AMERICAN INSTITUTE OF CROP ECOLOGY
A RESEARCH ORGANIZATION DEVOTED TO PROBLEMS OF
PLANT ADAPTATION AND INTRODUCTION
WASHINGTON, 0. C.
AICE* SURVEY OF USSR AIR POLLUTION LITERATURE
Volume IX
GAS RESISTANCE OF PLANTS WITH SPECIAL REFERENCE TO
PLANT BIOCHEMISTRY AND TO THE EFFECTS OF MINERAL NUTRITION
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 RO1 AP00786
AIR POLUTION CONTROL OFFICE
i he
ENVIRONMENTAL PROTECTION AGENCY
APC
*AMERICAN INSTITUTE OF CROP ECOLOGY
809 DALE DRIVE
SILVER SPRING, MARYLAND 20910
1971
-------
PUBLICATIONS of the AMERICAN INSTITUTE OF CROP ECOLOGY
Net.
10
11
12
13
14
2)
24
25
KR AIN£-£colcyicol Crop Geography of the Ukraine end the
Ukrainian Agro-Qltraiic Analogues in North America
PCL-NO-^gricolrjro! Climatology of Poland and Its Agro-
CL;noKi_ V,atos' North America
C7cCKOSLOVAKIA-Agricul»urcl Clirnotology of Czechoslo-
vakia and Its Agro-Climatic Analogues in North America
YUGOiLAVIA-Agrici,! rural Climatology of Yugoslavia and Its
u-5re-Climarie Analogues in North America
GREECE-Ecological C^op Geography of Greece and Its Agro-
Climatic Analogues in Morrh America
ALSANIA-Ecologicol Plant Geography of Albania, Its Agri-
cultural Crop; 3"c Some North American Climatic Analogues
CHINA-Ecological Crop Geography of China and Its Agra-
Climatic Analogues in North America
GERMANY— Ecological Crop Geography of Germany and Its
A^ro-Climatic Analogues in North America
JAPAN (I )— Agricultural Climatology of Japan and Irs Agro-
Climofic Analogues in North America
FINLAND- Ecological Crop Geography of Finland and Its Agro-
Climatic Analogues in North America
SWEDEN-Agriculrural Climatology of Sweden and Its Agro-
Oimatic Analogues in North America
NORWAY-Ecologicaf Crop Geography of Norway and Its Agro-
Climatic Analogues in North America
SIBERIA-Agricgl rural Climatology of Siberia, Irs Natural Belts,
and Agro-Climatic Analogues in North America
JAPAN (2)— Ecological Crop Geography and Field Practices of
Japan, Japan's Natural Vegetation, and Agro-Climatic
Analogues in North America
RYUKYLJ ISLANDS-Ecologicol Crop Geography and Field
Practices of the Ryukyu Islands, Natural Vegetation of the
Ryui-yus, and Agro-Climatic Analogues in the Northern
Hemisphere
r-HENOi-OGY AND THERMAL ENVIRONMENT AS A MEANS
OF A PHYSIOLOGICAL CLASSIFICATION OF WHEAT
VARIETIES AND FOR PREDICTING MAT.URITY DATES OF
WHEAT
•',Bcseo' on Data of Czechoslovakia and of Some Thermally
•-nslagouT Areas of Czechoslovakia in the United States
Pacific I^Jonhweit)
"liEAT-CLIMATE RELATIONSHIPS AND THE USE OF PHE-
NOLOGY lh> ASCERTAINING THE THERMAL AND PHO-
TOTHERMAL REQUIREMENTS OF WHEAT
(Ba.ad on Daso of North America and Some Thermally Anal-
•>?&;., Areas of North America in the Soviet Union and in
-
A COMPARATIVE STUDY OF LOWER AND UPPER LIMITS OF
Tf.v>r-cRATUR!: !N MEASURING THE VARIABILITY OF DA Y-
OEG'=E SUMMATIONS OF WHEAT, BARLEY, AND RYE
BARLEY-CLIMATE RELATIONSHIPS AND THE USE OF PHE-
NOLOGY IN ASCERTAINING THE THERMAlMND PHO-
TOrKEWAL •'ECJIREMENTS OF BARLEY
Wlt RELATIONSHIPS AND THE USE OF PHENOL-
OGY ir-J ASCERTAINING THE THERMAL AND PHOTO-
T^r'MAL REQUIREMENTS OF RY"c
f ,G,'-iCvLTURAL ECOLOGY IN SUBTROPICAL REGIONS
.-/IC'OCCO, ALGERIA, TUNlSIA-Phyjicol Environment and
Agr.-.'jlrure . ...
LiSYA c-nd EGVPT-Physkal Environment and Agriculture. . .
J: ;iON OF SOOTH AFFICA-Physica) Environment and Agri-
c-jlfuie, v/rth Spir;-.il Reference to Winter-Rainfall Regions
AUST.-M1 A- Physical Environment and Agriculture, With Spe-
-io( ."rOrence 10 vVinter-Ramfdlf Regions .....
26 S. E. CALIFORNIA and S. W. ARIZONA-Physieal Environment
and Agriculture of the Desert Regions
27 THAILAND—Physical Environment and Agriculture
28 BURMA-Physical Environment and Agriculture
28A BURMA—Diseases and Pests of Economic Plants
286 BURMA-Climote, Soils and Rice Culture (Supplementary In-
formation and a Bibliography to Report 28)
29A VIETNAM, CAMBODIA, LAOS-rhysical Environment and
Agricul ture
29B VIETNAM, CAMBODIA, LAOS-Diseases andPestsof Economic
Plants
29C VIETNAM, CAMBODIA, LAOS-Climatological Data (Supple-
ment to Report 29A)
30A CENTRAL and SOUTH CHINA, HONG KONG, TAIWAN-
Physical Environment and Agricul ture S20.00*
308 CENTRAL and SOUTH CHINA, HONG KONG, TAIWAN-
Major Plant Pests and Diseases
31 SOUTH CHINA-lts Agro-Climatic Analogues in Southeast Asia
32 SACRAMENTO-SAN JOAQUIN DELTA OF CALIFORNIA-
Physicol Environment and Agriculture
33 GLOBAL AGROCLIMATIC ANALOGUES FOR THE RICE RE-
GIONS OF THE CONTINENTAL UNITED STATE
34 AGRO-CLIMATOLOGY AND GLOBAL AGROCLIMATIC
ANALOGUES OF THE CITRUS REGIONS OF THE CON-
TINENTAL UNITED STATES
35 GLOBAL AGROCLIMATIC ANALOGUES FOR THE SOUTH-
EASTERN ATLANTIC REGION OF THE CONTINENTAL
UNITED STATES
36 GLOBAL AGROCLIMATIC ANALOGUES FOR THE INTER-
MOUNTAIN REGION OF THE CONTINENTAL UNITED
STATES
37 GLOBAL AGROCLIMATIC ANALOGUES FOR THE NORTHERN
GREAT PLAINS REGION OF THE CONTINENTAL UNITED
STATES
38 GLOBAL AGROCLIMATIC ANALOGUES FOR THE MAYA-
GUEZ DISTRICT OF PUERTO RICO
39 RICE CULTURE and RICE-CLIMATE RELATIONSHIPS With Spe-
cial Reference to the United States Rice Areas and Their
Latitudinal and Thermal Analogues in Other Countries
40 E.WASHINGTON, IDAHO, and UTAH-Physical Environment
and Agriculture
41 WASHINGTON, IDAHO, and UTAH-The Use of Phenology
in Ascertaining rhe Temperature Requirements of Wheat
Grown in Washington, Idaho, and Utah and in Some of
Their Agro-Climatically Analogous Areas in the Eastern
Hemisphere
42 NORTHERN GREAT PLAINS REGION-Prelirainory Study of
Phenological Temperature Requirements of a Few Varieties
of Wheat Grown in the Northern Great Plains Region and in
Some Agro-Climatically Analogous Areas in the Eastern
Hemisphere
43 SOUTHEASTERN ATLANTIC REGION-Phenological Temper-
ature Requirements of Some Winter Wheat Varieties Grown
in the Southeastern Atlantic Region of the United States and
in Several of Its Laritudinally Analogous Areas of the Eastern
and Southern Hemispheres of Seasonally Simitar Thermal
Conditions
44 ATMOSPHERIC AND METEOROLOGICAL ASPECTS OF AIR
POLLUTION-A Survey of USSR Air Pollution Literature
45 EFFECTS AND SYMPTOMS OF AIR POLLUTES ON VEGETA-
TION; RESISTANCE AND SUSCEPTIBILITY OF DIFFERENT
PLANT SPECIES IN VARIOUS HABITATS,IN RELATION TO
PLANT UTILIZATION FOR SHELTER BELTS AND AS BIO-
LOGICAL INDICATORS-A Survey of USSR Air Pollution
Literature
(Continued on inside of back cover)
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AICE* SURVEY OF USSR AIR POLLUTION LITERATURE
Volume IX
GAS RESISTANCE OF PLANTS WITH SPECIAL REFERENCE TO
PLANT BIOCHEMISTRY AND TO THE EFFECTS OF MINERAL NUTRITION
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 - APC
AIR POLUTION CONTROL OFFICE
of the
ENVIRONMENTAL PROTECTION AGENCY
*AMERICAN INSTITUTE OF CROP ECOLOGY
809 DALE DRIVE
SILVER SPRING, MARYLAND 20910
1971
-------
TABLE OF CONTENTS
Page
PREFACE v
Maps of the USSR
Orientation vii
Climatic* Soil and Vegetation Zones viii
Major Economic Areas ix
Major Industrial Centers x
Principal Centers of Ferrous Metallurgy and Main
Iron Ore Deposits xi
Principal Centers of Non-Ferrous Metallurgy and
Distribution of Most Important Deposits of
Non-Ferrous Metal Ores xli
Principal Centers of the Chemical Industry and of
the Textile Industry xiii
Principal Centers of Wood-Workingf Papert and Food
Industries xiv
Main Mining Centers , xv
Principal Electric Power Stations and Power Systems ... xvi
PHYSIOLOGICAL-BIOCHEMICAL PRINCIPLES OF THE GAS RESISTANCE
OF PLANTS
V. S. Nikolayevskiy 1
EFFECT OF MINERAL NUTRITION ON THE GAS RESISTANCE OF
FORAGE GRASSES
V. M. Yatsenko, V. S. Nikolayevskiy, V. V. Firger,
and V. V. Suslova 28
EFFECT OF MINERAL NUTRITION ON CERTAIN PHYSIOLOGICAL-
BIOCHEMICAL CHARACTERISTICS AND GAS RESISTANCE OF
FORAGE GRASSES
V. M. Yatsenko and V. S. Nikolayevskiy 34
EXPERIENCE IN THE USE OF THE BIOCHEMOLUMINESCENCE METHOD
FOR DIAGNOSING THE GAS RESISTANCE OF PLANTS
V. S. Nikolayevskiy and A. G. Miroshnikova 50
iii
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Page
GAS RESISTANCE AND CERTAIN BIOCHEMICAL CHARACTERISTICS OF
ETIOLATED AND GREEN PLANTS OF FORAGE GRASSES
V. S. Nikolayevskly and A. T. Miroshnikova 54
EFFECT OF MINERAL NUTRITION ON THE METABOLISM OF CARBON-14
COMPOUNDS AND ON GAS RESISTANCE OF FORAGE GRASSES
V. V. Firger and T. B. Karpova 69
METABOLISM OF CARBON-14 COMPOUNDS IN FORAGE GRASSES AND
THE EFFECT OF SULFUR DIOXIDE ON IT
V. S. Nikolayevskiy, V. V. Firger, and
G. A. Vaseva 82
EFFECT OF SULFUR DIOXIDE ON PIGMENTS OF FORAGE GRASSES
V. S. Nikolayevskiy and V. V. Suslova 92
iv
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PREFACE
In the USSR, the rapid development of various industrial enterprises
and the extensive use of the internal combustion engine for motor vehicle
transportation contribute greatly to massive qualitative changes in the
habitat of living organisms through an ever increasing pollution of air,
soil, and streams. In many industrial areas and in large cities new
environmental conditions are being created which can sustain the existence
of no plant life at all or only of the most pollution-resistant plant
species or some of its variant forms.
The resistance of plants to pollution varies with a whole complex of
factors, primarily with type of emission, growth conditions, the physi-
ological-biochemical and morphological characteristics of a given plant
species, the developmental stage of the plants, and the nature of the ex-
posure. The combination of these factors appears to govern to a large
degree the resistance of a plant species, as well as of its ecotypes, and
its varieties or cultivars.
For plant scientists, two approaches are available to counteract the
noxious effects of pollution, namely:
(a) One approach rests on the recognition that plant species differ
in their resistance to a given pollutant or to certain combinations
of pollutants, and also that differences in resistance to pollution
have been found to exist in individual plants or varieties within
certain species. It should be possible, therefore, to obtain by selec-
tion plants resistant to certain pollutants and thus suitable to
areas where such pollutants prevail. It is self-evident that in the
assessment of environmental conditions prevailing in a given area the
nature of air pollution must be considered in relation to all the
other potent physical and biological stresses and factors governing
the overall problem of plant selection and plant adaptation.
(b) The second approach rests on the observation that certain agro-
technical field practices, notably a judicious supply of mineral
nutrients, increase the resistance of plants to air pollution.
The present volume consists largely of reports dealing with (1) an ex-
tensive survey of the principal advances in research on the gas resistance
of plants and of the causes of plant vulnerability to gas, and (2) a number
of investigations conducted in laboratories and on field plots of the Botan-
ical Garden of Perm1 University (a) in reference to the feasibility of regu-
lating the gas resistance of plants by means of certain nutritional elements,
-------
(b) in reference to a number of biochemical and physiological indicators
that can be used to solve the problem of diagnostics of the gas resistance
of plants and to characterize species differences in their gas resistance,
and (c) to determine the minimum-permissible norms of certain air pollutants
for individual species of plants.
Perm', where these investigations were conducted, is an area lying in
the highly industrialized Ural Region, which occupies a most important po-
sition in the industrial economy of the USSR. The local vegetation of
Perm1 and its environs, as well as that of the whole industrialized Ural
Region and that of a great many other highly industrialized areas and
regions of the USSR is badly affected by the increasing contamination of
the natural environment with the toxic emissions from numerous industrial
enterprises and power plants.
Some background information on the distribution of the Soviet industry's
production machine may be of interest in connection with that country's
present and potential pollution problems and investigations. The planned
distribution of production in the Soviet Union favors effective exploitation
of the natural resources of the USSR, especially in its eastern areas where
enormous natural resources are concentrated, and has led to the creation of
large industrial centers and complexes of heavy industry in many of the
country's economic areas (see page ix). The many diverse climatic conditions
of the country and its major economic areas as well as the geographical dis-
tribution of the Soviet Union's principal industrial and mining centers and
of its principal electric power stations and power systems can be seen from
the various maps presented as background material in this volume.
It is hoped that the selected papers will be conducive to a better
appreciation of some of the air pollution investigations conducted in the
USSR. As the editor of this volume I wish to thank my co-workers in the
Mr Pollution Section of the Institute for their valuable assistance.
M. Y. Nuttonson
September 1971
vi
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ADMINISTRATIVE DIVISIONS
S.S.R.
1. R.SF.S.R
2 KarelO'Finnish S.S.R
3. Estonian S.S.R
4. Latvian SS Ft
5. Ltthuarxan S S R
6. White RuMianSS.R.
7. Ukrainian &SR
8. Moldavian S.S.R.
9. Georgian S S R
10. Armenian S.S.R.
11. Aieroaydrhan S S.R.
12. KaiakhSS.R.
19. Uzbek S.S R
14. Turkmen S.S R
15. Tadlhik SS.R.
16- Kirgu S.S.R.
A Komi ASSft
kaya ASSR
jyj ASSR
hskjya ASSR
kaya ASSR
ya ASSR
ia ASSf»
nstax* ASSP
",'• •^-!.,1 *iSR
niKa.a ASSft
kaya ASSR
ka»a ASSR
evanikaya ASSft
Buryat '•' • „ -.1. • ASSR
Ya»-utska»a ASSR
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CLIMATIC ZONES AND REGIONS* OF THE USSR
X'..?::rARClc OCEAN^/-?
OKHOTSK
Zones: I-arctic, II-subarctic, Ill-temperate, IV-subtropical
Regions: 1-polar, 2-Atlantic, 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,
L2-monsoon forests, 13-Pacific forests, 14-Atlantic-continental steppe,
15-continental steppe West Siberian, 16-mountainous 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,
2A-mountainous Pamir-Alay
(After B. P. Alisov, "Climate of The USSR", Moscow 1956)
SOIL AND VEGETATION ZONES IN THE U.S.S.R.
viii
(After A. Lavrishchev, "Economic Geography
of the U.S.S.R.", Moscow 1969)
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MAJOR ECONOMIC AREAS OF THE U.S.S.R.
XVI Kazakhstan
XVII Cenlral Asia,
XVIII Byelorussian
PLANNED DISTRIBUTION OF INDUSTRIAL PRODUCTION IN ORDER
TO BRING IT CLOSER TO RAW MATERIAL AND FUEL SOURCES
An example of the planned distribution of industrial production in the USSR is the creation of large
industrial centers and complexes of heavy industry in many of the country's economic areas: the North-West
(Kirovsk, Kandalaksha, Vorkuta), the Urals (Magnitogorsk, Chelyabinsk, Nizhny Tagil), Western and Eastern
Siberia (Novosibirsk, Novokuznetsk, Kemerovo, Krasnoyarsk, Irkutsk, Bratsk), Kazakhstan (Karaganda, Rudny,
Balkhash, Dzhezkazgan).
Large industrial systems are being created - Kustanai, Pavlodar-Ekibastuz, Achinsk-Krasnoyarsk,
Bratsk-Taishet and a number of others. Ferrous and non-ferrous metallurgy, pulp and paper, hydrolysis and
saw-milling industries are being established in the Bratsk-Taishet industrial system. The Achinsk-Kras-
noyarsk industrial system is becoming one of the largest centers of aluminum and chemical industries, and
production of ferrous metals, cellulose, paper, and oil products.
Construction of the third metallurgical base has been launched in Siberia, and a new base of ferrous
metallurgy, using the enormous local iron and coal resources, has been created in Kazakhstan. A high-
capacity power system is being organized in the same areas. Non-ferrous metallurgy is being further
developed in Kazakhstan, Central Asia and in Transbaikal areas. The pulp and paper, as well as the timber,
industries are being developed at a fast rate in the forest areas of Siberia and the Far East.
Ferrous metallurgy is also developing in the European part of the country by utilizing the enormous
iron ore resources of the Kursk Magnetic Anomaly and the Ukrainian deposits. Large new production systems
are under construction in the North-West, along the Volga, in the Northern Caucasus and the Ukraine.
(After A. Lavrishchev, "Economic Geography
of the U.S.S.R.", Moscow 1969)
ix
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THE MAJOR INDUSTRIAL CENTERS OF THE USSR
Main centres of ferrous metallurgy
" " " non-ferrous metallurgy
O Centres ol chemical industry
(After A. Efimov, "Soviet Industry", Moscow 1968)
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PRTNCTTPAT. r.RNTRBS (VF FFBBOIK;
rolling
Smelling of ferroalloys
Mining of:
iron ores
coking coal
manganese ore's
MAIM IRON ORE DEPOSITS IN THE U.;...;,.;,,,
(After A. Lavrishchev, "Economic Geography of
the U.S.S.R.", Moscow 1969)
xi
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PRINCIPAL CENTERS OF NON-FERROUS METALLURGY IN THE U.S.S.R.
zi^&SrSSW
DISTRIBUTION OF MOST IMPORTANT DEPOSITS OF NON-FERROUS METAL ORES
Ni Nickel ores
B Bauxites
H Nephelines
A Aluniles
M Mercury ores
Pt Platinum
•I Copper ores
o Tin ores
Complex orei
xii
(After A. Lavrishchev, "Economic Geoeranhv
of the U.S.S.R.", Moscow 1969)
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PRINCIPAL CENTERS OF THE CHEMICAL INDUSTRY IN THE U.S.S.R.
Chemical industry (different branches]
Oil-refining industry
Production of synthetic rubber
O -Production of mineral fertiliser
PRINCIPAL CENTERS OF THE TEXTILE INDUSTRY IN THE
Orekhovo-
Zuyevo
PjvlovsVy Posad
s«,Pukh0v«L_^0,£-
Figures on Ihe map show.
1 Vy\hny t> Rzhev 12 V
Volochok 7 (),-,!!.! 13 Tbiliji
B Vy,i;md 14 Kirovabad
3 Pikov 9 Klmliy IS
4 Kalii>in 10 $n
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PRINCIPAL CENTERS OF WOOD-WORKING AND PAPER INDUSTRIES IN THE U.S.S.R.
'^$5?K?.°moW^g^^=
^<£^.^^mf^^^
Industry:
Timber-sawing and wood-working
@ Paper
C ^ Principal lumbering areas
SH? Foresls
JO Q 500 10001
PPINCIPAL CENTERS OF THE FOOD INDUSTFY IN TrlL U.S.S.F.
(After A. Lavrishchev, "Eoononiio Geoeraohv
of the U.S.S.R.", Moscow 1969)
xiv
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THE MAIN MINING CENTERS OF THE USSR
Oil refining
Oil pipes
Gas pipes
Power stations
(After A. Efimov, "Soviet Industry", Moscow 1968)
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PRINCIPAL ELECTRIC POWER STATIONS AND POWER SYSTEMS IN THE U.S.S.R.
Principal Eleclric Power Stations
Thermal Hydro-power
QC in pperation
^* under construclion
" and planned
@* ^^ Groupi o( electric
power itah'oni
Operating alomic eleclric power stations
Areas ol operation of single power grids
European par! of the U.S.S.R.
Central Siberia
Areas of operation of integrated power grids
North-Weil'
and Wesl
Northern Kazakhstan
Central Asia
YM*»h|'
1 Baltic
2 Narva 10 Dnieproges
3 Keg urn 11 Kakhovka.
4 Plavinas 12 Starobeshevsk
5 NovayaByelorimkaya 13 Zuyevikaya
6 Dubossary 14 Shterovka
7 Kanev 15 Krasnodar
8 Kremenchug 16 Kashira
•j^^xvTTr^v.'.'.'..-.•.•. •'-'-'•'• •'•'•'•'•4-'-'-','.','f-',,v,-/..v-;,';d',',,vf'.:V &?—j-,
Figures mdicale following power stations; LV-l/Vladivcrrto
9 Dnieprodzerzhinik 17 Shatura I'.'.'f— " —-
18 Eleklrogorsk
1? Ivankovo
20 Th'e 22nd C.P.S.U. Congress
HEPS on the Volga
21 The Lenin HEPS on Ihe Volga
22 Chardarinskaya
23 Chirchik-Bozsu
24 Nurek
25 Ragunikaya
26 Var;ob
27 Toktogul
28 Alamedi
360 0
360km
(After A. Lavrishchev, "Economic Geography
of the U.S.S.R.", Moscow 1969)
-------
PHYSIOLOGICAL-BIOCHEMICAL PRINCIPLES OF THE GAS RESISTANCE OF PLANTS
V. S. Nikolayevskiy
From Ministerstvo Vy^iego i Srednego Spetsial'nogo Obrazovaniya RSFSR.
Permskiy ordena trudovogo krasnogo znameni gosudarstvennyy universitet im.
A. M. Gor'kogo. Uchenye Zapiski No. 222. "Gazoustoychivost1 Rasteniy".
Vypusk 1. Perm', P. 5-33, (1969).
Present State of The Problem of Gas Resistance of Plants
The problems of composition of atmospheric air and of effect of toxic
gases on living organisms were debated even by ancient thinkers: Lucretius
describes places near Avernus, Italy, which caused overflying birds to suf-
focate and fall. He gave a completely materialistic explanation for this
phenomenon: poisonous gases escaping from the ground can either spoil the
air or dilute it, causing birds to lose their lift. Areas around Avemus
are rich in sulfur emissions (Mirtov, 1961).
Air pollution as a social problem arose at the beginning of the 14th
century (Halliday, 1962), but the first studies were started at the begin-
ning of the second half of the 19th century. An inevitable by-product of
the development of industry and of civilization in the industrial areas of
the countries of Europe, America and Asia was the appearance of polluted
air and the formation of unproductive and poisoned land, where completely
natural phytocoenoses (association of plants) were either seriously damaged
or completely destroyed.
Enterprises of the ferrous and nonferrous metallurgy, as well as the
chemical industry and the thermal electric power stations emit daily enormous
amounts of various noxious gases (F£ , HF, d.2, S02, CO, ^65, ^63, ^S,
unsaturated hydrocarbons, metal oxides, etc.) and cause considerable damage
to vegetation and to the animal world over vast surrounding areas. Major
nonferrous metallurgical enterprises discharge up to 500 tons of sulfur diox-
ide per day, and each year, the emission of sulfur by industry and furnaces
on the earth amounts to 60 million tons (Katz, quoted by Tomas, 1962).
The composition of noxious gases depends on the design of the enterprise
and on the raw material it processes. The amount of gas discharged is de-
termined by the output of the enterprise. The gas concentration in the
ground layer of air depends on a whole set of weather-climatic and physical-
geographical conditions prevailing in the region and also on the height of
discharge of the gases, as well as on the hermetic tightness of the equipment
and on the processes of dilution and sedimentation (Ryazanov, 1961 b). A
combination of a lower gradient, calm, and an increased atmospheric humidity
— 1 —
-------
produces so-called temperature inversions (Ryazanov, 1961 b; Timofeyeva,
Sadilova, and Kuperman, 1964), during which the gas clings to the ground,
poisoning and destroying everything on its surface. The higher the temp-
erature gradient, the lower the relative humidity of air, the higher the
barometric pressure, and the more intense the solar radiation the
stronger are the vertical air currents, and the greater the angle of open-
ing of the smoke plume, and hence, the more vigorously proceeds the process
of gas dilution.
The degree of the adverse effect of gases and the damage they cause to
agriculture and forestry are very considerable. Thus, in the vicinity of
major nonferrous enterprises and chemical plants, the radius of the air pol-
lution zones is 17-25 km. In some cases, the effect of the gases is detected
at a distance of 40 km or more from the smoke pollution source.
In view of the fact that atmospheric air is a major source of carbon
nutrition of green plants, its pollution has a detrimental effect on their
vital activity. Modern man has artificially created a new ecological factor
that in its impact on plants is no less important than such factors as the
drought or the salinization of soils. For example, the chief cause of the
massive death of coniferous plantings in cities is the heavy pollution of
air caused by the motor vehicle traffic.
Accumulations of gases and dust hang steadily in the atmosphere in the
form of a cap over large cities and over industrial areas. During the morn-
ing and evening hours, the visibility is greatly reduced. At many industrial
enterprises and in the cities of the Soviet Union ("Timofeyeva, Sadilova,
Kuperman, 1964; Nikolayevskiy, Kazantseva, 1966; Il'kun, 196-8), Poland
(Paluch, 1968), the U.S.A. ( Tomas , 1962), and other countries, the air is
polluted with a complex mixture of gases. In the U.S.A. and in England, the
formation and action of smog (a mixture of gas, dust, and fog) has been de-
scribed. A peculiar third type of smog occurring in the Ukraine was de-
scribed by G. N. Il'kun (1968).
The disappearance of vegetation in the vicinity of industrial enter-
prises as well as cases of massive poisoning and death of people, birds,
and animals (63 persons died in Maas, Belgium in 1930; in Donora, Pennsyl-
vania [U.S.A.], in 1948, 6000 people were taken ill and 20 died; in London,
England, in December 1952, a death rate increase of 4000 people as a result
of smog was recorded) have attracted the attention not only of public health
personnel but also of agricultural and forestry experts, botanists, and
phys iologists.
The first studies of the gas resistance of plants were initiated in
Germany in 1861 (Wentzel, 1963), in Russia in 1900-1910 (Nelyubov, 1900,
1910; Sabashnikov, 1911), and in the U.S.A. in 1915-1920 (Fortunatov, 1958).
Studies of the gas resistance of plants in the USSR were particularly
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expanded in the pre- and postwar years. Halliday (1962) estimates that
during the postwar period, a total of about 150 papers devoted to the
problem of gas resistance of plants are published every year throughout the
world. In a number of countries of Central Europe there are periodic inter-
national conferences once every two years devoted to the problem of the
"Influence of Polluted Air on Forests". The last conference was held on
9-14 September 1968 in the city of Katowice, Poland (Ed. note: The proceedings
have been published largely in German and Polish). In the USSR an analo-
gous conference was first held in Sverdlovsk in 1962 at the initiative of
the Commission on the Conservation of Nature of the Ukrainian Branch of
the Academy of Sciences, as well as of the Ural University and the Botani-
cal Garden of the Biology Institute. This conference was held in connection
with successful studies of the gas resistance of plants carried out at the
copper smelting plants of the Central Ural by V. V. Tarchevskiy (1964 a, 1964).
In 1964 and 1966, the second and third conferences were held in Sverdlovsk,
and in 1968, the first Ukrainian conference took place in the city of Donetsk.
In view of the fact that the section of the botanical garden of the
Institute of Biology of the Ukrainian Branch of the Academy of Sciences
which dealt with the topic "Gas Resistance of Plants" was closed in 1966,
we were compelled to transfer our studies to the botanical garden of Perm1
State University (in the city of Perm') where the laboratory of "Experimental
Ecology and Plant Acclimatization" was established. In the Ural, this is the
only botanical garden conducting research on the problem of "influence of
polluted air on the life of plants growing in cities and industrial centers".
Most of the papers in the present volume represent studies conducted by
the laboratory staff in 1967 and 1968.
The investigation of the gas resistance of plants from the ecological
standpoint is being conducted at the Bashkir State University by Yu. Z. Kulagin
(1964 a, b, 1965, 1966 a, b, 1968). In addition, the gas resistance of plants
is being studied in botanical gardens of Dnepropetrovsk State University
(Babkina, 1968 a, b; Gayevaya, 1962), Kiev (Il'kun, Silayeva, Mironova, 1968),
Donetsk (Panov, Rubtsov et al., 1968), Minsk (Getko, 1968), Karaganda
(Sitnikova, 1964, 1966), at the Ukrainian Scientific Research Institute of
the Academy of Municipal Services, Sverdlovsk (Kazantseva, 1965) , at Donetsk
State University (Negrutskaya, 1968; Negrutskiy, 1968), at the Krivoy Rog
Pedagogical Institute (Dobrovol'skiy, 1968), and other institutions.
As was shown by the conferences held in Sverdkovsk (1962, 1964, 1966)
and Donetsk (1968), these studies are being conducted by individual enthus-
iasts and groups of researchers and thus far there are no major institutions
(with the exception of the Scientific Research Institute of Zabrze, Poland)
which have done any extensive or thorough scientific research on the various
aspects of the complex problem of conservation of nature, and on gas resis-
tance of plants in particular. The necessity of establishing an institute
of conservation of nature in the Ural Region is evident to all, since it is
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here that the effect of man on nature is manifested in its most varied forms.
Characteristics of Toxic Compounds and Permissible Air Pollution Norms
Industrial enterprises pollute the air with a great variety of toxic
compounds. The diversity of the raw materials being processed and of the
products obtained is matched by that of the ingredients polluting the bio-
sphere. To date, there has been no scientifically developed classification
for the toxic compounds polluting the air. Such a classification may be
arrived at on the basis of two principles: 1 - physical and chemical compo-
sition of the compounds, and 2 - nature of their toxic action on plants.
From a physical standpoint, they can be divided into gases (simple and
mixed) and vapors and dust (aerosols). Although, thus far, the chemical
mechanism of, their toxic action on plants has been little studied, these
compounds can be arbitrarily divided into the following categories, taking
into account their chemical properties: 1) acid gases (fluorine, chlorine,
sulfur dioxide, carbon monoxide, carbon dioxide, nitrogen and phosphorus
oxides, hydrogen sulfide); 2) vapors of acids (hydrochloric, hydrofluoric,
sulfuric and sulfurous, etc.); 3) oxides of metals (lead, arsenic, selenium,
magnesium, zinc, etc.); 4) alkaline gases (ammonia); 5) vapors of metals
(mercury); 6) various organic gases and carcinogens (saturated and unsatu-
rated hydrocarbons, phenol, carbon tetrachloride, etc.).
The composition of dust may include the most varied compounds formed
during processing of raw materials of both acid and alkaline character, and
also various salts, unburned coal particles, soot, etc.
The nature of the action of various acid gases on plants is apparently
similar (Tomas, 1962; Nikolayevskiy, Kazantseva, 1965) and consists in modi-
fying the physiological-biochemical processes as a result of acidification of
the cell protoplasm. This causes an interruption of photosynthesis, an en-
hancement of the action of oxidizing enzymes, and the onset of photodynamic
oxidations. The action of S(>2 on plants will be examined in more detail below,
and also in a number of other articles.
Acid vapors cause the appearance of scorching or scalding on the surface
of leaves. In this case, the veins of the leaves as well as the generative
organs are damaged.
Like acid gases, ammonia penetrates the leaf mesophyll, and its injurious
effect is due to the alkalinization of the cell environment.
The effect of mercury vapor (Kroker, 1950) and metal oxides on plants
has been little studied, but it may be postulated that the mechanism and
chemistry of their damage to the plants differs from that of acid gases.
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Some metal oxides, for example magnesite dust (Kulagin, 1964 b), cause the
death of woody plants not only because of plugging up the stomata and affect-
ing the photosynthesis, but also as a result of a peculiar salinization of
the soils, the interference with the mineral nutrition of the plants, and
the reduction in the activity of useful microflora.
Hydrocarbons cause epinasty in plants (Nelyubov, 1900, 1910; Kroker,
1950) and other serious impairments of form development processes. A marked
toxicity for plants is shown by many carcinogens (carbon tetrachloride, phenol),
The nature of their action on plants is unknown.
Thus, the toxicity of compounds for plants depends on their chemical
nature. In the action of these compounds one can distinguish (Nikoleyevskiy
and Yatsenko, 1967) both a direct chemical action and a nonspecific action
consisting in a change of the pH of the cell medium. This causes a change of
the ionic equilibrium, as well as a change in the activity of the enzymes
(their action becomes disordered), there is also a decrease in the stability
of proteins and the biocolloids of the cytoplasm and the organelles of the
cell decrease, and the character of plant metabolism is disturbed. The dealth
of cells and tissues may be regarded as the result of advanced and irrever-
sible processes of hydrolysis and photodynamic oxidations.- The action of
individual gases (chlorine) also has its specific aspects, apparently related
to the characteristics of their direct chemical action.
It may be postulated that the nonspecific nature of action is character-
istic of many other compounds (metal oxides, organic acids, phenol, etc.)
that change the pH of the protoplasm. Gas mixtures and smog (Kroker, 1950;
Tomas, 1962; Il'kun, 1968) are marked by a unique and specific character of
action.
In the present paper it is practically impossible to describe all the
toxic compounds, since their assortment increases each year, and the study
of the influence of gases on plants and animals does not keep pace with the
increase in air pollution.
Public health organizations have established permissible air pollution
norms for many toxic compounds (Ryazanov, 1961 b; Uzhov, 1962). The maximum
single permissible norm for pollution of air with sulfur dioxide is 0.5 mg/m3,
and the mean daily norm is 0.15 mg/m3. They are respectively 0.03 mg/m3 and
0.01 mg/m3 for fluorine and 0.1 and 0.03 mg/m3 for chlorine.
Since the mechanism of toxic action of many gases is different in auto-
trophic and heterotrophic organisms (Kroker, 1950), it may be postulated that
for the former, the permissible air pollution norms of the All-Union State
Sanitation Inspection will be unacceptable. Opinions on the permissible air
pollution norms for plants are contradictory. V. A. Ryazanov (1961 b) holds
that they are similar for animals and plants (862), while other investigators
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(Tomas, 1962; Kyabinin, 1962; Nlkolayevskiy, Kazantseva, 1966) believe that
the permissible air pollution norms for plants should be much lower than
for animals.
While single norms of permissible air pollution have been established
for man and animals, they cannot be single for plants of different sensi-
tivity. Moreover, because of the change in the sensitivity of plants to
gases in ontogenesis, the permissible norms even for a given species should
be different at different stages of growth and development. The permissible
air pollution norms for plants should be determined at the stages of heading
and blooming, since at these stages the plants are least resistant to gases.
The fundamental solution to the problem of purity of atmospheric air,
soil and water can be arrived at technologically, but even the most modern
filters cannot completely eliminate the pollution of the environment. The
development of new industries and new technological processes, the use of
new types of raw materials, and the further development of motor transport
will also, in the future, cause serious pollution of the biosphere by toxic
compounds. On the other hand, the total utilization of waste at the majority
of industrial plants has thus far proven economically unprofitable. Thus
the biological method of purification of atmospheric air is of enormous im-
portance. Green vegetation, soil, and the water surface of seas, which
absorb the toxic compounds from air, prevent the poisoning of heterotrophs.
According to the data of Zh. A. Medvedev and Ye. A. Fedorov (1956), in a
closed space, up to 90% of sulfur dioxide is absorbed by plants in one hour,
and the remainder by the soil and the surface of various objects.
Calculations show that the toxic compounds discharged into the atmos-
phere are enough to produce in the biosphere, for 23 years, a concentration
above the maximum single permissible norms.. If one considers that a sub-
stantial part of these compounds spreads through the atmosphere in the
vicinity of the earth's surface (up to 1 km), the indicated concentration can
be formed in the course of a single year. It is only because these compounds
are utilized by plants, the soil, and the water surface of reservoirs that
humanity is saved from self-poisoning, and thus life on earth is able to
continue.
The concentrations of fluorine, chlorine and sulfur dioxide at a number
of enterprises in the Ural (Yefimova, 1964; Timofeyeva, Sadilova, Kuperman,
1964; Sadilova, 1964) exceed the permissible norms set by the All-Uhion State
Sanitation Institute by a factor of 5-10 or more.
General Characteristics of the Noxious Action of Acid Gases
Acid gases have a diverse influence on biogebcoenosis. Essentially,
any of its components is exposed to the action of a new anthropogenic factor.
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Even inanimate objects such as rocks, buildings and equipment are corroded
by acid gases.
Acid compounds in the soil lead to the formation of salts that are
readily washed out and to the loss of important basic elements (Ca, K and
others) by the soil. Despite the incompetence of Wielera's soil decalci-
fication theory (Krasinskiy, 1950), many researchers (Wentzel, 1954;
Nikolayevskiy, 1964) have noted that gases reduce the fertility of soils,
thus decreasing the viability and the gas resistance of plants. The noxious
effect of acid compounds on the soil involves not only the washing out of
important cations and a decrease of the soil absorbing complex, but also an
impairment of the activity of the microflora and other organisms in the
soil. We observed (Nikolayevskiy, 1964) at copper-smelting plants that the
surface layer of imported fertile soil (up to 1 cm) under the influence of
gases and pyrite dust in the course of 1-2 years becomes similar in agro-
chemical properties to the poisoned local soils. After swelling, seeds of
many plants die on such soils (Nikolayevskiy and Suslova, in press). The
absence of vegetation causes a considerable water and wind erosion of the
soils, converting-vast areas into industrial deserts.
Acid gases cause an impairment of growth (Krasinskiy, 1953; Gol'dberg,
1956; Lemke, 1961), a retardation or acceleration of plant development
(Babushkin, 1955; Antipov, 1957, 1960; Yatsenko, Nikolayevskiy, Firger, and
Suslova, 1968), a decrease of the yield (Kroker, 1950; Lemke, 1961; Thomas,
1962), the appearance on leaves of necroses differing in form and color
depending on the types of plants and gases, and a change in the color of
generative organs (Mirtov, 1961). Under the influence of S(>2, a change is
observed in the color of flowers of dame's rocket and of ageratum. This
phenomenon is caused by the acidification of the protoplasm under the in-
fluence of SOo. Anthocyan changes its color under the influence of the pH
of the medium.from blue to pink and even red.
The poisoning of soils impairs root formation in plants (Ryabinin,
1962 a).
The shedding of leaves and needles in woody species (Krasinskiy, 1937,
Nikolayevskiy, 1964) produces a sparse, openwork crown. In some species,
a repeated change of foliage may lead to a disturbance in the correspondence
of the morphophysiological rhythms to seasonal climatic conditions. Under
the influence of gases, the plants' resistance to ecological factors de-
creases, and conversely, the latter may considerably influence the gas
resistance of plants.
Different species of plants have different sensitivities to acid gases.
For this reason, a change in the composition of phytocoenoses is observed
in the vicinity of industrial enterprises: there is a disappearance of
nonresistant species (blue grass, bedstraw, majanthemum, common chickweed,
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plantain, geranium, buttercups), while there is the preservation and a
wider distribution of the resistant ones (Tatarian lettuce, toadflax,
prostrate knotweed, meadow fescue, cinquefoil goosefoot, yarrow, shepherd's-
purse, coltsfoot, horsetail). The quality of fodder grasses in meadows
decreases. Pastures become unsuitable for cattle grazing.
In natural forest stands, one observes a decrease of the bonitet (by
1-2 classes) of the density of the stand, a decrease in the annual increment,
a lowering of soil fertility, and an increase in the activity of pests
(Jahnel, 1954; Lemke, 1961; Schneider and Sierpinski, 1968).
In the vicinity of aluminum plants, domestic cattle come down with
fluorine cachexia because of the high fluorine content of the plants (Bla-
gosklonov et al. , 1967). Medical personnel have observed a decrease in
the immunobiological reactivity of the organism in the juvenile population
living in industrial areas (Navrotskiy,1960), resulting in a considerable
spreading of infectious diseases. In animals (Battan, 1967), smog causes
disturbances of the respiratory and cardiac activities. One of the causes
of the widespread occurrence of cancer diseases today is thought to be the
pollution of air with various carcinogenic agents.
Nature of the Noxious Effect of Acid Gases on Plants
Acid flue gases cause particularly serious and varied damage to green
autotrophic organisms, since the assimilation of S(>2 instead of C02 quickly
leads to the accumulation of toxic products in cells and to a serious im-
pairment of many physiological-biochemical processes. The influence of acid
gases on plants may be divided into direct action on the assimilative appara-
tus and indirect action via poisoning of the soils, deterioration of the
conditions of mineral nutrition, and damaging effect on root systems
(Nikolayevskiy and Yatsenko, 1968).
The degree of injury and hence damage because of gases depends on the
toxicity of the gas, its concentration and duration of action, and the com-
bination of weather-climatic conditions and species of plants. Most toxic
for plants are.F£, C12, S02, and smog (Tomas, 1962). V. Kroker (1950) gives
the following sequence of gas toxicity for plants: Cl2>S02>NH3>HCN>H2S. On
the other hand, their toxicity for animals and man is almost the opposite:
HCN>H2S>Cl2>S02>NH3. This distribution of the degree of toxicity of gases
for plants and animals is undoubtedly due to the nature of their action on
organisms. Compounds acting on the nervous and cardiac systems (HCN and
H2S) are more toxic for animals, whereas compounds decomposing the cell
proteins and bio colloids and disturbing the autotrophic character of metabo-
lism are more toxic for plants.
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Vogl, Bortitz and Polster (1965) distinguish five degrees of injury to
plants by sulfur dioxide depending on its concentration and the duration of
the absorption of gas by the leaves: lack of injuries, latent injuries, or
chronic, acute, and catastrophic injuries. Depending on the plants' sensi-
tivity, under similar environmental and air pollution conditions, different
species will have different injuries.
Under the influence of sulfur dioxide, necrosis blotches of different
colors and shapes appear on plant leaves (Shabliovskiy, 1937). A character-
istic feature is the fact that they are located on leaves between the veins,
and in the case of grasses and conifers, at the tips of the leaves and
needles. The color and shape of the necroses depend on the plant species,
stages of growth and development, age of the leaves, and type of gas and its
concentration in air. Young leaves have darker blotches and old leaves,
lighter ones.
The fall of 4- to 5-year-old needles and frequently three-year-old
needles is observed in coniferous species. Death of the latter (with the
exception of the larch) in cities and at industrial enterprises is due not
only to the injury of the needles, but also to a reduction of the assimilative
surface and to carbohydrate deficiency (Nikolayevskiy, 1966). The increased
survival rate of leaf-bearing species under these conditions is explained by
the regeneration of the leaf apparatus.
The assimilative organs of plants are subject to greater injury, whereas
the generative organs, stems, pedicles and veins of leaves, i.e., organs
which do not participate appreciably in photosynthesis, are injured much less.
Under the influence of acid gases, the yield of the vegetative mass and seeds
decreases, and the seeds are frequently of lower quality and undersized.
Toxic gases and dust sharply reduce the productivity, quality, and
density of plantings, and decrease the increment of trees in height and diam-
eter (Ilyushin, 1953, 1961; Gol'dberg, 1956). American authors (Tomas, 1962)
propose formulas for calculating the yield loss of agricultural crops on the
basis of the concentrations of gases, time of their action, and vulnerability
of leaves.
i
Acid gases and more rigorous microclimatic conditions prevailing in
the areas of industrial plants cause an increase of xerophytization in the
structure of leaves (Nikolayevskiy, 1966 b, 1967). Under industrial condi-
tions, plants have smaller leaves and a larger thickness of the epidermis;
the size of the cells and stomata decreases, and the number of stomata per
mm2 of leaf increases. The reduction of photosynthesis and formation of
metabolites lead to a depression of the elongation stage and formation of
small cells and leaves. On the other hand, xerophytization of leaves in
plants frequently causes an increase in gas resistance (Isachenko, 1938;
Nikolayevskiy, 1967).
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Sulfur dioxide penetrates the leaf to a considerable extent through
the stomata (Lotfild, 1921; Ivanov, 1936; Shabliovskiy, 1937; Krasinskiy,
1950; Jahnel, 1954; Nikolayevskiy, 1963; Kisser, 1968). Under the influ-
ence of S02» the regulation of the movement of stomata is disturbed, the
degree of opening of the stomato decreases (Nikolayevskiy, 1964 b; Vogl,
1964), the intensity of motion of the protoplasm increases, and the perme-
ability is enhanced (Morkovin, 1901). The functioning of the stomatal
apparatus of plants accounts for their slight vulnerability at night,
during dry spells, and in winter in coniferous species, since the stomato
are closed at that time.
Initially, the injuries are concentrated in the spongy parenchyma
near the stomata (Fortunatov, 1958). Acid gases cause the breakdown of
lamellae, swelling and destruction of plastids, piasmolysis, and cell
dehydration (Bredeman, 1933; Jahnel, 1954; Fortunatov, 1958; Il'kun et al.,
1968). The cell nucleus is more resistant to gases than the plastids. In
plants exposed to industrial conditions, the ventilation of the spongy
parenchyma decreases, and sometimes one layer of palisade tissue falls out.
Using a tagged gas, Godzik (1968) found that woody plants differ
markedly in their accumulation of S^Q2~ Autoradiography showed a nonuni-
formity in the absorption of the gas by the leaf. Sulfur dioxide concentrates
in the leaf, epidermis, in the coniferous needle endodermis, and in the water-
conducting systems.
Acid gases cause serious disturbances of the water regime of plants.
Under the influence of S02 transpiration first increases, then decreases
(Morkovin, 1901; Kisser, 1968) as a result of an impairment of the water
supply (Jahnel, 1954). Under the influence of chlorine (Kisser, 1968), the
transpiration decreases at first, then increases.
In view of the specific character of the necroses formed on leaves
(edges in dicotyledons, tips of needles in conifers and of leaves in grasses),
it has been postulated that they are due to the water regime of the plants
(Kisser, 1968; Nikolayevskiy and Suslova, in press). Toxic compounds move
with the water current along the vessels and toward the edges and tips of
the needles and leaves, and concentrate, reaching lethal doses.
Acid gases cause the interruption of photosynthesis in plants (Ivanov,
1936; Thomas and Hill, 1937; Jahnel, 1954). The degree of influence of
gases on photosynthesis and the nature of the aftereffect depend on the con-
centration of the gases. The action of S(>2 disturbs the synthesis of sugars;
pigments, proteins, polysaccharides, and vitamins break down (Krasinskiy,
1950; Jahnel, 1954; Holte, 1958; Fortunatov, 1958).
Particularly contradictory are the views held by researchers concerning
the influence of acid gases on plant respiration: some (Zheleznova-Kaminskaya,
-------
1953; Ilyushin, 1953; Ryabinin, 1962 b) postulate that respiration and the
respiratory quotient increase, while others (Thomas and Hill, 1937; Kras-
inskiy, 1950) hold that S02 has no appreciable influence on plant respiration.
In our view, the contradiction is due to the lack of reliable experiments.
In the majority of cases, the respiration was studied after the lapse of a
considerable period of time since the time of action of S02- Respiration, as
a process most labile and sensitive to changes in the environment, must
undoubtedly undergo some profound changes. We established (Nikolayevskiy,
1968) that S02 alters the chemistry of respiration in plants. In resistant
species (summer-cypress, Eastern poplar, box elder), S02 enhances the role of
the pentose monophosphate oxidation route, whereas in nonresistant species
(European white birch, balsam poplar), the role of the Krebs cycle in res-
piration is enhanced. In view of the major adaptive significance of respira-
tion and its role in the immunity of plants (Rubin, Artsikhovskaya, 1960;
Turkova, 1963; Rubin, Ladygina, 1966) it would be difficult to represent it
as neutral toward acid gases.
Under the influence of acid gases, the activity of enolase, phosphatase,
amylase, and catalase is reduced (Krasinskiy, 1950; Fortunatov, 1958;
Nikolayevskiy, Suslova, 1968), while the activity of peroxidase and poly-
phenol oxidase is increased. We were the first to confirm experimentally
(Nikolayevskiy and Miroshnikova, 1968) K. Noack's theory (1920, 1924) con-
cerning the formation of photodynamic oxidation processes under the influence
of S02.
Sulfur dioxide causes a sharp increase in oxidation processes in both
etiolated and green plants, and only concentrated gas causes inhibition of
luminescence and respiration. Thus, slight S02 concentrations cause an
increase in respiration and ultrafaint luminescence of plants and an increase
in the activity of oxidizing enzymes. High S02 concentrations cause a de-
pression of luminescence, discontinuation of respiration, and a rapid death.
Acid gases and poisoned soils at industrial plants cause a loss of
germinating power in seeds and a disturbance of the rhythms of growth and
development. Usually, the initial stages of development of plants are delayed
(in woody plants - the swelling of buds, in forage grasses - sprouts, third
leaf and tillering), and the subsequent stages are accelerated (Yatsenko et al.
1968; Suslova and Nikolayevskiy, 1968; Firger and Nikolayevskiy, 1968). The
duration of the vegetative stage is shortened in some plants (Antipov, 1957),
and in others increases as a result of second and sometimes multiple leaf
formation. The decreases in the yielding capacity of forage grasses is directly
related to the degree of influence of gases.
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Theories of the Gas Resistance of Plants
By gas resistance (Krasinskiy, 1950), the majority of investigators
mean the ability of plants acted upon by gases to preserve their growth,
development, and ornamental properties.
Many authors have proposed different hypotheses regarding the causes
of the vulnerability and gas resistance of plants. Individual physiological
or biochemical indices or processes are very frequently used in the explana-
tions. At the same time, the diversity and complexity of the mechanism and
chemistry of the effect of acid gases on plants are ignored. This effect is
due to the complexity of the constitution of living matter (plants) and the
marked qualitative diversity between different species as well as within a
species.
The compilation of assortments of gas-resistant plants should have as
its theoretical basis an exact knowledge of the physiological-biochemical
processes responsible for the differences in the vulnerability and gas
resistance of plants. It is particularly interesting and important to know
the chemistry of conversion of sulfur dioxide in plants and its influence
on the metabolism, the oxidation-reduction processes, and the stability of
cell proteins and biocolloids.
i
Haselgof and Lindau (1903) held the cause of necroses and death of
plants to be the interaction with carbohydrates in the leaves, but it was
noted that the stronger plants are injured in the morning, when the leaves
contain less carbohydrates.
Schreder, Reus and Wieler (1912) held that gases act on plants indirectly,
via the soil. They identified the gas resistance with the ability to grow
on poisoned soils. We have established (Nikolayevskiy and Suslova, in press)
that on poisoned soils, even nonresistant species can germinate, whereas
resistant ones may die. Consequently, the survival rate of plants must not
be identified with their gas resistance. On the other hand, despite a con-
siderable indirect damage done by poisoned soils, the cause of injury and
death of the plants in this case appears to be different.
K. Noack (1920) was the first to use the laws of photochemistry in
connection with gas resistance. Sulfur dioxide blocks photosynthesis, but
it does not affect the binding of luminous energy by pigments. The latter
gives rise to photodynamic oxidation processes that result in the formation of
burns on leaves.
Dorris (quoted by Tomas, 1962) held that S02 causes the oxidation of
chlorophyll. The formation of pheophytin leads to the death of the leaf
tissue.
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According to Bleasdale (1932), sulfur dioxide disturbs the equilibrium
in the content of the SO, anion and of sulfhydryl groups in plants. This
leads to a disturbance of metabolism and to leaf injury.
Nemec (1958) held that the accumulation of sulfur dioxide by leaves
promotes the absorption of cations of toxic metals (Zn, Al, Cu, Pb, As,
etc.) from the soil. The latter cause: the conversion of chlorophyll into
pheophytin and the discontinuation of photosynthesis. The death of plants
is the result of both poisoning by these compounds and carbohydrate de-
ficiency.
A major contribution to the development of the gas resistance theory
was made by N. P. Krasinskiy (1937, 1939, 1950) and his students (Knyazeva,
1950; Guseva, 1950, and others). He proposed that gas resistance be
divided into three arbitrary types: 1 - biological resistance consisting
in the ability of plants to rapidly regenerate the damaged foliage, a
feature characteristic of rapidly growing woody species; 2 - anatomical-
morphological resistance, because of leaf structure characteristics that
slow down the rate of gaseous exchange and accumulation of toxic gases;
3 - physiological-biochemical resistance connected with the physiological-
biochemical characteristics of plants. Yu. Z. Kulagin (1968) distinguishes
a fourth type of resistance - anabiotic gas resistance. In the state of
anabiosis, for example, coniferous species are not damaged by gases in
winter.
N. P. Krasiriskiy (1950) showed that plants containing a large quantity
of readily oxidizable organic substances are more extensively damaged as
a result of photodynamic'oxidation processes. He used this to confirm
K. Noack's hypothesis (1920). Krasinskiy (1950) ascribed a major importance
in gas resistance to the reducing properties of the cell content and to
the chemistry of metabolism. He considered gas resistance to be a system-
atic index of the species, genera, and families of plants. The cruciferae
family were found to be the most resistant, and legumes, the least resistant.
V. M. Babkina (1968) holds that evolutionarily young species are more resis-
tant than the old, ancient ones. Gurkal (1959) attributes this to differ-
ences in the nature of metabolism. Plants that are evolutionarily progress-
ive have an unspecialized metabolism (a greater variety of products of
metabolism) and are thus better able to tolerate the action of S02. Ancient
species have a specialized type of metabolism (Blagoveshchenskiy, 1950) and
are therefore less resistant to gases.
The available literature data and reviews (Krasinskiy, 1950; Kroker,
1950; Tomas, 1962; Jahnel, 1954) and also the results of our studies make
it possible to state some preliminary views on the mechanism by which plants
are injured by acid gases and on the causes of differences in the gas
resistance of plants.
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The degree of injury to plants by sulfur dioxide depends on the rate
of absorption of the gas. Injuries in plants occur after a lethal level of
sulfur compounds has been reached (Ivanov, 1936; Krotova, 1957; Jahnel, 1954).
The anatomical-morphological structure of leaves (Rnyazeva, 1950) and the
function of the stomatal apparatus (Nikolayevskiy, 1963, 1964 b), which pro-
mote an increase in the intensity of gas exchange, are at the same time the
cause of the high vulnerability of plants to damage from sulfur dioxide.
The high rate of gaseous exchange in woody plants is provided by the high
degree of aeration of the spongy parenchyma of the leaf (Knyazeva, 1950;
Nikolayevskiy, 1966 b, 1967) the lack of distinct mechanical coatings (cuticle,
hairs), a long period of open stomata in the course of 24 hours, a ratio of
the height of palisade tissue to that of spongy tissue of less than one,
and the presence of stomata on both sides of the leaf.
In contrast to literature data (Knyazeva, 1950; Fortunatov, 1958) we
observed a statistically significant inverse correlation between the quantity
of stomata and the vulnerability (Nikolayevskiy, 1966 b, 1967; Nikolayevskiy
and Firger, in press). A large number of fine stomata and a high degree of
regulation of their movements indicate an increased ecological flexibility of
gas-resistant plants. The leaves of gas-resistant woody plants (box elder,
European spindle tree, snowberry) are subject to a whole series of character-
istics which slow down the rate of gaseous exchange, namely, the presence of
pubescensce or additional coatings (silverberry), a lesser aeration of spongy
parenchyma, the magnitude of the ratio hn/hr, a decrease in the degree of
opening of the stomata during the day.
As established by Ye. I. Rnyazeva (1950) and us, the role of the anatom-
ical-morphological structure of leaves in the gas resistance of plants makes
it necessary to postulate that differences in gas resistance are due to dif-
ferent rates of absorption of acid gases. The same conclusion was reached
by Tomas, (1962).
According to the data of Krotova (1953), the lethal level of sulfur
accumulation by leaves of woody plants is lower in gas-resistant species
than in nonresistant ones (maple 0.3-0.4%, poplar, dogwood, ash 0.75-0.9%).
In forage grasses (Nikolayevskiy and Suslova, in press), in contrast, the
lethal level of sulfur accumulation is frequently higher in the more, resis-
tant species. This characteristic of plants serves as the basis for select-
ing species with an increased gas-absorption capacity (Kulagin, 1968; Getko,
1968) for the purpose of making possible a greater sanitary-hygienic role of
plantings. On the other hand, one can postulate a deeper and theoretically
important relationship between the vulnerability of plants and the intensity
of photosynthesis and respiration, metabolism, and direction of biochemical
processes.
Some researchers (Krotova, 1953) have assumed that resistant plants
are characterized by a higher intensity of photosynthesis. First, they
- 14 -
-------
proceeded from the fact of the low gas resistance of coniferous woody
species, which have (Ivanov, 1936) a low photosynthesis, and, second,
they assumed that the products of photosynthesis (carbohydrates) are
utilized by the plant for detoxieating* the sulfur compounds. Our studies
(Nikolayevskiy, 1963, 1967) showed a direct correlation between the vulner-
ability and the'rate of photosynthesis (r = 0.79). Resistant species
(box elder) are characterized by a decreased rate of photosynthesis, and
nonresistant species (European white birch, Manchurian walnut, etc.) - by
a faster photosynthesis rate.
Subsequently, the cause of the low gas resistance of coniferous species
to the action of gases was cleared up (Nikolayevskiy, 1966 a). It was found
that the coniferous species were more resistant than the leaf-bearing ones
to a single action of even high S(>2 concentrations, with the exception, of
the genus Larix. Their low.survival rate in cities is because the coniferous
needles have a long lifespan and hence are exposed to the action of gases
for a longer time. A reduction of the assimilative surface (because of
shedding of the old coniferous needles) leads to the death of the plants as
a result of carbohydrate starvation.
The dependence of the gas resistance of plants on the intensity of
gaseous exchange is confirmed, on the one hand, by the anatomical-morpholog-
ical structure of the assimilative organs of the plants, and on the other
hand, by examples of reduction of plant vulnerability in the shade, the
absence of injury at night and during the winter season, and also the absence
of injuries in organs and parts not taking any appreciable part in photo-
synthesis (petioles, leaf veins, young shoots, and the reproductive organs).
The low intensity of photosynthesis explains the high resistance of succulents
(echeveria, sedum). According to literature data (Brilliant, 1951), the
intensity of photosynthesis in the cruciferae is 1.5 times lower than in
legumes. Tomas (1962) and Il'kun (1968) also hold that differences in the
gas resistance of plants are primarily due to differences in the rates of
absorption of sulfur dioxide.
An important role in gas resistance is played by the direction of the
physiological-biochemical processes. N. P. Krasinskiy (1950) showed that
resistant plants are characterized by increased reducing properties of the
protoplasm and a higher content of glutathione. V. A. Guseva (1950) observed
that resistant plants are characterized by a lower oxidation-reduction poten-
tial (ORP) and aerobicity index rH2. Our studies (Nikolayevskiy, 1965 a)
confirmed Guseva's data (1950). In addition, a direct relationship was ob-
served between the vulnerability of plants to sulfur dioxide and the content
of ascorbic acid in the leaves. It is well known that a high ascorbic acid
content correlates in plants with a high oxidation activity (Rubin, Spiri-
donova, 1940). According to the data of L. Michaelis (1936) and T. F. Lupareva
* [Translator's note: "intoxicating" in Russian original.]
- 15 -
-------
(1958)-, an equality of the oxidation and reduction processes in plants is
observed when index rH2 = 13-14. For gas-resistant woody plants, rH2 is
found to be 13-15, and for nonres1stant ones, 16-20. Hence, resistant
plants are characterized by a lower intensity of photosynthesis, a low
ascorbic acid content, and a relatively high activity of reduction processes-.
Experiments (Nikolayevskiy and Suslova, 1968) established that under the
influence of SC>2 and I^SO^ the resistant plants show, at first, an increase
of the reducing properties, while in the case of nonresistant species one
finds an increase in the oxidizing properties of the protoplasm. Krasinskiy's
hypothesis (1950) concerning the role of oxidizable substances in gas resis-
tance requires an experimental biochemical verification. Many physiologists
do not accept his "oxidizability" index because of the roughness of the
determination and biochemical vagueness.
A study of the content of substances oxidizable with 0.1 N KMnO^ in
leaves by Krasinskiy's method (1950) confirmed his conclusions concerning
the dependence of gas resistance on the quantity of oxidizable substances.
The correlation coefficient for 47 leaf-bearing woody plants was 0.54 at
X = 7.7>3. It was postulated (Nikolayevskiy, 1963) that the content of
mr
oxidizable substances in leaves is related to the intensity and direction
of the physiological processes. It thus becomes possible to understand,
on the one hand, the causes of the diverse gas resistance of plants (differ-
ences in photosynthesis rates) and, on the other hand, the causes of the
relationship between the vulnerability of plants and the quantity of sub-
stances in leaves oxidizable by potassium permanganate. Species differences
in the intensity of photosynthesis in plants are of course related to the
activity of oxidation-reduction enzymes. It may be assumed that the activity
of photodynamic oxidations in plants under the influence of S02 will depend
on the activity of the oxidizing enzymes of plants.
Of major importance for the gas resistance of plants should be the
nature of metabolism (Nemec, 1957). Gurkal (1959) observed that resistant
forms of Norway spruce contained much higher amounts of ct-pinene in their
needles. The positive role of pentose phosphate type oxidation in the
drought resistance (Abrarov and Petinov, 1964) and immunity (Rubin and
Zeleneva, 1964) of plants is explained by the formation of compounds with
increased reducing properties. Apparently, the same role is played by the
apotomic oxidation route in the respiration of gas resistant plants under
the influence of S02 (Nikolayevskiy, 1968).
The chemistry of transformation of sulfur dioxide in plant leaves is
complex and thus far unclear, since S02 may be simultaneously an oxidant and
a reductant. It is possible that the nature of the transformation of S02
and its influence on the metabolism of plants of different resistances will
be different. It is thought (Kroker, 1950; Yang and Mbu, 1961; Tomas, 1962)
that S02 is oxidized to the sulfate and is partially tied up by organic
- 16 -
-------
compounds. As a result, the toxicity of SOo decreases by a factor of 30.
Simultaneously with the oxidation of sulfite, sulfurous and sulfuric acids
are formed, acidifying the protoplasm (Krasinskiy, 1950).
In the case of sulfur deficiency in the soil (Kroker, 1950), the
sulfur is rapidly included in the composition of amino acids and proteins
in reduced form. This accounts for the beneficial effect of SO- on plants
in areas of the USA with a low sulfur content of the soil.
The reduction of sulfate proceeds in light (Doman, 1957) through
oxidation of the carbohydrates (Sabinin, 1955; Kaliniyevich, 1959). Nemec
(1957) and Tomas (1962) emphasize that carbohydrates also have a beneficial
influence on the gas resistance of plants.
Thomas and Hill (1937) observed that sulfate is excreted into the
soil by the root systems of plants that have been exposed to S02. It is
possible that this is the way in which plants can regulate the sulfur content
of tissues and avoid toxic concentrations.
A study of the water regime showed that woody plants resistant to S02
(box elder, elder, etc.) are characterized by an increased water content
and water-holding capacity, a low concentration of cell content, and a mod-
erate content of bound water (Nikolayevskiy, 1965 b). For plants not
resistant to sulfur dioxide (white birch, Siberian crabapple, etc.), the
opposite is true.
A great similarity is frequently observed in the action of certain
acid gases on green plants. Rohmeder and Schonborn (1965) note that plants
resistant to SO^ display a high resistance to fluorine. This was also
observed by us (Nikolayevskiy and Yatsenko, 1968; Firger and Nikolayevskiy,
1968) at the Perm1 Chemical Plant in 1966-1967.
According to the studies made by Ye. N. Kazantseva (1965), forage and
herbaceous ornamental plants resistant to fluorine are characterized by
similar anatomical-morphological and physiological characteristics, as are
also the S02~resistant woody plants. This appears to be a manifestation of
the above-indicated nonspecificity(of the action of acid gases on plants,
leading to a disturbance of the dark reactions of photosynthesis and to the
appearance of photodynamic oxidation processes in light (Nikolayevskiy and
Kazantseva, 1966).
The survival rate of plants under industrial conditions is determined
not only by their gas resistance, but also by the action of other ecological
factors: frosts, droughts, or soil salinization. Ilyushin (1953), Krotova
(1957), Nikolayevskiy (1964), and Kulagin (1965) hold that drought- and
frost-resistant plants also display a higher gas resistance. The drought
resistance of the silverberry is explained by the presence of special
- 17 -
-------
umbellate excrescences of the cuticle which cover entirely both surfaces
of the leaf and protect it from overheating; they also decrease the evapor-
ation of moisture. The high resistance of the silverberry to S02 and F is
explained by the fact that the excrescences of the cuticle slow down the
rate of gaseous exchange and absorption of toxic compounds.
Of interest is the fact that the water-holding capacity is a universal
indicator of the resistance of plants to various extreme factors (low temper-
atures, droughts, gases). Hence, the resistance of plants requires the pres-
ervation of the structure and functions of the proteins and biocolloids of the
protoplasm and organelles of the cell.
In our view (Nikolayevskiy, 1965 a), the striking analogy in the resis-
tance of plants of three families to different factors is not accidental.
N. P. Krasinskiy (1950) observed a high resistance of the cruciferae and low
resistance of legumes to SC^- Cereals occupy an intermediate position in
their resistance to S(>2. The same was observed in the case of their tolerance
of ionizing radiations (Vasil'yev, 1962; Preobrazhenskaya, and Timofeyev-
Resovskiy, 1962).
The sensitivity to ionizing radiation is associated with a high intensity
of vital processes (Vasil'yev» 1962). The analogy in the behavior of plants ,
'toward the action of different factors may be attributed to the fact that
the gas and radiation resistance is based on the intensity and direction of
the physiological-biochemical processes. At the basis of the biochemical
processes of injury to plants by x rays (Dubinin, 1961) and gases (K. Noack,'
1920; Krasinskiy, 1950) lies a disturbance of the processes of accumulation
of luminous energy and photosynthesis, and the appearance of similar photo-
dynamic oxidations.
Yu. Z. Kulagin (1965, 1966 a) successfully developed an ecological
concept of gas resistance, whereby the degree of plant resistance is determ-
ined, in addition to everything else, by the presence or absence of critical
periods. If the critical period in plants happends to coincide with the
action of a gas, even relatively resistant species may die, and conversely,
nonresistant plants which have excaped injury during critical periods may
survive.
Experience in compiling assortments of gas-resistant plants (Nikolayevskiy
and Kazantseva, 1965; Nikolayevskiy and Firger, 1969, in press) and their
comparisons with similar lists of other authors (Shabliovskiy, 1937; Isachenko,
1938; Krasinskiy and Knyazeva, 1950; Zheleznova-Kaminskaya, 1953; Deroyan,
1957; Kuntsevich and Turchinskaya, 1957; Bulgakov, 1958; Vanifatov, 1958;
Podzorov, 1961; Gayevaya, 1962; Ryabinin, 1962; Sitnikova, 1966; V. M. Babkina,
1968) show that it is impossible to obtain universal lists. The physico- ;
geographical conditions of individual regions may exert a substantial influence
on the anatomical-morphological and physiological-biochemical characteristics
- 18 -
-------
and gas resistance of plants. A considerable geographical variability of
gas resistance is observed in the box elder. Under the conditions of the
Ural and Moscow regions, the box elder is considered to be a gas-resistant
species, but in Karaganda, Transcaucasia, the Donets Basin and Dnepropetrovsk,
it was found to be nonresistant. These areas differ in their moisture regime.
Drought conditions (Karaganda, Dnepropetrovsk) complicate the water regime
of plants and decrease the resistance of the protoplasm proteins, causing a
decrease in gas resistance.
The literature gives indications of the important ro],e of mineral nutri-
tion in the gas resistance of plants (Nemec, 1957; Kazantseva, 1965; Kisser,
1968; Yatsenko et al., 1968), and of the moisture regime (Jahnel, 1954),
illumination, and temperature (Kroker, 1950; Krasinskiy, 1950).* A given com-
bination of ecological conditions may foster an increase of the gas concen-
tration in the ground layer of air. In addition, by reducing the general
vital capacity of plants, frosts and high temperatures may bring about a high
injury rate even in the case of low gas concentrations.
Thus, the resistance of plants to acid gases depends on the following
conditions: (a) combination of the complex of ecological factors at the time
of the action of gases, (b) biological characteristics of the species and
their ecological flexibility, (c) stages of plant growth and development,
(d) anatomical-morphological structure of the leaves, (e) intensity of gaseous
exchange, (f) stability of the proteins and biocolloids 'of the cell, (g) degree
of disorganization caused by the gases in metabolism and enzyme activity,
(h) nature and intensity of photodynamic oxidation processes, and (i) the
buffering capacity of the cell content and other factors.
For certain specific gases and geographical areas, many authors recom-
mend different assortments of gas resistant plants for use around industrial
enterprises. More than 20 lists of recommended plants have been proposed
in the USSR. The gas resistance criteria used are (a) the plant vulnerability
index (Krasinskiy, 1937, 1950; Antipov, 1959), (b) the survival rate under
unfavorable conditions (Bulgakov, 1958), (c) the quantity of oxidizable sub-
stances in the leaves (Vanifatov, 1958; Antipov, 1968), and (d) the effect
of gases on the physiological and growth processes of plants (Sitnikova, 1964,
1966).
» Some of the other papers presented in this volume, which deal with studies on the gas resistance of
plants in relation to the problems of mineral nutrition, metabolism of C1* carbon compounds under the
influence of SCL, and influence of the gas on the pigment systems and ultrafaint luminescence of
plants, nay serve as a supplement to the survey reported in this paper.
- 19 -
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Conclusion
The present study gives a survey of the principal advances in research
on the gas resistance of plants in the USSR and in the countries of Europe
and America. In a survey of this kind it is practically impossible to give
a detailed review of all the published studies dealing with this topic.*
We have attempted to mention the key points that are conducive in some measure
to the elucidation of the causes of plant vulnerability and gas resistance.
Since the beginning of these investigations, important data - both from the
theoretical and the practical point of view - have been obtained which are
conducive to the clarification of the causes of the diverse resistance of
plants. Thus, the influence of toxic compounds on the anatomical-morphological
structure of leaves and the physiological-biochemical characteristics of
plants have been studied. Also, the species of plants resistant to various
compounds in different physico-geographical zones have been ascertained.
Many investigators have made a serious contribution to the development of the
theory of plant resistance to acid gases. In several countries (U.S.A.,
USSR, Poland), investigators have undertaken detailed studies of the chemistry
of injuries to plants by acid gases as well as of the chemical mechanism of
transformation of S02 in plants and its influence on the metabolism of car
bon compounds with the aid of modern isotopic methods of analysis using
and C^. Major advances have been made in the study of the resistance of
plants to sulfur dioxide, while at the same time insufficient attention has
been given to the study of the resistance of plants to chlorine, hydrogen
sulf'ide, hydrocarbons, carcinogenic compounds and to the other ingredients
which pollute the atmospheric air. There have been practically no investi-
gations into the influence of the chemical compounds on the microflora of
soils.
Of major importance for a successful diagnosis and study of the gas
resistance of plants is the establishment of direct indicators characteriz-
ing the degree of plant resistance. We have given considerable attention
to the development of methods for studying the gas resistance of plants.
Two methodical studies have been especially devoted to this problem.
Research carried out in the last two years has shown that under the
influence of fertilizers, forage grasses do not always clearly display the
relationship between the change in the conteftt of ascorbic acid, oxidizable
substances, water-holding capacity, and gas resistance.** Apparently, these
indicators do not reflect the direct relationship to the gas resistance, but
are indirect, describing the intensity of physiological-biochemical processes
and the stability of protoplasmic biocolloids. This once more underscores
the necessity of studying the mechanism and chemistry of injury to plants by
- Some of the remaining studies of this volume supplement the present paper to some extent.
•* See the paper of Yatsenko and Nikolayevskiy in this volume.
-------
gases and the causes of the diverse gas resistance of plants. The establish-
ment of direct indicators of gas resistance will aid in a correct appraisal
of the resistance of individual plant species.
The. present day status of the problem of plant gas resistance in the
USSR and other countries (Transactions of the 6th International Conference
on the Problem of "Effect of Smoke Pollution of Air on Forests," 1968) and
the lack of a unified, generally accepted theory of plant gas resistance
point up the urgent necessity of:
1 - Studying the physiological-biochemical bases of the gas resistance
of plants;
2 - Studying the chemistry of transformation of toxic compounds in
plants and the mechanism of their detoxication*;
3 - Developing methods of increasing the gas resistance of plants;
4 - Studying the sanitary-hygienic role of plants.
)
On the basis of our studies, to characterize the gas resistance, we can
recommend the determination of certain indicators correlating with the
resistance:
1 - Anatomical-morphological structure of leaves (number of stomata,
thickness of epidermis, aeration of spongy tissue, daily dynamics of the
movement of stomata), which characterize the rate of absorption of acid
gases;
2 - Ascorbic acid content, which characterizes the rate of metabolism
and the activity of oxidation processes;
3 - Water-holding capacity, characterizing the hydrophily and resistance
of proteins and biocolloids of the protoplasm;
4 - Quantity of oxidizable substances in the leaves, using N. P. Kras-
inskiy's method (1950);
5 - Vulnerability of plants to gases under both industrial and experi-
mental conditions;
6 - Determination of the lethal and sublethal doses of absorption of
sulfur dioxide by means of a tagged gas (s35o2).
We have developed a new method of determining the aeration of the spongy
parenchyma of leaves by means of vacuum infiltration.
[Translator's note: "intoxication" in the Russian original. ]
21 -
-------
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- 23 -
-------
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EFFECT OF MINERAL NUTRITION ON THE GAS RESISTANCE
OF FORAGE GRASSES
V. M. Yatsenko, V. S. Nikolayevskiy, V. V. Firger, and V. V. Suslova
Perm1 University
From Akademiya Nauk Ukrainskoy SSR. Tsentral'nyy respublikanskiy botan-
icheskiy sad. Donetskiy botanicheskiy sad. Materialy Pervoy Ukrainskoy
konferentsii "Rasteniya i promyshlennaya sreda". Izdatel'stvo "Naukova
dumka", Kiev, p. 88-94, (1968).
A successful planting of greenery on the grounds of industrial enter-
prises discharging noxious gases is not dependent solely on sound selection
of an assortment of gas-resistant plants. Of late, research men and practi-
cal gardeners have come to the conclusion that it is necessary to develop
special agro-technical methods of fertilizer application in order to produce
more viable and resistant plantings and compositions C^useva, 1950; Krasinskiy,
1950; Wentzel, 1959; Materna, 1963). Traditional agro-technical methods of
plant cultivation, borrowed from the practices of farming and forestry, have
-proven relatively unsuccessful and inadequate. The highly unfavorable ecolog-
ical conditions for plants at industrial sites make it necessary to test old
methods (Guseva, 1950; Materna, 1963) as well as, in the main, new agro-
technical methods (Bobko and Fortunatov, 1958) and ways of growing plants
and increasing their gas resistance.
In 1967, we attempted to use a field method for studying the influence
of the elements of mineral nutrition on the growth, development and gas
resistance of forage grasses (a) under the conditions prevailing at two in-
dustrial enterprises (a chemical plant emitting S02, F, HF, and nitrogen
oxides, and a nonferrous metallurgical plant discharging Cl2 and 802) and
(b) under experimental conditions at the Botanical Garden of Perm University.
We used an 8-treatment experimental scheme (control, N, P, K, NP, NK, PK, NPK).
The resistant species chosen for this investigation was meadow fescue (Festuca
pratensis Huds) and the non- res is tent species chosen were timothy grass
(Phle.um pratense L.). .
Prior to sowing, fertilizers were applied to the plots in the following
proportions: K - 9g, N - 4g, and P - 6g per 1 m2. At the Botanical Garden,
the plants (10 in each test glass) were fumigated with sulfur dioxide in
chambers (SO- concentration from 0.03% to 0.4% by volume, 1 hour). The vulner-
ability was determined by linear measurement pf necroses in the second leaf
(from the top) in ten replications. At the end of the summer, the yield '
(plant height and number of leaves) was calculated for each treatment. In '/
addition, foliar nutrition (fertilizer spray applications) tests were conducted
at the Botanical Garden to determine the effect of the same fertilizers and of
- 28 -
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certain physiologically active substances on the gas resistance of the
plants. Since the effect of fertilizers in a field experiment may be
levelled by the action of other factors, a periodic dressing of the plants
with fertilizers was carried out at the Botanical Garden according to the
same scheme and norms, starting on 12 July and their influence on the vul-
nerability and on certain biochemical characteristics of the plants (quantity
of substances undergoing oxidation and of ascorbic acid, oxidation-reduction
potential, and content of organic acids) was determined.
The studies showed that in the course of the growing period, the gas
resistance is maximum at the stage of emergence of the seedlings and at
the stage of 3 leaves, then decreases, and slightly increases at the end
of August. This is apparently due to the intensification and character of
metabolism in the middle of summer in the fescue and in the stages of booting,
heading, and blossoming of timothy grass. It is very likely that the differ-
ences of gas resistance between meadow fescue and timothy grass are due to
the biological characteristics of the growth and development of these species.
During the first year, the fescue ends its growth with the tillering stage
and blossoming takes place in the second year, whereas the timothy grass
completes during the first year its full life cycle. Because of an intensi-
fication of metabolism, the gas resistance of timothy grass is sharply re-
duced during the heading and blossoming stages.
In meadow fescue at the chemical plant, the course of the developmental
stages lags 1-3 days behind the Botanical Garden plants for the emergence of
seedlings stage; 6-9 days for the third-leaf stage, and 9-12 days for the
tillering stage. Thus, under the Influence of acid gases and poisoned soils,
the growth and development of the fescue at the chemical plant are retarded,
and this lag increases at each stage. In timothy grass, a lag also takes
place during the first three stages of growth and development (emergence
1-2 days, third leaf 5-9 days, tillering 5-9 days). However, a substantial
acceleration of development (6-12 days) is observed as early as during the
booting stage. The acceleration of development in timothy grass is preserved
during the heading stage in treatments N, P, K, and NK. A retardation of
development recurs in the remaining treatments and stages.
Whereas in the Botanical Garden, timothy grass passed through all the
growth-development stages up to the ripening of seed in all the treatments
of the experiment, such was not the case at the chemical plant where timothy
completed its life cycle only in treatments N, P, K, and NK. In the remain-
ing treatments it did not go on to the formation of the reproductive organs.
Hence, as in the case of the meadow fescue, under the influence of acid
gases and poisoned soils, a retardation of development is observed in timothy
grass during the first three stages near the chemical plant, followed by a
substantial acceleration of its development during subsequent stages. In the
control timothy plants, growing near the chemical plant, the life cycle can-
not be completed in a one-year period. In treatments with fertilizers, the
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plants also did not reach the blossoming stage. Nitrogen, phosphorus,
potassium, and nitrogen-potassium cause an acceleration of the growth and
development at the heading stage and the termination of the life cycle of
timothy grass in one year.
Table 1
Influence of Mineral Nutrition on the Gas Resistance of Forage Grasses
Treat-
ments
"
Vulnerability of Plant
Meadow fescue
Botanical
garden
Control ICO
N 113
P 109
K 155
Up 109
Bk 39
pK 132
Npk 15^,7
Chemical
plant
100
100
125
%
'*7,7
CO?
129
128
Metal-
lurgical
plant
100
64,3
101
92
-------
influence exerted on the plants by the poisoned soil of the industrial areas.
Analyses showed that the soils near the chemical plant have a substantially
reduced content of .mobile forms of nitrogen, phosphorus and potassium.
An experimental study of the influence of mineral fertilizers on gas
resistance as determined from the growth and development stages showed that
in meadow fescue, even in the course of a single stage (tillering), a suc-
cession of the positively acting nutrient elements is observed in different
calendar periods, i.e., June-July-August. In timothy grass, the same
phenomenon can be observed in accordance with the growth and developmental
stages. In meadow fescue, depending on the calendar periods, N, P, K, NP,
and NK were found to have a positive effect on the gas resistance, and in
the case of timothy grass, NP, N, K, and NK. The succession of nutrient
elements having a positive effect on gas resistance 'in plant ontogenesis is
completely regular. The mineral requirements of plants are a function of
their growth and development and of the biological characteristics of the
species.
The positive effect of fertilizers on plant growth (height of shoots,
length of leaves) also changes in plant ontogenesis. In the fescue, at the
beginning of summer, an increase in leaf size was observed in all the treat-
ments. In the middle of summer, the fertilizers caused a decrease in stem
growth and an increase in leaf growth, whereas at the beginning of August
the fertilizers decreased the leaf growth, and in the middle of August the
positive effect of fertilizers on the growth of the plants was manifested
again.
In timothy grass at the beginning of summer, almost all the treatments
produced a positive effect on the growth of the plants. This was followed
by a depression of shoot growth and an enhancement of leaf growth in treat-
ments N, P, and K.
In gas resistance of fescue in the first year of life (tillering),
the most important role is played by nitrogen (Botanical Garden) and potas-
sium (chemical plant), and by nitrogen in the case of timothy grass. The
positive role played by nitrogen in the gas resistance and also in the growth
and development of plants is explained by the important part which it plays
in their protein metabolism.
Experiments with foliar nutrition showed that the gas resistance in
fescue is increased by potassium, and in timothy grass by N, NK, and K.
A positive effect in fescue was produced with potassium ferricyanide and
citric and malic acids; and in timothy grass, in addition, by oxalic acid*
NaOH, boron, molybdenum, and hydroquinone. A negative effect was produced
in both species by P, PK, NPK, CaC03, ZnS04, MgS04, FeS04, KMn04, and H2S04.
The positive influence of organic acids on the gas resistance of forage
- 31 -
-------
plants can be explained by their important role in plant respiration (tri-
carboxylic acid cycle). The adverse effect of sulfates and H2SO. on plants
can be explained by the additional negative influence of the SO^ anion.
Potassium permanganate is a strong oxidant, and hydroquinone, a strong
reductant. That is why the former increased the vulnerability of the plants,
while the latter decreased it in timothy grass. The positive effect of po-
tassium ferricyanide on plant resistance may be attributed to its role as
an inhibitor of oxidation processes of sulfhydryl groups of many enzymes and
proteins.
Conclusions
1. Under the influence of acid gases and poisoned soils, the development
of fescue and timothy growing near a chemical plant is retarded.
2. Fertilizers can be used to regulate the growth and development of
plants and their gas resistance.
3. The role of individual nutrient elements (N, P, K and their combin-
ations) depends on the biological characteristics of the species, as well as
on gas resistance, on the stages of growth and development, on the nature of
soils, and on the weather and climatic conditions of the season. A positive
influence of nitrogen on the gas resistance of forage plants was observed in
connection with its important role in protein metabolism.
4. The gas resistance of plants can be increased by means of physi-
ologicklly active compounds and fertilizers applied by the foliage nutrition
method. Potassium ferricyanide, hydroquinone, NaOH, and malic and citric
acids were found to have a positive influence.
- 32 -
-------
LITERATURE CITED
BodKO E.B., 4>oprynaTOB M.K., 1958. OHUT xwuimecKOti <5opb-
Ga c ycKxaKHea JiecoHaca/r,«eHan B ropoacxoK oScTanoBKe. Hasec-
s-un T.C.X.A., » 6.
rycesa B.A. 1950. BjinnHne limiepanfcHoro nKTamw HB OKHCJIH-
Te^BHO BOCCTaHOBKICJlLHUri PC3UI1 It raSOyCTOitHHBOCTB paCTCHHfi.
U.-P.
EpacKucicHtt H.n. 1950. TeopeiiiiecKJie OCHODU nocTpoemin
pacTGHnii. Tau xe.
Katerna J. 1963. ZTy5ovani odolnosti drcvin proli
Acinkum kourovych plynu hngenim. "Pt-ace vyatumn. tiatot-u
Icon. CSSR". IS63 N 26.
ffcntsel K.P. I?'J9. £ur Dodenbeelnfluseiinc durch
ifinotriallc lAift*oi-unrelnigungen tmd Dungung In
Rouchschadenologen inoboaoudere mit Kolu Foret. uaa Holzniit,
14,H8.
- 33 -
-------
EFFECT OF MINERAL NUTRITION ON CERTAIN PHYSIOLOGICAL-BIOCHEMICAL CHARACTERISTICS
AND GAS RESISTANCE OF FORAGE GRASSES
V. M. Yatsenko and V. S. Nikolayevskiy
From Ministerstvo Vysshego i Srednego Spetsial'nogo Obrazovanlya RSFSR.
Pennskiy ordena trudovogo krasnogo znameni gosudarstvennyy universitet im.
A. M. Gor'kogo. Uchenye Zapiski No. 222. "Gazoustoychivost1 Rasteniy".
Vypusk 1. Perm1, p. 69-84, (1969).
Alongside with the selection of an assortment of resistant plants,
the development of methods of increasing the gas resistance of plants
assumes a great importance since many industrial enterprises have grounds
where not only woody but also forage plants are unable to grow. Acid gases
and toxic dust are accumulated by the soil, decreasing its fertility (Wentzel,
1959) and curtailing the activity of the useful microflora (Kulagin, 1964).
V. Kroker (1950) noted that the roots of plants sometimes suffer more ser-
iously from industrial pollutants than do the leaves. Inhibition of the
growth of root hairs can be observed on poisoned soils (Ryabinin, 1965).
All this impairs the mineral nutrition of plants and leads to the accumu-
lation of certain toxic elements - Zn, Pb, Cu (Nemec, 1958) and to a decrease
in the friability and resistance of plants to extreme factors (Kulagin, 1968).
! •.
In order to improve the conditions of growth and development of plants,
practical gardeners change the topsoil regularly, i.e., once every 2 to 3
years, at the copper-smelting plants of the Middle Urals (Krasnoural'sk,
Kirovograd, Revda). We established earlier (Nikolayevskiy, 1964) that
within 2-3 years fertile soils imported to the copper-smelting plants become
quite similar in their agrochemical properties to those of the local poisoned
soils.
In the Soviet Union and abroad (Guseva, 1950; Mashinskiy, 1953; Nemec,
1957; Freebairn and Taylor, 1960; Kazantseva, 1965; Ryabinin, 1965; Kisser,
1968) studies were conducted on various methods of increasing the gas resist-
ance of plants. Soil neutralization (liming), application of fertilizers
to the soil (K, P, N, Ca), leaf nutrition, and the action of various compounds
such as OP-7, sucrose, urea, and potassium ascorbate were found to have a
positive effect. The literature contains data to the effect that plants are
capable of adapting to the action of acid gases (Levitskaya, 1955; Bulgakov,
1958). On the other hand, methods of growing the plants (Mashinskiy, 1953)
and the experience of practical gardeners indicate that the gas resistance
of plants can be increased by means of advanced agricultural field practices.
The role of individual elements of mineral nutrition (N, P, and K) in
plant life varies. The biochemical role of nitrogen in plants is related to
- 34 -
-------
the important part that it plays in protein and nucleic metabolism, which
determines the plants' physiological-biochemical and genetic characteristics.
The anion of phosphoric acid enters into the composition of many important
organic compounds (phospholipids and phosphatides, ATP and ADP, etc.). It
participates in many energetic processes that take place in the cells. Potas-
sium in plants regulates the ionic and water regime of the cell and the perme-
ability of the cell membranes. Both the individual elements of mineral nutri-
tion and their definite combination are important for the plants as a function
of their growth and development. The deficiency of certain elements in the
soil causes an impairment of metabolism and of the form-development processes,
and this may be reflected in the gas resistance of plants.
Our studies (Yatsenko, Nikolayevskiy, Firger, and Suslova, 1968), both
under experimental conditions in the university botanical garden and under
field conditions (at industrial plants) showed that nitrogen fertilizers in-
crease the gas resistance of plants, whereas phosphorus and complete fertil-
izers (NPK) decrease it. Of great interest is the study of the nature of the
influence of fertilizers on the physiological-biochemical characteristics of
plants for the purpose of determining the mechanism of their effect on gas
resistance. With the exception of an article by Guseva (1950), the literature
offers no data on this problem. However, Guseva does not give the results
for all types of fertilizers, but only for two forms of nitrogen, fertilizers
(NaN03 and NH^SO^).
In 1967, in the botanical garden, we employed a field method in a study
of the influence of mineral nutrition on certain physiological-biochemical
characteristics and gas resistance of forage grasses.
Method of Investigation
Two species of forage grasses differing in gas resistance were pre-
selected for the studies: the meadow fescue (Festuca pratensis Huds) as the
resistant species and timothy (Phleum pratense L.) as the nonresistant species.
The seed was sown on poor sandy soils on May 7, 1967 in accordance with an
8-treatment scheme: control, N, P, K, NP, NK, PK, and NPK in three replica-
tions. The size of the plots was 2 m x 2 m. Before the seeding, fertilizers
were applied in the proportion of N 4 g, P 6 g, and K 9 g per m*. Starting on
July 10, the plants were watered once a week with solutions of fertilizers in
the same proportions; this was followed by a study in relation to (a) their
vulnerability in a gas chamber, (b) the quantity of oxidizable substances in
the leaves, according to the method of N. P. Krasinskiy (1950), (c) the con-
tent of ascorbic acid, after Sapozhkova (1966), (d) the total acidity, after
Yermakov and Arasimovich (1952), the oxidation-reduction potential on LPU-0.01,
and (e) the water-holding capacity, after Nichiporovich (1926).
35 -
-------
Effect of Mineral Nutrition on the; Vulnerability
Phases, Dates,
Treatments
Tillering
lol
t
si
|i
£S
§,|
II
Booting
= 1
ll
— CO
— !
Average Plant \
Vulnerability \
ll
«2
and
Heading
I
I
Average Plant
Vulnerability
Timothy
Control
N
P
K
NP
NK
PK
NPK
Control
N
P
K
NP
UK
PK
NPK
20,0
23,0
43,4
25.0
18.0
45.5
34.0
590
13.2
12.0
15.0
39.0
5.0
17.0
33.0
29.0
75.7
82.0
81,4
81.8
88.5
87,2
91.8
95?
74.0
67.5
73.7
G9.2
79.0
67.5
81.0
77.5
85.3
89.0
85.3
81,5
88,8
89,2
90.6
843
82,0
87,8
86.0
84.2
79.2
81.2
91.7
88.0
60.3
64.7
70.0
628
65.1
74,0
72.1
795
56,4
55,8
58.2
64,1
54,4
55,2
C8.6
64.8
64,9
57,6
78,2
604
57,2
60,7
67.2
671
60.1
69,4
47.2
53.4
62,1
55,0
64.9
57,5
92,0
97.0
95,0
905
97.5
96,0
97,0
93.7
48,6
75,4
79,8
75,3
80.5
80,5
77,5
74.8
71,3 76.1
69,5 74.7
78,0 83.7
46.7 65.9
76.4
77.0
72,7 76.5
70.2 78.1
64.3 75.0
22,9 43.9
58.0 67,6
48.2 .58,4
43,7 57.4
53.4 65.3
40.7 58.7
47.9 63.4
•15,5 59,3
58,8
80.7
54,4
76,2
61.7
87.6
90.9
91.4
76,7
78.9
68,7
69.4
63,6
53,6
71,2
50,9
77.8
87,9
73,3 89,2
77.9 96,3
68.7 70.3
66.0 92.4
67,2
94,1
90.9
54,1
45,1
55,4
91,0
74,8
81.1
76,2
71.7
73.4
81,9
88,0 91.0
90,1 9f.:,8
Meadow Fes-
63.4
55.3
64,4
35,2 64,3
59.4
44,3
61,2 59,8
56.2 75,7
66.0
74,9
64.7
59,8
62,8
56,3
55.8
58,9
67.7
63.9
- 36 -
-------
Table 1
of Forage Grasses to Sulfur Dioxide
SC>2 Concentrations
Blooming
Is1
Ripening
>o
So"
— in
>
$•0
— CO
II
0.-H
IS
Aftermath
IMCO
C-IWJ
55
o
&£
3 «•
Grass
97,2 32,4 49,4 41,4 55.0 17,9 25.2 35,4 57,2 33,9 10.9 18,3 18,1 15,8
96,7 72,3 45,2 74,2 72,1 18,0 39,4 17,3 77,3 38.0 16.3 36.1 50,0 34,1
95,9 32,1 82,3 55,3 66,4 10,3 78,6 24,7 35,4 37,2 — - — —
51,2 27,0 85,0 50,5 53,4 7,9 51,0 19,4 27.2 26,4 47,2 43.2 26,0 38,8
73,2 70,8 47,8 11,2 50,8 3,1 48,4 17,0 38,7 26,8 35,3 40,1 28,6 34,7
84,6 31,1 97,3 33,6 61,6 4,4 41.5 33,3 59,0 34.6 — — —
82,7 19,4 80,0 36.5 54,6 21,0 74,4 55,0 58.0 52,1 8,9 17,1 17,3 14,4
79,5 33,9 94,3 11,8 54,9 37,2 9I,,3 57,7 76.4 65.6 40,0 21,6 18,1 26.6
cue - tillering
67,3 41,5 15,6 65,3 47,4 5,2 53,8 71,4 39,9 42,6 30.8 34.5 11,5 25,6
73,0 36,4 19,5 70,5 49,8 4,8 39,4 71,2 57,2 43,2 24,8 43,0 50,1 39.3
71,9 48,6 22.0 56,9 49,8 3,2 10.7 22,9 54.4 22,8 — — —
71,4 55,2 55,5 18.6 50,2 7,4 45,2 19,8 77,1 37,4 46,2 49.8 16,1 37.4
57,2 43,6 30,8 52,4 46,0 3,7 9,2 74,7 52,3 35,0 41.3 25,3 45.2 37,3
5I.C 42,7 31,8 18,2 36,1 11,8 12,4 47,8 3.0 18,8 — — —
57.G 33,6 48.4 48,6 47,0 15,1 26,8 64,1 44,9 37.7 36,6 24.2 12.1 24,4
K3.1 51.9 48,8 30.5 48,6 18,6 49,2 17,6 74,7 40.0 35,0 11,2 17,1 21.1
- 37 -
-------
We selected these indicators because earlier studies (Krasinskiy,
1950; Guseva, 1950; Nikolayevskiy, 1967) established a correlation between
these indicators of plants and their gas resistance. The 862 concentration
in the polyethylene gas chamber was preselected so as to bring out the
differences in the vulnerability of the two types of forage grasses. Sulfur
dioxide was obtained through the interaction of a weighed amount of NA2SO.J
with excess 50% I^SO^. The amount of ^£§03 was calculated from the equation
Na2S(>3 + H2S04 = Na2S(>4 + S02 + K^O. To obtain an initial S02 concentration
of 1/4000 by volume, 0.118 g of Na2S03 was taken; for 1/2000, 0.236 g; for
1/1000, 0.472 g; for 1/500, 0.964 g of Na2S03.
The vulnerability was determined as the ratio of the damaged length of
the leaf to the total length in percent. In each treatment of each of the
two species, the vulnerability was determined in ten plants in the first
three leaves from the top of the shoot. The results were treated statis-
tically by Leont'yev's method (1961).
Results of the Investigation
The species of forage grasses whose gas resistance we studied differ
in the biology of growth and development. The meadow fescue is a perennial
winter grass, whereas the timothy grass is a "dvuruchki"* (Fedbrov, 1968).
For this reason, in the first year of life in the case of the meadow fescue,
the tests were conducted only during its tillering stage, while in the case
of the timothy grass tests were conducted also during the stages of booting
(shooting), heading, blooming., and ripening, and in both species after the
hay has been cut, i.e., during the period of young aftermath.
In order to bring out more clearly the role of mineral nutrition in the
gas resistance of the plants, the latter were sown on poor sandy soils
(Table 2).
As is evident from Table 2, the soils of the test area are relatively
acid and characterized by an insufficient content of nitrogen and potassium,
but are comparatively well-supplied with phosphorus.
The weather-climate conditions of the summer of 1967 (Fig. 1) were
favorable for the growth and development of forage grasses, with the excep-
tion of August, when an increased solar radiation and irregular precipitation
* [Editor's note: Fedorov (1968) states that on the basis of the pattern of their growth-development
perennial grasses can be divided into two groups, namely:
(a) winter grasses — defined as those grasses which do not bear fruit during the year they.
are sown, and which do not produce a second bloom after hay cutting!
(b) the so-called "dyuruchfei" — defined as those grasses which can bear fruit during the year
they are sown, and which can produce a second bloom after the hay has been cot in early summer.
The growth-development pattern of these grasses resembles that of the grain crops.]
-------
were observed. It is known from literature data (Ivanov, 1936; Kroker,
1950; Krasinskiy, 1950) that an increased solar radiation promotes a
greater vulnerability of plants to sulfur dioxide. That is why a change
in the vulnerability of the plants (Table 1) was observed in the meadow
fescue even during a single stage of growth and development (tillering).
The decrease in the gas resistance of the meadow fescue during its first
year of life may also be attributed to an increase in the calendar age of
the leaves.
Table 2
Agroehemical Characteristics of Soils of the Test Area
No. of Soil
Cross Section
•fcg
.c'l-H
ll
pH
-p
c
o X
'&£
O-HCT
if!
4>
0)
i«*
CO ID-H
rH«&
HJ O q>
2£*
g
•H
g^jR
+J to*
™'g
0)
eo 3
t< bo
e§"8
-------
3 " a 3i it 2' *» •* » >' •« »« u 10 It SO If ft Si
May
June
July
August September October
Fig. l. Meteorological conditions of the vegetation period of 1967
in Perm1.
Notation: 1 - relative humidity of air, #; 2 - mean daily air
temperature, °C; 3 - duration of solar radiance, hours;
4 - precipitation, mm.
during the tillering stage in the meadow fescue. At the beginning of summer,
an increase in the size of the plants was brought about by P, NK, PK, and
NPK, and an increase in leaf size was noted in all the treatments. In the
middle of summer, the fertilizers caused a retardation of stem growth and a
more vigorous leaf growth. At the beginning of August, the fertilizers
caused a decrease in the rate of leaf growth, and in the middle of August,
the positive effect of fertilizers on the growth of shoots (N, K, UK) and
leaves (N, NP, NPK) reappeared.
The gas resistance of the meadow fescue during its first year of life
(tillering) was greatly influenced by nitrogen. The positive role of
nitrogen in gas resistance and also in the growth and development of the
plants is explained by its major importance in the protein and nucleic me-
tabolism of the plants.
In the course of the annual cycle, the resistance of the timothy
(Table 1) first increases toward the booting stage, then decreases (heading,
blooming) and again increases for a short time during ripening. Hence, the
gas resistance of the plants is not a constant indicator, and varies with
the growth and development.
If one compares the action of fertilizers on the gas resistance of the
timothy according to the stages of growth and development, one finds that
nitrogen increases the gas resistance during the first 2-3 days after the
- 40 -
-------
application of fertilizers at all the stages except heading, and that a
decrease in resistance occurs on the 4th-5th day. Phosphorus caused a
decrease in the resistance of the timothy at all the stages except booting
(2nd day) and ripening.
The influence of potassium on the timothy was similar in character to
that of nitrogen. Among mixed fertilizers, a positive effect on the gas
resistance of the timothy was displayed by NP and NK (blooming and ripening),
and PK (heading). The positive influence of fertilizers on the stem growth
of the timothy is observed at the beginning of tillering, and on the leaf
growth, at the stages of tillering, heading, and blooming. Various fertil-
izers had a positive influence on the gas resistance of the fescue and
timothy, but in both species we noted the positive role of potassium, and in
the fescue, that of nitrogen as well. These fertilizers cause a similar
increase in stem and leaf growth in both species.
Table 3 shows the dependence of the vulnerability of the plant leaves
on the age, i.e. , leaf position irt reference to height of plant, under
the influence of fertilizers.
In the control treatment, the timothy shows a distinct basipetal direc-
tion in the vulnerability of the leaves; the fescue shows it less distinctly.
Under the influence of fertilizers applied on 29 July 1967 by spraying*, the
timothy retained the differences in leaf vulnerability according to leaf
position in reference to height, whereas in the meadow fescue, the leaf vul-
nerability increased in the acropetal direction (with the exception of po-
tassium). This was also observed when the fertilizers were introduced into
the soil, but in this case the exception in the meadow fescue was NPK.
Statistical treatment of the differences in the vulnerability of the
plants under the influence of the fertilizers showed that in the timothy,
close-to-significant differences were produced by N, K, NK and PK, and in the
meadow fescue, by N, NP, NK, and PK.
The change in the amount of oxidizable substances in the leaves of
forage grasses during ontogeny and the changes taking place under the influ-
ence of the fertilizers are presented in Table 4.
It is evident from Table 4 that during the summer, the content of
oxidizable substances increases with age in the meadow fescue (tillering
stage) and in the timothy, when the latter enters the reproductive stage
(heading, blooming), whereas these substances decrease during the ripening
stage. In the fescue, even during the tillering stage, the amount of oxi-
dizable substances increases, and in the second half of August, decreases
again. The aftermath of both species was found to be more resistant than
* [Editor's note: foliar nutrition. J
- 41 -
-------
Table 3
Vulnerability of Leaves of Different Ages Acted Upon by Fertilizers and S02
Date
•H I
• CDC
Treatments
Control
N
P
K
SP 1 NK 1 PK
NPK
Timothy
29/VII
1/VH1
1
2
3
4
I
2
33.8
25,7
41.7
75,3
30.4
45,6
38,2
,41,0
40.7
53,3
28,9
38,6
35,8
55,3
28,7
—
—
Meadow
29/VII
1/VIH
1
2
3
I
2
8.4
13.6
11,5
58,5
56.6
25.8
13.1
14,7
28.4
40.7
32.5
8.6
—
' —
—
44.7
30,8
51,0
89,0
22.3
31,6
fescue
5.5
12,0
14,0
20,8
13.9
33.3
30,4
45.1
61,4
35.4
89,0
20.7
12,2
—
17.7
16.1
20.7
25.1
47.7
—
—
—
43,0
21.4
—
—
—
32.8
47.5
54.1
82.8
28.6
45.4
14,7
15.2
—
8.9
3.0
43.2
52.6
58.2
64.3
28.3
34.7
18,5
16.9
—
33.2
55,5
the adult plants. In the timothy, the amount of oxidizable substances and
the vulnerability are greater than in the fescue.
A positive effect, i.e., a decrease in the content of oxidizable sub-
stances in the timothy was observed under the influence of K. The same
treatment also produced a higher gas resistance in the timothy. In the
meadow fescue, a similar effect was produced by K, whereas N, PK and NPK
caused an increased vulnerability and raised the content of oxidizable
substances. The change in the latter under the influence of fertilizers
does not always correspond .clearly to the change in vulnerability.
In ascorbic acid content, the timothy differs markedly, from the fescue
(Table 5). As in the case of woody plants, the nonresistant species -
timothy - is characterized by a higher ascorbic acid content than the fescue.
In both species, the ontogeny shows a direct relationship between the ascorbic
acid content and the vulnerability of the plants to sulfur dioxide. However,
as in the case of the content of oxidizable substances, the change in the
ascorbic acid content under the influence of fertilizers is not always clearly
related to the change in the vulnerability of the plants. In the timothy, a
decrease in the amount of ascorbic acid and vulnerability is detected only
under the influence of NP.
- 42 -
-------
Table
Effect of Mineral Nutrition on Differentiated Oxidizability.
Stages
and
Periods
Control
Water-
Soluble
Substance.
Water-
Insoluble
Substance
*j
Vulnerabi
N ^
8
.35
Sf*
J!
o> o in
$
Vulnerabi
K
(A
r Water-
Soluble
| Substance
ul
• Water-
Insoluble
Substance
*
i
1 Vulnerabi
NP
at
Water-
' Soluble
Substance
in
was
r-ta
-P mo
$
r~t
| Vulnerabi
PK
in
Water-
Soluble
Substance
OT
Water-
Insoluble
Substance
4J
Vulnerabi
NPK
(A
Water-
Soluble
Substance
in
Water-
Insoluble
Substance
Vulnerabi
1
u>
1
Tillering
7— 10/VI1
Booting
12— U7VII
Heading
18— 25/VIt
Blooming
28/VII—
9/VIII
Ripening
10— 17/VII1
Aftenaath
22-31/VIII
Iill«ring
7— 10/Vl'l
12— I4/VII
18-25/VII
28/VII—
9/VIII
10— 17/V
Aftermath
22-31/VIII
Timothy
3,6
4,3
7,2
3.8
3,8
4,6
1.3
2.5
3.4
3.3
2.6
2.2'
78,4
71,3
55,1
55,0
33,9
19,3
3,8
3.9
6.1
2.4
.3,3
3,6
1.1
1.0
2.6
2,8
3.0
2,1
77,3
69,5
72,8
50,2
38,0
22,1
3,8
3.4
5.6 .
2,8
3,9
2.9
0.8 75,4 5,0 1,4 77.2 4,1
I.I 46.3 4.6 -1.2 76.1 3.5
2,8 47,8 3,8 3.0 68,4 6.1
2,2 11,8 2,8 2.4 6,3 3,6
2,5 42,2 3,8 2,4 33,8 4,1
2,6 32.6 2,9 1,8 28.8 3,2
1,6
1.8
3.2
2.4
2.0
2,6
81,2
70,2
56,8
2(5,4
50,8
14.3
4.4
4.1
5,4
4.8
4.3
2.3
1.4
M<
2,8
3.2
2.5
2.4
80.4
, 84.3
62,4
13,2
(>5,5
27,9
Meadow fescue
3.0
2,7
2.4
2.9
2,0
2,1.
0.9
1.1
1.3
2.7
2,2
1,8
54.4
49.8
61.0
11.8
55,0
24,8
3,3
2.9
3.2
2,7
2,5
2,4
0.7
1,1
1.4
2.3
1.9
1,9
72.4
68,4
54,0
15,1
55,9
33,2
3.5
3.6
4.0
1,9
2.5
2,2
0.9 64,4 4.4 0,9. 71,3 4,1
1.3 56.6 3,1 1,0 58,5 3,0
1.5 53,3 2,1 1,4 58,3 2,9
1,8 6,6 3,0 2,4 6,5 2,2
1,5 47,4 2,1 1,8 45.4 2.4
1,9 42,0 2,1, 1,9 33,4 2,2
0.6
0.9
' 1.5
2,9
1.8
1,8
71.2
59,6
56.9
16,8
45.3
27,6
3.8
3.1
3.0
2.8
1.9
2,3
0.5
1.0
1.3
2,3
2.2
1,9
66,2
48,2
64.6
18.8
47.2
20,5
-------
Table 5
Effect of Mineral Nutrition on the Ascorbic Acid Content of Leaves.
Stage'
and
Periods
Control
• o
t
W
<«< •
Vulnera-
bility -1
N
O *rt *
O-H
Ul O
«*.<< -
Vulnera-
bility .
K
Ascorbic
4s&
ft
NP
1 Ascorbic
Acid
&
33
9 *H
> ft
PK
Ascorbic
Acid
i*
13.
:> .0
NPK
o
• H
t,
OH
010
<«
1
I
»H
s».Q
" Timothy ' x
Tdllering
7-10/VH
Booting
12— 14/VII
Heading
18-25/VII
Blooding
9/VIIf
Ripening •
10— I7/VI1I
Aftermath
22-31/VIU
Tillering
7 -10/VII
I-J-H/VII
18-25/Vll
a*/Vli—
D/VIII
10-1 7/V
27,5
27,9
30,2
31,4
17,4
21,3
20,5
14,5
14,7
12,9
12,2
64,9
65,0
73,8
42.7
38.9
19,0
54,4
49.8
61,8
29.6
55.0
32,0
«19,2
28,5
24,8
19,7
21,8
17,0
20,0
13,9
12,0
31,9
57,6
75,1
82,4
39,4
44,7
27,7
Meadow
72,4
(58,4
57,8
35,1
55,9
24,0
\
21,0
27,5
23,3
15,4
20,3
fescue'
16.0
13,7
11.2
10,9
13,3
60,4
61,2
77,0
24.7
32,5
33,7
64,4
56.6
57,0
10,6
47.4
27,0
22,4
30,4
25,3
17,5
21.2
17,8
15,2
12,9
12,3
10,6
57,2
68,9
69,9
5,2
34.7
28.8
I
71.3
58.5
53,6
21.8
45.4
24,0
21,4
31,7
23,7
19.2
-
21.0
16.5
13.5
10.6
11,3
10.2
67,2
80,6
76,7
24,7
C2.5
14.9
71,2
59.0
63.2
27.4
45.3
30,0
23,7
30,1
22,4
21.8
20,4
17.0
13.5
12.3
10.4
11,8
67,1
77,8
81,1
21,2
75, U
2(5,0
Cifl.2
48.2
(i8.0
22.7
47.2
12.3
22.2
13,5
3(5,6
17.3 36,8 14,2 35,7 14.3 24.5 l.r..:» 19.K
-------
Table 6
Effect of Mineral Nutrition on Total Acidity.
Stage*.
and
Periods
Control
fi %
*«
K
31
O 0'
EH td
PK
E-l •*
1 >>
U -P
4) *p4
5,7!
3 J3
NPK
O O
^ jj>
q?*H
CrH
a' a
Timothy
Tillffring
7— 10/Vlf
Booting
12— 14/VII
Heading
18-25/VII
1 ' Blooming
.p. 28/V1I—
w 9/VIII
1 Ripening
10-l7/\nil
Aftermath
22— 31/VII1
3.3
1,0
4,7
0,8
0,6
1,1
92,0
58,8
55,1
55.0
39,3
19.1
3.0
1.5
2.0
0,5
0.8
2.0
97,0
80,7
74.8
•50.2
41.7
27.7
3,0
1.0
1.2
0,5
0.5
2,0
90,5
76,0
47,6
33,1
32.5
33,7
3,3
1.0
1.3
0,4
0.6
1.8
97.5
GI.7
G8.4
12.4
34,7
28,8
3.6
1.3
1.9
0,6
0.5
1,8
97,0
90,7
56.8
26.4
(i().7
14.9
2.8
1.3
1,3
0.5
0,9
1,6
93.7
91,4
62.4
13,2
51.7
26,0
Meadow fescii'e' *
Tillering
7— 10/VII
12— 14/VII
18-25/VII
28/VII—
9/VII1
10-I7/V
Aftermath
22— 3I/VIII
2.1
1.6
0.6
0.5
0,5
1.9
48,6
7(5,7
47,8
29,6
55,0
22.2
2.8
2.1
0,0
.0.5
0,6
2,3
75.4
78,9
40,8
35,1
55,9
36,6
3.4
1.8
0.5
0.4
0.5
2,0
75.3
(»,4
41,9
10,6
39,8
36,8
3.6
I.I
0.5
0.4
0.4
2,0
«0.5
(i,'l,G
57,3
21,8
53.7
35.7
2.8
1.8
0.5
0,6
0,5
2,1
77.5
71.2
44,9
27,4
45.3
24.5
2.8
2.1
0,6
0.4
0,6
2.0
7-1.8
50,9
59,0
22.7
47,2
19,8
-------
Table 7
Dynamics of the Oxidation-Reduction Potential and Gas Resistance During the Vegetation Period Under
the Influence of Fertilizers.
&
I
Stages
and
Periods
. Control
I'll
Eh
rlla
i
g>>
•s-s
3-H
>.o
N
pll
Eli
HI,
i
2 >>
aa
5 a
K | NI>K
pl-l
Eh
rH8
S
4 J
3-3, P"
3>r4 1
> A
Hh
rl-U
A,
15
3 *rl
f> .0
Heading
I7-24/VI1 (5.81
Ripening
10—J7/VIII 5.61
Aftermath
21—30/VIII 6,10
Tillering
17-2-J/VII 0.-I7
27/VII—9/VIII «M4
10-17/VI11 0,22
Aftermath
2I-3J/V1H 5,96
0,273 22,91 73,8
0,280 23,62 42,7
(UN I8.IX) 46,3
0,2-14 20,66 19,0'
0,259 21,84 64,7
0,258 21,74 21,7
0.248 21,00 62.6
Timothy
7,00 0,285 23,79 82.4
6,73 0.271 22,73 39,6
5.76 0.218 19,08 47,3
6,16 0,247 20,83 27,7
Meadow fescue
6.61 0,263 22,28 59.8
0,42 0,257 21,63 22.2
6,25 0,250 21,10 55,3
7,05 0,292 24,11 80,1 7,13 0,295 24.32 81,1
6.51 0,263 22.05 24.7 6,62 0.207 22.40 21,2
6,10 0.242 20,55 23,3 6,30 0,255 21,35 07,0
6,17 0,244 20.71 33,7 6,29 0.256 21.37 2G.O
6,80 0,276 23,07 56,3 6.60 0,261 22.1A ftj.Q
6.54 0,262 22,08 20.6 6.42 0.257 21.03 29/J
6,18 0.245 20,80 32,5 (>,42 0.200 21.74 33.4
0,233 19,93 22,2 6,23 0,250 21,04 36.6 6,00 0.235 20,08 36.8 6.28 0,255 21.30 19.8
-------
Table 8
Effect of Mineral Nutrition on the Water-Holding Capacity of Leaves
(Percent of Water Held for 2 Hours).
Control
Waters-Holding
[Capacity, %
Vulnerability
N
Water->Holding
Capacity, %
Vulnerability
K
Water-Holding
Capacity, %
Vulnerability
NP
Water-Holding
Capacity, %
Vulnerability
PK
Water-Holding
Capacity, %
i
Vulnerability
NPK
Water-Holding
Capacity, %
Vulnerability
Timothy
11 July — Tillering Stage
39.9 92,0 35,4 97,0 49,8 90,5 42,2 97,5 49,1 97,0 52,0 93,7
13 July —Booting Stage
62,0 58.8 70.9 80,7 59,6 76,2 61,9 6l',7 63,9 90,9 67,4 91.4
Meadow fescue
11 July — Tillering Stage
45.3 48,6 46,4 75,4 55,9 79,8 44,2 75,3 48,2 80,5 44,9 80.5
13 July.— Tillering Stage
58.6 22,9 61.0 58.0 72,9 43,7 57,9 53,4 52,9 47,9 62,0 45.5
In forage grasses, a decrease of the total acidity with age is observed
during the ontogeny. The resistant species - meadow fescue - is characterized
by a lower total acidity than the timothy. At the same time, the seasonal
dynamics of the total acidity and vulnerability are reversed as compared with
the differences between the species, i.e., a decrease of total acidity in both
species is associated with an increased vulnerability. Fertilizers caused an
increase of the total acidity in the meadow fescue and a decrease in the
timothy. No distinct relationship was observed between the change of the
total acidity and vulnerability under the influence of the fertilizers.
A study of the oxidation-reduction potential showed that in the meadow
fescue and timothy, the values of the protoplasm pH, Eh and rH2 do not re-
flect any relationship to the gas resistance.
The water-holding capacity, which characterizes the resistance of proto-
plasmic biocolloids, shows a direct correlation to the gas resistance of
plants (Nikolayevskiy, 1967). In the meadow fescue and timothy (Table 8), a
direct relationship was observed between the gas resistance of the plants
and their water-holding capacity (control, 11 and 13 July). There was a sig-
nificant difference in water-holding capacity between the species as well.
In the meadow fescue, an increase of the water^holding capacity was caused by
K, and in the timothy, by K (11 July), PK, and NPK. However, the increase in
the water-holding capacity under the influence of the fertilizers is frequently
associated with an increase in vulnerability; this is difficult to explain,
- 47 -
-------
considering the correlation between the water-holding capacity and vulner-
ability of plants (Nikolayevskiy, 1967).
Conclusions
1. The studies established the feasibility of regulating the gas
resistance of plants by means of mineral nutrition. A positive effect was
obtained in forage grasses by the application of nitrogen and potassium.
The positive effect of nitrogen is apparently due to its important role in
protein metabolism, and that of potassium, to its role in the regulation
of the permeability and ionic regime of the protoplasm.
2. During ontogeny, because of a change in the plants' requirements
for the various nutritional elements, a regular succession of the types of
fertilizers that have a positive effect on the gas resistance is observed.
3. .As established by a number of investigators, the indicators of
plant gas resistance (oxidizability, ascorbic acid, water-holding capacity.
oxidat-ion-reduction potential) can be used to characterize species differ-
ences in the gas resistance of forage grasses.
Under the influence, of fertilizers, the change of these indicators
does not always exactly coincide with the change in the gas resistance.
This makes it necessary to postulate that they are connected only indirectly
with the mechanism of plant resistance.
- 48 -
-------
LITERATURE CITED
B y .1 r a K o B M. B. 1958. OHUT oaeJiciicimsi r. Kpacnoypa.ii.CKa. C6. «Ma-
Tcpiia.ibi no osc.ienoiiiKO ropoAoo Vpa.ia», own. I, VHMH AKX. Cuepa.ioBCK
Fyccsa B. A. I960. B.iiiflinie Miiiicpa.iwioro iniTaiiun na OKiic.iHTeJibno-
noccTaiioBiiTc.ii.Hbifi pc>KiiM ii rasoycToiViiiBOCTb pacTcimfl. C6. ^IbiMoycToii-
•IHBOCTl. paCTCIIHH II AblMOyCTOUMHnUC aCCOpTHMCHTU». AKX PCCP H FopbK.
yii-T. MocKsa — FopbKiifi.
E p M a K o B A. C., A p a c H M o B H M B. B. H Ap. 1952. MCTOHH CIIO.XHMH-
•iccKoro nccne.iOBaiiiin pacTcuiifi. Ccjtbxosriia.
H B a H o D A. A. 1936. 4»i3iio;ionifl pacreniifi. Foc.'iec6yMii3AaT.
KpaciiHCKiifi H. n. 1950. TcopcTH'tccKiie ocnoau nocrpoeiinfl accop-
TUMCIITOB rasoycToft'iiinbix pacTeiniii. C6. «JluMoycTOfi'ii!BOCTi> pacTeiiiiii « flti-
MoycTofiHiiBbie accopniMeiiTH*. AKX PCCP 11 FopbK. yn-T, MocKBa — Topb-
XHH.
Ky.ianiii K). 3. 1964. flbiMonuc OTXOJIH sanoaa ManieaiiT H
.ICCOB ae.npiion soiiu ropo^a CBTKH (lOxiiufi Ypa.i). C6. «O.xpa«a npiipoAbi >ia
Vpa.ie», Bbin. 4, VAH, CaepA-iOBCK.
KyjiarHii K). 3. 1968, O raaoycTOiVntBocTH apcDccuwx pacreiniM H 6no-
.••onmccKofi OMHCTKC arMocuafl npoMbiui.iciiiiocrw, A's 5.
eAopo n A. K. 1968. Biio^onin Miioro.icTinix rpan. HS.I-BO «Kofloc», M.
51 u c n K o B. M., H n K o .1 ac B c K H ft B. C., * n p r e p B. B., C y c .1 o -
aa B. B. 1968. B.inmiue Miiiicpa.iwioro niiTamin iia ra.ioycTOiViiiBocTb rawu-
nux Tpan. MaTcpna.;n.i nepBofi VKpaHiicKoii Koii^cpciiuiiii «PacTcnna n npo-
MMiii.icininn cpcaa». «HayK«na ,a>'MKa», KHCB.
Kisser J. 19C8. Physlologischc Plroblcme dcr Einwirkung von Luftvc-
unrcinigungen auf die Vegetation. Matcrialy VI Micdzynarodowcj konfercn-
cji. Kotowice.
N e m c c A. 1957. Studio o kourovych skodach na icsinch porostech
v okoli papirun v Ceske Kamcnici. Lcsnictvi, N I.
Nemec A. 1958. Vliv konri1 a popilku na intoxikari smrkovich porostu.
Lcsnictvi, N 5.
-------
EXPERIENCE IN THE USE OF THE BIOCHEMOLUMINESCENCE METHOD
FOR DIAGNOSING THE GAS RESISTANCE OF PLANTS
V. S. Nikolayevskiy and A. G. Miroshnikova
Perm' University
From Akademiya Nauk Ukrainskoy SSR. Tsentral'nyy respublikanskiy botan-
icheskiy sad. Donetskiy botanicheskiy sad. Materlaly Pervoy Ukrainskoy
konferentsii "Rasteniya I promyshlennaya sreda". Izdatel'stvo "Naukova
dumka", Kiev, p. 115-120, (1968).
The hypotheses of K. Hoaka and N. P. Krasinskiy (1950) explain the
formation of gas burns on leaves by oxidation processes in chlorophyll-
bearing plant tissues. Krasinskiy*s proposed method of determining the
amount of oxidizable substances in plants in an acid medium, which he iden-
tified with the activity of oxidation processes in plants under the influ-
ence of sulfur dioxide and light, is not very reliable. This method is
biochemically primitive. While it enables one to Judge the nature of the
oxidation processes in plant cells under the influence of sulfur dioxide
it does not permit an identification of the biochemical composition of the
substances undergoing oxidation or the ascertaining of the sequence and de-
gree of. their oxidation.
The appearance and development of the new biochemoluminescence method
in biophysics (Tartusov, Ivanov, and Petrusevich, 1967) opens up new oppor-
tunities for the study and interpretation of the biochemical aspect of oxi-
dation processes in plants under the influence of sulfur dioxide. A set-of
electronic instruments with a high sensitivity FEU-42 photomultiplier are
used to record spontaneous and induced oxidation processes in plants. The
extremely faint luminescence detected by the photomultiplier is amplified
with an USh-2 amplifier and recorded with a pulse counter.
According to present day understanding (Tarusov et al., 1967), the
ultrafaint luminescence of plants is due to spontaneous oxidation processes
in the cell biolipids. Many oxidation processes in cells have been found
to be associated with luminescence. Despite the fact that only a small
number of the molecules that participate in the reaction, luminesce, the lumin-
escence reflects the kinetics of the given reaction. Consequently, the
measurement of chemoluminescence will permit the determination of kinetic
parameters and the interpretation of the mechanism of oxidation processes.
It was found (Agaverdiyev, Tarusov, 1965) that the luminescent activity of
plants reacts sharply to the slightest shifts in ecological conditions
(temperature, composition of air, humidity, etc.).
- 50 -
-------
Since the ultrafaint luminescence is recorded by the photomultiplier
only in etiolated plants, we attempted to study the differences in the
ultrafaint luminescence of plants and certain biochemical characteristics
(12 species of forage plants) and also the change of ultrafaint luminescence
under the steady influence of different concentrations of sulfur dioxide.
In the latter case, use was made of 4-10-day etiolated sprouts of meadow
fescue and timothy grass, which differ markedly in gas resistance in the
green state. The first species is resistant to SO,, and the second is non-
resistant. We studied the vulnerability of green fs02 concentration 1/400
by volume) and etiolated (concentration 1/500) plants aged 8 days, as well
as the amount of substances oxidizable by 0.1 N KMnO^ by Krasinskiy's method
(1950), the intensity of ultrafaint luminescence based on green and dry
weight and on a single plant, and the water content of the plants. The ultra-
faint luminescence was determined in the course of 10 sec in four replica-
tions. In the study of the influence of various S02 concentrations (0.001%,
0.01%, 0.1%, 1%, 10%, 100%) on the luminescence of plants, we recorded first
the luminescence before the action S02 (control), and then from the start of
the action of the gas for 10 minutes at 30 sec intervals. The plants were
grown in Petri dishes on filter paper.
The studies showed that the plants in the etiolated and green states
have different gas resistances. It was found that in order to observe clear-
cut differences in vulnerability to the gas, the etiolated sprouts require
an S0~ concentration 8-10 times greater than the green sprouts. On the other
hand the high resistance to SOy in plants in the green state also is not
always retained in etiolated plants (Table 1).
According to Krasinskiy (1950), there is no distinct relationship between
the gas resistance of green and etiolated sprouts and the content of oxidiz-
able substances.
An inverse relationship was observed between the gas resistance of
etiolated sprouts of the cereals and their ultrafaint luminescence, based
on 1 g of green and dry substance, and a direct relationship to the lumines-
cence was found based on one plant (Table 1). Hence, the stronger the free
oxidation in tissues, the lower the resistance of the plants to sulfur dioxide.
A direct dependence between the luminescence and gas resistance in the calcula-
tion based on one plant can apparently be explained by differences in the size
and mass of a single plant in different species (the gas-resistant plants had
larger sprouts). Under the influence of S02, the ultrafaint luminescence of
plants changes as a function of the gas concentration, and hence, as a function
of the amount of sulfur dioxide absorbed by the sprouts. Even a minimum S02
concentration (0.001%) activates the luminescence. As the S02 concentration
rises (to 10%), the activation of ultrafaint luminescence in the plants in-
creases. Concentrated sulfur dioxide (100%) causes a depression of the lumin-
escence of sprouts. A depression of the luminescence of sprouts with 100% S02,
as well as its activation at lower gas concentrations (10%, 1%, 0.1%), show a
- 51 -
-------
certain relationship to the vulnerability of the plants. In the resistant
species, meadow fescue, there is a smoother depression of luminescence
with 100% S02, and a greater activation by the gas in concentrations of 10%,
1%, and 0.1%. In the less resistant species, timothy, grass, a sharper de-
pression of luminescence with 100% SO, and a relatively weaker activation of
luminescence with 10%, 1%, and 0.1% S02 were observed. '
Table 1
Some Characteristics of 8-day Sprouts of Forage Grasses and Their Gas Resistance.
Species
Vulnerabil-
ity of Green
Sprouts in %
of Leaf
Length
_,
Yulnerabil-
lty of Eti-
nlni-oH
wJLdl/CU
Sprouts in
% of Leaf
Length
Amount of Oxidizable
Substances in 1 g of
Green Weight of
Etiolated Sprouts
Water-
Soluble
Substances.
Meadow fescue 2 8,8 . 0,65 ' '
Pasture ryegrass 15,5 6,5 0,59
English ryegrass 6 6,3 0,55
Common fescue 5- 14,7 —
Awnless brome grass 23 - -
Red-top fiorin ~ i00 O*6*
Orchard grass -9,8 14,4 0,7
Keadow foxtail 26 9*3 0,3
timothy grass 38 15,1 .1,1
Creeping quackgrass 20 7,9 0,3
Water-
Insoluble
Intensity of Ultrafaint
Luminescence, Pulses
per 10 sec, Based on:
1 g of
Green
iBJEht.
.0,15 296
.0,24 325
0,20 12$
-Li6
274
1 g of
Dry
3020
2742
29aO
1630
3023
0,18 625 ol245
0,18 516
0,12 3x8
0,15 306
0,18 304
4618
4756
2839
3666-
1
Plant
-
1,79
1,72
1,67
0,82
2,5
2,14
2,22
0,9
0,62
3,11
The above studies showed that the biochemoluminescence method can be
used to record the effect of toxic gases (SO^) in concentrations below
1 ppm (0.0001%) on etiolated plants. Hence, this method of diagnosing the
toxic effect of gases on plants is more than 10 times as sensitive as the
method of influence on photosynthesis, proposed by Thomas and Hill (1937).
Conclusions
t
1. It was found that etiolated plants of forage grasses are more
resistant to 802 than green plants. Damage to etiolated plants by sulfur
dioxide requires concentrations 10 times as high as those required for green
plants.
2. In etiolated plants there is no relationship between the amount of
substances oxidized by 0.1 N KMnO^ in 1 gram after Krasinskiy (1950) and the
vulnerability of plants to sulfur dioxide.
- 52 -
-------
3. Under the influence of S02 (cone, from 0.001% to 10%), the lum-
inescence in etiolated plants increases. The activity of the luminescence
is more pronounced in the resistant species - fescue, and less pronounced
in timothy grass. Concentrated sulfur dioxide depresses the luminescence
of both species.
LITERATURE CITED
I. ArasepAKCB A.CI., Tapycos B.K., 1965. Cjjopxcsatfafi xe-
'..!«[iecuie;inKH creS^e;*, naifcintu B saBHCimocTH OT TeMncparypii.
H3i.;^, ?.IO, B.2.
2< KpacHHcxnR H.n. 1965. TeopeiHHoaor«
-------
GAS RESISTANCE AND CERTAIN BIOCHEMICAL CHARACTERISTICS
OF ETIOLATED AND GREEN PLANTS OF FORAGE GRASSES
V. S. Nikolayevskiy and A. T. Miroshnikova
From Ministerstvo Vysstego i Srednego Spetsial'nogo Obrazovaniya RSFSR.
Permskiy ordena trudovogo krasnogo znameni gosudarstvennyy universitet im.
A. M. Gor'kogo. Uchenye Zapiski No. 222. "Gazoustoychivost1 Rasteniy".
Vypusk 1. Perm', p. 115-131, (1969).
The mechanism of oxidation processes in plants under the influence of
S02 has not been studied thoroughly thus far. Despite the considerable time
that has elapsed since the publication of the works of K. Noack (1920), who
assumed that sulfur dioxide causes the development of photodynamic oxidation
processes in light, the character of these processes is unclear nor are we
too clear as to the biochemical composition of the "oxidizable" substances
determining the gas resistance of plants after N. P. Krasinskiy (1950).
For this reason, neither Krasinskiyfs term "oxidizability" nor his method
of determination have attained a serious consideration among physiologists
and biochemists.
At the present time it has been established that S0£ depresses the
photosynthesis of plants (Ivanov, 1936; Thomas and Hill, 1937), increases
the respiration and the respiratory quotient (Zheleznova-Kaminskaya, 1953;
Ryabinin, 1962), inhibits the action of the enzymes enolase, phosphatase,
amjrlase, and catalase (Fortunatov, 1968; Nikolayevskiy, 1968; Nikolayevskiy
and Suslova, 1968), and enhances the activity of terminal oxidases, peroxi-
dase and polyphenol oxidase; the oxidation of sulfide is accomplished at
the expense of carbohydrates, according to Hazelhorf and Lindau (cited by
"Krasinskiy, 1950). Under the influence of S0£, an enhancement of the
reducing properties and of the pentose phosphate oxidation route in respir-
ation has been observed in resistant plants. In nonresistant plants, an
enhancement of the oxidizing properties and of the Krebs cycle was observed
(Nikolayevskiy and Suslova, 1968; Nikolayevskiy, 1968). The advantage of
the apotomical method of oxidation in the resistance.of plants to extreme
factors is explained by the important role of pentoses and hexoses in the
utilization of toxic products. Hence, resistant plants are characterized by
a greater buffering capacity of the protoplasm and a faster oxidation of
S02 to the sulfate, so that the toxicity of SOj is reduced almost 30-fold
(Tomas, 1962). In nonresistant plants, the oxidation of the anion of S02
appears to be retarded.
Many indicators of plant gas resistance such as the content of oxidiz-
able substances, ascorbic acid, oxidation-reduction potential, water-holding
- 54 -
-------
capacity and others proposed by various authors (Krasinskiy. 1950;
Nikolayevskiy, 1964) are only indirect and correlational, and thus do
not always reliably characterize the resistance of plants to S02.
Great possibilities for the study of the mechanism and chemistry of
photodynamic oxidations in plants were opened up by the development of a
new biophysical method of recording ultrafaint luminescence. The study of
the mechanism of oxidation processes under the influence of S02 will make
it possible to find direct indicators for an exact evaluation of the resis-
tance of plants. It has been found that the activation of luminescence is
the result of the oxidation of a substrate, and a decrease of luminescence
signifies a decrease of the oxidative processes or the closeness of the
object to death. It is postulated that the source of ultrafaint radiation
is the spontaneous oxidation of cell lipids (Zhuravlev, Polivoda, and
Tarusov, 1961).
Dark chemiluminescence makes it possible to estimate the general level
of physiological activity of cells and tissues (Vladimirov and L'vova, 1965)
the change in the state of certain functional systems, the viability of
certain tissues (B. N. Tarusov, I. I. Ivanov, and Yu. M. Petrusevich, 1967),
the mitotic activity of cells, the frost resistance, heat resistance, and
gas resistance of many plants (Gasanov et al., 1963; Agaverdiyev et al.,
1965; Agaverdiyev and Tarusov, 1955; Nikolayevskiy and Miroshnikova, 1968),
and the antitumor and radiation-protection properties of certain substances
(Tarusov et al., 1968). The biochemiluminescence method may be used to
ascertain (a) the temperature range of the life of various plant species
(Veselovskiy, 1963; Agaverdiyev, Doskach, and Tarusov, 1965), (b) the sub-
lethal and lethal concentrations of toxic gases for plants (Nikolayevskiy
and Miroshnikova, 1968), and (c) the various links in the biochemical chain
of oxidation processes under the influence of acid gases.
Making use of the new method we attempted to study the mechanism of
the action of sulfur dioxide on etiolated and green plants and to determine
the possibility of diagnosing the gas resistance of various plants.
Method of Investigation
The studies were conducted on etiolated and green plants of the meadow
fescue (Festuca pratensis Huds.) (a resistant species) and timothy (Phleum
pratense L.) (a nonresistant species). The plants were grown in Petri dishes
on filter paper at t = +20°C. Plants aged 4, 6, 8, 10, and 12 days were
used for the experiments. Alongside with the ultrafaint luminescence, inves-
tigations were made of the amount of oxidizable substances after N. P. Kras-
inskiy (1950), the content of ascorbic acid after Sapozhkova (1966), and the
vulnerability of the plants to sulfur dioxide (Nikolayevskiy and Suslova,
in press). The isotopic method was used for an exact determination of the
- 55 -
-------
amount of gas absorbed. The plants were gassed with 0.35% sulfur dioxide
having a specific activity of 2 microcuries per liter in the course of
1 hour. Twenty-four hours later, the vulnerability and activity of
were determined in 10 plants on a B-2 instrument using an SBT-7 end-window
counter at a voltage of 380 volts. The determination of the ultrafaint
luminescence was carried out on a quantum-measuring device consisting of a
highly sensitive photomultiplier FEU-42, USh-2 amplifier, PP-15 radiometer,
and VS-22 high-voltage rectifier. Plants weighing 3-5 g were placed in a
beaker which was connected via micro-vacuum pumps to a closed system con-
taining S02 in concentrations of 100%, 10%, 1%, 0.1%, and 0.001% by volume.
The luminescence without the gas was recorded first (control), then the
gas was introduced, and the change of luminescence was recorded for 10 minutes
(experiment). In the work with green plants, the spontaneous luminescence
was measured first (it was usually at the level of the background), then the
plants were illuminated with an OR-19 illuminator for 5 sec, and the lumin-
escence was recorded again. After the luminescence fell to the minimum
(4 minutes), sulfur dioxide was introduced (100%, 10%, 1%, 0.1%, and 0.01%
by volume), the luminescence being measured at the same time. After the
luminescence caused by the action of SOn decayed, the plants were again
illuminated with light from OR-19 (5 seconds), and the new luminescence was
recorded. • *
Results of Investigation
Two species of forage grasses differing sharply in resistance in both
the green and the etiolated state were selected by preliminary experiments.
Earlier, we established (Nikolayevskiy and Miroshnikova, 1968) a certain
inverse relationship between the gas resistance of etiolated eight-day
sprouts of cereals and their ultrafaint luminescence per one gram of green
and dry matter, and a direct relationship with the luminescence per plant.
It is known from literature data (Krasinskiy, 1950; Nikolayevskiy,
1963, 1965) that the amount of substances oxidizable with a 0.1 N solution
of KMnO^ and the ascorbic acid content of plants are indicators of their
gas resistance. In resistant plants there are less oxidizable substances
and ascorbic acid per gram of green or dry substance. We attempted to study
the reliability of these indicators for green and etiolated sprouts of
forage grasses in relation to their age and gas resistance (Table 1).
It is evident from Table 1 that the vulnerability of plants of both
species in the etiolated and green states increases regularly from the
four-day to the eight-day age, then decreases. Both etiolated and green
sprouts of the meadow fescue were found to be 2-3 times more resistant 'than
those of the timothy. Etiolated plants of both species were found- to be
more resistant than green ones. The fact that the retention of species dif-
ferences in the vulnerability of the plants is pronounced even in the etio-
lated state indicates that the gas resistance is a constant species indicator.
- 56. -
-------
Table 1
Some Biochemical Characteristics of Sprouts of Forage Grasses and Their
Gas Resistance in Relation to Age.
>>
<0
Q
0)
So
«*
Etiolated Plants
Amount. of Su
ces~0j
Eiaizao
ostan-
le
with 6.1 n boiu- 1
tion of KMnOi per 1
graaiof green
Substances -a
rdi
o a
£
li
r+
fi S
^
o»
«-g
EH O
o
o
• rl
•£,
i/i u>
£
'rH
•rl
1
(U
5
^
Green Plants
Amount c
Oxidizat
if Sutu
le wii
rtances
,h 0.1
N Solution of KMnOi
per Gram of Green
Substance
r-IO
Oq)
C/}H
I&4
(,
a
.pr-l O
1
Jr4H
C.X1-P
li
S
•H <1J
d 43
^s
E-tO
^
•a!
O
• rl
(4
M M)
•a B
£
3
t:
2
i
Meadow Fescue
4 0,95
6 0.85
8 1,1
10 0,85
12 0,35
0,15
0.4
0.45
0,65
0,2
1.1
1,25
1.55
1.5
0,55
0,23
0,17
0,18
0.19
0,16
2.8 1.25
3.4 0,9
7,7 0,9
5,1 1,45
4,2 —
0,25
0,65
0.4
0,35
—
1.5
1.55
1.3
1.8
—
0.64
0.37
0,23
0,27
—
7.1
21,7
18,5
10
—
Timothy
4 0.85
6 0.7
S 0,85
10 0.9
12 0,22
0.1
0,8
1,3
1.65
0.3
1.05
1.5
2,15
2,55
0,52
0,15
0,18
0,19
0.20
0,10
6,8 2,05
10.6 1.05
14.6 0.95
12,1 1.25
17.0 —
0.2
0,95
0.55
0,45
—
2.25
2,00
1.50
1.7
—
0,29
0,50
0.20
0.39
—
19.0
26,5
24,4
14,1
—
On the other hand, this makes it possible to study the resistance of plants
by means of the biochemiluminescence method. The retention of species dif-
ferences in the gas resistance of plants in the etiolated state makes it
necessary to assume the presence of a relationship not only to photosynthe-
sis, but also to some other species characteristics, in particular, to the
stability and buffering capacity of the cell content, i.e., proteins and
biocolloids of the protoplasm and organoids.
The amount of ascorbic acid in etiolated plants increases with age up
to 10 days, and decreases in the 12-day sprouts. In green plants, there is
no distinct pattern of change in the content of ascorbic acid with age. In
both cases, the meadow fescue contained less ascorbic acid (with the excep-
tion of the four-day age) than the timothy, this being in accord with our
conclusion (Nikolayevskiy, 1964) that the gas resistance of plants is depend-
ent on the ascorbic acid content. The latter is higher in green plants,
which is consistent with their greater vulnerability as compared to etiolated
plants.
- 57 -
-------
In the water-insoluble fraction, the amount of substances oxidizable
by a 0.1 K KJ&iO^ also increases with age. In' the meadow fescue, their
content is less than in the timothy in the etiolated and green states, this
being in accord with Krasinskiy's conclusions (1950). However, in the
etiolated timothy, the content of oxidizable substances was found to be
higher than in the green timothy.
Thus, the indicators of gas resistance of plants proposed by Krasinskiy
(1950) and us (Nikolayevskiy , 1964), i.e., the amount of substances oxidiz-
able by a 0.1 N solution of KMnO^ and ascorbic acid can indeed characterize
the gas resistance in individual species of plants.
In another experiment (Table 2) we attempted to study the vulnerability
and accumulation of sulfur in plants by means of the isotopic method. The
plants were gassed with labeled sulfur dioxide (S3502) of low specific
activity.
It is evident from Table 2 that, as in the preceding case, the etiolated
plants are more resistant than the green ones, and the meadow fescue is more
resistant than the timothy in both cases. The vulnerability of the plants
changes with age: the four and ten day old sprouts were injured less, and
the six and eight day ones more. It was found that the activity of live and
dry plant leaves per gram of dry substance was directly proportional to the
vulnerability of the plants. Hence, the nonresistant species accumulate
S3502 up to the lethal level more rapidly, and are therefore more extensively
injured by gases.
Table 2
Absorption of
by Plants of Forage Grasses.
VI
.3
o
a
01
£ .
3
Du-
HI
CO
Q
10
•g
3
a.
o
o>
St
•a:
Etiolated Plants
•jft
£
X
rH
• H
%
tt
1
>
4 6.8
6 10.6
| 8 14,6
4 10 12.1
tH
12 17,0
4 2.8
*« 6 34
0 §
'II 8 7.7
S£ 10 5.1
12 4.2
01
c
o
?*§
• rl CD
s.s
«<«-5
9.0
8.6
21
12
13.6
24
18
27
17
22
c
CM
#1
!&
•
^•S
~£
•sS
<< o
Green Plants
•;*•
£
.-a
rH
-H
g
ft
1
>"
20600 19
35185 26,5
331818 24,4
148333 14.1
145831 14.1
57561 7,1
139047 21,7
215652 18,5
87037 10
17G428 4,2
CD
s
1$
•HO.
>
4* S
-------
A new, interesting phenomenon was observed (Table 2). It was found
that in the green plants of the timothy, whose vulnerability is on the
average greater than that of the meadow fescue, the activity of S35 in the
leaves was 30% lower per plant, and 19% lower per gram of dry weight. This
was also observed in a comparison of the etiolated plants. Even in the
case of the same activity, the fescue is injured by sulfur dioxide less than
the timothy. Hence, the lethal limit of accumulation of sulfur dioxide in
the meadow fescue is higher than in the timothy. This may be attributed to
a higher stability of the proteins and biochemical structures in the cell
and a higher metabolism in the meadow fescue than in the timothy.
We verified the activity of sulfur in plant roots and ungerminated
seeds. It was noted that the amount of labeled sulfur was 2-3 times greater
in the roots of green plants than in those of etiolated ones. It may be
assumed that green plants can transport the absorbed sulfur in the form of
the SO^ anion into the roots and thence into the soil, thus reducing its
content in the leaves. A similar observation was made by Thomas and Hill
(1937). In the meadow fescue, the ability to redistribute and eliminate
sulfur from the leaves is greater than in the timothy.
Statistical treatment of the data showed that the coefficient of vari-
ation v for the activity of $35 in the plants was almost always higher than
the same indicator for the vulnerability. On the other hand, the coefficient
of variation and the accuracy of the experiment (P) for both indicators (vul-
nerability and activity) vary almost synchronously, i.e., the changes of v
and P for the vulnerability indicator coincide with the change of their activ-
ity. The accuracy of the experiment: fluctuates within the range from 3 to- 18%.
The differences in vulnerability with age are significant in both species or
border on significance, whereas differences in activity are only slightly
significant in most cases. The statistical differences in vulnerability be-
tween the etiolated and green plants are significant in the fescue and only
slightly significant in the timothy. The differences in activity are only
slightly significant. Thus, even in the presence of slight differences in
the absorption of sulfur dioxide by the plants and in relation to age, signifi-
cant differences in vulnerability are observed in the majority of cases.
The observation of differences in vulnerability in etiolated plants makes
it possible to assume that a change in the ultrafaint luminescence of plants
under the influence of SC>2 will characterize their gas resistance. Figures
1-3 illustrate the effect of sulfur dioxide and C02 of different concentra-
tions on the ultrafaint luminescence of etiolated plants in relation to their
age. From the literature (Agaverdiyev and Bynov, 1968) it is known that 100%
GOo intensifies the ultrafaint luminescence of cultivated cereals. We were
the first to show (Nikolayevskiy and Miroshnikova, 1968) that S02 causes an
intensification of the luminescence of etiolated plants even at negligible gas
concentrations (up to 0.001%).
- 59 -
-------
It is evident from Fig. la that in the 8-day etiolated sprouts of
forage grasses, a sharp depression of luminescence is observed under the
influence of 100% SO^, and a less marked depression under the influence of
100% C&2' !n the meadow fescue, we even observe at first a slight activa-
tion of luminescence under the influence of 100% C02. In wheat of the
Moskovka variety, as should be expected (Agaverdiyev and Bynov, 1968), a
strong activation of luminescence by 100% carbon dioxide and a slight acti-
vation by pure sulfur dioxide are observed. Differences in the nature of
the action of carbon dioxide and sulfur dioxide may be regarded as the result
of the greater chemical activity of S02 as compared with that of (X^ and the
greater strength of the acids I^SO^ and 112803 as compared with l^COq, which
are formed in the plants upon absorption. This is clearly evident ^Figs. 2-3)
from the effect of low 862 concentrations on the plants. Sulfur dioxide in
concentrations of 10% or less caused an activation of luminescence. The de-
tection of a similarity in the effect of the two gases on the ultrafaint
luminescence of sprouts provides one more proof of the universal and nonspecific
character of the action of gases on plants, manifested in a change of the
oxidation-reduction processes under the influence of acidification of the cell
protoplasm, and shows that the use of the term "acid gases" is justified
(Nikolayevskiy and Kazantseva, 1966; Nikolayevskiy and Yatsenko, 1968).
24001
.1600-
B
io \ Z 1 6 6 (O
Tine of Experiment, Minutes
Fig. 1 (a, b). Effect of 10C# S02 and 10C# CO on the ultrafaiat luminescence of etiolated
sprouts of forage grasses eight days old.
Notations 1 - meadow fescue - S02; 2 - meadow fescue - CO^; 3 -timothy -
SO,; 4 - timothy - CO-; 5 - red clover - SO,; 6 - Red clover - COa; 7 - Moskovka
whiat - S025 8 - Hoskfcvka wheat - C<>2.
- 60 -
-------
tloo
too
too
3 0
10% SO.
Meadow Fescue
Timothy
\
I i i » 4 it
H.
| -
a wo
rvx.
"^Ov
Y ^
'• -« N
i '-••<*'
{"*•*• + '"~'
? . \"
f j * « • w f « « « • w
5^ Time of experiment i~ miriu€es"
Fig. 2 (a, b, c, d). Effect of different concentrations of SO, on
the ultrafaint luminescence of etiolated sprouts of forage grasses
of different ages.
Notation: a - meadow fescue - 1C# SC^; b - timothy - 1$ SO,;
c - meadow fescue - $ S02; d - timothy - 1% S02j 1 - age four
days; 2-6 days; 3-8 days; 4-10 days; 5-12 days; 6-8 days
(control).
Sulfur dioxide (10%) causes a strong activation of ultrafaint lumin-
escence in etiolated plants of both species, which rapidly changes into
depression (Fig. 2 a, b). With age, the activity of luminescence under the
influence of 10% S02 increases in the plants. The character of the change
in the activity of the luminescence of sprouts under the influence of 10%
S0? with age (Fig. 2 a, b) coincides with, the change of their vulnerability
to the gas (Table 2). At the same time, the activation of luminescence in
the fescue iss always greater than in the timothy. This may undoubtedly be
attributed to differences in their gas resistance. Their regularity is also
observed in the action of other S02 concentrations (Fig. 2 c, d, 3 a, b, c,
d, e, f).
The experiments have shown that lower sulfur dioxide concentrations also
cause an increased luminescence of etiolated sprouts of forage grasses. In
these cases, the peak of the luminescence occurs somewhat later after the
- 61 -
-------
start of the action of S02. This is also noticeable during exposure to
0.001% sulfur dioxide (Fig. 3 e, f). i
too
*00
30.
Time of experiment, minutes
Fig. 3. (a, b, c, d, e, f). Effect of different SCfe concentrations
on the ultrafaint luminescence of sprouts of forage grasses.
Notations a - meadow fescue - O.J# SOgj b - timothy - O.J# S02;
c - meadow fescue - 0.0# S02, d - timothy - 0.0]* S02j 1 - age 4
days; 2 - fr daysj 3-8 daysf 4-10 daysj 5-12 daysj 6-8 days
(control)*
- 62 -
-------
A check determination of the luminescence of sprouts of forage grasses
in the course of 10 minutes showed that transplanting the plants into
beakers and their slight desiccation causes a certain depression of spon-
taneous luminescence.
Thus, the forage plants, which differ in their resistance to sulfur
dioxide, differ also in the activation of luminescence by the gas in the
etiolated state. Since in the etiolated state a vigorous vital activity is
observed in these plants when they are from 4 to 10-12 days old, (at the ex-
pense of the reserved substances stored in the seeds) one observes that the
greatest vulnerability of the plants to the gas and the greatest activation
of luminescence takes place precisely during the middle of this period
(6-8 days).
With the aid of the above-described setup one can diagnose the re-
sistance of plants by using etiolated sprouts. In this case, the degree of
resistance can be determined from the relative magnitude of the flash of
luminescence under the influence of the gas. This setup can also be used to
determine the permissible norms of air pollution for individual species of
plants on the basis of the appearance of the luminescence flash.
In cities and industrial centers, sulfur dioxide acts on the green
plants. Therefore, from a scientific point of view, it is important to find
out whether SO2 causes an intensification of the oxidation processes and the
luminescence flash in green plants, and to determine how it affects the
accumulation of luminous energy by the plant pigments. To this end, we
studied the absorption of luminous energy in 5 seconds of illumination on
the basis of the emission of light in the dark. When the luminescence of
green plants fell to the minimum (4-5 minutes) , SO, was used to determine
the luminescence flash, as in the case of etiolated plants. After the flash
had decayed, a 5-second illumination was applied again, and the binding of
luminous energy was determined from the magnitude of the emission of light
relative to the first light flash. The results of these studies are illus-
trated in Fig. 4.
It is evident from Fig. 4 that a five-second illumination causes the
emission of light in the dark, strongest in the case of sprouts 6-8 days
old and considerably weaker in 10-day plants. The intensity of light emis-
sion in the meadow fescue is somewhat higher than in the timothy. In the
4-8-day meadow fescue and 4-6-day timothy, 100% SC^ causes a luminescence
flash in the dark. The luminescence intensity of green plants under the
influence of 100% S02 is much greater in the meadow fescue. Sulfur dioxide
apparently causes a dissociation of the light and dark reactions of photo-
synthesis, and the luminous energy accumulated by the pigments is liberated,
producing a luminescence flash.
- 63 -
-------
•>l
'. ,".
It
• •!':
i *
o j'i
1 * 5
o ;
•~ i
* i
•• .-. i
2- /ft
a ' "=V*
•-i 'i.j\.
§-.. ~-
— ..
g Llight j»
*»% SO,
Meadow Fescue
•
a
t
jsL •
I Timothy
•
I
i
I
t A
— 7 i I i » "i" *J » "S • 1 « *
1 tight 5" Eight y I Light 5"
Xk *.
Mm
o
in.
0).
?i
-P
—3 T
Light 9' t'
• I * ' *• Min
Light 5 "
^--^r£*"~g
Light 5"
1
• 1 i
Light 5"
'Min
Fig. 4. (a, b, c, d). Effect of light (5 seconds) and SOg on the
ultrafaint luminescence of green sprouts of forage grasses.
Notation: a - meadow fescue - 10$ S02; b - timothy - 100J& S02;
c - meadow fescue - lOJt SO.; d - timothy - ICJt S02| 1 - age 6 days;
2-8 days; 3-10 days. *•
Subsequent irradiation of the plants with light (5 seconds) causes
another flash of radiation in the dark, but it id substantially weaker
(by a factor of 2-4) than the first light flash.
Thus, when the green plants are placed in the dark, they emit light.
Under the influence of 100% SO^, the green plants produce a luminescence
flash in the dark, but after the action of the gas, the plants absorb less
luminous energy when irradiated again. This may be attributed to an S02-
induced inhibition of the photochemical processes by which luminous energy
is bound by the plants. At the same time, the phenomenon of incomplete
depression of the second luminescence flash by sulfur dioxide la only aeemingly
so. Luminous energy is absorbed and emitted in the dark not only by pigments,
but also even by the cellulose membranes of cells. Apparently, they are the
ones responsible for the second luminescence flash.
- 64 -
-------
Timothy
Light 5"
1
k
/\
I
/ v_ .
i • :
ight 5" 3fa
_J
b
\
\ «
\
* — ^.__
* 1 * *
Light 5"
Light 5" jb. Light 5»'
Fig. 5 (a, b, c, d). Effect of light and S02 on the ultrafaint
luminescence of green sprouts of forage grasses.
Notation: a - meadow fescue - 1# SO,; b - timothy - 1% 902?
c - meadow fescue - 0.0$ S02; d - timothy - 0.01& 1 - age 6 days;
2-8 days; 3-10 days.
Under the influence of 10% sulfur dioxide (Fig. 4 c, d), a change in
the luminescence of the plants similar to the one produced by using 100%
S02 was observed, but in the timothy, the gas causes a fainter luminescence
than in the meadow fescue. At the same time, the luminescence flash after
the second irradiation with light was stronger than in the meadow fescue.
This cannot be attributed to the inequality of the vegetative mass in the
experiment, since the mass of the timothy was always smaller. The effect of
1% S07 on green plants (Fig. 5 a, b) is similar to the action of the 10% gas.
Under the influence of 0.1% and 0.01% S02, no luminescence in the dark is
observed in green plants of the meadow fescue and of timothy, and a second
irradiation with light causes an even greater luminescence flash than the
first flash in the meadow fescue, and it lasts longer in the timothy.
Obviously, 0.1 and 0.01% S02 causes an increase in the binding of luminous
energy by the pigments and other microstructures of the cell in green plants.
Thus experiments with green plants showed that high sulfur dioxide
concentrations (100%, 10%, 1%) cause a flash of ultrafaint luminescence which
is strongest in the meadow fescue and weaker in the timothy. Lower S02 con
centrations (0.1 and 0.01%) do not cause a luminescence flash in green plants
- 65 -
-------
in the dark. Concentrated gas depresses the binding of luminous energy
by the plants, whereas low gas. concentrations increase it.
Conclusion
Etiolated and green plants of the meadow fescue and timothy show
statistically significant differences in their vulnerability to sulfur
dioxide at nearly all ages (4, 6, 8, and 10 days). This makes it possible
to use the biochemiluminescence method for studying the gas resistance of
plants.
In etiolated and green plants of both species, a direct relationship
was confirmed between the amount of oxidizable substances, ascorbic acid,
and vulnerability to sulfur dioxide.
By using the isotopic method it was found that the timothy (a nonre-
sistant species) accumulates SC^ up to lethal concentrations more rapidly,
this being responsible for the greater vulnerability of these plants. On
the other hand, the lethal limit of accumulation of S02 in the timothy is
lower than in the meadow fescue (a resistant species).
Under the influence of S02 (concentrations from 10% to 0.001% and
lower),: a luminescence flash (intensification of oxidation processes) was
observed in etiolated plants. The greatest activation of luminescence and
its shorter period (2-3 minutes), changing into depression, is observed at
an S02 concentration of 10%. A lowering of the SO* concentration causes a
decrease of the luminescence peak, but there is a simultaneous increase in
the duration of the flash to 5-7 minutes.
The degree of influence of 80^ on the ultrafaint luminescence of plants
depends on their age: the greatest effect is observed at the age of greatest
physiological activity (6-8), when the resistance of the plants to S0~ is
lower. One hundred percent sulfur dioxide causes a substantial depression of
the luminescence of etiolated plants.
It is possible that the nature of the ultrafaint luminescence of etio-
lated plants under the influence of SO. and CC^ is similar. Differences in
the action of 100% S02 and (X>2 on etiolated plants result from their differ-
ent chemical activities. This discovered phenomenon is proof of the non-
specific nature of the action of "acid" gases on plants.
In experiments with green plants it was found that SO, (concentrations
of 100%, 10%, and 1%) causes a luminescence flash (intensification of oxida-
tion processes) which is greater in the meadow fescue than in the timothy.
Sulfur dioxide gas in concentrations of 0.1 and 0.01% causes no luminescence.
Under the influence of S02, an inhibition of photochemical reactions of binding
-------
of luminous energy is observed in the former case, and in the latter case
the gas causes an activation of these reactions.
The biochemiluminescence method may be used to solve problems of
diagnostics of the gas resistance of plants and to determine the minimum
permissible norms of air pollution for individual species of plants.
LITERATURE CITED
AraacpAiieB A. 111. 11 Bun OB O. A. 1967. O B.iiuiiiiiii yr.iexiic.ioro
rasa na CBCpxc.iafioc na-iyieiiiie pacTeniiu. V-4. aan. FlepMCK. yn-ra. As 157.
A r a u e p ;t u c B A. 111., Tapycon B. II. I9G5. Cucpxc.iaOaa xeMii.noMH-
Kccuenuwi CTcG.ieii iniiciiimu B ManiiciiMocrii OT Tc.MnepaTypu. /Kypn. «Bno-
jj}U3iiKa», r. 1.0, ULIII. 2.
AraoepAHCB A. UI., J3. o c K o i fl. E.. T a p y c o B E. H. 1965. Cocpx-
c.ia6oc H3.iyiteime pacTeimii npn noiiii/Keiiuii reMnepaiypw. AOK.I. All CCCP,
As 4.
B CCC.IOB c KM fi B. A.. Ccxa.MODa K. H.. Tapycoa B. H. 1963.
K'Bonpocv o MexamnMc cncpxc.iaOou choiiTniiuofi .iiuMiiiieciimiiiun opramoMoo.
>Kypll. c K c -i e 3 n o D a - K a M e H c K a n M. A. 1953. Peay.ibraTbt ititTpojyKmin
XBOfuiux 3K3OTOB B Jleiiiuirpa.ic H cro oKpccriiocrsix. tpvju 601. nn-ia IIM.
B. J\. KoAtapOBa. AH CCCP. ubm. 3.
H Baiioa JI. M. 1936. Qiiaiio.ioriifl piicTciiiu'i. roc.iecCy.Mi!3jiaT.
Kasaimcoa E. H. 1965. PaaoycToiViiiBocrb aeKopatiiBiibix pacieiuift D
yc.ioBiiflx npo»3BO,icrBciiiib!x TCppiiTop'iifi a.no.MiuiiicBofi iipo.MUtij.ieiiiiocTii ypa-
Jia. ABTopccJi. KaiiA. Aiicccpr.. CncpxiOBCK.
K p a c n n c K u A FI. n. 1950. CO. «/lbiMoycToi)AH, nun. 31, CBCPJVIOBCK.
H UKo.i a esc XH ft B. C. 1964. HcKOiopwe aiiaTO.MO-4>ii3iiOviorimecKiie
ocoCciuiocTii Apeaeciiwx pacTeinii'i B ycaosunx Mcaenvianiubiiofl npoMbuii.ien-
HOCTII CpejHcro Vpa-ia. KanaiiAaT. Aiiccepr.
H HKO.T acBCKiifi B. C., Cyc.no B a B. B. 1965. B.iiifliuie cepiuicroro
rasa H cepiioii KIIC.IOTM na ^wi3Ho.ioro-6HoxnMiiMccKiie npoucccu B .nicrbflx pae-
VM. 3an. HepMCK. yn-ia, sbin. 175.
- 67 -
-------
H n K o -i a e a c K H H B. C. 1966. B.iii&mie ceptiircToro aiirn.apiiaa na | awn. 5, CBCPA^OBCK.
HiiKo.iaeBCKiiH B.' C., Cyc.ioaa B. B. (B nciaTH). FaaoycTofmH-
oocTb ii iiexoTopbic ocotieiuiocTH TiOMOineinifl S^Oj rasomiw.MH rpasaMH.
H H K o .1 a e B c x H ft B. C., C y c Ji o B a B. B. 19C8. raaoycToiimiBocTb AS-
KopaniBiiux H raaomibix pacTeimfi B yc.noBHnx KOMomiaTa UBeruoft Meia.TJiyp-
mii Tlpeaypa.ibfl. MaTepun.iM TlepBou yKpaimcKOii Koiicpepeiimm «PacTeiiHH H
itpoMbiui.ieiuiaH cpeAa», , KHCB.
Hll KO.ia CBCKHfl B. C. 1968. Po.lb IICKOTOpUX OKIIC.IllTevlbllU.X CHCTeM
B Awxainni u rasoycToii'iiiBOCTH paCTcinift. Oiuiio.ioniH pacTeimfi, T. 15, Bbin. 1.
H UK o.i aeocKii ft B. C. n flueiiKo B. M., 1968. Sarpnaiieuue aiMoc-
4>epHoro Boaayxa npeAnpnaTiiaMii u ropoaax Tlpcaypa.ibH 11 cocroaiuie MX ose-
eiuiji. y, «HayKOBa ,iyMKa», KHCB.
T a p y c o B B. H., H a a H o B H. H., n c T p y c c n n i KD. M., 1967. Cflcpx-
c.iaCoc cnc'icinic Ciio.ioni'iecKiix CIICTCM. Hxvno .\\OCKOB. yn-ta.
P n 6 it H H H. 19G2. B.IIIJIIIIIC iipoMi.iiu.iciiuwx raaoa ua pocr Aepcoben H
"KvcTapiiiiKou. Botaii. /Kypita.1.. ^. 47, awn. 3.
C a n o /K K o B a E. B. 19C6. OnpeAwiciuie acKopCiinoDoA KIIC.IOTW. A.
«Konccpuii. H oiioinccyiii. npoM-crb*, K* 5.
T a p y c o B B. H., n o .1 u B o ,1 a A. H., >K y p a B .1 e B A. H. 1961. «Bno-
i]m:. II.. /Ky pa ii.icn^A. H., lion on A. H. 1965. BIIO.IIOMII-
ii. Tp. cir.inoniiy.M;! MOIIII. HSA-HO «IIayKa».
Tapycon B. II.. HaauoD H. H., flcrpyccBim K>. M.. Tano-
HCIIKO B. H. I9CC. Mcc-TCflOBaintc aiiTiioKiic.niTcnbiioff aKTiinnocTii CIIO.DUIH-
AOB pii:i.iiiKciiCBa.
opry n aToa II. K. 1958. Kpimi'iccKiiii oOsop aMcpiiKaiicKiix paOoT no
B-iiis!i;iio iip'oMMiiMouuux .IWMOB H raaoB na jieca. Zlowiaaw TCXA, sun. 36.
Nonck K. 1920. Untcrsuchungen ubcr lichtkatalylischc Vorgangc vor
physiologischer Bcdciitung. Zcitschr. Bot. Bd. 12.
Thomas M. D. and Hill G. R. 1937. Relation of sulfur dioxide in the
atmosphere to photosyutcsis and respiration of alfalfa. Plant. Pliysiol,
t. 12. N- 2.
- 6*-
-------
EFFECT OF MINERAL NUTRITION ON THE METABOLISM OF CARBON-14 COMPOUNDS
AND ON GAS RESISTANCE OF FORAGE GRASSES
V. V. Firger and T. B. Karpova
From Ministerstvo Vysshego i Srednego Spetsial'nogo Obrazovaniya RSFSR>
Permskiy ordena trudovogo krasnogo znameni gosudarstvennyy universitet im.
A. M. Gor'kogo. Uchenye Zapiski No. 222. "Gazoustoychivost1 Rastenly".
Vypusk 1. Perm', p. 85-97, (1969).
Because of a considerable pollution of air and soil, woody plants can-
not grow at the nonferrous metallurgical plants of the Urals. Herbaceous
plants are also disappearing from vast territories both in and around the
plants (up to 1-2 km and more), baring the soil to erosion. Even the sow-
ing of the hardiest herbaceous plants frequently fails to provide, there,
a reliable and lasting ground cover. Increasing the gas resistance of
plants by various methods will make it possible to grow greenery on such
grounds and prevent their water and wind erosion.
The methods of increasing the gas resistance of plants can be arbi-
trarily divided into agrotechnical ones (agrotechnical cultivation and
maintenance practices), chemical (regulating mineral nutrition, effect of
various chemical compounds), and biological (creation of resistant culti-
vated phytocoenoses).
Already the first investigators (Wieler, cited by Krasinskiy, 1950;
Ryabinin, 1952) indicated the possibility of indirect damage to plants by
acid gases through poisoning and causing deterioration of the soil. That
is why it is undoubtedly possible to increase the gas resistance of plants
by regulating and improving their mineral nutrition and also through the
effect obtained by means of spraying with various physiologically active
and other compounds.
A number of authors (Kazantseva, 1965; Kisser, 1968; Nemec, 1958; Yat-
senko, Nikolayevskiy, et al., 1968) established the positive role of nitro-
gen and potassium, and also of liming the soil, in the gas resistance of
plants.
Attempts have been made (Bobko and Fortunatov, 1958; Guseva, 1950;
Freebairn and Taylor, 1960) to increase the gas resistance of plants by
means of spraying with various compounds and by means of washing off the
toxic compounds from the leaves with various solvents. The mechanism of
injury to plants by acid gases (Noack, 1920; Ivanov, 1936; Krasinskiy, 1950;
Kroker, 1950; Nikolayevskiy, 1964) is related to a disturbance of photosyn-
thesis in the plants.
- 69 -
-------
As of this time, neither in the USSR nor in other countries, have any
studies been made on the role of photosynthesis chemistry in the gas resis-
tance of plants, or on the effect of the elements of mineral nutrition and
of the physiologically active compounds on the metabolism of carbon com-
pounds and on plant resistance. Our investigations were aimed at studying
these problems in two plant species differing in their gas resistance:
the meadow fescue (resistant species) and timothy (nonresistant species).
Method of Investigation
In the spring of 1967, these species were seeded on poor sandy soil.
In August, after mowing, the plots (2 m x 2 m) were watered with solutions
of fertilizers according to an 8-treatment scheme: control, N, P, K, NP,
NK, PK, and NPK. The fertilizers were applied in the following proportions:
N 4 g, P 6 g, and K 9 g per 1 m^. On the second day (28 August) and 5th day
(24 August), after watering, the effect of the fertilizers on the vulner-
ability of the plants to sulfur dioxide and on the metabolism of carbon-14
compounds was studied in reference to the same treatments.
The vulnerability of the plants was determined by gassing them in poly-
ethylene chambers. Ten plants of each species in each treatment were placed
in test tubes into a chamber for 1 hour, at an initial SO^ concentration'of
1/3000 by volume. The vulnerability was determined after 24 hours. It was
determined as the ratio of the injured portion of the leaf to the total
length of the leaf in percent. The introduction of labeled carbon dioxide
(c!^02-l% with a specific activity of 2 ]ic per 1 ml) into the leaves and
radiochromatographic analysis were carried out by following the procedure
described by A. T. Mokronosov (1966)1 The activity of the samples was read
off on a B-2 instrument with an SBT-7 counter. All the studies involving
the use of labeled compounds Were carried out at the radiochemistry labor-
atory of the Natural Sciences Institute of Perm' State University.
Results of Investigations
The vulnerability of the plants was determined, at different times in
addition to the above-indicated two days. With the age of the aftermath,
the vulnerability of the meadow fescue decreased, and that of the timothy
increased. This is apparently due to age-connected changes in the physi-
ological-biochemical processes and in the gas resistance of plants during
their renewed growing after mowing. Whereas at the start of the experiments
(21-23 August) the fescue plants were even less resistant to S02 than the
timothy, one observed already on 24 August that their vulnerability began to
approach the indicators characteristic of adult plants. It is also possible
that both in the control and in the treatments with fertilizers, the gas
resistance of the forage grasses was affected by the weather conditions of *
- 70 -
-------
that period. While a relatively dry and warm weather prevailed after the
first watering with fertilizers, cloudy and cool weather prevailed after
the second watering.
Data on the vulnerability of plants to sulfur dioxide and on the inten-
sity of photosynthesis are shown in Table 1. It is apparent from the data
that the meadow fescue, with the exception of the control treatment (28 August),
is characterized by a lower intensity of photosynthesis than in the case of
timothy. On the second day after the application of fertilizers, an increase
in the resistance of the fescue was observed in treatments K, NP, and PK, and
in all the treatments for timothy. On the fifth day after watering with
fertilizers, a positive effect was observed in the fescue in all the treat-
ments, and in the timothy, in all the treatments except N. No definite
relationship was observed between the change in the intensity of photosyn-
thesis (Table 1) and the vulnerability of the plants to sulfur dioxide under
the influence of the fertilizers.
Statistical treatment of the data on the vulnerability of plants to
sulfur dioxide showed that significant or close-to-significant differences
appeared in the fescue under the influence of nitrogen and potassium, and in
the timothy under the influence of K, P, and PK (24 August).
Let us consider the nature of metabolism of carbon-14 compounds in both
species in the control and under the influence of fertilizers on the 2nd and
5th day after their application (Table 2).
No substantial differences were observed in the percentage distribution
of the activity of C^ in the alcohol fraction, starch, hemicellulose and
proteins in the fescue and timothy in exposures of 10" and 30" + 4.5". Some
differences in the 51 exposure were observed in the content of radiocarbon
in the alcohol-soluble fraction and starch: in the fescue, as compared with
the timothy, the activity of the alcohol fraction is lower and higher in
starch.
On the average, about 90% of carbon-14 is concentrated in the alcohol
fraction (sugars, amino acids and organic acids), 3-9% in the starch, 1-4%
in hemicellulose, and 0.2-2.9% in the proteins. This was also observed on
the 5th day after the application of fertilizers (Table 3).
Let us consider the percent distribution (Tables 4 and 5) of the
activity of carbon-14 in the forage grasses in their different groups of
organic compounds (sugars, amino acids, and organic acids) and in individual
products of photosynthesis under the influence of fertilizers and in relation
to the change in the gas resistance of the plants. In the control plants of
the fescue and timothy in cloudy weather after 10 sec of exposure, the compo-
sition of the photosynthesis products formed is nearly the same. The distri-
bution of labeled carbon over the compounds is also similar. In the meadow
- 71 -
-------
Species
Date
Intensity of Photosynthesis. 'Pulses per Minute
Control -
.a
in
S 0
& CD
& M
a o
1 "*
.a
01
IM
SJR
o
•In J5
HJi
N
.a
-------
Table 1
and the Vulnerability of Forage Grasses to Sulfur Dioxide.
K
in
• H
Photosynthes
in 10 sec
in
• H
Photosynthes
in 30 sec
>>
Vulnerabilit
% of Length
NP
in
•H
Photosynthes
in 10 sec
1
VI
0)
JS
•po
Is
I Vulnerabilit
' 46 6f Length
PK 1
w
Photosynthes
in 10. sec
m
Photosynthe:
in 30 sec
Vulnerabilil
% of Length
NPK
in
Photosynthe
in 10 sec
i
.3
«i
|o
I*
O* -H
^
J->
1 Vulnerabili
1 # of Length
15320 20480 4,6 15260 21150 4,8 13360 19810
75,0 105.0 26,0 61,5 109,0 26,6 65.3 101
21030 24360 23.0 22170 20900 22.3 17310 21440
115.0 83 113,0 127,0 81.0 90.0
107,0
15322 36529 15,0 20991 24843 18.0 24-145
102
15067
3.1 18480 2I1HO 22.0
17.2 90.0 109,0 123.0
9,7 25540 19770 23.6
35,0 130,0 93.5 85.5
20,0 21021 32310
68,0 81.5 57,5 93.0 56,0 69,0 108.0 34,0 77,0 92,0 72;3
52322 47657 18.0 27765 18818 20,0 22138 34309 26,0 30164 33030
21,0
81,0
6.0
148.0
85.0 47,4 78,0 33,6 53 63.0 61.0 68,0 86.0 50.0 15.8
dioxide, a substantial incorporation of C1* in the sugars (raffinose), and
a decrease in the labeled C14 in the amino acids (glycine, alanine). Almost
the same was observed under the influence of NPK. Here the labeled carbon
was more extensively included in sucrose (5*). An increase in the gas re-
sistance of the meadow fescue occurred on the second day after watering with
K, NP, and PK (Table 4). In these treatments, the incorporation of C14 in
the sugars of the meadow fescue increased (K - sucrose, NP - sucrose and
fructose-1, 6-diphosphate, PK - sucrose), the incorporation of the labeled
C14 in the amino acids decreased (K and PK - alanine) in the NP treatment,
and the labeled C14 increased in glycine. At the same time, the C14 content
of organic acid decreased.
In the timothy, the vulnerability of the plants to sulfur dioxide de-
creased in all the treatments under the influence of the fertilizer, particu-
larly that of PK. As compared to the control, this treatment is characterized
by an increase of the labeled C14 in the sugars (sucrose) and a decrease in
organic acids (PGA and malic acid).
On the whole, in the case of a positive effect of the fertilizers, the
meadow fescue is characterized by a more homogeneous distribution of C^
over the individual compounds than the timothy.
On the fifth day after the application of fertilizers (Table 5), all
the treatments showed a positive effect on the increase of the gas resis-
tance of the plants with the exception of nitrogen in timothy. In the meadow
- 73 -
-------
Table 2
Distribution of Carbon-l4 Activity in Forage Grasses According to Fractions
nf Poyf.il i^cT-g an 4-tia Sj»r»«nH Day. ?fi tllglist. 1067.
»i
.a
I
Treatment.
Acjji^qfta, of Hydrolyzates,. Percent of Initial Y^ue
PGA, Sugars,
Amino Acids.
and Orranic
Acids
10"
5.'
Starch
10"
5'
Hemi-
Gellulose
10"
5'
Proteins
10*
5'
Total
10» 1 5'
Control
*
3
§
•s
1
K
f
N
K
. NP
PK
NPK
Control
N
K
NP
1 PK
! NPK
87,2
90,1
91,0
90,8
89,0
93,5
92,7
92,5
87.0
93.2
91.3
90,2
86.5
87.0
95,0
85,0
'86,0
79,0
88.5
91.5
89,2
91.9
87,6
89.0
5,2
5,7
6,2
3.6
4.1
1'°
3.0
3,26
6,3
2,2
4,8
4,5
9,3
7,7
—
9,7
7.8
11,7
7.5
5.5
6.5
3,65
5,4
5,8
3.2
0^6
0.7
3,2
3,1
0.4
1,2
0,8
1,8
2.1
1.1
1.6
1,3
1.0
—
2,8
1.3
2,05
0,75
0.6
1.7
1,8
2.3'
1,2
2.8
2,5
2,2
1,4
3.5
1,5
2.75
2,5
4,05
1,7
2,3
3,1
2,5
3,7
4.75
2,0
4,2
5,7
2.2
2,3
2,3
2.3
4,1
3.4
99,0
99.26
100
99,0
99,7
99.4
99,65
99,16
99,15
99.2
99,5
99.4
99,6
99,4
99,75
99.5
99,3
98.45
98,95
99.9
100,4
99,65
99,4
99,4
fescue, the greatest positive effect on the gas resistance of the plants
was produced by N and K, and in the timothy, by K and NPK. In these cases,
in the fescue we observed (Table 5) an increase of the labeled C1* in the
sugars (raffinose), a decrease of C1* in the amino acids (a- and 3- alanine),
and the preservation of the relative activity of organic acids. In the
timothy, potassium and NPK caused a decrease of labeled carbon in the sugars
and an increase in the amino acids (51 exposure) and organic acids.
Thus, in the presence of increased solar radiation (24 August), on the
fifth day after the watering, the role of the fertilizers in the gas resis-
tance of the plants and their influence on the metabolism of carbon-14 were
different than in cloudy weather (28 August). The relationship between the
gas resistance of the plants and the chemistry of photosynthesis can be
understood if one considers the weather conditions on the days of the experi-
ment. The difference in the chemistry of photosynthesis in the plants
(control treatment) is manifested most clearly during hot, sunny weather.
- 74 -
-------
Table 5
Distribution of Carbon-lA Activity in Forage Grasses on the Fifth Day After
Application of Fertilizers.
in
•H
o
&
OT
Treatment
Activity ef Hydrolyzates. Percent of Initial Value
PGA, Sugars,
Amino Acids,
and Organic.
Acids
10"
5'
Starch
10"
•"
ftajni-
cellulose
10"
*
Proteins
10"
5'
Total
10" 5'
1
a
§
(0
a>
g
•o
CO
^
|
•H
6-1
Control
N
K
NP
PK
NPK
Control
N
K
NP
PK
NPK
—
92,5
93,0
91,7
91,3
87,0
- 88.8
93,0
92.2
91.8
92.1
91,8
90.2
90,0
90,9
89,0
89.5
89,2
87,0
93.6
92.1
91.7
92,7
91,2
—
3,3
1.6
1,7
2.9
5,25
.5.7
—
0.27
—
3.95
2,3
3.05
4.67
5.5
6,0
6,8
4,7
5.6
4,0
3.6
3,7
1,6
f.O
—
1.5
1,6,
2.0
3.05
4.7
2,0
—
3,3
—
1,0
1,7
3,16
1,6
0,9
2,55
0.53
3,2
3,9
2,1
2.2
0,9
2,05
2.3
—
2,0
3,0
1.9
2.15
2.06
0.2
—
3,9
—
2.15
3.6
3.65
2.8
2.1
1.7
2,9
2.5
2,7
2.6
1,5
2.7
2,6
2,95
—
99,3
99,2
97,3
99,4
99.0
96,7
_
99.67
—
99.2
99,4
98,86
99.07
99.4
99.25
99,73
99,3
99.2
102,3
99,3
99.0
98.85
100,45
Under these conditions (51 exposure), the fescue is characterized (Table 5)
by an increased formation of organic acids - 66% and a comparatively slight
formation of amino acids and sugars (13-14%). In timothy, on the other
hand, there is an increased synthesis of sugars (79.7%) and a very slight
formation of amino acids and organic acids (7-12%). At the same time, the
increase in the gas resistance of the meadow fescue is associated with an
enhanced synthesis of sugars and a decrease in the synthesis of amino acids
and organic acids, whereas in the timothy, it is due to an increase in the
synthesis of amino acids and acids of the Krebs cycle.
Earlier we observed (Nikolayevskiy, 1968) that in plants resistant to
S02 (box elder, summer-cypress), under the influence of the gas and light,
the role of the pentose phosphate by-pass in respiration increases, whereas
in nonresistant species (white birch, balsam poplar) the role of the Krebs
cycle increases in respiration. Thus, data on the role of oxidation systems
in the respiration of plants and on the chemistry of photosynthesis (Table 5)
- 75 -
-------
Metabolism of Carbon-l4 Compounds in Forage Grasses on the Second Day
Species and
Treatments.
Timothy, control
> » »
> > N
> » N
» » K
> > K
» » NP
» > NP
> . » PK
> > PK
> » NPK
» ' » NPK
Meadow fescue,
control
» > >
» > N
* » N
» K
» » K
> » NV?
» » NP
$•
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
> » PK 10"
> » PK 5'
» » NPK 10"
» » NPK 5'
p*02 rH
i-Hf* *
•H-p cd
3 s
C
-------
Table 4
After the Application of Fertilizers (28 August
Compounds and Classes^ Percent of Activity of Starting Spot -
Acids
C^-alanine
1 3 -alanine
a
•6
EH
Organic Acids
£
3
o
•3;
o
.3
rH
CD
S
Tartaric
Aeid
Pyruvate
EH
.3*
It
to to
•d
•8
EH
Vulnerability
4,7
:io,9
6,4
•6.2
4,9
4,8
5,8
•5.3
9.2
3,9
14,7
16
8.9
fifi
no
4,4
10,5
7,2
14,3
4,0
2,8
3.2
1,1
10.2
—
5,5
—
3.0
—
3.9
2,0
24,6
—
—
3,7
—
__ _
4.0
1.9
—
2.6
—
—
_
—
11.9
24.3
12,8
9.6
12,1
10,3
13,2
26,0
38.2
13,5
22,2
5,8
12.6
8,4
39.4
14,2
22,4
13.4
19,7
11.2
5.6
9.7
13,4
18.5
50,5
9,4
19,0
14.6
23.3
7.2
37.0
10,3
19,9
17,4
33,5
8.5
48.8
41.1
25.8
11.6
23,6
13,1
19,6
13.2
13,1
21.8
30,7
15.9
6,6
8.6
39,3
10,6
4.9
3.4
6,2
2,4
4,4
5,7
3,2
5,8
8,9
5,4
5,5
4,4
20,0
6.3
6,2
5,1
3.9
7,4
19,2
1.4
2,4 2,4 . —
9,2 - -
— 5,2 —
2.0 2.4 —
6,4 •- —
3,4 3,0 2,4
6,7 — —
— — —
4.0 — —
— — —
2.1 8.6 1,6
6.0 - —
6,7 - -
7.7 - -
2.1 — —
6.3 — 1.8
7.6 -
- - 7.3
9.8 5.4 —
6.0 - 2,6
10.3 — —
1.6 — 2,4
3,2 - -
— — —
61.9
29.8
63,5
29.6
34,6
20.1
49,0
13.2
28.2
23.1
49,0
20.3
64.4
54.2
33,4
24,1
51,2
26,7
41.0
26,9
27,3
33,2
53.1
.17.3
7.2
2.5
4.7
6.2
5.2
1.6
24.2
2.0
6.2
19,2
6.2
4.2
7.4
12.6
5.5
2.6
19.4
3.3
9.0
2.7
5.6
6.3
22.2
2.8
100
100
100
100.
100
100
100
10
100
100
100
100
100
100
100
100
100
100
100
100
100
100
too
100
18.0
25.4
4.6
4.8
,
3,1
22.0
27,7
23,0
23,0
22.3
9.7
23.6
- 77 -
-------
Metabolism of Carbon-Ik Compounds in Forage Grasses on the
Species and
Treatments
a
g.
£
4.JL
•£. S tn
3* *M
.«>.*
5.-Sw
Distribution of Cwbon-14 Over Indi-
Sug-
rT :
H.
If
$
o
•S
I
$
g
1
$
•g
1
Meadow Fescue
Control
>
>
>
.>
»
>
»
»
»
>
>
Timothy
»
>
>
>
>
>
»
>
»
>Ccntrol
> K
» K
» K
» K
» XP
» KP
» PK
» PK
» NPK
> NPK
Control
•Control
» N
> N
» K
> K
> PK
» PK
> NPK
> NPK
UT
5'
10"
5'
10"
5'
10*
5'
10"
5'
5'
10"
10"
5'
10"
5'
10"
5'
10"
5'
10"
5'
1831
1694
2822
1628
1590
3093
2242
1876
2562
1798
3504
5876
3446
1402
5333
1981
2121
4033
3152
2511
2.0
3,0
7.0
0,7
1,9
9.8
3,0
0,1
2.9
0,4
4,3
25.0
37,5
2.4
1.7
25.5
1.2
1.0
0.4
10,0
11.1
3,4
19.4
15.2
68.5
95
61.2
55,5
9.5
36.4
76.8
11.7
20.7
66.8
66.7
12.4
33.1
6.5
57,5
0,4
5,0
1.6
13
3.4
1.3
6.0
1.6
2.1
25
3.3
2.1
4.5
3.0
in relation to gas resistance yield a pattern similar to that of the depend-
ence of plant resistance to St^ on the chemistry of the oxidation-reduction
processes of photosynthesis and respiration. Our data attest to an inter-
linking and intimate coordination of metabolism which determine the resis-
tance of the plants.
The resistance of the aftermath of forage grasses to sulfur dioxide
changes with age. In the meadow fescue, this resistance is very low at the
start of the regrowth and high after the regrowth is completed. In timothy,
on the contrary, the resistance is high at the start and low at the end of
the regrowth period.
- 78 -
-------
Table 5
Fifth Day After Application of Fertilizers_(24_August 1967).
victual Compounds and Classes, % of Activity of Starting Spot
ars
r-i
a
-8
H
Amino Acids
.9
o
5
o
5
§
(-1
I
3
g
1
CQ
1
•8
el
Organic Acids
<
£
•a*
.•s-3
ex.
.8,1
H-r4
0) O
a«
r-t
a
•8
EH
•8
^#
sf
•11
.ss
1-1
ed
*
6-1
$
rH
.rt
1
§
rH
s
:»
14,3 3,3 9,8 — 13.1 66,0 — — 66,0 6.6 100 26.0
11,1 — 13,4 3,2 16,6 62,1 2,9 5.5 70.5 1.8 100 J2(>
30,7 0,15 0,13 0,1 0,55 68,0 — — 68.0 0.75 100
64.9 — 1,0 — 1,0 34.1 — - 34,1 — 100 ,,„
26.2
69.8
12,5
64,6
60,4
50,8
39.4
79,7
17,7
69,3
20.7
70.8
14,5
37,6
6,5
60,5
3,4
6,4
3,8
2.7
8.1
1.1
6,8
4,3
4.8
7.0
3.6
9.2
5.9
7.5
—
—
15.8
6.9
4,6
13,6
7.0
1,9
9,6
2.9
5.2
6.7
6.5
5.9
19.9
9.7
5.5
8.3
24.8
—
— '
—
5,4
1,8
—
—
—
—
2,6
3.3
1.7
30.3
—
7,6
44,0
13,3
8,4
16,3
20,5
4.8
16,4
7,2
10,8
13.7
12,7
18.4
27,5
47.5
5.5
15.9
26.7
11,6
66,3
8.4
14,4
42,4
27,2
8.2
60,7
9,4
54,9
8.8
49,6
H.l
77,8
15,2
I.I
2.0
—
0.7
1.8
—
10.9
1.2
0.2
3.1
2.3
— .
2.1
1.5
2.7
2.3
—
2,4
—
1.6
1.6
—
4.8
3.1
1,4
—
2.1
1.9
—
,2.0
1.6
5.6
27.8
16.0
66.3
10.7
17.8
42,4
42,9
12,5
62.3
12.5
59.7
10,7
51.7
14.6
82.1
23.1
2.0
0.9
12.8
8.4
1,3
2,0
1.4
0,6
10
4.5
7,3
0.1
6.3
0.3
5,9
0.5
100
100
100
100
100
100
too
100
100
100
100
100
100
100
100
100
Different fertilizer treatments had a positive influence on the gas
resistance of the forage grasses. At the same time, the positive effect
of the fertilizers depends on the weather conditions.
The nonresistant species, timothy, has a higher intensity of photosyn-
thesis than the meadow fescue (the resistant species).
A similar relationship between the gas resistance and a reduced intensity
of photosynthesis was first noted in woody plants by V. S. Nikolayevskiy
(1963).
- 79 -
-------
Conclusions
The control plants of meadow fescue and of timothy after a 10-second
and 5-minute exposure to ^-^2 show practically no differences in the
nature of metabolism of the main groups of organic compounds. Under the
influence of fertilizers, during a five-minute exposure with C^C^, the
polymerization rate of carbohydrates is somewhat higher in the fescue than
in the timothy: the percentage of the labeled carbon is higher'in the
starch and lower in sucrose.
Fertilizer treatments with a positive effect on the gas resistance of
plants cause different and serious changes in the synthesis of individual
products. The change in the chemistry of photosynthesis may be attributed
both to the indirect effect of fertilizers and periods following their
application, and to the species characteristics and the effect of weather
conditions. In the meadow fescue, in the presence of increased solar radi-
ation, the improvement in gas resistance under the influence of fertilizers
is associated with an increase in synthesis of sugars and a reduction in
the synthesis of amino acids and organic acids; on the contrary,' in the
timothy, an increase of the synthesis of the latter compounds is observed.
- 80 -
-------
LITERATURE CITED
Bo6KO E. B., tbopTynaroa H. K. 1958. Onwr xu.MmiccKoii 6opb6«
c ycbixaiincM .iccoiiaca/Kaennu B ropoACKofl oGcranoBKe. Ibnccnis TCX4,j\96,
P y c c B a B. A. 1950. B.IHJUIHC Mimepa.ibHoro niiraiina na OKiic.iHTe.iwio-
&occTanoBiiTe.ibiibifi pe/Kii.M H raaoycToihiiBocTb pacTemiii. C6. «^uMoycTofi-
iHBGCTb pacTcinifi 11 flbiMoycToii'iiiBbie accopTii.MciiTbi». AKX PC0CP u TopbK.
yn-T, MocKBa — FopbKiiH.
H B a H o B Jl. A. 1936. ^>K3Ho.iori!a pacTeHiifl. FocJiecoyMiisjiaT.
K a 3 a u u c D a E. H. FaaoycTofwHBocTb jeKopaniBiibix pacTcHiift B ycno-
nponsBoacTBCHUbix TCppiiTopufl a-noMUHiiCBoft npoMbiui.ieiiHOCTii Vpa.ia.
pccfepar xaHfl. AiicccpTamm. CBBPA-IOBCK.
K p a c ii u c K u A H. n. .1950. TeopeTimecKHe OCHOBBI nocrpoeniin accop-
THMCHTOB raSOVCTOl'mHDbJX paCTeHHH. C6. «ZlblMOyCTOH4)!BOCTb paGTe.lHfl H
JlblMOyCTOftMHBbie aCCOpTH»ICHTbI». AKX PC<1>CP H FopbK. yH-T, MoCKBa—
I'OpbKHH.
MoKpOHOcoB A. T. 1966. HeKoiopwe Bonpocu MetoAHKH npHMcHemia
Hsoiona yiviepo.a.a-14 A^H HayneHHa ii3iiaionm pacTciuifi>.
nun. 1, T. 15.
P n 6 it u H u B. A. 1962. BJIJIHHHG npoMUUiJieimux rason Ha pocr AepeBbca
u KycraoiiiiKOB. Botan. xcypn., T. 47, Nt 3.
flu'enKO B. M.. HHKO/taeBCKHfi B. C, nprep B. B., Cycjto-
B a B B. 1968. B.iiimnie Miinepa.ibiioro niiTaiinn na rasoycTofi'iiiBocrb raaon-
KWX rpan. Marcpiia-iH IlcpBofi VKpaiiHCKOfi KOH^cpciiuiiH cPacTcnun H npo-
Mbiui.iCHiiafl cpcAa». HSA-BO «HayKOBa AyMKa>, K»eB.
Kisser J. 1968. Physiologische Probleme der Emwjrkung von Luft-
verunrcinigungen auf die Vegetation. Materialy VI A\iediynarodo\vej konfc-
rCnCJNcmcc'A! 1958. Vliv kourc a popilku na intoxiknci smrkovich poroslu.
CSNoa'ck K. 1920. Untersuchungen fiber Uchtkatalytischc Vorgange vor
Physiologisclier Bedentung. Zeitsehr. Bot. Bd. 12.
- 81 -
-------
METABOLISM OF CARBON-14 COMPOUNDS IN FORAGE GRASSES
AND THE EFFECT OF SULFUR DIOXIDE ON IT
V. S. Nikolayevskiy, V. V. Flrger, and G. A. Vaseva
From Ministerstvo Vysshego i Srednego Spetsial'nogo Obrazovaniya RSFSR.
Permskiy ordena trudovogo krasnogo znametii gosudarstvennyy universitet im.
A. M. Gor'kogb. Uchenye Zapiski No. 222. "Gazoustoychivost1 Rasteniy".
Vypusk 1. Perm1, p. 57-67, (1969).
At the start of the 20th century, physiological-biochemical methods
became increasingly popular in studies of the gas resistance of plants.
They were used to reveal the characteristic physiological-biochemical
peculiarities of plants in connection with differences in gas resistance
(Krasinskiy, 1950; Kroker, 1950; Nikolayevskiy, 1964; Kazantseva, 1965;
Sitnikova, 1966; Kulagin, 1965, 1966; Il'kun and.Motruk, 1968; Kisser, 1968).
On the other hand, many of the established indicators of plant gas resistance
("oxidizability," water-holding capacity, ascorbic acid content, etc.)
apparently are not directly related to gas resistance. More often than not,
they are merely indirect reflections of differences in the vital activity
and gas resistance of plants. Considering the general biological importance
of the proteins and biocolloids of the protoplasm, and the importance of
the metabolism of organic compounds in the regulation of the life processes
of various biosysterns (cell, organism, population), one can assume their
important role in plant gas resistance as well.
The development of new isotopic methods of studying the metabolism of
organic compounds - radiochromatography (Arnon, 1959; Mokronosov, 1966) -
made it possible to initiate more refined biochemical studies of plant gas
resistance (M. D. Thomas et al., 1944 a, b; Harrison et al., 1944; Godzik,
1968). Still, differences in the nature of the metabolism of organic com-
pounds in plants differing in gas resistance, conversion and de to 3d. cat ion*
of acid gases by the plant, and change and disturbance of the processes of
photosynthesis and respiration under the influence of gases are still unclear.
We attempted to study the chemistry of photosynthesis in two species of for-
age grasses of different gas resistance, namely: that of the meadow fescue
(a resistant species) and of the timothy grass (a nonresistant species)'.
Procedure
Plants for the experiments were grown in pots on fertile soil. The
photosynthesis and the effect of sulfur dioxide on it were studied on cut
* [Translator's note: "intoxication" in Russian original.]
- 82 -
-------
plants 20 and 30 days old with the help of labeled carbon dioxide (1% C1402,
specific activity 2 yC per ml). The introduction of C1402 into the leaves
and the radiochromatography of organic compounds were performed according
to a method described by A. T. Mokronosov (1966). Sulfur dioxide (S3502)
with a specific activity of 10 uC in 1 %, was introduced by fumigating the
plants in polyethylene chambers. The vulnerability was determined as the
ratio of the damaged length of the leaf to the total length, expressed in
percent. The activity of the plant samples was read on a B-2 instrument
with an SBT-7 counter at a voltage of 380 volts.
The amount of substances oxidizable by a 0.1 N solution of KMnO^ was
determined by N. P. Krasinskiy's procedure (1950), the determination of
ascorbic acid content according to the method of Ye. V. Sapozhkova (1966).
and of the total acidity, by the methods of A. Ye. Yermakov, V. V. Arasi-
movich, and others (1952).
Experiments dealing with the study on the metabolism of carbon- 14 and
sulfur- 35 compounds were carried out in the following variants:
1. The plants were fumigated with sulfur dioxide (S35o2) for 10 minutes
and 1 hour. The fixation of the material was carried out with ethanol vapor
(for 3 minutes) after the fumigation and after 24 and 72 hours.
2. On the control and experimental plants (from the preceding variant),
photosynthesis was studied in a 100 ml chamber closed with mercury (Mokronosov,
1966). The exposures were 15sec C1402 , 1^ C^, and lm±n cl^Oj+Vn light.
3. Sulfur dioxide (S^5o2) was introduced into the cut plants in the
same photosynthesis chamber from a gas-washing bottle connected in parallel.
After S02, cl^02 was introduced into the chamber. Sulfur dioxide - 14% with
a specific activity of 8.3 yC per ml, C1402 - 1%, specific activity 2 yC per
ml.
4. In the last variant, the photosynthesis was studied in a mixture of
1% S350« and 1% C1402 for 5 and 15 seconds and 1, 5, and 10 minutes. The
specific activity of S3502 was 5.5 uC per ml, and that of C1402, 2 yC per ml.
Studies using isotopes of carbon and sulfur were carried out in the
radiochemistry laboratory of the Natural Sciences Institute of Perm1 State
University. The authors express their gratitude to V. Ye. Zhuravlev and the
laboratory staff for their cooperation in these studies.
Results of Investigation
In order to select the objects for the study of gas resistance, we first
tested 20-day old green sprouts of 9 species of plants (Table 1).
- 83 -
-------
It is evident from Table 1 that among the forage grasses, the lowest
vulnerability to gas injury is displayed by the meadow fescue, and the
highest by the timothy grass. Differences in the vulnerability of these
species were statistically significant in all five experiments.
lable 1
Vulnerability of Plants to Sulfur Dioxide
Species
1.
2.
5.
4.
5.
6.
7.
8.
9.
Meadow fescue
English ryegrass
Orchardgrass
Pasture ryegrass
Creeping wheatgrass
Aimless bromegrass
Meadow foxtail
Timothy grass
Red clover
Percent Damage to
Length of leaf
2.0
6.0
9.8
15.5
20.0
23.0
26.0
98.0
100.0
In the case of the fescue and timothy plants studies were made as to
the content of their oxidizable substances, ascorbic acid, and organic
acids, as well as on the effect of sulfur dioxide (Table 2).
The species differing in gas resistance - fescue and timothy - possess
certain biochemical characteristics (Table 2). The resistant species
(meadow fescue) is characterized by a reduced content of substances oxidiz-
able by potassium permanganate, a reduced ascorbic acid content, and a
lower total acidity. Under the influence of sulfur dioxide an increase in
the content of oxidlzable substances takes place, in contrast to Krasinskiy's
data (1950). It may be postulated that the change in the content of these
compounds also depends on the time elapsed after the beginning of the effect
of sulfur dioxide. During the first few hours, the latter apparently causes
depolymerization and an increase in the amount of oxidizable substances. As
time goes on, more extensive processes of oxidation and outflow of organic
compounds produce a decrease in the amount of oxidizable substances.
The ascorbic acid content decreases under the influence of SO- (Table 2).
This is particularly obvious in the calculation per gram of dry weight. The
decomposition of ascorbic acid under the influence of SO- is more marked in
the nonresistant species - timothy, and less marked in the fescue. As is
evident from Table 2, a more precise and reliable indicator of gas resistance
and of the effect of sulfur dioxide on plants is the ascorbic acid content.
-------
Table 2
I
8
Some Biochemical Characteristics of Forage Grasses and the Effect of Sulfur Dioxide on Grasses
Species
1
3
Ou
'S
faO
•s
.•y
'•§ ..
1
Or-t
>OL,
Control experiment
Amount. of Substan-
ces oxidizable by
0.1 N KMnOi. in
1 g of green
weight
01
rH
a
O.S
Total Con-
tent of
organic acids
in percent of
malic acid
c
3
S.
CJ
"o
-p
4)01
O.SC
jj
a
%
jf.f
01
c
01
2
o
0101
jj
Q
a
"o
4
01 01
a.*
Meadow fescue
Timothy grass
Meadow fescue
Timothy grass
20 days
20 days
30 days
30 days
9.7 0.95
48.4 0.85
17.5 0.50
50.8 0.90
1.75 2.65
1.90 2.75
0.90 1.40
1.55 2.45
4.6
5.3
2.7
3.4
7.0
6.2
3.4
4.2
0.022
0.029
0.020
0.026
0.33
0.33
0.26
0.22
0.55
0.70
0.55
1.20
1.30 2.15
2.40 3.10
1.00 1.55
2.30 3.50
4.6 4.0
3.7 2.5
2.5 2.1
3.0 1.7
0.069 0.53
0.028 0.25
0.024 0.21
0.039 0.22
-------
Comparison of the intensity of photosynthesis in the plants (Table 3)
shows that for all exposures, the rate of carbon dioxide absorption in the
meadow fescue, in comparison with timothy grass, is lower by an average
factor of 1,5. This confirms our conclusion (Nikolayevskiy, 1963) that the
intensity of photosynthesis is lower in the resistant species.
fable 5
Intensity of Photosynthesis in Forage Grasses, Pulses per min (200 ng
of Dry Weight)
Species'
Exposure
15 sec
C"02
Meadow fescue W25
Timothy grass 5630
1 min
C»0S
6052
9015
1 nin. -j- 4 min
C««O2 light
7011
13353
The absorption and redistribution of S3502 by the plants (Table 4)
showed that the build-up of 5^02 ^ *&& fescue continues for the duration
of 30-50 minutes, whereas in the timothy, the build-up takes place only dur-
ing the first 10 minutes. During the first 10 minutes of the experiment, the
timothy accumulates per gram of dry weight three times as much toxic gas as
the fescue. Even in long exposures, the amount of absorbed sulfur dioxide
in the timothy is greater than in the fescue. . The rate of outflow of toxic
compounds for 10 minutes was 3-5% of the accumulated amount.
Short periods of gassing and low S02 concentrations activate the
photosynthesis (absorption of C^-^O^) » whereas higher concentrations of the
gas and longer exposures to it cause a decrease in the intensity of photo-
synthesis.
Table k
Absorption of Sulfur Dioxide by The Plants, Pulses per Bin
(1 g of Dry Weight)
Species
Exposure to S^O^
10 nih
10 mil
-r23»lr
up to -,-
fixation
I to-
1 hr +
23 hr.
up to
fixation
1 hr +
72 hr
fixation
Meadow fescue
48630
45300 115750 122950
112600
Timothy grass
166800 156350 166043 —
Sulfur dioxide absorbed by the plants undergoes various chemical trans-
formations. According to M. D. Thomas, R. H. Hendricks, G. R. Hill (1944),
Krpker (1950), and Yang and Mou (1961), S02 in the plants is oxidized to
- 86 -
-------
sulfate (SO,) and is partially utilized in the synthesis of amino acids and
proteins. Our experiments (Table 5) showed that in the fescue up to 88%
and in the timothy up to 95% of the labeled sulfur is contained in the alco-
hol- soluble fraction, only slight amounts being present in starch, hemi-
cellulose, proteins, and cellulose. It may be assumed that in the latter
compounds sulfur is tied not chemically, but by adsorption and electro-
static forces.
Chromatography of the alcohol-soluble fraction in the system phenol-
water (80:20) and butanol - formic acid - water (75:13:12) showed that the
labled sulfur remains almost entirely in the starting spot. Slight indi-
cations of the advancement of the sulfur with monosaccharides were noted
only in the fescue. Since these forage grasses, when subjected to short-
duration exposures , showed no sulfur-containing compounds in any of their
organic products formed during photosynthesis, one may assume as well that
in the alcohol-soluble fraction, the sulfur is adsorbed on different organic
substances. It is also possible that a certain insignificant amount of
sulfur enters into the composition of readily water- and alcohol-soluble
proteins and their amino acids (cystine, methionine).
Since the incorporation of C- into the indicated groups of organic
compounds in the forage grasses is similar, we shall consider the differ-
ences in the metabolism of individual compounds (Table 6) . In short ex-
posures of the fescue the labeled carbon is concentrated in fructose-1,
6-diphosphate , sucrose, alanine, and citric acid. In long exposures
(I1 + V), the labeled carbon also appears in all the other compounds. In
the timothy, the labeled carbon concentrates in the sugars and alanine even
in long exposures. Thus, forage grasses which differ in gas resistance
also differ in the nature of the metabolism of organic compounds.
The resistant species (meadow fescue) is characterized by a high rate
of synthesis of various organic compounds at a comparatively low absorption
rate of C^02- T^e nonresistant species (timothy) is characterized by a
high intensity of gaseous exchange and low rate of transformation of organic
compounds. If we are to imagine that instead of (XL the plants will assimi-
late S02, as is sometimes the case at industrial enterprises, then the role
of the chemical processes of photosynthesis in the gas resistance of plants
becomes understandable. The oxidation of the anion S02 to the sulfate pro-
ceeds faster in the meadow fescue than in the timothy, thus decreasing the
toxicity of the sulfate. On the other hand, the great variety of the photo-
synthesis products (a more limited specialization of metabolism, according
to A. V. Blagoveshchenskiy, 1950) apparently plays an important part in
blocking the noxious action of sulfur dioxide.
Since labeled sulfur is not incorporated into any of the organic com-
pounds, in which carbon is incorporated at the same exposures, the data of
Table 6 on the activity of the products of metabolism in variants involving
- 87 -
-------
. can be attributed only to the incorporation of carbon C in them.
Table 5
Results of General Radiochemical Analysis of the Distribution
of Sulfur in
Experimental
Conditions
. Distribution of S*'-O2 over groups of
organic compounds H
Initial
lilry wfienb
6"
OH
•g-g
V*40
3Z
f.
g
cd
8
\ .
i-H
.sa
SO
^3
in
-S
o.
t,
tSo>
3 S
i-l VI
«-H
OE-I
•g
fr< O
10 min S3SO2
1 hr SM0,
1 hr SM02
+23 hr
10 min
10 nun
+23 hr
1 hr S*02
MeadQTt
48650 84,7 4.1 5,4 4.1 1,5 99,8
118905 84.7 1,9 6,5 .3.6 0.9 97.6
122950 88.2 2.4 2.8 5,1 1.1 100
Timothy Grass
166800 84,1 5,3 2,7 7,1 0,9 99,8
156350 94.9 1.4 0,9 2.7 0 99.9
166043 93,3 — — — —
Under the influence of short exposures to S02 (15" S3502 + 1'
the synthesis of complex sugars (sucrose, raffinose) and alanine is re-
tarded in the meadow fescue. In the meadow fescue, under the influence of
long exposures to S02 (51 S^02 + 1' C^02) , the synthesis of glucose,
fructose, raffinose, and asparagine is intensified, whereas that of sucrose,
a-alanine and citric acid is reduced. In the variant with an additional
four-minute illumination, a decrease in the activity of C^ is observed in
glucose, fructose, and particularly sucrose and 3-alanine (Table 6). Thus,
despite the action of sulfur dioxide, the synthesis of all the organic com-
pounds is preserved in the fescue.
In the timothy, a prolonged exposure to sulfur dioxide (51 s35Q2 + 1'
) causes more serious disturbances in the metabolism qf organic com-
pounds: the synthesis of fructose-1, 6-diphosphate , fructose, and citric
acid decreases considerably, and there is an acceleration in the synthesis
of glucose, sucrose (by a factor of 10), a-alanine, and malic acid. An
additional four-minute illumination causes C^ to concentrate in sucrose.
As in the control variant, the timothy synthesizes a limited number of com-
pounds, in contrast to the meadow fescue.
Even in very long exposures to SO,. (1 hour) , the meadow fescue retains
a close- to-normal metabolism of organic compounds despite the presence of
visible damage.
- 88 -
-------
Tible 6
Metabolism of Carbon C^ Compounds in Forage (i
oo
VD
Meadow fescue
Meadow fescue
Meadow fescue
Timothy
grass
Timothy
grass
Meadow fescue
Meadow fescue
Meadow fescue
Meadow fescue
Meadow fescue
Timothy
grass
Timothy
grass
Meadow fescue
Meadow fescue
Species
Exposure
Distribution 'of Cw Over Individual Compounds, Percent
Starting
Spot
i *
•POO.
oJ>o
8
o
o
3
r-t
C3
Fructose
Sucrose
Baffinose
ri CU
CO u
V) bO
cool
•JJC
o
OI<1>
CC
O.H
m.
•Eo
•HO
O
edO
6-10
15"C"0,
l'C'«02
rC'«0,+4' light
l'S»Oj-f
^ligh^
+4'J light
-f 4' light
1 hrv S0j+
1 hr SO,+
3.5
1.6
1 1.0
8.1
4,6
24,3
50,0
21,2
7,2
3.3
N.2
30,0
32.7
41,9
55.8
21.7
55,1
24.8
43,0
30.0
17.6
45.2
—
—
1.0
—
7.6
1,7
3,2
—
5.9
5.3
3.1
—
1.9
3.3
11,3
4.4
—
—
8.5
5,3
8.7
3.1
17,2
42,5
18.9
2.4
51.7
5.9
5.8
11.4
21,0
36,5
20.7
—
—
6,0
—
—
4.5
3.5
—
13.6
4.0
__
—
—
1.8
1.5
_
—
3.5
—
5.6
3.8
-_
34.0
18.4
13.9
6,3
10,0
8.1
7.0
8.0
II. •>
13,1
10
15.3
2.9
1,3
0,9
— 14.6 —
— — 2.0
3.4
4.3
4,5
2.7
0.7
3.7
100
100
100
100
100
100
100
100
100
100
100
12,2 30.2 44.6 — — 9.5 — — 3.7 100
16.7 31.1 — 5.0 6,2 4.2 5.0 28.0 — 3.6 - 100
10.2 17,5 — 4.6 43.7 10.0 2.5 11.5 — — — 100
-------
Conclusions
1. Forage grasses differing in gas resistance also differ in their
content of oxidizable substances and ascorbic acid. A resistant species
(meadow fescue) is characterized by a reduced content of oxidizable sub-
stances and ascorbic acid. Under the influence of S02» an increase in the
content of oxidizable substances is sometimes observed.
2. The meadow fescue as compared with timothy is characterized by a
reduced intensity of photosynthesis and a lower rate of absorption of SC^.
3. When plants are fumigated with sublethal and lethal doses, the
sulfur isotope concentrates in the alcohol-soluble fraction (up to 88-95%).
A slight activity is observed in starch, hemicellulose, and proteins, and
a very slight activity in cellulose.
4. Most of the sulfur accumulated in the plant appears to be in
inorganic form, since it cannot be separated by paper chromatography and
remains almost entirely in the starting spot.
5. The forage grasses studied differ in the nature of the metabolism
of carbon C-^ compounds under normal conditions in long exposures (5 minutes
or more). The meadow fescue is characterized by a great variety of synthe-
sizable substances (unspecialized type of exchange, according to A. V. Blag-
oveshchenskiy); on the other hand, the timothy synthesizes a limited number
of products (spezialized type of exchange).
6. Under the influence of S0«, definite changes in the metabolism of
carbon in forage plants are observed which are attributable to their gas
resistance.
90
-------
LITERATURE CITED
Ap iion C. 1959. HaoTomibie MCTOAM B 6iioxn.Miiii. M. Haa-Do wiioctp.
.•HIT-Dbl, M.
5.1 a r o B e m e H c K n A A. B. 1950. BIIOXHMIIMCCKIIC OCIIOBU auojiioiiiiOH-
noro npouccca y pacTcimfi. AH CCCP, M.—Jl.
E p M a K o B A. C., A p a c H M o B H M B. B. H ap. 1952. MCTO,IU BHOXIIMH-
MccKoro iicc.KMOBaimfl pacTeniifi. Ce.ibXO3ni3. M.
HjibKyn F. M., MoTpyK B. B. 1968. ii3iio-ioro-OiioxnM)i'iecKiie na-
pyuiciiiiH B- pacTeiiiinx, nbisbmacMbie aTMoc4>epiiUMii 3arpH3iiHTCJiJiMH. Mare-
pna.TH Ilepnoii yKpaimcKoit K0iic()cpciiuiiii «PaCTcm»i H npoMNUi.iemiaa cpe-
aa>. «HayKoua 4yMiCP H TopbK. yii-ra, Mo-
CKBa — FopoKiifi.
K p o K c p B. 1950. POCT pacTCimft. H3,i- Hiioctp. ;IHT., M.
Ky .1 a r H n K). 3. 1965. rasoycToiViiioocTb H aacyxoycioiVniBocTb .apeacc-
liwx nopo.i. Tpy,iw Ilu-ia Ciio.ioriui yAH, nun. 43.
Ky.iarnii IO. 3. 1966. Bo.tiiuft pc/KiiM H rasoycTofi'iiiBoCTb flpcBccsibix
pacTeniifi. C6. c-Oxpana npiipoaw na Vpa.iea. V<1>AH, nwn. 5.
M o K p o ii o c o B A. T. 1966. ricKotopue nonpocu MCTO.IIIKII npiiMc;ie;iii3
ii.-CTona yncpo.ia-14 A-IH iinyieinin (l»OTOciiiiTc:ia. SaniiCKii CncpA-tOBCKoro
e.ietiitfl EDO, Bbin. 4, Cncpi.ioncK.
H n K o .1 a c B c K H ft B. C. 196.1 O iioKasarc.mx raaoycToii'iimocTii .apo-
iiiiix pacroiuifi. Tpyji,i Hii-ra ono.i. VAH. nun. 31. Cncp.vioucK.
H H KOKypn.
«KoncepBiian H OBomccyuiiubiian npoM.», tf« 5.
Hur Jl., Moy fl. 1961. McTaGo.iimi coeAiiiicinifi ccpu. HSJ-BO imocip.
.HIT., M.
G o d z i c S. 1968. Pobicranie i rozmieizczenie SMOj w organach asymi-
lacyjnych kilku gatunkow drzew. Matcrialy VI Micdzynarodowci konterencji
«\Vplv\v zanicczyszczen powietrza na lasy. Katowice.
fi a r r i s o n B. F., T h o m a s M. D., H i 11 G. R. 1944. Radioautographs
showings the distribution of sulfur in wheat. Plant Physiology, vol. 19. N 2.
Kisser J. 1968. Physiologisch: Problemc dcr Eimvirkung von Luftverun-
reinigungen auf die Vegetation. Matcrialy VI Micdzynarodowei konfercncjt
«\Vplyw zanierzyszczen powietrza na lasy». Katowice.
Thomas M. D., H e n d r i c k & R. H., H i 11 G. R. 1944. Some chemi-
ca! reaction of SOZ after absorption by alfalfa and sugar beets. Plant
Physiology, vol. 19, N 2.x
Thomas M. D., Hendricks R. H. Bryner L. C. Hill G. R.
1944 A study cf-ual Sulfur metabolism of wheat, barley and corn using
n-dioactive Sulfur. Plant Physiology, vol. 19, N 2.
- 91 -
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EFFECT OF SULFUR DIOXIDE ON PIGMENTS OF FORAGE GRASSES
V. S. Nikolayevskiy and V. V. Suslova
From Ministerstvo Vysshego i Srednego Spetsial'nogo Obrazovaniya RSFSR.
Permskiy ordena trudovogo krasnogo znameni gosudarstvennyy universitet im.
A. M. Gor'kogo. Uchenye Zapiski No. 222. "Gazoustoychivost1 Rasteniy".
Vypusk 1. Perm', p. 99-114, (1969).
According to K. Noack (1920, 1925) and N. P. Krasinskiy (1950), a def-
inite role in the injury to plant leaves by sulfur dioxide is played by pig-
ments, which continue the absorption and accumulation of solar energy despite
the discontinuation of photosynthesis. These authors explain the formation
of necroses on leaves exposed to S02 by the development of photodynamic oxi-
dation processes the energy for which is supplied by the pigments. Along-
side the destruction of the cell content under the influence of SOn, the
pigments in the leaves are also destroyed by this gas. Acted upon by S02
and light, the chlorophyll in the leaves is converted into pheophytin (Jahnel,
1954; Nemec, 1958; Dorris, cited by Tomas, 1962). The decomposition products
of pigments apparently also participate in the photodynamic oxidation of the
cell substrate, since A. A. Krasnovskiy (1959) established that pheophytin is
also capable of binding luminous energy, and G. P. Brin (1959) and V. B. Yevs-
tigneyev (1962) showed that pheophytin is capable of sensitizing oxidation
processes. Pheophytin was also found to have even a greater oxidizing effect
than chlorophyll.
In order to determine the chemistry and mechanism of photodynamic oxi-
dation processes in plants under the influence of S02 in light, it is impor-
tant to study the influence of this gas on pigment systems. Thus far, no
detailed investigations along these lines have been made in the USSR or in
other countries. From very scanty literature sources (Krasinskiy, 1950;
Jahnel, 1954; Spaleny, Godny, and Marzhan, 1962) it is known that under the
influence of a steady action of acid gases, the chlorophyll content of plants
decreases. The degree of reduction of chlorophyll is directly related to
the vulnerability of the plants.
According to modern concepts (Sapozhnikov et al., 1962; Sapozhnikov,
1963; Khodzhayev, 1963), not only chlorophyll but also carotinoids partici-
pate in the absorption of luminous energy. It is assumed that carotinoids
(the system lutein-violaxanthin) participate in the transfer of oxygen in
one of the intermediate steps of photosynthesis.
Plant pigments act not only as acceptors of luminous energy, but also
participate in the regulation of growth and development. An increase of
xerophytization of leaves, reduction of growth and yield, and disturbance
of the stages of development, noted by several authors (Antipov, 1957;
- 9Z -
-------
Nikolayevskiy, 1964; Babkina, 1968), may be due not only to a reduction of
photosynthesis but also to a disturbance of the form-developing role of
pigments'.
The degree of resistance of plant pigments to the action of extreme
factors (light, temperature, acids) depends not only on the chemical struc-
ture of the molecule, but also on the type of their bonding with the protein-
lipoid complex in the plastids. In the active monomer form, chlorophyll is
less resistant to the action of light and acids than in the aggregate form
(A. A. Krasnovskiy, 1959).
By decreasing the pH of the protoplasm and organelles in plants,
sulfur dioxide enhances the enzymatic oxidation processes (Nikolayevskiy,
Suslova, 1968; Nikolayevskiy, 1968). This apparently causes the rupture of
labile bonds between the pigments and the protein-lipoid complex and the
formation of monomeric and even molecular forms of the pigments, causing a
decrease of their resistance to light. Evidence of the above is provided by
the appearance of necrotic blotches in plants under the influence of SC>2 in
light, and their absence from plants in the shade (Kroker, 1950; Krasinskiy,
1950; Jahnel, 1954). In experimental studies, in order to reveal the injuries
more rapidly, it is necessary that plants fumigated with sulfur dioxide always
be placed in direct sunlight.
From the theoretical,point of view, in order to determine the mechanism
and chemistry of the 862 action on green plants, it is important to study
the dynamics of the process of destruction of pigments and change of their
photodynamic role. It is more likely that the pigments are not destroyed
immediately after coming in contact with SO^, but gradually, during the
parallel effect of S02 and light. For this reason, necroses appear only 1 to
24 hours or more after the plants have been placed in direct light. Less
likely is the hypothesis that pigments are destroyed only during the final
stages of the death of cells. The elucidation of the mechanism by which
chlorophyll is revived (Bazhanova et al., 1964; Shlyk, 1965) in the process
of metabolism makes it necessary to assume an increase in the breakdown of
pigments and an impairment of their synthesis as a function of the SC^ con-
centrations, light, and time elapsed since the gassing of the plants.
Among the objectives of our investigation were those of studying (a) the
resistance of individual plant pigments to acidification of the protoplasm as
a result of accumulation of S02, and (b) the stability of the pigments as a
function of the stages of growth, development, and gas resistance of the
plants.
Method of Investigation
The studies were conducted on the meadow fescue (Festuca pratenis Huds) -
a gas-resistant species, and timothy (Phleum pratense L.) - a nonresistant
- 93 -
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species. These forage grasses were grown at the university's botanical
garden on poor sandy soil. During the tillering stage of the two species
and during the booting and blooming stages of timothy, the plants were
fumigated with sulfur dioxide in a 0.125 m3 polyethylene chamber for 1 hour.
The initial S02 concentrations (2.5 x 1(T5; 5 x 10~5; 1Q-4; 2 x 10~4;
4 x 10~4 by volume) were produced by reacting an exact amount of Na2S(>3
with sulfuric acid according to the equation
Na2S03 + H2S04 = Na2SC>4 + S02 + H20.
The composition of the pigments in the experimental and in the con-
trol plants was studied immediately after the gassing, and a second time
after 24 hours. The second and third leaves, counting from the top of the
plant, were used for the analysis. The pigments were separated by paper
chromatography according to Sapozhnikov (1964), and the quantitative determ-
inations were made on FEK-M. The calibration curves for FEK-M were plotted
by determining a series of pigment solutions on SF-4a and FEK-M. Since
after the fumigation and injury of the plants with sulfur dioxide a decrease
in the water content of the leaves was observed, it was necessary, in order
to compare the data for the first and second days of the experiments, to
determine the moisture content of the leaves and to take into account a cor-
rection for the change of the moisture content.
The vulnerability was determined by measuring the injured and total .
length of the leaf with a ruler. The ratio of the former to the latter (in
percent) was taken as the vulnerability.
Results of Investigation
Since the meadow fescue in its first year of life goes only through
the tillering stage, while the timothy goes through all the stages of growth-
development up to blooming and fruiting, the pigment composition and the
effect of sulfur dioxide on it were studied in the first species during the
tillering stage and in the second, during the stages of tillering, booting,
and blooming.
During the tillering stage (Table 1) in the meadow fescue, chlorophylls
a and b account for 82.6%, carotene for 1.3%, and xanthophylls, for 16.1%
of the total content of pigments in the leaves. Among the pigments entering
into the composition of xanthophylls, the predominant one is lutein - 48%.
The ratio chlorophyll a (|£} ±n ^ meadow fescue ±8 4>15;
chlorophyll b Xb
chlorophyll a + b /XaJMbs _ 4>y.
yellow pigments y. p.
xanthophylls , X. N , _ _
" r ( ) = J.J.U.
carotene Ncar.
- 94 -
-------
Thus, among chlorophylls, the reduced form, chlorophyll a, predominates,
and among yellow pigments, the oxidized forms - lutein, violaxanthin and
neoxanthin are predominant.
The pigment composition of the timothy during the tillering stage is
very close to that of the meadow fescue (Table 1), with the exception of
the ratio x' , which was found to be somewhat smaller.
car.
Hence, forage grasses which differ in their resistance to SO- show
no substantial differences in the pigment composition during their tiller-
ing stage.
In the timothy (Table 1), a change in the pigment composition of the
leaves can be detected during its growth and development. During the booting
stage as compared with the tillering stage, the amount of chlorophyll^ de-
creases somewhat, and the content of chlorophyll b_ increases, the total
chlorophyll content remaining the same. Thus the ratio Xa/Xb decreases to
3.6. A certain decrease also takes place in the ratio X/car. during the
booting stage, because of an increase in the carotene content. At the same
time, the total absolute content of pigments during the booting stage in-
creases by 34% as compared with the tillering stage, and increases by 79%
during the blooming stage.
During the blooming stage of timothy, the ratio of green pigments
Xa/Xb = 4.6 is once again restored. The ratio X./car. increases to 12.6.
Hence, in the timothy, a comparatively stable pigment composition is ob-
served during the ontogeny. The ratio X./car. changes only slightly, and
the ratio X-a+b/y.p. remains almost unchanged.
During the tillering stage, we chose such SC^ concentrations that it
would be possible to obtain a vulnerability of 0 to 100% in both species.
Subsequently, these SO^ concentrations were also used in other stages of
growth and development.
Figure 1 shows the change in the vulnerability of the meadow fescue
and timothy as a function of the stages of growth, development, and gas
concentrations. It is evident from Fig. 1 that during the tillering stage,
the leaves of the timothy accumulate 862 up to lethal 'doses more rapidly
than those of the fescue. Hence, S02 causes the poisoning and death of the
leaf cells and tissues more rapidly in the timothy than in the meadow fescue.
However, at S02 concentrations of 2.5 x 10~5 and 4 x 10~4, the vulnerability
of both species is similar: in the former case it is 0%, and in the latter,
92 and 93%.
As the timothy grows and develops, a regular increase in the vulner-
ability of leaves is observed at the same S02 concentrations (Fig. 1); this
may be explained, on the one hand, by an increase in the lethal effect of
- 95 -
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Table 1
Seasonal Dynamics of Pigments in Forage Grasses
I
VO
Species and
Stages
Amoxmt of Pigments- in ng i»r g of Green Weight
Chlorophyll
a
Chlorophyll
b
Chlorophyll
a + b
Carotene*...
'§
ll
•g»
Neoxanthin
Total xan-
thophyllis '
Total
Yellow
Pigments
A
4,6
4,7 13,0
4.6 8.7
4.5f> 7,3
4.3 12.0
-------
S02 concentration
Fig. 1. Change in the vulnerability of forage grasses as a
function of the stages of growth and development and concen-
tration.
Notation: 1 - meadow fescue - tillering; 2 - timothy - tiller-
ing; 3 - timothy - booting; k - timothy - blooming.
the toxic gas, and on the other hand,, by an increased rate of gaseous ex-
change- and hence, a greater accumulation of S(>2 during an equal time interval.
The decrease in the gas resistance of the timothy when the latter enters the
stages of booting and blooming becomes understandable if one considers that
an intensification of the physiological-biochemical processes takes place in
the plants during these stages. Earlier, we established (Nikolayevskiy , 1964)
a direct correlation between the intensity of photosynthesis and respiration
in woody plants and their vulnerability to sulfur dioxide.
Let us ' consider the effect of SCL on the pigment composition of the
meadow fescue according to the stages of growth and development and as a
function of the gas concentration and vulnerability of the plants (Table 2) .
For convenience of comparison, the tables give the content of pigments in
percent qf the control. In the meadow fescue during the tillering stage, the
lowest SC»2 concentration, 2.5 x 10"^, caused a decrease in the content of
carotene, violaxanthin, and neoxanthin on the day of the experiment. In addi-
tion, a slight increase was observed in the content of chlorophyll b_ and in
the ratio of the different pigments to each other Xa/Xb, Xa + b/y. p. X. /(car.),
close to those in the control. Because of the destruction of yellow pigments ,
the ratio
+ b
.y- P-
and
car.
increased somewhat in the experiment.
An SCL concentration of 5 x 10~5 caused on the first day a decrease in
•carotene content and increase in the content of violaxanthin and neoxanthin.
As before, an increase in the ratio X/car. was observed in this case. More
pronounced changes were observed in the experimental plants on the second day.
- 97 -
-------
Thus, as compared with the control plants, the content of pigments was
higher, namely: chlorophyll 9-26%, carotene 55%, lutein 30%, violaxanthin
92%, and iieoxanthin 132% higher than in the control. As a result, the ratios
of the pigments (Xa/Xb; Xa 4- b/y. p.; X/car.) decreased.
In the meadow fescue, an SO* concentration of 10~^ caused a reduction of
50% in chlorophyll a. and b_ and of more than 50% in yellow pigments. The caro-
tene content remained unchanged. As a result, the ratio X/car. decreased to
10.8. On the second day, a certain increase in xanthophylls was observed as
compared to the first day of the experiment. An S02 concentration of 2 x 10~4
on the first day caused in the meadow fescue a slight increase in the content
of chlorophyll a_ and b_, carotene, and neoxanthin and a decrease in the con-
tent of violaxanthin and lutein. On the second day, as in the preceding case,
an increase in the content of all the pigments, particularly marked in the
case of chlorophyll _a and b_, lutein, and neoxanthin, was observed.
The highest SO. concentration, 4 x 10~4, which caused an almost complete
destruction of the plants, decreased the contents of all the pigments on the
first day, but on the second day an increase of their content was observed.
The latter phenomenon may be explained by an increase in the extractability
of the pigments as a result of hydrolysis of the proteins in the protoplasm
and plastids.
Thus, in the meadow fescue during the tillering stage, the change in the
content of pigments stands in some direct relationship to the SOo concentra-
tion. The yellow pigments are destroyed more rapidly and more severely under
the influence of $$2* an^ among these, carotene is destroyed at a particularly
rapid rate.
On the second day after the action of SO., an increase in the content
of pigments was observed in the meadow fescue. The increase in the content
of chlorophyll b_, lutein and neoxanthin was more pronounced. The carotene
content at low S02 concentrations also increased, and at high concentrations
remained unchanged on the second day. Under the influence of SO-, the appear-
ance of pheophytin was observed in the plants as indicated by the chromato-
grams.
In the timothy during the tillering stage, the first SO- concentration,
2.5 x 10~5, caused a substantial destruction of chlorophyll £ and b_ and a
certain decrease in lutein and increase in neoxanthin content on the first
day of the experiment. The ratio Xa + b/y. p. decreased. In the timothy,
on the first day, the 5 x 10~5 S02 concentration caused a decrease in the
content of chlorophyll ^, lutein, and neoxanthin, and an increase in the con-
tent of chlorophyll b_ and violaxanthin. On the second day, the reduction in
the content of chlorophyll SL and b_ continued, and the lutein content increased.
In the timothy, the 10~^ SO^ concentration caused a destruction of chloro-
phyll b^, lutein, as well as of nteoxanthin, and an increase in the content of
- 98 -
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Table 2
Effect of S02 on the, Pigments of Forage Grasses
Sog Con-
centra-
tion
I!
Q*>
•l.
8S
EH «>
•H
.0
O5
.'5
^
Amount of Pigments in 1 g
of Green Weight, # of Control
rH
1
2 '
o
1-1
fi B
.-1
1
o
o
g^ol
g
a)
O 1 C_>
.9
0)
4>
S
^
4>
a
o
•rl
>"
1
§
g
01
Ratio
Xa
Xb
Ratio
Xa+b
y. p.
Ratio
X
car.
2.5X10-9 1 hr. 0 98
5X10-S Ihr.— 100
24 hr. 13.8 109
10-« 1 !"•• 46
24 hr- 37
.2X10-4 Ihr.— 114
24 hr. 60 147
4X10-4 Ihr.- 81
24 hr. 92 180
2XlO-» 1 hr. 0 59
5XIO-5 Ihr.— 95
24 hr. 41.1 83
10-« Ihr.— 116
24 hr<55,6 -
Fescue - tillering
108 55 94 64 69
95 73 99 129 120
126 155 130 192 232
55 100 46 35 44
88 57 75
110 108 88 89 111
255 — 130 93 160
81 — 99 81 81
205 - 235 230 174
Timothy - tillering
58 — 92 105 112
118 100 86 160 80
76 — 105 106 82,5
89.5160 81.5 114 61
— 208 116 124 102.5
2XlO-< 1 hr.— 104 103 81 163 162 95.0
3.65
3,34
4,3
4,5
3,95
3.4
45
4.1
4,1
4,25
6,0
3.5
3.6
3,6
3,5
3.1
3,95
4.0 "
5.65
4.55
4,25
4J
3.35
4.3
—
4.4
4,45
5.3
6.9
5.0
4.5'
6.2
4725
4-7
4,9
3,88
4,6
—
4-9
6,2
8.2
4,8
4,7
4,5
5,1
3.1
3-7
—
5.3
4,15
10.2
14.4
12.1
18,5
9.2
9.6
24.7
10,8
9,4
8.1
_
7.3
19.5
7.8
8.2
8,3
11,0
15.4
8.4
J4.7_
15.5
9,6
- 99 -
-------
Table 2 (Cont'd)
Effect .of SOp on the Pigments of Forage Grasses
S02Con-
tion
W»H
' EH «
Vulnerability,
Amount of Pigments in 1 g
of Green Weight, % of Control
Chlorophyll
a
Chlorophyll
b:
Carotene
.5
*
|
Violaxanthii
-
4*
2
Ratio .
Xa
Xb
Ratio .
Xa+b-.
y. p.
Ratio
X
•n
24 *r. 90.7 86,5 230 0 100 87,5 96,0 -f—
4X10-" 1 ;hr.— 81-6150 — 112 102 89
24 AT. 93,0 123 228 — 140 81 130
Timothy - booting
5XIO-5 1 -hr.— 79 98 — — 73 116
24 to. 58,3 60,6119 — 60 56,593
10-* 1 hr._ 100 70 56 109 84 77
24 hr. 82 64 58 — — 134 —
2X10-- 1 hr.— 96 100 100 68 83,5107
24 hr. 56 75 65,5 — 59 7& 70
Timothy - blooming
2X10-* I hr. — 68 50 — 70—80
24 hr. 82 108 100 — 95 93 72
5X10-5 1 hr. — 57,5 41 — 64 71.543
24 -hr.100 42.2 67.5 — 36 27.4 44
10-« 1 hr. — 108 89 71.8127 132 75.8
24 Iff. 100 56 89,5— 79 96.5150
Note. In the last three columns, the control is in the numerator and
the experiment is in the denominator.
A5_
I,/
7,4
3,7
_¥_
3,95
3.15
S3
3.25
3.0
4,1
3.3
4.4
4.2
2,9
3.3
4.16
5,7
4.03
4.4
5.6
7,8
3.7
2.4
4.0
3.7
—
3.8
4,6
4.1
. _,
4-9
5,1
5.3
4.8
5.5
—
5.6
5.1
3.6
4,1
3.9
4,2
14.0
9.2
13.5
5.8
_^
11.2
18.6
28,5
-7.0
5.6
10,0
13.2
22.0
14.3
8.2
13.1
11.6
- 100 -
-------
chlorophyll a_t carotene, and violaxanthin. On the second day, an increase
in the content of all the yellow pigments was observed. The 2 x 10~^ S02
concentration caused a decrease in the content of carotene and neoxanthin,
and an increase in the content of the remaining pigments. In the timothy,
on the second day, there was an increase in the content of chlorophyll b_
and a decrease in the content of the remaining pigments.
In the timothy, on the first day, the highest SO- concentration,
4 x 10~4, caused a decrease in the content of chlorophyll _a and neoxanthin.
At the same time, an increase in the content of chlorophyll b and lutein
was observed. On the second day, an increase in the content of all the pig-
ments except carotene and violaxanthin was observed.
Thus, in contrast to the meadow fescue, low S02 concentrations cause
more serious changes in the composition and ratio of the pigments in the
timothy than high concentrations. Chlorophyll a and b_ are destroyed more
extensively in the timothy by low S02 concentrations. High S02 concentra-
tions have a stabilizing effect on chlorophyll. However, after the action
of low S02 concentrations, 5 x 10~5, the chlorophyll content continues to
decrease on the second day, whereas its content increases after the action
of high S02 concentrations (4 x 10"^). In the timothy, in contrast to the
meadow fescue, low S0_ concentrations do not destroy the carotene on the
first day, and act most destructively on the second day. The effect of high
SO- concentrations on timothy and meadow fescue is similar. In the timothy,
S02 has a lesser influence on xanthophylls than in the meadow fescue. At
the same time, the increase in the content of xanthophylls on the second day
after the action of S02 is more pronounced in the meadow fescue than in the
timothy.
During the booting stage of the timothy, sulfur dioxide in the concen-
tration of 5 x 10~5 destroys lutein completely, chlorophyll ^ partially,
and chlorophyll b_ only slightly, and increases the content of neoxanthin.
On the second day, a decrease is observed in the content of chlorophyll .a,
violaxanthin, and neoxanthin, and an increase in the content of chlorophyll b_
and lutein.
In the timothy, on the first day, sulfur dioxide in the concentration
of 10"^ caused a decrease in the content of chlorophyll b_, carotene, violax-
anthin, and neoxanthin. On the second day, a decrease in the content of all
the pigments except violaxanthin was observed.
In the timothy, sulfur dioxide in the concentration of 2 x 10~^ caused
a stabilization of the content of chlorophyll a. and b_ and carotene, and a
decrease in the content of lutein and violaxanthin. On the second day, a
decrease in the content of all the pigments was observed.
Thus, during the booting stage of the timothy (as during the tillering
stage), chlorophyll, carotene, and lutein are more extensively destroyed by
- 101 -
-------
low S(>2 concentrations. The most stable of the yellow pigments during
this stage is neoxanthin.
During the blooming stage of the timothy, low SO* concentrations,
2.5 x 10~5, cause a more substantial destruction of chlorophylls JL and b_
and yellow pigments. On the second day, an increase in the content of the
pigments is observed.
The 5 x 10~5 S02 concentration causes a decrease in the content of all
the pigments in the timothy on the first and second day. However, the con-
tent of chlorophyll b_ increased on the second day. In the timothy, the
10~4 S02 concentration caused a decrease in the content of chlorophyll b_,
carotene, and neoxanthin and an increase in the content of chlorophyll _a,
lutein and vlolaxanthin on the first day; on the second day. there was a de-
crease in the content of all the pigments except neoxanthin.
Thus, during the blooming stage of the timothy, the stability of chloro-
phylls and xanthophylls is similar. On the first day of the experiment,
chlorophyll _a is more resistant to the action of S02» but on the second day
its resistance is lower than that of chlorophyll b_.
Discussion
The two species of forage grasses which we studied, meadow fescue and
timothy, show significant differences in gas resistance during the tillering
stage. Under similar experimental conditions, the same SO2 concentrations
(with the exception of below-sublethal and above-lethal concentrations) cause
a greater vulnerability in the timothy than in the meadow fescue. The char-
acter of the vulnerability curves of the plants as a function of SOo concen-
trations during the tillering stage is different: in the meadow fescue the
relationship is more direct, proportional, and in the timothy, almost loga-
rithmic. It should be noted that in the meadow fescue, the range of S02 con-
centrations between the sublethal and lethal doses is wider than in the
timothy. This undoubtedly indicates a greater resistance of the meadow fescue
to S02.
During the ontogeny of timothy, one observes in connection with an
intensification of the vital processes, a decrease in resistance and hence,
an increase in vulnerability in the presence of similar SO2 concentrations.
This reaffirms the dependence of the gas resistance of plants on the morpho-
physiological rhythms, on stages of growth and development, and on the inten-
sity and direction of the physiological-biochemical processes in the leaves.
Despite the marked biological nature of the growth and development) and
the physiological-biochemical (gas resistance) difference between the two
plant species which we studied - meadow fescue and timothy - the pigment
- 1Q2 -
-------
composition of the leaves during the tillering stage is very similar. It
is possible that during other stages (booting, heading, blooming), differ-
ences in the pigment composition exist, but to find them it would be neces-
sary to continue the studies with the meadow fescue during its second year
of life. In the timothy, an increase in the concentration of the pigments
is observed during ontogeny; this is important in connection with the inten-
sification of the life processes in plants entering the reproductive stage.
It is possible that the revealed increase of pigment concentrations in
the timothy is only seeming to be so as a result of a decrease in the
strength of the bonding between the pigment and the protein-lipoid complex.
If such be the case, then one of the causes of the increased vulnerability
of the timothy during the booting and blooming stages becomes clear: the
increase in the sensitivity of the pigments to S02 and light during the boot-
ing and blooming stages results from a decrease in the protective influence
of proteins. This is associated with an increase in the extractability of
the pigments.
At the same time, the ratio of the pigments in the timothy during the
booting and blooming stages remains almost exactly the same as during the
tillering stage. The stability of the ratio of the various pigments in
the leaves during ontogeny appears to be a hereditary characteristic devel-
oped by evolution.
The effect of SOn on the pigment composition of the plants during the
tillering stage varied in the plants studied. To some degree this can be
related to the biology of the species and their differences in gas resistance.
In the meadow fescue, a more direct proportional concentration dependence of
the destruction of chlorophyll and vulnerability of the leaves is observed.
In comparison with the timothy, in the fescue the xanthophylls are destroyed
more extensively, and chlorophyll is comparatively more resistant to SO-.
The fescue displays more clearly its ability to increase the content of pig-
ments in injured leaves. The mechanism of restoration of the pigments in
injured leaves is unclear. Two explanations are possible: 1 - intensifica-
tion of pigment synthesis and 2 - increase in the extractability of pigments
under the influence of S02. The latter hypothesis is more plausible, since
the greatest increase in the content of pigments was observed on the second
day after the action of high S0~ concentrations, which appear to cause some-
thing like a fixation of the plant tissues and a high vulnerability. In this
case, there can be no question of the existence of complex processes of bio-
synthesis of pigments. More substantial differences in the change of the
pigment composition of the leaves in the two species of forage grasses are
observed in the effect of low SCL concentrations, i.e., concentrations closer
to those acting on vegetation under industrial conditions.
There are some marked differences in the nature of the action of low
and high SO- concentrations on the pigments of plants. For the most part,
- 103 -
-------
low S0_ concentrations cause the destruction of many pigments even on the
first and second day (timothy) or an increase in the content of pigments
on the second day (meadow fescue). High SC^ concentrations seem to stabil-
ize (fix) the pigment composition. An increase in the content of the pig-
ments (xanthophylls in timothy) is sometimes observed. In the meadow fescue,
the increase in the content of pigments, with the exception of carotene, is
more pronounced on the second day. This type of effect of S0£ may be explained
by differences in the chemistry of decomposition of the pigments. It may be
postulated that low S(>2 concentrations cause an intensification of the enzy-
matic oxidation of pigments as a result of a slight acidification of the pro-
toplasm and intensification of the activity of hydrolytic enzymes. High S02
concentrations cause a considerable acidification of the pH of the protoplasm,
have an inactivating influence on hydrolytic enzymes, and promote a stabil-
ization of the pigment composition* In this case, by destroying the protein-
lipoid complex in the plastids, SO simultaneously causes an increase in the
extractability of the pigments, ana thus the values of the pigment content
which are obtained on the second day are too high.
During the booting and blooming stage of the timothy exposed to SO-, no
increase in the content of pigments is observed on the second day with the
exception of the 5 x 10~5 concentration (booting) and 2.5 x 10~5 (blooming).
This is obviously due to a change in the ontogenetic state of the plants.
It is possible that such an influence of SO2 on the pigments is also related
to the low regenerating capacity of the cellular structures in chronologically
old leaves. A further decomposition of the pigments at these stages on the
second day in the case of exposure to high SO* concentrations probably takes
place as a result of autolysis during the destruction of plastids.
Under the influence of SO , on the first and second day of the experi-
ment, pheophytin bands were observed on the chromatograms in cases of de-
crease of the chlorophyll content. In both cases, the destruction of caro-
tene and chlorophyll was due to oxidation processes induced by SOn. This
can be determined from the character and chemistry of the processes leading
to the formation of pheophytin from chlorophyll.
Conclusions
1. The forage grasses meadow fescue and timothy, which differ in gas
resistance, are also markedly different during the tillering stage from the
standpoint of the nature of the changes in their vulnerability in relation
to gas concentration. The fescue is characterized by an almost directly pro-
portional dependence, and the timothy, by .a logarithmic dependence.
2. During the ontogeny of the timothy, there is observed a regular de-
crease of the gas resistance and a decrease of the sublethal and lethal S02
concentrations as a result of intensification of the physiological-biochemical
- 104 -
-------
processes with the onset of the reproductive stage.
3. Timothy shows during ontogeny a general increase in the concen-
tration of the pigments without any appreciable change in their proportions.
This may be due to an increase in the extractability of the pigments at the
end of the vegetative period.
4. Under the influence of S02, certain differences in the destruction
of individual pigment systems are observed in the plant species studied
during their tillering stage. As compared to the fescue, xanthophylls in
the timothy are destroyed more extensively, whereas the stability of chlor-
ophyll is about the same. It may be assumed that the bonding between the
pigments and the protein in the meadow fescue is stronger than in the
timothy.
5. Low SC>2 concentrations apparently cause an intensification of the
enzymatic oxidation of the pigments, whereas high SO^ concentrations, by
inactivating the enzymes, cause a stabilization of the content of the pig-
ments .
6. During the stages of booting and blooming of the timothy, a lack
of restoration of the content of pigments was observed on the second day
after exposure to S0_. This can be regarded as a confirmation of our views
concerning the change in the ratio of pigment forms differing in stability
(aggregate and monomeric forms) during ontogeny and under the influence of
so2.
- 105 -
-------
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-------
46 THE SUSCEPTIBILITY OR RESISTANCE TO GAS
AND SMOKE OF VARIOUS ARBOREAL SPECIES
GROWN UNDER DIVERSE ENVIRONMENTAL
CONDITIONS IN A NUMBER OF INDUSTRIAL RE-
GIONS OF THE SOVIET UNION-A Survey of USSR
Air Pollution Literature
47 METEOROLOGICAL AND CHEMICAL ASPECTS
OF AIR POLLUTION; PROPAGATION AND DIS-
PERSAL OF AIR POLLUTANTS IN A NUMBER OF
AREAS IN THE SOVIET UNION-A Survey of USSR
Air Pollution Literature
48 THE AGRICULTURAL REGIONS OF CHINA
51. MEASUREMENTS OF DISPERSAL AND
CONCENTRATION, IDENTIFICATION, AND
SANITARY EVALUATION OF VARIOUS AIR
POLLUTANTS. WITH SPECIAL REFERENCE TO
THE ENVIRONS OF ELECTRIC POWER PLANTS
AND FERROUS METALLURGICAL PLANTS
-A Survey of USSR Air Pollution Literature
52 A COMPILATION OF TECHNICAL REPORTS ON
THE BIOLOGICAL EFFECTS AND THE PUBLIC
HEALTH ASPECTS OF ATMOSPHERIC
POLLUTANTS - A Survey of USSR Air Pollution
Literature
49 EFFECTS OF METEOROLOGICAL CONDITIONS
AND RELIEF ON AIR POLLUTION; AIR CON-
TAMINANTS - THEIR CONCENTRATION,
TRANSPORT, AND DISPERSAL-A Survey of USSR
Air Pollution Literature
53 GAS RESISTANCE OF PLANTS WITH SPECIAL
REFERENCE TO PLANT BIOCHEMISTRY AND TO
THE EFFECTS OF MINERAL NUTRITION - A
Survey of USSR Air Polution Literature
50. AIR POLLUTION IN RELATION TO CERTAIN
ATMOSPHERIC AND METORO LOG) CAL
CONDITIONS AND SOME OF THE METHODS
EMPLOYED IN THE SURVEY AND ANALYSIS
OF AIR POLLUTANTS-A Survey of USSR Air
Pollution Literature
Reprints from various periodicals.
A INTERNATIONAL COOPERATION IN CROP IMPROVEMENT
THROUGH THE UTILIZATION OF THE CONCEPT OF
AGROCLIMATIC ANALOGUES
(The Use of Phenology, Meteorology and Geographical
Latitude for the Purposes of Plant Introduction and the Ex-
change of Improved Plant Varieties Between Various
Gauntries.)
B SOME PRELIMINARY OBSERVATIONS OF PHENOLOGICAL
DATA AS A TOOL IN THE STUDY OF PHOTOPERIODIC
AND THERMAL REQUIREMENTS OF VARIOUS PLANT
MATERIAL
*C AGRO-CLIMATOLOGY AND CROP ECOLOGY OF THE
UKRAINE AND CLIMATIC ANALOGUES IN NORTH
AMERICA
D AGRO-CLIMATOLOGY AND CROP ECOLOGY OF PALES-
TINE AND TRANSJOP.DAN AND CLIMATIC ANA-
LOGUES IN THE UNITED STATES
E USSR-Some Physical and Agricultural Characteristics of the
Drought Area and Its Climatic Analogues in the United States
f THE ROLE OF BIOCLIMATOLOGY IN AGRICULTURE WITH
SPECIAL REFERENCE TO THE USE OF THERMAL AND
PHOTO-THERMAL REQUIREMENTS OF PURE-LINE VARI-
ETIES OF PLANTS AS A BIOLOGICAL INDICATOR IN
ASCERTAINING CLIMATIC ANALOGUES (HOMO-
CLIMES)
•Out of Print.
Requests for studies should be addressed to the
American Institute of Crop Ecology, 809 Dale
Drive, Silver Spring, Maryland 20910.
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