Ernst Mayr: Biologist-historian

Ernst Mayr's historical writings began in 1935 with his essay Bernard Altum and the territory theory and have continued up through his monumentalGrowth of Biological Thought (1982) and hisOne Long Argument: Charles Darwin and the Genesis of Modern Evolutionary Thought (1991). Sweeping in their scope, forceful in their interpretation, enlisted on behalf of the clarification of modern concepts and of a broad view of biology, these writings provide both insights and challenges for the historian of biology. Mayr's general intellectual formation was guided by the GermanBildung ideal, with its emphasis on synthetic and comprehensive knowledge. His understanding of how to write history was inspired further by the example of the historian of ideas Arthur Lovejoy. Some strengths and limitations of this approach are explored here through attention to Mayr's treatment of the French biologist J.-B. Lamarck. It is contended that Mayr's contributions to the history of biology are not restricted to his own very substantial historical writings but also include his encouragement of other scholars, his development of an invaluable archive of scientific correspondence, and his insistence that historians who write about evolution and related subjects acquire an adequate understanding of the principles of Darwinian biology.This paper was originally delivered at the biennial meeting of the International Society for the History, Philosophy, and Social Studies of Biology, held in Brandeis in July 1993, in the special session organized by John Greene on Ernst Mayr's contributions to systematics, evolutionary theory, and the history and philosophy of biology. The paper is presented here with only slight modifications of the original, oral presentation. As indicated in the text, a full assessment of Mayr's historical work, including situating that work in the context of Mayr's other work and contemporary developments in the history of science, would require a much more extensive study than I have been able to undertake here.     

Ernst Mayr as community architect: Launching the society for the study of evolution and the journalEvolution

Ernst Mayr's contributions to 20th century biology extend far beyond his defense of certain elements in evolutionary theory. At the center of mid-century efforts in American evolutionary studies to build large research communities, Mayr spearheaded campaigns to create a Society for the Study of Evolution and a dedicated journal,Evolution, in 1946. Begun to offset the prominence ofDrosophila biology and evolutionary genetics, these campaigns changed course repeatedly, as impediments appeared, tactics shifted, and compromises built a growing coalition of support. Preserved, however, were designs to balance the community and journal with careful equation of status and explicit partitioning of responsibilities within the working coalition. Choice terms such as cooperation and unity carried a strong political message. Mayr's editorship ofEvolution provides a superb example of these balancing efforts. The mid-century infrastructural activities described herein also represented aggressive attempts to leverage control across several layers of community. Leaders of these campaigns sought: (1) to promote evolutionary studies as a modernized research discipline and place it at the center of American biology, (2) to promote evolutionary studies within existing disciplines — e.g. systematics, genetics, and paleontology, (3) to foster certain research styles within evolutionary studies, and (4) to emphasize certain solutions to prominent research questions. Throughout, Mayr interjected his priorities, tactics and energy.     

The uncertain foundation of neo-Darwinism: metaphysical and epistemological pluralism in the evolutionary synthesis

The Evolutionary Synthesis is often seen as a unification process in evolutionary biology, one which provided this research area with a solid common theoretical foundation. As such, neo-Darwinism is believed to constitute from this time onward a single, coherent, and unified movement offering research guidelines for investigations. While this may be true if evolutionary biology is solely understood as centred around evolutionary mechanisms, an entirely different picture emerges once other aspects of the founding neo-Darwinists’ views are taken into consideration, aspects potentially relevant to the elaboration of an evolutionary worldview: the tree of life, the ontological distinctions of the main cosmic entities (inert matter, biological organisms, mind), the inherent properties of self-organizing matter, evolutionary ethics, and so on. Profound tensions and inconsistencies are immediately revealed in the neo-Darwinian movement once this broader perspective is adopted. This pluralism is such that it is possible to identify at least three distinct and quasi-incommensurable epistemological/metaphysical frameworks as providing a proper foundation for neo-Darwinism. The analysis of the views of Theodosius Dobzhansky, Bernhard Rensch, and Ernst Mayr will illustrate this untenable pluralism, one which requires us to conceive of the neo-Darwinian research agenda as being conducted in more than one research programme or research tradition at the same time.     

On the combined analysis of proximate and ultimate aspects in diel vertical migration (DVM) research

Although evolutionary ecologists agree that proximate and ultimate aspects are two sides of one coin, they are seldom interested in studies on physiological and behavioural mechanisms at the base of ecological phenomena. Nevertheless, these mechanisms are objects of selection and evolved to realise adaptive significances. This paper is a plea to bring both fields closer together, and, by means of an example of Diel Vertical Migration of Daphnia, some proximate and ultimate aspects are discussed. It is argued that light changes, not fish kairomone, is the primary cause for an individual to swim downwards at dawn and upwards at dusk. However, what is called a causal factor might differ when ecosystems or individuals are studied. In addition, causality in ecology is not simple, and has the character of a `set of necessary conditions'. To illustrate the importance of proximate analyses in DVM, two basic response mechanisms are discussed: Photobehaviour system 1 and 2. The physiological character of these systems leads to a fixed type of migration or to a phenotypically induced DVM, respectively. The adaptive significance of the first might be a reduction of the hazardous effects of UV radiation and of the second a lowering of mortality due to visually hunting predators.     

The importance of the “Gegenbaur school” for German morphology

Summary The school Carl Gegenbaur cultivated at Heidelberg (1873–1901) was critical to the history of German morphology in multiple ways. During and after his lifetime, the school carried out detailed comparative anatomical and embryological investigations in an evolutionary framework, thereby contributing substantially to the project of vertebrate evolutionary morphology. Its members also defended their mentor when his ideas came under attack. After his death, they labored to perpetuate his program and his memory in the increasingly unwelcoming environment of medical education and research. While the senior members of the school did this largely through institutional means-seeking to place Gegenbaur sympathizers in academic and editorial positions-its junior members absorbed some of the criticisms of the school to develop a modified, more functional approach to evolutionary morphology. The school thus kept the Gegenbaur program alive and active in the German-speaking lands for over fifty years. This paper is drawn largely from Nyhart (1995), esp. chapter 7 (supported by NSF award no. 8910873). Information not otherwise documented derives from this book.     

How evolutionary biology challenges the classical theory of rational choice

A fundamental philosophical question that arises in connection with evolutionary theory is whether the fittest patterns of behavior are always the most rational. Are fitness and rationality fully compatible? When behavioral rationality is characterized formally as in classical decision theory, the question becomes mathematically meaningful and can be explored systematically by investigating whether the optimally fit behavior predicted by evolutionary process models is decision-theoretically coherent. Upon investigation, it appears that in nontrivial evolutionary models the expected behavior is not always in accord with the norms of the standard theory of decision as ordinarily applied. Many classically irrational acts, e.g. betting on the occurrence of one event in the knowledge that the probabilities favor another, can under certain circumstances constitute adaptive behavior. One interesting interpretation of this clash is that the criterion of rationality offered by classical decision theory is simply incorrect (or at least incomplete) as it stands, and that evolutionary theory should be called upon to provide a more generally applicable theory of rationality. Such a program, should it prove feasible, would amount to the logical reduction of the theory of rational choice to evolutionary theory.     

Cellular aging beyond cellular senescence: Markers of senescence prior to cell cycle arrest in vitro and in vivo

The field of research on cellular senescence experienced a rapid expansion from being primarily focused on in vitro aspects of aging to the vast territories of animal and clinical research. Cellular senescence is defined by a set of markers, many of which are present and accumulate in a gradual manner prior to senescence induction or are found outside of the context of cellular senescence. These markers are now used to measure the impact of cellular senescence on aging and disease as well as outcomes of anti‐senescence interventions, many of which are at the stage of clinical trials. It is thus of primary importance to discuss their specificity as well as their role in the establishment of senescence. Here, the presence and role of senescence markers are described in cells prior to cell cycle arrest, especially in the context of replicative aging and in vivo conditions. Specifically, this review article seeks to describe the process of “cellular aging”: the progression of internal changes occurring in primary cells leading to the induction of cellular senescence and culminating in cell death. Phenotypic changes associated with aging prior to senescence induction will be characterized, as well as their effect on the induction of cell senescence and the final fate of cells reviewed. Using published datasets on assessments of senescence markers in vivo, it will be described how disparities between quantifications can be explained by the concept of cellular aging. Finally, throughout the article the applicational value of broadening cellular senescence paradigm will be discussed.     

Integrating cellular senescence with the concept of damage accumulation in aging: Relevance for clearance of senescent cells

Understanding the aging process and ways to manipulate it is of major importance for biology and medicine. Among the many aging theories advanced over the years, the concept most consistent with experimental evidence posits the buildup of numerous forms of molecular damage as a foundation of the aging process. Here, we discuss that this concept integrates well with recent findings on cellular senescence, offering a novel view on the role of senescence in aging and age‐related disease. Cellular senescence has a well‐established role in cellular aging, but its impact on the rate of organismal aging is less defined. One of the most prominent features of cellular senescence is its association with macromolecular damage. The relationship between cell senescence and damage concerns both damage as a molecular signal of senescence induction and accelerated accumulation of damage in senescent cells. We describe the origin, regulatory mechanisms, and relevance of various damage forms in senescent cells. This view on senescent cells as carriers and inducers of damage puts new light on senescence, considering it as a significant contributor to the rise in organismal damage. Applying these ideas, we critically examine current evidence for a role of cellular senescence in aging and age‐related diseases. We also discuss the differential impact of longevity interventions on senescence burden and other types of age‐related damage. Finally, we propose a model on the role of aging‐related damage accumulation and the rate of aging observed upon senescent cell clearance.     

The ultimate and proximate mechanisms driving the evolution of long tails in forest deer mice

Understanding both the role of selection in driving phenotypic change and its underlying genetic basis remain major challenges in evolutionary biology. Here, we use modern tools to revisit a classic system of local adaptation in the North American deer mouse, Peromyscus maniculatus, which occupies two main habitat types: prairie and forest. Using historical collections, we find that forest‐dwelling mice have longer tails than those from nonforested habitat, even when we account for individual and population relatedness. Using genome‐wide SNP data, we show that mice from forested habitats in the eastern and western parts of their range form separate clades, suggesting that increased tail length evolved independently. We find that forest mice in the east and west have both more and longer caudal vertebrae, but not trunk vertebrae, than nearby prairie forms. By intercrossing prairie and forest mice, we show that the number and length of caudal vertebrae are not correlated in this recombinant population, indicating that variation in these traits is controlled by separate genetic loci. Together, these results demonstrate convergent evolution of the long‐tailed forest phenotype through two distinct genetic mechanisms, affecting number and length of vertebrae, and suggest that these morphological changes—either independently or together—are adaptive.     

The role of the amateur in mycology—what would we do without them!

The amateur's role in the history of mycology will be traced from its early roots deeply based in the collection of fungi for food to the amateur's present day activities. Attention will be drawn to the most important key amateur figures studying fungi solely or as a wider part of natural history in the eighteenth and nineteenth centuries, and the early part of the present century. How their work influenced the development of mycology will be demonstrated. It is often forgotten that the authorities attached to fungal names, including those we use in mycology on a daily basis, often belong to amateurs. The true professions of these amateurs, the net-work of correspondents they developed, etc. will be revealed and discussed in the context of an overall understanding of fungi as organisms. In addition the formation of mycological societies, and how they have become a focus of amateur activity and a source of accurate and disciplined information valuable to professional scientists will be plotted. The kind of work undertaken by amateurs I refer to will be demonstrated by documenting the activities of the British Mycological Society which celebrated its centenary in 1996. The continuing work of this band of workers and their counterparts throughout the world in the closing years of the millennium will be described. It is argued that they are nationally important and necessary resources, despite many governments or their advisers, especially the most influential ones, being under the delusion that systematics is not cutting edge science. It is also emphasized that amateurs will have an even more important role to play in the future as custodians of knowledge—until hopefully opinions change. The essential points of this paper were addressed to the 42nd Annual Meeting of the Mycological Society of Japan held at Kyoto University, Kyoto, on 16–17, May 1998.     

Emerging patterns of genetic variation in the New Zealand endemic scallop Pecten novaezelandiae

Both historical and contemporary processes influence the genetic structure of species, but the relative roles of such processes are still difficult to access. Population genetic studies of species with recent evolutionary histories such as the New Zealand endemic scallop Pecten novaezelandiae (<1 Ma) permit testing of the effects of recent processes affecting gene flow and shaping genetic structure. In addition, studies encompassing the entire distributional range of species can provide insight into colonization processes. Analyses of genetic variation in P. novaezelandiae (952 individuals from 14 locations, genotyped at 10 microsatellite loci) revealed a weak but significant regional structure across the distributional range of the species, as well as latitudinal gradients of genetic diversity and differentiation: estimates of migration rates supported these patterns. Our results suggest that the observed genetic structure and latitudinal gradients reflect a stepping‐stone model of colonization (north to south) and emerging divergence of populations as a result of ongoing limitations to gene flow and insufficient time to reach migration–drift equilibrium. The low levels of interpopulation and interregional genetic differentiation detected over hundreds of kilometres reflect the recent evolutionary history of P. novaezelandiae and stand in contrast to patterns reported for other evolutionary older species at the same spatial scale. The outcomes of this study contribute to a better understanding of evolutionary processes influencing the genetic variation of species and provide vital information on the genetic structure of P. novaezelandiae.     

The principle of complementarity in the design of reserve networks to conserve biodiversity: A preliminary history

Explicit, quantitative procedures for identifying biodiversity priority areas are replacing the often ad hoc procedures used in the past to design networks of reserves to conserve biodiversity. This change facilitates more informed choices by policy makers, and thereby makes possible greater satisfaction of conservation goals with increased efficiency. A key feature of these procedures is the use of the principle of complementarity, which ensures that areas chosen for inclusion in a reserve network complement those already selected. This paper sketches the historical development of the principle of complementarity and its applications in practical policy decisions. In the first section a brief account is given of the circumstances out of which concerns for more explicit systematic methods for the assessment of the conservation value of different areas arose. The second section details the emergence of the principle of complementarity in four independent contexts. The third section consists of case studies of the use of the principle of complementarity to make practical policy decisions in Australasia, Africa, and America. In the last section, an assessment is made of the extent to which the principle of complementarity transformed the practice of conservation biology by introducing new standards of rigor and explicitness.     

The interaction of multiple environmental stressors affects adaptation to a novel habitat in the natterjack toad Bufo calamita

The potential to adapt to novel environmental conditions is a key area of interest for evolutionary biology. However, the role of multiple selection pressures on adaptive responses has rarely been investigated in natural populations. In Sweden, the natterjack toad Bufo calamita inhabits two separate distribution areas, one in southernmost Sweden and one on the west coast. We characterized the larval habitat in terms of pond size and salinity in the two areas, and found that the western populations are more affected by both desiccation risk and pond salinity than the southern populations. In a common garden experiment manipulating salinity and temperature, we found that toads from the west coast populations were locally adapted to shorter pond duration as indicated by their higher development and growth rates. However, despite being subjected to higher salinity stress in nature, west coast toads had a poorer performance in saline treatments. We found that survival in the saline treatments in the west coast populations was positively affected by larger body mass and longer larval period. Furthermore, we found negative genetic correlations between body mass and growth rate and their plastic responses to salinity. These results implicate that the occurrence of multiple environmental stressors needs to be accounted for when assessing the adaptive potential of organisms and suggest that genetic correlations may play a role in constraining adaptation of natural populations.     

The adaptive legacy of human evolution: A search for the environment of evolutionary adaptedness

The growth of evolutionary psychology has led to renewed interest in what might be the significant evolutionary heritage of people living today, and in the extent to which humans are suited to a particular adaptive environment—the EEA. The EEA, though, is a new tool in the battery of evolutionary concepts, and it is important both that it is scrutinized for its utility, and that the actual reconstructions of the environments in which humans and hominids evolved are based on sound palaeobiological inference and an appropriate use of the phylogenetic context of primate evolution.     

The determinants of the molecular substitution process in turtles

Neutral rates of molecular evolution vary across species, and this variation has been shown to be related to biological traits. One of the first patterns to be observed in vertebrates has been an inverse relationship between body mass (BM) and substitution rates. The effects of three major life‐history traits (LHT) that covary with BM – metabolic rate, generation time and longevity (LON) – have been invoked to explain this relationship. However, most of the theoretical and empirical evidence supporting this relationship comes from endothermic vertebrates, that is, mammals and birds, in which the environmental conditions, especially temperature, do not have a direct impact on cellular and molecular biology. We analysed the variations in mitochondrial and nuclear rates of synonymous substitution across 224 turtle species and examined their correlation with two LHT (LON and BM) and two environmental variables [latitude (LAT) and habitat]. Our analyses indicate that in turtles, neutral rates of molecular evolution are hardly correlated with LON or BM. Rather, both the mitochondrial and nuclear substitution rates are significantly correlated with LAT – faster evolution in the tropics – and especially so for aquatic species. These results question the generality of the relationships reported in mammals and birds and suggest that environmental factors might be the strongest determinants of the mutation rate in ectotherms.     

关帝山林区退化灌木林转变为华北落叶松林对生态系统碳储量的影响

针对我国大量灌木林出现退化而宜林地又日益减少的现状,在适宜种植乔木的地区,将退化灌木林转变为乔木林被认为是一种可行的植被恢复方式。以关帝山林区退化灌木次生林转变而成的不同林龄（10、18、23、27年和35年）华北落叶松（Larix principis-rupprechtii Mayr.）林为研究对象,并以相邻的退化灌木次生林为对照,探究这种转变对生态系统碳储量及其组分的影响,将为我国开展造林/再造林、林业碳汇项目等工作提供科学依据和数据支撑。与灌木林相比,造林初期的生态系统碳储量及其组分均出现不同程度的下降。10年生华北落叶松林的生态系统碳储量相对于灌木林显著下降了32.9%（P < 0.05）,但并非所有组分的下降都显著（P < 0.05）。植被碳储量下降34.7%,其植被地上和地下碳储量分别下降5.4%和70.9%,但只有植被地下碳储量是显著减少的（P < 0.05）;凋落物碳储量下降42.8%,但并不显著（P=0.71）;土壤有机碳储量（0-50 cm）显著下降32.6%（P < 0.05）,其不同土层（0-10、10-30 cm和30-50 cm）的碳储量也都出现显著减少（P < 0.05）。林龄从10年到35年,华北落叶松林生态系统碳储量增加了1.6倍,植被及其组成（地上和地下）、凋落物、土壤有机碳及其不同土层（0-10、10-30 cm和30-50 cm）等的碳储量也随之不断增加,从而使得生态系统碳储量及其组分逐渐达到并全面超过灌木林。但是,不同组分要达到灌木林的碳储量水平,需要的时间存在较大差异：土壤有机碳库 > 植被地下碳库 > 植被地上碳库,其中深层土壤有机碳 > 表层土壤有机碳（0-10 cm）。     

The synthetic theory of evolution: general problems and the German contribution to the synthesis

Summary A metatheoretical and historiographical re-analysis of the Evolutionary Synthesis (the process) and the Synthetic Theory (the result) leads to the following conclusion: The Synthetic Theory is not a reductionistic, but rather a structuralistic theory with a limited range of relevant hierarchical levels. Historically the Synthesis was not a sudden event but a rational long-term project carried out between 1930 and 1950 by a large number of biologists in several countries. In the second part of our paper the contributions of several German biologists to the Synthesis are analyzed.