Darwin and His Pigeons. The Analogy Between Artificial and Natural Selection Revisited

The analogy between artificial selection of domestic varieties and natural selection in nature was a vital element of Darwin’s argument in his Origin of Species. Ever since, the image of breeders creating new varieties by artificial selection has served as a convincing illustration of how the theory works. In this paper I argue that we need to reconsider our understanding of Darwin’s analogy. Contrary to what is often assumed, nineteenth-century animal breeding practices constituted a highly controversial field that was fraught with difficulties. It was only with considerable effort that Darwin forged his analogy, and he only succeeded by downplaying the importance of two other breeding techniques – crossing of varieties and inbreeding – that many breeders deemed essential to obtain new varieties. Part of the explanation for Darwin’s gloss on breeding practices, I shall argue, was that the methods of his main informants, the breeders of fancy pigeons, were not representative of what went on in the breeding world at large. Darwin seems to have been eager to take the pigeon fanciers at their word, however, as it was only their methods that provided him with the perfect analogy with natural selection. Thus while his studies of domestic varieties were important for the development of the concept of natural selection, the reverse was also true: Darwin’s comprehension of breeding practices was moulded by his understanding of the working of natural selection in nature. Historical studies of domestic breeding practices in the eighteenth and nineteenth century confirm that, besides selection, the techniques of inbreeding and crossing were much more important than Darwin’s interpretation allowed for. And they still are today. This calls for a reconsideration of the pedagogic use of Darwin’s analogy too.     

Darwin’s Sublime: The Contest Between Reason and Imagination in <Emphasis Type="Italic">On the Origin of Species</Emphasis>

Recent Darwin scholarship has provided grounds for recognising the Origin as a literary as well as a scientific achievement. While Darwin was an acute observer, a gifted experimentalist and indefatigable theorist, this essay argues that it was also crucial to his impact that the Origin transcended the putative divide between the scientific and the literary. Analysis of Darwin’s development as a writer between his journal-keeping on HMS Beagle and his construction of the Origin argues the latter draws on the pattern of the Romantic or Kantian sublime. The Origin repeatedly uses strategies which challenge the natural-theological appeal to the imagination in conceiving nature. Darwin’s sublime coaches the Origin’s readers into a position from which to envision nature that reduces and contains its otherwise overwhelming complexity. As such, it was Darwin’s literary achievement that enabled him to fashion a new ‘habit of looking at things in a given way’ that is the centrepiece of the scientific revolution bearing his name.     

Artificial Selection and Domestication: Modern Lessons from Darwin’s Enduring Analogy

It is clear from his published works that Charles Darwin considered domestication to be very useful in exploring and explaining mechanisms of evolutionary change. Not only did domestication occupy the introductory chapter of On the Origin of Species, but he revisited the topic in a two-volume treatise less than a decade later. In addition to drawing much of his information about heredity from studies of domesticated animals and plants, Darwin saw important parallels between the process of artificial selection by humans and natural selection by the environment. There was resistance to this analogy even among Darwin’s contemporary supporters when it was proposed, and there also has been disagreement among historians and philosophers regarding the role that the analogy with artificial selection actually played in the discovery of natural selection. Regardless of these issues, the analogy between artificial and natural selection remains important in both research and education in evolution. In particular, the present article reviews ten lessons about evolution that can be drawn from the modern understanding of domestication and artificial selection. In the process, a basic overview is provided of current approaches and knowledge in this rapidly advancing field.

Separated at Birth: The Interlinked Origins of Darwin’s Unconscious Selection Concept and the Application of Sexual Selection to Race

This essay traces the interlinked origins of two concepts found in Charles Darwin’s writings: “unconscious selection,” and sexual selection as applied to humanity’s anatomical race distinctions. Unconscious selection constituted a significant elaboration of Darwin’s artificial selection analogy. As originally conceived in his theoretical notebooks, that analogy had focused exclusively on what Darwin later would call “methodical selection,” the calculated production of desired changes in domestic breeds. By contrast, unconscious selection produced its results unintentionally and at a much slower pace. Inspiration for this concept likely came from Darwin’s early reading of works on both animal breeding and physical ethnology. Texts in these fields described the slow and unplanned divergence of anatomical types, whether animal or human, under the guidance of contrasting ideals of physical perfection. These readings, it is argued, also led Darwin to his theory of sexual selection as applied to race, a theme he discussed mainly in his book The Descent of Man (1871). There Darwin described how the racial version of sexual selection operated on the same principle as unconscious selection. He thereby effectively reunited these kindred concepts.     

“A Great Complication of Circumstances” – Darwin and the Economy of Nature

In 1749, Linnaeus presided over the dissertation “Oeconomia Naturae,” which argued that each creature plays an important and particular role in nature’s economy. This phrase should be familiar to readers of Darwin, for he claims in the Origin that “all organic beings are striving, it may be said, to seize on each place in the economy of nature.” Many scholars have discussed the influence of political economy on Darwin’s ideas. In this paper, I take a different tack, showing that Darwin’s idea of an economy of nature stemmed from the views of earlier naturalists like Linnaeus and Lyell. I argue, in the first section of the paper, that Linnaeus’ idea of oeconomia naturae is derived from the idea of the animal economy, and that his idea of politia naturae is an extension of the idea of a politia civitatis. In the second part, I explore the use of the concept of stations in the work of De Candolle and Lyell – the precursor to Darwin’s concept of places. I show in the third part of the paper that the idea of places in an economy of nature is employed by Darwin at many key points in his thinking: his discussion of the Galapagos birds, his reading of Malthus, etc. Finally, in the last section, I demonstrate that the idea of a place in nature’s economy is essential to Darwin’s account of divergence. To tell his famous story of divergence and adaptation, Darwin needed the economy of nature.     

Adaptations: Using Darwin's Origin to teach biology and writing

Charles Darwin's On the Origin of Species is at once familiar and unfamiliar. Everyone knows that the Origin introduced the world to the idea of evolution by natural selection, but few of us have actually read it. We suggest that it is worth taking the time not only to read what Darwin had to say, but also to use the Origin to teach both biology and writing. It provides scientific lessons in areas beyond evolutionary biology, such as ecology and biogeography. In addition, it provides valuable rhetorical lessons—how to construct an argument, write persuasively, make use of evidence, know your audience, and anticipate counterarguments. We have been using the Origin in various classes for several years, introducing new generations to Darwin, in his own words.     

Extending Darwin’s analogy: Bridging differences in concepts of selection between farmers, biologists, and plant breeders

Darwin developed his theory of evolution based on an analogy between artificial selection by breeders of his day and “natural selection.” For Darwin, selection included what biologists came to see as being composed of (1) phenotypic selection of individuals based on phenotypic differences, and, when these are based on heritable genotypic differences, (2) genetic response between generations, which can result in (3) evolution (cumulative directional genetic response over generations). The use of the term “selection” in biology and plant breeding today reflects Darwin’s assumption—phenotypic selection is only biologically significant when it results in evolution. In contrast, research shows that small-scale, traditionally-based farmers select seed as part of an integrated production and consumption system in which selection is often not part of an evolutionary process, but is still useful to farmers. Extending Darwin’s analogy to farmers can facilitate communication between farmers, biologists, and plant breeders to improve selection and crop genetic resource conservation.     

Why Trees Are Important

The Tree of Life is the result of the interplay of changes in information and speciation. Almost 100 years after publication of Darwin’s Origin, the inception of Phylogenetic Systematics has resulted in a revolution in data inference. I briefly trace the development of this revolution and show examples of how data are interpreted relative to phylogenetic trees. I then provide brief discussions of how to read tree diagrams and the need to access the quality of phylogenetic inference.     

The Preface to Darwin’s <Emphasis Type="Italic">Origin of Species</Emphasis>: The Curious History of the “Historical Sketch” *

Almost any modern reader’s first encounter with Darwin’s writing is likely to be the “Historical Sketch,” inserted by Darwin as a preface to an early edition of the Origin of Species, and having since then appeared as the preface to every edition after the second English edition. The Sketch was intended by him to serve as a short “history of opinion” on the species question before he presented his own theory in the Origin proper. But the provenance of the “Historical Sketch” is somewhat obscure. Some things are known about its production, such as when it first appeared and what changes were made to it between its first appearance in 1860 and its final form, for the fourth English edition, in 1866. But how it evolved in Darwin’s mind, why he wrote it at all, and what he thought he was accomplishing by prefacing it to the Origin remain questions that have not been carefully addressed in the scholarly literature on Darwin. I attempt to show that Darwin’s various statements about the “Historical Sketch,” made primarily to several of his correspondents between 1856 and 1860, are somewhat in conflict with one another, thus making problematic a satisfactory interpretation of how, when, and why the Sketch came to be. I also suggest some probable resolutions to the several difficulties. How Darwin came to settle on the title “Historical Sketch” for the Preface to the Origin is not certain, but a guess may be ventured. When he first submitted the text to Asa Gray in February 1860 he called it simply “Preface Contributed by the Author to this American Edition” (Burkhardt et al., eds., vol. 8, 1993, p. 572; the collected correspondence is hereafter cited as CCD). In fact he had thought of it as being properly called a Preface much earlier, perhaps as early as 1856, as will be seen in what follows. It came to be called “An Historical Sketch of the Recent Progress of Opinion on the Origin of Species” only in the third English edition, April 1861. This is the title it retained thereafter, with the exception of an addition to the title in the sixth English edition, “Previously to the Publication of the First Edition of this Work” (Peckham, 1959, pp. 20, 59). The word “sketch,” on the other hand was one of two words Darwin commonly used in private correspondence to refer to the book that would later become the Origin, the other word being “Abstract,” and both signifying that Darwin thought of the work as being a resume rather than a full-fledged study (e.g., letter to J.D. Hooker, May 9 1856, CCD vol. 6 p. 106; letter to Baden Powell January 18 1860, CCD vol. 8 p. 41; letter to Lyell 25 June 1858, CCD v. 7, 1991, pp. 117–8; letter to Lyell May 1856, CCD, v. 6 p. 100). The most likely source of the title “Historical Sketch” for Darwin’s Preface is Charles Lyell’s Principles of Geology in which, beginning with the third edition (1834), Lyell added titles to his chapters, calling chapters 2–4 “Historical Sketch of the Progress of Geology” (Secord, in Lyell [1997], p. xlvii; for other uses by Lyell of this expression, cf. Porter, 1976, p. 95; idem 1982, p. 38; and Lyell, 1830 [1990], p. 30). Further parallels between Lyell’s Introduction and Darwin’s “Historical Sketch” in terms of content and strategy are suggested below.     

Why Darwin rejected intelligent design

As a Cambridge University undergraduate Charles Darwin was fascinated and convinced by the argument for intelligent design, as set forth in William Paley’s 1802 classic, Natural Theology. Subsequently, during his five-year voyage on HMS Beagle (1831–1836), Darwin interpreted his biological findings through a creationist lens, including the thought-provoking evidence he encountered during his historic visit to the Galápagos Islands in September and October 1835. After his return to England in 1836 and his subsequent conversion to the idea of organic evolution in March 1837, Darwin searched for a theory that would explain both the fact of evolution and the widespread appearance of intelligent design. His insight into the process of natural selection, which occurred in September 1838, provided this alternative explanation. Darwin’s Origin of Species (1859) exemplifies his skillful deployment of the hypothetico-deductive method in testing and refuting the arguments for intelligent design that he had once so ardently admired.     

Brocchi,Darwin, and Transmutation: Phylogenetics and Paleontology at the Dawn of Evolutionary Biology

Giambattista Brocchi’s (1814) monograph (see Dominici, Evo Edu Outreach, this issue, 2010) on the Tertiary fossils of the Subappenines in Italy—and their relation to the living molluscan fauna—contains a theoretical, transmutational perspective (“Brocchian transmutation”). Unlike Lamarck (1809), Brocchi saw species as discrete and fundamentally stable entities. Explicitly analogizing the births and deaths of species with those of individual organisms (“Brocchi’s analogy”), Brocchi proposed that species have inherent longevities, eventually dying of old age unless driven to extinction by external forces. As for individuals, births and deaths of species are understood to have natural causes; sequences of births and deaths of species produce genealogical lineages of descent, and faunas become increasingly modernized through time. Brocchi calculated that over 50% of his fossil species are still alive in the modern fauna. Brocchi’s work was reviewed by Horner (1816) in Edinburgh. Brocchi’s influence as a transmutational thinker is clear in Jameson’s (1827) “geological illustrations” in his fifth edition of his translation of Cuvier’s Theory of the Earth (read by his student Charles Darwin) and in the anonymous essays of 1826 and 1827 published in the Edinburgh New Philosophical Journal—which also carried a notice of Brocchi’s death in 1827. The notion that new species replace older, extinct ones—in what today would be called an explicitly phylogenetic context—permeates these essays. Herschel’s (1830) discussion of temporal replacement of species and the modernization of faunas closely mirrors these prior discussions. His book, dedicated to the search for natural causes of natural phenomena, was read by Charles Darwin while a student at Cambridge. Darwin’s work on HMS Beagle was in large measure an exploration of replacement patterns of “allied forms” of endemic species in time and in space. His earliest discussions of transmutation, in his essay February 1835, as well as the Red Notebook and the early pages of Notebook B (the latter two written in 1837 back in England), contain Brocchi’s analogy, including the idea of inherent species longevities. Darwin’s first theory of the origin of species was explicitly saltational, invoking geographic isolation as the main cause of the abrupt appearance of new species. We conclude that Darwin was testing the predicted patterns of both Brocchian and Lamarckian transmutation as early as 1832 at the outset of his work on the Beagle.     

Diatoms from two macro-tidal mudflats in Chignecto Bay,Upper Bay of Fundy,New Brunswick,Canada

Mudflat research is dispersed among several fields (ecology, sedimentology), each with its own focus and methodology. Consequently, although the volume of mudflat literature is considerable, our understanding of mudflat ecology remains fragmented. For example, little is known about the structure of microbial communities outside Western Europe. Here we present the first North American specific composition and densities of live mudflat diatoms and relate them to properties of their environment on two closely located flats. The two flats (Daniel’s and Buck’s) were similar until the mid–1980s. After this time the biological and sedimentary environment on Buck’s Flats began to change and resulted in a precipitous decline of the keystone invertebrate Corophium volutator (Pallas). The specific diatom composition on each of the two flats examined was still very similar. Tychoplanktonic diatoms were numerically dominant on both flats. The flats differed significantly in the relative contribution of epipelic diatoms, which were about an order of magnitude greater on Buck’s Flats. CCA analysis suggests that very few of these species exist within their optimal habitat. Some of the differences appeared small, but were statistically and biologically significant. Daniel’s Flats sediments had a 30% larger mean grain size, less water and organic carbon compared to Buck’s Flats sediments. Buck’s Flats had more variable depths of the oxygenated layer, often with anoxic inclusions throughout. Daniel’s Flats supported more C. volutator, while Buck’s Flats contained greater densities of diatoms. The importance of preserving environmental conditions (sedimentary and biotic) prevailing on flats such as Danielȁ9s Flat in order to foster populations of Corophium at a level necessary to support foraging migratory shorebirds is also discussed.     

The ‘philosophical grasp of the appearances’ and experimental microscopy: Johannes Müller’s microscopical research, 1824–1832

Romantic Naturphilosophie has been at the centre of almost every account of early nineteenth-century sciences, be it as an obstacle or as an aid for scientific advancement. The following paper suggests a change of perspective. I seek to read Naturphilosophie as one manifestation among others of a more general concern with the question of how experience enables the subject to acquire knowledge about objects. To illustrate such an approach, I focus on Johannes Müller’s early work. Here one finds two contrasting images of microscopical observation, its set-up, and the observer: the embryological study of 1830 demands a ‘philosophical grasp’ of the appearances. In contrast, the investigations of blood of 1832 are presented as a series of controlled experiments. I argue that an interpretation of this contrast in terms of an appropriation and casting aside of Naturphilosophie is not altogether convincing. Instead, both images of microscopy are manifestations of a more general problem, namely, the problem of exactly how subject and object came together in experience. I show how this concern not only shaped the methodological sensibilities particular to Müller’s embryology and the investigation of bodily liquids but also provided the epistemological principles and the target for his sense-physiological experiments. It bound Müller’s work together with Naturphilosophie and linked Naturphilosophie with other contemporaneous projects in philosophy. All of these enterprises sought to contribute to ongoing debates about how experience allowed the subject to acquire knowledge about the world.     

Brocchi’s Subapennine Fossil Conchology

The Italian geologist Giambattista Brocchi (1771–1826) is presented as a key figure in the historical period preceding young Charles Darwin’s first work on transmutational theory while on the Beagle. The brief biographical account focuses on Brocchi’s writings related to his analogy that species have births and deaths like individuals, and culminates in his most important work, Subapennine Fossil Conchology of 1814. Brocchi’s analogy as an original and fertile way to approach the fossil record was to influence Darwin’s first evolutionary thinking. Relevant passages of the book are presented for the first time in an English translation.     

<Emphasis Type="Italic">Ancient Hunters and Their Modern Representatives</Emphasis>: William Sollas’s (1849–1936) Anthropology from Disappointed Bridge to Trunkless Tree and the Instrumentalisation of Racial Conflict

During the first decades of the 20th century, many anthropologists who had previously adhered to a linear view of human evolution, from an ape via Pithecanthropus erectus(today Homo erectus) and Neanderthal to modern humans, began to change their outlook. A shift towards a branching model of human evolution began to take hold. Among the scientific factors motivating this trend was the insight that mammalian evolution in general was best represented by a branching tree, rather than by a straight line, and that several new fossil hominids were discovered that differed significantly in their morphology but seemed to date from about the same period. The ideological and practical implications of imperialism and WWI have also been identified as formative of the new evolutionary scenarios in which racial conflict played a crucial role. The paper will illustrate this general shift in anthropological theory for one particular scientist, William Sollas (1849–1936). Sollas achieved a synthesis of human morphological and cultural evolution in what I will refer to as an imperialist model. In this theoretical framework, migration, conflict, and replacement became the main mechanisms for progress spurred by ‘ ȁ8nature’s tyrant,’ natural selection.     

Thermodynamical interpretation of evolutionary dynamics on a fitness landscape in an evolution reactor,II

In our previous report [Aita, T., Morinaga, S., Hosimi, Y., 2004. Thermodynamical interpretation of evolutionary dynamics on a fitness landscape in an evolution reactor I. Bull. Math. Biol. 66, 1371–1403], an analogy between thermodynamics and adaptive walks on a Mt. Fuji-type fitness landscape in an artificial selection system was presented. Introducing the ‘free fitness’ as the sum of a fitness term and an entropy term and ‘evolutionary force’ as the gradient of free fitness on a fitness coordinate, we demonstrated that the adaptive walk (=evolution) is driven by the evolutionary force in the direction in which free fitness increases. In this report, we examine the effect of various modifications of the original model on the properties of the adaptive walk. The modifications were as follows: first, mutation distance d was distributed obeying binomial distribution; second, the selection process obeyed the natural selection protocol; third, ruggedness was introduced to the landscape according to the NK model; fourth, a noise was included in the fitness measurement. The effect of each modification was described in the same theoretical framework as the original model by introducing ‘effective’ quantities such as the effective mutation distance or the effective screening size.     

Experimenting with Transmutation: Darwin, the Beagle, and Evolution

Detailed analysis of Darwin’s scientific notes and other writings from the Beagle voyage reveals a focus on endemism and replacement of allied taxa in time and in space that began early in the journey. Though it is impossible to determine exactly when Darwin became a transmutationist, the evidence suggests that he was conversant with the transmutational ideas of Lamarck and others and testing (“experimenting” with) them—before he received a copy of Lyell’s Principles of Geology, vol. 2, in November 1832, in which Lyell describes and disputes Lamarck’s theory. To the two rhea species of Patagonia and the four mockingbird species of the Galapagos, we can now add the living Patagonian cavy (rodent) species, and its extinct putatively related species that Darwin collected at Monte Hermoso (Bahia Blanca) in the Fall of 1832, as a replacement pattern absolutely critical to the development of Darwin’s transmutational thinking. Darwin developed his first transmutational theory by adopting “Brocchi’s analogy” (Rudwick 2008)—i.e. that births and deaths of species are analogous to the births and deaths of individuals. Births and deaths of species, as of individuals, are thus explicable in terms of natural causes. Darwin explored these themes and the replacement of the extinct cavy by the modern species explicitly in his February 1835 essay (Darwin 1835a).

The Charms of Nature: Darwin on Meaning and Value

I review George Levine’s provocative and highly original book Darwin Loves You. Levine, whose “home discipline” is English Literature, offers a compelling interpretation of Darwin’s works, evaluating their content and Darwin’s prose style to identify a distinctly Darwinian attitude toward nature as a source of meaning and value. Levine believes that Darwin exemplifies the capacity to feel “enchantment” about the natural world, suggesting that, if Darwin’s example were followed, a “Darwinian re-enchantment of the world” would be brought about. This would offer a secular, non-supernatural basis for purpose, meaning, and value. I conclude with a few critical remarks about the scope and cogency of Levine’s proposal.     

Ludwik Fleck and the causative agent of syphilis: sociology or pathology of science? A rejoinder to Jean Lindenmann

In 1905 two different microbes were proposed to fill the vacant role of etiologic agent for syphilis, one, the Cytorrhyctes luis, by John Siegel, the other, Spirochaeta pallida, by Fritz Schaudinn. After gathering and reviewing the evidence the majority of medical scientists decided in favor of Schaudinn’s candidate. In a previous issue Jean Lindenmann challenged Ludwik Fleck’s suggestion that under suitable social conditions Siegel’s candidate could just as well have won acceptance by the scientific community (Lindenmann, 2001). To refute this counterfactual thesis, Lindenmann presented an asymmetric account of the dispute over the etiology of syphilis. He adopted the view of the proponents that Schaudinn’s spirochete had already been there in syphilitic lesions for centuries, only awaiting the discovery of an appropriate staining technique to be revealed. Here a more symmetric analysis of the episode will be attempted, paying serious attention to the arguments put forward by the spirochete’s opponents, who expatiated on the many possibilities of inadvertently creating artifacts through microscopic preparation and staining. The symmetric account that is presented in this rejoinder thus aims to trace the simultaneous construction of facts and artifacts. It will not, however, resurrect Fleck’s counterfactual thesis.