On Carl Gegenbaur’s theory on head metamerism and the selection of taxa for comparisons

Summary After the rejection of the idealistic “vertebral theory of the skull,” the basic idea of the theory had been taken up again by the German morphologist Carl Gegenbaur. He proposed a metameric ordering system of the vertebrate head based on a comparative investigation of selachians. This “segmentation theory of the vertebrate head” had a stimulating effect on comparative anatomy. Traces of the basic ideas are still inherent in the biological sciences. Despite the fact that Carl Gegenbaur proposed a rather phylogenetic approach, his systematic investigations remained on a typological level. It is argued herein, that the reason for that was his inability to perform his comparisons independent of a rather intuitive a priori hypothesis about phylogenetic relationships, thus leaving no possibilities for testing homologies. This is exemplified by Carl Gegenbaur’s choice of species for comparative work as well as his use of these species in comparative work. Choosing species for comparisons is discussed.     

The history of comparative anatomy in Jena — an overview

Summary Comparative anatomy and zoology both have long academic traditions in Jena. At first, the two subjects developed in parallel and had many similarites in research topics. This development is covered in the first part of the paper. The close relationship between the two subjects started to break apart when Carl Gegenbaur and Ernst Haeckel were active at Jena University. In 1865 Haeckel became the first full professor of zoology in Jena, and zoology became more independent from comparative anatomy. In the second part of our paper, we follow the developments in comparative anatomy in Jena from Gegenbaur’s immediate students up until the end of Hans B?ker’s tenure in Jena in the mid-1930s. Certain subjects are in focus throughout this period, for example vertebrate head morphology and development, (the “head problem”), the relationship between anatomy and biology, and evolutionary questions. Some of these subjects have remained important research topics in zoology and comparative anatomy in Jena until the present day.     

Post-Haeckelian comparative biology — Adolf Naef’s idealistic morphology

Summary The present study describes the conceptual framework of Adolf Naef’s idealistic morphology as presented at the onset of the 20^th century. According to Naef, Haeckel’s and Gegenbaur’s approaches towards a phylogenetic biology were insufficient. He made it clear that Haeckel’s ideas were based on typological morphology. Thus, Haeckel’s views on comparative biology pointed back to pre-Darwinian concepts. Naef’s consequence was not to work out his own evolutionary morphology but to systematize the earlier typological concept. Consequently, he separated comparative morphology from phylogenetic studies. This idea was adopted by Hennig and was even imported into modern cladism.     

Rational Love,Relational Medicine: Psychiatry and the Accumulation of Precarious Kinship

In north Indian psychiatry, clinical attentions to women’s symptoms often involve scrutiny of emotions related to marriage and its breakdown. In pharmaceutically oriented practice, relations are used to evaluate biologies, and drugs produce the truth about relations at the same time that they produce the truth about bodies. In the process, clinical practice often involves unmaking relations, generating loss, in certain instances, as a dire result. In this, a particular kind of clinical knowing emerges, engaging broad cultural and historical connections between love and madness more than definitions of right and wrong unions. In asking how disciplinary and relational modes of biomedicine converge, I argue that in north Indian psychiatry’s attentions to women, rather than enforcing normative configurations of “the family,” biomedicine grapples with the gendered fallout of kinship.     

Functional homology and homology of function: biological concepts and philosophical consequences

“Functional homology” appears regularly in different areas of biological research and yet it is apparently a contradiction in terms—homology concerns identity of structure regardless of form and function. I argue that despite this conceptual tension there is a legitimate conception of ‘homology of function’, which can be recovered by utilizing a distinction from pre-Darwinian physiology (use versus activity) to identify an appropriate meaning of ‘function’. This account is directly applicable to molecular developmental biology and shares a connection to the theme of hierarchy in homology. I situate ‘homology of function’ within existing definitions and criteria for structural assessments of homology, and introduce a criterion of ‘organization’ for judging function homologues, which focuses on hierarchically interconnected interdependencies (similar to relative position and connection for skeletal elements in structural homology). This analysis of biological concepts has at least three broad philosophical consequences: (1) it provides the grounds for the study of behavior and psychological categories as homologues; (2) it demonstrates that philosophers who take selected effect function as primary effectively ignore large portions of comparative, structural, and experimental research, thereby misconstruing biological reasoning and knowledge; and, (3) it underwrites causal generalizations, which illuminates inferences made from model organisms in experimental biology.

Are Animals Just Noisy Machines?: Louis Boutan and the Co-invention of Animal and Child Psychology in the French Third Republic

Historians of science have only just begun to sample the wealth of different approaches to the study of animal behavior undertaken in the twentieth century. To date, more attention has been given to Lorenzian ethology and American behaviorism than to other work and traditions, but different approaches are equally worthy of the historian’s attention, reflecting not only the broader range of questions that could be asked about animal behavior and the “animal mind” but also the different contexts in which these questions were important. One such approach is that represented by the work of the French zoologist Louis Boutan (1859–1934). This paper explores the intellectual and cultural history of Boutan’s work on animal language and the animal mind, and contextualizes the place of animal behavior studies within late-nineteenth-and early-twentieth-century French biology. I explore the ways in which Boutan addressed the philosophical issue of whether language was necessary for abstract thought and show how he shifted from the idea that animals were endowed with a purely affective language to the notion that of they were capable of “rudimentary” reasoning. I argue that the scientific and broader socio-cultural contexts in which Boutan operated played a role in this transition. Then I show how Boutan’s linguistic and psychological experiments with a gibbon and children provide insights into his conception of “naturalness.” Although Boutan reared his gibbon at home and studied it in the controlled environment of his laboratory, he continued to identify its behavior as “natural.” I specifically demonstrate the importance of the milieu of the French Third Republic in shaping Boutan’s understanding not only of animal intelligence and child education, but also his definition of nature. Finally, I argue that Boutan’s studies on the primate mind provide us with a lens through which we can examine the co-invention of animal and child psychology in early-twentieth-century France.     

“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.     

Defining and explaining culture (comments on Richerson and Boyd, Not by genes alone)

We argue that there is a continuum of cases without any demarcation between more individual and more cultural information, and that therefore “culture” should be viewed as a property that human mental representations and practices exhibit to a varying degree rather than as a type or a subclass of these representations and practices (or of “information”). We discuss the relative role of preservative and constructive processes in transmission. We suggest a revision of Richerson and Boyd’s classification of the forces of cultural evloution.     

Psychoontogeny and psychophylogeny: Bernhard Rensch’s (1900–1990) selectionist turn through the prism of panpsychistic identism

Toward the end of the 1930s, Bernhard Rensch (1900–1990) turned from Lamarckism and orthogenesis to selectionism and became one of the key figures in the making of the Synthetic Theory of Evolution (STE). He contributed to the Darwinization of biological systematics, the criticism of various anti-Darwinian movements in the German lands, but more importantly founded a macroevolutionary theory based on Darwinian gradualism. In the course of time, Rensch’s version of the STE developed into an all-embracing metaphysical conception based on a kind of Spinozism. Here we approach Rensch’s “selectionist turn” by outlining its context, and by analyzing his theoretical transformation. We try to reconstruct the immanent logic of Rensch’s evolution from a “Lamarckian Synthesis” to a “Darwinian Synthesis”. We will pay close attention to his pre-Darwinian works, because this period has not been treated in detail in English before. We demonstrate an astonishing continuity in topics, methodology, and empirical generalizations despite the shift in Rensch’s views on evolutionary mechanisms. We argue that the continuity in Rensch’s theoretical system can be explained, at last in part, by the guiding role of general methodological principles which underlie the entire system, explicitly or implicitly. Specifically, we argue that Rensch’s philosophy became an asylum for the concept of orthogenesis which Rensch banned from evolutionary theory. Unable to explain the directionality of evolution in terms of empirically based science, he “pre-programmed” the occurrence of human-level intelligence by a sophisticated philosophy combined with a supposedly naturalistic evolutionary biology.

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.     

Grandmothers,hunters and human life history

This paper critiques the competing “Grandmother Hypothesis” and “Embodied Capital Theory” as evolutionary explanations of the peculiarities of human life history traits. Instead, I argue that the correct explanation for human life history probably involves elements of both hypotheses: long male developmental periods and lives probably evolved due to group selection for male hunting via increased female fertility, and female long lives due to the differential contribution women’s complex foraging skills made to their children and grandchildren’s nutritional status within groups provisioned by male hunting.     

Toyama Kametaro and Vernon Kellogg: Silkworm Inheritance Experiments in Japan,Siam, and the United States, 1900–1912

Japanese agricultural scientist Toyama Kametaro’s report about the Mendelian inheritance of silkworm cocoon color in Studies on the Hybridology of Insects (1906) spurred changes in Japanese silk production and thrust Toyama and his work into a scholarly exchange with American entomologist Vernon Kellogg. Toyama’s work, based on research conducted in Japan and Siam, came under international scrutiny at a time when analyses of inheritance flourished after the “rediscovery” of Mendel’s laws of heredity in 1900. The hybrid silkworm studies in Asia attracted the attention of Kellogg, who was concerned with how experimental biology would be used to study the causes of natural selection. He challenged Toyama’s conclusions that Mendelism alone could explain the inheritance patterns of silkworm characters such as cocoon color because they had been subject to hundreds of years of artificial selection, or breeding. This examination of the intersection of Japanese sericulture and American entomology probes how practical differences in scientific interests, societal responsibilities, and silkworm materiality were negotiated throughout the processes of legitimating Mendelian genetics on opposite sides of the Pacific. The ways in which Toyama and Kellogg assigned importance to certain silkworm properties show how conflicting intellectual orientations arose in studies of the same organism. Contestation about Mendelism took place not just on a theoretical level, but the debate was fashioned through each scientist’s rationale about the categorization of silkworm breeds and races and what counted as “natural.” This further mediated the acceptability of the silkworm not as an experimental organism, but as an appropriately “natural” insect with which to demonstrate laws of inheritance. All these shed light on the challenges that came along with the use of agricultural animals to convincingly articulate new biological principles.     

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.     

Macroevolution via secondary endosymbiosis: a Neo-Goldschmidtian view of unicellular hopeful monsters and Darwin’s primordial intermediate form

Seventy-five years ago, the geneticist Richard Goldschmidt hypothesized that single mutations affecting development could result in major phenotypic changes in a single generation to produce unique organisms within animal populations that he called “hopeful monsters”. Three decades ago, Sarah P. Gibbs proposed that photosynthetic unicellular micro-organisms like euglenoids and dinoflagellates are the products of a process now called “secondary endosymbiosis” (i.e., the evolution of a chloroplast surrounded by three or four membranes resulting from the incorporation of a eukaryotic alga by a eukaryotic heterotrophic host cell). In this article, we explore the evidence for Goldschmidt’s “hopeful monster” concept and expand the scope of this theory to include the macroevolutionary emergence of organisms like Euglena and Chlorarachnion from secondary endosymbiotic events. We argue that a Neo-Goldschmidtian perspective leads to the conclusion that cell chimeras such as euglenids and dinoflagellates, which are important groups of phytoplankton in freshwater and marine ecosystems, should be interpreted as “successful monsters”. In addition, we argue that Charles Darwin had euglenoids (infusoria) in mind when he speculated on the “primordial intermediate form”, although his Proto-Euglena-hypothesis for the origin of the last common ancestor of all forms of life is no longer acceptable.     

The evolution of paired fins

Summary The Archipterygium is Gegenbaur’s most lasting contribution to the study of vertebrate limb evolution. This transformational hypothesis of gill arches to limb girdles, rays to fins, and proposal of a vertebrate fin-limb groundplan, is generally treated as a flawed alternative to the more widely accepted lateral fin-fold hypothesis of vertebrate limb evolution. When compared to the phylogenetic distribution and diversity of fins and limbs, both hypotheses fail. Dermal skeletal lateral folds, spines and keels originate repeatedly in vertebrate evolution, but paired fins with girdles originate at pectoral level and are anteroposteriorly restricted. Pelvic fins emerge later in phylogeny; pectoral and pelvic appendages primitively differ. Endoskeletal girdles never exhibit characteristics of gill arches. The emergent sequence of paired fin evolution depends upon phylogenetic hypotheses within which extant agnathan interrelationships are uncertain; positions of jawless fossil fish along the gnathostome stem are insecure; the fossil data set is patchy. However, certain features of the data set are robust. This has prompted a reconsideration of Gegenbaur’s hypothesized arch-girdle relationship, and an iterative homology between scapulocoracoid and extrabranchial cartilages is suggested. No transformation of arch to girdle is necessarily implied, but some signal of developmental relatedness is predicted.     

Overheated Rats,Race, and the Double Gland: Paul Kammerer,Endocrinology and the Problem of Somatic Induction

In 1920, Eugen Steinach and Paul Kammerer reported experiments showing that exposure to high temperatures altered the structure of the gonad and produced hyper-sexuality in “heat rats,” presumably as a result of the increased production of sex hormones. Using Steinach’s evidence that the gonad is a double gland with distinct sexual and generative functions, they used their findings to explain “racial” differences in the sexuality of indigenous tropical peoples and Europeans. The authors also reported that heat induced anatomical changes in the interstitial cells of the gonad were inherited by the heat rats’ descendants. Kammerer used this finding to link endocrinology to his long-standing interest in the inheritance of acquired characteristics. The heat rats supported his hypothesis that the interstitial cells of the double gland were the mechanism of somatic induction in the inheritance of acquired characteristics. The Steinach–Kammerer collaboration, Kammerer’s use of Steinach’s “puberty gland” to explain somatic induction, and his endocrine analysis of symbiosis reveal Paul Kammerer’s late career attempt to integrate endocrinology and genetics with the political ideals of Austrian socialism. With them he developed a bioethics that challenged the growing reliance on race in eugenics and instead promoted cooperation over competition in evolution. I relate his attempt to the controversies surrounding the interstitial cells, to the status of extra-nuclear theories of heredity, and to Kammerer’s commitment to Austromarxist social reforms during the interwar period. I am very grateful for the help of several archivists, including Valerie-Ann Lutz, Roy Goodman and Robert Cox at the American Philosophical Society Library, Arlene Shaner at the New York Academy of Medicine, Shawn Wilson at the Kinsey Institute Library, the staff at the Archives of the University of Vienna, and Yukiko Sakabe at the Austrian Academy of Sciences. Thanks are also due to Andreas Lixl, who gave very helpful advice on German language and to many colleagues, including three anonymous reviewers, Paul Silvia, Alyce Miller, and the late Gilbert Gottlieb, who provided valuable comments as readers of earlier drafts. My discussions with Veronika Hofer have been especially rewarding. The research was funded by National Science Foundation grant #0240151.     

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.     

Science through the internet: Researching, evaluating and citing websites

This article attempts to convey the joys and frustrations of skimming the Internet trying to find relevant information concerning an academic’s work as a scientist, a student or an instructor. A brief overview of the Internet and the “do’s and don’ts” for the neophyte as well for the more seasoned “navigator” are given. Some guidelines of “what works and what does not” and “what is out there” are provided for the scientist with specific emphasis for biologists, as well as for all others having an interest in science but with little interest in spending countless hours “surfing the net”. An extensive but not exhaustive list of related websites is provided.     

“Force-Talk” in Evolutionary Explanation: Metaphors and Misconceptions

The notion of “pressure” as an evolutionary “force” that “causes” evolution is a pervasive linguistic feature of biology textbooks, journal articles, and student explanatory discourse. We investigated the consequences of using a textbook and curriculum that incorporate so-called force-talk. We examined the frequency with which biology majors spontaneously used notions of evolutionary “pressures” in their explanations, students’ definitions and explanations of what they meant when they used pressures, and the structure of explanatory models that incorporated evolutionary pressures and forces. We found that 12–20 percent of undergraduates spontaneously used “pressures” and/or “forces” as explanatory factors but significantly more often in trait gain scenarios than in trait loss scenarios. The majority of explanations using “force-talk” were characterized by faulty evolutionary reasoning. We discuss the conceptual similarity between faulty notions of evolutionary pressures and linguists’ force-dynamic models of everyday reasoning and ultimately question the appropriateness of force-talk in evolution education.