoubtedly
one  of the factors attracting early
settlement  in  the area.  Other natural
resources of the drainage basin provided
the incentive for early settlement  and
substantial economic growth in the
region.  The timber  resources of the
Saginaw River and its tributaries were
rich and easily accessible, which led to
early development of the area and the
establishment of an industrial base. Once
cleared of its natural vegetation, much of
the land was found suitable for intensive
agricultural practices. Finally, the pres-
ence of subsurface resources,  primarily
petroleum and salt, made possible the
establishment  of one of  the midwest's
centers  of  chemical  industry.  Unfortu-
nately, the development of the Saginaw-
Bay City-Mid I and industrial complex and
intensive regional agriculture proceeded
at the expense of deterioration of water
quality  within Saginaw Bay. At the
present  time it  is one of the most
seriously modified parts of the Great Lakes
system.  During the past few decades
Saginaw Bay has been beset with water
quality  problems including  obnoxious
algal blooms, taste and odor problems in
municipal water supplies, and fish flesh
tainting. It should be pointed out that the
perturbation of primary  producer com-
munities in this region reflects the effects
of many factors.  The most  obvious of
these are the effects of excessive nutrient
loadings. At the present time the waters
of Saginaw Bay are probably the most
productive  in  the entire Great  Lakes
system.  The composition of the  phyto-
plankton and  benthic  algal flora also
reflects the effects of extreme conserva-
tive element  loadings.  Although these
loadings have apparently been decreased
to some degree in recent years, the flora
of the Bay still contains many elements
usually found in brackish water localities.
Finally, although not  experimentally
documented, certain population distribu-
tions within the Bay can most plausibly be
explained by direct toxic effects.
  Saginaw Bay is also an extremely dynamic
system. There are strong gradients in
almost all factors of physiological interest
between the lower Bay  and the open
waters  of  Lake  Huron.  As  might be
expected, these gradients are reflected in
the population and community responses
of the  phytoplankton  flora. Idealized
dilution gradients are grossly modified by
mass transport  of water masses  and
their entrained chemical constituents,
fauna, and flora into, and  away from, the
Bay. There appears to be considerable
selection among the  population compo-
nents of the assemblage(s) being trans-
ported.  Certain populations, primarily
blue-green algae, appear to be conserved,
in the sense  that their  abundance  is
highly correlated with the concentration
of biologically conservative chemical
elements being discharged from the Bay.
Other populations, primarily diatoms, are
apparently subjected to  much greater
losses during transport At the same time,
water masses from Lake Huron, containing
biological communities adapted to phys-
ical and chemical conditions found in the
open lake, are imported  into the Bay. The
purpose of this large scale investigation
was to  provide data on  phytoplankton
biovolume which would support a model
of processes occurring  in  the Bay.
Biovolumes  were estimated  and poly-
phosphate bodies were examined  in
many species.
  Parts  of this investigation were pre-
viously reported, namely:  studies  of
primary consumer organisms in Saginaw
Bay and southern Lake Huron (Gannon,
in prep; Stemberger et al. 1979), studies of
physical  and  chemical  conditions  in
Saginaw Bay (Smith et al. 1977), a study
of phytoplankton abundance and distri-
bution in southern Lake Huron (Stoermer
and Kreis, et al. 1980) a process oriented
model of Saginaw Bay (Biermann, et al.
1980). Although each of these  studies
was  the  responsibility of a different
laboratory, common sampling techniques
were  used. Pertinent data are reproduced
in this report, detailed methods are given
only  for  bivolume  calculations  and
polyphosphate body studies.
  In most instances phytoplankton samples
were  taken  by submersible  pump and
fixed immediately with 4%gluteraldehyde.
Samples were kept on ice in the dark until
they were processed into slides  by the
membrane clearing technique. Sampling
cruises were conducted at approximately
monthly intervals over a 2-year period.
  Population estimates were developed
from  1  mm  strip counts of randomly
selected areas of the slide preparations.
Populations  were enumerated and di-
mensions of  the taxa recorded. The
volume  data  were  converted  to an
estimate of carbon  biomass. Volume
density estimates of cell components (cell
wall, chloroplasts, vacuoles, storage
products, and remaining cytoplasmic
material) were obtained by electron
microscope morphometric  methods.
Estimates of volume density were deter-
mined using both paper profile cut-and-
weigh and grid-point-counting techniques.
  Eight taxa were selected for quantitative
stereological  analysis,  including  repre-
sentatives from the classes Cyanophyceae,
Bacillariophyceae,  Euglenophyceae, Cryp-
tophyceae, Dinophyceae,  and Prymne-
siophyceae. Seventy-five micrographs
were examined for each  taxon, except
where scarcity of a particular taxon in the
water samples necessitated a somewhai
reduced  sample size.  In  all  cases,
examination of coefficients of variation
and plots of  cumulative means and
variances indicated adequate sampling ol
the  material.  A transparent 0.5-cm
square sampling lattice was superimposed
over the  micrographs for quantitative
measurements.
  Cell  volume estimates were obtained
from  light microscopic examination of
cells from the same assemblages used for
quantitative stereological analysis.
  Both x-ray energy dispersive analysis
(to confirm the presence of phosphorus)
and  light  microscopy  were used in the
examination of polyphosphate bodies.
  The results of the study reported here
indicate  that the problems associated
with biovolume estimates can be both
understood and, in many cases, minimized.
The  inherent  cellular organization of
some species contributes significantly to
the error of biomass calculations currently
in use.  Corrected  cellular  volumes
determined from the morphometric data
by  subtracting  the  volume  of inert
structures from the total volumes were
found in this study. Table 1 indicates the
striking differences found in various taxa.

Table 1.    Percent of Total Cellular Volume
          Identified as "Metabolizing
          Biovolume" for Selected Taxa
A. flos-aquae
S. binderanus
F. capucina
E. viridis
C. erosa
P. lindemann:
Haptophyte sp. #1
Haptophyte so. #2
64.1
29.9
43.5
44.6
67.7
58.6
58.3
81.9
  Polyphosphate inclusion bodies were
widely found in diatoms and even in algae
in Saginaw Bay. The abundance of these
bodies corresponds to the usual pattern
of greatest total phytoplankton density,
and to the most probable area of excur-
sion of materials from the Bay.

Conclusions and
Recommendations
  1. The  use of electron microscopic
     morphometric analysis leads to a
     more accurate estimate of biovolume
     than other methods presently in
     use.
  2. The  ecological causes  and  conse-
     quences of polyphosphate storage
     in phytoplankton may be important

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     in evaluating impact of areas such
     as Saginaw Bay on the rest of the
     Great Lakes system.
  3. It was previously assumed  that
     polyphosphate  body formation was
     important  only in  prokaryotic or-
     ganisms.  The results of this study
     suggest that  the  mechanism  is
     present in most of the major physio-
     logical groups present in the Great
     Lakes. Notable exceptions were the
     Cryptomonads  and Dinoflagellates.
     Although induction was not attempted
     in species of these groups under
     experimental conditions, none of
     the wild populations examined
     contained polyphosphate  bodies.
     Representatives of all of the other
     major algal physiological groups
     did. Most of the species examined in
     the study are usually associated
     with eutrophied conditions in the
     Great Lakes and it is possible that
     luxury consumption of phosphorus
     is one of the factors which confers
     competitive  advantage on these
     populations.  Further  research will
     be needed to answer this question.
  4. These results  indicate that heavy
     metals may be  sequestered  in
     polyphosphate bodies.
  5. Meteorological conditions  have a
     strong influence on  the  eventual
     fate of materials entering the Bay,
     and on the  fate of phytoplankton
     populations generated  in the  Bay.
     Further research should be devoted
     to the effects  of conditions which
     force  strong advective events on
     processes in Lake Huron.
  chemical factors in Saginaw Bay (Lake
  Huron). U.S. Environmental Protection
  Agency, EPA-600/3-77-125. 143 pp.
Stemberger, R.S.,  Gannon, J.E.  and
  Bricker, F.J. 1979. Spatial and seasonal
  structure  of rotifer communities in
  Lake Huron. U.S.  Environmental  Pro-
  tection Agency, EPA-600/3-79-085.
  160 pp.
Stoermer,  E.F.,  and  Kreis, R.G.  1980.
  Phytoplankton composition and abun-
  dance in southern Lake Huron. Univ.
  Michigan,  Great Lakes Research Divi-
  sion, Special Report No. 65. 382 pp.
   E. F. Stoermer, L Sicko-Goad, and L C. Frey are with the University of Michigan.
    Ann Arbor. Ml 48109.
   Nelson A. Thomas is the EPA Project Officer (see below).
   The complete report, entitled "Effects of Phosphorus Loading on Phytoplankton
    Distribution and Certain Aspects of Cytology in Saginaw Bay, Lake Huron,"
    (Order No. PB 83-250 035; Cost: $14.50, subject to change) will be available
    only from:
          National Technical Information Service
          5285 Port Royal Road
          Springfield. VA 22161
          Telephone: 703-487-4650
   The EPA Project Officer can be contacted at:
          Environmental Research Laboratory
          U.S. Environmental Protection Agency
          6201 Congdon Blvd.
          Duluth.  MN 55804
                                               A-US GOVERNMENT PRINTING OFFICE 1983-659-017/7222
References
Bierman, V.J.,  Dolan, D.M., Stoermer,
  E.F., Gannon, J.E., and  Smith,  V.E.
  1980. The development and calibration
  of a spatially simplified  multi-class
  phytoplankton model for Saginaw Bay,
  Lake Huron. Great Lakes Environmental
  Planning  Study, Contribution No. 33,
  126 pp. Great Lakes Basin Commission,
  Ann Arbor,  Michigan.
Gannon, J.E.,  Bricker, F.J. In preparation.
  Spatial  and  seasonal structure of
  crustacean  zooplankton communities
  in Lake Huron. U.S.  Environmental
  Protection Agency.
Schelske, C.L, Moll, R.A., and Simmons,
  M.S. 1980.  Limnological conditions in
  southern  Lake Huron, 1974 and 1975.
  U.S. Environmental Protection Agency,
  EPA-600/3-80-74.  178 pp.
Smith, V.E.,  Lee,  K.W., Filkins, J.C.,
  Hartwell,  K.W., Rygwelski,  K.R.  and
  Townsend,  J.M.   1977.  Survey of

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