FISHERIES RESEARCH BOARD OF CANADA

Translation Series No. 1043

The problem of primary production in water bodies

By G. G. Vinberg and O. I. Koblents-Mishke

Original title: Problemy pervichnoi produktsii vodoemov.

From: Ekologiya vodnykh organizmov. Akademiya Nauk SSSR. Publ. by: Izdaterstvo "Nauka", Moscow. A collection of papers, No. 6, pp. 50-62, 1966.

Translated by the Translation Bureau (OK) Foreign Languages Division

Department of the Secretary of State of Canada

Fisheries Research Board of Canada Marine Ecology Laboratory

Dartmouth, N. S.

1968

23 pages typescript

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1

1 (50)

Paper No. 6 from: !Ecology of aquatic organisms!

G. G.Vinberg 0. I.Koblents-Mishke

(White Russian State University and Oceanology Institute of the Ac. of Sciences of the USSR)

Problems of primary production in water bodies

The primary production is the result of vital funct-

ions and activities of vast numbers of vegetable species

which, depending on their specific character, exist under

varied conditions. A lot of valuable information describing

the multiformity and specificity of adaptations developed

by higher and lower plant species has been accumulated by

biology. However, despite the diversity in morphological,

physiological and ecological properties of individual species

all vegetable organisms, without exception, play a definite

part in the biotic cycle by forming the first trophic level

in the utilization of solar radiation. In this respect, no

difference whatever exists between plant organisms.

nos-2oo— 10-9 1

2 (51)

Therefore, by measuring the vital function of all plants -

photosynthesis - and its intensity degree, we have the possibi-

lity of expressing magnitudes of primary production in strictly

objective and comparable units of matter and energy, quite ir-

respective of which vegetable organisms live under which given

conditions. It is precisely this special feature in research

methods for determining primary production which conferred on

them the outstanding position they hold in modern hydrobiology.

It is well known that studios in primary production

advanced considerably when the insuffiently sensitive oxygen

device for measuring the intensity of plankton photosynthesis

was supplemented by the radiocarbon method; the latter was pro-

posed in 1951 by the Danish plant physiologist and hydrobioLo-

gist Steeman-Nielsen.

Research into primary production ought to establish,

firstly, the magnitude of primary production characterizing a

given water body or region and, secondly, the factors and

conditions determining this magnitude.

The dependency of primary production on factors of lu-

minous energy, on the relation between photosynthesis intensi-

ty in the plankton and its chlorophyll or other vegetable pig-

ment content in various trophic and other circumstances, etc.,

is being assiduously investigated by field and laboratory methods.

Owing to the uniform and objective manner of expressing

primary production which can be given in weight units of the

assimilated carbon or in calories per space unit for a fixed

3 (si)

time, regardless of the species composition of the producers,

we gain the valuable working tool of regarding this magnitude

as the function of a limited number of variables; and these

are, in the first place, light and temperature conditions, pig- C .

ment content, trophlt possibilities favouring the development

of phytoplankton, factors of water mixing or water stability.

No other but this kind of functional interpretation of primary

production does justice to the phenomenon. On these lines, the

mathematical investigation apparatus suggests itself not only

as applicable, but as necessary; it is successfully utilized

in this domain by quite a few authors, such as Riley in USA,

Cushing, Tailing and Steel in England. So far, mathematical

solutions are given Cor particular cases only, but in our

opinion a theory is developing which, apparently, in a not too

distant future will permit to calculate the biological magni-

tude of primary production as a function of a certain number ( 5 2 )

of conditions.

The extent t(J which the research into circumstances

determining the primary production is considered to be of con-

sequence, is obvious from the fact that, for instance, at the

Oceanographic Section of UNESCO the problems are examined by

study groups of outstanding scientists invited for this purpose

from various countries. In October, the second session of the

group investigating conditions of luminous energy in the sea

was held in Moscow with the participation of the most prominent

hydro-opticians from USA, Sweden, France, and hydrobiologists

)4. (52)

working on problems of primary production: Steeman-Nielsen

(Denmark), Steel (Great Britain), Jeats (Australia), Saidjo

(Japan), as well as Soviet specialists on hydro-optics and

primary production of the plankton. A co-ordinated plan for

further investigations was drawn up (Report of SCOR-UNESCO

working group 17, 196i4).

Another international study group examined methods for

determining chlorophyll content in plînkton, and at the session

from June L. t;o June 6, 19614., in Paris, a recommendation on these

problems was adopted (Report of SCOR-UNESCO, 1960.

Systematic investigations concerning primary production

were originally developed in limnology, since measuring plankton

photosynthesis by the oxygen method is quite possible on great

stretches of freshwater bodies, due to the relatively higher

development of phytoplankton in this environment.

Studies completed up to 1959 were summarized by G.G.

Vinberg in a book , published in 1960 in Minsk, and they were

also contained in the compendium by Strickland which appeared

in the same year in Canada.

In the course of the la -st years, primary production

has been measured by many authors and in many water bodies. As

examples we mention the interesting results obtained by Y.Ï.

Sorokin, M.A.Salmanov, and I.V.Pototskaya (Pototskaya and

Tsyba, 1961) for the reservoirs of the Volga - Don cascade.

It WaS revealed that primary production increases in its move-

ment downstream from north to south. The highest primary

5 (52)

production ascertained in the reservoir of Tsimlyan (3570

kcal/m2 per annum) shows a correspondingly high fish production

(35,8 kg/ha. in 1961).

Abroad, too, a great number of research works concerning

primary production in the most varied pieces of water have been

published; among these we find, on the one hand, such investigat-

ions as measurements of photosynthesis underneath the ice in

the arctic lakes of Alaska (Hobble, 1964), primary production

in temporary water bodies of the Antarctic (Armitage, House,

1962), and on the other hand, there are studies in primary

production in tropical African lakes (Tailing, 1964).

In a general way, all these new data fell in line (53)

rather well with primary production limits established pre-

viously and characteristic of water bodies of varied types

(Vinberg, 1960). It seems that onlyaccording to data obtained

by the Danish authors Jonasson,A Mathiesen (1959), one should

raise the upper limit to 660 g C/m2 or 7000 kcal/m2 per annum.

By the way, such high magnitudes can, apparently, occur only

in a small number of lakes which have been artificially trans-

formed into eutrophie environments by the efforts of man. In

this context, it seems proper to mention that it is precisely

the primary production quantity for plankton which serves as

the best index for the initial effect of fertilizing water

bodies. Therefore, suitable methods in this direction acquire

a particular importance for working out rational and effective

forms for fertilizing fish-farming ponds (Vinberg and Lyakh-

novich, 1965).

6 ( 5 3)

It is of a particular and not only theoretical but also

practical interest that many data point to the high primary

production and the high oxygen content in turbid and polluted

water bodies. lt has been demonstrated more than once that in e „ ,

many cases the photos,ynthetic aeration in turbid' waters surpas-. e= 1; , ses the one of the atmosphere. It is to be regretted that in-

sufficient attention is being paid to this fact in calculations

of the so-called 'cur e of OXygen bend! for turbi'd waters and

the self-purifying ability of water bodies. Unable to enlarge

on these questions, we want to remarkoonly that research into

primary production is not less important for sanitary hydro-

biology than for investigations with regard to fishery (Vin-

berg, 1964).

In oceanology, the interest in primary production ex-

panded considerably in the course of the past decade, following

the elaboration of the radiocarbon method. During the last couple

of years, research into primary production was frequently sup-

plemented by the spectrophotometric determination of chloro-

phyll content in the plankton.

In Denmark, the country of origin of the radiocarbon

method, systematic observations of primary production have

been conducted for several years (Steeman-Nielsen, 1964). At

least five main centers for these investigations exist in the

U.S.: two on the Pacific, two on the Atlantic coast, and one

on the Hawaiian Islands. In this connection, important work

3s being done in Canada on an expertly-equipped station at

7 ( 53)

Nanaimo (Antio and others, 1963) as well as at the University

of Vancouver (Gilmartin, 1964). Actively working centers for

the study of primary production exist in Australia and Japan;

other countries, too, (France, Norway, etc.) are enged in this

research.

In the USSR, the primary production of plankton in the

northern seas was occasionally investigated by the oxygen method

as early as in the I -thirties by P.P.Shirmov. The radiocarbon

method was utilized for the first time in 1957 by Y.I.Sorokin

and 0.1. Koblents-Mishke on the resealitch ship tVityazt in the

Pacific Ocean. Ever since, such work is regularly conducted on

the cruises of the 1 Vityaz , . These investigations were also

among the tasks of the tObt during her cruises into the Ant- (511)

arctic, and of the ISedov , in the equatorial Atlantic. Since

1960, the Biological Institute of Southern Seas in Sevastopol

(Z.Z.Finenko, T.M.Kondratteva) has joined in studies for pri-

mary production in the Black, Azov, and Red Seas, in the Medi-

terranean and in the Atlantic Ocean.

These last years, the Atlantic Research Institute of

Pelagic Fisheries and Oceanography (the AtlantNIRO) has begun

to apply the radiocarbon study method for primary production

in the equatorial part of the Atlantic Ocean (V.D.ChmyrI);

the Baltic NIRO (A.K.Yurkovskii) does the same in the Baltic

Sea and in the Atlantic Ocean, where lately co-workers of the

All-Union Research Institute of Pelagic Fisheries and Ocean-

ography (the VNIRO, V.V.Volkovinskii and V.N.Bessonov) and

8 ( 54)

members of the Pacific Section of the Oceanography Institute

of the Academy of Sciences (the IOAN, G.G.Starodubtsev) joined

them in similar research.

Primary production in the sea is also studied by other

methods, for instance, according to fluctuations in oxygen

content within 24 hours (M.V.Fedosov, V.N.Ivanenkov).

Activities in the study of primary production are not

only characterized by their great extent but also by their high

organizational level. Apart from the UNESCO working groups in

this direction, spoken of a few pages back, we want to enumerate

the following meetings; the conference on primary production

in Bergen (1957) convoked by the International Council for Sea

Research (Rapports et proc ès-verbaux, 1958); the Primary Pro-

duction Section of the I Oceanographic Congress in New York,

1959; the X Pacific Congress in Honolulu in 1960 (Report of

SCOR-UNESCO, 1961). The laboratory of Professor Doty at the

University of Hawaii conducts special international cruises

for working out and co-ordinating methods - 'of intercalibrat-

ion' - at which, in 1960 and 1961, Soviet scientists parti-

cipated (Koblents-Mishke, 1962 b). For similar purposes, the

Oceanology Institute of the Soviet Union conducted symposia

on the Black-Sea base of the Institute in 1959 and 1963. Re-

sulting from the 1959 symposium, the 'Methodical textbook... ,

was published (1960).

On the XIII, XIV, and XV Congresses of the Internatio-

nal Association of Limnologists, held in Helsinki, Vienna, and

9 (54)

Madison, a great deal of attention was devoted to primary

production.

In 1960, in Minsk, a conference on primary production

took place which assumed the character of an Ail-Union assembly

(see the coilection: 1 The primary production of seas and inland

waters,' 1961).

The scope of the work conducted in the seas brought it

about that as early as in 1962 over 120 cruises of ships of

various countries plowed the seas and conducted measurements

of primary production by the radiocarbon method. In the Pacific

Ocean alone, such measurements were taken at more than 3000

points up to 1962. These studies enabled Mr. 0.I.Koblents-

Mishke to delimitate three basic and two intermediate zones

differing as to the magnitude of primary production, i.e., as

to the effectiveness of utilizing the radiation energy which

reaches the sea surface. It was established that in the most (.55)

productive (eutrophic) waters, an average of 237 g C/m2 is

assimilated per annum, in less productive waters 91, and in

those wide stretches of the central ocean not more than 28

g C/m2 . The general distribution picture of the primary pro-

duction magnitudes revealed itself to be similar to the picture

of the biomass magnitude distribution of the phytoplankton.

However, the differences in the biomass of phytoplankton be-

tween the zones of differing productiveness are much more

clearly expressed than the differences in the average primary

production of these zones. Consequently, the biomass and the

10 (55)

species composition of the phytoplankton can be useful as

one of the biological indicators for waters of differing pro-

ductivity, but they cannot be utilized to express the quantita-

tive primary production and the effectiveness in radiation ener-

gy exploitation. This fact has once again disclosed that only

direct measurements of photosynthesis intensity of the phyto-

plankton can quantitatively characterize the primary production

and, at the present state of our knowledge, this method cannot

be replaced by data about the biomass of the phytoplankton or

by whatever other indices.

The general bulk of primary production in the Pacific

Ocean equals 10.109 T Chear. Extending this figure to the

whole area of the World Ocean, we get 22.10 9 T C, or 61 g C/m2

per annum. If, in accordance with the calculations of Steeman-

Nielsen, we assume that, due to respiration of the phytoplank-

ton distributed below the eutrophie layer, the pure (the effect-

ive) phytoplankton production is lower by 40%, then the primary

production would amount to 13.109 T C/year.

It is remarkable that this magnitude practically con-

curs with the estimation of primary production of the sea

given by Steeman-Nielsen and based on the first data ho obtained

during the expedition of the tGalatheal as early as in 1962.

A notion is prevalent that, on an average, the solar

energy affecting a unit of water surface is utilized with

greater effectiveness than the same solar energy by the ve-

getable cover on land. In accordance with this opinion, it is

11 (55)

sometimes asserted even in popular scientific literature that

in prospective production man can get a greater yield from 1 ha.

of the sea than from 1 ha. of land. As was already pointed out

(Vinberg, 1960), these views are based on greatly exaggerated

and erroneous evaluations of pelagic primary production which

have found a wide acceptance. The figures adduced above, which

indicate the average productiveness of the various pelagic

zones, correspond to the following magnitudes of the general

utilization of solar radiation energy: 0,33, 0,11, and 0,02%.

It is interesting that very similar figures have been obtained

for the main categories of ecosystems on land, namely: forest -

0,33, arable land - 0,25, meadows and pastures - 0,10, deserts

and ice - 0,01% (Duvigneaud, 1962). At the same time, forests (56)

occupy over 40% of the continents, while the eutrophie waters

of the ocean, lying mainly around the coasts, occupy a very

small part of the general ocean expanse. As a result, the uti-

lization of the general radiation energy - the energy distrib-

ution and the ocean expanses occupied by waters of differing

productivity taken into account - has been established as aver-

aging for the total surface of the sea near to 0,04%, while,

on the other hand, the analogous magnitude for the land ap-

proximates 0,1%, i.e., it is by 2,5 times higher.

No doubt, these data are but approximate evaluations

and subject to further adjustments, but there cannot either be

any doubt that the above conclusion about the lower average

effectiveness in energy utilization for regions of the globe

12 (56 )

covered with water will not be invalidated by future more ac-

curate determinations of the mentioned magnitudes.

Primary production is just the first step in the product-

ion process in bodies of water, and it is directly exploited by

man only on subsequent links or levels of energy utilization.

If we take the total volume of pelagic fish and other economi-

cally valuable objects landed by fisheries as being 50.106 T,

or 1,39 kg/ha. per annum, which will be achieved within just

a few years, and if we assume that the brutto caloricity of

1 g of wet weight equals 1 kcal, the general yield will be

50.10 12 kcal/year. Taking 1 g C as being equivalent to 9,36

kcal, we get 22.109

T C.9,36.106 kcal/T C=206.10 kcal. -

remind that the primary production amounting to 22•109T C was

established by the radiocarbon method and, therefore, it is

somewhat (e.g., by 10 - 20%) lower than the real gross primary

production. Hence we may assume that, at the actually achieved

level of fishing, it equals 0,02% of the gross primary pro-

duction of the sea.

lt is instructive to compare this magnitude, which has

kept growing so impetuously these last times, with the cor-

responding, much more stable data of the correlation between

primary production and fishing in lakes. One can assume that

the gross primary production of lakes exploited by fisheries

-oscillates between 500 - 3000 kcal/m2 per annum, and their

yield in fish and other products is 5 - 60 kg/ha. or 0,5 - 6

kcal/m2 a year, which represents 0,05% - 2% of primary production.

28 0,3

91 1,8

•.)

27 8,4 4

13 (56)

Adopting 2000 and 2 kcal/m2 per annum (20 kg/ha.) as typical

magnitudes, we get 0,1%. This quantity is quite unexpectedly

not very far removed from the 0,02% of the already achieved

level of exploitation of biological resources in the ocean

which, at present, provides 1% of the nourishment of man as to

caloricity (6 - 8% of albumen and about 25% of organic(animal)

albumen).

In order to mark out ways and means for a more effect-

ive utilization of the primary production in water bodies, it

is essential to know the norms and patterns of the production

process which determine in particular the share of the primary

production apt to be removed by man as the final product of a

given piece of water. We are only at the beginning of attacking

this grandiose task. For the time being, only some fragmnet- (57)

ary illustrations can be adduced, and one might indicate the

basic orientation for solving this problem. For instance, when

comparing the various zones of the Pacific Ocean as to their

primary production and the biomass of zooplankton (according

to data of V.G.Bogorov and M.E.Vinogradov), we get some inter-

esting results (see table 1).

Tablel

Productivity of various water types in different zones of the Pacific Ocean

np0AyKT1IBHOCTI, BOB. pa3nux TUBOB B oTpeablimx umax Tuxoro oKeana

Index of

productivity

Primary production g Cim2 yeiqz

nomass of mosoplankton g C/ra % of primary rroduction

Type of nH1 mv1 waters _ oligo- meso- eu- W01-X.* MC bIC rP xbimc CnipAulA, troph. troph. troph. _

14 (57)

As we see, an increase in the primary production is not

only attended with an increase in the biomass of zooplankton,

the ratio of this biomass to the primary production grows in

its turn. According to these data, the effectiveness in utiliz-

ing the primary production is also greater in eutrophie regions.

However, from a general point of view, it is unjusti-

fied"to compare production and biomass. This cannot give us more

than a starting point for our investigation but cannot help us

in establishing the norms and patterns of the production process.

In order to get nearer to the solution of this problem,

we ought to extend the functional approach, which turned out

to be so fruitful in the study of primary production, to the

subsequent production levels. This might appear unreal consid-

ering the seemingly boundless diversity of forms, functions,

and specific adaptations we are confronted with when examining

the animal world in bodies of water. Biologists. used to fix‘i

their attention first of all on well-defined bpecies differences,

are inclined to underestimate the fact that the entire immense

multiformity of adaptational devices is oriented towards en-

suring the requirements of a few basic functions of living

matter, which functions are subject to the general laws of life.

This is but natural in our contemporary production-

oriented biological ecology which, for its first province of

attention, has adopted the energetic research principle into

trophic relations and the trophic structure of ecological

systems.

( 57)

The foundation of this energetic principle is the ob-

vious and unquestionable assertion that the vital activity of

the total heterotrophic animal population in a body of water (58)

is performed at the expense of energy derived from primary pro-

duction, and that all production is inevitably attended with

a dissipation of energy parts from the preceding trophic level.

The rate of this utilization and dispersion of energy

is determined by the metabolism intensity; and the general vo-

lume of destruction - by the product total of metabolism inten-

sities in relation to the respective biomasses.

Consequently, as the first function to interest us, we

may take the metabolism intensity which can be expressed as

expenditure for metabolism, i.e., as a share of biomass expended

on energetic exchange (metabolism) within a time unit. For many

aquatic animals, e.g., for planktonic Crustacea, metabolism in-

tensity has been studiEd thoroughly enough. In other cases,

metabolism intensity and expenditure on metabolism, indicated

with a certain degree of accuracy, can be taken from already

ascertained correlations between metabolism intensity and in-

dividual size (Vinberg, 1962). Unfortunately, this so far

available information has not been generalized to a sufficient

degree and brought to the attention of hydrobiologists. The

second function, equally of a quite general character, is the

growth rate or the magnitude of increment. l(2\ •vv e wisn o 1hat the sum total of growth by in-

dividuals represents in itself the production of a population.

o 2 5

10 15 20 * 25' )),)

5 7

I()

'0

•:t )

1 3

Il 6

In* 21*

:u1

4 7

12* 17* 22* 27 * :t7

4 9

14 10 24 34

0

8 13

13*

16 (58)

By taking the sum total of accretion plus expenditure

on metabolism, we, obviously, obtain the quantity of assimilated

food, i.e., we find out how much nourishment must be assimilated

in order to ensure a certain production.

Ail this represents prefectly clear, elementary, and

trivial propositions; and it is just this fact which confers

on them their insuperable strength and significance. Many au-

thors, when examining the various factors of the biotic cycle

in a water body, are utilizing them. Thus, by way of an example,

Steeman-Nielsen (1962), discussing the possible correlation

between phyto- and zooplankton, presents the following table.

Table2

Growth of zooplankton per 24 hours at varying quanti-ties of assimilated food and varying expenditure on metabolism, acc. to Steeman-Nielsen (1962); (all magnitudes are in per cent of biomass per 24 hours)

Ta6mmua 2

CyToquidt npupocT aoonmanKTona npu pa3HOM KOMU4OCTOO ycnoenuori mum n paanmx TpaTax ila o6men, no CTumann-Hu.nhceuy (1962) (nce FICJWOM111,1

B uponeuTax OT 6HOMaCCM aa cyTKu)

Assimilated Yernemuls)

Bum food

5 6 7

Tpaul oGmen letabolism pendituro on 3 I •4 ' I I g 10 12 15

Y'' h 41 adde_d Quill. 24-hour

(1

IS

2 0 7 5 3

.,!t1 3 .2 33

1411M e ( (a111) ,M MV alle( Mv1K011,( A 1) 11 C 11011 H 1 Togel v)

Note : Figures marked with an asterisk are ex- plained in the text.

The table shows that accretion per 24 hours, expressed

17 ( 59)

In per cent of biomass, represents the difference between as-

similated food and expenditure on metabolism. This is correct,

but Steeman-Nielsen did not heed one fundamentally essential

circumstance. Expenditure on metabolism and increment are not

independent magnitudes. Added growthffias of necessity attended

with expenditures on metabolism 01 , and theycannot be less

than a certain minimal magnitude.

n and T are connected by the coefficient of the

utilization of assimilated food on accretion:

" K" r , or n --K T-- ri '

Even at embryonic growth K" is not higher than 0,7 -

- 0,8. Once this is established, it is easy to see that all

magnitudes in the left lower corner of table 2, which we marked

with an asterisk, are impossible and unreal.

This example goes to prove how indispensable it is to

take into account : the regularity of interdependence between

growth (production) and metabolism (destruction), which can be

denoted by the coefficient K" . Knowing K" and T we can

judge about the magnitude of production (n).. At present, a

rapid accumulation of information concerning the concrete mag-

nitudes of K" is going on. This is a special, a big, and as

yet unresolved question. Based on available data, we get the

impression that for natural populations K" , in many instances,

has a significant magnitude, for instance, 0,4. Extensive re-

search is still necessary, but it looks as though one can hope

18 (59)

that this problem might be elucidated within a relatively

short time.

The production of species populations might also be

calculated in another independent manner resorted to at present

by many hydrobiologists, namely: by computations of growth rate

and numerical strength of age classes or size groups of individ-

ual species populations.

A similar method, applied to zooplankton, was published

in 1961 by P.G.Petrovich, E.A.Shushkina, G.A.Pechent. It is

described in more deatil in a paper of Pechent and Shushkina

(1964). Production magnitudes computed by this method were

compared with magnitudes obtained according to metabolism in-

tensity, and they agreed fairly well.

On table 3, some data taken from the work of Vinberg,

Shushkina and Pechent (1965) are summarized.

Table 3

Primary production of plankton plus macrophytes and production of herbivorous zooplaneon in the lakes

of Naroch (in kcal/m' per annum) TaGmnua 3

liepsuquan npoRyKujin 11.113.11KTOIla n magpocpwroB It nponming paCTIITeMblIOMI,11010

30011.3allICT011a H 11a90 , 1illiCKIIX o3epax ICIC(7, At 2 an ro

Haroch ,Myastre- .30torin-Index Hapoq bwmp.

1.1.1(21aaTeMb 1111 .1,1)4WW ,1111 1 , »13PW

1)rimary nroduçtion (Zooprankton (predators excl.)

biomass production P B 5 of primary product.

pesotroph.etitroph 1 967 1 157

2,95 96,5 9,0 9 ,75

eutroph. 1800

6,0 160.9 26,8 9,0

67,8 11.3 4,4

In this case, too, the primary production in eutrophie

19 (59)

waters is not only greater, but it is utilized with a higher

degree of effectiveness by the zooplankton. We remark that it

would be premature to ascribe to this fact the validity of

a general law.

Although the adduced e.eamples are a result of exten- (60)

sive, painstaking and labour-consuming efforts, they are only

meant to demonstrate the peculiarities of that new stage in

developing the theory of biological productiveness which ori-

ginated in the works about primary production. The extension

of the same principles to the totality of the productive pro-

cess is dictated by nessessity; namely, the relentless logic

in the progress of science; and also by the tasks which pract-

ical requirements of the sanitary-technical and the fisheries-

economical exploitation of water bodies place before our

theories.

The only way to apprehend the norms and regularities

determining the final production needed by man is by knowing

how to express the separate stages of the production process -

the intermediate production on ail of its levels - in strict-

1y defined magnitudes. And only by knowing how to designate by

well-defined magnitudes the functional significance of a popu-

lation, can we understand its part in the process of the self-

purification of water bodies.

The international biological program for biciegi-cally

produetive limnology investigations,Was drawn up in 1963 and

1964 by a highly representative international committee; it

20 (60)

consisted of such authoritative limnologists as Roessler (USA),

Worthington (England), Siole (Federated German Republic), Rhode

(Sweden), Tonolli (Italy), and others. The program is based on

the consistent application of the energetic principle, and it

is in full harmony with the ideas developed here.

This, precisely, is the main orientation in the advance

of hydrobiology, and many limnologists and oceanologists follow

these lines in their work, which we couid corroborate with

many examples. It is particularly important and meaningful that

the priority in working out the basic propositions in this

direction belongs to Soviet scientists.

Research into primary production has come to occupy

such a prominent place in contemporary hydrobiology not only

because it possesses in itself and in many respects both a

theoretical and practical interest, but in particular for the

reason that it opens the way to go on from speculative con-

structions to a concrete study of all stages of the production

process, to pass from reasonings about the one or the other

aquatic organism and its significance to concretely express

its functional part in a water body by clearly determined

magnitudes.

By continuing along these lines, hydrobiologists will

be successful in the very near future in comprehending the

norms and patterns of biological productiveness, and in

g u

21 ( 61)

utilizing this knowledge for a more rational exploitation

of the biological resources contained in water bodies.

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