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Symposium 19 Good Ideas at the Time: Historians Look at Ecology

2008; Ecological Society of America; Volume: 90; Issue: 1 Linguagem: Inglês

10.1890/0012-9623-90.1.142

2327-6096

Matthew K. Chew,

Environmental Philosophy and Ethics

Ecologists' working knowledge of our own disciplinary history often consists of the classic papers we were taught (and now teach), local institutional lore, and the recent literature we review for our own publications. The ESA's approaching centennial in 2015 provides an incentive to look deeper and reflect on the culture of ecological science in America. Why bother? In his introduction to the 2006 ESA symposium "What Makes an Ecological Icon?" Aaron Ellison suggested, "losing the historical context for our work, and the disappearance from contemporary literature of carefully garnered data and results, can lead to unnecessary repetition of research." Indeed, but the preface to Robert McIntosh's 1985 classic, The Background of Ecology: Concept and Theory, included the statement, "Ecologists should be free of what one ecologist called the "Tyranny of the Present," not simply because knowledge of past events is interesting, but because ignorance of the past makes for redundance [sic] at best and confusion at worst." McIntosh's unnamed ecologist echoed Ralph Waldo Emerson's 1836 essay "Nature," and Frankenstein author Mary Shelley's 1822 diary; both of them probably encountered the "Tyranny of the Present" in writings attributed to Marcus Tullius Cicero (106–40 BCE). And so it goes. The past does not assert itself; we must pursue it as it recedes, in order to recognize the resurgence of abandoned or discredited ideas under new names, and the extent to which both internecine politics, institutional personality, and individual charisma have determined what was heard, published, and cited. Such influences are not unique to ecology, but ecology is far from immune to them. I organized "Good Ideas at the Time" with the softest possible focus in order to accommodate investigators with a wide range of backgrounds and interests. All I demanded was a historical approach to an ecological subject. Although Mark Hineline could not attend, the remaining presentations still offered a sampler of the scholarship undertaken by members of organizations like the International Society for the History, Philosophy and Social Studies of Biology (ISHPSSB), the History of Science Society (HSS), the Philosophy of Science Association (PSA), and the Society for Social Studies of Science (4S). Ecology has attracted lesser followings in those societies than (e.g.) genetics or evolutionary and developmental biology, but there are a few of us at every meeting, and we are awash in interesting material. How many eyes does horror have? How many times will terror strike? They were born in that tragic time that Science made its great mistake…now from behind the shroud of night they come, a scuttling, shambling horde of creatures destroying all in their path. Or so the poster for the 1972 film Night of the Lepus warns the audience (Fig. 1). However, ironically, "Science's great mistake" was born not out of an insane desire for knowledge, but from the attempt to control "a plague of rabbits" without employing poisons. Thus, in "ecodoom" films the practice of science upsets nature's natural balances, and the application of more science to restore those balances often simply makes things worse. But what do these portrayals of "mad" science tell us about the relationship between science and nature? Promotional poster for the film Night of the Lepus (1972) For one thing these images reflect the gendered science/nature relationship: In Mel Brooks' parody Young Frankenstein, Frankenstein (Gene Wilder) and corpse are hoisted toward the lightning-filled sky while Frankenstein proclaims that he wants to "penetrate into the very womb of impervious nature herself." Mad science images are also employed in public debates about scientific research, its ethics, and resultant regulation. Finally, children's drawings reflect the stereotype. In addition to glasses and wild hair, the scientists (generally middle-aged, white males) work in laboratories plastered with "Top Secret" signs (reviewed in Frayling 2005:12–16). In its origins, modern science was, at least in some cases, defined to exclude areas of specific interest to women. A case in point is Ellen Swallow Richards, who wrote to Ernst Haeckel asking for permission to use his term "oekologie" (the study of organisms in their environment) for a new science to develop positive relationships between humans and their environment. She was criticized for engaging in applied work, rather than pure research (Breton 1998:59–60). Hrdy (1986:120) argued that "[…] the possibility that the empathy for other females subjectively felt by women researchers may have been instrumental in expanding the scope of sexual selection theory" to include research on female behavior. This history leads to continuing tension around the question of whether women would lead the scientific enterprise in new directions. However, filmmakers have been less reluctant to ascribe different traits to female scientists. In the first type of film, the female scientist is mad because she uses science to protect nature. In Carnosaur, Jane Tiptree (Dianne Ladd) genetically engineers a virus causing women to birth dinosaur eggs, thus destroying the human species because of humanity's destruction of nature. Tiptree, who is feminized in comparison to the other female characters, with manicured nails, plucked brows, and makeup, is positioned as a vengeful Mother Nature. In the second case, the female scientist avoids madness by changing her affinity from nature to humans through romance or maternity. In Kingdom of the Spiders, scientist Dianne Ashley (Tiffany Bolling) and veterinarian Robert "Rack" Hansen (William Shatner) fight human-eating tarantulas. Initially, Ashley is allied with nature: instead of reacting with disgust to the presence of a spider in her bedroom, she gently picks it up and strokes it before releasing it outside. Ultimately, she abandons her alliance with nature and her scientific role; she spends the last part of the film consoling Hansen's niece. In the final case, the female scientist is mad because she has no empathy for nature. In Humanoids from the Deep, Dr. Susan Drake (Ann Turkel) genetically engineers fast-growing salmon to replenish a depleted fishery, producing the humanoids that mate with human women. Although Drake helps the town defeat the humanoids, she returns to her lab to deliver a humanoid baby (product of rape) that kills its mother during its birth. There is the clear potential for further madness, since her scientific curiosity has released another humanoid. Thus, ecodoom films emphasize the tensions among science, nature and gender. And low-budget horror films are ideal examples, because as Adam Simon, director of Carnosaur, explains when comparing his film to Jurassic Park: "We could be smarter, in some ways, because when you make a $100 million movie, you're making a corporate product that has to please millions of people, so no matter how beautifully you do it, it has to be somewhat debased on the level of ideas. We could be more political than they could be. And we could be grosser" (Biodrowski 1993:23). James ("Jack") Justus followed Kasi's multimedia opening act with a look at the "Emergence of the Stability–Diversity–Complexity (SDC) Debate of Community Ecology, 1955–1975." The SDC debate has persisted as a central focus of theoretical ecology for half a century. The debate concerns the deceptively simple question of whether there is a relationship between the complexity and/or diversity of a biological community and its stability. Claims that there is a positive relationship have a long history in ecology. The "balance of nature" was a staple of the schools of natural philosophy from which biology emerged, long before the term "ecology" was even coined. Some early ecologists such as Fredrick Clements and A. J. Nicholson continued this tradition by attempting to derive the existence of a "natural balance" in biological populations from organismic metaphors and analogies with physical systems. With the possible exception of Lotka and Volterra, not until Robert MacArthur's first publication, while still a graduate student of G. E. Hutchinson in 1955, was the inchoate question initiated as a scientific debate with an interesting theoretical argument about food webs. From 1955 to the early 1970s, the prevailing view among ecologists was that diversity and/or complexity were important, perhaps principal causes of community stability. Robert May, a physicist turned mathematical ecologist, confounded this view using mathematical models of communities that seemed to confirm the opposite, that increased complexity jeopardizes stability (May 1974). Between 1955 and 1975 the SDC debate opened, a strong consensus emerged, and this consensus then dissolved (see Justus 2008). The praise May's work received for its mathematical rigor and the criticisms it received for its seeming biological irrelevance thrust the SDC debate into the ecological limelight; but subsequently, different analyses seem to support conflicting claims and indicate an underlying lack of conceptual clarity about ecological stability, diversity, and complexity. To understand what the SDC debate is about and the conceptual and methodological issues it raises, James began by analyzing the seminal works by MacArthur (1955) and Elton (1958) that helped transform poorly formulated questions about a "natural balance" to a scientifically precise hypothesis, and consolidated ecologists' opinions. His analysis revealed the formidable theoretical and empirical challenges of evaluating stability–diversity–complexity relationships. For example, MacArthur (1955) has been misinterpreted as demonstrating a positive relationship between a particular type of stability (i.e., resistance to change following perturbation) properties like the number of links in a food web.. Although MacArthur suggested some intuitive reasons to expect such a relationship, the portion of his analysis taken to demonstrate it actually presupposes it—a fact that MacArthur clearly understood. Charles Elton also believed there was a positive relationship between the stability of a community and its structural complexity. Elton's Oxford research was concerned with empirical and practical ecological projects, such as detailed biological surveys and improved rodent control methods. He was cautiously skeptical of the biological relevance of mathematical models of biological communities like those used by MacArthur. Elton (1958) focused instead on empirical evidence that seemed to show that more diverse communities were more resistant to invasion by exotic species than others, and experienced fewer and less severe population fluctuations than others. Although Elton (1958) and MacArthur (1955) are commonly cited as analyzing the same relationship between stability and diversity, their analyses presuppose slightly different stability concepts, given Elton's focus on fluctuations and invasion resistance. Moreover, the six types of evidence for a positive stability–diversity relationship Elton proposed have proved uncompelling. James turned next to the first experimental test of stability–diversity relationships using wild cabbage conducted by David Pimentel (1961) in fallow fields outside Ithaca, New York, even as Elton's book went to press. The limitations of Pimentel's study—for example, that diversity was measured as species richness or the difficulties of distinguishing the effects of different kinds of richness (plant vs. insect)—reveal the formidable empirical challenges involved in evaluating stability–diversity–complexity relationships. A point apparently not widely appreciated is that Pimentel (1961) appears to be the first ecologist to recognize that the statistical portfolio effect may explain why more species-rich biological communities seem to contain more constant abundances. Although it was probably the first experimental study of the SDC debate, Pimentel's empirical work received much less attention than MacArthur's theoretical analysis. This likely reflected the transformation of ecology into a more mathematical and theoretical discipline occurring at the time. Largely through the work of G. E. Hutchinson and his students (MacArthur most importantly), mathematical ecology became both more sophisticated and more prevalent. During the 1960s and 1970s, ecologists were increasingly concerned with formalizing and theoretically systematizing ecological concepts, and the SDC debate was extended by two important attempts to more precisely define its conceptions: Lewontin's (1969) analysis of the relationship between ecological and mathematical concepts of stability and Hurlbert's (1971) incisive critique of the concept of ecological diversity. Robert May's (1974) analysis of mathematical models of biological communities epitomized this approach, and it brought greater mathematical rigor and sophistication to the SDC debate. May's influential work upended the popular belief among ecologists that "diversity begets stability"— but probably should not have. May presented a variety of theoretical results, but the most important was an inverse relationship between stability (understood as local stability), species richness, and frequency and intensity of species interactions in randomly constructed linear models of biological communities. Although this stability concept differs significantly from those of MacArthur, Elton, and Pimentel, this theoretical result and others seemed to demonstrate that high species richness, and frequency and intensity of species interaction, precludes rather than enhances community stability. DeAngelis (1975) and Lawlor (1978) criticized May's work for being biologically implausible, but it nonetheless changed many ecologists' opinions about the SDC debate. This history reveals that different concepts of stability, diversity, and complexity were employed by different ecologists involve in the SDC debate. Given this conceptual diversity, it is unsurprising that the results they found often differed and were sometimes incompatible. The methodological limitations of Pimentel's study, the difficulty of interpreting and extrapolating from the theoretical work of MacArthur and May, and that the kinds of evidence Elton suggested in favor of a positive stability–diversity relationship ultimately proved inconclusive, also indicate the formidable challenges posed by the SDC and explain why the debate remains unresolved. I followed Jack Justus with an account of another, perhaps theoretically vaguer but very persistent and influential idea in Ecology and the De-natured World. A few weeks before the 93rd ESA Annual Meeting, several ecologists were quoted in The New York Times complaining that human encroachment was incrementally destroying ecological research sites (Nijhuis 2008). This occurrence dovetailed with a pair of questions I was already engaged with, one for ecology as a discipline (but which I could shed some light on) and one I thought I could answer outright. The first, for all of us to ponder and debate, is why do we, a priori and categorically, disqualify modern humans from being part of nature and susceptible to normal ecological analysis? Robert May sharpened the point for me during his conference keynote address by exclaiming "This is us, not some natural event!" I am but a recent addition to the centuries-long queue of scientists and philosophers pondering the nature of "naturalness," but until the status of Homo sapiens is normalized, ecological science has a problem: we pervade nature as we know it. Recent work (e.g., Foley et al. 2005, Halpern et al. 2008) illustrates just how little of the biosphere can be considered unaffected by human activity. Insisting that real ecological relationships only occur, and can only be properly studied, where humans have no discernable influence seems to be a drastic case of scientific self-marginalization. That introduces the second question: how long have ecologists (and more specifically, American ecologists) known that humans were such a pervasive influence? I answered that question using quotations and illustrations from Marsh (1864, 1874), Hooker (1867), Pound and Clements (1900), Clements (1905, 1916, 1936), Adams (1913), Shelford (1913, 1963), Shelford et al. (1921, 1926), Elton (1930, 1958), Clements and Shelford (1939), Tansley (1944), and Bates (1956) to show that by the time the science of ecology coalesced in the 1890s ecologists already knew that humans were global, pervasive ecological influences, and we never forgot; returning us to question one, now recast. Knowing that human ecological influence is pervasive, why do we insist on treating putatively "natural" study areas as if they were the only places we can study real ecology (Fig. 2)? Map showing North American natural areas identified during the early 1920s by the ESA Committee on the Preservation of Natural Conditions. (A composite of original figures 6 and 7 in the Naturalist's Guide to the Americas, 1926). My answer challenges the ESA membership to pause for a reality check. Many of the authors I quoted, back to Pound and Clements in 1900, admitted that they could not systematically comprehend the whole range of human influences or incorporate those effects into their hypotheses. Their solution, by design or default, was to ignore the technologist in the room and study ecology in remote or isolated places where human influence could (they presumed) be ignored. Somewhere along the line, and already detectable in the writings of Elton (1943) and Leopold (1948), avoiding confounding human influences metastasized into axiomatic misanthropy. It took a new skill set and much research and reflection before I grasped how fundamentally the assumption that human activity distorts normal phenomena had sculpted modern ecology. Carelessly equating "anthropogenic change" with "ruin" sours our hypotheses, observations, and interpretations of ecological phenomena. Every study conceived and justified and designed with reference to damage or distortion "begs" its own conclusions. Neither sincere regret nor righteous indignation about the state of the world can repair flawed logic, or produce from it good science. The overwhelming extent of human ecological influence may not seem like something to celebrate for biophiles with a "sense of place." I "get" that; most of my own favorite places are changing drastically, and in ways I wouldn't have hoped to witness. Nevertheless, when I remove the nostalgiatinted glasses provided by my mentors and ecology's icons, I find life under the régime de l'homme no less worth investigating, understanding, and even appreciating. The entirely independent "Concepts and Questions" article by Ellis and Ramankutty (2008) subsequently appearing in Frontiers in Ecology and the Environment suggests that I am not alone in this assessment. I promised an exclamation point about repeating history: during my talk, I mentioned a fact that always seems to bemuse ecologists: in July 1945, after much heated discussion among ESA officers, the whole membership voted to revoke Victor Shelford's longstanding, de facto authority as Chairman of its "Committee on the Preservation of Natural Conditions" to "take direct action designed to influence legislation" on behalf of the whole Society. Late that year, Shelford began the process of founding a splinter group called the Ecologists' Union. Many senior ecologists are aware of "what happened next," but during a pregnant pause, no one in the audience admitted knowing that an internecine, ESA squabble had thereby given birth to an offspring now called The Nature Conservancy. The "revelation" generated enough of a buzz that a day later, a young ecologist who had attended the meeting (but not Good Ideas at the Time) mentioned it to me as we waited to disembark from an airliner in Phoenix. But the real exclamation point is that Sara Tjossem revealed exactly the same fact in her presentation two years ago at the "Icons" symposium at our Memphis meeting, and it was mentioned in the ensuing Bulletin writeup (Ellison 2006). The past does not assert itself. We must pursue it as it recedes. I'm sorry to report that I have been unable to reestablish contact with presenter Victor Cassidy since the meeting. Rather than attempting to summarize Victor's presentation, which followed mine, I recommend visiting his web site 〈http://www.victorcassidy.com〉, which includes photos and a sample chapter from his book Henry Chandler Cowles: Pioneer Ecologist (Sigel Press, 2007). You'll get the idea. That completes the opening acts. The headliner was Frank N. Egerton, who is undoubtedly Ecology's current Dean of History; in 2007 the ESA acknowledged this by honoring him with a Distinguished Service Award. Frank is responsible for "Good Ideas at the Time" to the extent that he put me up to organizing it, offered helpful comments on drafts of the symposium proposal and contributed his presentation, "Homage to Frederic E. Clements: The History of Studies on Plant Succession." Clements seems to have been the first historian of an aspect of ecology, with a 24-page "General Historical Summary" to his huge monograph, Plant Succession 1916. His survey began with a study on peat bogs (1685), but subsequent research has pushed back the date of the earliest bog studies to the mid-1500s, which included observations on bogs flowing down hill. The earliest diagram having ecological relevance is probably one by John Honohane (1697) illustrating such a flow. Clements found many examples of observations on plant succession in the scientific literature that he summarized or quoted, often in his own translations. Students of plant succession included Georges Buffon (1742) and Carl Linnaeus (1749). A Genevan living in London, Jean Deluc, in the early 1800s, was probably first to use the term "succession" to describe changes in vegetation. The Danish naturalist, Japetus Steenstrup (Fig. 3) was the founder of paleoecology (1842). He studied cross-sections of ponds that filled with vegetation (Fig. 4) and realized that different layers of plant remains provided a successional history of the plants of the area. He published before the discovery of past ice ages, but Scandinavian investigators in the later 1800s used the ice age evidence to help understand plant succession. Swedish botanist Rutger Sernander may have been first to diagram the succession of species in an area (1894), almost two decades before diagrams were used to describe food chains and webs. However, once used for food chains and webs, they persisted much longer than did diagrams in plant ecology. Other ecologists indicated that plant successions tend toward a climax formation, but not all agreed with Clements' theory that there was only one climax for a region and that it was determined entirely by climate. Nineteenth century Danish naturalist Japetus Steenstrup, the founder of paleoecology. Steenstrup's pond paleostratigraphy, as represented by Clements in p 1916. Historians like Frank Egerton need no convincing that history is important. Ecologists may feel they are too busy making history rather than studying it; but among all the sciences ecology's collective past may be peculiarly important to its collective present. Very few early chapters are now wholly irrelevant to our ongoing investigations. We continue to dispute concepts as fundamental as our basic units of study, first because they are complex and take time to work out, but also because they are complicated both by our participation in the systems we study, and by our perceptual, cognitive, and emotional limitations as organisms. We ignore early ecology at our peril, not only because we may unnecessarily repeat prior research, but also because we are recognizably reacting to many of the same problems, in the same ways, as our forbears. We can still learn not only from their solutions, but from the way they experienced and framed the problems they solved.