The Trophic-Dynamic Aspect of Ecology Author(s): Raymond L. Lindeman Reviewed work(s): Source: Ecology, Vol. 23, No. 4 (Oct., 1942), pp. 399-417 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1930126 . Accessed: 30/01/2012 10:50 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. http://www.jstor.org THE TROPHIC-DYNAMIC ASPECT OF ECOLOGY RAYMOND L. LINDEMAN OsbornZoological Laboratory,Yale University Recent progressin the studyof aquatic community. A more "bio-ecological" food-cycle relationships invites a re- species-distributionalapproach would appraisal of certain ecological tenets. recognize both the plants and animals Quantitative productivitydata provide as co-constituentsof restricted"biotic" a basis for enunciating certain trophic communities,such as "plankton com- principles, which, when applied to a munities," "benthic communities,"etc., series of successional stages, shed new in which membersof the living commu- light on the dynamics of ecological nity "co-act" with each other and "re- succession. act" with the non-livingenvironment (Clementsand Shelford,'39; Carpenter, "COMMUNITY" CONCEPTS '39, '40; T. Park, '41). Coactions and are consideredby bio-ecologists A chronologicalreview of the major reactions effectorsof succession. viewpointsguiding synecological thought to be the dynamic The trophic-dynamicviewpoint, as indicates the followingstages: (1) the paper, emphasizes the static species-distributionalviewpoint; adopted in this (2) the dynamic species-distributional relationshipof trophicor "energy-avail- withinthe community- viewpoint,with emphasis on successional ing" relationships phenomena;and (3) the trophic-dynamic unit to the processof succession. From allied viewpoint. From either species-distri- this viewpoint,which is closely to approach butional viewpoint,a lake, forexample, Vernadsky's"biogeochemical" might be considered by a botanist as (cf. Hutchinson and Wollack, '40) and containingseveral distinct plant aggre- to the "oekologischeSicht" ofFriederichs a primfiary gations, such as marginal emergent, ('30), a lake is consideredas floating-leafed,submerged, benthic, or ecological unit in its own right,since all phytoplankton communities, some of the lesser "communities" mentioned which might even be considered as above are dependent upon other com- food "climax" (cf. Tutin, '41). The associ- ponentsof the lacustrine cycle (cf. ated animals would be "biotic factors" figure1) fortheir very existence. Upon of the plant environment,tending to furtherconsideration of the trophic cycle, limit or modifythe developmentof the the discriminationbetween living organ- community" aquatic plant communities. To a strict isms as parts of the "biotic nutri- zoologist,on the otherhand, a lake would and dead organismsand inorganic "environment" seem to contain animal communities tives as parts of the The roughlycoincident with the plant com- seems arbitrary and unnatural. munities,although the "associated vege- difficultyof drawing clear-cut lines be- non- tation" would be consideredmerely as a tweenthe livingcommunity and the the part of the environment'of the animal living environmentis illustratedby difficultyof determiningthe status of a ' The term habitatis used by certain ecologists slowly dying pondweed covered with (Clements and Shelford, '39; Haskell, '40; T. periphytes,some of which are also con- Park, '41) as a synonym for environmentin the usual sense and as here used, although Park tinuallydying. As indicatedin figure1, points out that most biologists understand " hab- much of the non-livingnascent ooze is itat" to mean "simply the place or niche that rapidly reincorporated through "dis- an animal or plant occupies in nature" in a species-distributionalsense. On the other hand, be hoped that ecologists will shortly be able to Haskell, and apparently also Park, use " environ- reach some sort of agreement on the meanings ment" as synonymouswith the cosmos. It is to of these basic terms. 399 400 RAYMOND L. LINDEMAN Ecology,Vol. 23,No. 4 solved nutrients" back into the living magnitude, i.e., the biotic community "biotic community." This constantor- plus its abiotic environment. The con- ganic-inorganiccycle of nutritive sub- cept of the ecosystemis believed by the stance is so completelyintegrated that writerto be of fundamentalimportance to consider even such a unit as a lake in interpretingthe data of dynamic primarilyas a biotic communityappears ecology. to forcea "biological" emphasis upon a more basic functionalorganization. TROPHIc DYNAMICS This concept was perhaps first ex- pressedby Thienemann('18), as a result Qualitativefood-cycle relationships of his extensive limnologicalstudies on the lakes of NorthGermany. Allee ('34) Althoughcertain aspects of food rela- expresseda similar view, stating: "The tions have been known for centuries, picturethan finallyemerges . is of a many processes within ecosystems are sort of superorganismicunity not alone stillvery incompletely understood. The between the plants and animals to form basic process in trophicdynamics is the biotic communities,but also betweenthe transfer of energy from one part of biota and the environment." Such a the ecosystemto another. All function, conceptis inherentin the termecosystem, and indeed all life,within an ecosystem proposedby Tansley ('35) forthe funda- depends upon the utilizationof an exter- mental ecological unit.2 Rejecting the nal source of energy, solar radiation. terms "complex organism" and "biotic A portionof this incidentenergy is trans- community,"Tansley writes, "But the formedby the processof photosynthesis more fundamentalconception is, as it into the structureof living organisms. seems to me, the whole system(in the In the language of communityeconomics sense of physics),including not only the introducedby Thienemann ('26), auto- organism-complex,but also the whole trophic plants are producerorganisms, complexof physicalfactors forming what employingthe energyobtained by photo- we call the environmentof the biome. synthesisto synthesizecomplex organic . .It is the systemsso formedwhich, substances from simple inorganic sub- fromthe point of view of the ecologist, stances. Although plants again release are the basic units of nature on the face a portionof this potentialenergy in cata- of the earth.. These ecosystems,as bolic processes,a greatsurplus of organic we may call them,are of the most vari- substanceis accumulated. Animals and ous kinds and sizes. They form one heterotrophicplants, as consumerorgan- category of the multitudinousphysical isms, feed upon this surplus of potential systems of the universe, which range energy,oxidizing a considerableportion from the universe as a whole down to of the consumed substance to release the atom." Tansley goes on to discuss kineticenergy for metabolism, but trans- the ecosystem as a category of rank formingthe remainderinto the complex equal to the "biome" (Clements, '16), chemicalsubstances of theirown bodies. but points out that the term can also Following death, every organism is a be used in a generalsense, as is the word potential source of energy for saproph- "community." The ecosystemmay be agous organisms (feeding directly on formallydefined as the systemcomposed dead tissues), which again may act as of physical-chemical-biologicalprocesses energysources for successive categories active within a space-time unit of any of consumers. Heterotrophic bacteria im- 2 " and fungi, representingthe most The ecological system composed of the bio- portant saprophagous consumption of coenosis + biotop " has been termed the holocoen by Friederichs ('30) and the biosystemby Thiene- energy,may be convenientlydifferenti- mann ('39). ated fromanimal consumersas special- October,1942 TROPHIC-DYNAMIC ASPECT OF ECOLOGY 401 SolarRadiation SolarRadiation A External DissolvedNutrients N j Internal A, PhytoplanKters / Pandweids Bact eria > -* 2 tZooplantkters~z OOZ Brawsers--4- -N - \/'l X A3 OmPmdoos Benic PmaWk A3 Betd A3GgQ1 Geerlaized PredatorsnS~~~~~~wimfniiod-yl Predatorsredainhp aftors A3dmanA AIG. na4 ized decomposers3of organic substance. solved nutrientsonce more in resynthe- Waksman ('41) has suggested that cer- sizing complex organic substance, thus tain of these bacteria be furtherdiffer- completingthe food cycle. entiated as transformersof organic and The careful study of food cycles re- inorganic compounds. The combined veals an intricate pattern of trophic action of animal consumersand bacterial predilectionsand substitutionsunderlain decomposerstends to dissipate the po- by certainbasic dependencies;food-cycle tential energy of organic substances, diagrams, such as figure1, attempt to again transformingthem to the inorganic portray these underlyingrelationships. state. From this inorganic state the In general,predators are less specialized autotrophicplants may utilize the dis- in food habits than are theirprey. The 3 Thienemann ('26) proposed the term reducers ecological importanceof