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Natural Selection: Domains, Levels, and Challenges
In this work, George C. Williams--one of evolutionary biology's most distinguished scholars--examines the mechanisms and meaning of natural selection in evolution. Williams offers his own perspective on modern evolutionary theory, including discussions of the gene as the unit of selection, clade selection and macroevolution, diversity within and among populations, stasis, and other timely and provocative topics. In dealing with the levels-of-selection controversy, he urges a pervasive form of the replicator-vehicle distinction. Natural selection, he argues, takes place in the separate domains of information and matter. Levels-of-selection questions, consequently, require different theoretical devices depending on the domains being discussed. In addressing these topics, Williams presents a synthesis of his three decades of research and creative thought which have contributed greatly to evolutionary biology in this century.
224 pages, Paperback
First published June 1, 1992
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March 11, 2025
A WIDE-RANGING DISCUSSION OF ASPECTS OF EVOLUTIONARY THEORY
George Christopher Williams (1926-2010) was an evolutionary geologist. He wrote in the first chapter of this 1992 book, “Successful biological research in this century has had three doctrinal bases: MECHANISM (as opposed to vitalism), NATURAL SELECTION (trial and error, as opposed to rational plan), and HISTORICITY. This last is the recognition or the role or historical contingency in determining properties of the earth’s biota… Mechanism implies that only physico-chemical processes are at work in an organism. Every vital function is performed by material machinery that can in principle be understood from a physical and chemical examination. The opposed doctrine of vitalism maintains that the observable machinery has but limited autonomy and is controlled by a purposive entity peculiar to living organisms. The history of much of biology in the 19th and early 20th centuries can be viewed as a gradual retreat of vitalism and (final?) triumph of mechanism.” (Pg. 3)
He continues, “An open mind must be kept on the issue of vitalism, especially for neural phenomena. There could be no scientific finding more important than the demonstration of a mental principle, not itself physical, but capable of altering cause-effect relations in neural machinery. Such a mechanism would at one stroke compromise mechanism and make possible a scientific study of the mind-body relation… which has justifiably engaged philosophers at least since Descartes.” (Pg. 4)
He adds, “The second basis of modern biology [after Mechanism] is the assumption that the Darwinian process of natural selection accounts for all aspects of the adaptation of an organism to a particular way of life in a particular environment. Natural selection is a system of corrective feedback that favors those individuals that most closely approximate some best available organization for their ecological niche… Biologists today use it … to explain the great generation-to-generation stability of a species’ characteristics, or the near absence of evolution. They assume that an organism is already close to some maximum achievable level of adaptation…” (Pg. 5)
He summarizes, “This book is written from a belief in the continuing validity of the three conceptual foundations of biology, and from a conviction that these three are sufficient at the most basic level… I assume that the same three principles, and no others, will be found to apply everywhere in the universe where life has arisen. This is [Richard] Dawkins’ idea of ‘universal Darwinism.’ Of course any of my three basic principles might be wrong. I suggested as much about mechanism when I discussed the possibility of an immaterial but physically active principle of mentality… I predict that no extraterrestrial biota will ever be found in which the universal principles of mechanism, natural selection, and historicity will not prevail.” (Pg. 7-8)
He states, “Information can proliferate and be edited by natural selection only if the selection affects the information at a greater rate than competing processes such as mutation and drift. A given package of information (codex) must proliferate faster than it changes, so as to produce a genealogy recognizable by some diagnostic effects. A good test of the susceptibility of an entity to natural selection is whether its history can be modeled realistically by a dendogram… the branching lines in the figure could represent the history of a recognizable ‘meme’ (Dawkins’ cultural analog of the gene)… A branching could represent the successful spread of the idea from one family to another.” (Pg. 13)
He suggests, “Genes and memes differ in mode of transmission: genes exclusively from parent to offspring, memes between any associated individuals… There is no more reason to expect a cultural practice transmitted between churchgoers to increase churchgoers’ fitness than there is to expect a similarly transmitted flu virus to increase fitness.” (Pg. 15)
He argues, “Discussion of higher levels of selection have been obfuscated in recent years by the term ‘species selection’ [e.g., Steven Stanley, Stephen Jay Gould]… which implies that there is something special for natural selection, about the species level of classification… If I am right that all levels of genealogical inclusiveness from local gene pools to classes and phyla may be subject to selection, the term ‘clade selection’ … is a great improvement.” (Pg. 24) [Later, he states, “the idea of punctuated equilibrium of [Niles] Eldredge and Gould… is a complex and controversial topic, and I prefer to neglect it except for the brief treatment in chapter 8… Pg. 53]
He acknowledges, “A purely practical consideration is the paucity of information about characters, individual or collective, in organisms known only as fossils. Suppose that, during the Cretaceous, one family of snails started with one genus and ended it with a hundred, while another went from a hundred to zero… It is surely of value here to recognize that the first family had… gene pools superior to those of the second, even though is most unlikely that we will ever know whether it was from coding for more cooperative foraging or for broader temperature tolerance.” (Pg. 27)
He asserts, “In my opinion the recognized microevolutionary processes that form the heart of the neoDarwinian synthesis are an adequate description of the evolution taking place in any Mendelian population… Having taken that position, I must also take another. The microevolutionary process that adequately describers evolution in a population is an utterly inadequate account of the evolution of the Earth’s biota. It is inadequate because the evolution of the biota is more than the mutational origin and subsequent survival or extinction of genes in gene pools… This position is not equivalent to a belief in the efficacy of clade selection. A purely random flux of cladogenesis and extinction would inevitably have important macroevolutionary consequences… S.J. Gould… has consistently emphasized the importance of chance extinction in the evolution of the earth’s biota.” (Pg. 31)
He summarizes, “I hope I have adequately shown that clade selection does operate, at as yet unknown levels of frequency and intensity. It must also be true that there is a strong stochastic element in phylogenetic success and failure, and that chance events can have major effects through all subsequent history.” (Pg. 37)
He notes, “The main theoretical challenge is currently not the reality of natural selection of these entities (transposable elements, outlaw genes, driving chromosomes, etc.), but rather with the general suppression of their short-term selfishness and the usual fairness of cell division and Mendelian heredity. This prevalent ruliness of DNA behavior means that selection at the organismic or higher levels must be much stronger…” (Pg. 41-42)
He rejects the concepts of ‘unity of type’ and ‘Bauplan’… [under which] any major group of organisms …are said to share a basic body plan, and it is supposedly difficult for the evolutionary process to alter this plan. The idea is misguided and dispensable… Having one head, one tail, and one heart seem to be normal conditions for vertebrates but… there is no reason to think of them as parts of a developmentally inflexible ‘Bauplan.’” (Pg. 87-88)
He contends, “I am sure I could get a monkey… to compose Hamlet’s soliloquy in a few minutes. I need only impose a program of artificial selection that would preserve everything the computer-monkey did that made its cumulative effort resemble Hamlet’s words more closely, and reject all changes that decreased the resemblance. The idea of producing some specified text by the editing of randomly placed symbols may not be an obvious analogy to the Darwinian process.
He observes, “The question of evolutionary rate in relation to available geological time is indeed a serious theoretical challenge, but the reason is exactly the opposite of that inspired by most people’s intuitions. Organisms in general have not done nearly as much evolving as we should reasonably expect. Long-term rates of change… are almost far slower than they theoretically could be. The basis for such expectation is to be found most clearly in observed rate of change in environmental conditions that must imply rapid change in the intensity and direction of selection in nature.” (Pg. 128)
He notes, “a large proportion of Recent species are essentially identical to their Pliocene ancestors … I find it puzzling that no shift in selection pressures in the last 100 million years would have produced any noteworthy change in a structure as complex and informative as a fish skeleton, which ought to reflect even very subtle changes in locomotor or trophic adaptations.” (Pg. 131-132)
He asks, “Why are there no viviparous [live-bearing] turtles?... There [are] no obvious impediment[s] to egg retention in turtles that would not apply to snakes and lizards, which have evolved viviparity many times… It is easy to imagine that the preadaptations for the evolution of internal fertilization would often be lacking… The universality of internal fertilization with total absence of viviparity in the turtles and birds seems most mysterious.” (Pg. 140-141)
This book will be of keen interest to those seriously studying evolutionary theory.
George Christopher Williams (1926-2010) was an evolutionary geologist. He wrote in the first chapter of this 1992 book, “Successful biological research in this century has had three doctrinal bases: MECHANISM (as opposed to vitalism), NATURAL SELECTION (trial and error, as opposed to rational plan), and HISTORICITY. This last is the recognition or the role or historical contingency in determining properties of the earth’s biota… Mechanism implies that only physico-chemical processes are at work in an organism. Every vital function is performed by material machinery that can in principle be understood from a physical and chemical examination. The opposed doctrine of vitalism maintains that the observable machinery has but limited autonomy and is controlled by a purposive entity peculiar to living organisms. The history of much of biology in the 19th and early 20th centuries can be viewed as a gradual retreat of vitalism and (final?) triumph of mechanism.” (Pg. 3)
He continues, “An open mind must be kept on the issue of vitalism, especially for neural phenomena. There could be no scientific finding more important than the demonstration of a mental principle, not itself physical, but capable of altering cause-effect relations in neural machinery. Such a mechanism would at one stroke compromise mechanism and make possible a scientific study of the mind-body relation… which has justifiably engaged philosophers at least since Descartes.” (Pg. 4)
He adds, “The second basis of modern biology [after Mechanism] is the assumption that the Darwinian process of natural selection accounts for all aspects of the adaptation of an organism to a particular way of life in a particular environment. Natural selection is a system of corrective feedback that favors those individuals that most closely approximate some best available organization for their ecological niche… Biologists today use it … to explain the great generation-to-generation stability of a species’ characteristics, or the near absence of evolution. They assume that an organism is already close to some maximum achievable level of adaptation…” (Pg. 5)
He summarizes, “This book is written from a belief in the continuing validity of the three conceptual foundations of biology, and from a conviction that these three are sufficient at the most basic level… I assume that the same three principles, and no others, will be found to apply everywhere in the universe where life has arisen. This is [Richard] Dawkins’ idea of ‘universal Darwinism.’ Of course any of my three basic principles might be wrong. I suggested as much about mechanism when I discussed the possibility of an immaterial but physically active principle of mentality… I predict that no extraterrestrial biota will ever be found in which the universal principles of mechanism, natural selection, and historicity will not prevail.” (Pg. 7-8)
He states, “Information can proliferate and be edited by natural selection only if the selection affects the information at a greater rate than competing processes such as mutation and drift. A given package of information (codex) must proliferate faster than it changes, so as to produce a genealogy recognizable by some diagnostic effects. A good test of the susceptibility of an entity to natural selection is whether its history can be modeled realistically by a dendogram… the branching lines in the figure could represent the history of a recognizable ‘meme’ (Dawkins’ cultural analog of the gene)… A branching could represent the successful spread of the idea from one family to another.” (Pg. 13)
He suggests, “Genes and memes differ in mode of transmission: genes exclusively from parent to offspring, memes between any associated individuals… There is no more reason to expect a cultural practice transmitted between churchgoers to increase churchgoers’ fitness than there is to expect a similarly transmitted flu virus to increase fitness.” (Pg. 15)
He argues, “Discussion of higher levels of selection have been obfuscated in recent years by the term ‘species selection’ [e.g., Steven Stanley, Stephen Jay Gould]… which implies that there is something special for natural selection, about the species level of classification… If I am right that all levels of genealogical inclusiveness from local gene pools to classes and phyla may be subject to selection, the term ‘clade selection’ … is a great improvement.” (Pg. 24) [Later, he states, “the idea of punctuated equilibrium of [Niles] Eldredge and Gould… is a complex and controversial topic, and I prefer to neglect it except for the brief treatment in chapter 8… Pg. 53]
He acknowledges, “A purely practical consideration is the paucity of information about characters, individual or collective, in organisms known only as fossils. Suppose that, during the Cretaceous, one family of snails started with one genus and ended it with a hundred, while another went from a hundred to zero… It is surely of value here to recognize that the first family had… gene pools superior to those of the second, even though is most unlikely that we will ever know whether it was from coding for more cooperative foraging or for broader temperature tolerance.” (Pg. 27)
He asserts, “In my opinion the recognized microevolutionary processes that form the heart of the neoDarwinian synthesis are an adequate description of the evolution taking place in any Mendelian population… Having taken that position, I must also take another. The microevolutionary process that adequately describers evolution in a population is an utterly inadequate account of the evolution of the Earth’s biota. It is inadequate because the evolution of the biota is more than the mutational origin and subsequent survival or extinction of genes in gene pools… This position is not equivalent to a belief in the efficacy of clade selection. A purely random flux of cladogenesis and extinction would inevitably have important macroevolutionary consequences… S.J. Gould… has consistently emphasized the importance of chance extinction in the evolution of the earth’s biota.” (Pg. 31)
He summarizes, “I hope I have adequately shown that clade selection does operate, at as yet unknown levels of frequency and intensity. It must also be true that there is a strong stochastic element in phylogenetic success and failure, and that chance events can have major effects through all subsequent history.” (Pg. 37)
He notes, “The main theoretical challenge is currently not the reality of natural selection of these entities (transposable elements, outlaw genes, driving chromosomes, etc.), but rather with the general suppression of their short-term selfishness and the usual fairness of cell division and Mendelian heredity. This prevalent ruliness of DNA behavior means that selection at the organismic or higher levels must be much stronger…” (Pg. 41-42)
He rejects the concepts of ‘unity of type’ and ‘Bauplan’… [under which] any major group of organisms …are said to share a basic body plan, and it is supposedly difficult for the evolutionary process to alter this plan. The idea is misguided and dispensable… Having one head, one tail, and one heart seem to be normal conditions for vertebrates but… there is no reason to think of them as parts of a developmentally inflexible ‘Bauplan.’” (Pg. 87-88)
He contends, “I am sure I could get a monkey… to compose Hamlet’s soliloquy in a few minutes. I need only impose a program of artificial selection that would preserve everything the computer-monkey did that made its cumulative effort resemble Hamlet’s words more closely, and reject all changes that decreased the resemblance. The idea of producing some specified text by the editing of randomly placed symbols may not be an obvious analogy to the Darwinian process.
He observes, “The question of evolutionary rate in relation to available geological time is indeed a serious theoretical challenge, but the reason is exactly the opposite of that inspired by most people’s intuitions. Organisms in general have not done nearly as much evolving as we should reasonably expect. Long-term rates of change… are almost far slower than they theoretically could be. The basis for such expectation is to be found most clearly in observed rate of change in environmental conditions that must imply rapid change in the intensity and direction of selection in nature.” (Pg. 128)
He notes, “a large proportion of Recent species are essentially identical to their Pliocene ancestors … I find it puzzling that no shift in selection pressures in the last 100 million years would have produced any noteworthy change in a structure as complex and informative as a fish skeleton, which ought to reflect even very subtle changes in locomotor or trophic adaptations.” (Pg. 131-132)
He asks, “Why are there no viviparous [live-bearing] turtles?... There [are] no obvious impediment[s] to egg retention in turtles that would not apply to snakes and lizards, which have evolved viviparity many times… It is easy to imagine that the preadaptations for the evolution of internal fertilization would often be lacking… The universality of internal fertilization with total absence of viviparity in the turtles and birds seems most mysterious.” (Pg. 140-141)
This book will be of keen interest to those seriously studying evolutionary theory.
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