PB 198 063
AICE SURVEY OF USSR AIR POLLUTION LITERATURE.
VOLUME III. THE SUSCEPTIBILITY OR RESISTANCE
TO GAS AND SMOKE OF VARIOUS ARBOREAL SPECIES
GROWN UNDER DIVERSE ENVIRONMENTAL CONDITIONS
IN A NUMBER OF INDUSTRIAL REGIONS OF THE SOVIET
UNION
M. Y. Nuttonson
American Institute of Crop Ecology
Silver Spring, Maryland
December 1969
NATIONAL -ECHNICAL INFORMATION SERVICE
Distributed ,.. Mo foster, serve
and promote the nation's
economic development
and technological
advancement.'
U.S. DEPARTMENT OF COMMERCE
This document has been approved lor public release and sale.
-------
AICE* SURVEY OF USSR AIR POLLUTION LITERATURE
Volume III
THE SUSCEPTIBILITY OR RESISTANCE TO GAS AND SMOKE
OF VARIOUS ARBOREAL SPECIES GROWN UNDER DIVERSE ENVIRONMENTAL CONDITIONS
IN A NUMBER OF INDUSTRIAL REGIONS OF THE SOVIET UNION
Edited By
M. Y. Nuttonson
The material presented here is part of a survey of
USSR literature on air pollution
conducted by the Air Pollution Section
AMERICAN INSTITUTE OF CROP ECOLOGY
This survey is being conducted under GRANT 1 R01 AP00786-01 APC
THE NATIONAL AIR POLLUTION CONTROL ADMINISTRATION
'AMERICAN INSTITUTE OF CROP ECOLOGY
809 DALE DRIVE
SILVER SPRING, MARYLAND 20910
1970
-------
BIBLIOGRAPHIC DATA
SHEET
1. Report No.
APTD-0637
3. Recipient's Accession No.
4. Title
subt.tie
Survey of USSR Air Pollution Literature
Volume III - The Susceptibility or Resistance to Gas and Smoke
of Various Arboreal Species Grown Under Diverse Environmental
r.nnHiHnnc in a Niimhor nf TnHuctHal Rpninnc. nf thp Sovipt Unii
Report Date
December 196
-------
This report was furnished to the
Air Pollution Control Office by
the American Institute of Crop
Ecology 1n fulfillment of Contract
No. AP00786-01.
-------
TABLE OF 'CONTENTS
Page
PREFACE v
USSR ORIENTATION MAP vlii
USSR MAP OF CLIMATIC ZONES AND REGIONS lx
/, THE EFFECT OF INDUSTRIAL GASES AND INDUSTRIAL SMOKE
ON VARIOUS FOREST SPECIES,
P. S. Pogrebnyak 1
- THE INFLUENCE OF SMOKE AND GAS ON THE FLOWERING AND
FRUITING OF SOME TREES AND SHRUBS/ ,
V. G. Antipov 3
f THE EFFECT OF SPECIFIC INDUSTRIAL GASES ON THE GROWTH OF
SOME TREE SPECIES ..'
V. G. Antipov 9
t; REACTION OF TREES AND SHRUBBERY TO AIR POLLUTION IN THE
FOREST-PARK BELT OF MOSCOW AND MEASURES FOR
EXTENDING THE LIFESPAN OF PLANTS ,
A. P. Shcherbakov and I. F. Cherednichenko 13
CURRENT CONDITIONS AND SCIENTIFIC PROBLEMS IN STUDYING
THE INJURIOUS EFFECTS OF INDUSTRIAL POLLUTANTS ON
PLANTS AND IN DEVELOPING METHODS FOR CONTROLLING
THEM IN THE URALS
S. A. Mamaev 23
{' THE DRAINAGE OF TEMPORARILY SWAMPED SOILS IN RELATION TO
PINE PLANTINGS GROWING UNDER CONDITIONS OF
INDUSTRIAL AIR POLLUTION ;
N. V. Podzorov 33
f- THE EFFECT OF INDUSTRIAL SMOKES AND GASES UPON CONIFEROUS
U FORESTS GROWING UNDER CONDITIONS OF INCREASED
HUMIDITY IN THE MOSCOW REGION ("PODMOSKOV'E"),
V. G. Antipov '. 39
| , SOME PECULIARITIES OF THE SUSCEPTIBILITY OF SCOTCH PINE
SPROUTS TO SULFUR DIOXIDE INJURY,
S. A. Mamaev and V. S. Nikolaevskiy AS
GAS RESISTANCE OF PINE AND BIRCH'
Yu. Z. Kulagin '. 51
iii
-------
Page
THE EFFECT OF SULFUR DIOXIDE ON WOODY PLANTS UNDER THE
ENVIRONMENTAL CONDITIONS PREVAILING IN THE
SVERDLOVSK REGION,
V. S. Nikolaevskiy ................................... 58
CHARACTERISTICS OF PHOTOSYNTHESIS AND OF SOME OTHER PROCESSES
IN CONNECTION WITH SMOKE AND GAS RESISTANCE OF TREES
AND SHRUBS (
A. S. Sitnikova ...................................... 6
INDICATORS OF GAS RESISTANCE OF ARBOREAL PLANTS/
(According to Investigations Conducted in The City of
Krasnoural'sk)
V. N. Nikolaevskiy ................................... 70^-
THE ACTIVITY OF CERTAIN ENZYMES AND GAS RESISTANCE OF
WOODY PLANTS J
V. S. Nikolaevskiy 95
VARIATION IN THE OXIDIZABILITY OF THE CELL CONTENT AS ONE
OF THE INDICATORS OF GAS RESISTANCE IN PLANTSJ
V. G. Antipov and I. I. Chekalinskaya 100 /V
EFFECT OF SULFUR DIOXIDE ON THE ENZYMATIC ACTIVITY OF TREE
LEAVES . < r
V. S. Nikolaevskiy 109 (J
iv
-------
PREFACE
Much of Che background material presented in the preface to
Volume II of this series is repeated here in view of its relevance to
the present volume.
Contamination of the natural environment constitutes a major problem
in all industrial regions of the Union of Soviet Socialist Republics
(USSR). The country's industry and transport are continually bringing
about massive qualitative changes in the habitat of man and vegetation
through an ever increasing pollution of air, soil, and streams. In
recent years there has been a greater awareness of the immense problems
of air and water pollution on the part of the urban and rural administra-
tive agencies as well as on the part of various research institutes of
the USSR. There is a mounting demand there to maintain a high quality
physical environment. Protective measures against the pollution threat
are gradually taking shape. Much relevant air pollution research data
are being developed and are apparently put to good use in some parts of
this vast and diverse country.
Reports of investigations brought together in this volume deal with
a number of aspects of the relationship of air pollution and vegetation.
A considerable number of these investigations have been conducted in vari-
ous industrial regions of the USSR, regions that are geographically far
apart from each other and subject to distinctly different natural and man-
made environmental conditions.
Many of the investigations reported in this volume relate to the urban
and rural areas particularly affected by air pollution and a number of
these areas are situated in widely separated industrial regions, the main
ones being:
The Ural Region of RSFSR
The Moscow Region of RSFSR
The Leningrad Region of RSFSR
The Belorussian SSR
The Azerbaizhan SSR
The Kazakh SSR.
The problem of selecting suitable plants for new environmental con-
ditions — whether natural or man-made — has challenged man throughout
the centuries of his migration. Selection of plants for introduction and
establishment in a new environment may be a rather simple hit-or-miss
undertaking, or it may become a painstaking process requiring a knowledge
of plant responses to such potent factors of the physical environment as
photoperlod, light intensity, temperature, humidity, soil moisture, and
soil fertility. It is rather widely accepted now that air pollution, where-
ever it occurs, must be taken into account as another highly potent factor
-------
in the physical environment of plants. The nature of air pollution has
to be considered In the assessment of the environmental conditions pre-
vailing in a given area in relation to the overall problem of plant se-
lection and plant adaptation. A plant, irrespective of Its genetic po-
tentialities and elasticity of adaptation to environmental conditions,
responds to the various stresses of its environment. These include the
stresses of air pollution together with all the other potent factors of
the atmosphere as well as of soil and those of the biological environ-
ment of a plant, notably, pests and diseases, and the quality of manage-
ment or abuse by man.
As will be seen from the data of the papers presented in this volume
a considerable number of studies are being conducted In the USSR in refer-
ence to susceptibility or resistance of different plant species and their
ecotypes to various phytotoxic air pollutants In different parts of the
country.
It must be borne in mind that the data presented in this volume re-
late to many diverse environments in a vast land area; that the USSR ex-
tends for about 7,000 miles from west to east and 3,000 miles from north
to south; and that the country covers a wide range of climatic and soil
conditions throughout much of its north-south and west-east extent. In
this.connection, a brief outline of the very general natural features of
the USSR may be desirable. Lowlands and plains dominate the landscape
of the major portion of the country. Its landscape can be roughly de-
scribed as one consisting of broad latitudinal climate-vegetation-soil
belts of the lowlands and plains and of narrow, vertical climate-vegeta-
tion zones of the highlands and mountains. Each of the broad latitud-
inal belts is distinct from the other in the major features of its clim-
ate, vegetation, and soils, though within each latitudinal belt there is
a decrease in the annual precipitation as one proceeds from west to east.
The latitudinal belts include the nearly barren and treeless tundra In
the extreme north, where the winters are severe, the summers, short and
cool, and where precipitation Is very limited. There follow the belts
of the taiga or coniferous forests, mixed forests, woodlands, forest
prairie or forest steppe, the steppe, and the semi-desert. Finally in
the extreme south, east of Caspian Sea, there are the dry deserts, hot
in summer and cold in winter, and, along the southern reaches of the
Black Sea in Transcaucasia, there is a relatively limited area, humid and
more or less subtropical, which is subject to mild winters, hot summers,
and heavy precipitation.
A considerable number of surveys and studies presented in this vol-
ume deal with the effect of industrial atmospheric pollutants on various
plant species grown in forests, in public parks, in gardens, in street
plantings as well as for sanitary-protection purposes in different cities
and localities at various distances from industrial developments, and
situated differently in respect to their directional position from speci-
fic pollutant sources. The adverse effects of air pollution on some of
the indigenous and introduced plant species — effects often leading to
complete plant destruction, such as for example, the death of ornamental
plantings in urban parks and gardens and the death of forest plantings -
- are described in some of the studies. Susceptibilities to specific
phytotoxic air pollutants and Injury symptoms In different species and
their ecotypes, are also discussed.
vi
-------
A number of papers bring out the fact that the response to air pollu-
tion differs with the plant species as well as with the ecotypes within a
species and that the growth stage of the plant, notably the age of the leaf,
Is an Important factor In determining Its sensitivity to air pollutants.
Much attention is given in quite a few of the papers to the resistance of
various plants to different gases and to smoke, to some of the environmental
conditions as well as to some physiological indices in relation to smoke and
gas resistance of plants, and to the problem of plant selection for resist-
ance to smoke and gas pollution.
The data of several studies and surveys presented in this volume sug-
gest that on the basis of experimental plot tests and field observations
the genetic susceptibility or resistance of certain plant species and eco-
types have been determined in a number of areas. The use of the resistant
plants in industrial regions as shelter belt plantings around residential
areas and in urban parks, gardens, and street plantings merits consideration.
The same is true (a) in reference to rural areas subject to air pollution
where shelter belt plantings are used as a means of protecting farm crops,
forests, and other vegetation, and (b) in reference to certain cultural
practices that appear to be conducive to a lessening of plant sensitivity
to phytotoxic air pollutants.
It is hoped that the papers selected for presentation in this volume
will permit an assessment of some of the USSR studies dealing with the
manifold interrelationships of air pollution and vegetation. There is a
possibility that the usefulness of such studies would be greatly increased
if supplementary detailed information could be assembled and analyzed with
regard to the specific environmental conditions under which each set of
the data reported were developed. Such information, properly organized
and analyzed, would permit a more precise identification of specific envi-
ronmental responses of a given species or ecotype or variety grown in a
given area under specific pollutant and time-concentration relations. It
would also permit a clearer visualization of comparable climate and soil
conditions in North America. This may facilitate the verification of
plant responses and plant adaptability and may also make possible the util-
ization of some of the USSR plant material under similar ecological condi-
tions in North America.
As the editor of this volume I wish to thank my co-workers in the
Air Pollution Section of the Institute for their valuable assistance.
Special thanks are due to Dr. M. Hoseh, who as the principal translator
rendered valuable service in connection with many phases of this survey.
M. Y. Nuttonson
Silver Spring, Maryland
May 1970
vii
-------
U.S.S.R.
ORIENTATION
-------
CLIMATIC ZONES AND REGIONS* OF THE USSR
OKHOTSK
Zones: I-arctic, II-subarctlc, Ill-temperate, IV-subtropical
Regions: 1-polar, 2-Atlantlc, 3-East Siberian, 4-Pacific, 5-Atlantic,
6-Siberian, 7-Pacific, 8-Atlantic-arctic, 9-Atlantic-continental forests,
10-continental forests West Siberian, 11-continental forests East Siberian,
12-monsoon forests, 13-Pacific forests, U-Atlantic-continental steppe,
15-continental steppe West Siberian, 16-snountainous Altay and Sayan,
17-mountainous Northern Caucasus, 18-continental desert Central Asian,
19-mountainous Tyan-Shan, 20-western Transcaucasian, 21-eastern Transcau-
casian, 22-mountainous Transcaucasian highlands, 23-desert south-Turanian
24-mountainous Pamir-Alay '
* After B. P. Alisov
ix
-------
THE EFFECT OF INDUSTRIAL GASES AND INDUSTRIAL SMOKE
ON VARIOUS FOREST SPECIES
P. S. Pogrebnyak
From P. S. Pogrebnyak. Obshchee lesovodstvo. Vtoroe izdanie. Moscow,
Izdat. "Kolos", 1968, p. 215-17.
The air over large industrial centers contains gaseous and particu-
late matter vented by the heating units and smokestacks of industrial
plants. This matter which remains suspended and does not precipitate
over a long period of time, consists of sulfur oxides, nitrogen oxides,
carbon monoxide, tarry substances, soot, cement dust, and the like.
Industrial exhausts are injurious to vegetation, particularly to
evergreen trees and especially to conifers. Coal dust clogs the stomata
and deprives them of elasticity; cement dust alkalizes the cell content
of leaves and hydrolyses (eats away) their integumental tissue. Sulfur
dioxide is particularly injurious to leaves; at a content of 0.00001%
by volume in the air its effect is noticeable on plants. Penetrating
the cells through the stomata and by osmosis through the integumental
tissues of the leaves, sulfur dioxide forms strong acids, sulfurous and
sulfuric, which lower the pH of the cell content to 2.0-2.5 and depress
the cytoplasm*. The large scale desiccation of coniferous species in
forests near industrial plants is a widespread phenomenon in Western
Europe. In our country it has not attained such large proportions. How-
ever, foresters have to consider the problem of protection, prevention,
and treatment of forests damaged by gases vented into the air by chemical
and metallurgical plants.
Coniferous species are affected by poisonous gases more than other
species. The perennial leaves of the majority of the evergreen conifers
are poisoned continuously and are usually dead in the second year. The
species that normally shed their leaves annually and remain bare during
winter (larch, deciduous trees, and to a large extent arborvitae) get rid
of the greater part of absorbed poisons. Thus, these species, being bare
in winter, are capable of withstanding better the severe industrial gas
pollution of the air.
The poisoning by sulfur dioxide starts with the yellowing of the
leaf tips, and the appearance of reddish spots, and ends with the color
change of leaves or needles to brownish red, an indication of necrosis.
In the case of evergreen conifers exposed to moderate but constant poison-
ing, the current year needles appear normal, those of the previous year
sick, and older needles fall off. Damaged trees suffer partial loss of
their current year leaves along the edges of the crown, and this in turn
*[Translator's note: The Russian translates as "depress the plasm."]
- 1 -
-------
causes thin foliage. The greatest damage to evergreen conifers is caused
in summertime, during the period of vigorous development of the needles
and of intensive photosynthesis. During winter, even large concentrations
of injurious admixtures in the air do not cause noticeable damage.
Moss and lichen are among the most sensitive indicators of air pollu-
tion; they disappear even at small concentrations of sulfur dioxide in the
air. Moss on old straw-thatched roofs disappears even near such small
gas venters as garages and small coal-operated power plants. Recent investi-
gations showed that spraying pine boughs, slightly damaged by sulfur dioxide,
with a 2% solution of sodium carbonate containing a small quantity of
permanganate restores the normal reaction of cell sap.
There are various degrees of resistance of tree species to industrial
gases, as compiled by several authors (Schroder and Reuss, Stoklaza,
Wislicenus, Grohman, Gerlach, and others). These standards are frequently
contradictory because the observations on the basis of which they were com-
piled were made under differing conditions, and because in most tree species
the vulnerability differs with age. Thus, a young fir is insensitive to
industrial gases but after the age of AO it becomes most sensitive, along
with spruce, and Swiss stone pine. Larch and those evergreens having dove
gray needles, such as Colorado spruce, Engelmann's spruce, Serbian spruce,
as well as arborvitae, junipers, lodgepole pine, and false cypress, should
be considered resistant and are recommended for growing under conditions of
moderate air pollution. Eastern white pine is considered resistant to
smoke (Cermak, 1960). The.dove gray color is generally attributable to a
wax deposit on the surface of the needle, the thin layer of wax protecting
the leaf from the penetration of injurious gases and smoke. Very sensitive
and non-resistant are spruce, fir, and yew, particularly when they grow
old. Leafy species arranged according to their increasing degree of resist-
ance are: linden, ash, beach, maple, elms, alder, poplar, birch, and oak.
Of these, the last three are fully resistant and the first three are sensitive.
For conditions existing in Karaganda, the resistance arranged in decreas-
ing .order is: pinnate elm, Russian olive or oleaster, balsam poplar, box
elder, Tatartan or Tartar honeysuckle, lilac, Siberian pea shrub, or peat
tree, European white birch, and Scotch pine (A. S. Sitnikova, 1966). Under
conditions of Azerbaidzhan, to the above should be added: pistachio, almond,
willow-leaved pear, corkbark elm, and others. From this it is apparent that
gas resistance is to some degree correlated with drought resistance.
It is therefore obvious that the problem of protecting forests against
noxious gases primarily concerns conifers, exclusive of larch. One of the
most important ways of coping with this problem is to select gas- and smoke-
resistant varieties and types of woody species.
The dust concentrations of forest air is negligible. In suburban for-
ests it amounts to I/14th of the average dust content in the atmosphere
(Neuwirth, 1965). Forests are natural filters for dust and noise.
Literature
["ran^lator's note: Unfortunately the author did not provide any bibliography;
therefore, no sources of the references mentioned are available.]
- 2 -
-------
THE INFLUENCE OF SMOKE AND GAS ON THE
FLOWERING AND FRUITING OF SOME TREES AND SHRUBS
V. G. Antlpov
Central Botanical Garden of the Academy of Sciences of the Belorussian SSR
From Akad. Nauk BSSR. Belorusskoe Otdelenle Vsesoyuznogo Botanicheskogo
Obshchestva. Sbornik Botanlcheskikh Rabot, Vyp. II, (Minsk, 1960), p. 167-172.
The question of the influence of smoke and gas discharged from
industrial enterprises on the flowering and fruiting of plants has been
relatively little studied. E. Haselhoff and G. Lindau (1903), accord-
ing to the observations of Fr. Nobbe and A. Shtokhardt, describe the
influence of coal smoke on wheat. They wrote that if the action of the
smoke coincided with the time of the flowering of the wheat, then the
stamens would dry up, resulting in empty or poor embryos of very light
(weight) seed. The side of the spike facing the source of smoke was
particularly affected. According to the observations of A. Shtokhardt,
in a fruit orchard exposed to coal smoke, as early as the next day there
were evident traces of damage to the leaves, and the young green fruit,
especially plums, fell off.
Survey data concerning the effect of smoke on the fruiting of trees
under the conditions of the Donbass were assembled by Kh. M. Isachenko
(1938).
The results of a study of seeds (in accordance with G.O.S.T. 2937-47),
collected on an industrial enterprise, were published by us in 1957.
During the course of two growing periods we made additional observa-
tions of the flowering and fruiting of trees and shrubs, in three different
industrial chemical enterprises.
Industrial Enterprise A
From a mill, situated 50 to 75 meters from the plantings, gas was
given off in an unorganized and continuous manner. The concentrations of
the gases in the area of the plants were as follows (in g./m.3):
methane - 0.01084,
ethylene - 0.00262,
acetone - 0.00175,
acetic acid - 0.0011
Likewise, the plants were affected by flue smoke from a heat and electric
power plant (HEP) situated 200 meters from the plantings. The concentrations
- 3 -
-------
of these (gas) discharges were not established. The relief of the land
was uniform; the trees and shrubs were not shielded by buildings from the
effect of smoke and gas.
Industrial Enterprise B
The distance from the smoke flue of HEP to the plantings ranged from
10 to 100 meters, the main pollutants were large particles of coal, carbon
black, soot, and ashes. The relief of land was uniform; the plants were
not protected by buildings from the effect of pollutants.
Industrial Enterprise C
From a mill, situated 75 to 100 meters from the plantings, ethyl
chloride, ethylene, ethyl ether (diethyl ether), and benzene were discharged
in an unorganized and continuous manner. The concentrations of ethyl chloride
and ethylene in the area of the plantings were as follows (g./m?):
ethyl chloride - 0.0116,
ethylene - 0.0081;
the concentrations were not determined for the rest of the gases. The
relief of the land was uniform and the plants were not closed off by build-
ings from the effect of gas.
An assessment of the abundance of flowering and fruiting of trees and
shrubs, growing in the given industrial enterprises, was done visually
according to a six-degree scale:
0 - no flowers or fruit
1 - very weak flowering and fruiting
2 - weak flowering and fruiting
3 - satisfactory flowering and fruiting
It - good flowering and fruiting
5 - abundant flowering and fruiting
The results of the investigations are presented in Table 1.
As can be seen from Table 1, out of the 25 species of trees and shrubs
examined In the 3 industrial chemical enterprises, the majority was observed
to have quite satisfactory flowering.
Significant quantitative decreases were observed in the flowering of
common birch, penduculate oak, snowy mespilus, and cluster bird cherry.
-------
Table 1
Speoies
1
Siberian pea tree
Caoagana arborescens L.
Eurppeau barberry .
Berb&ns vulfjans Lao.
White birch
Betula £ubescens Ehrh.
European white birch
Betula verrucosa Ehrh.
Common English hawthorn
Crataegus oxyacantha L.
Red-berried elder
Sambucus raoemosa L.
Russian elm _ ,,
Ulmus laevis Pall.
Jartar dogwood
ornus aloa L.
Pedunculate oak
Quercus robur L.
!'artar honeysuckle
.lOnicera tatarica L.
Garden serviceberry
Analaiichier rotundifolia
(lam.) Dun.-Cours.
Hedge cotoneaster _ . , ..
Cotoneaster lucida Sohlecht.
Norray, maple. ^
Acer platanoides L.
Small-leaved linden
Tuia cordate fill.
Speckled alder .
Alnus incana ( LJ Hoench.
Japanese rose
Rosa rugose thunb.
Mountain ash .
Sorbus aucu^ria L.
Sorbaria sorbifolia (L.) A.
Br.
Black currant
Ribes nigrum L.
Balsam poplar . .
Populus balsamifera L.
Mongolian poplar
PopDlus sufivfeolens Fisoh.
Pojulus trenula L.
Cluster bird cherry x . .
Padus racemose CLefc.; Gilib.
SjeeK mock orange .
Phil.'idephus coronarius L.
European ash
Fraxinus excelsior L.
Industrial Enterprise
ABC
d
b.
2
c
0
c
0
It
0
~ 5
4
0
4
2
3
0
3
.•a
I
3
5
0
5
0
1
0
5
3
0
3
0
0
0
2
h
8
c
k
k
2
1
5
4
2
0
1»
3
0
3
4
2
•U
1
5
2
1
0
5
4
0
0
4
0
0
1
2
1
Plover
6
0
3
1
2
1
3
3
3
5
0
4
1
0
£
&
7
0
1
0
0
0
2
1
2
0
0
1
0
0
Notes
8
Catkin seriously diminished
Majority of the fruit fell off prematurely
fruit clung for some time, but fell
prematurely
Cones of small dimensions
Fruit clung for sons time, but fell
prematurely *
Note: Dashes indicate absence of the given species on the grounds of the enterprise.
- 5 -
-------
In the Siberian pea shrub and common ash, the usual flowering was
absent in Enterprise C only, while in the Russian elm and small-leaved
lime, flowering was absent in Enterprise A only. Among all the examples
complete absence of flowering in all enterprises occurred only with the
balsam poplar.
Gas and smoke exhibited a stronger influence on the fruiting of
trees and shrubs than on flowering. Under the given conditions, for a
number of species, flowering might be observed but not setting of the
fruit as for example in the case of the Japanese rose and the trembling
poplar, in Enterprise A; as well as in the case of the Russian elm and
the black currant In Enterprise B; and In the case of the round-leaved
servlceberry, the Urals false spirea, and the cluster bird cherry In
Enterprise C. In the other species fruit setting took place but the
fruit fell unripened (white birch, common birch, pedunculate oak and
Mongolian poplar).
The following species showed satisfactory fruiting under the given
conditions: European barberry, common hawthorne, red berry elder, ledge
cotcneaster, Norway maple, speckled alder, mountain ash, and sweet mock
orange.
There was a significant decrease of fruit in the following species:
Tartar dogwood, small-leaved lime, Mongolian poplar and the common ash.
In the white birch, this diminishing was expressed In the number of
female flower clusters, as well as In their dimensions while in the
speckled alder, the decrease was basically in the dimensions of the female
flower cluster (Figure 1). Its male catkins had normal dimensions.
"-" •. Fig- 1: Speckled alder (Alnus
Incana (L.) Moench.)
a - Ripe cones with fruit (In
Enterprises A and C)
which developed under the
influence of smoke and gas.
b - Control
Fruiting was completely absent on all enterprises in the case of the
birch, Russian elm, pedunculate oak, round-leaved servlceberry, Japanese rose,
Urals false spirea, black currant, balsam poplar, and the cluster bird
cherry.
- 6 -
-------
Better flowering and fruiting of trees and shrubs was observed in the
enterprises where the basic air pollutants were not gases but carbon black,
ashes, and soot. In these enterprises, supplementary observations on the
flowering and fruiting of shrubs were made. For this survey 50 flowers
(more or less, depending on availability) were recorded from various parts
of the crown of each shrub. The resulting observations are given in
Table 2.
Table 2
Lower layer of crown
Upper layer of crown
Variety
V
CO
Jj
p
l-i
0)
g
W-l
*w
o
J^
4-i
*^
4J
g
3
o-
-
50
50
50
12
-
-
-
_
*
4J
•H
3
4-4
C
01
Ol
O
-
17
19
6
2
-
-
-
_
4-1
•H
3
U
n i
01
a
*
-
13
17
5
-
-
-
-
_
in
^4
01
(^
14-1
o
s^
4J
*p4
4-1
5
3
c-
12
50
50
50
50
-
-
-
_
4-1
T-l
3
^4
<4-(
C
01
(_l
0
5
15
12
4
17
—
-
-
_
4-1
>r4
3
p
>4-l
01
a.
2
2
1:
11
4
12
-
-
-
Side
Southern Northern
in
u
01
§
g-i
14_|
o
^
4J
•^
w
3
O*
15
50
50
50
-
15
50
-
50
4J
*H
3
^4
IU
C
01
01
M
0
-
26
22
18
_
_
16
—
9
4-1
•i-l
3
ii i
01
a
S
-
18
18
-
-
_
14
—
9
en
M
0)
|
y-i
IM
o
^
4-1
, j
4J
cd
3
o-
50
50
50
50
50
12
50
25
50
40
•H
•4-1
C
01
01
CJ
21
23
23
7
19
_
17
2
12
4-1
•H
3
*•"
0)
a
*
16
19
21
2
14
_
12
_
12
Note: Gas and smoke acted from the southern side of the crown.
- 7 -
-------
The observations indicated that the smoke and gas discharged from
industrial concerns have different effects upon the flowering and
fruiting of different species of shrubs. The common hawthorne and the
red berry elder did not react to the gas and smoke. The Tarter honey-
suckle, the European barberry, and the (game) round-leaved serviceberry
showed that on the sides of the crowns that were facing the sources of
the smoke and gas there were fewer flowers, less fruit setting, and a
greater drop in the immature fruit set than was the case on the opposite
side, in spite of the fact that this was the southern, more lighted part
of the crown. This was especially apparent In the Tartar dogwood where,
out of SO observed flowers (in the upper part of the crown) 18 formed
fruit sets and all of these dropped before ripening.
Thus, gas and smoke, discharged by industrial concerns, had little
noticeable effect on some individual species of trees and shrubs, while
With other species the effect was shown in the decrease of the flowering
and in the number of fruit set, as well as in the premature dropping of
the young fruit.
Literature
Antipov, V. G. Deystvie gazov, vybrasyvaemykh promyshlennyini
predpriyatiyami, na semen a derev'ev i kustarnikov. Bot. zhurn.,
t. 42, No. 8, 1957.
Isachenko, Kh. M. Vliyanie zadymlennosti na rost i sostoyanie
drevesnoy rastitel'nosti. "Sovetskaya Botanika" No. 1, Izd.
AN SSSR, 1938.
Haselhoff, E., Lindau, G. Die Beschad!gung der Vegetation durch Rauch,
Berlin, 1903.
- 8 -
-------
THE EFFECT OF SPECIFIC INDUSTRIAL GASES
ON THE GROWTH OF SOME TREE SPECIES
V. G. Antlpov
Central Botanical Garden of the Academy of Sciences of the Belorussian SSSR.
From Belorusskoe otdelenie Vsesoyuznogo botanicheskogo obshchestva.
Botanika. Issledovaniya. Vyp. V. (Minsk, 1963), p. 150-154.
The injurious effect of some chemical substances on plant growth is
known in the literature. Salts of heavy metals (copper, lead, silver, etc.)
and such organic substances as ether, chloroform, toluene, ethylene, etc.,
arrest plant growth. The concentration of injurious gases exerts a decisive
influence on the growth increment of plants. Thus, the majority of even
the most poisonous substances taken in small concentrations may exert a
stimulating rather than a depressing effect on growth (N. A. Maksimov, 1948).
We have made a study of the effect of gases from a chemical plant on
the annual increment of shoots. The concentration of the exhaust gases was
constant and known. In the area where the trees grew, the concentrations
were: methane, 0.0184; ethylene, 0.00262; acetone, 0.00175; and acetic acid,
0.0011 g./ra.3.
In selecting experimental plants their origin, age, and location were
taken into consideration. The plants selected were the healthiest ones,
10-20 years old, except the elm, which was older. In measuring the growth
Increment, the close and regular interdependence of the biological character-
istics of the various orders of branching were taken into account (E. I. Gus-
eva, 1951). The average total .was taken from 10 measurements made on like
stories and branching order.
Measurements made on control plants growing in the absence of gases
showed that in all studied species the annual growth increment decreased
as the branching order increased. This consistent pattern of behavior did
not apply to the elm, which can be attributed to it's greater age and to
the fact that in old trees the regularlity of growth increment changes
appreciably.
The regularity of the annual growth increment was disturbed also in
trees and shrubs growing on the site of the industrial enterprise. Particu-
larly great deviations occurred in the southern parts of the crowns, facing
the source of exhaust gases (Fig. 1). Shoots of the branching order I lost
more of the annual increment on that side of the crown than the others. In
many cases, the increments in the orders II and III were higher than in
order I. No such cases were observed in the control plants. This indicates
that industrial exhausts in the indicated concentrations hampered the
annual growth of those shoots in which the growth processes are most intense.
- 9 -
-------
N
N
N
Fig. 1 Growth increment of shoots
A - Crataegus oxyacantha L. (English Hawthorn); B - Cornus
alba L. (Tartarian dogwood); C - Quercua robur L. (English
oak); S - Southern side of tree crown; N - Northern side;
a - Growth increment on the industrial site; b - Control.
Roman numerals indicate branching order.
As early as 1911 V. V. Sabashnikov wrote about the effect of industrial
gases on the annual growth rings: "it was noticed that the annual rings of
trees exposed to a constant action of factory exhausts become narrow and
irregular to such a degree that a cross section through the tree makes possi-
ble the determination of the approximate year when the adverse effect of the
factory started*"
The effect of industrial gases on the thickness of annual growth of
trees were studied at a chemical plant which produced nitrates. A comparison
of cross sections from the trunks of poplars and birches grown on the site of
this enterprise and in a location not exposed to gases shows that the annual
rings of trees grown on the site of the enterprise produced periodic formations
of concentric thickening and thinning zones, which were unrelated to climatic
changes and were the result of the variable composition and quantity of the
deleterious exhaust gases.
On the industrial site 10 dead poplar trees (Populus balsamifera L.)t
20 birches (Betula oubescens Ehrh.). and a willow (Salix fragilis L.) were
studied. A comparison graph is drawn on the basis of the study (a) of two
birches that died on the industrial site and (b) of one control sample.
Except for the gases, the growth conditions and the state of the trees were
relatively uniform. Also the climatic and soil conditions could not exert a
telling influence. Therefore, a sharp decrease in the growth of certain
years can be attributed only to the injurious effect of industrial gases.
As
can be seen from the graph (Fig. 2), in the control specimens (C)
- 10 -
-------
the annual growth, on the north side of the tree, was, during all the years,
to some degree not unlike that of the growth on the south side, though some-
what smaller. The growth curve of trees grown on the industrial site have
shown a similar relationship. As the production of nitrates increased, the
random venting of nitrogen oxides increased, thereby causing wide-spread
destruction of vegetation. An analysis of a cross section through the
trunk shows that an old tree (A) was more resistant in the first few years.
15*0
1950
B fS
(950
1910
1350
Fig. 2 Annual increment of trunk diameter
of European white birch
A and B - On industrial site; C - Control;
N - Northern side of tree trunk;
S - Southern side
Only in the third year, after the volume of injurious exhaust gases
increased, did the increment in the thickness of the trunk decline appreci-
ably; but a young tree (B) reacted faster and in the following year its
growth declined greatly. In the old tree, in the early years of the gas
exposure, the increase in the trunk's diameter on the northern and that on
the southern side were equalized (the gases acted on the southern side).
Later, however, the growth increase on the northern side exceeded that on
the southern side, which is not characteristic of trees growing under normal
conditions. In the young tree the growth increment on the northern side of
the trunk exceeded that on the southern side in the second year. In the
next two years the growth of this young tree increased somewhat. This was
apparently because the dying-off of the tree crown on the southern side left
the entire undamaged root system to contribute to the growth on the northern
- 11 -
-------
side of the trunk. The decline of growth began in the fourth year. The
absence of a similar phenomenon in an old tree may be explained by its
greater resistance to change in growth pattern.
When the production of nitrates stopped and the venting of nitrogen
oxide ceased, the old tree could not recover its previous state and died
after two years, whereas the young tree could not recover for two years
but later showed a considerable growth. The curves giving the diameter
increment of the tree were equalized. In the last few years, in connection
with a technological change of the manufacturing process, the composition
of the gases changed and their concentration increased. The trees perished
en masse. In more resistant species the growth increment declined. Thus,
for example, the balsam poplar (Populus balsamifera L.) at the age of 20
had a height of 9 m. and for 16 years the average annual growth increment in
its diameter was 8 mm. , whereas in the last 4 years it was only 0.6 mm.
At present, the tree is completely dead.
Variations in the growth increment of annual rings can serve as index
of the injurious effect of various industrial gases on plants. For such
an objective, young trees are preferable because they are more responsive
to changes of surrounding conditions and recover more rapidly after being
affected by gas. The older trees react more slowly, take longer to recover,
and as a rule eventually dry up. These differences may be related to the
prevalence of different kinds of gas resistance (N. P. Krasinskiy, 1950)
at definite ages; in the case of old trees the nature of resistance being
anatomical, morphological, and physiological (less oxidation of the cell
content), whereas in young trees the biological resistance to gases is
greater.
Literature
Guseva» E- !• Biologicheskiy metod izucheniya zakonomernostey rosta i
plodonosheniya tsitrusovykh I drugikh plodovykh rasteniy. Tr. In-ta
fiziologii rasteniy im. K. A. Timiryazeva. 1951, t.VII, vyp. II.
Krasinskiy, N. P. Dymoustoychivost1 rasteniy i dymoustoychivye assortimenty.
Sb. rabot, Gorkiy-Moskva, 1950.
Maksimov, N. A. Kratkiy kurs fiziologii rasteniy. Ogiz, sel'khozgiz, 1948.
Sabshnikov, V. V. Vliyanie Kamennougol'nogo dyma na okruzhayushchuyu
rastitel'nost'. Boleznl rasteniy, 1911. No. 3-4.
Rusnow, P. Uber die Vorstellung von Rauchschaden im Nadelwald. Wien, 1910.
- 12 -
-------
REACTION OF TREES AND SHRUBBERY TO AIR POLLUTION
IN THE FOREST-PARK BELT OF MOSCOW
AND MEASURES FOR EXTENDING THE LIFESPAN OF PLANTS
A. P. Shcherbakov and I. F. Cherednlchenko
Forestry Laboratory of the Cosplan of the U.S.S.R.
From Akad. Nauk SSSR Ural. Filial. Ural. Cos. Univ. im. A. M. Gor'kogo.
Okhrana prirody na Urale. (Sverdlovsk, 1964) 4: 151-162.
The present work was undertaken at the suggestion of the Moscow Soviet
and at the behest of the Presidium of the U.S.S.R. Academy of Sciences,
with the cooperation of the Sanitary and Epidemiological Station of Moscow
and the Park Service of the Moscow Soviet. The investigation was carried
out in a number of Moscow suburban parks and forests. The main difficulty
was to find in areas free of air pollution plantings that would be similar
in all their characteristics (i.e., their species composition, age of
plants, and soil characteristics, as well as in their hydro-geological con-
ditions and grass stand, etc.) to those of the plots selected for observa-
tion in the air-polluted eastern areas of the park-forest belt of Moscow.
These plantings were to be used as control or check plots in the study.
Only a carefully worked out method of approach to the study of this
biologically complicated problem will enable us to identify the cause of
forest destruction. The latter is the result of many factors, such as in-
dustrial discharges, changes in the composition or make-up of standing
timber caused by untimely and incorrect felling, as well as by compaction of
the soil or by changes in the hydrological conditions, etc.
Several areas were chosen for investigation. In the smoke-polluted
eastern, northeastern and southeastern areas the sections and trees chosen
were as follow: (1) in the suburban Moscow district of the forest-park
administration, compartment no. 50 - a 70 to 80-year-old and a 16-year-old
pine, compartment no. 53 - a 60 to 70-year-old and a 12 to 14-year-old
spruce; (2) in the Mytishchinskiy forest-park administration, compartment no.
19 - a 70 to 80-year-old spruce; (3) in the Kuchinskiy forest-park administra-
tion, compartment no. 59 - a 60 to 70-year-old spruce; (4) in the Balashikhin-
skiy forest-park administration, compartment no. 101 - a 70 to 80-year-old
pine. In the non-smoke-polluted western and northwestern areas, the following
trees were chosen in the various forest preserves: (1) in the Khlyupinskoe
forest preserve - a 70 to 80-year-old pine and a 16-year-old pine; (2) in the
Sharapovskoye forest preserve - a 60 to 70-year-old spruce and a 12 to 14-year-
old spruce; (3) in the Stepanovskoye forest preserve - a 70 to 80-year-old
spruce; (4) in the Nikolina Gora - an 80-year-old pine. The prevailing wind
direction in Moscow is from west to east.
The growth of industrial establishments and large enterprises of factories
and mills, as well as the development in the last few decades of automotive
transport in Moscow, as in many other large cities in the U.S.S.R., caused the
- 13 -
-------
discharge into the air of millions of tons of gas, smoke, dust, and other
pollutants. Accumulated smoke forming a dense, dirty smog can often be
seen over large industrial cities. Quite frequently this dense dirty fog
hangs over the city at a height of 500 meters, and the ultraviolet rays
have difficulty in penetrating it even in the heat of summer. In most
cases, these rays do not reach the earth's surface. In all large industrial
cities of the world, because of the man-made smog, insolation decreased
during the last 50-60 years by 10-30% as compared with the early part of
the century. In this connection it is important to remember that full
natural sunshine is the best enemy of bacteria.
The chemical admixtures vented into the air by industrial enterprises
and automotive transport also affect negatively the vegetation. A single
thermal power station discharges into the air an average of up to 30 tons
of noxious gases per day, which is equivalent to 50 tons of sulfuric acid.
Oxidation of motor fuel containing tetraethyl lead results in a discharge
into the air of world capitals of up to 15 tons of lead daily in the form
of minute particles. Each automobile exhausts into the air an average of
6-10 cubic meters of gas. In large cities, millions of cubic meters of
noxious substances are discharged into the air (Table 1).
According to the data of many years of records of the Moscow Sanitary
and Epidemiological Station (Gabinova, Vasil'eva, Popov, 1961), the maxi-
mum concentration of sulfur dioxide in one of the easterly suburbs
(Izmaylovskly Park) occurs In November-February, and the lowest In May-July.
In this connection should be noted that the annual average concentration of
S02 for the period 1955-1960 dropped sharply in this area, apparently
because of a general lowering of the S02 content in the air of Moscow dur-
ing this period and of a general improvement in the city air conditions.
Similar conditions were found during our investigations of the Moscow
forest-park belt (Mytishchlnskiy, Podmoskovnyy, Kuchinskiy, and Balashi-
khinskly forest-parks). Thus, during May and June neither sulfur dioxide
nor nitrogen oxides were found, whereas in October and November they
reached a high level at all points, and at some points their concentrations
were near lethal (Table 2).
These data Justify the conclusion that the most damaging concentrations
of gases affect the trees during the autumn-winter season. However, the
injury to vegetation by gases is less in winter than in summer, particularly
to broad-leaf species because they drop the foliage. It is equally clear
that greater gas damage is to be expected to conifers than to broad-leaf
species. Meteorological conditions can augment the damaging action of in-
dustrial smoke and gases. Fog, dew, precipitation, and a high relative
humidity of the air enhance the action of gaseous air admixtures on plants.
During autumn and winter air layers at low heights from the land surface
contain products of incomplete combustion, which combine with droplets of
fog or dew to form condensation nuclei. Separately, the air pollutants
(carbon monoxide, carbon dioxide, sulfur dioxide, tars, etc.) are less
dangerous and less aggressive than in combination. In a fog, all these
substances concentrate and form additively an injurious mixture, more
dangerous to animals and plants. Wind direction and intensity can also
increase their adverse effect on vegetation.
- 14 -
-------
Table 1
Content of some chemical admixtures in the air of smoke polluted (A) and
smoke-free (B) areas of the Moscow forest-park belt.
Place of sampling
A. Podmoskovnyy
forest-park, com-
partment 50,
80-year-old pine
B. Khlyupinskiy for-
est preserve, near
Zvenigorod, 70 to
80-year-old pine
A. Podmoskovnyy
forest-park, com-
partment 53,
spruce
A. Balachikhinskiy
forest-park, com-
partment 101, 80
to 90-year-old
pine
B. Nikolina Gora,
sanitarium, thin
stand of pines ,
75 to 80 years old
Date
26/V
14/VIII
2/VI
26/V
14/VTI
30/V
29/VIII
7/IX
Time
of day
10-12
-
11-13
13-15
13-15
14-15
16-18
11-12
11-12
i
9/VI
1/IX
15-17
12-14
so2
0.15-0.36
0.3-0.42
Not found
H ii
0.42
Not found
0.48-0.72
0.15-0.48
Not found
0.42-0.72
Nitrogen
oxides
Not found
0.11-0.23
Not deter-
mined
ii ii
0.11-0.23
Not found
0.117-
-0.117
Not found
0.12-0.12
0.117-
Active
chlorine
0.037-
-0.058
Not found
ii it
0.019-
-0.037
Not found
ii ii
it ii
ii it
ii it
0.01-0.03
-0.936
Chlorides
Cl ion
Not deter-
mined
Not found
0.13-0.20
Not deter-
mined
0.12-0.12
0.03-0.05
0.12-0.12
Not found
0.13-0.23
Not deter-
mined
Remarks (where air
samples were taken)
Glade near edge of
thinned 80-year-old
pine stand
Same place
Glade behind forest
Clearing in forest
Near edge in open spacu
In forest, 20-30 m.
from forest edge, near
a railroad track in an
open space
Open space near forest
Open space between sin-
gle trees at height of
15 m.
Open space on shore of
r. Moskva, near bridge
Same place
-------
1 (Continued)
Place of Sampling
park, compartment
59-67, 80-yea
spruce, compa
85--70, 80-yea
spruce
Sharapovskoye
preserve, com
80-85, spruce
Time Nitrogen
Date of day SC^ oxides
Active Chlorides
chlorine Cl Ion
Remarks (where air
samnlos wore taken)
spruce
Same pi
12, adj
spruce
10, spruce
rest-
ment
_ _i j
r— old
rtment
r-old
forest
ipartment
y forest-
ment 19,
-old spruce
omp a rtment
year-old
ompartment
to 19,
ompartment
16/ VI
17/VI
2/VI
9/VT
30/V
17/VIII
5/IX
24/X
12-14
15-16:3
15-17
12-14
11-13
11-12
11-13
Not found
5 0.3-0.42
Not found
n n
it n
0.3-0.42
Not found
0.45-0.81
No
0.
No
••
II
0.
No
0.
-0.234
it n
-0.234
0.31-0.62
Not found 0.17-0.12
0.1-0.1
Not found
0.01-0.01
Not found
0.0-0.12
0.13-0.20
0.13-0.37
0.13-0.27
0.03
0.12-0.12
0.13-0.13
Small glade Ln forest
near "Korovly prud"
cow pond
Open space near forest
edge
Inside the forest on a
small glade near the
forest edge
Felling area
Near forest edge, at an
Intersectional clearing,
Inside forest
Open space near dense
forest
Intersectional clearing. In
the crown of trees, at a
height of 15 m. Edge of
compartment
Open space near dense for-
est stand
Table 2
Content of some pollutants in the air of the Moscow forest-park belt (mg./m.3)
Place of sampling
Podmoskovnyy forest-park
section 50
section 53
Kuchlnskly forest-park
Balachikhinskiy forest-park
Mytishchinkskiy forest-park
so2
May- June
0.15-0.30
Not found
n ti
it it
Aug. -Oct.
0.38-0.42
0.42
0.37-0.42
0.48-0.72
0.48-0.81
Nitrogen oxides
May-June
Not found
ii n
n n
n n
it ti
Aug . -Oc t .
0.170-0.23
0.150-0.23
0.205
0.117
0.31 -0.62
-------
In viewing the effect of air pollutants on vegetation, two aspects
can be brought out:
1. The protective role of green plantings as a shield blocking the
penetration of noxious air downwind into the depth of the stand. Dealing
with this problem are the interesting works of V. F. Dokuchaeva (1959)
and others, which recommend the planting of shelter belts for the protec-
tion of more valuable species against smoke and noxious gases.
2. The negative physiological effect of mechanical and chemical air
pollutants on vegetation. This effect manifests itself in two ways.
a. The direct effect of gases on the assimilation apparatus of the
leaves: as a result of absorption of 862 through the stomata assimilation
is disturbed, photosynthesis lowered, chloroplast iron inactivated, respira-
tion frequently increased (the respiration coefficient exceeds 1), vitamin B
destroyed, an unfavorable protein and carbohydrate balance sets in, and the
accumulation of silicic acid and strontium in the leaves increases.
This is followed by the appearance of external symptoms of disturbed
metabolism, injury to leaf tissue, leaf necrosis, all of which affect the
growth of the leaf and the increment of the tree's height and diameter.
Our observations in this respect coincide with those of foreign authors.
We have shown that under the influence of smoke and gases the conifer
needles grow more intensively in length than in thickness , whereas the re-
verse is observed in the leaves of broad-leaf species. In the initial
growth stage of a leaf, gases stimulate the growth In width while in subse-
quent stages they arrest growth, thus preventing the leaves from attaining
normal shape. This leads to an increase in the xeromorphy of the leaf's
structure. In addition, we observed in a young, 16-year-old pine, in the
Podmovokovnyy forest-park (compartment 50) that the tips of the needles
turned as a rule bright yellow, below which the needles were of a faded,
drab dark green color and foreshortened. At the same time, in a similar
stand of pines (Khlyupinskoe preserve) where the air is free of pollutants,
the needles showed no sign of disease and the annual increment in height
and diameter was appreciably greater than of the pines in the Podmoskovnoe
forest-park.
Data in Table 3 lead to the conclusion that the needles and the growth
buds of spruce and pines growing in the smoke-polluted areas of the Moscow
forest-park belt are greatly affected by air pollution. In all cases, includ-
ing young and old trees, the absolute green and dry weight and length of the
needles drop, the average linear length increment of apex shoots declines,
the average weight of apex and lateral buds diminishes, all of which is con-
nected with a weakening of the synthesizing activity of the leaves and of
the growth processes. An inverse relationship was observed in 1961 in the
number of needles per unit length of shoot, that is, the number of needles
per cm. on trees growing in smoke-polluted areas was appreciably greater
than on trees growing in pure air. The needles on the apex and lateral
shoots were broomlike, tightly bunched and appreciably shorter.
More convincing proof was obtained in spruce stands (80 to 85-year-old)
growing in smoke-polluted and unpolluted areas of the Moscow forest-park
belt. One example is quoted in Table 4.
- 17 -
-------
Table 3
Blometric data of pine and spruce needles of 1961 growth in smoke polluted1(A) and smoke-free (B)
areas of the Moscow forest-park belt.
Place of sampling
(sample trees)
I
A. Mytischinskiy for-
est-park, 70 to 75-
year-. ;ld aprme,
compartment 19
B. Stepanovskiy for-
est preserve, 70-
year-old spruce
30
1 II
A. Kuchinskiy forest-
park, 80-year-old
spruce, compart-
ment 59-57
B. Sharapovskoe for-
est preserve, 80
to 85-year-old
III
A. Podmoskovskiy for-
est-park , 80 to 90-
year-old pine, com-
partment 50
B. Khlyupinskoye forr-
est preserve, 80-
vear-old nine
Raw weight
of 1000
needles
g-
5.35
10.94
8.83
10.18
15.94
32.75
Z of
contr.
48.9
100
86.7
100
48.7
100
Air-dried
wt. of 1000
needles
g-
3.84
5.39
4.41
4.70
7.88
16.90
Z of
contr.
71.3
100
93.8
100
46.7
100
Moisture
in needles
Z
28.2(+?)
52.0
50.0
•
55.0
50.3
47.0(?)
Average
length of
needle
cm.
1.1
1.4
1.15
1.32
3.42
5.4!
Z of
:ontr.
78.6
100
87.1
100
62.8
100
Number of
needles
per 1 cm.
28.8
24.2
29.3
25.8
35.0
25.5
Z of
contr.
119.0
100
113.5
100
137.2
100
Average in-j
crement of
apex shoot
for 1961
cm.
5.8
6.5
4.0
9.5
1.8
2.7
Z of
contr.
89.2
100
42.1
100
66.7
100
Average wt.
of apex bud
mg.
99.4
114.4
80.0
135.4
10.5
65.0
Z of
contr.
87.0
100
59.1
100
19.1
100
Average wt.
of bud of a
lateral 1961
shoot
mg.
28.7
32.0
27.5
31.7
_
_
Z of
contr.
90.0
100
86.8
100
_
_
-------
Table 3 (Continued)
Place of sampling
(sample trees)
IV
A. Balashikhinskiy for-
est-park, 80 to 90-
year-old pine
B. Nikolina Gora, 75
to 80-year-old pine
V
A. Podmoskovskiy for-
est-park, 12-year-old
pine, compartment 50
B. Khlyupinskiy forest
i preserve, 12 to 14-
,_ year-old pine
vo
Raw weight
of 1000
needles
g-
19.35
32.39
57.55
60.28
% of
contr.
66.0
100
95.5
100
Air-dried
wt. of 1000
needles
g-
9.58
14.68
26.52
28.13
% of
contr.
65.2
100
94.2
100
Moisture
in needle;
%
53.2
55.0
53.0
54.6
Average
length of
needle
cm.
3.8
6.15
6.6
6.5
% of
contr.
61.8
100
100
100
Number of
needles
per 1 cm.
28.2
31.0
12.8
9.2
% of
contr.
91.0
100
139.0
100
Average in
crement of
apex shoot
for 1961
cm.
3.2
6.0
34.6
43.4
% of
contr
53.3
100
79.7
100
Average wt.
of apex bud
mg.
28.0
72.0
237.2
38.58
% of
ontr.
38.9
100
61.5
100
Average wt.
of bud of a
lateral 196i
shoot
mg.
3.0
24.5
86.6
140.2
% of
contr.
12.3
100
61.8
100
Table 4
Comparative growth increments of spruce plantings in smoke-polluted and smoke-free
areas of the Moscow region
Smoke-polluted plot
Spruce stand, 80 years old,
Kuchinskiy forest-park,
194 trees
Smoke- free plot
Spruce stand, 85 years old,
Shorokhovo, Zvenigorod
district, 210 trees (control
of Kuchin plantings)
Number of trees, %
withered
8.1
0.54
half
withered
76.5
1.0
healthy
15.5
98.5
Annual increment in height , cm.
1961
12.0-43%
21.0
1960
7.0-72%
25.0
1959
8.5-73%
31.0
1958
8.0-56.7
18.5
1957
i n 9 z.i L
21.0
-------
It follows from the quoted data that noxious gas pollutants reduce
the number of healthy trees 6-fold and the number of withered and dried-
up trees rose more than 85-times as compared with stands in pure air.
The annual height increment for the last 5 years declined 43-73%. Accord-
ing to data of senior forest pathologist in the Moscow forest-park adminis-
tration, Yu. A. Troyanovskaya, the large number of weakened and drying-up
trees in Kuchino greatly contributed to the development of numerous and
powerful sources of severe infestation of insect pests. These include
forest pests which attack the trunk of the tree (such as various bark
beetles and destructive woodborers of the Scolytidae [Ipidael spp. , as well
as the Norway spruce weevil, Norway spruce black long-hornet beetles, etc.)
At the same time, in a control or check area of 0.5 hectare of a spruce
stand, these pests were encountered only on 1-2 trees.
b. The indirect action of noxious gases in the air appears also
through the soil. Although the soil exerts to some extent a buffering
action, systematic and prolonged acidification changes the course of chemical
processes. The soil grows poorer in bases, its gas equilibrium is upset,
its microbiological activity is disturbed, and it accumulates soluble salts
of aluminum and strontium. This upsets the normal plant nutritive regime
(or status) of the soil. Ultimately, all the particles of smoke, soot,
dust, and gases polluting the air end up in the soil. Some of these, such
as silicon, fluorine, lead, copper, zinc, chromium, strontium, and others,
in small doses, are necessary microelements but in large amounts they are
toxic. Heavy metals penetrate the soil to a depth of 25 centimeters. The
soil absorbs a large quantity of these substances, as well as smoke, soot,
and dust. A good example can be seen near Moscow, in the Podolssk region
at the edge of the town of Pavshino, where cement mills are located; the
soil and the vegetation around them are covered with a thick layer of gray
dust. True, the floating dust is not as detrimental to soil and plants as
are the smoke and gases but in combination with them its action is very
aggressive.
It should be noted that the deleterious effect of chemical and mechanical
air pollutants is more pronounced on forest than on tilled soils, regardless
of the higher buffer capacity of the former and their higher humus content.
The tilled soils are plowed and fertilized every year. It is known that
the addition of mineral fertilizer (calcium and magnesium) and the tilling
are factors in overcoming the ill effects of smoke and gas. Smoke pollution
causes continuous acidification of forest soils. The acid components of
smoke and gases are not buffered by forest soils and cause more structure
degradation and lower fertility than in tilled soils.
Various kinds of trees and shrubs exhibit different degrees of biological
resistance to noxious chemical and mechanical impurities that pollute the air.
The most resistant of the conifers is larch because it changes its needles
every year. Next is pine, which changes its needles every 2 or 3 years;
next to the pine Is fir, which changes its needles every 3-5 years; and
finally comes spruce, which changes its needles once in 7 years. Some author-
ities consider the fir as the most susceptible to noxious air pollutants.
- 2C -
-------
Some investigators consider four groups of trees in relation to their
gas resistance: highly resistant, slightly susceptible, susceptible, and
very susceptible. These investigators have used the following criteria:
premature fall of leaves or of needles, appearance of spots on leaves,
bareness of tree tops, lowered resistance to pests, and arrested increment
of height and of diameter (Paprzycki, 1960). We have used the same cri-
teria in estimating the degree of injury caused by smoke and gas to the
growth of spruce and pine stands in the eastern sections of the Moscow
forest-park belt.
Many investigators contend that the resistance of trees to smoke gases
changes with the local conditions and with the age of trees. Young spruce
appear to be more resistant to gases. Clear signs of gas damage were found
only in the barling stage. Low quality, thin stands were usually found on
poor soils but this differed with the age of trees. In a 90-year-old spruce
and pine stand, IV and II bonitats, the spruce died out completely, while
the pine was well preserved. In another place, in a 60-year-old pine stand,
the Scotch pine was less resistant to SC>2, HF, and arsenical compounds
than the Weymouth pine. Under the influence of sulfur oxides, the pine pro-
duces bunched shoots on its top and forms a multi-apex crown. These signs
appear to a lesser extent in the Weymouth pine. The gas-resistant pine
varieties have a darker needle color and a larger angle at which the needles
are fastened to the shoots, as compared with the less resistant varieties
having a paler needle color, short needles, and a small angle of needle
attachment (Pelz, 1958). It can be considered as proven that a high resist-
ance to S02 and SO^ have been found in the following species: Persian wal-
nut, spruce, pine, ash, beech, woadwaxen, oats, alfalfa, lentil, heliotrope,
primrose, sweetpea, and nasturtium; high resistance to Cl and HC1 is charac-
teristic of Persian walnut, cherry, and grapevine; and high resistance to
F and HF has been found in the grapevine, apricot, and gladiolus. Particu-
larly sensitive to smoke gases are some fungi (Oidium and Microsphaer guezina).
Least sensitive to smoke gases are: oak, hornbeam, linden, maple, elm, birch,
alder, willow, poplar, mulberry, hawthorn, and elder (Antipov, 1957).
It was shown that when the leaves of apricots contain 3-10 mg. of fluorine
per 100 g. of their dry weight, the tree shows definite signs of injury,
whereas when they contain chlorine 200-300 g. dry leaves no injury was
noticeable (Translator's note: There is an obvious misprint in the preceding
phrase which renders the text unclear). Fluorine not exceeding 0.1 mg. and
chlorine not over 5-10 mg. per 100 g. of dry weight of leaves are considered
tolerable for the normal growth of trees.
Lately, considerable work has been done in the technique of estimating
noxious admixtures in the air. It can he based either on the external
symptoms of plants or on analysis of leaves (high contents of S02 and wax).
In other cases bio-indicators have been used with considerable success,
namely: gladiolus for the detection of fluorine; pine (Pinus taeda) capable
of accumulating sulfates, without showing signs of damage; bluegrass —
sensitive to general air pollution; and sunflower as well as mazzard cherry —
sensitive to sulfur oxides.
- 21 -
-------
In conclusion, It Is necessary to point out possible means for increas-
ing the resistance of tree stands under conditions of smoke gas pollution
and particularly, in the eastern part of the Moscow forest-park belt. The
advisability of using chemical fertilizers for this purpose is emphasized
in literature. Our own experience of many years in studying the physiology
of soil plant nutrition confirms this view. Systematic application of
mineral fertilizer to stands subject to pollution by acid gases and smoke
can, without doubt, greatly improve the chemical and physical conditions of
the forest soils, as well as their raicroflora and the nutrition of grasses
and woody plants. Here, of utmost importance are lime and lime-magnesia
(limestone, dolomite, marl, etc.) augmented by nitrates and phosphates. It
is advisable to apply in autumn, every 3-5 years, 2-3 tn./ha. of lime (CaO),
dolomite or marl; this is a suitable, effective, and economical procedure
in the agro-technology of forest-park management. In the following spring
should be added 100 kg./ha. of ammonium nitrate; this is best broadcast and
then worked in with rakes, hoes, or mattocks. Young trees can also be sprayed
with a 0.5% urea solution, which will greatly improve their growth.
At any rate, adopting a system of fertilizing the forest-park belts of
the cities is a powerful means for improving the structure and fertility of
the soil as well as for improving the life activity and productivity of tree
stands and their Increasing resistance to pests and to the ravages of city
smoke gases.
Literature
Antipov, V. G. Vliyanie dyma 1 gazov vybrasyvaemykh promyshlennymi
predpriyatiyami na sezonnoe razvitie derev'ev i kustarnikov.
"Bot.zh.", vyp. 42, no. 1, 1957.
Dokuchaeva, V. F. Rol1 drevesnykh nasazhdeniy v obezpylivanii atmosfernogo
vozdukha. "Zhurnal gigieny, epidemiologii, mikrobiologii i immuniteta"
(Chekhoslovakiya), 1959, vyp. 3, no. 2, str. 238-246.
Ershov, M. F. Fotosintez chistykh i zapylennykh list'ev vyaza melkollstnogo
i lipy melkolistnoy. "Dokl. AN SSSR" t.112, vyp. 6, 1957.
Zhokhov, L. I., Pery, G. V., Davidovich, E. M., Gabinova, Zh. L., Vasil'eva,
A. A. , Popov, B. V. Vliyanie zadymleniya atmosfernogo vozdukha na
zelenye nasazhdeniya. "Gorodskoe khozyaystvo Moskvy", no. 5, 1961.
- 22 -
-------
CURRENT CONDITIONS AND SCIENTIFIC PROBLEMS IN STUDYING
THE INJURIOUS EFFECTS OF INDUSTRIAL POLLUTANTS ON PLANTS
AND IN DEVELOPING METHODS FOR CONTROLLING THEM IN THE URALS
S. A. Mamaev
Institute of Biology, Academy of Sciences U.S.S.R., Ural Branch
From Akad. Nauk SSSR Ural. Filial. Ural. Cos. Univ. im. A. M. Gor'kogo.
Okhrana prirody na Urale. (Sverdlovsk, 1964) 4:7-18.
As the tempo of industrialization in the Urals accelerates, industry
exerts a constantly increasing diverse and profound effect on the natural
resources of the region, including that of vegetation. This occurs through
various factors, in various ways, of which the most significant is that of
industrial pollution. Numerous ash piles from power stations, plants and
factories, and large tracts covered by wastes from mining, occupy consider-
able areas in the Urals. On them and near them there is usually no vegeta-
tive cover, or else the vegetation found there comprises useless grasses,
trees, and shrubs. Many industrial enterprises dump their waste water into
rivers and lakes, thereby destroying aquatic and littoral vegetation.
Finally, the smokestacks of an enormous number of industrial developments
belch ash, soot, sulfur dioxide, and other gases which strongly pollute the
air and soil around them and create unfavorable conditions for plant growth
on large areas extending over hundreds of square kilometers. The magnitude
of this process is exemplified by the city of Sverdlovsk where the boilers
(of power stations) alone discharge into the air 400 tons of ash and over
100 tons of sulfur dioxide per day. In addition, various enterprises, auto-
motive traffic, and railroads contribute their share to air pollution.
At the present, there are no accurate figures concerning the negative
effect of industrial pollutants on vegetation. However, for the larger
Ural Region including Perm, Sverdlovsk, Chelyabinsk, and Orenburg districts,
Bashkir and Udmurt A.S.S.R. our preliminary estimates of damaged areas are
that these extend over 500,000 hectares.
It should be kept in mind that similar conditions obtain primarily and
in a particularly acute form in population centers, i.e. . in areas where
vegetation is especially valuable to man.
All this means that the time is ripe to address ourselves seriously to
the task of eliminating the deleterious consequences of air and soil pollu-
tion destructive of vegetation. However, in the Ural Region as well as in
many other areas of the country, the problem was not investigated scientific-
ally until recently, and if some attempts in this direction were made, they
were one-sided and dealt with specific issues rather than with the overall
problem. It can be assumed that the scientific and theoretical bases of
the problem were not worked out, its main points are not clearly different-
iated, and guidelines for such investigation were not formulated.
- 23 -
-------
In defining the basic tasks of the problem of the Interrelation of
vegetation and industrial pollution it is advisable to start with the
source of pollution affecting the environment and life of the plant.
Under the influence of industrial pollution, a whole series of factors
affecting plant life change simultaneously, even though the changes may be
of different magnitude. In examining the most Important of these, we
shall attempt to describe briefly the extent to which these problems were
investigated in the Ural Region and to suggest means for combatting their.
adverse effect on plant life.
It should be pointed out at the very beginning that in so short a
communication it is not possible to touch upon the effect of radioactive
pollution on plant life. This is a rather special problem at the present
and requires a separate treatment.
Changes in the Gaseous Composition of Atmospheric Air
The air of industrial areas has an Increased concentration of extraneous
gases. The composition of these gases varies and depends on the type and
nature of local industry. In almost all cases, the content of sulfur dioxide
increases to 0.5-1.5 mg./m.3, or more, within a radius of 1-2 km. from the
source (according to V. At Ryazanov the permissible concentration is
0.37 mg./m.3) and the carbon monoxide content increases to 20-35 mg./m.3.
(near metallurgical plants). Near aluminum and, frequently, near copper
smelters, the fluorine content of the air increases to 0.4 mg./m.3 in a
radius of 10 km.
The concentration of carbon dioxide (C02) also Increases. In the air
appear vapors of sulfuric acid, hydrogen chloride, ammonia, nitrogen oxides,
and the like. Most of these substances penetrate through the stomata into
the intercellular spaces, and precipitate as liquid acids on the surfaces
of leaves and stems, causing burns of various degrees of severity. In the
former case, the essence of the injurious effect of acid gases, according
to N. P. Krasinskiy, consists of photodynamic chlorophyll action. But this
does not exhaust the damage, as was pointed out by Krasinskiy himself (1950).
In the latter case, the burns are the result of a raechanical digestion of
leaf tissue by the strong acids. The different types of injuries must be
considered separately, regrettably, however, this is not always done by
those who Investigate gas resistance of plants.
Plant physiologists are faced by a serious task, namely, to study the
causes of gas resistance of plant organisms without restricting themselves
to the oxidation theory of the cell content. In the last few years, the
botanical gardens of the Institute of Biology of the Ural Branch of the
U.S.S.R. Academy of Sciences began some work in this field. V. S. Nikolaev-
skiy confirmed the assumption of N. P. Krasinskiy that the quantity of
oxidizable substances in the leaves of a number of woody plants may be
accepted as an index of resistance to acid gases. Resistant species, e.g. .
box elder, contain less oxidizable matter than species severely injured,
such as dwarf apple and European white birch. Also, changes are noticeable
- 24 -
-------
In the course of the growing period. The quantity of oxldizable substances
increases 1.5-2 times toward autumn; this is in agreement with an increased
leaf vulnerability to gases toward the end of summer. The established cor-
relation of gas resistance with the intensity of life processes is also sig-
nificant: the more resistant plants usually show a lower intensity of photo-
synthesis and respiration. There are also some preliminary data relative
to the great importance of the anatomy and morphology of the leaves to the
gas resistance of woody plants.
Other investigations on the gas resistance of plants (predominantly
woody) were also carried out in the Ural Region. Among these should be
mentioned the work of the Ural Institute of the U.S.S.R. Academy of Munici-
pal Economy, the work of some verdant-plant specialists in such cities as
Krasnoural'sk (M. V. Bulgakov), Berezniki, as well as in the Chelyabinsk
district. However, the purpose of these investigations was not the physio-
logical or anatomical causes of gas resistance. Generally, they gave an
overall evaluation of the resistance of plants to air and soil pollution.
Recently, Yu. Z. Kulagin studied the physiology of plants growing under
conditions of increased gas pollution.
High Dust Content of the Air
In the southern reaches of the Urals, naturally occurring wind erosion
brings about a high dust content of the air.
In the central Urals and in the forest belt of the southern regions
dust pollution of the air is caused essentially by power stations and coal-
fired central generating plants. Usually, the dust carries small particles
of coal, ash, and frequently, depending on the nature of the industry,
particles of silica, aluminum oxides, caustic magnesite, and compounds of
sulfur, lead, fluorine, copper, arsenic, and zinc. Dust pollution in some
areas of the Urals was studied in detail by the Institute of Industrial
Hygiene and Occupational Diseases (Sverdlovsk).
Dust of a particle size 0.1-100 microns settles at a distance of
several scores of meters to several kilometers from its source. The quanti-
ty of settled dust is quite considerable and may reach several hundred grams
per square meter per year. The suspended dust consists of particles 0.001-10
microns and its content in the air is 0.4-0.6 mg./m.3 (Thomson, 1959).
Dust settled on leaves interferes with the normal life processes by
sealing the stomata, causing the leaves to overheat, and shading off the
light. This, in turn, decreases the intensity of photosynthesis and causes
a 16-27% decline in the accrual of vegetative mass (Ershov, 1959). Poisonous
oxides of lead, fluorine, copper, and the like in the dust causes poisoning
and death of leaves.
In the Ural region valuable studies on the effect of dust pollution on
woody plants have been carried out for a number of years by the Institute of
Biology of the Ural Branch of the U.S.S.R. Academy of Sciences and the Bashkir
State University.
- 25 -
-------
Most of the Investigators concentrate on the Interaction of the leaf
and the dust particles, skirting the phenomena of stimulation and of depres-
sion of shoot growth. The same is true of the study of the effect of gases
on growth. Yet, a growing young shoot interacts actively with various
chemical agents. This is attested in the numerous works dealing with the
action of gtbberellin. Thus, our experiments with woody plants showed that
a bud of a leafy species is more sensitive to a gibberellin solution
placed on the bud's integument than are needle plants.
It should be taken into account that among the dust particles and
gases in the air surrounding cities and towns are many physiologically active
substances and irritants.
It should also be noted that city air contains more carbon dioxide, which
:is beneficial for the photosynthesis of plants.
The problems of combatting dust and gas pollution of air remain for
the present unsolved, not only in the Ural Region but in our country as a
whole as well as in most foreign lands. Sparging of tree tops and of lawns
and flowers is practiced in many cities. This is quite effective when
dealing with a non-cementing dust and in small areas. However, when the
dust forms a dense film on the leaves and shoots it becomes necessary to use
various solvents, the composition of which depends on the substances in the
dust.
The improvement of the soil, which in turn improves the vitality of the
plant, is also of great importance. Plant selection is another means of pro-
tection against the noxious effect of gases and dust. Forms and varieties
should be selected which, because of anatomical structure of the leaf and
shoot integument, retard the penetration of noxious substances into the
parenchyma. Also, plants capable of recovery after injury should be chosen.
ChanRe in the Physical and Chemical Properties of Soil
Here seven basic trends are to be distinguished: a) the formation of
dumps of various kinds; b) the addition to the soil of various particles,
settling out from smoke and as result of wind-blown dust from the dumps;
c) the effect of industrial waste water; d) the soil pollution in the park-
ing areas of auto-transport; e) the fills and dug ground resulting from con-
struction excavations; f) the pollution by garbage in settled areas; g) the
compaction of. soil in areas frequented by people (parks and suburban forests).
The last two trends are not directly related to industrial pollution. How-
ever, it is hard to separate results caused by industrial pollution from
those caused by household waste or by compaction by people because all
these negative phenomena appear simultaneously. On some of these problems
there exists a voluminous literature. For the Ural Region there exist the
works of the Ural State University (V. V. Tarchevskiy) on dumps and those
of the Institute of Municipal Economy (E. T. Mamaeva) on excavated soil, as
well as other works.
- 26 -
-------
Of the most urgent problems in making the cities verdant the most
vital is that of finding the means to reclaim ash piles and dumps of mine
overburden. These wastes affect adversely not only vegetation but, primarily,
the inhabited areas in many cities and towns in the Urals. Therefore, the
studies concerning classification of waste piles and dumps, soil formations
under these conditions, and the biology of plant groups growing under con-
ditions of an excess of some minerals in the soil and of insufficient moisture
and nitrogenous nutrients are of special significance.
The changes occurring in soils under the influence of excessive dust
precipitated from smoke is not studied sufficiently. Although reliable data
are scarce, it can be assumed that the upper soil horizons are enriched with
microelements and that soil acidity and microflora have changed.
The questions of soil formation in cities in conditions of extensive
pollution and soil compaction are of special significance. Here frequently
come into play rules entirely different from those governing natural soils.
It must be noted that, for the present, soil scientists are as yet not too
much concerned with the problems of soil formation in areas of intensive
human activity, although this problem is becoming increasingly urgent and
decisive in the process of soil condition changes.
There is very little material dealing with the suitability of different
soil groups of various degrees of contamination: a) for growing lawns and
ornamental plant species utilized in landscaping and in making cities ver-
dant; and b) for various trees and shrubs grown in forested areas. Informa-
tion concerning the effect of salt accumulation in soils or the effect of
various quantities of slag, rubbish, or wastes on plants is either totally
lacking or quite contradictory.
The effect of industrial pollution on the vegetation of suburban parks,
gardens, and greenswards is combined with the action of profound physical
changes resulting from compaction and from disturbance of natural runoff.
It is frequently overlooked that injury to the soil by noxious gases and
dust from various industries occurs simultaneously with impeded aeration of
the soil and with disturbance of its structure. It is our opinion that
studies should stress the need to clarify the complete role of the various
factors conducive to the change of plant environment.
The effect of industrial waste water was sufficiently well studied in
relation to its action on fish life of a body of water. As is known, the
fish assets of the rivers and lakes of the Urals are greatly diminished,
particularly in the last decades and even during the last few years, as
results of dumping industrial waste waters. However, little work has been
done in ascertaining the effect of industrial waste in so far as it brings
about changes in plant life of the watersheds and shore areas, and the
resistance of various plant species under the new environmental conditions.
Changes in the Biological Environment of the Plant Habitat
The increased air and soil pollution by industrial wastes greatly affects
the living organisms interacting with plants (particularly affected are the
- 27 -
-------
symbiotic organisms living in areas of maximum pollution). This, in turn,
disturbs the natural biological links and disrupts biocoenosis. Depending
on the group of bio-organisms, the basic changes occurring in areas of
Industrial pollution can be thus schematized as taking place: a) in the
species composition of soil flora and fauna; b) in the species composition
of pathogenic microorganisms; c) in the species composition of insects,
whether injurious to plants, carrying pollen, or carrying disease, also
d) in the composition of birds and mammals feeding on plants, on insects
and other arthropods, or on worms coming in contact with plants.
The importance of these organisms in plant life is well known, although
many facets of their interrelationship are as yet obscure. This emphasizes
the significance of the dislocations in the biocoenosis caused by the
extinction or suppression of some microorganisms and other life forms. There
are at the present a number of investigations that enable one to draw con-
clusions as to some details relative to changes in the composition of soil
bacteria and fungi as well as of lichens as a result of gas and dust pollu-
tion of the air. A great amount of work has been done by the medical and
sanitary service in studying water and air pollution in cities. However,
these investigations did not deal with problems related to the changes in
the biotic environment of plants.
Thus, this important field of research is wide-open. Of paramount
importance is the study of the effect of industrial pollutants on the compo-
sition, ecology, and mode of life of numerous pests and parasites among
the microorganisms and insects.
There are some data for example, which suggest that seed of many species
of trees and shrubs grown in intensely air-polluted surroundings are less
subject to diseases. It should be pointed out that their germination and
growth vigor were higher (investigations of V. G. Antipov, Leningrad, 1957).
Preliminary studies also show that some pest insects are less often encoun-
tered in cities.
At the same time it should be remembered that plants weakened by
adverse city conditions may be more subject to various diseases and pest
attacks. Thus, in the verdant plantings of Sverdlovsk as well as of other
cities there is a widespread occurrence of the following: bacterial canker
of balsam poplar; powdery mildew of gooseberries; fungal and bacterial dis-
eases of gladioli, dahlias, and phlox. Such pests as aphids, leaf moths,
sawflles, fruit moths, and the like are also of wide occurrence there. When
studying the ecology of living organisms in polluted areas, the enormous
effect of various chemicals used In forestry, landscaping and fruit growing
for combatting pests and weeds should not be overlooked.
Many of our higher plants and, above all, the widely distributed
conifers, such as pine, fir, and larch, have on their roots mycorrhiza.
Numerous studies have shown that destruction of mycorrhiza fungus weakens
the host plant and may cause its withering. The mycorrhiza fungus is very
sensitive to soil conditions and it is affected by soil pollution. We are
thus faced with the necessity of investigating the complex effects of un-
favorable plant growth conditions encountered near cities, towns, and
- 28 -
-------
Industrial enterprises. It is quite possible that many cases of Injury to
plantings, particularly, to conifers, are directly attributable to the
destruction of mycorrhiza fungus. This Is pointed out In great detail by
Yu. Z. Kulagln (1961) in his work on the greenbelt of the city of Satki.
In working out measures to prevent the withering of conifers, the proper
conclusions must be drawn from the above in order to ameliorate the soil
environment for the plants by means of the establishment of favorable
conditions for the survival of raycorrhiza.
Care of the soil is basic in combatting the ill effects of industrial
pollution. The water-air properties of the soil should be improved, its
fertility increased, favorable conditions for soil microorganisms cre-
ated, soil reaction adjusted as needed, and the harmful effect of pollutants
neutralized by addition of suitable chemicals and adsorbents.
Improving plant nutrition will lead to the general recovery of the
plants and will increase the resistance of their above-ground parts to
gas and dust.
Regrettably, these methods for preventing or neutralizing the adverse
effects of pollution have not been studied sufficiently as yet in the Urals.
The green plantings in the cities are fertilized but not much else is
practiced as yet in the maintainence and care of their soil environment.
Lately, the Institute of Municipal Economy and the Botanical Garden
of the city of Sverdlovsk started a study of the effect of fertilizer on
plant habitats in areas grossly polluted by industrial waste. Preliminary
data obtained from the plots established by the Botanical Garden in the
city of Revda indicate that fertilization improves greatly soil conditions
and enables plant life in areas where otherwise none could be sustained.
It is sometimes said that combatting pollution is a temporary task
inasmuch as its severity declines. Indeed, as technical means of control
of noxious gases and dust at their source improve, as the very technological
processes of industries improve, and as transport as well as industry are
to change to other fuels, the air and soil pollution will diminish.
However, it will never cease entirely because part of the waste will always
be carried out beyond the confines of industrial units. Also, new industries
will arise for which effective waste control will not yet be available. The
increase in population also contributes to various forms of pollution.
Undoubtedly, there must be a constant increase of the areas taken up by mine
wastes, excavations, fills, and disturbed soil resulting from the accelerated
tempo of building activity, as well as from increased volume and scale of
mining operations and of factory building.
Consequently, the importance of counteracting pollution will not diminish.
It is, therefore, imperative to augment the scientific research work in this
area in order to discover and to ascertain the regularity and natural devel-
opment of plant resistance on the basis of which there could be developed
methods of combatting the injurious effects of noxious gases, dust, and soil
pollution.
-------
In our opinion, these then are the concrete problems facing the
scientific Institutions:
1. Formulation of a general theory of gas and dust resistance of plants.
To this end it is necessary to carry out systematic studies of the physiology,
anatomy, and morphology of at least three groups of plants: anglospermous
herbaceous; angiospermous deciduous; and gymnospermous evergreen woody plants.
The latter must be paid special attention. On the basis of these studies,
methods should be worked out for combatting the ill effects of air pollution
and for protecting plants from gas Injury.
2. Investigation of soil formation and plant cover of dumps, ash piles.
excavation waste, disturbed ground, and fills. This should comprise soil,
geobotanical, and microbiological studies, combined with testing of various
species, varieties, and forms of plants under these conditions.
3. Investigation of plant nutrition under conditions of increased salt
content, and of changed water and air conditions, and in the presence of
extraneous and frequently poisonous admixtures. The results of these investi-
gations combined with the ones mentioned in the preceding Item 2 will make
it possible to formulate effective recommendations for reclamation of dis-
turbed territories, by biological means.
4. Elaboration of plant and plant group classification under conditions
of heavy air and soil pollution^ The first preliminary stage of such classi-
fication is available, now in the data of numerous studies conducted by various
investigators.. A more exact, and scientifically well-grounded, classification
will be developed upon completion of more thorough and detailed studies.
5. Methods of evaluating air and soil pollution in relation to the
suitability of the polluted environment for growing various species of plants.
For the present, in most cases there is a lack of reliable indicators and
data concerning the gas composition in the air or of the degree of salinity
and pollution of soil, on the basis of which recommendations could be made
relative to the selection and production of suitable plants. It is unknown
at which concentrations of the various gases, salts, and the like, begin
the unfavorable effects on plant growth in relation to the environmental con-
ditions and age of plants. The development of leaf diagnosis, widely used in
agriculture for determining plant needs (Magnitskiy, 1958; Childers , 1960,
and others), may be useful in this connection.
6. Investigation of the effect of industrial pollution on microorganisms
and fauna in verdant plantings of suburban belts as related to possible
changes in the biocoenosis.
The foregoing enumerated problems constitute some of the major basic
tasks facing investigators. Their solution requires much effort. Of great
significance is the problem of coordinating the scientific work dealing with
the investigations of the effect of industrial pollution on plants. Such
coordination should be applied within the framework of an entire geographical
and economical region, such as the one constituted by the Ural Region. This
- 30 -
-------
coordinating task should be vested, in our opinion, in the commissions for
the preservation or conservation of nature at the various branches of the
U.S.S.R. Academy of Sciences. In this particular case, we have in mind the
Ural Branch, which has already started such studies. The individual
commissions for nature preservation of the various branches of the Academy
should be united into a suitable body embracing the entire country. Such
a complex scientific problem cannot be undertaken without properly organ-
ized coordination. In tackling the problem of pollution effects a special
role, in our opinion, should be undertaken by the botanical gardens.
Botanical gardens are singularly equipped to undertake such investi-
gations by virtue of a number of reasons. In the first place our botanical
gardens are located in large industrial centers where generally one finds that
the ecological environment is severely disturbed. There is, therefore, the
opportunity to study the types of pollution directly on the territory of
the botanical gardens, and also extensive experimentation can be carried out
there. Secondly, only the botanical gardens provide conditions conducive to
long-term observations of plants, soils, and fauna, while excluding extran-
eous factors and their effects which frequently disturb and indeed impede
such work in other places. And, thirdly, botanical gardens usually have a
staff of scientifically trained personnel capable of experimental work, and
have the necessary equipment and apparatus. In addition, scientists from
other institutions lacking experimental facilities can join in the work of
the botanical gardens. The ties between the botanical gardens, institutions
of higher learning, and laboratories of scientific institutions should be
strengthened.
In the cities of the Ural Region there are four botanical gardens - two
in Sverdlovsk, and one each in Perm and Ufa. Only one of these, in Sverdlovsk,
carries on work on the effect of industrial pollutants on vegetation. At the
present time, the Botanical Garden of the Institute of Biology of the Ural
Branch of the U.S.S.R. Academy of Sciences carries on research along the
following lines: a) studies of the physiological and anatomical resistance of
woody plants to acid gases; b) selection of woody and flowering ornamental
plants endowed with greater viability under conditions of industrial pollution;
and c) means for improving soil conditions for plants growing on polluted
areas (in cooperation with the Ural Institute of the Pamfilov Academy of
Municipal Economy). These studies, which began 2-3 years ago (1961-1962),
are of a rather limited scope. Moreover, a sizeable part of this work is
carried on outside the basic science program of the Botanical Garden. The
extent of this work must be enlarged.
In conclusion let us note:
1. Within the last few years, work has been undertaken in the Ural Region
on the deleterious effect of industrial pollution on vegetation and on devis-
ing methods to overcome its results. Some of the institutions engaged in
this work are: the Institute of Biology of the Ural Branch of the USSR Academy
of Sciences and its Botanical Garden; the Ural Institute of the RSFSR Academy
of Municipal Economy; the A. M. Gorki Ural State University: the Bashkir State
University; the Sverdlovsk Institute of Labor Hygiene and Occupational Dis-
eases; the forestry and landscaping organizations of Sverdlovsk and Perm
- 31 -
-------
districts, and other institutions. However, these studies are largely
fragmentary, are mostly concerned only with certain plant groups, and,
what is most important, are conducted on none too high a scientific level.
2. The problem of plant protection against pollution is complex, and
its solution is dependent on the analysis of the effect of various aspects
of industrial pollution on plants, and on their subsequent synthesis in
order to evolve a general theory of plant resistance.
To work out the theoretical bases it is necessary to attract a number
of research institutions and to establish in the Ural Region a science cen-
ter dealing with this urgent problem.
Literature
Antlpov, V. G. Deystvle gazov, vybrasyvaemykh promyshelennymi predprlyatiyami,
na semena derev'ev i kustarnikov. "Bot.zh.", No.8, 1957.
Ershov, M. F. Vliyanie pyli na rost rasteniy. "Bot.zh." No.6, 1959.
Krasinskiy, N. P. Teoreticheskie osnovy postroenlya assortimentov gazoustoy-
chivykh rasteniy. Sb. Dymoustoychivost' rasteniy 1 dymoustoychlvye
assortlmenty. M., 1950.
Kulagln, Yu. Z. Ob ustoychivostl drevesno-kustarnikovykh porod k deystviyu
magnezitovoy pyli v rayone g. Satki. Sb. "Voprosy razvitlya lesnogo khozy-
aystva na Urale", vyp. 2, trudy In-ta biologil, vyp.25, Sverdlovsk, 1961.
Magnitskiy, K. P. Polevoy kontrol1 pitanlya rasteniy. M., Izd-vo "Znanie",
1958.
Mineral'noe pitanie plodovykh 1 yagodnykh kul'tur. M., Sel'khozgiz, 1960.
Ryazanov, V. A. Sanltarnaya okhrana atmosfernogo vozdukha. M., Medglz, 1954.
Tomson, N. M. Sanltarnaya okhrana atmosfernogo vozdukha ot zagryazneniya.
M., Medglz, 1959.
- 32 -
-------
THE DRAINAGE OF TEMPORARILY SWAMPED SOILS IN RELATION TO
PINE PLANTINGS GROWING UNDER CONDITIONS OF INDUSTRIAL AIR POLLUTION
N. V. Podzorov
Director, Okhtinsk Experimental Forest
From Izvestlya Vysshykh Uchebnykh Zavedeniy.
p. 27-31.
Lesnoy Zhurnal. 1967, No. 2,
During the last few years many pine stands have died in the Okhtinsk
forest resort, bordering on the city of Leningrad. Extensive damage is done
there to the conifers (a) by the smoke gases of the city's industrial enter-
prises situated near the forest and also (b) by swamping of forested areas.
The damage was greatest among pine stands growing in areas of longleaf
sphagnum moss* near the city limits. In similar pine stands at a distance
from the city the amount of dead wood was smaller.
When areas of pine growing in longleaf sphagnum mosses within the zone
of intensive air pollution were drained, the number of dead pine trees
dropped sharply (Table 1).
Table 1
Distance from
pollution
source, m.
No. of
square,
sections
plots
Distance.
from dr
age dit
.
from drain-
itch
No. of
pines.on
ixperi-
mental
.plots
in I960
No. of.
Dead pines byvgrowth
category, %
II
in
IV
300-1900
(1st experi-
mental area;
(sejfion
With the drainage of pine stands in longleaf sphagnum areas distant from
city limits (from 1900m. to 2650m.) the drop-off in the number of desicated
pine trunks changes less radically.
To study the effect of drainage of soils having temporary excess moisture
on pine stands exposed to industrial air pollution, experimental plots were
laid out in the longleaf sphagnum area, three years after the drainage.
* [Translator's note: The Russian term used "dolgomoshnikov" is not too
clear. Literally it means "of long mosses" and probably refers to long-sphagnum
moss peats of the peat-bug soils.]
- 33 -
-------
These experimental areas were subdivided into 20-meter-wide plots. On each
such plot were no less than 250 recorded trees. During this study, the
drainage ditches on the experimental plots were properly maintained.
The experimental plots were located in sections IS and 18, and in squares
38, section 10.
Between 1960 and 1964 the dead trees on the experimental plots were
counted. The results are summarized in Table 1.
The results showed that the number of dead pines Increased with the dis-
tance from the drainage ditch. On experimental plot "a" the percentage of
dead pines was 17.A, whereas on plots "b" and "c" the percentage was 33.99 and'
39.76%, respectively.
The total number of dead pines in the experimental plots of the 2nd area,
which was further removed from the sources of pollution, was smaller than in
the 1st experimental area. Here, too, the number of dead trees increased
with the distance from the drainage ditch.
On plots "c" and "f" located at a distance of 60-100m. from the drainage
ditch, there is an appreciable increase of dead trees of the "growth category"
I and II.
A morphological and physiological study was made of the pine needles, as
well as an examination of the morphological symptoms of cones and seed and of
the principal phlsiologlcal indices of the seed. The quantity of needles
damaged by industrial smoke varied with the distance from the drainage ditch.
The healthier needles were in all cases somewhat longer.
In the plantings that we investigated, the needles of trees growing near
the drainage ditch, after being washed free of the dustlike industrial pollu-
tion, were dark green and, at times, gray-blue, while the needles of trees
growing at a distance of 80-100 m. from the drainage ditch were light green.
The same is true for pine regrowth. The needles on new branches on distant
trees appeared in irregular clusters, whereas on trees near the drainage
ditch the needles were evenly distributed.
The weight of the needles on experimental plots situated at a considerable
distance from city limits was in all cases greater but even in these plots it
increased with the proximity of the trees to the drainage ditch.
The llfespan of pine needles was also affected by the distance from the
drainage ditch. The needles on trees in plots located 20-AO m. from the drain-
age ditch lasted for 2 years, whereas needles on trees in plots at a distance
of 80-100 m. from the ditch stayed on the trees for 1 year only. Beyond the
city limits, the pine needles on trees along the drainage ditch stayed for
3 years, whereas on trees at a distance from the drainage ditch the pine needles
stayed for only 2 years. The decrease of the life span of needles, as well as
their lesser weight and lesser abundance, are caused not only by the adverse
effect of swamping of the plantings but also by the gradual accumulation of
- 34 -
-------
sulfur compounds in the pine needles until the cumulative effect of the
sulfur compounds becomes lethal causing weakening and drop of needles.
Needles taken from trees growing at various distances from drainage
ditches were analyzed in order to study the dynamics of sulfur accumulation
in the mesophyll. The analyses showed considerable accumulation of sulfur
in the needles (Table 2).
Table 2
Distance from
source of pollu-
tion
m.
300-1900
(1st test area)
(araF&M)
Experimental
plots
a
b
0
d
e
f
Distance from
drainage ditch,
1.
0-20
40-60
60-100
0-20
40-60
60-100
Percent of sulfur in tree. needles (numerator;
undamaged; denominator: with signs of damage)
1-year old
0.202
0757
0.233
0.244
0.396
0.186
O734l
0.207
0.234
0736T
2-year old
0.240
07351
no needles
ii M
0.207
0734T
0.277
07333
0.260
07577
3-year old
no needles
ii n
ii ii
0.303
0.393
no needles
ii ti
m&mam
It was found that the needles on trees growing near drainage ditches
contain less sulfur than those on trees further away. Needles on trees
growing on plots "d", "e", and "f" contained appreciably less sulfur than
on trees growing on "a", "b", and "c". In all cases, the sulfur content of
2- and 3-year-old needles was higher than in 1-year-old ones.
Fifteen trees with approximately equally developed crowns and growing
under similar conditions were selected on each experimental plot in order
to study the effect of drainage on the fruit-bearing of pines growing on
longleaf sphagnum-covered terrain. Cones were collected from the southern
side of the middle section of the crown in the first days of February 1962.
For each plot the collected cones (200) were graded according to color, size,
shape, structure of seed wing, and the ratio of length to diameter. The
distance from the drainage ditch did not greatly affect the color or the
structure of the seed coat, or shell. There were some differences in the
length, diameter, and weight of the cones. The results are presented in
Table 3.
Pine cones growing on trees near the drainage ditch (0-20 m.) were
generally larger and their average weight greater. No appreciable increase
- 35 -
-------
Table 3
Distance
from source
of pollu-
tion
ro.
300-1900
(1st test
field)
1900-2650
(2nd test
field)
Experi-
mental
plots
a
b
c
d
e
f
Distance from
drainage ditch
m.
0-20
40-60
60-100
0-20
40-60
60-100
Average
cone
length
cm.
3.1
2.1
2.0
3.6
2.2
2.2
Average
cone
dlam.
cm.
1.7
1.3
1.4
1.8
1.3
1.3
Average
cone
weight
g-
6.3
4.8
4.7
6.8
5.0
5.1
No. of
cones in
1 kg.
162
217
216
152
193
210
in size and weight was noticed in cones taken from trees at a distance of
40-100 m. from the drainage ditch. The effect of distance from the drain-
age ditch on the size and weight of cones taken from trees on the 2nd experi-
mental area, which is further removed from the source of air pollution, is
more pronounced. However, no significant increase in size and weight of these
cones in comparison with those from the 1st experimental area was noticed
at a distance of 60-100 m. from the drainage ditch.
The seeds in individual cones were counted by a modified G. N. Nezabudkin
method. A skillet was placed on a gas ring and heated, then a cone was
placed in it, and first turned on its side and then placed on its base.
After one or two minutes the cone started opening and the scales, softened
by heating, were readily picked with tweezers. Only fresh cones are suitable
for this treatment, as the scales of dried cones are brittle and hard to
pluck for seeds. Fifty cones from each plot were studied. The results are
summarized in Table 4.
The number of seeds in cones from trees growing near the drainage ditch
increased and the percentage of seedless cones decreased. The number of
seeds in cones did not increase noticeably in samples taken from plots "b"
and "c". However, the number of cones bearing no seeds rose. Also noticeable
was an increase in the number of normally developed wings having no seed
(up to 4%). The cones from pine trees on plot "d" in area 2 had a noticeable
increase of seed, the number of seedless cones decreased. On plots "d" and
the normally developed wings having no seed amounted only to 1.5-2X.
"e"
- 36 -
-------
Table 4
Distance
from source
of pollu-
tion
m.
300-1900
(1st test
field)
1900-2560
(2nd test
field)
Experimental
plots
a
b
c
d
e
f
Distance
from drain-
age ditch
m.
0-20
40-60
60-100
0-20
40-60
60-100
Percent of cones having
number of seeds
3
10
16
17
4
7
7
5
14
22
21
10
6
7
7
10
16
16
12
16
27
11
22
12
11
21
31
21
15
24
12
11
23
16
15
24
4
-
-
18
8
-
Percent
of cones
without
seeds
16
22
24
12
16
23
An Increase in the number of seeds and a decrease in the percentage of
seedless cones taken from trees on plots "d" and "e" in the 2nd area, which
is farther from city limits, is the result not only of the effect of
drainage but also a result of the decrease of the degree of the adverse
effects of the industrial discharge.
In the 1st area, 150 cones contained 945 ripened seed, or an average of
6.3 seeds per cone. Cones from the 2nd area yielded 8.3 seeds per cone.
The seed content of cones varied greatly - from 0 to 24. Cones from the
1st area contained up to 8.6% empty seeds, while cones from the 2nd area
contained up to 6.1%. As the distance from the drainage ditch decreased,
the number of empty seeds declined 1.5-2.4%.
In the 1st area, the number of cones from some trees growing near the
drainage ditch increased. In the 2nd area, no difference could be observed
in the number of cones on trees growing either adjacent to the drainage ditch
(0-20 m.) or at a distance from it. It is, therefore, permissible to con-
clude that draining excessively moist forest areas is most beneficial to
the fruit bearing of those pines which grow in areas polluted by industrial
exhausts.
Mo special regularity was noticed in the length and width of seeds
collected from various experimental plots. There was some difference in
the weight of seed and also in some of the most characteristic physiological
indices of the sowing quality of the seed, depending on the proximity of
pine trees to the drainage ditch. The results are shown in Table 5.
- 37 -
-------
Table 5
Experimental
plots
a
b
c
Distance from
drainage ditch
ra.
0-20
40-60
60-100
Weight of
1000 seeds
g.
6.3
5.6
5.1
Percent of
absolute
germination
9R.2
97.3
96.2
Sprouting vigor
in 7 days , %
65.2
63.8
63.1
Our data indicate that the weight of 1000 seeds from trees growing near
the drainage ditch increases. Also, the germination and the vigor of seed
sprouting increase somewhat. The average rest period of all the studied
seeds was 7.4 days.
In summarizing the results of our observations it can be concluded that
the effect of polluting gases of the same concentration and duration will
affect pine trees differently, depending on the changes in the soil moisture
conditions.
Three years after drainage, the adverse effect of smoke decreases.
This is expressed in the decrease of the intensity of sulfur accumulation
In the pine needles, in the increase in the weight of the needles, cones
and seeds, as well as in the increase in the growth of needles and in the
improvement of the most characteristic physiological indices of the seed.
- 38 -
-------
THE EFFECT OF INDUSTRIAL SMOKES AND GASES UPON CONIFEROUS FORESTS
GROWING UNDER CONDITIONS OF INCREASED HUMIDITY
IN THE MOSCOW REGION ("PODMOSKOV'E")
V. G. Antipov
Central Botanical Garden of the Academy of Sciences of the Belorussian SSR
From Akad. Nauk SSSR. Byulleten' Glavnogo Botanicheskogo Sada. Vypusk 46
(Moskva, 1962) p. 41-46.
In the Journal "Lesnoe Khozyaystvo" No. 7 for 1960, E. V. Lugovoy
published an article, "The Influence of Gases and Dust on Coniferous
Plantings in the Moscow Region". The author considered that the increase
of moisture in the air, as a result of the introduction and the utilization
of water reservoirs in Moscow, had a favorable effect on the growth of
forest species and neutralized the adverse effects of industrial smokes and
gases on forest vegetation, including the coniferous species. In particu-
lar, he indicates that the common pine trees do fairly well in Moscow in
the vicinity of the factory "Krasnyy Bogatyr1". However, the Commission
to Investigate Moscow City Parks of Culture and Recreation (Sokol'nicheskiy
and Izmailovskiy) by 1949 came to the conclusion that the basic reason for
the destruction of woody vegetation, mainly coniferous species, is the
adverse effect of industrial gases ("Work of the Scientific-Technological
Commission ...", 1949). Meanwhile, the Moscow lake and Moscow's main
reservoirs, created with watershed water (Khiraktnskoe, Kliaz'minskoe,
Pialovskoe, Uchinskoe, and Ikshinskoe), were already filled with water in
1937 and partially so in the subsequent three years. This, however, did
not have any noticeable effect on the condition of the tree plantings in
Sokol'nicheskiy and Ismailovskiy parks. In Sokol'niki an insignificant
number of old pine trees were left only in the northern part of the park.
According to data of the head forester L. A. Kashcheev, in the general area
of Izmailovskiy Park, which extends over 1180 hectares, 120-year-old common
pines occupied in 1931 65% of the forest-covered land area (34 sections out
of 39). Many of the forest sections there were entirely covered by the
common pine. In the park at that same time, an intensified replacement of
pine by linden was noted. Thus, from 1931 to 1954, in the central part of
the park, closer to the city and to industrial enterprises, the area of
pine repopulation decreased by 85.96 hectares, and, in the part of the
forest more removed from the city, by 30.39 hectares. At that time there
occurred also a massive desiccation of firs, and the area they covered was
reduced during that period by 63%. A definite correlation between the
destruction of pines and their distance from industrial enterprises was
noted. Thus, at a distance of 2km from factories the decrease in pines per
hectare consisted of 97 n>3 of timber, but at a distance of 5km it was 23.4 m3.
The Commission of the Central Administration of Forest Economy and the
Field-Protection Planting, having conducted in July 1955 an investigation of
the desiccated plantings in the Moscow region, found that in the industrial
regions of Balashikh, Sokol'niki, and in the Izmailovskiy Forest, a massive
- 39 -
-------
loss of trees took place (30-AO m^ per hectare), while in similar pine
stands of Malakhovskiy and Sosnovoborskiy Forests, further from the
factories and mills, the annual loss of trees was insignificant (1-2 m3 per
hectare [Timofeev, 1956]). The map of Izmailovskiy Forest, published from
material of the Fifth Moscow Aerial Photo, Forest Structure Expedition,
graphically shows the dynamics of change of the areas over 13 years
(c. 1946-1959). Pine plantings are, however, well preserved in the north-
eastern part of the park, a part most remote from the industrial enterprises
(see figure).
Change in areas under pine plantings
in Moscow's Izmaylov park:
a - 1946; b - 1959
The accumulation of sulfates, by the assimilating mechanism (Bredemann,
1933; Pelz, 1959), is one of the verified symptoms of damage to plants from
sulfur dioxide. Analysis of coniferous needles of trees that grew up in the
neighborhood of a large industrial complex showed that at a distance of 2 km
in the direction of prevailing winds they contain 0.63X sulfur; at a distance
of 3 km, 0.54%; at 4 km, 0.36Z; and at 9 km, 0.08% (Rlazanov, 1954).
Analogous data were obtained from the analysis of the needles of thorny firs
in various regions of Moscow and its environs (Abramashvlli, 1957), and also
in our study of the reasons for the massive destruction of pine trees in the
Cheliuskintsev cultural and recreational park in the city of Minsk, which
is situated in the direction of the prevailing winds, about 2-5 km from
heavy industrial enterprises emitting a significant quantity of smoke and
gases.
- 40 -
-------
In view of the report by M. S. Gol'dberg (1949) (Table 1), it is
impossible to consider the increase of humidity as a factor conducive
to air purification.
Table 1. The effect of humidity upon the concentration of
atmospheric pollutants (according to Gol'dberg, 1949)
Pollutant
Change in intensity factor
Effect upon pollutant
concentration
SO,
Fluctuations of air humidity
Cloudy weather
Fog
High humidity and fog
Rain
No effect upon
concentration.
Clear cut rise.
Sharp, large increase.
Concentration increases,
dispersion slowed down.
No noticeable effects.
CL2
Relative humidity raised
from 50 to 98%
Concentration decreases,
number of positive
tests drops.
Soot
Fluctuations of air humidity
Cloudy weather
Fog, calm
No connection observed.
Concentration increases.
The same.
Dust
Increased humidity
Fog
Rain (screened according
to Lifman-Lizechang)
Increased concentration
of suspended dust.
Concentration of suspended
dust increases 2-4 times
over normal.
Sharp increase In number of
dust particles on horizontal
plates during first day of
lain and a decrease in the
following days; increase
occurs due to an increase
in soot particles.
- 41 -
-------
The concentration of contaminants in the air (except C12) are most
sharply increased during fog. Observations carried out in the Urals showed
a direct dependence between the concentration of soot and sulfur dioxide,
on the one hand, and the humidity of the air, on the other (Riazanov, 1957).
Thus, many investigations have shown that with the increase in humidity
there is an increase in the concentration of damaging smoke and gases. These
concentrations can often reach toxic proportions, sufficient to cause not
only chronic damage invisible to the eye, but also a sharp toxic effect
leading directly to the destruction of the plants. For example, in lowlands,
where the fog hangs on, the sharp toxic effect on green foliage from the
anhydrides of sulfuric and sulfurous acids in sufficiently strong concen-
trations (Kuz'min, 1950) can be observed. With the rise in temperature as
well as with an increase of humidity in the air and an increase in the
amount of precipitation the process of forest desiccation is increased
(Il'lushin, 1953).
In drought years, sulfurous anhydride causes less damage to plants
than in wet years (Lent, 1959). The increase in humidity is not only con-
ducive to the formation of acidic fogs, but also prevents the closing of
the stoma. The latter creates conditions which are beneficial for the
penetration into leaf hollows of air saturated with acidic gases. Plants
that have wilted are more resistant to the effects of sulfur dioxide
(Kroker, 1950), while the limited consumption of water increases; for
example, the resistance of tomatoes to "smog" damage, i.e., to fog saturated
with hydrocarbons and oxidants, characteristic of Los Angeles (Koritz and
Went, 1953).
Earlier observations by foreign investigators had already established
that humid and foggy weather increases the damage to plants by the smoke
gases of metallurgical works. These observations were confirmed by experi-
mental methods (Holmes, Franclin, and Gould, 1915) and extended to other
acid smokes (Cristiani, Gautier, 1925). It was established that the main
damaging effect was caused by acid gases in dissolved form.
Experimental studies have shown that the moist parts of plants are
more quickly damaged than the dry parts, not only by more soluble acidic
gases such as hydrogen fluoride but also by sulfur dioxide.
However, Zimmerman and Hitchcock, 1956, having carried out experiments
on approximately 40 species of plants for their comparative sensitivity to
sulfur dioxide and hydrogen fluoride, found no visible correlation between
the quantity of stoma and sensitivity. A deposit of soot and smoke leads
to the loss of illumination, and this causes the paling of chloroplasts and
a decrease in the productivity of photosynthesis (Krasinskiy, 1950). Thus,
the negative effect of soot and smoke on plants is not subject to doubt.
It Is difficult to study the physiological effects upon plants of
fog-borne sulfuric acid and vapors of sulfuric acid and sulfur trloxide
(SOj) because these substances are ordinarily found in the air along with
sulfur dioxide, and their isolation is quite difficult. However, to consider
- 42 -
-------
the anhydride of sulfuric acid harmless, as does engineer E. V. Lugovoy
(1960), Is without Justification. More than that, the massive poisoning
of the population, which has been observed in a number of industrial
regions in Belgium (In the Maas valley - 1930), In the USA (the city of
Donor - 1948), and in England (London - 1952) in periods of stable, anti-
cyclonic weather with temperature Inversion and fog, has been ascribed
by some authors to the joint effect of aerosols of sulfuric acid and
sulfur dioxide, as a result of which the overall toxic effect is increased
(Ryazanov, 1957). Characteristically, a sharp Increase in sulfuric acid
content of the air is observed during fogs.
At the present time, data are being received indicating that sulfurous
anhydride is most dangerous in the presence of water vapor and surface-
active dust, particularly soot, when it oxidizes into sulfuric anhydride
and forms sulfuric acid. This concurs with the higher resistance of plants
in dry weather. The toxicity of sulfur gases is to some degree also Increased
if carbon monoxide, with admixtures of aldehydes and especially ozonides,
are present. The toxicity of sulfur gases is greatly increased by the
presence of nitric oxides (Lent, 1959).
Ammonia and sulfur dioxide are the constant byproducts of widely
distributed types of industry. Ammonia is given off during the process of
making fertilizers, nitric acid, ammonium salts, and during the dry distill-
ation of coal, etc. Hydrogen sulflde is given off during the processing of
inorganic sulfur compounds from oil refining processes, from the manufacture
of artificial silk, etc. Even more so, it is impossible to consider incidental
the sulfurous anhydride that occurs under industrial conditions wherever
there takes place burlning of substances containing sulfur: in non-ferrous
metallurgy (the smelting of copper, zinc, nickel, etc.) and in fuel-power
plants that burn coal and fuel oil. The following data give an idea of the
amount of contamination in these cases. For example, the TETs-3 (heat and
electric power plant) in Minsk daily emits into the atmosphere such a quantity
of sulfurous anhydride that it is the cause for the massive dying of the
pine trees in the Chelyusklntsev cultural and recreational park, which is
located 5 km from the source of emmission.
The metropolitan parks in Sokol'niki and Izmailov are in good condition
only because during the course of more than twenty years, the susceptible
plants were replaced with more gas-resistant ones. But for these measures,
many areas (in these parks) would now be wastelands unfit for the recreation
of the working people.
Concrete measures for the preservation of valuable forest stands around
industrial centers of the country can be planned only in the course of many
years of scientific investigations that take into consideration the effects
of different industrial enterprises in relation to the kind and quantity of
their waste discharge, and in relation to other ecological factors as well
as the biological peculiarities of different species of plants.
- 43 -
-------
Literature
Abramashvlll, G. G. Vliyanie zagryaznenly atmosfernogo vozdukha na khvoynye
nasazhdeniya. -Gigiena i sanitariya, 1957, No. 4.
Voroshilov, lu. I., Nedotko, P. Ispol'zovanle mlneral'nogo topllva 1
Izmenenie prirodnoy sredy.- Okhrana prirody 1 zapovednoe delo v SSSR,
1960, No. 6.
Goldberg, M. S. Sovremennoe sostoyanie voprosa ob izuchenii zagryazneniya 1
samoochlshcheniya atmosfernogo vozdukha, V sb.: "Zagryaznenle 1 samo-
ochishchenle vneshney sredy", 19A9.
ll'yushin, I. P. Usykhanie khvoynykh lesov ot zadymleniya. M.-L. 1953.
ftrasinskiy, N. P. Dymoustoychivost* rasteniy 1 dymoustoychivye assortimenty.
Sbornik rabot. Gor'kiy - M., 1950.
Kroker, V. Rost rasteniy. M., NL., 1950.
Kuz'min, M. K. Deystvle dyma na rastitel'nost'. - Lesnoe khozyaystvo, 1950, No. 6,
Lugovoy, E. V. Vliyanie gazov i pyli na khvoynye nasazhdenya Podmoskov'ya.
Lesnoe khozyaystvo, 1960, No. 7.
Ryazanov, V. A. Sanitarnaya okhrana atmosfernogo vozdukha. M., Izd-vo min-va
kotranun. khozyaystva, 195A.
Ryazanov, V. A. 'Novye dannye po eksperimental'nomu obosnovaniyu predel'no
dopustimykh Kontsentratsiy atmosfernykh zagryazneniy. V sb.: "Predel'no
dopustimye kontsentratsii atmosfernykh zagryazneniy", 1957, No. 3.
Timofeev, V. P. Vozstanovlenie khvoynykh lesov Podmoskov'ya. - Lesnoye
khozyaystvo, 1956, No. 11.
Trudy nauchno-tekhnicheskoy koraissii po obsledovaniyu moskovskikh gorodsklkh
parkov kul'tury i otdykha. 1949. M. Rukopis'.
-------
SOME PECULIARITIES OF THE SUSCEPTIBILITY
OF SCOTCH PINE SPROUTS TO SULFUR DIOXIDE INJURY
S. A. Mamaev and V. S. Nikolaevskiy
Institute of Plant and Animal Ecology of the Ural Branch of the Academy of Sciences, U.S.S.R.
From Akad. Nauk SSSR Ural. Filial. Trudy Instituta Ekologii Rasteniy i
Zhivotnykh. Fiziologiya i ekologiya drevesnykh rasteniy, Vol. II
Materialy II Ural'skogo Soveschaniya, (Sverdlovsk, 1968), p. 203-207.
Among specimens of a given botanical species there occur a large
variety of morphological variations and biological peculiarities as well
as differences in physiological-biochemical indices and in chemical compo-
sition. These intraspecific differences may, directly or indirectly,
condition dissimilar resistance of the individual plants to unfavorable
external environmental conditions. It is well known that biotypes exist
which are more or less resistant to high soil salinity (Strogonov, 1962)
or more or less frost-resistant, etc. In the verdant plantings of many
industrial cities of the Urals we have observed in lilacs, poplars and
other species, a differentiation of plants in the degree of their sus-
ceptibility to injury from toxic gases. In addition, V. Kroker (1950)
presents data indicating that individual varieties of garden roses are
more resistant to mercury vapor than others. However, data on intra-
specific differentiation of plants according to the degree of gas resist-
ance are very scarce, although such data would be of great scientific
and practical interest. For example, the lack of records concerning
intraspecific variability may possibly explain the numerous discrepancies
in scales of various authors when evaluating the degree of gas resistance
of a number of species. In light of the above, we have endeavored to
clear up certain aspects of plant variability in relation to the degree
of their gas resistance. Experiments were carried out to determine the
effect of sulfur dioxide on the sprouts of pine seeds that were obtained
from different maternal specimens. These were taken from plants growing
together in the pine forest of Chaykovskiy industrial forest administra-
tion of the Perm region. For the germination, the seeds used were from
cones collected near the middle part of the tree crowns. The germination
tests were carried out on filter paper in petri dishes, during a period
from the 14th of August to the 14th of September, 1964. Into each dish
were placed 50 seeds. Thirty-day-old pine sprouts were subjected to fumi-
gation with sulphur dioxide in a gas chamber for a period of 17 hours
(from 4 p.m. on the 14th of September to 9 a.m. the 15th of September).
The initial concentration of gas was made up earlier and was balanced out
at 1.62 x 10~5 [Tech. lid.'s Note: No units given for this quantity.] of
sulfur dioxide by volume. The degree of injury was estimated according to
the percentage of the yellowed part of the cotyledon: n**2l. x 10°^ where
1
Ll is the length of the yellowed part of the cotyledon in millimeters, 12
the overall length of the cotyledon, and// is the percent of injury.
- 45 -
-------
From data reported earlier (Kroker, 1950; Nikolaevskiy, 1964), it is
known that the water supply to a plant plays an important role in gas
resistance. A worsening of conditions of water supply affects the dynamics
of the types of water in the plant, as well as the stomatal apparatus of
leaves (Alekseev, 1948) which in turn is reflected in the gas resistance
of the plant. For a clarification of the role of the water regime in our
experiments, part of the plants were subjected to fumigation with sulfur
dioxide at the same concentration for a period of 17 hours after drying
for two days following cessation of watering.
As shown in Table 1, there is no definite correlation between the
morphophysiological characteristics of the parent tree, and its seed, and
susceptibility of the sprouts to gas injury.
The only correlation established was that of an inverse correlation
9f susceptibility to gas injury and the degree of development of the sprout.
For example, the correlation coefficient, r, of the length of the cotyledon
and the degree of damage for family No. 7 is 0.18, for No. 11 it is 0.24,
and No. 12 is 0.26, while the ratio r/mr was the same from 0.8 to 1.5.
Within the limits of one family a great variability of injury of the
sprouts was observed as a result of their unequal conditions, different
micro-environments, and, perhaps, also due to genetic segregation. There-
fore, the coefficient of variation, C, for the entire group reaches a
magnitude of 39.6%, and within the limits of the separate families the
value was even higher (tree No. 7, 42.7% and tree No. 12, 81.8%). It can
be supposed that a considerable variation in extent of injury, both in the
sum of all lots and within the limits of the families, depended on the
degree of development of the plant. The larger plants generally were more
resistant to sulfur dioxide, while the progeny of one family varies greatly
in resistance depending on the size of the seedlings. In connection with
this, it was observed that an inverse correlation existed between the
extent of injury and the size of the cotyledon (within each family).
Water provision also influences the degree of injury to the sprouts.
An experiment, carried out with the same plants, showed a decrease in
seedling damage of around 15.8%, if the sprouts had not been watered for
the previous two days. Additional experiments are necessary to elucidate
the reasons for the water effect. It is possible that individual differences
in moisture provision, as well as in the vigor of seedling development, may
be responsible for differences in resistance to sulfur gas exhibited by
different families. On the other hand, a worsening of the condition of water
supply results in a decrease of the degree of stomatal openings, which could
also affect the susceptibility of the sprouts to gas injury (Alekseev, 1948;
Nikolaevskiy, 1964).
Of considerable interest is the mechanism of the action of harmful gases
on the pigments of the plants. Insofar as it has been reported in the liter-
ature (Krasinskiy, 1950; Noack, 1924; Jahnel, 1954), in the green plants the
pigments are among the first ones to be affected by acidic gases. Hence, a
study was made of the content of the various pigments in the cotyledons of
- 46 -
-------
Table 1
Characteristics of pine sprouts obtained from seeds of various trees and
sulfuric anhydride damage in relation to experimental variables.
(I - without drying the sprouts, II - with drying)
Number
of the
trees
2
7
9
11
12
17
18
22
32
34
Measure-
ment of
the trees
/_s
-" 0) -^
4-1 0) 4-> 4-1
r* 4-J Q) C
oo Q) S a)
T* B m u
24
22
26
26
26
26
27
28
27
26
33
28
26
37
35
32
42
38
32
30
Fruiting
Strong
Weak
Strong
Weak
Strong
Ave rage
Weak
Strong
Strong
Very strong
Ave rag
Average
weight
of one
seed
(milli-
grams)
5.58
5.63
4.61
5.22
4.68
5.54
4.83
4.42
5.85
4.31
Color-
ation
of the
seeds*
BB
MB
BB
BB
Br
G
Br
Bl
G
Bl
Germin-
ation-
of the
seeds
I
72
56
86
42
58
34
92
88
70
90
68.8
II
92
38
96
36
48
38
82
76
76
84
66.8
Average
number of
cotyle-
dons
on one
sprout
I
4.90
5.10
5.42
4.97
5.50
5.47
5.64
5 . 10
6.03
5.65
5.38
II
5.70
4.62
5.50
4.82
5.50
5.60
5.20
5.48
6.13
6.14
5.46
Average
length
of coty-
ledon
(milli-
meters)
I
16.1
17.8
18.7
16.8
15.0
16.5
18.9
14.5
18.9
17.2
17.0
II
14.9
19.4
18.5
18.5
13.6
16.6
19.4
15.5
21.7
16.3
17.4
Average
length of
damaged
parts of
cotyledons
(milli-
meters)
I
4.07
3.92
7.82
12.07
6.37
7.25
6.87
9.90
5.08
6.22
6.95
II
4.97
4.10
2.42
11.40
4.47
2.48
5.20
3.06
2.24
3.33
4.36
Damage of the
length of the
cotyledon in %
I
26.6
22.0
41.7
71.2
42.5
43.9
36.4
68.2
26.9
36.2
41.6
II
33.2
22.1
13.0
62.8
34.0
15.4
28.2
18.4
10.3
20.5
25.8
*Abbreviation meanings: BB - black-brown, MB - motley-brown, Br - brown,
G - gray, Bl - black
-------
pine sprouts that were fumigated with sulfuric acid vapor for 17 hours,
as well as of the sprouts of the controls. The results of the experiment
are shown in Table 2.
As a result of the effect of the sulfuric acid fumes, the main damage
was done to the upper part of the cotyledon. In an average injury of the
length of the cotyledons (30.850 chlorophyll A and B were completely destroyed,
in the damaged as well as in the undamaged part. The same occurred with
carotene. In the damaged part of the cotyledon, violaxanthin was also
totally destroyed, while in the undamaged portion only about one-half
(56.7%) of it was destroyed. In both parts of the cotyledon chlorophyll
A and B were changed into phaeophytin, a band which was well delimited on
'chromatograms. Similar reports may be found in the literature (Krasinskiy,
iVSO; Jahnel, 1954) on the conversion of chlorophyll into phaeophytin under
the influence of acidic gases.
According to the data of several authors (Sapozhnikov, 1937, 1963;
Eydel'man, Khodzhaev, 1963), xanthophylls are formed upon oxidation of caro-
tene. Since we did not observe an increase in xanthophylls, we can surmise
that if a transformation of carotene into xanthophyll occurred; than some of
the forms of xanthophyll are also subject to oxidation. This was apparent
in the case of violaxanthin: in the damaged part of the cotyledon it was not
detected, but in the undamaged part it remained at 56.7%. Evidently, a
greater oxidizing capacity resulted in the above phenomenon as a consequence
of sulfuric acid fumes.
As a result of the influence of the sulfuric acid fumes, the ratio of
chlorophyll A to chlorophyll B, carotene to xanthophyll, and chlorophyll to
carotene decreased and approached zero, but the ratio of lutein to violax-
anthin Increased. Consequently, under the influence of strong oxidants in
plants, first chlorophyll A and B are destroyed; from among the carotenoids -
- the most reduced was carotene, and from among xanthophylls - violaxanthin.
The most resistant is lutein, possibly due to additional transformation of
violaxanthin into lutein upon oxidation (Popova et al., 1964; Sapozhnikov
et aL, 1962).
Conclusions
1. A variability of gas resistance of 30-day-old pine seedlings to
sulfur dioxide was determined both for the entire group of plants as well
as within the limits of separate families. One of the causes of the differ-
ential plant damage by sulfur dioxide was found to be due to differences in
the micro-environment of the habitat (i.e., water availability) and probably,
individual peculiarities of the organism.
2. Under the influence of sulfuric acid fumes, in cotyledons of pine
sprouts, chlorophyll A and B are converted to phaeophytin, carotene is
entirely destroyed, and violaxanthin is partly destroyed. The most resist-
ant to the action of sulfuric acid fumes is lutein. Neoxanthin is less
resistant than lutein.
- 48 -
-------
Table 2
The content of pigments in cotyledons of the pine tree
which have been fumigated by sulfur dioxide.
Pigments and their correlation
in cotyledons
Before
fumigation
After
fumigation
milligrams in 1 gram of
raw moist weight
After fumi-
gation, % of
the control
Upper half of cotyledon
Chlorophyll A
Chlorophyll B
Carotene
Xanthophylls :
Lutein
Violaxanthin
Neoxanthin
Ratio:
Chlorophyll A to Chlorophyll B
Carotene to xanthophyll
Lutein to violaxanthin
Chlorophyll to carotene
0.457
0.245
0.013
0.112
0.102
0.091
1.86
0.04
1.09
2.21
0
0
0
0.116
0
0.083
0
0
c-o
0
0
0
0
103.5
0
91.0
_
_
_
-
Lower half of cotyledon
Chlorophyll A
Chlorophyll B
Carotene
Xanthophylls :
Lutein
Violaxanthin
Neoxanthin
Ratio:
Chlorophyll A to Chlorophyll B
Carotene to xanthophyll
Lutein to violaxanthin
Chlorophyll to carotene
0.463
0.206
0.021
0.093
0.067
0.119
2.24
0.07
1.39
2.23
0
0
0
0.091
0.038
0.064
0
0
2.40
0
0
0
0
97.7
56.7
53.8
—
_
_
^
- 49 -
-------
Literature
Alekseev, A. M. Vodnyy rezhlm rastenlya 1 vllyanle na nego zasukhl.
Kazan', Tatgosizdat, 1948.
Krasinskiy, N. P. Teoretlcheskie osnovy postroeniya assortimentov gasoustoy-
chivykh rasteniy. - Dymoustoychivost1rasteniy i dymoustoychivye
assortimenty. Gor'kly - M. Izd-vo Gor'kovskogo gos. un-ta, 1950.
Kroker, V. Rost rasteniy. M. Izd-vo Inostr. lit., 1950.
Nikolaevskiy, V. S. Nekotorye anatomo-fiziologicheskie osobennosti drevesnykh
rasteniy v svyazl s Ikh gazoustoychlvost'yu v usloviyakh medeplavil'noy
promyshlennosti Srednego Urala. Avtoref. dlss. Sverdlovsk, 1964.
Popova, I. A., Bazhanova, N. V., Sapozhnikov, D. I. Nekotorye osobennosti
svetovogo prevrashchenlya ksantofillov v izolirovannykh khloroplastakh.-
Bot. zh., 1964, No. 6.
Sapozhnikov, D. I. Prevrashchenle karotlna v ksantoflll pri fotoreduktsil
ugol'noy kisloty.- Blokhlmiya, 1937, t.II, vyp. 5.
Sapozhnikov, D. I. Karotlnoldy, kak uchastnikl perenosa klnloroda prl
fotoslnteze. -Pervyy Vsesoyuznyy blokhltnicheskiy s'ezd (tez. dokl.).
M. Izd-vo AN SSSR, 1963.
Sapozhnikov, D. I. Eydel'man, Z. M., Bazhanov, I. A., Maslova, T. G.,
Popova, 0. F. K voprosu ob uchastli karotlnoldov v protsesse fotoslnteza.
-Tr. Bot. In-ta 1m. V. L. Komarova AN SSSR, Eksperlm. hot., 1962, vyp. 15.
Stroganov, B. P. Soleustoychlvost1 rasteniy M., Izd-vo AN SSSR, 1962.
Eydel'man, Z. M., Khodzhaev, A. S. 0 dinamlke svetovoy reaktsii vzalmoprevrash-
chenlya ksantofillov v svyazl s kharakterom biosinteza pigmentov
plastid prl zelenenli. -Pervyy Vsesoyuznyy biokhimicheskiy s'ezd
(tez. dokl.), M. Izd-vo AN SSSR, 1963.
Jahnel H. Uber physlologlsches Elnwirkung von Schwefeldlox auf die Pflanzen.-
Wiss. Zs. Techn. Hochschule, 1954, Bd 4, No. 3.
Noack K. Untersuchungen uber die Elnwirkung Schwefelsaure auf die Pflanzen.
Berlin, 1924.
- 50 -
-------
GAS RESISTANCE OF PINE AND BIRCH
Yu. Z. Kulagin
Bashkir University
From Akad. Nauk SSSR Ural. Filial. Ural. Cos. Univ. im. A. M. Gor'kogo.
OkTTrana prirody na Urale. Vol. A, (Sverdlovsk, 1964) p. 115-122.
Under the conditions prevailing in the industrial areas of the Urals,
the study of smoke resistance of plants and, above all, of trees, is moti-
vated not only by the interest in landscaping of industrial and residential
areas but also by the needs of forestry. The withering of coniferous moun-
tain forests, primarily of pine stands, on tens of thousands of hectares,
caused by smoke from factories, poses an urgent problem of forest restora-
tion. However, the severe climatic and the unique forest-growing conditions
of the mountain slopes of the Urals greatly limit the number of species of
trees and shrubs capable of forest formation. Because of this, in the
selection of smoke-resistant species of trees and shrubs, the major attention
must be devoted to the local tree flora. In the present work we shall
analyze the gas resistance of the European white birch and Scotch pine in the
environs of the city of Karabash and its copper smelter.
The natural conditions of the eastern foothills of the Southern Urals
favor two species as forest plants on its mountain slopes: the Scotch pine
and the European white birch. Birch-pine forests covering the foothills were
designated by B. P. Kolesnikov (1961) as characteristic for the mountain for-
est areas of the East Ural Province where Karabash is located. The European
white birch and the pine are characterized by many authors (Krasinskiy, 1937;
Krasinskiy and Knyazeva, 1950; Kuntsevich and Turchinskaya, 1957; Vanifatov,
1959) as being highly susceptible to gas and incapable of growing in smoke-
polluted areas. Yet, some other investigators (Ilyushin, 1953; Bulgakov,
1961) suggest the possibility of using the birch for the purpose of making
verdant smoke-polluted areas. N. G. Krotova (1957, 1959) considers almost
all the major forest trees, with the exception of pine, suitable for reforest-
ation of smoke-contaminated areas in the surroundings of Moscow.
Such contradictions in assessing smoke resistance of trees can be explained
to some degree by the differences of regimen, i.e., the pattern of occurrence
and the nature of smoke pollution in the different areas investigated. Indeed,
under conditions of constant or prolonged minor smoke pollution with a sulfur
dioxide concentration of less than 1 mg./m.3, all deciduous species are found
to be gas-resistant, whereas coniferous species with perennial needles suffer
severely and wither, as a result of the detrimental cumulative effect of
sulfur compounds, which gradually accumulate in the perennial needles. Under
conditions of periodic, strong gaseous attacks of short durations, with a
sulfur dioxide concentration of 30-50 mg./m.3 or more, the leaves of all
species drop. The ability of a species to survive is related to the ability
of its defoliated shoots to withstand the adverse environmental factors and
the ability to renew their foliage next season.
- 51 -
-------
Westerly winds prevail in the areas of the Urals under consideration.
Inasmuch as wind carries the smoke, the areas located northeast, east, and
southeast from the industrial enterprise can be considered as permanently
smoke-affected, whereas the areas located south, west and north of the
plant are subject to intermittent smoke attack. Investigations have shown
that the permanently smoke-affected area, which includes the so-called
"Zolotaya Gora" (the Karabash forests of the Kyshtym Forest Reserve), is
completely deforested. Thirty to sixty years ago, the local pine stands
were subjected to clear-cutting to satisfy the needs of the gold mines, the
copper smelter, and of the local population. The heavy damage to the pines
caused by smoke from the smelter was among the reasons for clear-cutting of
the forests. In the years that followed, heavy smoke pollution slowed down
and hindered the revival of forest vegetation. The western slopes facing
the smelter are also devoid of grass cover. Only the following sparsely
growing plants were recorded: sweet William, purple stonecrop, garden
burnet, bittersweet, yellow bedstraw, clover lupine, drug Solomon1s-seal,
low meadow rue, quack grass, reed grass, and others. On the less smoke-
affected eastern slopes, in areas where fresh soils are found, bunches of
quack grass and reed grass covering not over 30% of the area and growing up
to 40 cm. in height develop on gently sloping ground. It .Is only at the
foot of the eastern slope that there begin to appear solitary specimens of
European white birches.
In the area of intermittent smoke, the picture is quite different. In
places where pines were subjected to clear-cutting twenty years ago there
appeared white birches, which at the present are 17-20 years old. Their
height varies from 4 to 6 meters and their close stand is 0.5-0.7 m. The
ground vegetation covers 30-50% of the area and includes slender and Kentucky
bluegrass, reed grass and chee reed grass, clover lupine, northern bed-
straw, blueberry, libanotis, and others. The wide distribution of the
European white birch under these conditions is attributable to ecological
factors.
It should be noted that in the birch groves, located west of the smelter,
blighting by gases occurs as a rule in the second half of the growing stage
(July-August). During this period, the weather is often stormy with frequent
rain, low barometric pressure, and variable winds. By this time, birch shoots
have completed their growth; and the formed buds are well-adapted to the late
summer and fall-winter season and have a high drought and frost resistance.
According to the "Miassovo" meteorological station (Zharikov, 1959) lo-
cated in the Ilmen nursery near Karabash, July has an average precipitation
of 98 mm. or around 20% of the total annual precipitation. In some years,
July precipitation amounts to 30% of the annual. The average for May is 48 mm.
or 10%; for June, 57 mm. or 14%; and for August, 59 mm. or 12.4% of total
annual precipitation. It should be noted that in July there occur heavy and
long-lasting rains accompanied by sharp drops In barometric pressure and by
northerly and easterly winds. It is during this very time that the smoke
hovers close to the ground and affects the vegetation.
- 52 -
-------
In clear weather of antlcyclonic type with high barometric pressure,
the gases emanating from the smelter stacks rise upwards and gradually dis-
sipate in the air. Even if these gases do reach forests and ground vegeta-
tion at considerable distance from the smelter, their concentration is so
small as to be harmless. During cyclonic weather accompanied by rain, the
noxious gases hug the ground and thereby destroy vegetation. Even though
the birch crown be gravely damaged and prematurely defoliated, the birch
begins its normal growth the next season and the leaves unfold from the buds
which survived through the winter. Table 1 gives the characteristic growth
of European white birch standing 2 km. southwest from the smelter and
gravely (70-100%) damaged by sulfur dioxide in the second half of summer for
the last three consecutive years. One can assume that the crown of the birch
was damaged also in other years. In spite of that, there are no dead branches
in the crown and the growth of the current year is satisfactory.
Table 1: Characteristics of a typical 19-year old European white
birch repeatedly damaged by sulfur dioxide during the second half
of summer (July)
Height of
trunk ,
m.
Trunk
diameter
(at 1.3 m.)
cm.
5.5
5.6
Diameter
of crown,
m.
Height
of crown
start
m.
1.5
0.3
Height of increment of trunk, cm.
1961
24.5
1960
8
1959
27
1958
20
1957
13
It should also be noted that in some years, e.g., in 1960, gas attacks
on vegetation in the westerly direction occurred also in June, at the time
when the growth of the shoots was not yet complete, and the buds were not com-
pletely formed. Inspecting a damaged plot located in compartment 38 of the
Agardyash forest of the Kyshtym Reserve in 1961, it was found that the crown
of damaged birches withered. It was noticed that in birches severely damaged
by gas abundant new growth sprouted on the lower part of the trunk of trees.
In years when birches were severely damaged, they showed no "biological gas
resistance" (a term used by N. P. Krasinskiy 1937-1950), i.e. . they did not
develop new leaves immediately following the damage to their crown. This may
be related to the fact that the summer climate in this part of the Urals is
less favorable then in the central belt of European U.S.S.R. where Krasinskiy
conducted his investigations.
The pine is capable of gas resistance similar to that of the birch. The
pines in compartment 49 of the previously mentioned Reserve, occasionally
affected by gases, can serve as an example. In July 1960, vigorously growing
6- and 13-year-old pines, hitherto free of gas injury, were severely damaged
by sulfur dioxide. However on trees with healthy root systems and fully
formed buds, the remaining coniferous needles (Tables 2 and 3) were sufficient
to make possible normal sprouting in 1961. Thus, the pine after a single,
severe gas injury in the second stage of its vegetation, accompanied by the
loss of a considerable part of its foliage, is capable of renewing its growth
next season provided its buds survived during the winter.
- 53 -
-------
Table 2:
Characteristics of typical 6-year-old pines
damaged by sulfur dioxide in July 1960.
Tree
no.
1
2
3
4
5
Height,
cm.
Trunk
diara.
cm.
60
1.9
63
1.3
82
3.2
84
3.1
104
2.3
Year of
trunk in-
crement
1961
1960
1959
1958
1961
1960
1959
1958
1961
1960
1959
1958
1961
1960
1959
1958
1961
1960
1959
1958
Incre-
ment
cm.
8
3.5
16.0
9
14
23
7
11
11
15
14
22
18
32
13
15
15
40
13.5
13
Extent of
d 'image to
needles , %
0
100
0
0
0
95
77
0
90
0
0
0
68
0
0
0
71
0
0
Table 3:
Characteristics of typical 13-year-old pines
damaged by sulfur dioxide in July 1960.
Tree
no.
1
2
3
4
5
Height,
cm.
Trunk
diam.
cm.
190
5.6
195
4.5
250
6
256
6.2
267
6.4
Year of
trunk in-
crement
1961
1960
1959
1958
1957
1961
1960
1959
1958
1957
1961
1960
1959
1958
1957
1961
1960
1959
1958
1957
1961
1960
1959
1958
1957
Incre-
ment
cm.
54
49
16
21
25
18
50
27
30
16
48
57
43
35
42
40
55
46
40
40
55
54
66
16
26
Extent of
damage to
needles, %
0
75
11
—
— —
0
67
36
-
—
0
65
27
28
—
0
58
46
-
—
0
14
10
-
—
-------
The data on smoke resistance of birch and pine, quoted in this paper,
do not fit in with the widely accepted views advanced by N. P. Krasinskiy
and his co-workers (1937-1950). As is known, N. P. Krasinskiy suggests
three types of gas resistance: physiological, raorpho-anatomical, and bio-
logical. The physiological gas resistance is determined by the physiological
and biochemical properties of the plant, while the morpho-anatomical is
determined by those structural elements of the leaves which control gas
exchange and, consequently, the penetration of gases into the mesophyll.
The biological gas resistance consists of the ability of plants to renew
rapidly parts and organs damaged by gas and thus, regain their ornamental
aspects. N. P. Krasinskiy emphasizes that this type of gas resistance is
characteristic of rapidly growing trees and shrubs. E. I. Knyazeva (1950)
asserts: "...when speaking of biological gas resistance it should be remem-
bered that its extent is not constant for any one species. Biological gas
resistance can be strong in the first half of summer and then drop to almost
nothing by the end of summer. This is explained by the cessation of growth of
our trees and shrubs in the second part of summer. Plants continuing to grow
until autumn retain their biological gas resistance to the end of the growing
period. Because biological gas resistance is tied to the plant's ability to
sprout rapidly new leaves and shoots to replace the damaged ones, it depends
in large measure on the rate of growth. The latter is in a sense an index of
biological gas resistance."
Without stopping to analyze the first two types of gas resistance, let us
consider the so-called biological gas resistance in an ecological frame of
reference. It is clear from the above that a species capable of rapid regenera-
tion of its foliage during the same growth stage is endowed with high gas resist-
ance, whereas species devoid of this ability have a low gas resistance. From
this point of view, N. P. Krasinskiy and E. I. Knyazeva consider box elder,
poplar, birch, barberry, and alder buckthorn endowed with appreciable gas resist-
ance. Because of it, box elder is considered as a completely gas-resistant plant
and is recommended for planting in strongly smoke-affected areas, notwithstand-
ing that its foliage is severely damaged by sulfur dioxide. From this point of
view, the conifers, e.g., pine, spruce, and fir, are considered totally unsuit-
able for planting in such areas, because of their lack of biological gas resist-
ance, although their needles, because of their anatomical and morphological
characteristics, are but slightly affected by sulfur dioxide.
Our observations show that in conditions prevailing in the eastern foot-
hills of Southern Urals, the birch is devoid of the so-called biological gas
resistance. Defoliation of birches by gases in the first half of the summer
leads to withering of the crown during summer drought or during severe winter
conditions. Birch groves located west from the source of smoke are as a rule
subject to smoke damage in the second half of their growth stage when they are
devoid of leaves and display a high drought and frost resistance; this explains
the successful growth of birches under conditions of intermittent gas attack.
Therefore, the faster the completion of growth of shoots and the formation of
buds in the crown, the higher is the resistance of the tree. Thus, of special
interest are littleleaf linden and Siberian pea shrub, both having a short
growth period and a high drought resistance of the shoots. Extended growth of
shoots does not contribute to greater resistance of plants in the Ural region
- 55 -
-------
under discussion, because severe winter conditions kill these shoots,
although they may show a greater biological gas resistance during the
vegetative season.
Conclusions
1. Gas resistance of woody plants depends in great measure on the
forms of smoke pollution and on local climatic conditions. The smoke in-
jury to forests and their destruction under the conditions existing in the
Southern Urals is intimately connected with the zonal climatic conditions,
and, above all, with the weather during the growth season.
2. Two of the most important forest forming species - European white
birch and partly Scotch pine - can thrive satisfactorily under conditions
of intermittent severe gas attacks of short duration. Their gas resistance
is conditioned by the occurrence of these attacks in the second half of the
vegetative season. Normally, rainy weather, together with changeable winds
and low barometric pressure, occurs in July and pushes the noxious gases
down close to the ground.
3. The growth and formation of shoots is almost complete during May
and June; the absence of the so-called biological gas resistance but high
drought and frost resistance of defoliated birch and pine shoots insures
their viability in late summer, autumn and winter, and resurgence of vege-
tative processes the following season.
A. In Karabash, for green plantings located to the south, west and
north of the smelter European white birch should be considered most import-
ant. For aforestation of the mountain slopes littleleaf linden and Siberian
pea shrub may be used.
- 56 -
-------
Literature
Bulgakov, M. Vyrashchivanie berezy na pochvakh, otravlennykh otkhodami
proizvodstva. Sb. "Obmen opytom po zelenomu stroitel'stvu", Sverdlovsk,
1961 (Ural'skiy n.-i. in-t AKKh im. K. D. Pamfilova).
Vanifatov, D. N. Okislyaemost' kletochnogo soderzhimogo kak pokazatel'
gazoustoychivosti rasteniy. Referaty dokladov nauchnoy konferentsli po
ratsionalizatsli lesnogo khozyaystva i agrolesomelioratslt Kazakhstana,
Alma-Ata, 1959 (Kazakhskiy s.-kh. in-t).
Zharikov, S. S. Klimat rayona Il'menskogo zapovednika i sopredel'nykh
prostranstv Yuzhnogo Urala. Trudy Il'menskogo gos. zapovednika im.
V. I. Lenina, vyp.7, Izd. UFAN SSSR, Miass, 1959.
Ilyushin, I. R. Usykhanie khvoynykh lesov ot zadymleniya. M.-L., Goslesbura-
izdat, 1953.
Knyazeva, E. I. Gazoustoychivost* rasteniy v svyazi s ikh sistematicheskim
polozheniem i morfologo-anatomicheskimi i biologicheskimi osobennostyami.
Sb. "Dymoustoychivost1 rasteniy i dymoustoychivye assortimenty", Gor'kiy -
Moskva, 1950 (Gor'kovskiy gos.un-t i AKKh im. K. D. Pamfilova).
Kolesnikov, B. P. Lesorastitel'nye usloviya i lesokhozyaystvennoe rayonirovanie
Chelyabinskoy oblasti. Trudy In-ta biologii UFAN SSSR, vyp.26,
Sverdlovsk, 1961.
Krasinskiy, N. P. Ozelenenie promploshchadok dymoustoychivym assortimentorn. M. ,
Izd-vo "Vlast1 Sovetov", 1937.
Krasinskiy, N.P. i Knyazeva, E. I. Dymoustoychivye assortimenty. Sb. statey
"Dyraoustoychivost1 rasteniy i dymoustoychivye assortimenty". Gor'kiy -
Moskva, 1950 (Gor'kovskiy gos. un-t i AKKh im. K. D. Pamfilova).
Krotova, N. G. Vliyanie zadymleniya vozdukha na sosnu v Lesnoy opytnoy dache
sel'skokhozyaystvennoy akademii im. K. A. Timiryazeva i meropriyatiya po
sozdaniyu ustoychivykh nasazhdeniy. Avtoreferat kand. dissertatsii,
Fondy Mosk. ordena Lenina s.-kh. akademii im. K. A. Timiryazeva. M. , 1959.
Kuntsevich, I. P. i Turchinskaya, T. N. Ozelenenie fabrichno-zavodskikh plosh-
chadok i promyshlennykh poselkov. Izd. MKKh RSFSR, M., 1957.
- 57 -
-------
THE EFFECT OF SULFUR DIOXIDE ON WOODY PLANTS UNDER
THE ENVIRONMENTAL CONDITIONS PREVAILING IN THE SVERDLOVSK REGION
V. S. Nikolaevskiy
Institute of Biology* Ural Branch, USSR Academy of Sciences
From Akad. Nauk SSSR Ural. Filial. Ural. Cos. Univ. im. A. M. Gor'kogo.
Okhrana prirody na Urale. (Sverdlovsk, 1964) 4:123-132.
An important aim in the investigation of the effect of various gases
on plants is to study the physiological and biochemical reasons for the
different degrees of resistance of plants to these gases, as well as to
reveal and to ascertain the type of plants most resistant to the condi-
tions of industrial air and soil pollution prevailing in a given area.
Of equal importance is the study of methods for increasing the resistance
or plants to gases.
Although the first investigations of gas resistance of plants were
undertaken long ago both in our country (1900-10) and abroad (1860-70) ,
there is still much in this area which remains unclear. The physiological
processes in plants under the influence of acid gases are still almost
unknown. Nor is there a scientifically based biochemical and physiological
theory that would explain the reasons for plant damage caused by gases, or
a complete understanding of the different degree of susceptibility of
various plant species.
V. Sabashnikov (1911), N. P. Krasinskiy (1937-40), V. Krocker (1950),
I. K. Fortunatov (1958), and M. A. Zheleznova-Kaminskaya (1951, 1953)
summarized the work of various authors concerning gas resistance of plants.
They all consider that acid smoke gases affect photosynthesis, respiration,
transpiration, as well as the water regime of plants and also cause acidifi-
cation of the protoplasm and a change in the ionic balance, which affect
the stability of the biocolloids in the plasm.
Many of our investigators (Krasinskiy, 1937, 1940, 1950; Krocker, 1950;
Ryabinln, 1962), as well as foreign Investigators, established that under
the Influence of acid gases 1) the cell sap becomes acidified, 2) the intens-
ity of photosynthesis declines, 3) the respiration first drops, and then
Increases, according to Vlller (1905), while, according to Krasinskiy (1950),
respiration remains unchanged or increases, 4) the Intensity of metabolism
is lowered, and the redox potential changes, 5) sulfur and other elements
accumulate in the leaves, 6) the amount of ascorbic acid declines, and
7) the drain of assimilators ceases (Viller, 1905).
In severe Injuries, the chloroplasts lose their bright color, swell,
and even adhere to the cell wall. In damaged cells the chlorophyll changes
to pheophytln.
A most general theory of gas injury to plants was advanced by Noack (1920)
and developed by N. P. Krasinskiy (1949-50). This theory states that acid
- 58 -
-------
gases disturb and arrest the activity of enzymes connected with photo-
synthesis. Chlorophyll, deprived of its basic function to assimilate
carbon dioxide from the air, continues to accumulate photoenergy, acts
photodynamically, destroying itself and the substances surrounding it.
At the same time there is the formation of phenophytin.
Of great significance to plants is the permeability of protoplasm
and its buffer capacity as well as the stability of the plasma biocolloids
and of the enzyme complex.
In our study of gas resistance we confined ourselves to investigating
photosynthesis and respiration, water conditions, pH, Eh, and rl^ of cell
sap, the movement of stomata, and the quantity of readily oxidizable sub-
stances. In addition, we studied the anatomical properties of leaves and
the dynamics of their susceptibility to injury in woody plants growing under
conditions of the copper smelting industry of the Central Urals (Krasnoural'sk,
Kirovgrad, and Revda), as well as in a laboratory gas chamber where cut
branches of plants were used.
In the first stage of this work, in 1959-60, the physiological indices
were studied for five species: balsam poplar, dwarf apple, European white
birch, aspen, and box elder. The investigation was carried out under con-
ditions of intermittent gassing (200 m. from smelter) and under conditions
free of gases (trees in a forest glade of about 500 m. diameter and at a
distance of 7 km. from the smelter). In the forest glade study the naturally
growing aspen and European white birch were utilized. In addition, in the
fall of 1959, there were planted balsam poplar, dwarf apple, and European
white birch. All analyses were carried out in quadruplicate, three times
daily (at 7 A.M., 1 P.M., and 7 P.M.).
According to literature data (Krasinskiy, 1950; Vanifatov, 1959;
Bulgakov, 1959, 1961; lonin, 1961) as well as according to our own observa-
tions of the degree of susceptibility of leaves to injury, by the Krasinskiy
(1950) method, the trees under study can be grouped as follows: mildly suscep-
tible (box elder); moderately susceptibile (balsam poplar and aspen); and
very strongly susceptible (dwarf apple and birch).
The investigations established a relationship between the resistance
of woody species to acid gases and the intensity of photosynthesis and
respiration. The species having a higher intensity of photosynthesis and
respiration are injured to a greater extent by acid gases (dwarf apple and
birch).
The extent of daily variations of readily oxidizable substances in the
leaves is apparently related to the intensity of physiological processes
(photosynthesis, respiration, and others). The variations during the day
are insignificant (2-4 ml./I g. of green weight) for the mildly susceptible
box elder, whereas for the strongly susceptible dwarf apple and birch the
variations were 8.5-19.5 ml.
- 59 -
-------
It was also found that the gas resistance of the investigated species
is correlated to the total water content in the leaves. The leaves of the
gas-resistant box elder had a higher water content. The concentration of
the cell sap was lowest in the mildly susceptible box elder (9-10%) , where-
as in the dwarf apple and birch it was one and a half times higher and
more.
In this article we shall examine also the effect of sulfur dioxide on
the performance of the stomatal apparatus and on some physiological indices:
photosynthesis, pH, and the concentrations of cell sap and of oxidizable
substances in the leaves. The dynamics of the stomata movement in the course
of the day was studied following the penetration of alcohol, benzene, and
*ylol into the intercellular spaces of leaves. To facilitate the compari-
son of stomata movement in various species, the following designation was
adopted: first degree - implies the penetration of xylol only,, (the stomata
are practically closed); second degree - refers to the penetration of xylol
and benzene, (stomata half open); and third - refers to the penetration of
all three (which implies that the stomata are fully open). Taking into
account the degree of stomata opening we shall proceed to analyze the
stomata movement in leaves of woody plants growing under conditions of Inter-
mittent gassing (in a city park) and under conditions free of gas (in a for-
est) — see Fig. 1, a and b.
Under environmental conditions free of gas (Fig. 1, b) leaves of
naturally growing aspen and birch were studied. In addition, the previous
year balsam poplar and European white birch were planted in a forest clear-
ing. Obviously, data on trees transplanted into forest environment cannot
be accepted as controls but they will enable us to draw some other conclusions,
In the aspen and birch growing naturally in the forest, the average opening
of the stomata in June was higher than in the transplanted trees. The dynam-
ics of the stomatal movement in the transplanted trees indirectly confirms
the conclusions of A. M. Alekseev (1948) that plants growing in conditions
of dry soil reduce the extent of their stomata openings. Weakening of the
root system induced by transplantation apparently creates conditions resembl-
ing those characteristic of dry soils. The stomata of the naturally growing
aspen and birch showed no substantial difference in movements that could
influence the rate of their gas metabolism. In all the species growing
under forest conditions noticeably enlarged stomatal openings were observed
in the fall.
Under conditions of intermittent gassing (Fig. 1, a) the average opening
of the stomata of the poplar, aspen and dwarf apple was larger than in the
box elder and birch. Considerable reduction in the stomatal movement of the
birch may be attributed to a deterioration of the tree caused by the gas -
a condition that could also be determined from the appearance of the tree's
crown.
Observations of the performance of the stomatal apparatus of the woody
species during the summer (Fig. 1, a) show that toward the end of the grow-
ing season there was even a greater reduction in the stomatal movement of
the box elder - a species of low susceptibility to gas injury. The degree
- 60 -
-------
of stomatal opening of the European white birch also decreased, but this
was apparently due to the considerable gas injury sustained by the tree.
The regulatory apparatus of the stomatal movement of trees exposed to
intermittent gassing (Fig. 1, a) reacts differently in different species
while under conditions free of gas (Fig.1, b), the stomata openings enlarge
in the autumn, as was to be expected from literature data (Alekseev, 1948).
0)
a
o
CO
00
c
1-1
c
OJ
K
VI
vn.
, Fig.l Change in the average stomata openings during the
growing season by month (monthly averages based on
5-7 days' observations).
a - city park; b - forest
1 - European white birch; 2 - box elder; 3 - dwarf apple;
4 - aspen; 5 - balsam poplar; 6 - European white birch,
transplanted into the forest glade.
The difference in the movement of the stomata of the trees (Fig. 1,
a and b) we are inclined to attribute to the intermittent action of acid
gases. This is particularly noticeable in the birch. The greater opening
of the birch stomata in July is related to a change of its foliage, which
resulted from gas burns. The opening of the stomata of the new foliage on
the birch as yet not affected by gassing is the same as in the control
plants. It is quite likely that the data for the poplar in autumn (Fig. 1, a)
are somewhat higher because the poplar and the aspen had changed their foliage.
It should also be taken into account that conditions of microclimate and
soil in the city and in the forest are not identical. According to L. B. Lunts
(1952), the city microclimate is characterized by a greater daily amplitude
of air temperature variations, a higher air and soil temperature (A-16°C, de-
pending on the soil cover), and a lower relative air humidity. According to
our data (Nikolaevskiy, in press) the relative air humidity in the city is
lower than in the forest, in July by 14.5% and in September by 6.6%.
- 61 -
-------
According to the data of I. N. Rakhteenko and L. A. Krot (1960) the
soil moisture in cities is lower because of a greater surface runoff caused
by excessive compaction, by asphalt, concrete, and cobblestone paving.
Consequently, the dynamics of stomata motion could be affected by the less
favorable climatic and soil conditions in the city as well as by gas.
A study of the stomata movement in the course of summer helps in deter-
mining certain causes of leaf injury to the experimental species. Ivanov
(1936) and Krocker (1950) suggested a possible connection between the move-
ment of the stomata and the susceptibility of leaves to injury of the various
species. Our investigations (Nikolaevskiy, in press) confirm this view.
The severity of leaf injury must then depend on the quantity of acid
gas absorbed per unit of dry weight of leaves. The rate of gas absorption
depends on the quantity and size of the stomata openings per unit surface
pf leaf as well as on the presence of intercellular spaces. According to
the Stefan Law, a decrease in the stomata openings by half practically does
not affect the rate of gas exchange between atmospheric air and the inter-
cellular spaces (Maksimov, 1958).
A further decrease in the stomata openings will result in a smaller
but not proportional gas exchange. Therefore, a daily variation in the
openings from 3 to 2 (as previously explained) cannot be considered signifi-
cant for the rate of gas exchange. A lowering of the openings from 2 to 1,
or even lower, will affect the gas exchange. Therefore, we cannot discern
differences in the rate of gas exchange in the poplar, aspen and dwarf apple
during the summer (Fig. 1, a) but there is a discernible lowering of it in
the box elder when compared with the others. It can be assumed that a lower-
ing in the rate of gas exchange in the box elder under conditions of periodic
gassing contributes to a decrease in the intake of noxious gases, thereby
preventing graver leaf injury.
In addition to these field investigations, we set up in March experi-
ments to study the effect of sulfur dioxide on tree leaves. First, the
leaves were forced out in a hothouse. Branches were then placed in flasks
containing tap water. The leaves of various species were then fumigated in
a plexiglas gas chamber, with different concentrations of sulfur dioxide
and for different periods of time. Analyses were made the day following
fumigation.
We studied the effect of sulfur dioxide on the movement of stomata by
the infiltration method, pH changes of leaf sap potentiometrically, photo-
synthesis by the method of L. A. Ivanov and L. N. Kossovich (1946), the
quantity of readily oxidizable substances in the leaves by the N. P. Krasin-
skiy method (1950) , and the concentration of dry matter in the cell sap
refractometrically. The results are given in Table 1.
The results show that under the influence of S02, the stomata after
fumigation were less open than in control plants, even one day later.
A mere 10 minutes' exposure to an S02 concentration of 1/8000 by volume
(experiment No. 1) caused a decrease in the extent of stomata openings.
- 62 -
-------
Table 1
Effect of SO? on leaves of woody plants based on the results of investigations in a gas chamber.
Species
Exp.tfl
Hairy lilac
Balsam poplar
Dwarf apple
European white birch
Exp. #2
Balsam poplar
Dwarf apple
European white birch
Exp. #3
Balsam poplar
Dwarf apple
European white birch
Exp. #4
Balsam poplar
Dwarf apple
European white birch
Degree of
stomata openings
Exp.
Concen
2
1
2
1
Control
tration of
3
2
3
2
Visible photo-
synthesis ,
mg. CC>2 per
1 dm./hr.
Exp.
S02 -
+0.5
-1.3
-4.0
-9.2
Control
1/8000, du
+1.6
+0.7
+2.3
+21.2
Concentration
of cell sap, %
Exp.
ration
12.0
13.0
11.0
13.5
Control
10 min. ,
12.0
13.5
14.0
14.5
pH of cell sap
Exp.
illumii
5.9
6.3
6.1
6.2
Control
ation 15,
6.4
6.5
6.3
6.5
Oxidizable matter
per 1 g. of raw
wt. of leaves
Exp.
000 lux
20.1
19.5
17.1
23.5
Control
17.4
30.7
28.0
28.4
Damage to
leaf sur-
face, %
-
Concentration of S02 - 1/4000, duration 20 min., illumination 8,000 lux
2
2
0
3
3
2
+0.9
-1.9
-9.2
+3.5
+0.8
+3.3
-
^
^
^
—
^
10
80
Concentration of S02 - 1/4000, duration 1 hr. , illumination 10,000 lux
2
1.5
2
3
3
2
-1.2
-4.2
-11.0
+3.6
+2.5
+1.9
_
_
Concentration of S02 - 1/4000, duration 1 hr. , i
2
2
1.5
3
3
2
-1.6
-1.4
-5.6
+0.3
+1.2
+0.4
-
-
5.4
5.5
5.7
6.1
6.1
6.3
-
50
40
100
.lumination 32,000 lux
5.6
5.8
5.8
7.2
6.6
6.4
—
-
100
100
100
-------
Data in Table 1 confirm our conclusion concerning the effect of
intermittent gassing on the dynamics of stomatal movement in woody plants
during the summer. In all cases where the leaf injury exceeded 5% of the
surface area, photosynthesis was arrested, and the respiration during
light exposure increased. S02 provoked particularly strong respiration
in birch and dwarf apple.
Sulfur dioxide caused acidification of the cell sap; it was weak
when the fumigation was brief and the SC>2 concentration low (experiment
No. 1) and stronger in extended fumigation and higher S02 concentration
(experiments No. 3 and 4). Simultaneously, there was a decrease in the
readily oxidizable substances in the leaves. An exception was lilac in
experiment No. 1. In this instance the photosynthesis did not drop and
the amount of oxidizable substances increased.
Lowering of the amount of oxidizable substances in the leaves of dam-
aged plants was apparently caused by an increased respiratory intensity.
It should be noted that in the experimental plants respiration greatly
varied in time. Because of this, data obtained by us for a decrease in
oxidizable substances in the leaves of experimental plants show no good
correlation with transpiration.
Data of foreign investigators show that sulfur dioxide in concentrations-
of 0.0000002-0.000001 by volume is capable of lowering the intensity of
photosynthesis, however, after fumigation, photosynthesis is restored. Con-
centrations of 0.000007 or higher, and with the duration of fumigation for
approximately 1 hour, bring about considerable injury to the plant (alfalfa)
- an injury that is expressed in a substantial reduction of photosynthesis
throughout the greater part of daylight during a period of 8-9 days..
Our experiments were conducted at concentrations of 100-200 times that
used by the American investigators. This is the reason why we were able to
observe complete cessation of photosynthesis, even on the second day after
fumigation, and considerable respiration in daylight. As seen in Table 1,
following fumigation there is a decrease in the concentration of cell sap
in the leaves.
Of equal interest are the data obtained in the study of the effect of
SO2 on leaves of branches cut from woody plants and treated in a gas chamber
in sunmer. The experiment was carried out for 30 minutes on July 17, 1960,
at a S02 concentration of 1/4000, under 25-30 thousand lux illumination.
Unlike the winter experiments, the analyses of the leaves were made the
same day, 4 hours after fumigation. (Table 2).
As in the previous experiments (Table 1), here too can be pointed out
that under the influence of S02 (Table 2) the stomata openings decreased,
the pH of the cell sap was lowered, and the quantity of oxidizable sub-
stances declined. The concentration of cell sap in the leaves 4 hours
after fumigation increased. It can be assumed that under the influence of
S02 there is during the first few hours an increase in the concentration of
cell sap due to sudden hydrolysis, and that on the following days there is
- 64 -
-------
a lowering in the concentration resulting from transpiration and absence of
photosynthesis.
Table 2
Effect of sulfur dioxide on leaves of woody plants.
Species
Poplar
Birch
Apple
Elm
Ash
Aspen
Box eldet
Changes of some indices under the influence of SQ2
Stomata
openings
Exp.
3.0
2.0
1.0
1.0
3.0
2.0
1.0
Control
3.0
2.5
2.0
2.0
3.0
2.0
1.0
Concentration
of cell sap
Exp.
-
16.0
20.8
20.0
17.6
24.0
-
Control
15.6
15.2
18.8
19.2
15.0
21.0
14.5
pH of cell sap
Exp.
5.08
A. 52
5.40
5.72
5.98
5.62
4.0
Control
6.20
6.08
6.22
6.02
6.05
5.92
4.05
Oxidizable matter
per 1 g. of raw
wt. of leaves
Exp.
24.8
36.0
44.0
22.6
25.6
23.4
25.0
Control
27.0
45.0
46.5
26.0
28.0
26.3
25.2
Our anatomical studies showed that under city conditions the intermittent
gassing and perhaps also the poorer microclimatic and soil conditions, bring
about an increase of xerophilization of the leaves. This is expressed in a
decrease in the size of leaves, an increase in the number of stomata per
1 mm.2, a decrease of their size, a decrease in the thickness of the cancell-
ous parenchyma, an increase in the thickness of the epidermis, and also, in
the case of strongly susceptible species (dwarf apple and birch), the loss
of one layer of palisade tissue.
Phenological observation of the development and drop of leaves of woody
plants reveals that under city conditions, and particularly under the influ-
ence of gas, the growing period is shortened. In the case of the box elder
which is more gas-resistant, the leaves drop later than those of the less
resistant species — the dwarf apple and the birch.
Conclusions
1. Under the influence of acid gases, and particularly under simultane-
ously acting urban conditions, the five investigated species: box elder,
balsam poplar, aspen, European white birch, and dwarf apple, showed a dis-
tinct reduction in the openings of their stomata in autumn; the extent of
this reduction differed for the different species. In the absence of gas-
sing, the reverse was true.
- 65 -
-------
2. The smallest opening of the stomata during the vegetative period
was observed in the box elder. This could be the reason for the decrease
in the rate of gas exchange, which is conducive to a reduction in the vul-
nerability of the leaves to noxious gases.
3. High concentration of sulfur dioxide causes considerable narrowing
of the stomata of the leaves of woody species and upsets the normal course
of physiological processes. Thus, the intensity of photosynthesis drops
and respiration is observed in daylight. In addition, acidification of
cell sap and a decrease in the quantity of oxidizable matter is also observed.
4. The greater damages, which manifest themselves in appreciable
acidification of the cell protoplasm and in an extensive disturbance of
photosynthesis, are observed in the less resistant species - dwarf apple and
birch. These disturbances appear to a lesser extent in the more resistant
species - poplar and lilac.
Literature
Alekseev, A. M. Vodnyy rezhim rasteniya i vliyanie na nego zasukhi. Kazan',
Tatgosizdat, 1948.
Bulgakov, M. V. Opyt ozeleneniya g. Krasnoural'ska. Sb. "Materialy po
ozeleneniyu gorodov Urala", vyp.l. Sverdlovsk, 1958.
Vanifatov, D. N. Okislyaemost* kletochnogo soderzhimogo kak pokazatel'
gazoustoychivosti rasteniy. Referaty dokl. Nauchnoy konferentsii po
ratsionalizatsii lesnogo khozyaystya i agrolesomelioratsii Kazakhstana,
Alma-Alta, 1952.
Zhfrleznova-Kaminskaya, M. A. Rezul'taty introduktsii khvoynykh ekzotov v
Leningrade i ego okrestnostyakh. Sb. "Introduktsiya rasteniy i zelenoe
stroitel'stvo", M.-L., Izd-vo AN SSSR, 1953.
Ivanov, L. A. Fiziologiya rasteniy. M., Gostlesizdat, 1936.
lonin, V. M. , Koltasheva, V. F. Ozelenenie sanitarno-zashchitnykh zon. V
kn. "Rekomendatsii po ozeleneniyu gorodov", Izd. UNII AKKh im. Pamfilova,
Sverdlovsk, 1961.
Krasinskiy, N. P. Ozelenenie promploshchadok dymoustoychivym assortimentom.
Izd. AKKh im. Pamfilova, 1937.
Krasinskiy, N. P. 0 fiziologlcheskoy sushchnosti gazoustoychivosti rasteniy.
Uch.zap.Gor'kovskogo gos. un-ta, vyp. 9, Gor'kiy, 1940.
- 66 -
-------
Kraslnskiy, N. P. Dymoustoychlvost1 rasteniy 1 dymoustoychivye assortimenty.
Uch.zap. Gor'kovskogo gos. un-ta, vyp.14, Gor'kiy, 1949.
Krasinskly, N. P. Teoreticheskie osnovy postroeniya assortimentov gazoustoy-
chivy kh rasteniy. Sb. "Dymoustoychivye rasteniya 1 dymoustoychivye
assortimenty". Izd. Gor'kovskogo gos. un-ta, Gor'kiy-Moskva, 1950.
Kroker, V. Rost rasteniy. Izd-vo Inostr.lit., M., 1950.
Lunts, L. B. Zelenoe stroitel'stvo. M., Goslesbumizdat, 1952.
Maksimov, N. A. Kratkiy kurs fiziologii rasteniy. M. , Sel'khozgiz, 1958.
Nikolaevskiy, V. S. Ekologo-fiziologicheskie Issledovaniya gazoustoychivosti
drevesno-kustarnikovykh porod v usloviyakh g. Krasnoural'ska. V pechati.
Nikolaevskiy, V. S. 0pokazatellakh gasoustoychivostl drevesnykh rasteniy po
issledovaniyam v g. Kraenoural'ske. V pechati.
Nikolaevskiy, V. S. Osobennosti anatomicheskogo stroeniya list'ev u razlichnykh
po gazoustoychivosti drevesnykh porod. V pechati.
Rakhteenko, I. N. i Krot, L. A. Ekologlcheskie osobennosti rosta i razvitiya
nekotorykh drevesnykh dekorativnykh rasteniy v usloviyakh gorodskogo
ozeleneniya i v estestvennykh usloviyakh. Byull. In-ta biologii AN
BSSR, Minsk, 1960.
Ryabin, V. M. Vliyanie promyshlennykh gazov na rost derev'ev i kustarnlkov.
"Bot.ah.", t. 47, No. 3, 1962.
Sabashnikov, V. Vliyanie kamennougol'nogo dyma na okruzhayushchuyu rastitel'nost1
"Bolezni rasteniy", No. 314, 1911.
Fortunatov, I. K. Kriticheskiy obzor amerikanskikh rabot po vliyaniyu promyshlen-
nykh dymov i gazov na rasteniya. Dokl. Sel'skokhozyaystvennoy akademli im.
Timiryazeva, vyp. 36, M., 1958.
- 67 -
-------
CHARACTERISTICS OF PHOTOSYNTHESIS AND OF SOME OTHER PROCESSES
IN CONNECTION WITH SMOKE AND GAS RESISTANCE OF TREES AND SHRUBS
A. S. Sitnlkova
Karaganda Botanical Garden of the Kazakh SSR Academy of Sciences
From Akad. Nauk SSSR Ural. Filial. Ural Cos. Univ. im. A. M. Gor'kogo.
Okhrana prirody na Urale. (Sverdlovsk, 1964) 4:133-135.
In the conditions of the Karaganda region, where drought and dry winds
are frequent, the establishment of green plantings is of exceptional impor-
tance.
However, along with the local adverse conditions of soil and climate,
there is also the air pollution factor hampering green plantings in the
industrial cities and in industrial areas as a result of the gas, smoke and
dust prevailing there. The effect of gas, smoke, and dust on growth, on
development, and on the physiological processes of plants has been insuf-
ficiently studied. Investigations as to carbohydrate exchange, photosyn-
thesis, and other physiological processes are very limited as yet.
Our objective was to study the processes of growth and plant develop-
ment, photosynthesis, and the dynamics of carbohydrate and chlorophyll
accumulation, under various conditions of growing and of gas pollution.
For three years (1959-1961) we investigated the following plants: European
white birch, pinnate elm, common lilac, Tartarian honeysuckle, box elder,
balsam poplar, oleaster, Scotch pine, and Siberian pea shrub. The plants
under investigation were 3-5 years old.
The plants were studied in a number of locations subject to different
growth conditions, which can be characterized as follows:
Central Ore Concentration Mill. Five stacks located 200 m. from the
plantings discharge continuously smoke and soot. The concentration of CO
is 30.0-117.6 mg./m.3 and of S02, 0.28-1.33 mg./m.3. The air is dust-laden
and on certain days the dust concentration is 4.7 g./m.3. The industrial
site slopes toward the plantings, which are not protected by buildings from
gas and smoke. The soil of the area is dark chestnut, overlaying a mottled,
$ley, saline clay.
Karaganda Metallurgical Plant. Plantings in the vicinity of the factory
are located 300 m. from the blast furnaces and are constantly exposed to
gases and to smoke. The soil of the site is dark chestnut, of the solonchak
type, modified and polluted by construction rubble.
Karaganda Botanical Garden. Plantings in the Garden are at a distance
of 2 km. from a coal cut. In this area there is considerable gas in the air -
- CO, up to 117 mg./m.3 and S02, up to 0.25 rag./m.3 on some days; quite fre-
quently, the concentration of these pollutants is insignificant. The area of
-------
the Garden is level, and the plants are not shielded from smoke and gases.
The dust content of the air is negligible. The soil is dark chestnut,
heavy loam over calcareous clay.
During the growing period, observations were made on growth and
development of the plants, dynamics of chlorophyll and carbohydrate accumu-
lation, intensity of photosynthesis, and oxidizability of cell sap. Carbo-
hydrate analysis was done by the Bertrand method. Three fractions were de-
termined — monosaccharides, disaccharides, and polysaccharides. Chlorophyll
was determined colorimetrically, in an alcohol extract. Intensity of photo-
synthesis was determined by the S. V. Tageeva method. Leaves for analyses
were taken from the middle layer of the crown on the southwestern side.
During the summer, the observations were made in five periods at intervals
of 15 to 20 days, exactly at 9 and 11 A.M. and at 1 and 3 P.M. Oxidizability
of the cell sap was determined by the N. P. Krasinskiy method. As a result
of this investigation the following was found:
1. The development of plants was affected by conditions under which
they grew. In the proximity of the Ore Concentration Mill and the Metallurg-
ical Plant, the growth-development phases of box elder, oleaster and Siberian
pea shrub were accelerated.
2. Gas and dust pollution near the Concentration Mill and the Metallurg-
ical Plant intensified the growth of lateral shoots in the balsam poplar,
oleaster and pinnate elm.
3. In all the plants under study the chlorophyll declined toward the
end of the growing season. In the common lilac, oleaster, and pinnate elm
growing near the Concentration Mill or the Metallurgical plant the chloro-
phyll content rose.
4. The intensity of photosynthesis varied in the studied plants
ontogenically and in the course of a day.
The highest indices of photosynthetic intensity for most of the studied
plants were recorded during the morning hours. Appreciable gas and smoke
pollution hampered photosynthesis. However, the same conditions enhanced
photosynthesis in the oleaster and in the pinnate elm. This can be accepted
as manifestation of adaptability of these species to existing conditions.
5. From the point of view of oxidizability of cell sap, the studied
plants can be ranged in decreasing order of their gas resistance: pinnate elm,
oleaster, balsam poplar, box elder, Tartarian honeysuckle, Siberian pea shrub,
common lilac, European white birch, and Scotch pine. Of the studied plants
the most promising for green plantings on industrial sites in the Karaganda
coal basin are: pinnate elm, oleaster, and balsam poplar. Box elder, Tartar-
ian honeysuckle, Siberian pea shrub, and common lilac are of moderate resist-
ance. The European white birch and Scotch pine are low-resistant species.
- 69 -
-------
INDICATORS OF GAS RESISTANCE OF ARBOREAL PLANTS
(According to Investigations Conducted in The City of Krasnoural'sk)
V. N. Nikolaevskiy
Institute of Plant and Animal Ecology, Urals Branch of the USSR Academy of Sciences
From Akad. Nauk SSSR. Ural. Filial. Trudy Institute Biologii. Vip. 31,
(Sverdlovsk, 1963) p. 59-79.
In our country the first studies on the effect of poisonous gases upon
plants are attributed to the beginning of the twentieth century (Nelyubov,
1900-1910; Sabashnikov, 1911). They were interrupted during the period of
World War I and the [Russian] Civil War, but were resumed during the years
of national industrialization. A great interest in selecting gas-resistant
plants is developing at the present time in connection with the great spread
of industrial and residential construction specified by the Seven-Year Plan
for the growth of the Soviet national economy.
Visual, subjective methods of evaluating the degree of gas resistance
in plants are characteristic of the majority of the projects carried on up
to the present time. The resistance of plants has been determined by the
percentage of damage to the leaf surface (Krasinskiy, 1950) or by the
ability of transplanted trees to acclimatize (Bulgakov, 1958-1961). The
assortments of gas-resistant plants compiled on the basis of the above
method of approach are not universal: there are considerable contradictions
among them, and their employment under different geographical and soil-
climate conditions often yields negative results.
N. P. Krasinskiy (1937, 19AO, 1950) has conducted major studies on gas
resistance. Proceeding from the fundamental biochemical lav of S. L. Ivanov
(1926) and the conclusion drawn by A. V. Blagoveshchenskiy (1950) on the
biochemical bases of the evolutionary process in plants, Krasinskiy holds
that the gas resistance of plants is connected with the systematic position
of a species [This probably means the relative classification of a species
within a morphological or taxonomic system—Trans.]. He substantiates this
connection with data on the damage to plants by gases and the amount of
oxidizable substances in the leaves of 18 families of plants. N. P. Krasinskiy
(1950) developed methods for studying the gas resistance of plants.
The essential shortcoming in several previous studies (lonln, Koltasheva,
1961) is the fact that the gas resistance of plants was studied without tak-
ing into account the physiological and biochemical processes that take place
in leaves under the effects of gases and without regard for the influence of
the external environment upon the condition of the plants and their resistance
to gases. The compilation of the assortments of gas-resistant plants on the
basis of leaf damage and the conclusions of Krasinskiy (1950) does not resolve
the questions of the dependency of gas resistance upon the external environ-
mental conditions and upon the condition and the stage of development of the
plants. Therefore, they do not answer the practical needs for increasing the
gas resistance of plants by selection, raising, maintenance, fertilization, etc.
A certain exception Is presented by the work of V. A. Guseva (1950),
which showed the possibility of Increasing the gas resistance of plants by
using fertilizers, and by the work of M. V. Bulgakov (1961).
- 70 -
-------
Other studies by N. P. Krasinskiy (1950) and V. A. Guseva (1950)
on the reducing strength, the content of glutathione and ascorbic acid, and
the oxidation-reduction potential of plants these being of considerable
interest for understanding the gas resistance of plants were in the
nature of preliminary surveys and did not attain any further development.
The gas resistance of arboreal species, in our opinion, can be found
to be dependent upon a great number of factors. Among these factors we
can note the following: the special anatomical characteristics of the struc-
ture of the leaves and the activity of the stomas, the physiological properties
of the cell protoplasm (the buffer capacity of the cell contents, the permea-
bility and viscosity of the protoplasm), the quality and stability of enzymes
and proteins, the intensity and biochemical direction of the physiological
processes in the leaves, the water regime, etc.
In the first step of our studies in a short three-year period it was,
of course, impossible to encompass all the points in one study; therefore,
we had to limit ourselves to the study of a few of the physiological processes:
photosynthesis, respiration, transpiration, the activity of the stomas, the
quantity of easily oxidizable substances, the water holding capacity of
the leaves, the pH of the cell contents and the concentration of dry matter in
the leaves. We also studied the special anatomical features of the structure
of the leaves.
The results of the first year's work led us to a conclusion concerning
the gas resistance of arboreal plants as a function of the intensity of photo-
synthesis and concerning the existence of a certain correlation between the
gas resistance and the winterhardiness of plants (Sergeev, 1953 and 1961;
Nikolaevskiy, in press).
The intensity of photosynthesis was studied by the method of L. A. Ivanov
and N. L. Kossovich (1946); stoma activity was studied by the method of infil-
tration (alcohol, benzene, xylene); and transpiration was studied by rapidly
weighing on torsion weights. The concentration of dry matter was determined
by the refractometric method in the juice squeezed out by a hand press from
leaves killed by steam. The amount of substances oxidizable by a decinormal
solution of potassium permanganate was accounted for by N. P. Krasinskiy's
method (1950). The pH of the cell contents was studied with an LP-5 potentio-
meter. The determination was made by small glass and calomel electrodes in the
aqueous squeezings from the leaves (2-3 g. of leaves were crushed into 10 ml.
of distilled water).
A few of the methods we used are not distinguished by great accuracy.
This applies in particular to the determination of transpiration (Sveshnikova,
1959) and pH. However, even though they do not have a high absolute accuracy,
these methods were the only possible ones for us under field conditions,
because of their lesser complexity and laboriousness, taking into account the
great number of other tasks going on at the same time. Besides, we hoped to
obtain at least some comparative data of various plant species growing under
identical conditions, without claiming an absolute precision of the data.
- 71 -
-------
All determinations were run three times per day: at 7 A.M., 1 P.M. and
7 P.M. with four repetitions, but transpiration with eight to ten repetitions.
In order to ascertain the effect of meteorological conditions upon the
physiological processes, we considered the atmospheric humidity (by an August
psychrometer), the daylight exposure (by a luxmeter), and measured air tempera-
ture at the top of the tree.
Simultaneously we made use of the data of the meteorological station at
the Krasnoural'sk municipal park administration, submitted by M. V. Bulgakov.
The subjects of the study were trees selected in a public garden located
200 m. from the gas sources and, therefore, subject to their effects to a
great degree. The trees were 12 to 15 years old. All the studies were con-
ducted during the summer on five arboreal species: aspen [Populus tremulal.
balsam poplar [Populus balsamifera]. box elder [Acer negundo]. Siberian crab
apple [Pyrus baccata], and European white birch [Betula verrucosa]. The leaves
used for testing were taken during the summer from the same trees from
their south side in the middle of the crown.
The control trees used were aspens and European white birches of natural
origin in the Krasnoural'sk Forest, located 7 km. southeast of the factory.
In addition to these, balsam poplar, Siberian crab apple, and white birch
were planted in the same place in the autumn of 1959. Therefore, the results
observed on these transplanted trees cannot be considered as controls, but
they characterize the condition of arboreal species after transplanting.
The control trees were located in a large forest glade and were not sub-
ject to any specific effect of forest conditions.
Meteorological Conditions During the Growing Period
The meteorological conditions during the growing period are not very
favorable for the normal growth and development of plants. Spring in Kras-
noural'sk is usually lingering and cold, which is in general characteristic
of the Central Urals. Late spring frosts occur even up to the middle of June.
Snow falls on the budding leaves at the end of May or the beginning of June
almost every year.
Growth of arboreal species begins almost one week later than in Sverdlovsk,
i.e., at the end of the first half of May. Late frosts and snow bring about
no less damage to the leaves of the arboreal species than do gases. In two or
three days after a snowfall, there appear on the leaves blotches reminiscent
in color and form of actual gas burn.
In 1961 during a snowfall on the fifth of June, we were able to conduct
a series of experiments on the effects of fallen snow on the leaves of arboreal
species. These data allowed us to notice the sharp decrease in the intensity
of the physiological processes in leaves as expressed by the decrease in con-
centration of cell sap and the shifting of pH to the alkaline side. In the
arboreal species more resistant to late spring frosts, the decrease of these
Indices occurs at a lesser degree than in the nonresistant species.
- 72 -
-------
Early autumn frosts In Krasnoural'sk occur In the second half of August,
which, Just as do acid gases, leads to earlier losses of decorative quality,
to the cessation of growth, and to the deterioration In the condition of the
urban verdant plantings (Table I).
Table 1
Meteorological data for the 1960 growing period In Krasnoural'sk*
Month
May
June
July
August
First half of
September
Total
Number of days
Clear
2
1
2
5
1
11
Partly
cloudy
18
20
1A
19
8
79
With
precipi-
tation
11
9
15
6
5
A6
Precipi-
tation,
mm.
30. A
A7.5
A2.7
16.8
16.0
153. A
Relative
humidity,
%
72
82
76
90
80
Average
temp. ,
°C
6.6
16. A
16.1
13.6
11.6
1A.2
Maximum
wind
speed
m. /sec.
8
10.5
7
5
— —
The growing period was characterized by relatively low maximum tempera-
tures and sufficient precipitation. But in July and August of certain years,
there occur dry periods, and the temperature of air in the soil may reach,
according to our observations, A5 to A8°C. (26-27 July 1960).
The number of cloudy and rainy days comprises 34% of the total number
of days of the growing period. In the remaining 2/3 of the period, increased
sunlight is observed (clear or partly cloudy days), i.e., such weather con-
ditions that are conducive to maximum gas damage when gas pollution occurs;
as indicated by L. A. Ivanov and also by V. Kroker (1950).
According to the data of the meteorological station of the Krasnoural'sk
Municipal Park Administration, the number of days with gas pollution during
1960 was found to be as follows:
June 1 day
July 7 days
August 6 days
September (1/2 mo.)....! day
The maximum concentration of harmful gases in the vicinity of the indus-
trial plants in Krasnoural'sk is very high. According to data by L. V. Titnofeeva
(1958), the concentration of harmful acid gases (S02, F) in the vicinity of
copper smelting concerns is 8 to 10 times higher than the permissible standards
set by sanitary regulations. In addition, the microcllmatic conditions for
arboreal species deteriorate significantly in the cities and in industrial
centers. The air temperature in July-August at the surface of cobblestone
- 73 -
-------
pavement is 4 to 16eC higher, and 8 to 23° C higher on asphalt pavement, than
at the surface of lawns and the tops of arboreal species (Lunts, 1952).
Taking into account the particularly severe microclimatic conditions
for the verdant plantings in the city — namely, the effect of asphalt sur-
facing of sidewalks and streets upon the temperature of the air and the great
amount of dust and of gas pollution — it becomes evident that the environ-
mental conditions are extremely unfavorable for the growth of urban plantings.
This is also reflected in the gas resistance of arboreal species.
Results of the Studies
It must be taken into account that with some species we could not adhere
strictly to the methods we used at the beginning of the work. The deviations
lie in the fact that the birch in the public park renewed its leaves at the
beginning of July, and the poplar, at the end of July; this was reflected in
all the physiological indices. In the birch the changes began from the
twentieth of July, and in the poplar, after 2 August. In addition, we could
not maintain uniformity of the environmental conditions for the crab apple.
Thus, in the middle of the day the crown of the crab apple was shaded by the
poplars standing close by, which could have influenced, to a certain degree,
the indices obtained and the susceptibility of the leaves of a given tree to
gas damage.
Before we proceed to the analysis of the processes studied, it Is neces-
sary to evaluate the gas resistance of each of the arboreal species and the
change in resistance during the year (Figure 1).
June
July
August September
During the summer a regular Increase of leaf damage by harmful gases
was observed. This1 is connected with the distribution of days in the month
during which there was the effect of gas on the verdant plantings of the city.
- 74 -
-------
In accordance with the scale proposed by N. P. Krasinskiy (1950) and
on the basis of the means for the growing period (Table 2), the degree of
susceptibility to damage of the arboreal species is classified in the
following manner:
Box elder very low (less than 5%)
Balsam poplar low to medium (10-20%)
Aspen medium (20-30%)
White birch medium-to-high (30-40%)
Siberian crab apple low-to-mediurn (10-20%)
In the Siberian crab apple trees growing in open areas, the degree of
leaf damage was close to that indicated for the birch. This suggests that
the two species have similar low resistance to gases.
The intensity of damage caused by gases depends, when other conditions
are equal, upon the concentration of the gases in the atmosphere and the
duration of exposure. Therefore, on the basis of data that we obtained on
the susceptibility to damage of the leaves of arboreal species as well as on
the basis of data found in literature (Krasinskiy, 1950; Vanifatov, 1959;
Knyazeva, 1950), it can be briefly summarized that the arboreal species
studied have been classified as (a) lightly-damaged (box elder), (b) medium-
damaged (poplar, aspen), and (c) heavily-damaged (crab apple, birch).
We shall begin the characterization of the physiological conditions of
the arboreal species studied with the description of the way the stomas
function. The dynamics of their activity during the day were studied on
the basis of the penetration of alcohol, benzene, and xylene Into the inter-
cellular channels of the leaf. In order to facilitate comparison, we substi-
tuted arbitrarily numerical symbols for the designations of the infiltrating
liquids; at the same time, these symbols represent the relative sizes of
the stoma apertures. Since xylene (1) can penetrate even into stomata that
are almost closed, the number 1 designates the minimum sizes of the stoma
apertures. Benzene can penetrate half-open stomata (2) and alcohol penetrates
completely open stomas (3).
This method subsequently facilitates the treatment of materials and
the comparison of the operation of the leaf stoma apparatus both during a
day and during the entire year.
Only xylene can penetrate the intercellular channels of leaves of
arboreal species when the stoma apertures are only slightly open (1 and less).
The amount of liquids which can penetrate into the intercellular channels of
the leaves increases in proportion to the stoma aperture; all three chemical'
compounds penetrate fully open stomata.
In studying stoma activity, we evaulated the degree of penetration of
alcohol, benzene, and xylene into the intercellular channels of a leaf as
good, medium, and poor. Poor penetration of benzene simultaneously implies
good penetration of xylene.
- 75 -
-------
Table 2: Bean monthly indices of the intensity of physiological processes in the
species in the forest and in the public garden.
Species
Actual photo-
synthesis,
06. C02
per dn^/hr.
Respira-
tion,
ng. COg
per
da^/hr.
Transpir-
ation, g.
per
dnrVhr.
Water
content
of the
leaves,
Aoount of
substances
oxidizable
by KUnOjt
per g.
green wt.
caused
by
gases,
Actual photo-
synthesis,
OB. C02
per dn^/hr.
Respira-
tion,
ng. C02
per
dn^/hr.
Transpir-
ation, g.
per
dn^/hr.
Water
content
of the
leaves,
Amount of
substances
oxidizable
by KJtnOit
per g.
green *t.
Public Park
Box elder
Poplar
Birch
Crab apple
Box elder
Poplar
Crab apple
Box elder
plar
'rab apple
@px elder
plar
Aspen
Birch
Crab i
apple
••June
1C
July
August
No data
September
0.7
1.2
-------
Table 2: (Continued)
Species Actual photo- Respira- Transpir- Water Amount of Damage Actual photo- Respira- Transpir- Water Amount of
synthesis, tion, ation, g. content substances caused synthesis, tion, at ion, g. content substances
iag. CC»2 ng. C02 per of the oxidizable by ng. C02 ng. COj per of the oxidizable
per dm^/hr. per dn^/hr. leaves, by KHnO^ gases, per dm?/hr. per dm^/hr. drn^/hr. leaves, by KHnC>4
dn^/hr. % per g. % % per g.
green wt. green wt.
Average data for the growing period
Box elder
Poplar
Birch
Crab apple
6«1
J:l
B'.5
1 a
11
5 6
1 190
Hi
6948
74. «
1:!
62:$
21.6
1:8
49:8
t.Q
g:?
15:0
6 2
\\
7 4
l.p
i:i
A» f
0 737
1^
° **
75. 5
1:1
61:2
21.2
^-1
5^:5
Clear Cloudy
Forest June
Poplar
Native birch
Transplanted
birch
Crab apple
Poplar
Native birch
Transplanted
Crab apple
AlpeSr
Native birch
Transplanted
birch
Crab apple
Poplar
Aspen
Native birch
Transplanted
Crabapple
14.7
4.6
9.0
1
8.8
fcl
12.5
9.8
9.2
1
h 1
I'l
1.6
1:|
1.4
1.8
I'l
2.5
II
0.42
II
0.67
8:8
0.254
0.59
0.492
If
68.9
11
65.4
ffif
59 .9
61.1
H
32.3
II
46.8
|:t
30.8
62.1
—
—
July
—
—
September
—
—
—
M
9.5
ILL
4.0
—
—
No Data
—
—
M
34.0
ii:i
11.6
5.6
Average data for the growing period
62.8
65.1
47.2
—
7.5
1*
6:9
1.2
?'i
x!
0:9
2.4
11
0.275
0.328
0.326
11
65.7
60.6
60.6
|:J
29.5
48.4
41.2
-------
Hence, the degree of stoma aperture Is represented by the following
arbitrary symbols:
Poor penetration
Medium penetration
Good penetration
Xylene
0.0
0.5
1.0
Benzene
1.0
1.5
2.0
Alcohol
2.0
2.5
3.0
We shall analyze the activity of the stomata in the leaves of the
arboreal species under conditions of periodic emmissions of gases
(Figure 2a) and without them (Figure 2b). The mean monthly data are
derived from 5- to 7-day observations.
It is evident from Figure 2 that the average dimensions of stoma
aperture in the poplar, crab apple, and aspen in the public park are
greater than in the box elder. The degree of stoma aperture in the birch
is less than half of the maximum possible dimensions. This means that
there is a considerable regulation of stoma movements and this is explained
as being the result of the deterioration during recent years of the tree's
condition under the effect of gases. The deterioration can be seen even
from the external appearance of the crown.
2 . -~ 5
June
July September June July September
Pig. 2t Change in the degree of average stoma aperture during the growing
period.
(a) Public park| (b) forest (arbitrary numbers are the sane as in Figure l).
In the aspen and birch growing naturally in the forest, the daily average
degree of stoma aperture is greater than that of the trees in the public park.
The average stoma aperture dimensions for the entire growing period are smaller
In the crab apple and poplar trees transplanted to the forest in 1959 than they
are in the trees in the public park. In the transplanted birch, however, the
stoma aperture in the forest and in the park are of similar dimensions. This
can be explained (1) by the effect of transplanting upon stoma activity in the
trees transplanted to the forest and (2) by the deterioration of the condition
of the tree in the case of the birch in the public park.
- 78 -
-------
In the public park the regulation of stoma activity is most noticeable
in the box elder and the birch while in the forest it is most noticeable
in the transplanted poplar, crab apple, and birch trees.
Increases in the degree of stoma aperture in the birch exposed to the
effect of gases can be observed in July (Figure 2). This is explained by
the replacement of its leaves. If there were no replacement of the leaves,
the curve indicating the change in the degree of stoma aperture in the
birch evidently would be parallel on the graph to the one for the box elder.
The same behavior of the stoma apparatus in the arboreal species differing
in their susceptibility to injury can be explained by the fact that the
decreasing sizes of stoma aperture in the box elder is probably a character-
istic of the species, and in the birch, it is the result of considerable
damage caused by the poisonous gases and by the continuous yearly deterior-
ation of the tree. This is also substantiated by the performance of the
stoma of the birch trees not exposed to gas emissions during the summer.
The study of stoma activity during the summer is conducive to some
clarification of the reasons for the different susceptibility to leaf dam-
age of the studied species. In a number of papers (Ivanov, 1936; Wislicenius,
1914; Jahnel, 1954) there are references to a possible connection between
stoma activity and the degree of susceptibility to leaf damage in different
species.
The intensity of the damage to leaves by gases must depend upon the
amount of absorbed acid gas per unit of dry or green weight of the leaves;
whereas the rate of gas absorption depends upon the number and sizes of stoma
apertures per unit of leaf surface and upon the presence of the intercellular
channels. The reduction of stoma aperture by one-half, according to Stefan's
law for stomata, affects hardly, if at all, the rate of gas exchange between
the air and the intercellular channels of the leaf (Maxlmov, 1958). Further
narrowing of stoma apertures causes a regular but nonproportional reduction
in the gas exchange rate. This means that with the variation of size In the
stoma apertures during the day within a range of from 3 to 2 (in our arbitrary
symbols) we cannot speak of differences in the gas exchange rate. The reduc-
tion of the stoma apertures from 2 to 1 and less will cause decreases in the
rate of gas exchange.
Considering the foregoing statements, we cannot speak of essential
differences in the gas exchange rate In the poplar, aspen, and crab apple
during the summer, but in the case of box elder one can observe a substantial
decrease in Its gas exchange rate when comparing it with that of the other
trees (see Figure 2). It can be assumed that the reduction of the gas exchange
rate in the box elder under the effect of gases is conducive to a decrease in
absorption of the harmful gases and to an increase in Its resistance.
In the young birch leaves subject to periodic exposure to gas emissions,
the average size of the stoma apertures is half of their possible size
(July-2, Figure 2a). This, according to Stefan's law, does not restrict the
rate of gas exchange, is conducive to a significant absorption of sulfur
dioxide and leads to considerable leaf damage. During the fall in these same
birch leaves, a substantial decrease in the stoma apertures is observed when
compared either to the summer stoma apertures data or to the stoma data of the
- 79 -
-------
birch not exposed to gas emissions. The decrease in the gas exchange rate
during the fall observed in the birch under exposure to gas emissions (see
Figure la) is explained by the effect of the considerable gas damage to the
leaves.
A comparison of the function of the stoma apparatus in the leaves
(Figure 2a, 2b) shows that, at the end of the growing period a decrease in
the average size of stoma aperture is observed in all species in the public
park, while in the forest the reverse is true because the stoma aperture
increases. An exception is the July data for the birch and poplar. An
especially sharp increase in the degree of stoma aperture is observed towards
fall in the arboreal species transplanted to the forest. Such difference in
the behavior of the stoma apparatus between the leaves of the arboreal species
in the public park and in the forest evidently developed as a result of the
effect of the poisonous gases in the former and the absence of gases in the
latter case.
The intensity of the basic physiological processes (photosynthesis,
respiration) is, to a significant degree, connected with the functioning of
the stoma apparatus. The monthly indices of the Intensity of actual photo-
synthesis are based on averages derived from 3- to 4-day observations.
It can be seen from Figure 3 and Table 2 that the greatest intensity
of photosynthesis in the public park takes place during clear weather in
the birch, poplar, and crab apple, and the least intensity is found in the
box elder. If we consider the intensity of photosynthesis in July (when we
observed the greatest number of days with the presence of gas and one
could have expected the presence or absence of the connection between photo-
synthesis and the susceptibility to leaf damage due to gases) then it is
clear that according to their intensity of photosynthesis the species can be
classified from highest to lowest as follows: birch, crab apple, aspen, pop-
lar, and box elder. It can be seen from Table 2 that all the species, except
the crab apple, have the same sequence of susceptibility to gas damage in
July.
Consequently, the species characterized by a higher intensity of photo-
synthesis are damaged to a greater degree by gases. This substantiates our
preliminary conclusion concerning the dependency between the intensity of
photosynthesis and gas resistance.
In cloudy weather (Figure 3c), because of the different light require-
ments of arboreal species, there is no such close connection, but it is evident
here as well that in July the highest intensity of photosynthesis is observed
in birch and crab apple while the lowest is observed in poplar and box elder.
It can be seen from Figure 3b that birch and crab apple in the forest
hold first place in their intensity of photosynthesis. Consequently, the
indices of yearly average intensity of the photosynthesis of th'e trees grow-
ing in the public park and in the forest are quite similar. The photosyn-
thesis indices obtained for birch in the forest were high because of the
younger calendar age of the leaves.
- 80 -
-------
W
2L
^
June July Sept* >
Juno
July sept.
in
•8
'*
/fl
6
6
fNot included on original
copy — Trans.'!
N/7 \
June July August Sept.
Fig. 3: Change in the intensity of photosynthesis during summer.
(a) Public pork, clear; (b) forest, clear: (c) public park, cloudy: (d) forest, cloudy
CGrepn missing — tronS.j . (Arbitrary symbols are thS sate as in Figure 1.)
Some scientists (Ivanov, 1936; Kroker, 1950; Wlslicenius, 1914) assume
that the degree of the damage in plant leaves caused by acid gases is depend-
ent upon the intensity of photosynthesis. They note that under conditions
that are conducive to the increase in the intensity of photosynthesis, the
plants are more damaged by acid gases. The authors point out that a prelim-
inary storage of plants in the dark for two or three hours before exposure to
gas emissions contributes to the reduction of susceptibility to leaf damage
because of gases.
Wieler (1905) indicated a greater susceptibility to injury in the case of
the coniferous needles of a two-year-old pine than of those of a one-year-old
pine. The reason for this effect he considered to be the increased intensity
of photosynthesis. This is confirmed by the data of A. V. Savina (1941),
- 81 -
-------
whose data indicate that the intensity of photosynthesis in the needles of
a two-year-old pine is 1.5 to 2.5 times higher than that of the one-year-
old pine.
According to current concepts (Kretovich, 1961; Kolesnikov, 1959),
biochemical processes of plants during photosynthesis and during respira-
tion are closely interrelated. Therefore, the study of respiration in
the arboreal species concurrently with the study of photosynthesis is of
considerable interest.
2 3
Fig. 4: Change.!
. longe.in
intensity of
respiration
during the
summer.
S. i.)
June
July
Sept.
June
July Sept.
The data on intensity of respiration shown in Figure 4a and Ab indicate
that with arboreal species we find the same interdependence between respira-
tion and gas resistance as obtains between photosynthesis and gas resistance.
During clear weather the average intensity of respiration for the growing
period of all arboreal species studied, with the exception of crab apple, is
analogous in sequence to that for photosynthesis. This can be seen from the
following figures: box elder 1.8 mg., poplar and aspen 2.3 mg. , birch 2.7 mg. ,
and crab apple 2.0 mg. per dm^ per hour. The arboreal species have also the
same sequence for susceptibility to gas leaf damage.
It has been observed that the minimal intensity of respiration for all
arboreal species is during the middle of the summer. Towards the fall the
intensity of respiration continues to decline or increases slightly. The
data shows that birch growing in the forest (natural and transplanted,
Figure 4b) as well as in the public park constitutes an exception, seemingly
because of the differences in the calendar age of the leaves.
Respiration, of course, goes on during photosynthesis and its relative
share of participation in actual photosynthesis, expressed in percentage has
been found to equal 29.5% for the only slightly susceptible to gas damage
box elder and 21 to 23% for the greatly susceptible to gas damage species of
birch and crab apple.
- 82 -
-------
The position developed by several investigators (Ivanov, 1946;
Blagoveshchenskiy, 1950) concerning the relationship of the physiological
and biochemical processes as well as their intensities to the systematic
classification of plant species makes it possible to understand the inter-
dependence between intensity of photosynthesis and of respiration, and of
gas resistance of plants. Actually, if the magnitude of gas damage depends
upon the extent and intensity of photo-oxidizing processes in the leaves
when photosynthesis is stopped by the effect of acid gases (Noack, 1920;
Krasinskiy, 1950; Jahnel, 1954), then the intensity of photo-oxidation must
depend upon the photosynthetic and the enzymatic activity of the protoplasmic
structures, which we are able to judge to a certain degree by the intensity
of photosynthesis and respiration.
Consequently, on the basis of the indices of photosynthesis and leaf
respiration in arboreal species, even without exposure to gas emissions,
we can assume possible injury to the leaves of arboreal plants in the case
of periodic effects of gas emissions.
Thus, the arboreal species that are only slightly susceptible to damage
by acid gases (box elder) are characterized by a low intensity, and those
greatly susceptible (birch, crab apple), by a high intensity of photosyn-
thesis and respiration during their growing period.
As shown by L. A. Sergeev (1961) , the species having a reduced intensity
of photosynthesis and respiration are more winterhardy. Here, in our opinion,
is indicated some analogy between hardiness of plants and their resistance
to acid gases.
The results of studies of transpiration in arboreal species suggest
that transpiration is the least reliable or stable index. The instability
of transpiration is explained (Sveshnikova, 1959) by its essential dependency
upon the meteorological conditions. In the poplar, birch and crab apple
growing in the public park, the data obtained in clear weather are close to
the average per year intensity of transpiration, and the highest intensity
of transpiration was found in the box elder (Figure 5a and Table 2). During
cloudy weather, the intensity of transpiration decreases in all species.
In the three species mentioned above, the indices of transpiration are rather
close (Figure 5c). In the case of box elder, when compared with other species
in the public park, a more significant decrease of the transpirational water
consumption was found during the cloudy weather.
The intensity of transpiration of trees in the forest (Figure 5b) is
smaller in comparison with the same species in the public park, the only
exception being aspen. The decreasing intensity of transpiration in the
transplanted species in the forest is the result of the transplantation.
The determination of leaf water content was made by drying the leaves to
a constant weight at temperatures of 100 to 102°C. Leaves that were taken
during the summer for the study of transpiration were utilized for the water
content determination.
- 83 -
-------
It can be seen from Figure 6a that the crab apple and the aspen have
lower leaf water contents during the summer than do the other species,
while the box elder leaves have the highest water content. In the case of
birch in the public park, the increase of the total moisture in July can
be explained by a change of its leaves [renewal of leaves: translator's note].
If the change of leaves is not to be considered, then the average leaf water
content in the birch is lower than in the poplar, and this is also verified
by the change of the leaf water content in the natural birch in the forest.
Consequently, the box elder, which is only slightly susceptible to
damage by gases, is characterized by an increased total water content of
its leaves.
In the natural birch growing in the forest, the leaf water content is
close to that of the birch planted in the public park; in the aspen, the
leaf water content is higher. In trees newly planted in the forest, with
the exception of birch, the leaf water content is higher than in the public
park trees. This fact is explained by the effects of transplantation
(Figure 6b).
With all arboreal species the total water content in the leaves decreases
as autumn approaches, but in the box elder this decrease is less pronounced
than in other species.
The susceptibility of the leaves to damage by gases in the species
studied is well correlated with the total water content of the leaves. The
box elder, which is resistant to the gases, is characterized by a high water con-
tent , while the medium and the highly susceptible to gas damage species
(poplar, aspen, birch, and crab apple), are characterized by a low leaf
water content.
Concurrently with the study of photosynthesis, respiration, and trans-
piration that take place during the day determinations were also made of
the amount of oxidizable substances in the leaves (Figure 7). The quantita-
tive study of the easily oxidizable substances In the leaves was made in
order to verify some of N. P. Krasinskiy's (1950) conclusions and also in
order to study the relationship between the dally course of photosynthesis
and the quantitative change of oxidizable substances in the leaves. To make
sure that the changes in the leaf water content during the day do not alter
the indices calculated per kg. of the green weight, the leaves from the
public park were taken in distilled water to a chemical laboratory at the
factory. Prior to taking samples for weighing and pulverization, leaves
were carefully squeezed out between sheets of filter paper.
The quantity of oxidizable substances in the plant leaves (as shown in
Figure la, 7b, and 7c) characterizes the resistance of plants to gases. In
the box elder, which is only slightly susceptible to damage by gases, the
amount of oxidizable substances is 1.5 to 2 times lower than in the birch
and the c-ab apple. During summer, the amount of oxidizable substances in
the leaves increases, but this increase varies with different species, as can
be seen from Figure 7a. Towards autumn the increase of oxidizable substan-
ces in the box elder leaves is less intensive than in the poorly resistant
-------
species, i.e., in the crab apple and the birch. Their content is lower in
the leaves of the forest trees than in the leaves of the trees growing in
the public park.
t,8
',*
0,8
(.
S.
f •• \V
^' -^
June
July
Sept,
Jane
July
Sept.
+>
.5
a.
in
s
*/,*
<,2
lo-
ot
0,6
0,*
0,7
June July August Sept.
Fig. 5» Transpiration of the arboreal species during summer.
(a) Public park, clear; (b) forest, clear; (c) public park, cloudy.
(Arbitrary numbers are the same as in Figure 1.)
N. P. Krasinskiy's (1950) identification of the quantitative indices of
the easily oxidizable substances in the leaves with the character and degrees
of photo-oxidations arising in living plant cells under the effects of acid
gases has raised serious objections among physiologists.
Of course, the chemical processes of the oxidation of dead plant substrate
can not be identified with the biochemical processes in living plant cells
because, first, we deal here with a completely different condition of matter
and, second, upon the acidification of the substrate to pH 1 or lower, the
picture is deliberately distorted; the latter is because of the fact that the
- 85 -
-------
amount of oxldizable substances increases as a result of the oxidation of
difficultly oxidizable or nonoxidizable substances which are found under
normal conditions in plant cells.
Proceeding from the fundamental biochemical law of S. L. Ivanov
(1926), we cannot be sure that with a substantial acidification of plant
substratum there will develop with different plant species an identical
and monotypical increase in the amount of oxidizable substances. The basis
of the method for determining the amount of oxidizable substances, as
recommended by N. P. Kraslnskiy (1950), ignores these important aspects.
\ -S
June
July Sept.
June
July
August Sept,
Fig. 6: Change of leaf water content during the summer.
(a) Public park, clear; (b) forest, clear; (c) public park, cloudy.
(The arbitrary numbers are the same as in Figure 1.)
Apparently, he assumed that upon acidification of the cellular substrate,
there will be a similar increase in the oxidizable substances in different
plant species and, as a result of this similar background of the assumed
increase of oxidizable substances, the differences in the content of
oxidizable substances in the leaves of the different species will be retained.
- 86 -
-------
In view of the above observations, it appears to be necessary to be
cautious in the use of the data obtained by this method.
70
60
50
10
• JO
3 20
d
5
•
50
JO
20
10
Fi
a.
June
5
July Sept.
^<3
June
July Sept.
June
(a)
(The
July August Sept.
7< Change in the amount of leaf substances oxidized by a 0.IN solution
InO^ per g. of green weight. ~
Public park, clear; (b) forest, clear; (c) public park, cloudy.
arbitrary numbers are the same as in Figure 1.)
To ascertain the reliability of the data obtained, we determined the
amount of oxidlzable substances in the poplar and the crab apple, conduct-
Ing tests with ten repetitions. Statistical analysis of these data makes
it possible to note that the average error of the arithmetical mean is
- 87 -
-------
+1.5 ml., and the accuracy of the experiment Is within the limits of +1.3
to 2.1% (crab apple - +1.3%; poplar = ±2.1%).
Consequently, the data obtained by this method are fairly reliable and
possess sufficient accuracy for the purpose of our investigations.
It is self-evident that with only 4 repetitions (Figure 7 and Table 2 ) the
accuracy will be lower. To avoid random errors in the data analyzed for each
day of the experiment we grouped the data obtained on a monthly basis and
derived the average values. Thus, the average data in Table 3 are derived
from 5- to 7-day determinations.
Table 3 : Fluctuations of the amount of oxidizable substances in the
leaves during the day - for the growing jperiod of 1960
Species
Box elder
Balsam poplar
Aspen
White birch
Siberian crab apple
Box elder
Balsam poplar
Aspen
White birch
Siberian crab apple
Box elder
Balsam poplar
Aspen
White birch
Siberian crab apple
Box elder
Balsam poplar
Aspen
White birch
Siberian crab apple
Amount of oxidizable substances
per g. of green weight, mg. 0.1N. KMnO^
at
7
A.M.
at
1
P.M.
at
7
P.M.
Average
amount of
oxidizable
substances
Amplitude of
fluctuations in
the amount of
oxidizable
substances
per day
Fluctuations
%of daily
average
amount of
oxidizable
substances
Public Park, June
16.2
20.7
21.7
26.3
37.3
18.8
28.2
34. 3
24.2
47.7
18.2
22.5
24.3
30.6
37.5
17.3
28.6
33.8
19.7
43.7
19.1
24.1
23.4
28.4
41.7
17.7
22.4
23.1
28.4
39.2
2.9
3.4
2.6
4.3
4.4
16.4
15.4
11.3
15.1
11.2
Public Park, July
18.4
29.8
38.2
28.0
45.4
18.2
28.9
35.4
24.0
45.4
1.5
1.6
4.4
8.3
4.0
8.2
5.6
12.4
34.5
8.8
Public Park, September
29.1
43.5
43.2
48.6
50.6
26.8
37.8
47.6
53.3
69.9
25.6
42.2
42.4
44.5
60.4
27.2
41.2
44.5
48.8
60.3
3.5
5.7
5.2
8.8
19.5
12.9
13.8
11.7
18.0
32.4
Average Data for the Growing Period
—
—
—
—
— —
—
—
—
—
— —
—
—
—
—
— —
—
—
—
—
— —
2.64
3.57
4.06
7.15
9.13
12.4
11.6
11.8
22.6
17.5
- 88 -
-------
It is evident from the average monthly data in Table 3 that significant
fluctuations in the amount of oxidizable substances can be observed during
the day. The smallest fluctuations are observed in the box elder and the
greatest in the crab apple and the birch.
In the balsam poplar in July, a substantial decrease in the amplitude
of the fluctuation of the amount of oxidizable substances can be observed.
This can be explained by the effect of considerable gas damage and by the
decrease in the intensity of photosynthesis in July to 7.5 mg. C02 per
dm^/hour as against 11.9 mg. in June.
Parallel with the absolute figures of fluctuations of the amount of
oxidizable substances, the relative numbers of these fluctuations are given
in the table as a percentage in relation to the arithmetical mean.
The percentage of fluctuations in the amount of oxidizable substances
in the leaves indicates that these fluctuations are 2-10 times or more
greater than the experimental accuracy (2.1%). Therefore, the fluctuations
during the day in the amount of oxidizable substances in the leaves of
arboreal species are explained by the intensity of physiological processes:
photosynthesis, respiration, and other factors.
A comparison of the average monthly data for photosynthesis (Table 2)
with the fluctuations in the amount of oxidizable substances during the same
months leads to the conclusion that the box elder, which is more resistant
to gases, is characterized by a decreasing intensity of photosynthesis and
respiration. At the same time, it has the lowest amount of oxidizable sub-
stances in its leaves and the smallest amplitude of fluctuations of this
during the day. Birch and crab apple, which are not resistant to gases,
are characterized by the highest intensity of photosynthesis and respiration
and also by considerably larger amounts of oxidizable substances and a
greater amplitude of their fluctuation in the leaves during the day.
According to the average data for the growing period, the species most
resistant to gases are characterized by a smaller amplitude of fluctuations
of the amount of oxidizable matter and by smaller amounts of this matter.
The concentration of dry matter in the leaves (Table 4), which was
derived by the refractometric method, varies in the box elder between 9.3
and 12.7%, 14.3 - 21.4% in the birch, and 19.5 - 23% in the crab apple.
Therefore, the box elder, which is more resistant to harmful gases, is
characterized by a reduced concentration of the dry matter. In the nonre-
sistant species of birch and crab apple, the concentration of this matter
is 1.5 to 2 times greater than in the box elder. In the young birch leaves
(August 1 and 4) the concentration of dry matter is lower than in the old
leaves, which correlates well with their higher gas resistance. During the
summer the concentration of dry matter in the leaves increases.
In the forest trees, the concentration of dry matter in the leaves is
1 to 2.5% lower than in the trees of the public park.
The measurement of the pH of the cell contents (Table 4) indicated that
in all the public park species the effect of the 862 accumulation from the
- 89 -
-------
atmosphere causes quite significant decreases of pH. It is of interest to
note that in crab apple and birch, which are nonresistant to gases, the
decrease of pH in the public park in comparison with the forest is 0.85 -
0.97 while in the somewhat more gas-resistant poplar and aspen, the pH of
the cell contents in the public park trees is 0.15 - 0.20 less than in the
trees growing in the forest.
During the autumn a noticeable shift of pH to the alkaline side can be
noticed in the cell content of all species in the public park and in the
forest. This can be explained probably by the irreversible seasonal pro-
cesses in the aging of leaves of the arboreal plants. In the box elder the
lowest pH of the cell contents observed was 4.2 - A.65. The highest pH of
the cell contents is found in the aspen and the birch (6.85 - 6.A7).
Table 2 shows the monthly dynamics of the change in leaf damage. If
we compare these data with the monthly number of days in which gases from
the industrial plant were enveloping the city, then the time relationship
of the susceptibility of leaves to gas damage on these days is clearly
shown. In the poplar and aspen, the gas damage progressively increases,
and in the beginning of August the damaged poplar leaves are shed. The
damaged leaves of the aspen do not drop until fall, thereby affecting
adversely its ornamental aspects. The damaged box elder leaves turn yellow
rapidly, fold, up and fall off, but the leaves freshly damaged by the gases
are scalded or blighted over no more than 5% of their surface. In the birch
the increase in susceptibility to damage during the fall is explained by
the replacement of its leaves in the middle of summer.
In estimating leaf damage caused by gases in the individual species,
we could not help becoming aware of the shortcomings and inadequacy of the
method introduced by N. P. Krasinskiy (1950). This method suggests that
in the revised estimation of gas damage, only the percentage of damaged
area of the leaves is considered. In comparing the individual species
(birch and crab apple), it is often observed that even when leaf damage is
approximately the same in surface area it can be quite substantially different
with respect to the number of burned leaves expressed as percentage of all
the leaves in the crown. The determination of the number of damaged leaves
in percentage makes it possible to estimate more precisely the degree of
resistance of the individual species and to discover differences not noted
at first glance in the resistance of several species with fairly similar
susceptibilities to damage.
Conclusions
1. In the box elder, which is resistant to sulphur dioxide, the
smaller stoma apertures can be the reason for the decrease in the rate of
gas exchange and can thus contribute to the reduction of the susceptibility
of the leaf to damage by poisonous gases.
2. In all the arboreal species investigated (box elder, balsam poplar,
aspen, white birch and Siberian crab apple) under exposure to harmful gases
(S02, F), a reduction, in various degrees, of the stoma apertures is observed
as autumn approaches. With the absence of the gases, the reverse effect is
observed.
- 90 -
-------
Table 4: Concentration of dry matter in the leaves and pH of
the cell contents
Species
Concentration of dry
matter, %
at
7
A.M.
at
1
P.M.
at
7
P.M.
average
for the
day
pH of cell contents
at
7
A.M.
at
1
P.M.
at
7
P.M.
average
for the
day
June 27, Public Park
Box elder
Poplar
Aspen
Birch
Crab apple
Box elder
Poplar
Aspen
Birch
Crab apple
Box elder
Poplar
Aspen
Birch
Crab apple
Box elder
Poplar
Aspen
Birch
Crab app le
9.3
17.5
18.0
18.6
19.5
9.3
19.0
19.7
21.4
11.8
10.8
19.5
19.5
18.5
19.0
9.8
18.6
19.1
19.5
20.1
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
—
August 1, Public Park
11.8
15.8
20.5
14.3
21.5
11.7
18.4
18.8
14.7
21.1
11.8
20.0
18.4
16.1
20.2
12.7
17.8
19.1
16.5
22.3
13.7
17.2
17.4
14.9
21.1
12.4
17.1
18.9
14.6
21.2
—
—
—
—
—
—
—
—
—
—
_—
—
—
—
—
__
—
—
—
—
August 4, Public Park
11.8
20.0
19.8
14.3
19.8
12.1
19.2
19.1
15.6
21.1
4.8
5.8
6.4
5.2
5.2
3.9
5.2
5.9
5.3
5.2
3.9
5.8
6.2
6.0
5.8
4.2
5.6
6.2
5.5
5.4
September 4, Public Park
13.2
19.3
16.7
16.8
21.5
12.7
16.8
19.0
19.2
23.0
12.5
18.4
17.8
16.0
20.7
12.8
18.1
17.8
17.3
21.7
4.6
6.4
6.6
6.0
6.0
4.6
6.4
6.8
—
6.2
4.7
6.3
6.5
6.9
5.9
4.6
6.3
6.6
6.9
6.0
3. The amount of easily oxidizable substances in the leaves of
arboreal species has been found to be a function of their resistance to
acid gases. In the more resistant species (box elder), the amount of
these substances is lower than in the heavily susceptible to damage species
(crab apple, birch).
4. The amount of easily oxidizable substances in the leaves of
arboreal species increases 1.5 to 2 times as autumn approaches, which cor-
relates with the increased susceptibility of the leaves to damage by gases.
5. In arboreal species the magnitude of the fluctuations during the
- 91 -
-------
day in the amount of easily oxidizable substances upon exposure to gas
emissions is related to the intensity of photosynthesis and respiration.
In the more resistant species, small fluctuations (2 to A ml.) in the
amount of easily oxidizable substances are characteristic; in the less
resistant species, these fluctuations are greater (8.5 to 19.5 ml.).
6. The resistance to acid gases of the arboreal species studied
depends upon the intensity of photosynthesis and respiration. In the box
elder, which is more resistant, a lower intensity of photosynthesis and
respiration is observed.
7. The gas resistance of the species studied appears to be correlated
w;Lth and depended upon the total water content in the leaves. For arboreal
species resistant to gases, a higher water content of the leaves is
characte ri s t i c.
8. From among the arboreal species studied the species box elder,
which is resistant to gases, has been found to have during the summer the
lowest concentration of dry matter in the sap of the leaves killed by steam.
9. Under the effect of acid gases, the pH of the cell contents of
the leaves decreases. The decrease of pH is a function of the gas resist-
ance of the species: a larger decrease of pH occurs in the less resistant
species than in the more resistant.
10. Under the effect of the gases, an accelerated aging of the leaves
occurs; this causes an earlier termination of growth in the species under
periodic exposures to gas emissions in comparison with those trees not
affected by the gases.
11. The study of the physiological processes in the leaves showed that
these processes enable us to judge the degree of resistance of arboreal
species to gases. Moreover, the physiological studies are conducive to
clarification of the reasons for the variations in susceptibility to leaf
damage in arboreal species.
- 92 -
-------
Literature Cited
Blagoveshchenskiy, A. V. Biokhimicheskie osnovy evoliutsionnogo proteessa u
rasteniy, M., Izd-vo AN SSSR, 1950.
i
Bulgakov, M. V. Opyt ozeleneniya g. Krasnoural'ska. Mat-ly po ozeleneniyu
gorodov Urala, vyp. 1, Sverdlovsk, 1958.
Bulgakov, M. V. Vyrashchivanie berezy na pochvakh, otravlennykh otkhodami
proizvodstva. Sb. "Qbmen opytom po zelenomu stroitel'stvu", Sverdlovsk.
Izd, UNIX AKKh RSFSR, 1961.
Guseva, V. A. Vliyanie tnineral'nogo pitaniya na okislitel'no-vosstanovitel'nyy
rezhim i gazoustoychivost1 rasteniy. Sb., "Dytnoustoychivost1 rasteniy i
dyraoustoychivye assortimenty". Gor'kiy-Moskva, izd. Gor'kovskogo un-ta,
1950.
Ivanov, S. L. Osnovnoy Biokhimicheskiy zakon evoliutsii veshchestva v orgazizmakh.
Tr. prikladnoy botaniki i selektsii, t. XVI, M., 1926.
Ivanov, L. A., N. L. Kossovich, Polevoy roetod opredeleniya fotosinteza v
assimilyatsionnykh kolbakh. Bot. zh. t. 31, vyp. 5, 1946.
Ivanov, L. A. Svet i vlaga v zhizni nashikh drevesnykh rasteniy. "Timirya-
zevskiye chteniya", No. 5, M., Izd-vo AN SSSR, 1947.
lonin, V. M. i V. F. Koltasheva. Ozelenenie sanitarno-^ashchitnykh zon. Sb.
"Rekomendatsii po ozeleneaiyu gorodov". Sverdlovsk, Izd. UNII AKKh
RSFSR, 1961.
Kroker (Crocker), V. Rost rasteniy. M. Izd-vo inostr. lit., 1950.
Krasinskiy, N. P. Ozelenenie promploshchadok dymoustoychivym assortimentom.
M., Izd. AKKh SSSR, 1937.
Krasinskiy, N. P. 0 fiziologicheskoy sushchnosti gazoustoychivosti rasteniy.
Gor'kiy, Uch. zap. Gor'kovskogo un-ta, vyp. 9, 1940.
Krasinskiy, N. P. Metody izucheniya gazoustoychivosti rasteniy. Sb. "Dytnoystoy-
chivost1 rasteniy i dymoustoychivye assortimenty". Gor'kiy-Moskva, Izd.
Gor'kovskogo un-ta, 1950.
Krasinskiy, N. P. Teoreticheskie osnovy postroeniya assortimentov gazoustoy-
chivy kh rasteniy. Sb. "Dyraoustoychivost' rasteniy i dymoustoychivye assort-
imenty". Gor'kiy-Moskva, Izd. Gor'kovskogo un-ta, 1950.
Knyazeva, E. I. Gazoustoychivost1 rasteniy v svyazi s ikh sistetnaticheskim
polozheniem i morfologo-anatomicheskimi i biologicheskimi osobennostyarai.
Tarn -zhe.
- 93 -
-------
Kretovich, V. L. Osnovy biokhimii rasteniy. M., Izd. Vysshey shkoly, 1961.
Kolesnlkov, P. A. Vzaimosvyaz' mezhdu dykhaniem i fotosintezom. "Uspekhi
sovremennoy blologii", vyp. 47, No. 3. 1959.
Lunts, L. B. Zelenoe stroitel'stvo. M. Goslesbumizdat, 1952.
Maksimov, N. A. Kratkly kurs fiziologii rasteniy. M., Sel'khozgiz, 1958.
Nelyubov, D. N. Geotropizm v laboratornom vozdukhe. Izv. Rossiyskoy Akademii
nauk, 1910.
Nikolaevskiy, V. S. Ekologo-fizlologicheskie issledovaniya gazoustoychivosti
drevesno-kustarnikovykh porod v usloviyakh g. Krasnoural'ska, V pechatl.
gergeev, L. A. Vynoslivost1 rasteniy. M.-L. Izd-vo "Sov.nauka", 1953.
Sergeev, L. A., K. A. Sergeeva i V. K. Mel'nikov. Morfo-fiziologicheskaya
periodichnost' i zimostoykost' drevesnykh rasteniy. Ufa. Izd. Bash FAN
SSSR, 1961.
Sveshnikova, V. M. Izuchenie transpiratsii u rasteniy v estestvennykh uslov-
iyakh proizrastaniya, "Polevaya geobotanika", t. 1, M., Izd-vo AN
SSSR, 1959.
Savina, A. V. Izuchenie vliyaniya rubok ukhoda na svetovoy rezhim i energiyu
assimilyatsii vsosnovom nasazhdenii. Tr. VNIILKR, vyp. 21, 1941.
Sabashnikov, V. Vliyanie kamennougol'nogo dyma na okruzhayushchuyu rastitel'
nost1. "Bolezni rasteniy", No. 3-4, 1911.
Timofeeva, L. V. Vrednye vybrosy medeplavil'noy promyshlennosti na Srednera
Urale. V sb. "Voprosy gigieny truda, profpatologii i promyshlennoy
toksikologii". Izd. Ministerstva zdravookhraneniya RSFSR i Sverdlovskogo
in-ta gigieny truda i profpatologii, Sverdlovsk, 1958.
Noack, K. Untersuchungen uber die Finwirkung shwefler Saure auf die Pflanzen,
1920.
Wieler, A. Untersuchungen uber lichtkatalytischeVorgange von physiologischer
Bedeutung. Zeitschrift Botanik. Bd. 12. 1905.
Uislicenius, H. Sairanlung von Abhandlungen uber Abgase und Rauchschaben.
Berlin, 1914.
Jahnel, H. Physiologisches uber Einwirkung von Schwefeldioxid auf die Pflanzen.
Wissenshaftliche Zeitschrift, v. 4, No. 3, 1954.
- 94 -
-------
THE ACTIVITY OF CERTAIN ENZYMES
AND GAS RESISTANCE OF WOODY PLANTS
V. S. Nikolaevskiy
Institute of Plant and Animal.Ecology of the> Ural Branch
of the Academy or Sciences of the USSR
From Akad. Nauk SSSR. Ural. Filial. Trudy Institute Ekologii Rasteniy i
Zhivotnykh. Fiziologiya i ekologiya drevesnykh rasteniy. (Materialy II
Ural'skogo soveschaniya) II. (Sverdlovsk, 1968) p. 208-211.
The nature of the chemical action of sulfur dioxide on green plants
is still not clear. The known hypotheses (Noack, 1924; Krasinskiy, 1950)
give only general ideas concerning the mechanism of the damaging action
of photodynamic oxidation processes. Many have observed that sulfur
dioxide in weak concentrations inhibits photosynthesis while not affecting
respiration (Thomas and Hill, 1937; Kroker, 1950); others have shown that
it at first lowers respiration and then increases it (Wieler, 1905); and
still others have reported that respiration, under the influence of sulfur
dioxide gas, increases or remains constant (Krasinskiy, 1950). Our investi-
gations (Nikolaevskiy, 1964) have shown that, with noticeable plant
damage, photosynthesis ceases, while respiration is intensified several
times in light. These apparent contradictions are probably owing to the
fact that the respiration was studied on plants that were exposed to
different intensities of sulfur dioxide. According to the literature,
sulfur dioxide may be considered an inhibitor of the dark reactions of
photosynthesis. The works of Thomas and Hill (1937) confirm this view,
and have shown that weak concentrations of sulfur dioxide gas influences
photosynthesis in the same manner as does a passing cloud. After removal
of the action of the gas, photosynthesis returns to normal. There is still
little information in the literature concerning the role of enzyme systems
in gas resistance. The first and not-so-successful attempt in this direction
was undertaken by N. P. Krasinskiy (1937). It seems improbable that such an
active anion [sic] as sulfur dioxide would not exhibit an influence on
oxidation-reduction enzymes. It is true that, owing to its precipitating
action on proteins, sulfur dioxide gas occupies one of the first positions
in a series of lyotropic anions (Maksimov, 1958). According to the notion of
A. L. Kursanov (1940), during the precipitation of the plant's proteins, the
bond is broken in the enzyme-protein complex and the hydrolytic enzyme
activity is increased.
It has been shown that plants in the process of ontogenesis undergo a
change in terminal oxidase (Turkova, 1963), which could be represented by
various enzymes: cytochrome oxidase, polyphenol oxidase, peroxidase, and
others. Their role is to assist in accomplishing the final stage of respira-
tion — the union of the hydrogen of the oxidized substrate with the acid.
The above mentioned enzymes, in addition to dehydrogenases, are most often
considered in a study of the relation of respiration to the phenomenon of
plant resistance to various unfavorable environmental conditions.
- 95 -
-------
The resistance of plants to abiotic factors Is related to the lability
of metabolism (Slsakyan, 1940). At present, several types of plant respira-
tion are known; glycolytic, hexosemonophosphatic, and several others.
There is evidence, for example, that apotome respiration plays an important
role in the resistance of plants to infections (Rubin, 1960). It can be
supposed that the roles of the first-mentioned two types of respiration
will be different in relation to gas resistance.
We have studied the activity of peroxldases and polyphenol oxidases as
described by A. N. Boyarkin (1951, 1954) and catalases according to the
method of A. N. Bakh and A. I. Oparin (which was borrowed from Val'ter et al. ,
1957) on leaves of the box elder tree Acer negundo L. (resistant to sulfur
dioxide gas) and of the European birch Betula verrucosa L. (non-resistant
tb Sulfur dioxide gas). The dynamics of the activity were investigated dur-
ing the course of two months (July, August). Also studied was the effect of
different concentrations of sulfur dioxide gas on the susceptibility of plants
to injury in relation to enzyme activity.
During the summer the leaves of box elder are characterized by a low
catalase activity and a high peroxidase activity, while in the case of birch
the reverse was true, that is, there was a high activity of catalase and a
low peroxidase activity (Fig. 1). This agrees very well with data (Mikhlin,
1960; Turkov, 1963) that indicate a higher activity of cafalase during the
lower activity of peroxidase. During the fall, the birch begins to exhibit
polyphenoloxidase activity, while the elder displayed this phenomenon later.
/» n n a
a n n *i i a na
n a
i
July
August
Fig. 1 Activity of catalase (a) and
peroxidase (b) in 1964 in leaves.
1 - common birch; 2 - box elder
- 96 -
-------
In experiments with artificial fumigation of foliated branches of the
box elder and common birch by sulfur dioxide, interesting data were also
obtained. In Fig. 2 is represented the change of enzyme activity under the
influence of sulfur dioxide gas in relative indices (percent of control).
Depending on the concentration of sulfur dioxide gas and the susceptibility
to injury, the activity of the two enzymes of the elder and birch was changed
in different ways. In the elder, a weak concentration of sulfur dioxide gas
induced a slight decrease in catalase activity and a corresponding increase
in the peroxidase activity. The birch, under the influence of those same
concentrations of sulfur dioxide gas, showed a significant decrease in cata-
lase activity, but peroxidase activity remained almost constant. With an
increase in susceptibility to injury, the two mentioned enzymes of these
species underwent a change in activity. The activity of the catalase never
achieved its peak level. This is in agreement with the data (Ostrovskaya,
Bershteyn, 1953; Mikhlin, 1960) concerning the decrease of catalase activity
of plants under the influence of the anions. The activity of peroxidase in
birch under the influence of sulfur dioxide at first slightly decreased,
then sharply increased (at injury of 50% and more) and again decreased. The
increase in peroxidase activity in the elder began earlier (at 20% injury).
In Fig. 2, it is apparent that plants having different degrees of gas resist-
ance react differently to sulfur dioxide gas. These phenomena are evident
2
*j
c
o
u
U
<
710
20ff
700
ISO
no
to
0 tO 29 M *f U 80 10 tO 90 (Off
Damaged section of leaf, %
Fig. 2 Influence of sulfur
dioxide gas on the activity of
catalase (a) and peroxidase (b)
on the day of fumigation of the
leaves.
1 - common birch
2 - box elder
even if injury is not obvious. Furthermore, according to the character of
the curves of enzyme activity, it can be assumed that the elder possesses
more labile enzymic systems than does the birch. A similar conclusion was
drawn by us earlier on the basis of studies of the water regime and anatomi-
cal structure of leaves (Nikolaevskiy, 1964). At the end of August, a signif-
icant increase in activity of polyphenoloxidase (from 6-13 times) under the
influence of sulfur dioxide was observed.
The nature of the simultaneous study of activity of the two enzymes
(catalase and peroxidase) and of the degree of leaf injury allows for a
supposition of their mutual interdependence. It is known that peroxidase
- 97 -
-------
cannot replace catalase (Mlkhlin, 1960), but published data indicate that
the activation of peroxidase may be connected with catalase inhibition.
In the presence of weak or unseen damages, the activity of enzymes on
the second and third day after fumigation again approaches normal. With
high injury (upward of 20-30%), the activity of the catalase on the second
and third day remained constant or continues to decrease. At the same time,
the activity of the peroxidase in the box elder, after three or four days,
returned to normal, but in the birch continued to decrease.
Conclusions
1. Arboreous plants differing in gas resistance (box elder and common
birch) are characterized by differences in seasonal dynamics of enzyme
activity (catalase and peroxidase).
2. Under the influence of sulfur dioxide, even with unseen leaf injury,
there occurs a change in enzyme activity. The weakening of the enzyme system
is more critical for the less resistant species — the common European birch.
3. Sulfur dioxide inactivates catalase and in a determined interval of
susceptibility to injury, the activity of peroxidase and polyphenoloxldase
increases.
4. The activation of peroxidase apparently enables plants to resist
poisoning by peroxides.
Literature
Boyarkin, A. N. Bystryy metod opredeleniya aktivnosti peroksidazy.-Biokhimiya,
1951, t. 16, vyp. 4.
Boyarkin, A. N. Bystryy metod opredeleniya aktivnosti polifenoloksidazy.-Tr.
In'ta fiziologii rasteniy im. K. A. Timiryazeva AN SSSR, 1954, t. 8, vyp. 2.
Val'ter, 0. A., Pinevich, L. M. , Varasova, N. N. Praktikum po fiziologii
rasteniy s osnovami biokhimii. M.-L., Sel'khozgiz, 1957.
Krasinskiy, N. P. Ozelenenie promploschadok dymoustoychivym assortimentom.
M. , Izd-vo AKKh SSSR, 1937.
Krasinskiy, N. P. Teoreticheskle osnovy postroeniya assortimentov gazoustoy-
chivy kh rasteniy.-Dymoustoychivost1 rasteniy i dymoustoychivye assortimenty.
Gor'kly -M., Izd-vo Gor'kovskogo gos. un-ta, 1950.
Kroker, V. Rost rasteniy. M. , Izd-vo inostr. lit., 1950.
- 98 -
-------
Literature (Continued)
Kretovich, V. L. Osnovy biokhimll rasteniy. M. , Izd-vo "Vysshaya shkola", 1961.
Kursanov, A. L. Obratlmoe deystvie fermentov v zhivoy kletke. M.-L., Izd-vo
AN SSSR, 1940.
Maksimov, N. A. Kratkiy kurs fiziologii rasteniy. M. , Sel'khozgiz, 1958.
Mikhlin, D. M. Biokhimiya kletochnogo dykhaniya. M., Izd-vo AN SSSR, 1960.
Nikolaevskiy, V. S. Nekotorye anatomo-fiziologicheskie osobennosti drevesnykh
rasteniy v svyazi s ikh gazoustoychivost'yu v usloviyakh medeplavil'nykh
kombinatov Srednego Urala. Avtoref. diss. Sverdlovsk, 1964.
Ostrovskaya, L. K., Bershteyn, B. I. Vliyanie nitratov na katalaznuyu
aktivnost1 tkaney.-Voprosy biokhimii azotnogo i mineral'nogo pitaniya
rasteniy. Kiev, Izd-vo AN USSR, 1953.
Rubin, B. A. Dykhanie i ego rol* v immunitete rasteniy. Timiryazevskie
chteniya, 19. M., Izd-vo AN SSSR, 1960.
Sisakyan, N. M. Biokhimicheskaya kharakteristika zasukhoustoyshivosti rasteniy.
M. , Izd-vo AN SSSR, 1940.
Turkova, N. S. Dykhanie rasteniy. M. , Izd-vo MGU, 1963.
Noack K. Untersuchungen uber Einwirkung Schwefelsaure auf die Pflanzen.
Berlin. 1924.
Thomas, M. D., Hill, G. R. Relation of sulfur dioxide in the atmosphere to
photosynthesis and respiration of alfalfa.-Plant Physiol., 1937, vol. 12,
No. 2.
Wieler, A. Untersuchungen uber lichtkatalytische Vorgange von physiologischer
Bedeutung.-Zs. Bot., 1905, Bd 12.
- 99 -
-------
VARIATION IN THE OXIDIZABILITY OF THE CELL CONTENT AS ONE OF
THE INDICATORS OF GAS RESISTANCE IN PLANTS
V. G. Antlpov, I. I. Chekalinskaya
Central Botanical Garden of the Academy of Sciences of the Belorussian SSR
From Akad. Nauk SSSR Ural. Filial. Komls. po Okhrane Prlrody. Rastltel'nost'
1 promyshlennye zagryaznenlya. Okhrana prlrody na Urale. Vol. V,
(Sverdlovsk, 1966) p. 29-35.
Introduction
According to our calculations, there Is at the present time Information
concerning the gas resistance of more than 450 species and varieties of trees.
There is, however, considerable divergence in the determination of the degree
of gas resistance of individual species and varieties, and particularly of the
widespread ones. This impedes the practical use of these data. The contra-
dictions are particularly great within the limits of a genus. While the spruce
and the fir are considered by all the authors as of low resistance, and the yew
and arborvitae of high resistance, the equally widespread genera such as pine,
larch, and juniper are sometimes subject to contradictory views. Still more
confused is the picture of the deciduous trees.
Similar contradictions are found also within a species. Thus, among the
coniferous species the widely distributed Austrian pine and Mugho Swiss moun-
tain pine are considered by some authors to be very gas-resistant; by others,
moderately resistant, and by still others as only slightly resistant. The
same holds true in regard to broad-leaved trees. Thus, the summer oak, white
willow, Norway maple,Tartarian maple, black alder, common lilac, balsam poplar,
and black poplar are considered by some authors as the most resistant and by
others as the least resistant species.
A step in the right direction was the division of the injurious gases into
groups according to their effect on plants, and the selection of plants for
such groups. Thus, V. Krocker (1950) subdivided the gases into physiologically
active, slightly injurious (ethylene, propylene, and carbon monoxide), and
injurious (hydrocyanic acid, mercury vapors, sulfur dioxide, ammonia, chlorine,
and hydrogen sulfide). N. P. Krasinskiy (1937) separates acid gases (sulfur
dioxide and sulfur trioxide, chlorine, and hydrogen chloride) and non-acid gases
(ammonia). Adams and Pullmann (1956) as well as N. D. Thomas (1962) consider
that the greatest damage to plants is caused by sulfur dioxide, fluorine-
containing compounds, and smog. There are also other groupings.
The selection of plants according to their reaction to the pollutants gave
interesting data in many cases. According to the studies of the K. D. Pamfilov's
Academy of Municipal Economy, blackthorn and Virginia birdcherry showed greater
resistance to chlorine than to other gases, whereas silverberry and woollyleaf
mock orange were less resistant (Kuntsevich and Turchinskaya, 1957).
- IOC -
-------
Part of this investigation was devoted to the study of gas resistance
within a species. Special attention was devoted to elucidating the effect
of various ecological and weather conditions on the vulnerability of plants,
and, to a lesser degree, to the gas resistance of plants as it relates to
the daily and annual life cycle and to general ontogenesis of woody plants.
Thomas and Hill (1935) along with other investigators (among them our
own N. P. Krasinskiy) considered that the toxicity of sulfur dioxide is
largely attributable to its reducing properties.
N. P. Krasinskiy (1937, 1950) suggested differentiation of three kinds
of gas resistance in plants: biological, morphological-anatomical, and
physiological. According to his point of view the effect of smoke gases
should reflect particularly on cells having a greater oxidizability of
their water-insoluble substances, i.e.. the basic components of the proto-
plasm. In gas-resistant plants the oxidizability of the cell content is
lower and in gas-sensitive plants it is higher.
M. A. Zheleznova-Kaminskaya (1953) considered the extent of total
oxidizability of the cell content basic in the selection of gas-resistant
coniferous plant material for the city of Leningrad. M. D. Thomas (1962)
pointed out the changes in the gas resistance of plants during a day and
noticed that the leaves are sensitive to the action of sulfuric anhydride
in the morning. N. P. Krasinskiy (1950) found that in the course of growth
the total oxidizability usually increases with the age of the leaf.
There are indications of changes in the resistance of trees depending
on their age. Thus, 50 to 80-year-old Norway maple, oak, aspen, and linden
were less damaged than 5 to 10-year-old trees (Krasinskiy, 1937). Pelz
(1956) noted conversely that young trees are less damaged than old or
middle-aged ones. We have also found greater resistance of young common
pine in plantings in the Chelyuskintsev part of the city of Minsk.
Experimental Part
In setting up the investigations we decided to elucidate the variability
of the physiological gas resistance in diurnal, annual, and life cycle of
the plants' growth and development, in order to establish periods of greater
or lesser gas resistance. Here we are presenting preliminary data, which
are largely still under investigation. The research started in 1960. The
method used by N. P. Krasinskiy (1950) was utilized to determine the total
as well as the differential oxidizability of the cell content. Analyses
were made at an average of 10-day Intervals and the phenophase in the
plant development was then fixed by the method of the Botanical Institute.
During the spring samples were taken more frequently. Nine species (5 broad-
leaved and 4 coniferous) 25-30 years old were used in the experiments. The
selected species grew predominantly on the grounds of the Central Botanical
Garden of the Belorussian Academy of Sciences. Of the gas-resistant (so con-
sidered by many authors) broad-leaved species we used the Eastern poplar and
of the coniferous — the Colorado spruce. Of the low-resistant broad-leaved
species littleleaf linden, red ash, and pubescent birch were used; of conif-
erous — the Norway and the Eastern white pine; and the English oak, about
- 101 -
-------
whose gas resistance there are contradictory opinions, was also used.
In the 1960 experiments carried out with broad-leaved trees, it was
noticed that in the determination of oxidizability in the first half of
the summer the filtration of the extract took only several minutes,
whereas the same operation in the second half of the summer took several
hours. This is explained by the accumulation of starch. Keeping the
leaf extract exposed to air caused its oxidation and consequently led to
errors. Thus, in the Eastern poplar the oxidizability of water-soluble
substances after 20 hours decreased to 1/3 and that of the water-insolu-
ble substances to 1/2. In the red ash there was some increase. In the
English oak and littleleaf linden the oxidizability of water-soluble
substances in the middle of the day rose somewhat and then dropped insig-
nificantly. The oxidizability of water-soluble substances gradually
declined and reached 25% in 24 hours, regardless of whether the extract
was kept in light or in the dark. This shows that the change in oxidiza-
bility cannot be ascribed to photoreactions. As the.method of the investi-
gation was refined, the change in oxidizability of the cell content was
studied in relation to developmental phases of the plant and in relation
to weather and soil conditions. No appreciable variations in the oxidiza-
bility of the cell content were observed in connection with weather con-
ditions (sunny or cloudy skies, or sleet), except that the oxidizability
of water-soluble substances in the pubescent birch growing on poor soil
rose by 30% during sleet. It is noteworthy that this was not the case with
birches growing in rich soil. In the case of red ash growing in rich as
well as in poor soil, the oxidizability of water-soluble substances in the
same day dropped by approximately 15%. Soil conditions had no significant
effect on the oxidizability of the cell content of littleleaf linden, red
ash, and Berlin poplar. The variations did not exceed 5Z.
The most interesting data (see Table) were obtained in determining
the differential and total oxidizability of cell content in the various
developmental phases of the plant in the course of a growing period. The
least oxidizability was observed in leaves in their initial development
stages (L? and L.3) before they reached normal size and maturity. As the
leaf matures, the oxidizability, particularly of the water-insoluble sub-
stances, increases rapidly. The maximum Is reached in the English oak and
pubescent birch when all the leaves attain normal size and maturity (1,5 and
L6). In the red ash, Eastern poplar, and littleleaf linden the maximum Is
reached during the time from the onset of leaf turning to complete yellowing
of leaves. Then the oxidizability of the cell content drops rapidly but
does not reach the Initial level characteristic of the period when leaf
buds open.
In the course of a growing period the oxidizability of both water-
soluble and water-insoluble substances nearly doubles in littleleaf linden,
Eastern poplar, and English oak. In the red ash, the oxidizability of the
water-soluble substances increased more than three times; in the pubescent
birch it increased less significantly.
- 102 -
-------
Average total and differentiated oxidizability of cell content of leafy species
at various developmental stages of the leaves in 1961.
I
I-"
o
Oxidizability
L2
L3
L*
L5
L6
LT1
LT2
LT3
LDl
LD2
LD3
average
Littleleaf linden
Water-soluble
substances
Water-insoluble
substances
Total
No. of determinations
1.28
2.35
3.63
4
1.63
2.73
4.36
8
1.8
3.47
5.27
8
1.8
3.9
5.7
4
2.1
4.15
6.27
8
i
2.2
5.15
7.35
8
2.5
5.7
8.2
4
Eastern poplar
Water-soluble
substances
Water-insoluble
substances
Total
No. of determinations
1.22
2.12
3.34
8
1.46
2.5
3.96
8
1.85
2.75
4.60
16
-
-
-
-
2.1
3.4
5.5
12
2.4
4.0
6.4
4
2.0
3.5
5.5
4
-
-
-
-
-
-
-
-
2.2
5.25
7.45
8
2.0
4.5
6.5
4
1.94
4.13
6.07
T-56
1.7
2.85
4.55
8
-
-
-
-
-
—
-
-
-
—
-
-
1.81
3.01
4.82
T-60
Red ash
Water-soluble
substances
Water-insoluble
substances
Total
No. of determinations
1.13
1.75
2.88
4
1.86
4.1
5.96
8
1.9
5.4
7.3
12
_
-
-
-
2.1
5.7
7.8
12
2.35
5.9
8.25
8
2.8
6.0
8.8
4
2.65
5.15
7.80
8
—
-
-
-
_
-
-
-
—
-
-
-
2.11
4.85
6.96
T-56
(Continued)
-------
(Continued)
Average total and differentiated oxidizability of cell content of leafy species
at various developmental stages of the leaves in 1961.
Oxidizability
L< | U
lA
L5
Lb
LTl
LTZ
L13
LD1
LD*
LD-* I average
Water-soluble
substances
Water-insoluble
substances
Total
No. of determinations
Water-soluble
substances
1 Water-insoluble
£ substances
*• Total
1 No. of determinations
Pubescent birch
—
_
-
-
1.56
4.78
6.34
12
_
_
-
_
_
-
-
1.78
5.43
7.31
32
1.5
4.0
5.5
4
1.3
3.8
5.1
4
1.2
3.5
4.7
4
_
_
-
-
English oak
1.13
3.25
4.38
4
1.5
5.81
7.31
8
1.7
6.1
7.8
4
_
-
-
-
2.3
6.56
8.96
16
_
_
-
-
—
_
-
-
1.46
4.3
5.76
T-56
_
_
-
-
2.12
4.5
6.26
12
2.0
4.14
6.4
4
1.75
3.75
5.50
8
1.78
4.91
6.69
T-56
I/ - leaves in growing state; L.3 - leaves of normal size, not yet mature; L^ - leaves mature, yet not of
normal size; L5 - most of leaves mature, on upper part of shoots, still young leaves; L*> - all leaves of normal
size and maturity; LTl - appearance of autumn coloring; LT? - approximately half of the leaves turned; LT^ - all
leaves turned; LD* - onset of leaf drop; LD^ - approximately half of the leaves dropped; LJ)3 - most of the leaves
dropped.
-------
The oxidizability of the cell content of conifers was studied through-
out the year. The lowest total differential and oxidizability were found
in year-old needles, it was somewhat higher in two-year-old needles, and
the highest in older needles. Needles in their first year of life have the
lowest oxidizability, and it gradually increases until the growth of the
needles is complete.
Variations in the extent of oxidizability during the growth period
were observed in the needles from the first to the last year of their
growth. During the summer, a period of greatest photosynthesis in the
needles, an increase in the oxidizability of the cell content was observed
in the Colorado spruce; the increase was greater for the water-insoluble
substances than for the water-soluble. Toward autumn, when photosynthesis
activity of the needles slackens and the old needles begin to drop off,
the oxidizability decreased to 30% in the needles of all ages. With the
advent of cold, it again increases to an extent exceeding that of the summer
period. A certain relationship between the temperature of the air and
oxidizability was observed. Thus, in January 1961, the temperature in
Belorussia was very low (down to -408C). During that period the oxidizability
was very high, it decreased somewhat with increasing temperature and remained
stable until spring.
The oxidizability curve of water-insoluble substances of the Eastern
white pine is very similar to that of the Colorado spruce; however, its
oxidizability in the summer period is appreciably higher than in winter.
In January it shows a great increase in oxidizability. The oxidizability
of water-insoluble substances remained almost constant throughout the
year, except for short periodic increases in summer and winter.
The Scotch pine and the Eastern white pine have much in common, except
that in the former in December there was a sharp drop in the oxidizability
of water-insoluble substances by almost a third and then a sharp increase
— more than twofold — reaching a maximum in the middle of January, followed
by a gradual increase. These fluctuations in the oxidizability of the
water-insoluble substances are most pronounced in the two-year-old needles.
Discussion of Results
From the above it is clear that if oxidizability of the cell content
be accepted as an indicator of gas resistance of plants, then it is neces-
sary to consider the constantly changing gas resistance. There are some
indications of a connection between the fluctuations of the processes within
the plant and changes in its surrounding environment; this relation, however,
requires further study. It is quite possible that underestimation of the
phenophase through which the plant was passing caused contradictory data
on conifers presented by M. A. Zheleznova-Kaminskaya (1953). According to
her data, the oxidizability of the cell content of the needles in the
different species of the Pinaceae cannot serve as an indicator of their gas
resistance. Thus, in Abies sibirica Ldb.. Pinus svlvestria L.. and Pinus
peuce Gris., the oxidizability of the cell content is relatively low, and
yet, these species are not gas-resistant; indeed, N. P. Krasinskiy (1950)
considers the Scotch pine a nonrecommended species. M. A. Zheleznova-
- 105 -
-------
-Karainskaya points out that the high gas resistance of the larches cannot
be explained by physiological gas resistance because the oxldizability of
the cell content in the larch needles is highest of all conifers.
N. P. Krasinskiy (1939) brought out the question of the effect of
the age of leaves on the total oxidizability. He pointed out, in particu-
lar, that in the needles of the conifers the total oxldizability increases
with age (according to his data it is higher in the needles of the preced-
ing year than of the current year). We have shown that with age there is
particularly an Increase in the oxidizability of water-insoluble substances.
According to N. P. Krasinskiy, it is these substances that are dominant in
determining the gas resistance of plants.
N. P. Krasinskiy cites a number of Investigations of total oxidizability
in leaves of various age of essentially herbaceous plants. Time of taking
samples is not stated but it is known that they were always taken in the
morning (1950, p. 88). This apparently accounts for the contradictory recom-
mendations made by him. Thus, in 1937 he included among the gas-resistant
species European privet, and in 1950 he placed it among the non-resistant.
Silverberry and hedge cotoneaster were placed by him in 1937 among the
recommended species and in 1950 in the nonrecommended group, and so on.
In the numerous determinations on which Krasinskiy based his conclusions
concerning the relation between gas resistance of plants and their system-
atic position he did not take into account changes in oxidizability. This
possibly is the reason for the considerable fluctuation in oxidizability with-
in an individual genus, which Krasinskiy indicates.
Apparently, N. P. Krasinskiy did not consider the results that he
obtained to be reliable, because the selection of plants recommended by him
for use in industrial areas is chosen on the basis of visual observations
of a large number of Industrial enterprises and on the basis of laboratory
experiments with smoke exposure rather than on the basis of determining the
oxldizability of the cell content.
Seasonal variations in the oxidizability of the cell content observed
in trees is in agreement with the observations of Bobrov (1955) made on
Kentucky bluegrass. He suggests the use of bluegrass as an indicator of
air pollution by industrial gases and notes that the greatest injury is
sustained by mature leaves with reactive stomata. Old and young leaves are
less susceptible to smoke.
Conclusions
1. Changes In the oxldizability of cell content and, particularly, of
water-insoluble substances is connected with the developmental phases of
leaves.
2. Life activity of plants is connected with an increase in the oxidiza-
bility of the cell content, and a lowering in life activity of the plant leads
to a decrease in its oxldizability.
- 106 -
-------
3. In conifers there is a connection between air temperature and
the oxidizability of the cell content, and particularly, of the water-
insoluble substances.
4. Acceptance of the oxidizability of the cell content and, par-
ticularly, of its water-insoluble substances as an indicator of the
degree of gas resistance of plants should take into account considerable
variations.
5. Selection of woody plants should take into account the duration
of the gas resistance period of the particular species in the course of
its growth. The duration of distinct phenophases of the plant could
serve as an indicator of such periods.
Literature
Zheleznova-Kaminskaya, M. A. Rezul'taty introduktsii khvoynykh ekzotov v
Leningrade i ego okrestnostyakh.- Introduktsiya rasteniy i zelenoe
stroitel'stvo. vyp.3. Pod red. S. Ya. Sokolova. M.-L., Izd.-vo
AN SSSR, 1953.
Krasinskiy, N. 0. Ozelenenie promploshchadok dymoustoychivym assortimentorn.
M., 1937. (AKKh im. K. D. Pamfilova)
Krasinskiy, N. P. 0 fiziologicheskoy sushchnosti gazoustoychivosti rasteniy.
Uch.zap. Gor'kov. gos. un-ta, 1939, vyp. 9
Krasinskiy, N. P. Teoreticheskie osnovy postroeniya assortimenta gazoustoy-
chivy kh rasteniy.- Dymoustoychivost' rasteniy i dymoustoychivye
assortimenty. Pod red. N. P. Krasinskogo. M.-Gor'kiy, 1950 (Gor'kov.
gos.un-t i AKKh im. K.D. Pamfilova)
Kroker, V. Rost rasteniy. M. Izd-vo inostr.lit., 1950.
Kuntsevich, I. P., I. N. Turchinskaya. Ozelenenie fabrichno-zavodskikh
ploshchadok i promyshlennykh poselkov. M., 1957 (M-vo kommunal'nogo
kh-va RSFSR)
Tomas, M. D. Vliyanie zagryaznenlya atmosfemogo vozdukha na rasteniya.-
Zagryaznenie atmosfemogo vozdukha. Zheneva, 1962 (Vsemirnaya
organizatsiya zdravookhraneniya OON)
Adams, D. F., M. S. Pullmann. The effects of air pollution on plant life.-
Arch. Industr. Health, 1956, vol. 14, no. 3
- 107 -
-------
Literature (Continued)
Bobrov, R. A. The leaf structure of Poa annua with observations on its
smog sensitivity in Los Angeles county.-Amer. J. Bot. , 1955, no. 5
Pelz, E. Gasformige Luftverunreinigungen und Holzartenwahl in Gebieten
rait Industrierauchschaden.- Forst und Jagd, 1956, no. 8
Thomas, M. D., G. R. Hill. Absorption of sulfur dioxide by alfalfa and
its relation of leaf injury.-Plant Physiol., 1935, no. 10
- 108 -
-------
EFFECT OF SULFUR DIOXIDE ON THE ENZYMATIC ACTIVITY OF TREE LEAVES
V. S. Nlkolaevskiy
Jnstitut of Biology, Ural Branch of the Academy of Sciences, U.S.S.R.
From Akad. Nauk SSSR. Ural. Filial. Komis. po Okhrane Prirody. Rastitel'nost'
i promyshlennye zagryazneniya, Okhrana prirody na Urale. V (Sverdlovsk, 1966)
p. 19-23.
Many Investigators (Thomas and Hill, 1937; Krocker, 1950) noted that
sulfur dioxide in weak concentrations disturbs photosynthesis without affect-
ing respiration. Wieler (1905) pointed out that S02 first depressed respira-
tion and then heightened it. N. P. Krasinskiy (1950) thought that under the
influence of S0£ respiration either does not change or that it increases.
Our investigations (Nikolaevskiy, 1962, 1963, 1964) have established
the relationship between the gas resistance of woody plants and the intensity
and direction of physiological and biochemical processes within their leaves.
A study of the various forms of resistance led to the conclusion that the
resistance Is determined on the one hand by a group of factors connected with
the anatomical, physiological and biochemical properties of the plants and
on the other hand by the degree of resistivity and of lability of the metabo-
lism under adverse conditions.
In the five varieties of studied trees: ash-leaved maple, balsam poplar,
aspen, Siberian crab apple, and European white birch, a correlation was
established between gas resistance, lowered Intensity of photosynthesis and
respiration, and a lower oxidation-reduction potential. This relationship
may be explained by the fact that a decrease in the intensity of gas exchange
and metabolism is generally connected with a decrease in enzymatic activity
in plants (Siskyan, 1954). When photosynthesis is blocked or stopped, the
adsorbed solar energy finds a natural outlet in the oxidation of the
constituent and reserve organic substances and components of the cell.
Considering that most of the enzymes have reversible properties (Kursanov,
1940; Oparin and Gel'man, 1952), great damage should be expected in plants
with an increased enzymatic activity under normal conditions.
The varying effect of S02 on photosynthesis and respiration of plants
leads to the assumption that not all of the enzyme systems are affected by
the action of acid gases. The photodynamic action of chlorophyll under the
influence of S02 and of the cessation of photosynthesis, established by
Noack (1924) and N. P. Krasinskiy (1940), leads to the assumption that under
such conditions some photosynthetic reactions in the dark are inhibited.
Light reactions occurring upon accumulation of light energy are apparently
not affected by S02- A study of the effect of S02 on the enzymatic activity
of plants will, in our opinion, help in the understanding of the essence of
biochemical processes causing plant damage.
In 1963 we have studied the activity of catalase, peroxidase, and poly-
phenol oxidase in ash-leaved maple and in European white birch. The choice
- 109 -
-------
was motivated by the substantial differences in their physiological, bio-
chemical, and anatomical indices as well as by their reaction to acid gases
(Nikolaevskiy, 1962, 1963). In the terminology of N. P. Krasinskiy (1950 b)
ash-leaved maple belongs to the slightly affected varieties, while the
white birch belongs to the greatly damaged varieties. The activity of the
enzyme was studied from August 15 to September 25. For the same period
the effect of various concentrations of S0£ on the same enzyme was also
studied. In all, 14 fumigations were applied, in which the concentration
of S02 was 2.5x10-4 and 2xlO~5.
l. 0.01 in L
soln.
I attttuu.
September
xunxn
August
/ isatiutf
August , September
Enzynatio activity of plantsi
a- oatalasei b-
1.- European
/ a a nuts
September
maple
Catalase was studied gasometrically, peroxidase and polyphenol oxidase
by the D. M. Mikhlin and Z. S. Bronovitskaya (1949) method. Leaves for
analysis were collected at 9 A.M. from the same trees, on the southern side,
from the middle of the crown. The trees were 15-18 years old. Fumigation in
the gas chamber was carried out on cut twigs. As control, twigs placed in
tap water alongside the gas chamber were used. Analyses were repeated
2-4 times.
On the graphs are.given changes in the enzymatic activity of ash-leaved
maple and white birch during August and September 1963. Catalase activity in
the maple was lower than in the birch by 32.5%. Approximately the same differ-
ence was noticed in the activity of peroxidase and polyphenol oxidase in the
same species. Taking into consideration that enzymatic activity (Sisakyan,
1954) as well as physiological properties (Ivanov, 1946) in plants constitutes
characteristics of a spe-ies, we may clarify the possible causes of the
relationship between the gas resistance in plants and the intensity of photo-
synthesis. In our view, the differences in the enzyme activity in the maple
and in the birch confirm the assumption that the intensity of photodynamic
oxidation under the influence of SOj and of light is proportional to the
oxidation-reduction activity of enzyme systems in plants. Indeed, ash-leaved
maple is characterized b" a lower intensity of its gas exchange and of the
- 110 -
-------
Changes in the enzyme activity of tree leaves under the influence of sulfur dioxide
Plant
Absolute act iv
averages for A
September
L'ata-
lase
Poly-
phenol
oxidase
ity
ugust-
Feroxi-
dase
Results of fumigation of trees with S02
Concentration of 503 2.5x10"*
experiment
"CaCa-
lase
Holy-
phenol
oxidase
Heroxi-
dase
Control
Cata-
lase
Poly-
phenol
oxidese
Peroxi-
dase
Concentration of S02 2x1(7"^
Experiment
Cata-
lase
Poly-
phenol
oxidase
Peroxi-
dase
Control
tfata-
lase
Poly-
phenol
oxidase
Peroxi-
dase
Maple
Birch
First day after fumigation
29.4
43.5
3.5
4.6
3.3
4.7
10. 2»
38.1
4.5
~9T8
2.3
175.0
5.3
T08.0
2.9
98.0
4.2
150.0
26.8
100.0
45.8
100.C
1.6
100.0
4.9
100.0
3.0
100.0
2.8
T3oTo
29.5
85.2
34.4
"787E
5.1
79.6
6.7
155T5
1.0
38.5
2.1
T05T5
34.7
100.0
43.8
ToO
6.4
100.0
5.3
ToO
2.6
100.0
2.0
Toc~o
Second day after fumigation
Maple
Birch
—
-
•
-
—
-
8.6
cgTs
2.7
~&To
2.5
66^0
3.2
78.0
1.7
IsT?
1.8
82.0
28.9
TooTo
44.8
100.0
3.8
TooTo
4.1
100.0
2.9
100.0
2.2
100.0
27.0
763
34.3
86.8
1.5
7174
3.4
141.0
2.8
16575
1.8
69.0
35.3
100.0
39.0
100.0
2.1
100.0
2.4
100.0
1.7
100.0
2.6
100.0
* Numerator — absolute value; denominator — percent of control.
-------
activities of the three studied enzymes (see Graph) , and consequently by
a lower vulnerability. Undoubtedly, the differences in the adsorption
rates of acid gases by the leaves of the maple and of the birch (Nikolaevskiy,
1963) are also determining factors.
Experiments with artificial fumigation of twigs in a gas chamber (see
Graph) showed that the activity of the three enzymes changes under the
Influence of SC^. Particularly clear disturbances were observed in catalase.
Decline in the activity of catalase in the maple and in the birch is connected
with their vulnerability. Small concentrations of S02 (2xlO~5) caused in
both species similar changes in the activity of catalase. Definite changes
under the influence of SC>2 took place in the activity of the other enzymes.
High concentrations of SC>2 caused an increase in the activity of polyphenol
oxidase in the maple and of peroxidase in the birch. On the second day of
fumigation, the activity of all the three enzymes dropped in both species.
Small concentrations of S02 (2xlO~5) cause no externally apparent damage to
the maple and birch but lowered the activity of catalnse and affected the
activity of the two other enzymes.
There are certain differences in the action of small concentrations of
SC>2 when compared with the effect of high concentrations. In this case, the
maple exhibited decreased activity of all three enzymes on the first day and
an increased activity of peroxidase on the second day.
In the birch, on the first day, along with a drop in the activity of
catalase occurred an increase in the activity of the two other enzymes.
And on the second day, unlike in the maple, the activity of polyphenol oxi-
dase increased and that of peroxidase dropped. The inhibiting role of anIons,
and particularly of S02 is indicated in the literature (Mikhlln, 1960;
Koklna, 1939).
Data in the table lead to the assumption that decreased activity of
catalase under the Influence of S02 is likely to contribute to an increase in
the oxidation processes and to the damage of plants (Mikhlin, 1960) because
of accumulation of organic peroxide.
There is no clear understanding of the effect of SC>2 on other enzymes.
It may be assumed that the nature of the effect of 802 on enzymes will be
different In plant species characterized by differences in gas resistance.
Further studies are needed to elucidate the role of different enzyme systems
on the gas resistance of plants.
Conclusions .
1. Leaves of the ash-leaved maple and those of the European white
birch differ not only in their anatomy and physiology but in the activity of
some oxidative enzymes as well.
2. A clear proportional connection with injury to leaves by S02 is
established in the case of catalase. Great vulnerability of birch leaves is
connected with a greater decline In its catalase activity, when compared with
the maple.
- 112 -
-------
3. The effect of small and of large concentrations of sulfur dioxide
on enzymes differ. In the former case the activity of the three studied
enzymes increased on the second day following fumigation, while in the
latter case it dropped.
Literature
Ivanov, L. A. Svet i vlaga v zhizii nashikh drevesnykh porod.- Timiryazevskie
chteniya, V. M. Izd-vo AN SSSR, 1946.
Kokina, S. I. Ob okislitel'no-vosstanovitel'nykh protsessakh u pustynnykh
rasteniy.- Bot.zh. 1939, t.24, no. 1
Krasinskiy, N. P. 0 fiziologicheskoy sushchnosti gazoustoychivosti rasteniy.-
Uch.zap. Gor'kov.gos. un-ta, 1939, vyp. 9
Krasinskiy, N. P. Metody izucheniya gazoustoychivosti rasteniy.- Dymoustoychi-
vost1 rasteniy i dymoustoychivye assortimenty. M.-Gor'kiy, 1950 a (Gor'kov.
gos.un-t i AKKh.im. K. D. Pamfilova)
Krasinskiy, N. P. Teoreticheskie osnovy postroeniya assortiraentov gazoustoy-
chivy kh rasteniy: - Tarn zhe. 1950 b.
Kroker, V. Rost rasteniy. M., Izd-vo inostr. lit., 1950.
Kursanov, A. L. Obratlmoe deystvie fermentov v zhivoy rastitel'noy kletke.
M.-L., Izd-vo AN SSSR, 1940.
Mikhlin, D. M. Biokhindya :kletochnogo dykhaniya. M., Izd-vo AN SSSR, 1960.
Mikhlin, D. M. i Bronovitskaya, Z. S. Yodometricheskiy metod opredeleniya
polifenoloksidazy i peroksidazy.- Biokhimiya, 1949, t.14, vyp.4
Nikolaevskiy, V. S. Ekologo-fiziologicheskie issledovaniya gazoustoychivosti
drevesno-kustarnikovykh porod v usloviyakh goroda Krasnoural'ska.- Doklady
pervoy nauchnoy konferentsii molodykh spetsialistov-biologov. Sverdlovsk,
1962 (In-t biol. UFAN SSSR)
Nikolaevskiy, V. S. 0 pokazatelyakh gazoustoychivosti drevesnykh rasteniy
(po issledovaniyam v gorode Krasnoural'ske).- Trudy In-ta biol. UFAN SSSR,
1963, vyp.31
Nikolaevskiy, V. S. Vliyanie sernistogo angidrida na drevesnye rasteniya v
usloviyakh Sverdlovskoy oblasti.- Okhrana prirody na Urale, IV.
Rastitel'nost' i promyshlennye zagryazeniya. Sverdlovsk, 1964 (UFAN
SSSR i Ur.gos.un-t)
Oparin, A. I. i Gel'man, N. S. K voprosu o svyazi dykhaniya s sinteticheskimi
protsessami u vysshykh rasteniy.-Dokl. AN SSSR, 1952, t.85, no. 6
- 113 -
-------
Literature (Continued)
Slsakyan, N. M. Biokhimiya obmena veshchestv. M., Izd-vo AN SSSR, 1954.
Noack, K. Untersuchungen uber die Einwlrkung schwefliger Saure auf die
Pflanzen. Berlin, 1924.
Thomas, M. D. a. Hill, G. R. Relation of sulfur dioxide in the atmosphere
to photosynthesis and respiration of alfalfa.- Plant Physiol. , 1937,
vol. 12, N 2
Wieler, A. Untersuchungen uber lichtkatalytische Vorgange von phyaiologischer
Bedeutung.- Zs Bot., 1905, Bd. 12
- 114 -
------- |