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The emergence of ecology from
natural history
Keith R. Benson
The modern discipline of biology was formed in the 20th century from roots deep in the natural-history
tradition, which dates from Aristotle. Not surprisingly, therefore, ecology can also be traced to natural
history, especially its 19th-century tradition emphasizing the adaptive nature of organisms to their
environment. During the 20th century, ecology has developed and matured from pioneering work on
successional stages to mathematically rich work on ecosystem energetics. By the end of the century,
ecology has made a return to its natural-history heritage, emphasizing the importance of the integrity of
ecosystems in considering human interactions with the environment.
Today, the field of biology includes a vast array of diver- like molecular biology, ecology emerged as a distinct
gent and unique subdisciplines, ranging from molecular area in biology only at the turn of the century but very
biology to comparative endocrinology. With very few quickly developed its own conventions of biological
exceptions, most of these specialty areas were created by discourse. Unlike molecular biology and several other
biologists during the 20th century, giving modern biology biological subsciplines, ecology’s roots are buried deep
1 within natural history, the descriptive and often romantic
its distinctive and exciting character . However, before
1900, the field was much different because even the term tradition of studying the productions of nature.
2
biology was seldom used . Indeed, most of those who
studied the plants and animals scattered over the earth’s Perspectives on the natural world before the
surface referred to themselves as naturalists: students of 20th century
3 Aristotle, the western world’s greatest philosopher who
natural history .
Perhaps the most popular form of contemporary biology included the natural world in his philosophical treatments,
is the subdiscipline most closely related to the natural- was the first to record observations about the natural his-
4
history roots of biology: ecology. As with many things tory of the earth’s plants and animals . However, his teleo-
currently enjoying popularity, however, the term ‘ecology’ logical world of designed and invariable types hardly placed
is often poorly understood and even more inaccurately a stress upon the reciprocal and dynamic relationships
used. Considered by convention to be synonymous with that exist between the biotic world and the earth’s physi-
‘natural’, ‘environmental’or ‘conservation’, it is frequently cal environment. Not surprisingly, given his assumption
used to refer to a personal perspective on the natural of the eternal nature of species,Aristotle did not stress the
world or to a political position concerning the use of adaptive character of fauna and flora, which is perhaps
nature. In fact, many of those who describe themselves as ecology’s cornerstone. In fact, adaptation did not appear
ecologically oriented, as having an ecological perspective as a biological notion until nature was reinterpreted as the
or as being interested in ‘saving the ecology of the land’ product of a historical and developmental process at the
have never bothered to take a university-level course in end of the enlightenment (18th century). Thus, for almost
ecology or to have examined in any depth a classic eco- 2000 years, naturalists considered the earth to have been
logical text. As a result, the exact nature and definition of created originally much as it was observed.
‘ecology’remains obscured by its popular usage. As part of the scientific revolution capped by Isaac
In part, some of the definitional misunderstanding comes Newton at the beginning of the 18th century, natural
from ecology’s biological lineage. Certainly, its subject philosophers opted to examine the natural world for
matter (the planet’s ecosystems) has a much greater reso- mechanical explanations of natural phenomena, often in
nance with the general public than, say, the arcane and terms of mechanisms they could either observe in nature
esoteric subject that is molecular biology. Nevertheless, or infer from nature. These explanations, best exem-
plified by the law-like behavior of Newton’s universal
Keith R. Benson gravitation, promised to provide precise and knowable
information about nature, usually in mathematical form.
Is currently a professor of medical history and ethics at the Uni- No longer bound to accept the natural world as a created
versity of Washington, where he serves as Director of the Program
in the History of Science, Technology and Medicine. He is coeditor given, the philosophes of the enlightenment soon began
of two books on the history of American biology and has written to apply the Newtonian method to the biotic world.
numerous articles on the development of the biological sciences in
the USA. He is also a past Executive Secretary of the History of The limitations of this application became apparent
Science Society. At present, he is completing a book on the history almost immediately. Bernard de Fontenelle expressed the
of marine biology in the USA.
krbenson@u.washington.edu futility of the age’s mechanistic orientation when he
0160-9327/99/$ – see front matter © 2000 Elsevier Science Ltd. All rights reserved. PII: S0160-9327(00)01260-0 Endeavour Vol. 24(2) 2000 59
proclaimed that two mechanical watches will sit side by The ‘Darwin of Germany’, Ernst Haeckel, devoured On
side forever without producing a third but, if two dog the Origin of Species almost as soon as it was published,
‘machines’are placed side by side, more dogs would soon becoming an immediate convert to the theory of descent
appear! Animal generation, along with a host of other by modification, as evolution theory was originally known.
8
biotic operations, seemed to resist the simplicity of the In his influential book Generelle Morphologie , Haeckel
5
mathematical treatment of invariable mechanical laws . stressed the Darwinian notion of change over time pro-
Nevertheless, the mechanical philosophy opened the duced by the dynamic relationship of organisms and their
door for fresh investigations of plants and animals, their natural environments. As a measure of the importance of
relationships to each other, and their relationships to the this relationship, Haeckel, who was fond of neologisms,
natural world. By the beginning of the 19th century, the coined the word oecologie, referring to the study of the re-
notions of structural analogy (transformism) and func- lationship of organisms to their surroundings. In his popu-
tional integrity (biogeographical lar 1876 English edition of his ideas,
distribution) that investigators re- By the end of World War II, History of Creation, he noted that
corded from their observations be- ecology had become Darwin’s doctrine of adaptation
gan to lead them to examine the his- thoroughly transformed from provided the law-like nature to
9
torical record of the earth’s fauna explain ecological relationships .
6 scientific natural history to
and flora . This was particularly Haeckel was not, however, the first
true as naturalists observed that ecosystem ecology ecologist, nor did he immediately
different landscapes of the earth’s spawn an ecological program in
surface with almost identical physical conditions had Germany. Instead, he served as one of the seminal figures
remarkably different resident populations of plants and in the 19th century to stress the growing appreciation that
animals. This was quite a surprise because, according to the relationship of plants and animals to their natural en-
the prevailing view of natural law, the same environ- vironments was historical and dynamic. Two other Euro-
mental conditions should produce nearly identical pean naturalists, Oscar Drude and Eugenius Warming, who
species. Yet, for example,Australia had endemic forms of were influenced by these same ideas, soon began to stress
life seen nowhere else on the globe. the study of pflanzengeographie (plant geography), noting
The new stress on the uniqueness of the forms of life the community structure of plant groupings that character-
along with the uniqueness of the landforms served as the ized specific landforms with specific environmental con-
fertile soil for what became ecological insights. But those ditions10. Remarkably, for there was not an equivalent scien-
who made these observations were not, per se, ecologists. tific community in the USA to match that in Europe, these
Instead, they were among the 19th century’s most accom- ideas were picked up by American naturalists at the end
plished naturalists. At the beginning of the 19th century, of the 19th century: Charles E. Bessey at the University of
the German adventurer Alexander von Humboldt waxed Nebraska and John Coulter at the University of Chicago.
eloquent about the characteristic physiognomic features
of the landscapes in South America, stressing how these Ecology’s early-20th-century roots
visible features (hence his reference to physiognomy) Neither Bessey or Coulter, however, is well known as an
depended in large part upon the environmental character- ecologist. Both were natural historians at their respective
istics that controlled the flora. Four decades later, Joseph institutions, trained in the traditional methods of natural
Dalton Hooker was to make similar observations in his history, emphasizing the naming, description and classifi-
travels to the Himalayas, New Zealand, Tasmania and cation of plants and animals. However, both were also well
Australia itself. read in the new trends of the emerging field of biology
Hooker’s more famous compatriot and colleague Charles and they knew of the implications that evolution theory
Darwin observed similar features during his famous voy- had on their own studies. Encouraging their students to
age aboard the ‘Beagle’, completed about a decade before pursue new research opportunities that stressed the new
Hooker’s voyages but receiving their most influential biological perspectives at the end of the 19th century,
7
reading after the publication of On the Origin of Species . Bessey led Frederic Clements to the work of Drude, and
11
In fact, the importance of Darwin’s influential work on Coulter directed Henry Chandler Cowles to Warming .
the development of ecology cannot be emphasized enough. Certainly, there may be other national claims to the
After all, Darwin was the first to stress forcefully that ani- origins of ecology, including the German one, but the role of
mals and plants were not perfectly adapted to their natural the USA in the development of ecology through these two
environments, as earlier naturalists had once believed. midwestern schools cannot be diminished. Coulter’s star
Instead, they represented only the best-adapted forms pro- pupil, Cowles, was soon working on the plant-community
duced at a particular place and time by selective forces, structure of the sand dunes along Lake Michigan,
which, in turn, chose the adaptation that was optimal for referring to his work as physiographic plant ecology. That
the conditions that then existed. When conditions (including is, he was interested in studying the relationship of plant
both biotic and abiotic factors) changed, so the adaptive communities to the underlying geological formation, a
needs also changed. Those plants and animals that in- relationship that he thought explained why the physiology
cluded an adaptive characteristic favorable to the new of the plant responded to the geological features of the
conditions would survive; those not so favorably equipped land, leading to characteristic geographical groupings of
would perish. plants.
60 Endeavour Vol. 24(2) 2000
Noting that the plant community and the environment Forbes’ emphasis on the dynamic equilibrium of all the
were in constant flux, Cowles was led to view natural components of the lake seemed to offer a different way
systems as being characterized by change: processes that to appreciate nature, one that stressed the natural world
were evident in the succession of floral community struc- as a system. Instead of succession stages, Forbes pointed
ture observed in the sand dunes12. Early in the 20th century, ecologists toward seeing the world as a vast array of inter-
Cowles transported his new ideas to a marine setting at dependent environments through which materials and en-
the University of Washington’s new laboratory at Friday ergy were constantly being cycled. Charles Elton adopted
Harbor (on the San Juan Islands in the Gulf of Georgia), this position in his influential book, Animal Ecology (1927),
where he taught the first course in marine ecology in the freeing ecologists from merely examining the physical
USA. His most famous student, Victor Shelford, was to factors of environments but also obligating them to search
continue his work on the West Coast, working on eco- for methods to evaluate the immense constellation of
16
logical investigations at Friday Harbor through the early factors that determine community structure .
1930s.
In Nebraska, Bessey’s protégé was Frederic E. Clements, The maturation of modern ecology
a student who came to understand the Drude version of Although Clements’ work continued to be enormously
community structure through his own survey of native influential, its stress on deterministic climax communities
vegetation in that State. First in the Phytogeography of drew increasing criticism from ecologists in the 1930s,
Nebraska (1900), then in Research Methods of Ecology especially from those who were interested in adding the
(1905) and finally in Plant Succession (1916), Clements study of animals to ecology. The English ecologist A.G.
laid out his influential ideas of plant-community struc- Tansley provided the most cogent attack in an article in
ture, succession stages of community development and the new journal Ecology in 1935, challenging his col-
the climax community,the ultimate goal of mature natural leagues to adopt the term ecosystem, a reference stressing
13
habitats . But Clements did much more than just provide the dynamic nature of community structure rather than
17
a programmatic design for the new field of biology. Clements’ goal-directed climax stage . When G. Evelyn
Supported by the Carnegie Institution of Washington, he Hutchinson and his student Raymond Lindeman provided
was able to develop laboratory facilities in ecology in clear, albeit highly complicated, mathematical models to
Colorado and California, where he depict the various interacting com-
provided empirical and experi- ecology and its professional ponent parts of the ecosystem, ecol-
mental demonstrations of his new practitioners are often ogy promised to become a fully
ideas, thus helping to popularize greeted as advocates for the mathematized and experimental
the new field of ecology. conservation or preservation discipline.
Most of these biological investi- Lindeman, in particular, contri-
gators were also well aware that of the natural world buted to this important change
they were breaking new ground in when he published a paper in 1942
the understanding of the dynamical relationship between that synthesized the work of Clements, Elton, Tansley and
plants and animals and their environments, thus contribut- his mentor Hutchinson by speaking of biogeochemical
ing to a new field of ecology. Clements referred to the cycling, energy flow through trophic levels and dynamic
18
ecologist as an outdoor physiologist, a reference to the ex- succession . Even more importantly, he saw the con-
citing laboratory-based area of physiology just emerging in tinuous cycling of material through the ecosystem as an
institutes throughout Germany and England and begin- energy-driven process that included producers (organisms
ning to appear in the new scientific universities in the USA. that fixed the energy from the sun), consumers and de-
Shelford referred to the new field as scientific natural composers, which cycled material back to the producers
history, again a reference to the experimental and labora- as energy from the sun continued its one-way flow through
tory (field-laboratory) approaches developed at Nebraska the ecosystem.
and Chicago that acted to move natural history in a By the end of World War II, ecology had become
14
different direction from its traditional museum heritage . thoroughly transformed from scientific natural history to
However, limitations to the new field cropped up almost ecosystem ecology. Nowhere was this more evident than
immediately. One problem was that it was difficult to in the publication of the ecologists’ Bible, Principles of
investigate animals with the same approaches used for Animal Ecology, written by W.C. Alee and his colleagues
plants, as the fauna did not remain fixed to the environ- at the University of Chicago during the War but only
19
mental substrate in the same way that the flora was fixed. published in 1949 . Although community structure and
Thus, the physical factors that provided the causal deter- succession still remained basic ecological principles, the
minates of community structure were more difficult to climax community was now treated as virtually syn-
establish. Second, some ecologists soon began to question onymous with mature community and most of the book
the goal-directed nature of Clements’ climax communi- dealt with the dynamic inter-relationships of ecological
ties, preferring to view the natural world as being in investigations. Even more important, the last chapter of
constant flux. the book featured a long discussion of the evolution of
Many of these ecologists were influenced by the re- interspecies integration and ecosystems, emphasizing the
publication of an article by Stephen Forbes from 1887, influence of the Hutchinson and Lindeman approach
15
which received its greatest reception in the 1920s . (both authors are heavily cited in the book).
Endeavour Vol. 24(2) 2000 61
Even more dramatic for the new direction in ecology was Notes and references
the publication of Eugene Odum’s new text, Fundamentals of 1 Rainger, R. et al., eds (1988) The Development of American
Ecology (1953), with its overt recognition of systems theory. Biology, University of Pennsylvania Press
2 The Frenchman Lamarck and the German Treviranus are
In addition to the mathematical modeling from Hutchinson generally credited as being the first to originate the word
and Lindeman, Odum also benefited from his experience biologie at the beginning of the 19th century, which they
with the Atomic Energy Commission and its Atoms for used to separate the living world from the world of the inert.
Joseph Caron has written about the origin of the word biology
20 in its modern context: Caron, J. (1988) ‘Biology’in the life
Peace project . Using radioactive materials, he was able to
observe and to measure the recycling of inorganic materials sciences: a historiographical contribution. Hist. Sci. 26, 223–268
throughout the ecosystem, leading him to borrow from 3 The term ‘natural history’is the time-honored way to describe
the investigation of the natural world, including plants, animals
physiology and to refer to the metabolism of the ecosystem. and geological specimens. This was the tradition pioneered
By the time the second edition of the book appeared (1959), by Aristotle, given its modern guise by Georges Buffon in
it was full of energy-flow diagrams, with arrows pointing his Histoire Naturelle (1749–1789) and memorialized in the
famous Museum d’Histoire Naturelle in Paris
in all directions to emphasize the inter-relatedness of the 4 One of Aristotle’s most widely cited works is Historia
myriad factors within an ecosystem. Soon, Odum became Animalium, which is the text upon which many later works
convinced that it was the complex interactions within the in natural history were based. His student Theophrastus
wrote an Aristotelian botanical work, Historia Plantarum,
ecosystem that provided its stability, protecting it from which provided the botanical analog for later naturalists.
perturbations in much the same way as homeostasis in The best work on Aristotle as a biologist is Grene, M.
organisms regulates the physiology of those systems. (1963) A Portrait of Aristotle, University of Chicago Press
5 Roger, J. (1963) Les Sciences de la Vie dans la Pensee
Francaise du XVIII Siecle, Armond Colin
Ecology and environmentalism 6 Larson, J.L. (1994) Interpreting Nature, Johns Hopkins
There were other perturbations that soon attracted Odum’s University Press
7 Darwin, C. (1859) On the Origin of Species, John Murray
attention. Rachel Carson published Silent Spring in 1961, a 8 Haeckel, E. (1866) Generelle Morphologie der Organismen,
damning exposé of the pesticide industry in general and the G. Reimer
use of DDT in particular. President Kennedy convened the 9 Haeckel, E. (1876) History of Creation, Appleton
10 Drude, O. (1896) Deutschlands Pflanzengeographie (1896);
first presidential commission on the environment, a commis- Warming, E. (1896) Lehrbuch der Okologischen Pflanzen-
sion that almost immediately recommended the prohibition geographie eine Einfuhrung in die Kenntniss der Pflanzenvereine
of DDT. However, there were other issues within the en- 11 Tobey, R. (1981) Saving the Prairies: The Life Cycle of the
Founding School of American Plant Ecology, University of
vironment, ranging from suffocating air pollution in Los California Press
Angeles to fiery water pollution on the Cayuhuga River in 12 Cowles, H.C. (1899) The ecological relations of the
Cleveland. Odum soon saw his role not just as an ecological vegetation of the sand dunes of Lake Michigan. Bot. Gazette
27, 95–117; Cowles, H.C. (1899) The ecological relations of
researcher but as an advocate to preserve the earth’s fragile the vegetation of the sand dunes of Lake Michigan. Bot.
ecosystems. In the third edition of his text (1971), he added a Gazette 27, 167–202; Cowles, H.C. (1899) The ecological
chapter on the environment and conservation, also explaining relations of the vegetation of the sand dunes of Lake
Michigan. Bot. Gazette 27, 281–308; Cowles, H.C. (1899)
how ecological principles, such as the food pyramid, could be The ecological relations of the vegetation of the sand dunes
used to explain how non-biodegradable pesticides (e.g. DDT) of Lake Michigan. Bot. Gazette 27, 361–391
take their toll on the ecosystem by accumulating at the top of 13 Pound, R. and Clements, F.E. (1900) The Phytogeography
of Nebraska, Botanical Seminar; Clements, F.E. (1905)
the pyramid. Thus, ecology now claimed a new niche, that Research Methods in Ecology, University Publishing
of informing the populace about environmental issues. Company; Clements, F.E. (1916) Plant Succession,
Ironically, ecology’s contributions to the environmental Carnegie Institute of Washington
14 The rhetorical power of the words ‘laboratory’and ‘experiment’
crisis may have caused it to become much more popular with needs to be stressed within this context. Natural history’s
the public and to return it, at least in the eyes of the general traditional methods were often seen to be somewhat dated,
public, back to natural history. That is, the stress in ecology especially in comparison to the methods of experimentation,
a major component of physiology. Thus, the closer the
on the integrity of natural systems has led many to consider ecologists could come to experimental or laboratory-based
the word ‘ecology’to be synonymous with ‘environmental’, methods, the closer they imagined themselves to be to the
‘conservationist’or even ‘natural’. Certainly, ecology and its cutting-edge aspects of modern 20th-century biology.
15 Forbes, S.A. (1887) The lake as microcosm. Bull. South
professional practitioners are often greeted as advocates for Africa Peoria 77–87
the conservation or preservation of the natural world. And 16 Elton, C. (1927) Animal Ecology, Macmillan
this characterization is often correct, for many ecologists and 17 Tansley, A.G. (1935) The use and abuse of vegetational
concepts and terms. Ecology 16, 284–307
their professional organizations (e.g. the Ecological Society 18 G. Evelyn Hutchinson was enormously influential in the
of America) have been quick to criticize societal practices development of modeling in ecology, but the seminal article
that harm the environment and have also been among the was written by his student Raymond Lindeman. The paper
was published posthumously as Lindeman, R.L. (1942) The
forefront of citizens arguing for the need to set aside vast trophic–dynamic aspect of ecology. Ecology 23, 399–418
areas of the environment for study and for preservation. 19 Allee, W.C. et. al. (1949) Principles of Ecology, W.B. Saunders
However, in large part, in addition to scientific support, 20 Odum’s father was a sociologist who borrowed heavily from
the work of sociologists at Chicago, the same individuals
these positions are taken for the same esthetic reasons that who were so influential on the Chicago school of ecology.
lead non-ecologists to protect the natural world. That is, our In addition, his brother Howard studied with Hutchinson at
experiences in nature seem to have a salutory influence on Yale, providing Eugene Odum with direct access to the new
dynamical model of ecology. Odum, E.P. (1953)
us as humans. Ecologists are those fortunate enough to have Fundamentals of Ecology, W.B. Saunders
adopted a profession that keeps them closely attached to Note: See Slobodkin, L.B. and Slack, N. (1999) George Evelyn
their historical roots in natural history. Hutchinson: 20th-century ecologist. Endeavour 23, 24–30
62 Endeavour Vol. 24(2) 2000
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