Environmental biology of fishes : emerging science

Henry A. RegierInstitute for Environmental Studies, University of Toronto, Canada

Theoretical and practical roles appear to be emerg-ing rapidly for the special kind of science to befostered by this journal . The editor is committed toadvocating `where possible wise use and mainten-ance of the global fish resources' and has chosento focus on `morphological, physiological andbehavioral aspects of development which areoriented ecologically ; ecology of marine andfreshwater taxocenes, populations or stocks andthe associated systems modelling ; evolution andsuccession in natural and cultural environ-ments . . .". It happens that I agree entirely that aclear emphasis on this kind of science is nowneeded. In what follows I have sought to justifythis belief using some quite general concepts andarguments .

The rationale has four basic components : threeare in the nature of temporal developmental se-quences and one is a conceptual framework in twodimensions . Current trends taken separately, oras sets to be interpreted within the framework, allimply a growing need for rapid progress in thestudy of the `environmental biology of fish' .

Because the concepts elaborated below are quitebroad and still imprecise, it seems inevitable thatsemantic problems will arise . Consider, for ex-ample, the word `science' . Though it is often desir-able that the word `science' be assigned narrow,carefully defined connotations, I have chosen touse it here in a broad sense . If the argumentadvanced in this paper were analyzed further itmight well be desirable to split `science' into sub-systems of complementary components . Such arefinement might well add meaning to the presentanalysis but I feel that it would not alter the basicconclusions . I trust this would also be the case with

* Invited editorial

Received 26.2 .1976 Accepted 30 .3 .1976

Env, Biol . Fish ., Vol . 1, No . 1, pp . 5-11, 1976

other words here used with broad and somewhatvague connotations .

Expanding role of biology : fishery resources

In this section and the following I have sketchedtwo pairs of interactive sequences . In each pairone sequence is a listing of somewhat idealizedstages in the `maturation' of a resource user'sinvolvement with, or impact on, a fishery resourceand the aquatic environment. The other sequencein the pair, in phase with the first in a special case,lists the conceptual foci for the kinds of sciencethat are particularly useful to resource or environ-mental management at different stages of 'ma-turity' . These pairs are sketched here not as normsof a perfectly co-ordinated sequence that shouldbe followed in an ideal world, but instead are pro-posed as broad inferences of what has happened invarious geographic regions of the real world duringthe past century .

Table 1 illustrates the case for a rather largeand well-behaved fishery and fish resource . Con-sider a fish resource system as in the shelf seas orsome of the world's great lakes, in which theharvestable surplus of ecological production is nothighly responsive to the stress of light to moderatefishing intensity . Abundant stocks of many of thelarger species can accommodate considerablefishery exploitation without the risk of collapse,though eventually any fish stock will collapse orfluctuate violently given excessive exploitation .These presuppositions in effect imply that thefishes' natural environment is neither highly 'un-stable' (in the sense of large-scale, unpredictablefluctuations), nor highly `stable' (in the sense thatenvironmental fluctuations almost never exceed asmall scale.) In turn the human fishery is 'well-behaved' in that the expansion and intensification

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Table 1 . Idealized sequence of states in fishery development or''maturation" processes, with identification of the kind ofbiological science most directly relevant to each stage

State of fishery

Relevant, i-nnovative biologicalresource use

information

I . Exploration, trial

Linnean taxonomy, geographic and seasonalfishing, adaptation distribution of larger individuals ofof gear

profitable stocks

2 . Initiate and expand (a) Identify life history phases of MPSs,systematic exploit- and their spatial and temporal distribu-ation of most

tions, especially the critical reproduc-profitable stocks

tive phases(MPSs)

(b) First rough assessment of potentialyield of MPSs

3 . Itensify fishing on (a) Population dynamics of MPSs to yieldMPSs to full utili- separate estimates of optimum or maximumzation ; expand to

sustainable yieldless profitable

(b) Identify and clarify strong ecologicalstocks (LPSs)

interactions among species (MPSs and LPSs)in order - to adjust separate yield es-timates on an ad hoc basis

4 . Full and intense

Dynamics of the fish taxocene and/orexploitation of all ecosystem to investigate overall profita-marketable stocks

bility of yield of alternative managementoptions

5 . Chronic overfishing, From whole-system theory based in part oneventually recog-

study of replicate systems, estimate thenized as such

likelihood of major irreversible conse-quences, .such as species extinction

6 . Resource

"Ecological engineering" to correctrehabilitation

major ecological deficiencies and deform-ities and/or "therapeutic ecology" to aidnatural recovery processes

of the harvest process occurs gradually over anumber of decades . The fishery does not redirectits preferences nor alter its basic methods rapidlyand haphazardly in time and space . These idealizedcharacteristics are not very different from theecological circumstances and exploitive history ofthe temperate shelf seas and temperate large lakesof the world prior to about 1950 .

The term `most profitable stocks,' used inTable 1, should be explained . Simply put, it is themaximum difference between what the fishingenterprise receives, for its landed catch of partic-ular stocks, less the corresponding costs of fishingfor them. Thus fishing for less preferred abundantstocks with inexpensive vessels and gear nearshore may be as profitable as fishing for other morehighly preferred stocks with large vessels and intri-cate costly gear far off shore . Use of the concept,most profitable stocks' thus permits a broad gene-ralization of the idealized sequence over bothinshore and offshore stocks whether or not theyhistorically progressed through the six stages ofTable 1 temporally in phase. Where an approx-imate temporal correspondence of this kind maybe demonstrated in the historical record, as withsome shelf-seas fisheries, the present analysisshould appear to be somewhat more realistic .

The Stages 1 to 6 in the column to the right of6

Table 1 are not to be considered discrete in thesense that activities listed in each begin and endin that particular stage . Rather the stages identifywhere in the sequence a particular activity is per-ceived as being most creative, innovative andpotentially useful . As the concepts and methodsbecome well-understood they are transferred intopractice and are thereafter accepted as part of theshared common background of the scientific pro-fessionals .

Table 1 focuses on relevant biological informa-tion and makes no mention of other kinds of in-formation (economic, technological, social) alsouseful to fisheries managers at the various stages .The relevant kinds of limnological and ocean-ographic information could easily be sketched byfishery ecologists. Corresponding sequences canalso be sketched for economics, fishery technology,and policy sciences that deal with regulatory pro-cesses .

The kind of `environmental biology' to be fos-tered by this journal tends to relate to the lower halfof Table 1, - to the more `mature' stages of theresource use sequence . Most of the large fisheriesof the world are now past Stage 3 . Unfortunatelymany fishery biologists are lagging at least onestage behind that which would be most directlyrelevant according to Table 1 . Very few are onestage ahead - where innovative science shouldbe - but more of this later .

Expanding role of biology : aquaticenvironments

Again consider a somewhat idealized case in-volving the effect of a growing human settlement,with surrounding geographic `resource shed', on alake or estuary nearby . As before, the aquatic eco-system is taken to be fairly unresponsive and canwithstand moderate stress without major disrup-tion. Assume that no fishing is permitted and noneoccurs. Instead the aquatic ecosystem is subjectedto the usual insults of sewage loading, shorelinerestructuring, bottom dredging, loading by atmos-pheric particulates from an industrialized society,heat loading, and biological loading with exoticspecies . A sequence of stages roughly comparableto those of Table 1 is shown in Table 2 .

A comparison of Tables I and 2 shows that Ihave defined the use periods in both cases so thatthe biological information periods within the two

Table 2 . Idealized sequence of states in the use of the aquaticenvironment as a source for abiotic resources, as a sink forwastes, etc ., with identification of the kind of science partic -ularly relevant to each state

State of aquaticecosystem abuse

1 . Build harbours andcanals, drainwastebearingsurface waters,withdraw water forlocal distribution

2 . Build dams andlevees, dredgeharbours, constructpiped sewage andwater systems,wide-spreadswimming andboating

Restructure shore-lines, drainmarshes, treatsewage and drinkingwater chemically,construct and tendbeaches and yachtbasins, loading byatmospheric partic-ulate, that fallinto the habitat

4 . Widespread andintense multipleuse, with someeffort made toseparate usersspatially and/ortemporally

Chronic overusewith majorundesirable inter-actions betweenvarious users'ecological impacts

6 . Habitatrehabilitation

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5 .

Relevant, innovative biologicalinformation

Linnean taxonomy to identify which of themost profitable fish stocks (MPSs), ifany, are seriously affected by determin-ing whether their spatial niches encom-pass the small areas affected

(a) Identify life history phases of MPSswith their spatial and temporal distribu-tions, particularly of the criticalreproductive phases, relative to heavilystressed habitats(b) First rough assessment of overallimpact on several of the MPSs

(a) Approximate assessment of the impactsof separate anthropogenic stresses on theMPSs(b) Joint assessment of the two or threemost destructive stresses to determinethe general nature of any ecologicalinteractions among them, to judge whetherecologically-oriented regulation isneeded

More precise assessment of separate andjoint effects of stresses to considervarious alternative use and managementoptions, which requires a fairly compre-hensive understanding of ecosystemstructures and processes

On the basis of whole-system theory,predict the demise of one valued eco-system component after another

Ecological engineering to correct defi-ciencies in ecological structures and/ortherapeutic ecology to aid naturalrecovery processes

sequences resemble each other . If it happens thatfishery use and environmental abuse proceed his-torically in phase through Periods 1 to 6, then thekind of biological (and other) science most rele-vant at each period is conceptually similar for boththe use and abuse cases . Where they proceed outof phase it might at first glance appear that thescience relevant to the particular stress processwhich is lagging can afford also to lag behind theother. But this overlooks the fact that a species'habitat and functional niches are closely inter-related and both are influenced by each of themajor types of anthropogenic stress . Hence which-ever stress process leads in the `maturationsequence' determines the type of biological per-spective most effective for gaining an under-standing of the ecological response to either orboth stress types .

Putting scientific insight to work

In Tables 1 and 2 I have sketched a `maturationsequence' of idealized fishery use and environ-mental abuse processes . A cynical or pessimisticobserver might have terminated the sequence atStage 5. A rejuvenescence is possible in some cases,as with the oligotrophication recovery process thatfollows when anthropogenic eutrophication pro-cesses are controlled .

Four kinds of ecological paradigms may be dis-cerned in the right hand columns of Tables 1 and 2 .These are :

1 . Linnean taxonomy, including natural history ;2. population ecology, including population

dynamics ;3. ecological systems, including taxocenes and

ecosystems ; and4. ecological engineering and therapeutic eco-

logy .These four paradigms are now at different phasesof practical development as may be illustrated withthe aid of Table 3 .

Four steps may be described by which a para-digm becomes progressively incorporated intothought and action . A, creative innovation ; B, edu-cational transfer; C, practical application ; andD, institutionalization . The activities particularlycharacteristic of each phase may be identified bythe forms that the then relevant publications havetaken. The characteristic media types are shownfor each phase in Table 3 . Turning the processaround, the existence and state of disseminationof various kinds of scientific media, at some pointin time, permit a diagnosis of the state of maturityof a particular paradigm at that moment.

As with previous tables, activities in Phase Ado not cease when activities in Phase B get wellTable 3 . Sketch of steps in the practical assimilation of amajor scientific perspective (paradigm), show inq the t ,

cf publications characteristic of different ohasrr .

PHASE A :

CREATIVE INNOVATIONJournal papersBihliographinsSymposiaMonographs

iPHASE B :

EDUCATION, TRAINING- Anthologies and reviews

TextbooksMethods manualsSample mapsPrototype data banks

PHASE L .

INSTITUTIONALIZATION

-Parad1em's handhookAnnual colloquiaSpecialized _journalsConstitution for aprofessional aSSociatInn

PHASE C :

PRACTICAL APPLICATION- Case histories

Requlatorv protocols andstandardsPractical manualsData banks

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underway. It is the locus of major innovativeactivity that progresses down the sequence, butbuilds on what has already been accomplished .

The last Phase (D) is of some particular interest .In order that the workers within a particular para-digm complete Phase D, they must become wellaware of the broader scientific and social signi-ficance of that paradigm . With this step they com-plete the self-regulatory loop (left-pointing arrow)and henceforth proceed to develop all four phasesinteractively and in consort. Also they create theinstitutional frameworks within which workerswithin that paradigm interact (right-pointing ar-row) with other workers operating from other per-spectives. As always with institutionalization,Phase D is both liberating and enslaving, but thisaspect is not of immediate interest in this paper .Again Table 3 is not proposed as a normative

framework of what should happen, but rather as asketch of what usually has happened with variousscientific paradigms (e.g. Newtonian, Linnean,Darwinian, Malthusian, etc .)

Let us consider the four ecological approacheslisted above, as applicable to large well-behavedaquatic resources and environmental systems, inthe context of Table 3 . Linnean taxonomy is fullymature; population ecology is moving into PhaseD; systems ecology is just into Phase C and begin-ning to explore Phase D ; ecological engineering andtherapeutic ecology are in Phase A with some pre-liminary trials in Phase B .

Current status of useful paradigmsa special case

If the contents of the preceding sections have somerealism and relevance for the kinds of aquatic sys-tems specified, then we may in turn use thesemodels to judge the state of the science as currentlyapplied to individual resource use - environ-mental abuse combinations within that aquaticsystems set .

Numerous large lakes, estuaries and marine bayswithin the industrialized temperate regions arenow stressed to the level of Phase 4 (Tables 1 and2). Some are well into Phase 5 and a few into Phase6, see below. The fully relevant science in thesecases would be that which focuses on ecologicalsystems behavior (taxocenes and ecosystems .) Butthis paradigm's development has only progressedabout half way through its own maturation se-quence. This appears to be a noteworthy lag .

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International attention is now being directedtoward rehabilitation of some grossly abused aqua-tic systems such as Lakes Erie and Ontario andthe Rhine River. But the science of ecologicalengineering and therapeutic ecology is only in itsinfancy .

Perhaps it is fair to say that biological scienceseems to be lagging with respect to those systems inwhich fisheries and other anthropogenic stresseshave intensified rapidly in recent decades . Somelag, though frustrating to practical people, isunderstandable.

Less easily understood is the widespread lack ofinterest, on the part of many environmental bio-logists, to transfer insights from study of one ormore aquatic ecosystems to others . If use-patternof a system appears to be moving from Phases 3 to4 (Tables 1 and 2), it should make sense to examinewhat others have found who are currently intim-ately involved with a system already in Phase 4 .What is usually needed, somewhat unfortunately,is not just some more of the same kind of sciencethat was adequate to Phase 3 : a shift to Phase 4calls for some degree of re-education into a para-digm that transcends what was adequate at Phase 3,i .e. from population dynamics to ecosystem dyna-mics. This may involve mental deprogramming aswell as reprogramming for those environmentalbiologists who have shown no interest in the over-all dynamic processes of the science .

More challenging yet : as the process sketchedin Tables 1 and 2 progresses to Phases 4, 5 and 6,the workers of other disciplines (economics,management sciences, oceanography, etc .) arebrought into direct discussion with environmentalbiologists . Sooner or later it becomes worthwhileto search for paradigms that are congruent withinthe various relevant disciplines, such that allworkers may share a single transcendent paradigm.

A transdisciplinary approach to environmentalscience and other 'metaproblems' is making rapidprogress. It may not be developing in the linearsequential mode sketched above for schools ofenvironmental biology . Groups here and there arestriving to structure the four major components ofFigure 1 more or less contemporaneously . Interest-ing proposals and trials may be identified for eachof the four `steps' . With the increased pace ofscientific evolution it may be that the whole setmay reach operational effectiveness much morequickly than has been the case with earlier initia-tives within the conventional disciplines . This more

comprehensive approach may in turn become a`discipline', but that is not a foregone conclusion .This topic goes beyond the immediate concern ofthis journal, but it should be of interest that thekind of environmental biology here encouragedmay well be 'pre-adapted' for application in thetransdisciplinary contexts and processes nowemerging .

A broader perspective on the real world

So far, the discussion has dealt with rather idealizedexpository models of sequences related to a specialtype of resource-environment system . We now turnto examine what other types might be identified .The opposite to large and well-behaved would besmall and poorly-behaved . Two other possibilitiesexist: small and well-behaved; large and poorly-behaved .The connotations of the word `behave' have

been expanded somewhat beyond what is permis-sible and some clarification is in order .

The desirable qualities or resources of an eco-system may fluctuate due to poor self-regulatorycapability, or because large natural external forcesact irregularly, or because anthropogenic stressesact to destabilize it . The term `variability' will beused to denote the temporal irregularities in aresource or quality. Both the amplitude and fre-quency of oscillations contribute to the measure ofvariability. Beyond this the term is not definedexplicitly ; i.e. so far as I know there is not now anexplicit mathematical expression for the conceptthat I have in mind. `Variability' seeks to encom-pass much of what contributes to a decision-maker's uncertainty as to precisely what actionshould be taken, even if a reasonably completeunderstanding of the ecosystem has already beenachieved .

The concept of large vs . small may be definedas the `expectation' or long-term mean, say ofmaximum sustainable yield of a resource, or of aparticular index of quality such as phytoplanktonproduction as a measure of eutrophy-oligotrophy .

Arranging these two concepts as a 2 X 2 matrixwe obtain Figure 1. A rather involved discussioncan be addressed to the various edges and spaces ofthe matrix, but would be out of place here .

Our idealized special case in previous sectionsfalls in Cell I of the matrix . Because individualresources or system qualities in Cell I are of great

Fig. 1 . Four major expectation and uncertainty classes forwhich decision-making rules are quite different with conse-quent differences among the kinds of science that are directlyrelevant to decision-making .

practical importance, it is almost inevitable thatthe sequence of practical use and relevant biologyshown in Table 1 and 2 will be traversed at least toPhase 4 . Phases 5 and 6 are guesses or extrapola-tions based on what may be discerned to havealready happened in systems that fall in Cells IIand III .

Turning to Cell II, each individual resource orquality is too small to justify individual attention .There are `economies of scale' in ecologicalscience, hence a research approach that is eco-nomically practical for a large resource (e .g. stock-by-stock population dynamics) is excessivelycostly relative to benefits with a small resource .For these and other reasons workers in smallresource and environmental systems have tendedto skip through steps 1 through 3 (Tables 1 and 2)rather rapidly, and settle down to relevant work insteps 4 and 6 . Some important scientific beginningsto Phase 6 may be found scattered throughout thenorth-temperate zone . With respect to very smallsystems, such as Chinese aquacultural ponds, awell-developed methodology has existed forcenturies, though it has apparently never been welldescribed scientifically .

Cell III relates to flood plains, shallow lakes ofhigh latitudes, and some waters intensely used orabused by man. Whether the cause of the highuncertainty is natural or anthropogenic, the mostrelevant biology appears to be that of Phase 5 or 6(Tables 1 and 2) . To address the issues using onlythe biology most relevant to Phases 1 to 4 (as theyapplied in Cell I), would involve relatively largeresearch costs .

Cell IV systems must be dealt with in the first

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Low

expectation

High

expectation

High

uncertainty III LV

I"",

uncertainty II

instance by policy and management sciences ;biological and other sciences would be used toprovide some predictive data for consideration inthe context of the policy and management sciences .An expanded Phase 5 (Tables 1 and 2) seems mostrelevant; Phase 6 is largely ruled out where theforces causing the uncertainty are beyond localcontrol .

From the perspective of this section I judge thatthe kind of science to be addressed in this journalis already sorely needed in a broad spectrum ofreal-life situations .

New decision-making processes call for`Environmental Biology'

The United Nations Conference on the HumanEnvironment (Stockholm, 1972) developed anaction plan with many recommendations. Formany and perhaps all countries of the world theimplementation of these plans implied some majorchanges in natural and local decision-making pro-cedures or protocols . Henceforth the naturalenvironment and renewable resources shouldreceive much more careful attention . To achievethis meant that decision-making procedures had tobe altered in major ways . Many of the techniquesthat had to be brought together into an effectiveoverall process had already been introduced on alocal or trial basis, separately, here and there .It became apparent that a simple assembly of all ofthese did not add up to what was needed . Morecomprehensive and far ranging policies andlegislation had to be introduced . This process,begun some years before 1972 in some countries,has not yet been completed. Nevertheless some ofthe more important procedural programs now inplace may well remain operative for many years .

In my own country, Canada, a number of pro-cedures related to the aquatic environment haverecently been introduced and expanded fromearlier forms. These include more comprehensiveenvironmental monitoring, regional planning, en-vironmental assessment and review, resource andenvironmental rehabilitation, as well as strategicand tactical planning . The planners and managersrequire information well-suited to their needs .Heretofore biologists and other scientists of theolder disciplines have had great difficulty com-prehending the needs of these new kinds of pro-fessionals. The expository frameworks of the

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previous sections may find application for thesepurposes .

It is highly unlikely that all the concepts of thispaper will find acceptance . Nevertheless it maywell be a foregone conclusion that a large needwill be explicitly and widely recognized for 'en-vironmental biology' as we define it in this journal .

Our opportunity

Many ecologists are pessimistic about the futurehealth of the natural environment-the biosphere-as direct and indirect human demands on it expand .Especially in industrialized countries, manyidentify growth of the world's human populationas the primary problem. As of 1976, this is a seriousmisdirection of emphasis by the people of indus-trialized countries . The priority challenge forus is to learn to live in a manner that is much lesswasteful of resources and less damaging to theenvironment . From what I understand of thedeeply-held value systems of many underdevelopedcountries, I now fear the overall effect on the bio-sphere of their growing populations less than I dothe large and growing impact of our own wastefulways. Of course each is a big problem in its ownregional context, - neither outranks the other inany ultimate sense .

I have had the opportunity to participate as adelegate to the Second World Food Congress ofthe Food and Agriculture Organization (TheHague, 1970), in various activities related to theUN Conference on the Human Environment(Stockholm, 1972), and as a national advisor atthe UN World Population Conference (Bucharest,1974), as well as in numerous international scientificconferences and symposia. From these experiences,I conclude that a widespread and fairly generalconsensus already exists in these matters, thoughviews diverge sharply on details of science andpolicy .

If there is a basis for real optimism, and I believethere is, then we should work enthusiasticallytowards a better day . It may be that many eco-logical matters will get worse before they getbetter, but they must get better eventually .

Environmental biologists tend to share a valuesystem that seeks an accommodation between manand nature. On these matters humanity simply can-not risk a short-sighted amoral `objectivity' accord-ing to which a scientist can justify any `experiment'

of any scope. The stakes are too high . If we err wemust err on the side of a conservatism of theshared values of `man in nature .'

To be an ecological conserver is not inconsist-ent with Mahatma Ghandi's statement `The earthhas enough for every man's need but not forevery man's greed,' and that of Dom HelderCamara `We must lift ourselves out of the sub-human situation of misery without falling intothe in-human situation of super-luxury .'

This is our larger opportunity . If the analysessketched in this paper make sense, then energeticresearch and practical action by `environmental

biologists' may be of some lasting benefit withrespect to this larger opportunity .

Acknowledgements

Among the many friends who have contributed tothe development of this paper are included E . K .Balon, P. J. Colby, A . S. W. deFreitas, G . R .Francis, J . A. Gulland, H . F. Henderson, S . R .Kerr, K. H. Loftus, J . J. Magnuson, F. D. Mc-Cracken, J. L. McHugh, J. M. Neuhold and R. A .Ryder. But none of them should be held respon-sible for the inadequacies of the present work .

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