История экологии: от Аристотеля до современности

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Ecology is a new science and considered as an important branch of biological science, having only become prominent during the second half of the 20th century. Ecological thought is derivative of established currents in philosophy, particularly from ethics and politics. Its history stems all the way back to the 4th century. One of the first ecologists whose writings survive may have been Aristotle or perhaps his student, Theophrastus, both of whom had interest in many species of animals and plants. Theophrastus described interrelationships between animals and their environment as early as the 4th century BC. Ecology developed substantially in the 18th and 19th century. It began with Carl Linnaeus and his work with the economy of nature. Soon after came Alexander von Humboldt and his work with botanical geography. Alexander von Humboldt and Karl Möbius then contributed with the notion of biocoenosis. Eugenius Warming's work with ecological plant geography led to the founding of ecology as a discipline. Charles Darwin's work also contributed to the science of ecology, and Darwin is often attributed with progressing the discipline more than anyone else in its young history. Ecological thought expanded even more in the early 20th century. Major contributions included: Eduard Suess’ and Vladimir Vernadsky's work with the biosphere, Arthur Tansley's ecosystem, Charles Elton's Animal Ecology, and Henry Cowles ecological succession. Ecology influenced the social sciences and humanities. Human ecology began in the early 20th century and it recognized humans as an ecological factor. Later James Lovelock advanced views on earth as a macro organism with the Gaia hypothesis. Conservation stemmed from the science of ecology. Important figures and movements include Shelford and the ESA, National Environmental Policy act, George Perkins Marsh, Theodore Roosevelt, Stephen A. Forbes, and post Dust Bowl conservation. Later in the 20th century world governments collaborated on man’s effects on the biosphere and Earth’s environment. The history of ecology is intertwined with the history of conservation and restoration efforts.

In the early Eighteenth century, preceding Carl Linnaeus, two rival schools of thought dominated the growing scientific discipline of ecology. First, Gilbert White a "parson naturalist" is attributed with developing and endorsing the view of Arcadian ecology. Arcadian ecology advocates for a "simple, humble life for man" and a harmonious relationship with humans and nature. In An Essay on the Principle of Population Malthus argues for the reining in of rising population through 2 checks: Positive and Preventive checks. The first raising death rates, the later lowers birthing rates. Malthus also brings forth the idea that the world population will move past the sustainable number of people. This form of thought still continues to influences debates on birth and marriage rates to this theory brought forth by Malthus. The essay had a major influence on Charles Darwin and helped him to theories his theory of Natural Selection. This struggle proposed by Malthusian thought not only influenced the ecological work of Charles Darwin, but helped bring about an economic theory of world of ecology.

It is often held that the roots of scientific ecology may be traced back to Darwin. This contention may look convincing at first glance inasmuch as On the Origin of Species is full of observations and proposed mechanisms that clearly fit within the boundaries of modern ecology (e. g. the cat to clover chain – an ecological cascade) and because the term ecology was coined in 1866 by a strong proponent of Darwinism, Ernst Haeckel. However, Darwin never used the word in his writings after this year, not even in his most "ecological" writings such as the foreword to the English edition of Hermann Müller's The Fertilization of Flowers (1883) or in his own treatise of earthworms and mull formation in forest soils (The formation of vegetable mould through the action of worms, 1881). Moreover, the pioneers founding ecology as a scientific discipline, such as Eugen Warming, A. F. W. Schimper, Gaston Bonnier, F. A. Forel, S. A. Forbes and Karl Möbius, made almost no reference to Darwin's ideas in their works. Others focused on the dominant themes at the beginning, concern with the relationship between organism morphology and physiology on one side and environment on the other, mainly abiotic environment, hence environmental selection. Darwin's concept of natural selection on the other hand focused primarily on competition. The mechanisms other than competition that he described, primarily the divergence of character which can reduce competition and his statement that "struggle" as he used it was metaphorical and thus included environmental selection, were given less emphasis in the Origin than competition. Despite most portrayals of Darwin conveying him as a non aggressive recluse who let others fight his battles, Darwin remained all his life a man nearly obsessed with the ideas of competition, struggle and conquest – with all forms of human contact as confrontation. Although there is nothing incorrect in the details presented in the paragraph above, the fact that Darwinism used a particularly ecological view of adaptation and Haeckel's use and definitions of the term were steeped in Darwinism should not be ignored. According to ecologist and historian Robert P. McIntosh, "the relationship of ecology to Darwinian evolution is explicit in the title of the work in which ecology first appeared." A more elaborate definition by Haeckel in 1870 is translated on the frontispiece of the influential ecology text known as 'Great Apes' as " ecology is the study of all those complex interrelations referred to by Darwin as the conditions of the struggle for existence." Elton influenced by Victor Shelford's Animal Communities in Temperate America began his research on animal ecology as an assistant to his colleague, Julian Huxley, on an ecological survey of the fauna in Spitsbergen in 1921. Elton's most famous studies were conducted during his time as a biological consultant to the Hudson Bay Company to help understand the fluctuations in the company's fur harvests. Elton studied the population fluctuations and dynamics of snowshoe hare, Canadian lynx, and other mammals of the region. Elton is also considered the first to coin the terms, food chain and food cycle in his famous book Animal Ecology. Elton is also attributed with contributing to disciplines of: invasion ecology, community ecology, and wildlife disease ecology.

George "G" Evelyn Hutchinson was a 20th century ecologist who is commonly recognized as the "Father of Modern Ecology". Hutchinson is of English descent but spent most of professional career studying in New Haven, Connecticut at Yale University. Throughout his career, over six decades, Hutchinson contributed to the sciences of limnology, entomology, genetics, biogeochemistry, mathematical theory of population dynamics and many more. Hutchinson is also attributed as being the first to infuse science with theory within the discipline of ecology. Hutchinson was also one of the first credited with combining ecology with mathematics. Another major contribution of Hutchinson was his development of the current definition of an organism's "niche" – as he recognized the role of an organism within its community. Finally, along with his great impact within the discipline of ecology throughout his professional years, Hutchinson also left a lasting impact in ecology through his many students he inspired. Foremost among them were Robert H. MacArthur, who received his PhD under Hutchinson, and Raymond L. Lindeman, who finished his PhD dissertation during a fellowship under him. MacArthur became the leader of theoretical ecology and, with E. O. Wilson, developed island biography theory. Raymond Lindeman was instrumental in the development of modern ecosystem science.

"What is ecology?” was a question that was asked in almost every decade of the 20th century. Unfortunately, the answer most often was that it was mainly a point of view to be used in other areas of biology and also "soft", like sociology, for example, rather than "hard", like physics. Although autecology (essentially physiological ecology) could progress through the typical scientific method of observation and hypothesis testing, synecology (the study of animal and plant communities) and genecology (evolutionary ecology), for which experimentation was as limited as it was for, say, geology, continued with much the same inductive gathering of data as did natural history studies. Most often, patterns, present and historical, were used to develop theories having explanatory power, but which had little actual data in support. Darwin's theory, as much as it is a foundation of modern biology, is a prime example. G. E. Hutchinson, identified above as the "father of modern ecology", through his influence raised the status of much of ecology to that of a rigorous science. By shepherding of Raymond Lindemann's work on the trophic dynamic concept of ecosystems through the publication process after Lindemann's untimely death, Hutchinson set the groundwork for what became modern ecosystem science. With his two famous papers in the late1950s, "Closing remarks", and "Homage to Santa Rosalia", as they are now known, Hutchinson launched the theoretical ecology which Robert MacArthur championed. Ecosystem science became rapidly and sensibly associated with the "Big Science"—and obviously "hard" science—of atomic testing and nuclear energy. It was brought in by Stanley Auerbach, who established the Environmental Sciences Division at Oak Ridge National Laboratory, to trace the routes of radionuclides through the environment, and by the Odum brothers, Howard and Eugene, much of whose early work was supported by the Atomic Energy Commission. Eugene Odum's textbook, Fundamentals of Ecology, has become something of a bible today. When, in the 1960s, the International Biological Program (IBP) took on an ecosystem character, ecology, with its foundation in systems science, forever entered the realm of Big Science, with projects having large scopes and big budgets. Just two years after the publication of Silent Spring in 1962, ecosystem ecology was trumpeted as THE science of the environment in a series of articles in a special edition of BioScience. Theoretical ecology took a different path to established its legitimacy, especially at eastern universities and certain West Coast campuses. It was the path of Robert MacArthur, who used simple mathematics in his "Three Influential Papers, also published in the late 1950s, on population and community ecology. Although the simple equations of theoretical ecology at the time, were unsupported by data, they still were still deemed to be "heuristic". They were resisted by a number of traditional ecologists, however, whose complaints of "intellectual censorship" of studies that did not fit into the hypothetico deductive structure of the new ecology might be seen as evidence of the stature to which the Hutchinson MacArthur approach had risen by the 1970s. MacArthur's untimely death in 1972 was also about the time that postmodernism and the "Science Wars" came to ecology. The names of Kuhn, Wittgenstein, Popper, Lakatos, and Feyerbrend began to enter into arguments in the ecological literature. Darwin's theory of adaptation through natural selection was accused of being tautological. Questions were raised over whether ecosystems were cybernetic and whether ecosystem theory was of any use in application to environmental management. Most vituperative of all was the debate that arose over MacArthur style ecology. Matters came to a head after a symposium organized by acolytes of MacArthur in homage to him and a second symposium organized by what was disparagingly called the "Tallahassee Mafia" at Wakulla Springs in Florida. The homage volume, published in 1975, had an extensive chapter written by Jared Diamond, who at the time taught kidney physiology at the UCLA School of Medicine, that presented a series of "assembly rules" to explain the patterns of bird species found on island archipelagos, such as Darwin's famous finches on the Galapagos Islands. The Wakulla conference was organized by a group of dissenters led by Daniel Simberloff and Donald Strong, Jr., who were described by David Quammen in his book as arguing that those patterns "might be nothing more than the faces we see in the moon, in clouds, in Rorschach inkblots". Their point was that Diamond's work (and that of others) did not fall within the criterion of falsifiability, laid down for science by the philosopher, Karl Popper. A reviewer of the exchanges between the two camps in an issue of Synthese found "images of hand to hand combat or a bar room brawl" coming to mind. The Florida State group suggested a method that they developed, that of "null" models, to be used much in the way that all scientists use null hypotheses to verify that their results might not have been obtained merely by chance. It was most sharply rebuked by Diamond and Michel Gilpin in the symposium volume and Jonathan Roughgarden in the American Naturalist. There was a parallel controversy adding heat to above that became known in conservation circles as SLOSS (Single Large or Several Small reserves). Diamond had also proposed that, according to the theory of island geography developed by MacArthur and E. O. Wilson, nature preserves should be designed to be as large as possible and maintained as a unified entity. Even cutting a road through a natural area, in Diamond's interpretation of MacArthur and Wilson's theory, would lead to the loss of species, due to the smaller areas of the remaining pieces. Simberloff, meanwhile, who had defaunated mangrove islands off the Florida coast in his award winning experimental study under E. O. Wilson and tested the fit of the species area curve of island biogeography theory to the fauna that returned, had gathered data that showed quite the opposite: that many smaller fragments together sometimes held more species that the original whole. It led to considerable vituperation on the pages of Science.

+ A list of founders, innovators and their significant contributions to ecology, from Romanticism onward. Notable figure Lifespan Major contribution & citation Antonie van Leeuwenhoek 1632–1723 First to develop concept of food chains Carl Linnaeus 1707–1778 Influential naturalist, inventor of science on the economy of nature Alexander Humboldt 1769–1859 First to describe ecological gradient of latitudinal biodiversity increase toward the tropics in 1807 Charles Darwin 1809–1882 Founder of the hypothesis of evolution by means of natural selection, founder of ecological studies of soils Elizabeth Catherine Thomas Carne 1817 1873 Geologist, mineralogist and philosopher who observed rural vs urban living, spatially and culturally, finding in country living the best attack on suffocating class divides, healthier living, and best access to natural education. Herbert Spencer 1820–1903 Early founder of social ecology, coined the phrase 'survival of the fittest' Karl Möbius 1825–1908 First to develop concept of ecological community, biocenosis, or living community Ernst Haeckel 1834–1919 Invented the term ecology, popularized research links between ecology and evolution Victor Hensen 1835–1924 Invented term plankton, developed quantitative and statistical measures of productivity in the seas Eugenius Warming 1841–1924 Early founder of Ecological Plant Geography Stephen Forbes 1844–1930 Early founder of entomology and ecological concepts in 1887 Vito Volterra 1860–1940 Independently pioneered mathematical populations models around the same time as Alfred J. Lotka. Vladimir Vernadsky 1869–1939 Founded the biosphere concept Henry C. Cowles 1869–1939 Pioneering studies and conceptual development in studies of ecological successionJan Christiaan Smuts 1870–1950 Coined the term holism in a 1926 book Holism and Evolution. Arthur G. Tansley 1871–1955 First to coin the term ecosystem in 1936 and notable researcher Charles Christopher Adams 1873–1955 Animal ecologist, biogeographer, author of first American book on animal ecology in 1913, founded ecological energetics Friedrich Ratzel 1844–1904 German geographer who first coined the term biogeography in 1891. Frederic Clements 1874–1945 Authored the first influential American ecology book in 1905 Victor Ernest Shelford 1877–1968 Founded physiological ecology, pioneered food web and biome concepts, founded The Nature Conservancy Alfred J. Lotka 1880–1949 First to pioneer mathematical populations models explaining trophic (predator prey) interactions using logistic equation Henry Gleason 1882–1975 Early ecology pioneer, quantitative theorist, author, and founder of the individualistic concept of ecology Charles S. Elton 1900–1991 'Father' of animal ecology, pioneered food web & niche concepts and authored influential Animal Ecology text G. Evelyn Hutchinson 1903–1991 Limnologist and conceptually advanced the niche concept Eugene P. Odum 1913–2002 Co founder of ecosystem ecology and ecological thermodynamic concepts Howard T. Odum 1924–2002 Co founder of ecosystem ecology and ecological thermodynamic concepts Robert MacArthur 1930–1972 Co founder on Theory of Island Biogeography and innovator of ecological statistical methods

Human ecology began in the 1920s, through the study of changes in vegetation succession in the city of Chicago. It became a distinct field of study in the 1970s. This marked the first recognition that humans, who had colonized all of the Earth's continents, were a major ecological factor. Humans greatly modify the environment through the development of the habitat (in particular urban planning), by intensive exploitation activities such as logging and fishing, and as side effects of agriculture, mining, and industry. Besides ecology and biology, this discipline involved many other natural and social sciences, such as anthropology and ethnology, economics, demography, architecture and urban planning, medicine and psychology, and many more. The development of human ecology led to the increasing role of ecological science in the design and management of cities. In recent years human ecology has been a topic that has interested organizational researchers. Hannan and Freeman (Population Ecology of Organizations (1977), American Journal of Sociology) argue that organizations do not only adapt to an environment. Instead it is also the environment that selects or rejects populations of organizations. In any given environment (in equilibrium) there will only be one form of organization (isomorphism). Organizational ecology has been a prominent theory in accounting for diversities of organizations and their changing composition over time.

The Gaia theory, proposed by James Lovelock, in his work Gaia: A New Look at Life on Earth, advanced the view that the Earth should be regarded as a single living macro organism. In particular, it argued that the ensemble of living organisms has jointly evolved an ability to control the global environment – by influencing major physical parameters as the composition of the atmosphere, the evaporation rate, the chemistry of soils and oceans – so as to maintain conditions favorable to life. The idea has been supported by Lynn Margulis who extended her endosymbiotic theory which suggests that cell organelles originated from free living organisms to the idea that individual organisms of many species could be considered as symbionts within a larger metaphorical "super organism". This vision was largely a sign of the times, in particular the growing perception after the Second World War that human activities such as nuclear energy, industrialization, pollution, and overexploitation of natural resources, fueled by exponential population growth, were threatening to create catastrophes on a planetary scale, and has influenced many in the environmental movement since then.

Environmentalists and other conservationists have used ecology and other sciences (e. g., climatology) to support their advocacy positions. Environmentalist views are often controversial for political or economic reasons. As a result, some scientific work in ecology directly influences policy and political debate; these in turn often direct ecological research and inquiry. The history of ecology, however, should not be conflated with that of environmental thought. Ecology as a modern science traces only from Darwin's publication of Origin of Species and Haeckel's subsequent naming of the science needed to study Darwin's theory. Awareness of humankind's effect on its environment has been traced to Gilbert White in 18th century Selborne, England. Raymond Pearl's population studies are a case in point. Ecology in subject matter and techniques grew out of studies by botanists and plant geographers in the late 19th and early 20th centuries that paradoxically lacked Darwinian evolutionary perspectives. Until Mendel's studies with peas were rediscovered and melded into the Modern Synthesis, Darwinism suffered in credibility. Many early plant ecologists had a Lamarckian view of inheritance, as did Darwin, at times. Ecological studies of animals and plants, preferably live and in the field, continued apace however.

When the Ecological Society of America (ESA) was chartered in 1915, it already had a conservation perspective. Victor E. Shelford, a leader in the society's formation, had as one of its goals the preservation of the natural areas that were then the objects of study by ecologists, but were in danger of being degraded by human incursion. Human ecology had also been a visible part of the ESA at its inception, as evident by publications such as: "The Control of Pneumonia and Influenza by the Weather," "An Overlook of the Relations of Dust to Humanity," "The Ecological Relations of the Polar Eskimo," and "City Street Dust and Infectious Diseases," in early pages of Ecology and Ecological Monographs. The ESA's second president, Ellsworth Huntington, was a human ecologist. Stephen Forbes, another early president, called for "humanizing" ecology in 1921, since man was clearly the dominant species on the Earth. This auspicious start actually was the first of a series of fitful progressions and reversions by the new science with regard to conservation. Human ecology necessarily focused on man influenced environments and their practical problems. Ecologists in general, however, were trying to establish ecology as a basic science, one with enough prestige to make inroads into Ivy League faculties. Disturbed environments, it was thought, would not reveal nature's secrets. Interest in the environment created by the American Dust Bowl produced a flurry of calls in 1935 for ecology to take a look at practical issues. Pioneering ecologist C. C. Adams wanted to return human ecology to the science. Frederic E. Clements, the dominant plant ecologist of the day, reviewed land use issues leading to the Dust Bowl in terms of his ideas on plant succession and climax. Paul Sears reached a wide audience with his book, Deserts on the March. World War II, perhaps, caused the issue to be put aside. The tension between pure ecology, seeking to understand and explain, and applied ecology, seeking to describe and repair, came to a head after World War II. Adams again tried to push the ESA into applied areas by having it raise an endowment to promote ecology. He predicted that "a great expansion of ecology" was imminent "because of its integrating tendency." Ecologists, however, were sensitive to the perception that ecology was still not considered a rigorous, quantitative science. Those who pushed for applied studies and active involvement in conservation were once more discreetly rebuffed. Human ecology became subsumed by sociology. It was sociologist Lewis Mumford who brought the ideas of George Perkins Marsh to modern attention in the 1955 conference, "Man’s Role in Changing the Face of the Earth." That prestigious conclave was dominated by social scientists. At it, ecology was accused of "lacking experimental methods" and neglecting "man as an ecological agent." One participant dismissed ecology as "archaic and sterile." Within the ESA, a frustrated Shelford started the Ecologists' Union when his Committee on Preservation of Natural Conditions ceased to function due to the political infighting over the ESA stance on conservation. Ecosystem science, both basic and applied, began to compete with theoretical ecology (then called evolutionary ecology and also mathematical ecology). Eugene Odum, who published a very popular ecology textbook in 1953, became the champion of the ecosystem. In his publications, Odum called for ecology to have an ecosystem and applied focus. The second event was the publication of Silent Spring. Rachel Carson's book brought ecology as a word and concept to the public. Her influence was instant. A study committee, prodded by the publication of the book, reported to the ESA that their science was not ready to take on the responsibility being given to it. Carson's concept of ecology was very much that of Gene Odum. As a result, ecosystem science dominated the International Biological Program of the 1960s and 1970s, bringing both money and prestige to ecology. Silent Spring was also the impetus for the environmental protection programs that were started in the Kennedy and Johnson administrations and passed into law just before the first Earth Day. Ecologists' input was welcomed. Former ESA President Stanley Cain, for example, was appointed an Assistant Secretary in the Department of the Interior. The environmental assessment requirement of the 1969 National Environmental Policy Act (NEPA), "legitimized ecology," in the words of one environmental lawyer. An ESA President called it "an ecological 'Magna Carta.'" A prominent Canadian ecologist declared it a "boondoggle." NEPA and similar state statutes, if nothing else, provided much employment for ecologists. Therein was the issue. Neither ecology nor ecologists were ready for the task. Not enough ecologists were available to work on impact assessment, outside of the DOE laboratories, leading to the rise of "instant ecologists," having dubious credentials and capabilities. Calls began to arise for the professionalization of ecology. Maverick scientist Frank Egler, in particular, devoted his sharp prose to the task. Again, a schism arose between basic and applied scientists in the ESA, this time exacerbated by the question of environmental advocacy. The controversy, whose history has yet to receive adequate treatment, lasted through the 1970s and 1980s, ending with a voluntary certification process by the ESA, along with lobbying arm in Washington. Post Earth Day, besides questions of advocacy and professionalism, ecology also had to deal with questions having to do with its basic principles. Many of the theoretical principles and methods of both ecosystem science and evolutionary ecology began to show little value in environmental analysis and assessment. Ecologist, in general, started to question the methods and logic of their science under the pressure of its new notoriety. Meanwhile, personnel with government agencies and environmental advocacy groups were accused of religiously applying dubious principles in their conservation work. Management of endangered Spotted Owl populations brought the controversy to a head. Conservation for ecologists created travails paralleling those nuclear power gave former Manhattan Project scientists. In each case, science had to be reconciled with individual politics, religious beliefs, and worldviews, a difficult process. Some ecologists managed to keep their science separate from their advocacy; others unrepentantly became avowed environmentalists.

Theodore Roosevelt was interested in nature from a young age. He carried his passion for nature into his political policies. Roosevelt felt it was necessary to preserve the resources of the nation and its environment. In 1902 he created the federal reclamation service, which reclaimed land for agriculture. He also created the Bureau of Forestry. This organization, headed by Gifford Pinchot, was formed to manage and maintain the nations timberlands. Roosevelt signed the Act for the Preservation of American Antiquities in 1906. This act allowed for him to "declare by public proclamation historic landmarks, historic and prehistoric structures, and other objects of historic and scientific interest that are situated upon lands owned or controlled by the Government of the United States to be national monuments." Under this act he created up to 18 national monuments. During his presidency, Roosevelt established 51 Federal Bird Reservations, 4 National Game Preserves, 150 National Forests, and 5 National Parks. Overall he protected over 200 million acres of land.

Ecology became a central part of the World's politics as early as 1971, UNESCO launched a research program called Man and Biosphere, with the objective of increasing knowledge about the mutual relationship between humans and nature. A few years later it defined the concept of Biosphere Reserve. In 1972, the United Nations held the first international Conference on the Human Environment in Stockholm, prepared by Rene Dubos and other experts. This conference was the origin of the phrase "Think Globally, Act Locally". The next major events in ecology were the development of the concept of biosphere and the appearance of terms "biological diversity"—or now more commonly biodiversity—in the 1980s. These terms were developed during the Earth Summit in Rio de Janeiro in 1992, where the concept of the biosphere was recognized by the major international organizations, and risks associated with reductions in biodiversity were publicly acknowledged. Then, in 1997, the dangers the biosphere was facing were recognized all over the world at the conference leading to the Kyoto Protocol. In particular, this conference highlighted the increasing dangers of the greenhouse effect – related to the increasing concentration of greenhouse gases in the atmosphere, leading to global changes in climate. In Kyoto, most of the world's nations recognized the importance of looking at ecology from a global point of view, on a worldwide scale, and to take into account the impact of humans on the Earth's environment.