Stephen Jay Gould  

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Stephen Jay Gould (September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was also one of the most influential and widely read authors of popular science of his generation. Gould spent most of his career teaching at Harvard University and working at the American Museum of Natural History in New York. In 1996, Gould was hired as the Vincent Astor Visiting Research Professor of Biology at New York University, where he divided his time teaching there and at Harvard.

Gould's most significant contribution to evolutionary biology was the theory of punctuated equilibrium, which he developed with Niles Eldredge in 1972. The theory proposes that most evolution is characterized by long periods of evolutionary stability, which is infrequently punctuated by swift periods of branching speciation. The theory was contrasted against phyletic gradualism, the popular idea that evolutionary change is marked by a pattern of smooth and continuous change in the fossil record.

Most of Gould's empirical research was based on the land snail genera Poecilozonites and Cerion. He also made important contributions to evolutionary developmental biology, receiving broad professional recognition for his book Ontogeny and Phylogeny. In evolutionary theory he opposed strict selectionism, sociobiology as applied to humans, and evolutionary psychology. He campaigned against creationism and proposed that science and religion should be considered two distinct fields (or "non-overlapping magisteria") whose authorities do not overlap.

Gould was known by the general public mainly for his 300 popular essays in Natural History magazine, and his numerous books written for both the specialist and non-specialist. In April 2000, the US Library of Congress named him a "Living Legend".

Contents

Scientific career

Gould began his higher education at Antioch College, graduating with a double major in geology and philosophy in 1963. During this time, he also studied at the University of Leeds in the United Kingdom. After completing graduate work at Columbia University in 1967 under the guidance of Norman Newell, he was immediately hired by Harvard University where he worked until the end of his life (1967–2002). In 1973, Harvard promoted him to professor of geology and curator of invertebrate paleontology at the institution's Museum of Comparative Zoology.

In 1982 Harvard awarded him the title of Alexander Agassiz Professor of Zoology. The following year, 1983, he was awarded a fellowship at the American Association for the Advancement of Science, where he later served as president (1999–2001). The AAAS news release cited his "numerous contributions to both scientific progress and the public understanding of science." He also served as president of the Paleontological Society (1985–1986) and of the Society for the Study of Evolution (1990–1991).

In 1989 Gould was elected into the body of the National Academy of Sciences. Through 1996–2002 Gould was Vincent Astor Visiting Research Professor of Biology at New York University. In 2001, the American Humanist Association named him the Humanist of the Year for his lifetime of work. In 2008, he was posthumously awarded the Darwin-Wallace Medal, along with 12 other recipients. (Until 2008, this medal had been awarded every 50 years by the Linnean Society of London.

Punctuated equilibrium

Early in his career, Gould and his colleague Niles Eldredge developed the theory of punctuated equilibrium, which describes the rate of speciation in the fossil record as occurring relatively rapidly, which then alternates to a longer period of evolutionary stability. It was Gould who coined the term "punctuated equilibria" though the theory was originally presented by Eldredge in his doctoral dissertation on Devonian trilobites and his article published the previous year on allopatric speciation.

According to Gould, punctuated equilibrium revised a key pillar "in the central logic of Darwinian theory." Some evolutionary biologists have argued that while punctuated equilibrium was "of great interest to biology generally," it merely modified neo-Darwinism in a manner that was fully compatible with what had been known before.

Other biologists emphasize the theoretical novelty of punctuated equilibrium, and argued that evolutionary stasis had been "unexpected by most evolutionary biologists" and "had a major impact on paleontology and evolutionary biology."

Comparisons were made to George Gaylord Simpson's work in Tempo and Mode in Evolution (1941). However Simpson describes the paleontological record as being characterized by predominantly gradual change (which he termed horotely), though he also documented examples of slow (bradytely), and rapid (tachytely) rates of evolution. Punctuated equilibrium and phyletic gradualism are not mutually exclusive, and examples of each have been documented in different lineages. The debate between these two models is often misunderstood by non-scientists, and according to Richard Dawkins has been oversold by the media.

Some critics jokingly referred to the theory of punctuated equilibrium as "evolution by jerks", which prompted Gould to describe phyletic gradualism as "evolution by creeps."

Evolutionary developmental biology

Gould made significant contributions to evolutionary developmental biology, especially in his work Ontogeny and Phylogeny. In this book he emphasized the process of heterochrony, which encompasses two distinct processes: neoteny and terminal additions. Neoteny is the process where ontogeny is slowed down and the organism does not reach the end of its development. Terminal addition is the process by which an organism adds to its development by speeding and shortening earlier stages in the developmental process. Gould's influence in the field of evolutionary developmental biology continues to be seen today in areas such as the evolution of feathers.

Selectionism and sociobiology

Gould was a champion of biological constraints, internal limitations upon developmental pathways, as well as other non-selectionist forces in evolution. Rather than direct adaptations, he considered many higher functions of the human brain to be the unintended side consequence of natural selection. To describe such co-opted features, he coined the term exaptation with paleontologist Elisabeth Vrba. Gould believed this feature of human mentality undermines an essential premise of human sociobiology and evolutionary psychology.Template:Citation needed

Against Sociobiology

In 1975, Gould's Harvard colleague E. O. Wilson introduced his analysis of animal behavior (including human behavior) based on a sociobiological framework that suggested that many social behaviors have a strong evolutionary basis. In response, Gould, Richard Lewontin, and others from the Boston area wrote the subsequently well-referenced letter to The New York Review of Books entitled, "Against 'Sociobiology'". This open letter criticized Wilson's notion of a "deterministic view of human society and human action."

But Gould did not rule out sociobiological explanations for many aspects of animal behavior, and later wrote: "Sociobiologists have broadened their range of selective stories by invoking concepts of inclusive fitness and kin selection to solve (successfully I think) the vexatious problem of altruism—previously the greatest stumbling block to a Darwinian theory of social behavior... Here sociobiology has had and will continue to have success. And here I wish it well. For it represents an extension of basic Darwinism to a realm where it should apply."

Spandrels and the Panglossian paradigm

With Richard Lewontin, Gould wrote an influential 1979 paper entitled, "The Spandrels of San Marco and the Panglossian Paradigm", which introduced the architectural term "spandrel" into evolutionary biology. In architecture, a spandrel is a triangular space which exists over the haunches of an arch. Spandrels—more often called pendentives in this context—are found particularly in classical architecture, especially Byzantine and Renaissance churches.Template:Citation needed

When visiting Venice in 1978, Gould noted that the spandrels of the San Marco cathedral, while quite beautiful, were not spaces planned by the architect. Rather the spaces arise as "necessary architectural byproducts of mounting a dome on rounded arches." Gould and Lewontin thus defined "spandrels" in the evolutionary biology context to mean any biological feature of an organism that arises as a necessary side consequence of other features, which is not directly selected for by natural selection. Proposed examples include the "masculinized genitalia in female hyenas, exaptive use of an umbilicus as a brooding chamber by snails, the shoulder hump of the giant Irish deer, and several key features of human mentality."

In Voltaire's Candide, Dr. Pangloss is portrayed as a clueless scholar who, despite the evidence, insists that "all is for the best in this best of all possible worlds". Gould and Lewontin asserted that it is Panglossian for evolutionary biologists to view all traits as atomized things that had been naturally selected for, and criticised biologists for not granting theoretical space to other causes, such as phyletic and developmental constraints. The relative frequency of spandrels, so defined, versus adaptive features in nature, remains a controversial topic in evolutionary biology.

An illustrative example of Gould's approach can be found in Elisabeth Lloyd's case study suggesting that the female orgasm is a by-product of shared developmental pathways. Gould also wrote on this topic in his essay "Male Nipples and Clitoral Ripples," prompted by Lloyd's earlier work.

Gould was criticized by philosopher Dan Dennett for using the term spandrel instead of pendentive, a spandrel that curves across a right angle to support a dome. Robert Mark, a professor of civil engineering at Princeton, offered his expertise in the pages of American Scientist, noting that these definitions are often misunderstood in architectural theory. Mark concluded, "Gould and Lewontin's misapplication of the term spandrel for pendentive perhaps implies a wider latitude of design choice than they intended for their analogy. But Dennett's critique of the architectural basis of the analogy goes even further astray because he slights the technical rationale of the architectural elements in question."

Evolutionary progress

Gould favored the argument that evolution has no inherent drive towards long-term "progress". Uncritical commentaries often portray evolution as a ladder of progress, leading towards bigger, faster, and smarter organisms, the assumption being that evolution is somehow driving organisms to get more complex and ultimately more like humankind. Gould argued that evolution's drive was not towards complexity, but towards diversification. Because life is constrained to begin with a simple starting point (like bacteria), any diversity resulting from this start, by random walk, will have a skewed distribution and therefore be perceived to move in the direction of higher complexity. But life, Gould argued, can also easily adapt towards simplification, as is often the case with parasites.

In a review of Full House, Richard Dawkins approved of Gould's general argument, but suggested that he saw evidence of a "tendency for lineages to improve cumulatively their adaptive fit to their particular way of life, by increasing the numbers of features which combine together in adaptive complexes. ... By this definition, adaptive evolution is not just incidentally progressive, it is deeply, dyed-in-the-wool, indispensably progressive."

Cladistics

Gould never embraced cladistics as a method of investigating evolutionary lineages and process, possibly because he was concerned that such investigations would lead to neglect of the details in historical biology, which he considered all-important. In the early 1990s this led him into a debate with Derek Briggs, who had begun to apply quantitative cladistic techniques to the Burgess Shale fossils, about the methods to be used in interpreting these fossils. Around this time cladistics rapidly became the dominant method of classification in evolutionary biology. Inexpensive but increasingly powerful personal computers made it possible to process large quantities of data about organisms and their characteristics. Around the same time the development of effective polymerase chain reaction techniques made it possible to apply cladistic methods of analysis to biochemical and genetic features as well.

Technical work on land snails

[[File:Cerion_watlingense_land_snail_shells_(modern;_northeastern_San_Salvador_Island,_eastern_Bahamas)_(15043406837).jpg|thumb|right|305px|Cerion shells from San Salvador Island, Bahamas.]] Most of Gould's empirical research pertained to land snails. He focused his early work on the Bermudian genus Poecilozonites, while his later work concentrated on the West Indian genus Cerion. According to Gould "Cerion is the land snail of maximal diversity in form throughout the entire world. There are 600 described species of this single genus. In fact, they're not really species, they all interbreed, but the names exist to express a real phenomenon which is this incredible morphological diversity. Some are shaped like golf balls, some are shaped like pencils. ... Now my main subject is the evolution of form, and the problem of how it is that you can get this diversity amid so little genetic difference, so far as we can tell, is a very interesting one. And if we could solve this we'd learn something general about the evolution of form."

Given CerionTemplate:'s extensive geographic diversity, Gould later lamented that if Christopher Columbus had only catalogued a single Cerion it would have ended the scholarly debate about which island Columbus had first set foot on in America.

Influence

Gould is one of the most frequently cited scientists in the field of evolutionary theory. His 1979 "spandrels" paper has been cited more than 5,000 times. In Paleobiology—the flagship journal of his own speciality—only Charles Darwin and George Gaylord Simpson have been cited more often. Gould was also a considerably respected historian of science. Historian Ronald Numbers has been quoted as saying: "I can't say much about Gould's strengths as a scientist, but for a long time I've regarded him as the second most influential historian of science (next to Thomas Kuhn)."

In a survey conducted in 2013 and 2014 among international experts on intelligence, Gould was the lowest rated among the important intelligence researchers by all criteria (quality and correctness; innovativeness and development of new ideas; impact in contributions and importance of oeuvre).

The Structure of Evolutionary Theory

Shortly before his death, Gould published The Structure of Evolutionary Theory (2002), a long treatise recapitulating his version of modern evolutionary theory. In an interview for the Dutch TV series Of Beauty and Consolation Gould remarked, "In a couple of years I will be able to gather in one volume my view of how evolution works. It is to me a great consolation because it represents the putting together of a lifetime of thinking into one source. That book will never be particularly widely read. It's going to be far too long, and it's only for a few thousand professionals—very different from my popular science writings—but it is of greater consolation to me because it is a chance to put into one place a whole way of thinking about evolution that I've struggled with all my life."

As a public figure

Gould became widely known through his popular essays on evolution in the Natural History magazine. His essays were published in a series entitled This View of Life (a phrase from the concluding paragraph of Charles Darwin's Origin of Species) from January 1974 to January 2001, amounting to a continuous publication of 300 essays. Many of his essays were reprinted in collected volumes that became bestselling books such as Ever Since Darwin and The Panda's Thumb, Hen's Teeth and Horse's Toes, and The Flamingo's Smile.

A passionate advocate of evolutionary theory, Gould wrote prolifically on the subject, trying to communicate his understanding of contemporary evolutionary biology to a wide audience. A recurring theme in his writings is the history and development of pre-evolutionary and evolutionary thought. He was also an enthusiastic baseball fan and sabermetrician (analyst of baseball statistics), and made frequent reference to the sport in his essays. Many of his baseball essays were anthologized in his posthumously published book Triumph and Tragedy in Mudville (2003).

Although a self-described Darwinist, Gould's emphasis was less gradualist and reductionist than most neo-Darwinists. He fiercely opposed many aspects of sociobiology and its intellectual descendant evolutionary psychology. He devoted considerable time to fighting against creationism, creation science, and intelligent design. Most notably, Gould provided expert testimony against the equal-time creationism law in McLean v. Arkansas. Gould later developed the term "non-overlapping magisteria" (NOMA) to describe how, in his view, science and religion should not comment on each other's realm. Gould went on to develop this idea in some detail, particularly in the books Rocks of Ages (1999) and The Hedgehog, the Fox, and the Magister's Pox (2003). In a 1982 essay for Natural History Gould wrote:

Our failure to discern a universal good does not record any lack of insight or ingenuity, but merely demonstrates that nature contains no moral messages framed in human terms. Morality is a subject for philosophers, theologians, students of the humanities, indeed for all thinking people. The answers will not be read passively from nature; they do not, and cannot, arise from the data of science. The factual state of the world does not teach us how we, with our powers for good and evil, should alter or preserve it in the most ethical manner.

An anti-evolution petition drafted by the Discovery Institute inspired the National Center for Science Education to create a pro-evolution counterpart called "Project Steve," which is named in Gould's honor. At a meeting of the executive council of the Committee for Skeptical Inquiry (CSI) in 2011 selected Gould for inclusion in CSI's "Pantheon of Skeptics" created to remember the legacy of deceased CSI fellows and their contributions to the cause of scientific skepticism.

Gould also became a noted public face of science, often appearing on television. In 1984 Gould received his own NOVA special on PBS. Other appearances included interviews on CNN's Crossfire and Talkback Live, NBC's The Today Show, and regular appearances on PBS's Charlie Rose show. Gould was also a guest in all seven episodes of the Dutch talk series A Glorious Accident, in which he appeared with his close friend Oliver Sacks.

Gould was featured prominently as a guest in Ken Burns's PBS documentary Baseball, as well as PBS's Evolution series. Gould was also on the Board of Advisers to the influential Children's Television Workshop television show 3-2-1 Contact, where he made frequent guest appearances.

Since 2013, Gould has been listed on the Advisory Council of the National Center for Science Education.

In 1997 he voiced a cartoon version of himself on the television series The Simpsons. In the episode "Lisa the Skeptic", Lisa finds a skeleton that many people believe is an apocalyptic angel. Lisa contacts Gould and asks him to test the skeleton's DNA. The fossil is discovered to be a marketing gimmick for a new mall. During production the only phrase Gould objected to was a line in the script that introduced him as the "world's most brilliant paleontologist". In 2002 the show paid tribute to Gould after his death, dedicating the season 13 finale to his memory. Gould had died two days before the episode aired.




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