Raymond Lindeman and The

Raymond Lindeman and the between late lake succession and early logical efficiency of energy transfer over Trophic-Dynamic Concept in terrestrial succession. For 5 years he and his this expanded time scale. It is here that wife extensively sampled the population of the analogy between the development of Ecology aquatic plants and phytoplankton, the grazing an organism to maturity and community Robert Edward Cook and predatory zooplankton, the benthic fauna changes during succession finds its full- of worms and insect larvae, the crustaceans, est expression; and it is the elaboration Summary. Lindeman's classic paper on and the fish; and through this they gained a of this metaphor which has provided energy flow in ecosystems was initially very intimate understanding of the movement continuing inspiration to community rejected for publication in Ecology. of nutrients from one trophic level to another. ecologists (4a). Thus, in his effort to in- Reviewers felt there were insufficient data to To integrate this knowledge of food-cycle tegrate ecological patterns of differing support the theoretical model and that dynamics with current principles of temporal scales, Lindeman reduced the theoretical essays were inappropriate for community succession, Lindeman created the trophic relations of a community to a Ecology. The paper was subsequently trophic-dynamic viewpoint presented in the common denominator, energy, and created accepted by Thomas Park, the zoological last chapter of his thesis: "The trophic- around this focus a theoretical structure editor, after correspondence with G. Evelyn dynamic viewpoint, to be elaborated in this yielding predictions with which future Hutchinson who indicated the importance of paper, emphasizes the relationship of energy- workers could design their own theory in the availing (food cycle) relationships within investigations. development of ecology. the community to the process of succession'' (1). In essence he was grappling The Writing In September 1941, Raymond Linde- with the problem of time scales and ar- man and his wife, Eleanor Hall Linde- guing the importance of short-term On 19 March 1941, a year before the man, arrived in New Haven to begin trophic functioning to an understanding acceptance of his paper by Ecology, work with G. Evelyn Hutchinson, a lim- of long-term dynamical changes, this Raymond Laurel Lindeman received his nologist teaching at Yale University. depended upon the quantitative assess- Ph.D. in zoology from the University With him Lindeman brought the last ment of the biological relations of the or- ofMinnesota. As part of the completion of the chapter of his Ph.D. thesis, awarded the ganisms found in those lakes; and being an Ph.D. requirements, Lindeman had previous spring from the University of integration that he believed was hindered by already submitted the first chapter of his Minnesota for a 5-year study of the biol- the terminological distinction between thesis for publication, and it would ap- ogy of a senescent lake, Cedar Creek autecology and synecology. Lindeman's pear early in 1941 (5). He had therefore Bog (1). This chapter underwent several paper begins with a discussion of community made the decision to publish the thesis as draft revisions and was published concepts; and drawing on the work of a series of papers rather than a single posthumously in the journal Ecology as "The Thienemann and Tansley, he stresses the monograph, thus separating the great trophic-dynamic aspect in ecology" (2), and it functional integration of organic and body of collected data (6) from the theo- subsequently became the foundation for much inorganic cycles of nutritive substances: "The retical treatment represented in the future work concerning the dynamic flow of ecosystem may be formally defined as the trophie-dynamic paper (7). energy in plant and animal communities (3). system composed of physical-chemical- Lindeman was assisted throughout This paper would have formed but another biologicai processes active within a space- much of the fieldwork and writing of his historical fragment in the structure of modern time unit of any magnitude, that is, the biotic thesis by his wife, Eleanor, whom he ecological thought were it not for the fact that community plus its abiotic environment. married in 1938 (8). In the spring of 1941 when first submitted for publication, it was The concept of the ecosystem is believed they made plans for the examination of rejected by the editor on the advice of two by the writer to be of fundamental impor sediment cores taken from Cedar Bog Lake referees who were prominent limnological tance in interpreting the data of dynarnic with Ray performing chemical spectroscopic ecologists. After a subsequent exchange of ecology." analysis and Eleanor identifying diatom letters between Thomas Park, editor, and There follows a lengthy discussion microfossils. A year earlier, Ray had met Hutchinson, a somewhat modified version of of trophic operations; here the most Edward Deevey (9) at the hydrobiology the manuscript was accepted. Because of the important intellectual contributions of meeting at Madison (10), who had suggested ecological importance of the work, and the the paper are created. Quoting from a that Raymond apply for a fellowship to work unusual circumstances of its publication, I will locally published set of lecture notes by with Hutchinson at Yale. This he did, and in recount here the story of its rejection and Hutchinson, Lindeman establishes a the- April 1941 he was awarded a Sterling ultimate acceptance (4). oretical model of nutrient cycling ex- fellowship for a year. He worked on Perhaps the most prominent problem pressed explicitly in terms of energy flow manuscripts drawn from his thesis through to which ecologists in the early decades symbolized by mathematical equations. He the spring and submitted a large paper on of this century addressed themselves then proceeds to calculate the values of the food cycle dynamics which formed the main was the structural and temporal organization appropriate terms from his own data and body of data supporting his general of plant and animal communities. This was those of others. The analysis of trophic conclusions in the thesis (6). The final particularly true for the underlying ecological relations in terms of energy leads easily to chapter of his thesis was undergoing re- processes that determined the patterns of concepts of biological efficiency, and visions and would become the future change seen in the distribution of organisms Lindeman arrives at several very general trophic-dynamic paper. and the succession of species following relations regarding the flow of energy in In late August the Lindemans arrived natural and human disturbance. Lindeman ecosystems. In the final section in New Haven and another revision of believed that the understanding of ecological of the paper these relations are brought the trophic-dynamic paper was immedi- succession in lakes over long periods of to the analysis of successional devel- ately begun, stimulated by conversations timechose for study the trophic (nutritional) opment, with particular emphasis on with Hutchinson and a recently com- relations of all the inhabitants of a shallow, rate-controlling processes and the eco- piled, but not widely circulated, manu- weedy body of water lying in the transition script of Hutchinson entitled "Lecture 2bjbjÏ2Ï2222222222222222222 20î22- that he had great respect for the view- notes on limnology" (11). Manuscript 3 X22- point of the referees but felt that they [see (7)1 was completed by the end of the X228222222222222222222222222222222ÿÿ were intolerant of opinions other than month, and the appearance of quotes ¤222222222ÿÿ¤222222222ÿÿ¤22222222222 their own. The paper presented "practi- from Hutchinson reflect the influence of 222222ˆ22222: 222222: 22: 222222: 222222: cal working methods for evaluating and Yale on Lindeman's ideas. He immedi- 222222: 222222: 22µ22222222222N integrating the complex processes acting ately set about catching up ì¥Á25@ 222222–222222–222222–222222–22t2 within many types of natural commu- 22ð¿22222222222228¥22 immediately much interested in the ideas, nities-methods whose value could be contributed some of his own, and generously tested by certain minor modifications of spent a lot of time with me in rounding them research programs. Because this ap- out. The Cedar Bog Lake data adapted proach has given reasonably satisfactory themselves beautifully for illustrating the results in preliminary application, I feel trophic principles. The greatest gaps seem to that other ecologists (not necessarily lim- be in obtaining adequate terrestrial data. nologists) should be given an opportu- Dr. Hutchinson very strongly urged that the nity to consider this viewpoint with re- essay be published as soon as possible and spect to their own problems" (16). He has sent it off to Park for Ecology [see (13)]. made plans to rework the manuscript I'm afraid you're going to say that I've and submit it to the Quarterly Review of hazarded a great deal of theory on very little Biology. information, and you may be right. I have a The referees of the paper were feeling, though, that at least some of the ideas Chancey Juday at Wisconsin and Paul are piquing enough to start some people Welch at Michigan, the two most promi- making ecological studies on the basis of nent limnologists in the country. In addi- productivity and efficiency, and that would be tion to specific minor criticisms of the quite gratifying even though some of the data, the referees had the following gen- hesitantly proposed "principles" turn Out to eral comments to make: be wrong. I should like very much to have you and Dr. JUDAY: A large percentage of the following Lawrence comment on this latest brain-child, discussion and argument is based on "belief, if you care to--even though it be a none-too- probability, possibility, assumption and gentle reprimand. I'm really very grateful for imaginary lakes" rather than on actual all the criticisms and encouragement you've observation and data. The chances are that the already given--and feel that many o f the author's beliefs and imaginary lakes would be good parts of the paper (if any) were due to very different entities if he had a background the stimuli given by yourselves and the of observations on fifty or a hundred of the spirited seminar discussions out at your home 10000 lakes claimed by the state of last year. Minnesota instead of on only one, and that a Hoping that you have enjoyed a fruitful special type. According to our experiences, summer, are in excellent health and not too lakes are "rank individualists" and are very much pursued by hare-brained graduate stubborn students, about fitting into mathematical formulae and I am, artificial schemes proposed by man .... Some Ray Lindeman of the "broad generalizations" mentioned in the paper are certainly very broad; so broad in Lindeman now devoted further work fact that they cannot be regardedas having to the analysis of microfossils and the re- much value. visions of manuscripts in the senescent lake series (14). W£LCH: I would raise the question of suitability for publication in Ecology. This The Rejection paperis admitted by the author to be an .essay, and while I do not wish to put myself In the middle of November, more than in the position of suggesting to you what your a month after he submitted the trophic- editorial policy should be, I would express dynamic paper to Ecology, Lindeman re- my own feeling that papers in the form of ceived the letter of rejection. Park wrote, general essays should ordinarily be excluded. "lit is] with some reluctance and distress It seems to me unfortunate if the space which that I feel forced to take this action... should be occupied by research papers is [I] found your paper stimulating ... partly consumed by "desk produced" papers [but] I am not really competent in this unless they be of a most unusual and field." Both referees had recommended significant kind. In my humble opinion this rejection because the paper was without kind of treatment is sufficient evidence and premature, and premature. Limnology is not yet ready for therefore not suitable for Ecology (15). generalizations of this kind. The basic Lindeman was very distressed and wrote background data for such a paper is far too fragmentary. If Dr. Lindeman could put this are that such a theoretical study is very Because I feel that Lindeman's paper will paper aside for ten years, then bring it out and desirable and that Ecology should cover the actively encourage certain important kinds of see how it looks in the light of what we hope whole field of the subject. I was very pleased investigation, I believe it should be published ill be the added accumulation of limnologi- to see that you printed Haskell's work, as soon as possible. The very fact that he has cai information, he might possibly con- although I disagree with about one half had to use fragmentary data indicates that gratulate himself that he deferred its of it, and sincerely hope that further without an orienting hypothesis, the need for publication. What limnology needs now most contributions from that most stimulating obtaining the required observational and ex- of all is research of the type which yields individual will appear. Far from agreeing perimental results has not been clearly envi- actual significant data rather than postulations with Referee 2 as to what limnology needs, I sioned. As I have indicated, it is quite beyond and theoretical treatments. feel that a number of far-reaching hypotheses the powers of one man to perform the investi- that can be tested by actual data and which, if gation himself. Even should none of his The Acceptance confirmed, would become significant generalizations ultimately hold, the work of generalizations, are far more valuable than an disproving them will provide important The lines of difference on the issue of unending number of marks on paper information that would probably be obtained publication were clearly drawn 4 days indicating that a quantity of rather unrelated in no other way, and all authors should be later when Hutchinson, writing to Park observations has been made. As an example, allowed to take comfort in the words of Sir on general editorial matters for Ecology, one of the things that has impressed me most Thomas Browne, "the certainty here-of let the enlarged specifically on the Lindeman in my study of Linsley Pond is the fact that arithmetick of the last day determine.., manuscript (17): the morphometry of the basin affects the although at last we misse the truth, we die vertical distribution of certain sub- notwithstanding in harmless and inoffensive I also received your letter about Lindeman's stances when the lake is stratified. The errors, because we adhere unto that, work in which you courteously ask for my conditions for this to be apparent are clearly whereunto the examen of our reasonS, and reactions as to the opinions submitted by the very special, though the underlying cause is honest enquiries induce us" (Pseudodoxia referees. I entirely understand your not probably of very general significance. Yet epidemica, Bk. 6 Chp. VI). wishing to publish the work, in the face of because no one has been sufficiently aware of You will realize that much of the material at such adverse comments. In view of the fact the theoretical aspects of heating and the end of Lindeman's paper had occurred to that Dr. Lindeman himself felt uncertain as to transport of chemical material in stratified me independently. As the biogeochemical its appearance, in spite of my favorable view, lakes, no cases have ever been published in treatise in which it is discussed progresses so he submitted the manuscript to two plant which analyses were made at close enough slowly and threatens to assume such monu- ecologlsts with whom he has studied, both of vertical and temporal intervals to permit any mental proportions, it seemed best to hand whom happen to be on our editorial board, judgment as to whether the phenomenon him the relevant material, to use as he saw fit. and who reported respectively that the work occurs in other lakes in North America. At This fact may in part explain my strong feel- "looks excellent" and is "definitely in good times I have felt quite desperate about the ings on the matter, but over and above such shape for publication." May I suggest that number of opportunities that have been feelings I hope I have made clear that an im- you communicate to the referees (I thihk that missed in the middle western regions for portant question of policy seems to me to be internal stylistic peculiarities reveal their obtaining data confirming or, disproving the involved. identity as men for whom I have great hypotheses that have been forced on us by personal regard and who have been most kind our little lake here. In genetics, experimental Park sent a copy of Hutchinson's com- to me on many occasions) this letter, except embryology, biodemography, and other ments to the two reviewers, who still ad- for the preceding sentence, that they may sciences where the phenomena are not spread hered to their earlier criticisms, and then realize that most of the specific points out over great ranges of time and space, it is wrote Lindeman, "if you care to revise challenged are matters for which I, rather possible for one worker to produce your manuscript in any way you see fit than Lindeman, am responsible. I am most hypothesis after hypothesis, discarding and resubmit it to me, I shall try to find anxious that the encouragement I gave him, those that are invalid after a few weeks' work an impartial referee who will . . . make to forward this paper for publication, shall not in the laboratory. In field ecology, it is neces- the final decision" (18). Ray agreed to re- prejudice his reputation as an ecologist. My sary to have data collected over many months submit the paper after Christmas, with own view is that, if the work is published, or years, and for comparative purposes stud- several revisions incorporating sugges- after the ten years or so suggested by Referee ies are needed on localities very widely sepa- tions of a number of ecologists to whom 2 have elapsed, Lindeman will feel that he rated in space. To obtain the kind of data re- he had sent the work (19). has played a very considerable part in a quired takes two or three years' work on a As Christmas approached Eleanor healthy reorientation of ecological research. single locality; to suggest that any one wrote to Ruth Patrick concerning a visit Before that time, however, he will need a individual should wait until he has completed to the Philadelphia Academy of Natural position somewhere and although I still think investigations on fifty or one hundred lakes is Science over the holidays to identify dia- it most desirable to publish the work, I do not ironical rather than practical. It is therefore toms (20), and Ray made plans to travel want my backing of it to be a handicap to most important that all ecologists should have to Dallas to attend the meetings of the him. the opportunity to acquaint themselves with American Association for the Advance- As regards the detailed comments of referee the theoretical possibilities that may guide ment of Science. He would deliver an ad- 1:... them in their collection of data, and that they dress at these meetings coauthored with · . . The second part of criticism 3 of Referee should consider it their duty to acquire an ob- Hutchinson (21) in which many of the 1 and the comments of Referee 2 virtually jective understanding of the significance of conceptual innovations of the trophic- reduce to a discussion of a) whether any potentially fruitful hypotheses that may dynamic paper were presented. He was theoretical work is legitimate in ecology, and be advanced. This is, of course, the normal also very busy producing the draft of b) whether Ecology should print it. My own procedure in astrophysics, an even more ex- manuscript 4 which he planned to send feelings, quite apart from Lindeman's paper, pensive and time consuming science. to Park. It was shortly after returning from Dal- many ecological terms and concepts quantitatively represented by mathemat- las that Ray underwent a "mild recur- found in An Ecological Glossary (27). ical relations, Lindeman was able to de- rence of the jaundice attack I had in This concern with the establishment of velop a number of predictions with 1937" which put him in the Yale-New an appropriate language of ecology was which the validity of the model could be Haven Hospital for 3 weeks (22). In the probably the inevitable result of the inde- assessed. middle of February he returned home but all pendent development of plant and ani- Third, the trophic-dynamic approach field and laboratory work was suspended. On mal ecology, as well as limnology and identified a fundamental dynamic pro- 13 January he had sent a copy of the revised environmental physiology. In its enthusi- cess, energy flow, with which the season- trophic-dynamic paper to Victor Shelford, a asm to solidify the classification of eco- al trophic relations of organisms could be well-known animal ecologist, and he was logical patterns into a nomenclatural de- integrated into the long-term process of waiting for his response before submitting it scription, the synthetic impulse in ecolo- community change. Guided by the analo- again to Ecology. Sheiford's letter came on gists could easily lose sight of the under- gy between developing organisms and March (23): lying processes determining those pat- the dynamics of succession, most plant terns. Thus the first important con- ecologists determined the correct classi- The paper seems to me to be very well sequence of Lindeman's paper was to fication of Communities by changes in written and on a very interesting topic. I have, stress the major role of trophic function, the abundance and distribution of spe- however, not specialized in lake metabolism particularly quantitative relations, in the cies assumed to be characteristic of par- and so am not able to offer suggestions in that determination of community patterns ticular developmental stages. Limnolo- field, and wish you luck in going forward through succession. gy, more than ecology in general, had with the idea. Second, Lindeman's paper established stressed the importance of productivity tbe validity of a theoretical orientation in in order that various types of lakes might Within a week Ray, had a new copy ecology. Although the foundations of fu- be set into a general classification based typed and sent to Park. He in turn sought ture ecological theory were being quietly in large part on the abundance (biomass) an impartial third referee in the person of established in the 1920's and 1930's (28), of plankton and bottom faunal commu- his colleague at Chicago; W. C. Allee, much of this work was considered of nities. This approach, however, failed to who was unenthusiastically neutral on little relevance to the "real world"; and consider the metabolic relations of these the manuscript (24). The controversy some 20 years would pass before its in- "superorganisms" (31); and by intro- surrounding the issue had become well fluence was felt (29). Up to this time the ducing energetics, Lindeman reduced known among a number of ecologists, major tradition in ecological studies in the processes of the food cycle to their and Allee's lack of strong support put the the United States was the description most basic component. The importance young editor of Ecology in a delicate po- and classification of plant and animal of this innovation was even acknowl- sition. On 23 March 1942, Park wrote to communities. Little truly theoretical edged by Paul Welch, one of the original Lindeman (25): work, involving the construction of referees of the paper. In the second edi- mathematical models, had been pub- tion of his book Limnology first pub- I have carefully considered your revised lished and incorporated into the body of lished in 1935, Welch added, in the chap- manuscript and am herewith accepting it for accepted ecological knowledge. This can ter on biological productivity, a special Ecology. I rather imagine that the original be clearly seen in the book Bio-Ecology section on trophic relations (32): referees will still object to certain of its basic which was published in 1939 and repre- premises but I think it best to publish your pa- sented the summation of all previous Somewhat recently, certain investigators, per regardless. Time is a great sifter in these ecological principles by the most emi- notably Lindeman (1942), have attempted to matters and it alone will judge the question. nent plant and animal ecologists of the analyze the events within a food complex in time, Frederick Clements and Victor terms of energy. Because of the great paucity The Significance for Ecology Shelford. The classificatory approach to of detailed information basic to dependable the description of communities utilized formulation of such concepts, any discussion It seems appropriate to note several the biome ("the great landscape types of at present is largely hypothetical and must be points of significance in the publication vegetation with their accompanying ani- regarded as suggestive only. Hints that these of Lindeman's paper. First, ecology at mals'') as its fundamental unit, and the complex interrelations may eventually yield this time represented the merger of a earlier developmental stages of such to mathematical analyses appear in the work number of rather independent lines of re- units represented the process of commu- of Lindeman and others. Entry into the spec- search; it was primarily derived from a nity succession. Underlying this whole ulative aspects of this subject will not be un- very empirical tradition of field investi- ecological approach was the metaphor of dertaken here. However, out of the pioneer~ gations somewhat systematic in nature the developing organism, and the stable ing work done thus far, there have arisen bio- and 19th-century natural history in climax community is explicitly consid- logical Conclusions which seem to have cer- which most generalizations were the in- ered "a complex organism, or superor- tain validity. ductive descriptions of data. There was ganism, with characteristic development immense interest in problems concerning and structure. As such a social organ- Welch then continues to report the gen- the classification of observed ecological ism, it was considered to possess char- eral conclusions of the frophic-dynamic patterns, leading to a prolific termi- acteristics, powers, and potentialities paper, a result that surely would have nology and the consequent conflicts of not belonging to any of its constitu- pleased Lindeman. opinion concerning nomenclature (26). ents or paris ... the community; as Finally, the critical role of Hutchinson During his last year at Minnesota, for in- noted above, is more than the sum of its in the development and publication of stance, Lindeman would gather with oth- individual parts, that it is indeed an orga- this paper must be mentioned (4). Much er graduate students in the home of Wil- nism of a new order" (30), By creating a Of modern ecology has grown from the liam Cooper to debate the meaning of the theoretical model of trophic interactions, communal relations he was able to es- tablish with those fortunate enough to oretical conclusions. The present contribution treatment and updating of the limnological work with him, and from the depth and does far more than this, as here for the first studies, see the appropriate re- endurance of his intellectual vision. time, we have the interrelated dynamics of a gional chapters in Limnology in North biocoenosis presented in a form that is ame- America, D. Fry, Ed. (Univ. of Wisconsin Illness and Death nable to a productive abstract analysis. The Press, Madison, 1963). question, for instance, arises, "What deter- 4. For more details, see R. E. Cook, in As the spring of 1942 developed, mines the length of a food chain?"; the an- preparation. Most of the information in this Lindeman's health was not improving; swer given is admittedly imperfect, but it is article has been drawn from the papers of R. and, in a letter to Don Lawrence in far more important to have seen that there is a Lindeman deposited in the Archives of April, he wrote, "I am desperately anx- real problem of this kind to be solved. That Syerling Library, Yale University. ious to get back to my own work, on the final statement of the structure of a bio- 4a.E.P. Odum, Science 195, 1289 (1977). which almost nothing has been done coenosis consists of pairs of numbers, one an 5. R. L. Lindeman, "The developmental since Christmas, and hope to spend at integer determining the level, one a fraction history of Cedar Creek Bog, Minnesota," in least a few hours a day on it soon. The determining the efficiency, may even give Am. Midi. Nat.'25, 101 (1941). This paper is trouble is obscure--hepatic cirrhosis of some hint of an undiscovered type of mathe- the first in a series of works entitled unknown etiology, with a possibility that matical treatment of biological communities. "Ecological studies of a senescent lake" of it may become progressively worse in Though Lindeman's work on the ecology and which five were finally published. This was spite of everything" (33). Lindeman had history of Cedar Bog Lake is of more than lo- not Lindeman's first publication which was another hepatic attack at the end of April cal interest, and will, it is hoped, appear of "Some affinities and varieties of and soon wrote his close friend, Charles even greater significance when the notes the planktonic rotifer, Brachionus Reif, "We hope to be at the University made in the last few months of his life can be havanaensis Rousselet," Trans. Am. Microsc. of Pennsylvania next.year, as I have a coordinated and published, it is to the present Soc. 58, 210 (1939). Other papers in the fellowship there, but(confidentially) paper that we must turn as the major contri- "senescent lake" series are: II. M. F. Buell there is a better than even chance I won't bution of one of the most creative and gener- and H. F. Buell, 1941. "Surface level survive the summer. My liver trouble ous minds yet to devote itself to ecological fluctuations in Cedar Creek Bog, has gotten irregularly worse, in spite of science. Minnesota," Ecology 22, 317 (1941); HI. R. the best doctors, and after 4 months is L. Lindeman, "Seasonal food cycle dynamics beginning to show visceral oedema. I ex- in asenescent lake," Am. Midi. Nat. 26, 636 pect to have an exploratory operation (1942); IV. "Experimental simulation of soon in the more or less desperate hope winter anaerobiosis in a lake," Ecology 23, I that they can find out what the cause is References and Notes (1942); V. R. L. Lindeman, "Seasonal and then try for a cure. Eleanor is work- distribution of midge larvae in a senescent ing at the Yale Library and should be 1. R. L. Lindeman, "Ecological dynamics in a lake," Am. Midi. Nat. 27,428 (1942). These able to continue if worse follows worse" senescent lake," thesis, University of papers, along with the trophic--dy- (34). Minnesota, (1941). A copy of this work may namic paper, comprise the published works of On 15 June, Ray 'underwent surgery be found in Kline Science Library, Yale Lindeman. For a recent addition appropriately and died within 2 weeks. In an adden- University, New Haven. considered in this series, see M. F. Buell, H. dum to the trophic-dynamic paper, 2. R. Lindeman, Ecology 23, 399 (1942). F. Buell, W. A. Reiners, "Radial mat growth Hutchinson wrote (35): 3. Indications of the importance of this paper on Cedar Creek Bog, Minnesota," Ecology While this, his sixth completed paper, was for the understanding of the functioning of 49, in the press, Raymond Lindeman died after a ecosystems can be found in, "Dynamics of 1198 (1968). long illness on 29 June 1942, in his twenty- production in aquatic communities," Ecol. 6. R. L. Lindeman, "Seasonal food cycle seventh year. While his loss is grievous to all Monogr. 16, 311 (1946); L.. Slobodkin, dynamics in a senescent lake," Am. Midi. who know him, it is more fitting here to dwell "Energy in animal ecology," Adv. Ecol. Res. Nat. 26, 636 ( 1942 ) 26 on the achievements of his brief working life. 1, 69 (1962): V. F. Gallucci, "On th~ 7. All cmxespondence, and draft manuscripts The present paper represents a synthesis of principles of thermodynamics cited in subsequent footnotes, may be found Lindeman's work on the modern ecology and in ecology," Annu. Rev. Ecol. Syst. 4, 329 in box 1, Lindeman Papers, in the Archives of past history of a small senescent lake in Min- (1973); R. E. Ricklefs, "Energy flow in Yale University. Four draft manuscripts exist nesota. In studying this locality he came to ecosystems,'' in Ecology (Chiron Press, in the Archives: manuscript 1 (February realize, as others before him had done, that Newton, Mass., 1973), chap. 41; R. B. 1941); manuscript 2 (March 1941); the most profitable method of analysis lay in Williams, "Computer simulation of energy manuscript 3 (September 1941), which is reduction of all the interrelated biological flow in Cedar Bog Lake, Minnesota, based on shortly after Lindeman arrived at Yale; and events to energetic terms. The attempt to do the classical studies of Lindeman," in manuscript 4 (January 1942) which was this led him far beyond the immediate prob- Systems Analysis and Simula- written after the rejection of manuscript 3. lem in hand, and in stating his conclusions he tion in Ecology, B. C. Patten, Ed. (Academic 8. Owing to a childhood accident, Ray was felt that he was providing a program for fur- Press, New York, 1971), p. 543; see also, L. partially blind in one eye. This made him ther studies. Knowing that one man's life at B. Slobodkin, "On the inconstancy of somewhat dependent upon Eleanor for best is too short for intensive studies of more ecological theories," in Growth by microscopic identification of specimens. She than a few localities, and before the manu- Intussusception, Ecological Essays in Honor was a trained biologist herself, having script was completed, that he might never re- of G. Evelyn Hutchinson, E. S. Deevey, Ed. graduated from the University of Minnesota turn again to the field, he wanted others to [published in Trans. Conn. Acad. Arts Sci. in 1939; and in the last year they spent think in the same terms as he had found so 44, 291 (1972)] for a revisionary assessment together, she examined diatoms in sediment stimulating, and for them to collect material of Slobodkin's own work. For a historical cores taken from Cedar Creek Bog. This that would confirm, extend, or correct his the- resulted in an unpublished manuscript coauthored with Ray, "Microfossils in the climax equilibriurn) and were principally and encouragement throughout the research sediments of a senescent lake, and their contributed by J. P, Carpenter, W. S. Cooper, and writing. This work was supported by a successional significance: a preliminary E. S. Deevey, A. Hodge$on, D. B. Lawrence, Harvard Graduate Society Award. report." Mrs. Eleanor Burns has been of H. J. Lutz, A. E. Parr, and V. E. Shelford. considerable help in my research. 20. Eleanor Lindeman to Ruth Patrick, 11 9. E. S. Deevey had completed his thesis November 1941. work onb"Typological succession in 21. G. E. Hutchinson and R. L. Lindeman, Connecticut lakes" at Yale and had taken a "Biological efficiency in succession" Bull. teaching job at Rice University in 1939. Ecol. Soc. Am. 22, 44 (1941). 10. University of Wisconsin, A Symposium 22. Lindeman to Eddy, 10 February 1942. on Hydrobiology (Univ. of Wisconsin Press, 23. Shelford to Lindeman, 11 March 1942 Madison, 1941). 24. T. Park, personal communication. Il. Lindeman refers to this work as "Recent 25. Park to Lindeman, 23 March 1942. Park advances in limn01ogy (in manuscript) was clearly a strong supporter of a more 1942." The manuscript was distributed by the theoretical approach to ecology [see, for Osborn Zoological Laboratory, Yale instance, W. C. Alice and T. Park University, and copies exist in Kline Science "Concerning ecological principles," Science Library and Yale Archives in the Hutchinson 89, 166 (1939); and T. Park, "The laboratory papers. population as a test of a comprehensive 12.R. L. Lindeman, letter dated 29 September ecological system," Q. Rev. Biol. 16,274 and 1941. Although at the University of 440 0941)] and he played an important role in Minnesota Lindeman was officially the bringing theoretical and laboratory population student of Samuel Eddy, a fish systematist studies into the main body of ecological and limnologist, it appears that he was more work. strongly influenced by William S. Cooper, a 26.This was a problem of considerable plant ecologist in the botany department who concern to many ecologists of the day. See, was the intellectual stimulus for informal for example, 2 book reviews by C. Elton meetings of graduate students and younger entitled "American ecology" and faculty to discuss ecological concepts and "Scholasticism in ecology," inJ. definitions. Anita. Ecol. 9, 148 and 151 (1940). 13.Hutchinson was a consulting editor of 27. J. R. Carpenter, An Ecological Glossary Ecology t this time and Thomas Park at the (Univ.of Oklahoma Press, Norman, 1938). University f Chicago was zoological editor. 28. A. J. Lotka, Elements of Physical Biology 14. he seasonal food cycle paper was (Williams & Wilkins, Baltimore, 1925) G. F. originally submitted to American Midland Gause, The Struggle for Existence (Williams Naturalist on 3 May 1941. In rewriting the & Wilkins Baltimore, 1934); V. Volterra, trophic-dynamic paper (manuscript 3), "Varazioni e fluttuazioni del numero Lindeman extracted part of the comparative d'individui in specie animall convivanti," discussion of productivity from the seasonal Mem. Accad. Lincei 2 (No. 16), 31 0926). food cycle paper and incorporated it into 29. L. B. Slobodkin, Growth the Regulation manuscript 3 to provide supporting evidence of Animai Populations (Holt, Rinehart and for some of the theoretical concepts. On 29 Winston, New York, 1961). September he wrote a letter to the editor of 30. F. E. Clements and V. E. Shelford, Bio- American Midland Naturalist requesting Ecology (Wiley, New York, 1939), pp. 20-21. changes in the seasonal food cycle paper; It should be noted that this organismic and, approach paralleled the whole conceptual full of confidence, he wrote in the revised framework being established in productivity section o( this work, "The developmental biology at this time; see D. H. theoretical aspects of apparent and true food- Harroway, Crystals, Fabrics and group efficiencies for Cedar Bog Lake is Fields (Yale Univ. Press, New Haven, 1975). discussed in detail in a forthcoming paper 31. G. E. Hutchinson, "Review of Bio- (Lindeman, 1942 c)." He was later (17 Ecology,"Ecology 21, 267 (1941). November 1941) to strike this reference with 32. P. Welch, Limnology (McGraw-Hill, the comment, "I have just decided that this New York, ed. 2, 1952), p. 350. theoretical paper is too premature and should 33. Lindeman to Lawrence, 13 April 1942. not yet be published for some time." 34. Lindeman to Reif, 16 May 1942. 15. Park to .Lindeman, 14 November 1941. 35. Addendum, G. E. Hutchinson, Ecology 26 32, 417 (1942). 16. Lindeman to Park, 18 November 1941. 36. I would like to thank Polly Winsor, 17. Hutchinson to Park, 18 November 1941. Pamela Parker, Donna Haraway, Ann Bum, 18. Park to Lindeman, 8 December 1941. Frank Salloway, and Sharon Kingsland for 19. Lindeman to Park, 13 December 1941. advice and comments on this manuscript. I Most of these suggestions involved relatively also owe a great debt of gratitude to Professor minor revisions (except for his concept of G. E. Hutchinson for his continual inspiration