The diving reflex and asphyxia: working across species in physiological ecology.

Beginning in the mid-1930s the comparative physiologists Laurence Irving and Per Fredrik (Pete) Scholander pioneered the study of diving mammals, particularly harbor seals. Although resting on earlier work dating back to the late nineteenth century, their research was distinctive in several ways. In contrast to medically oriented physiology, the approaches of Irving and Scholander were strongly influenced by natural history, zoology, ecology, and evolutionary biology. Diving mammals, they argued, shared the cardiopulmonary physiology of terrestrial mammals, but evolution had modified these basic adaptive processes in extreme ways. In particular, seals' remarkable ability to hold breath, lower metabolism, produce energy anaerobically, and resist asphyxiation, provided a sharp contrast with terrestrial mammals, including humans. This diving physiology was an extreme elaboration of a general regulatory mechanism that allowed seals and other diving mammals to remain active underwater for extended periods. The decrease in heart rate referred to as bradycardia or the "diving reflex" was highly developed in diving mammals, but also found in less developed form in many other organisms faced by asphyxia. It therefore served as a kind of "master switch" for lowering metabolism in diving, hibernation, parturition, drowning, and other physiological responses involving lack of oxygen. Studying bradycardia unified a wide diversity of physiological phenomena, while also providing a context for contrasting the physiological responses of various species, including humans. Conducted in the laboratory and the field, this research served as a bridge between a comparative physiological ecology focused on non-human species and a human-centered general physiology.

Bergmann's Rule, Adaptation, and Thermoregulation in Arctic Animals: Conflicting Perspectives from Physiology, Evolutionary Biology, and Physical Anthropology After World War II.

Bergmann's rule and Allen's rule played important roles in mid-twentieth century discussions of adaptation, variation, and geographical distribution. Although inherited from the nineteenth-century natural history tradition these rules gained significance during the consolidation of the modern synthesis as evolutionary theorists focused attention on populations as units of evolution. For systematists, the rules provided a compelling rationale for identifying geographical races or subspecies, a function that was also picked up by some physical anthropologists. More generally, the rules provided strong evidence for adaptation by natural selection. Supporters of the rules tacitly, or often explicitly, assumed that the clines described by the rules reflected adaptations for thermoregulation. This assumption was challenged by the physiologists Laurence Irving and Per Scholander based on their arctic research conducted after World War II. Their critique spurred a controversy played out in a series of articles in Evolution, in Ernst Mayr's Animal Species and Evolution, and in the writings of other prominent evolutionary biologists and physical anthropologists. Considering this episode highlights the complexity and ambiguity of important biological concepts such as adaptation, homeostasis, and self-regulation. It also demonstrates how different disciplinary orientations and styles of scientific research influenced evolutionary explanations, and the consequent difficulties of constructing a truly synthetic evolutionary biology in the decades immediately following World War II.

Camels, Cormorants, and Kangaroo Rats: Integration and Synthesis in Organismal Biology After World War II.

During the decades following World War II diverse groups of American biologists established a variety of distinctive approaches to organismal biology. Rhetorically, organismal biology could be used defensively to distinguish established research traditions from perceived threats from newly emerging fields such as molecular biology. But, organismal biologists were also interested in integrating biological disciplines and using a focus on organisms to synthesize levels of organization from molecules and cells to populations and communities. Part of this broad movement was the development of an area of research variously referred to as physiological ecology, environmental physiology, or ecophysiology. This area of research was distinctive in its self-conscious blend of field and laboratory practices and its explicit integration with other areas of biology such as ecology, animal behavior, and evolution in order to study adaptation. Comparing the intersecting careers of Knut Schmidt-Nielsen and George Bartholomew highlights two strikingly different approaches to physiological ecology. These alternative approaches to studying the interactions of organisms and environments also differed in important ways from the organismal biology championed by leading figures in the modern synthesis.

The origin and early reception of sequence databases.

Emerging areas of scientific research never arise in a social or intellectual vacuum, but must establish themselves in relation to well-established disciplines. This necessity poses challenges for scientists who must not only create a new disciplinary identity, but must also defend their research from criticism and even condescension from other scientists. The early use of sequence databases provides an excellent case study for examining the challenges facing novel sciences. The need for sequence databases grew out of protein sequencing in biochemistry beginning in the late 1950s. The rapid increase in the number of sequences made databases an attractive resource, but protein biochemists often considered building, managing, and doing research with databases a "second-rate" science. Similarly, computational biologists who used databases and digital computers to study evolutionary phenomena faced criticism from more traditional evolutionary biologists. In retrospect, one can see this early computational biology as laying important foundations for the bioinformatics, molecular evolution, and molecular systematics of today. However, within the context of the 1960s, establishing a scientific identity posed serious challenges for Margaret Dayhoff, Walter Fitch, and Russell Doolittle and other computational biologists who used computers and databases to investigate evolutionary problems.

Waiting for sequences: Morris Goodman, immunodiffusion experiments, and the origins of molecular anthropology.

During the early 1960s, Morris Goodman used a variety of immunological tests to demonstrate the very close genetic relationships among humans, chimpanzees, and gorillas. Molecular anthropologists often point to this early research as a critical step in establishing their new specialty. Based on his molecular results, Goodman challenged the widely accepted taxonomic classification that separated humans from chimpanzees and gorillas in two separate families. His claim that chimpanzees and gorillas should join humans in family Hominidae sparked a well-known conflict with George Gaylord Simpson, Ernst Mayr, and other prominent evolutionary biologists. Less well known, but equally significant, were a series of disagreements between Goodman and other prominent molecular evolutionists concerning both methodological and theoretical issues. These included qualitative versus quantitative data, the role of natural selection, rates of evolution, and the reality of molecular clocks. These controversies continued throughout Goodman's career, even as he moved from immunological techniques to protein and DNA sequence analysis. This episode highlights the diversity of methods used by molecular evolutionists and the conflicting conclusions drawn from the data that these methods generated.