The classical/balance controversy continued along these lines throughout the first half of the sixties. Then, at about the same time that the classical position lost its leading advocate, the balance position received striking new support from Harry Harris, and independently from Dobzhansky's former student Lewontin, and Lewontin's research partner, Jack Hubby.80 These developments served more to reorient the controversy than to end it — and the resulting “neoclassical”/balance controversy is different enough to be grist for another mill.Social policy considerations no (...) longer play a role in keeping the dispute alive. This particular respect in which the issues have changed is, as Diane Paul suggests in her contribution to this volume, as striking as any other.82 There is, however, little danger of our forgetting that this was once much more than just a narrowly technical controversy — the additional social policy issues were far too blatant.However, although blatant, they were by no means the only, or even the most important, issues. In choosing to concentrate on the social policy considerations, I do not mean to suggest that the empirical issues were irrelevant, or simple and straightforward, or otherwise uninteresting. That is by no means the case. What I have tried to show is that there was much more to the classical/balance stalemate than just the empirical underdetermination of the theoretical issues, and that the empirical issues cannot be treated adequately without taking into account the social policy considerations that were involved. Dobzhansky and Muller both appealed to the dangers of misguided social policy that might have resulted from prematurely resolving their controversy in the other's favor. They called for high empirical standards on those grounds, more than once seeking to forestall the resolution of their dispute in this way. (shrink)

The paper describes the context and the origin of a particular debate that concerns the evolution of phenotypic plasticity. In 1965, British biologist A. D. Bradshaw proposed a widely cited model intended to explain the evolution of norms of reaction, based on his studies of plant populations. Bradshaw’s model went beyond the notion of the “adaptive norm of reaction” discussed before him by Dobzhansky and Schmalhausen by suggesting that “plasticity” the ability of a phenotype to be modified by the environment (...) should be genetically determined. To prove Bradshaw’s hypothesis, it became necessary for some authors to identify the pressures exerted by natural selection on phenotypic plasticity in particular traits, and thus to model its evolution. In this paper, I contrast two different views, based on quantitative genetic models, proposed in the mid-1980s: Russell Lande and Sara Via’s conception of phenotypic plasticity, which assumes that the evolution of plasticity is linked to the evolution of the plastic trait itself, and Samuel Scheiner and Richard Lyman’s view, which assumes that the evolution of plasticity is independent from the evolution of the trait. I show how the origin of this specific debate, and different assumptions about the evolution of phenotypic plasticity, depended on Bradshaw’s definition of plasticity and the context of quantitative genetics. (shrink)

Experimental taxonomy was a diverse area of research, and botanists who helped develop it were motivated by a variety of concerns. While experimental taxonomy was never totally a taxonomic enterprise, improvement in classification was certainly one major motivation behind the research. Hall's and Clements' belief that experimental methods added more objectivity to classification was almost universally accepted by experimental taxonomists. Such methods did add a new dimension to taxonomy — a dimension that field and herbarium studies, however rigorous, could not (...) duplicate. Nonetheless, experimental techniques were never completely divorced from traditional taxonomic methods. In practice, all experimental taxonomists employed a combination of descriptive and experimental methods. Most researchers freely acknowledged a debt to traditional taxonomy. Furthermore, the greater rigor of twentieth-century taxonomy was not due entirely to experimentalism. Both the experimental and descriptive aspects of taxonomy were improved by the increased use of quantitative methods, particularly statistics.52From the beginning, a number of experimental taxonomists were interested primarily in classification. But many approached their research from fields other than taxonomy. These botanists were concerned primarily with ecological and genetic problems rather than with classification. There is little indication that they drew a sharp distinction: for example, taxonomic and cytogenetic conclusions were interwoven in Babcock and Stebbins' 1938 study of Crepis 53 (this was even more true of Babcock's final mongraph on the genus, published in 1947). Similarly, the extensive series of monographs, “Experimental Studies on the Nature of Species,” initiated by the Carnegie Institution group in 1940 combined ecological, cytogenetic, and taxonomic conclusions. Indeed, the significance of the major projects completed by experimental taxonomists was largely due to the fact that they were comprehensive studies rather than strictly taxonomic or cytogenetic.In a general sense, the primary motivation behind much of experimental taxonomy was evolutionary. Beginning in the second decade of the century Hall and Clements exhorted taxonomists to take an explicitly evolutionary perspective on research. Hall undoubtedly spoke for the majority of experimental taxonomists when he stated, “If there be anything at all to organic evolution, then taxonomy is dealing with the products of evolution and it is this that gives to taxonomy both its highest mission and its greatest responsibility.”54Aside from a common interest in evolution, however, the theoretical orientations of experimental taxonomists were varied. This diversity is strikingly illustrated by the evolutionary views of members of the Carnegie Institution research group. Experimental taxonomy was initiated by Clements as one aspect of his Lamarckian study of adaptation and speciation. In contrast, Hall's research was inspired by a broad concern for evolutionary problems. Hall rarely referred to specific evolutionary mechanisms; rather, he applied a general conception of evolutionary processes to deduce phylogenetic relationships. His later associates at the Carnegie Institution explicitly dissociated themselves from Clements' theoretical framework. The neo-Darwinian interpretations of adaptation and speciation presented by Clausen, Keck, and Hiesey could hardly have been more different than those of Clements. However, this major shift in theoretical orientation should not obscure significant similarities between the research of Clements and later Carnegie workers. In terms of research problems and methodology, the first volume of “Experimental Studies on the Nature of Species” was an extension of the Clementsian research program. Clausen, Keck, and Hiesey's monograph was the mature discussion of transplant experimentation that Clements had very tentatively initiated during the first decades of the twentieth century. The bond that linked the members of the Carnegie Institution research group to experimental taxonomists in general was one of shared methodology rather than common theoretical orientation. While Clements' evolutionary views were eventually repudiated, his enthusiasm for innovative experimental methods was shared by later workers.The development of experimental taxonomy faced significant problems. During the period 1920–1950 this area of botanical research remained a hybrid discipline. The aims and scope of experimental taxonomy were never articulated in a completely unified manner. Consequently, even among experimental taxonomists, there were disagreements over the relation of their research to other botanical endeavors. Even though experimental taxonomy had close ties with general taxonomy, a number of experimental taxonomists questioned the “taxonomic” nature of their research55. Even to the extent that this hybrid discipline could be identified as a branch of taxonomy, problems arose. Taxonomists, as we have seen, were justifiably skeptical of what appeared to be a rapid influx of untested methods and ideas. Experimental taxonomists were not merely incorporating well-accepted methods from ecology and cytogenetics; during the period 1920–1950 the fields from which experimental taxonomists borrowed were themselves undergoing major theoretical and methodological changes. Despite problems and conflicts, experimental taxonomists did contribute improvements to classification. Furthermore, they made significant contributions to plant ecology and evolutionary genetics.The development of experimental taxonomy indicates that twentieth-century botanists were not necessarily isolated in naturalist and experimentalist camps. The joint session of taxonomists, cytologists, and geneticists at the 1926 International Congress of Plant Sciences indicates communication among specialists fairly early in the century. The papers and commentaries presented during this session do not reveal the hostility and intolerance that supposedly characterized encounters between experimentalists and naturalists. Nor do they suggest incompatible conceptual worlds separating geneticists and taxonomists.Discussions between taxonomists and other specialists were not limited to a single international congress. Particularly during the 1930s discussions among specialists appear to have been fairly widespread. Groups such as the Biosystematists and the Society for the Study of Systematics in Relation to General Biology served as forums for discussion among biologists from a variety of disciplines. The naturalist-experimentalist dichotomy tends to obscure the broad research interests of a number of prominent twentieth-century botanists. Most of the experimental taxonomists cannot be characterized adequately as either naturalists or experimentalists. Traditionally trained taxonomists such as Hall, Keck, and Turrill throughout their careers participated in both experimental and herbarium research. And a number of specialists in fields other than taxonomy took an active interest in taxonomic problems, not necessarily limited to experimental aspects. For example, Anderson suggested a number of innovations to make herbarium collections more amenable to statistical analysis.This historical study of experimental taxonomy indicates a different relationship between experimentalism and taxonomy than that portrayed by the naturalist-versus-experimentalist dichotomy. F. E. Clements originated experimental taxonomy as a revolt against descriptive botany. In retrospect, this revolution was not vigorously waged and was not successfully completed. Experimental taxonomy was never an entirely experimental approach to botanical research. Even the most ardent advocates of experimentalism relied heavily on methods inherited from traditional taxonomy. Moderate exponents of experimental taxonomy stressed the compatibility of experimental methods, field observation, and herbarium techniques. Attempts to fuse cytogenetics, ecology, and taxonomy during the period 1920–1950 resulted in an impressive body of research. However, this fusion constituted neither a repudiation of descriptive botany nor a complete revision of taxonomic theory of practice. (shrink)

The distinction between taxonomic plant geography and ecological plant geography was never absolute: it would be historically inaccurate to portray them as totally divergent. Taxonomists occasionally borrowed ecological concepts, and ecologists never completely repudiated taxonomy. Indeed, some botanists pursued the two types of geographic study. The American taxonomist Henry Allan Gleason (1882–1975), for one, made noteworthy contributions to both. Most of Gleason's research appeared in short articles, however. He never published a major synthetic work comparable in scope or influence to (...) the ecological texts of Clements, Schimper, and Warming.Despite exceptions such as Gleason, most plant geographers throughout the twentieth century have emphasized the distinction between ecological and taxonomic plant geographies. Why have these distinct traditions developed? In his book Geographical Ecology, Robert MacArthur has suggested a psychological explanation for the dichotomy: “Unraveling the history of a phenemenon has always appealed to some people and describing the machinery of the phenomenon to others... The ecologist and physical scientist tend to be machinery oriented, whereas paleontologists and most biogeographers tend to be history oriented.”46Without necessarily rejecting MacArthur's explanation, my study suggests a more complex relationship between taxonomic and ecological plant geographies. At the turn of the century a group of botanists self-consciously defined a new area of botanical research. These ecologists defined their new discipline in opposition to what they believed was a moribund, nineteenth-century, natural-history tradition. They turned from historically oriented, descriptive, taxonomic plant geography to experimental physiology. The new ecological plant geography was to focus on communities rather than on species, on proximate environmental causes rather than on historical explanations, and on physiological experiments rather than on morphological descriptions.As we look back, much of the “revolt from morphology” was rhetorical. Ecologists never completely replaced species as units of distribution, nor did they set geography on an explicitly physiological basis. Indeed, much of early ecological research was, quite simply, descriptive. Plant communities were defined in terms of dominant species, representative life forms, or general physiognomy. The underlying physiological basis for community characteristics was more often assumed than demonstrated by experiments.Despite the fact that ecological plant geography was not a truly physiological specialty, it was significantly different from more traditional taxonomic plant geography. First, ecologists were less explicitly evolutionary in their approach than were taxonomists. Following Darwin, most taxonomic plant geographers viewed distribution in historical terms. In contrast, early ecologists tended to ignore the traditional geographic problems. Most ecologists were skeptical of historical explanations, emphasizing instead the proximate, environmental causes of distribution. While some nineteenth-century biogeographers had studied the correlation between climate and vegetation, twentieth-century ecologists focused much more sharply on the interactions between plant and environment. Plant ecologists did not place biogeography on a physiological basis, but by emphasizing physiology they laid the foundation for a more detailed understanding of adaption. This emphasis on physiology and environmental causation was a second distinguishing characteristic of ecological plant geography. Finally, the idea of the plant community, articulated by Eugenius Warming in 1895, provided ecologists with a unique perspective on the distribution of plants. For early ecologists, the community was more than an assemblage of species; it was an integrated unit. The distribution of these units became the major focus of ecological plant geography. Communities never completely replaced species as geographic units, and the distinction between flora and vegetation was often blurred. Nonetheless, ecologists were innovative in studying the distribution of structurally and functionally integrated groups of plants.In the twentieth century plant geography has occupied an anomalous position in biology. It has not developed into an autonomous discipline, nor has it been incorporated into the developing discipline of ecology. Ecologists and taxonomists have pursued fairly distinct styles of geographic research, with the result that two relatively independent approaches to the study of plant distribution have persisted. (shrink)

A standard norm of reaction (NoR) is a graphical depiction of the phenotypic value of some trait of an individual genotype in a population as a function of an environmental parameter. NoRs thus depict the phenotypic plasticity of a trait. The topological properties of NoRs for sets of different genotypes can be used to infer the presence of (non-linear) genotype-environment interactions. While it is clear that many NoRs are adaptive, it is not yet settled whether their evolutionary etiology should be (...) explained by selection on the mean phenotypic trait values in different environments or whether there are specific genes conferring plasticity. If the second alternative is true the NoR is itself an object of selection. Generalized NoRs depict plasticity at the level of populations or subspecies within a species, species within a genus, or taxa at higher levels. Historically, generalized NoRs have routinely been drawn though rarely explicitly recognized as such. Such generalized NoRs can be used to make evolutionary inferences at higher taxonomic levels in a way analagous to how standard NoRs are used for microevolutionary inferences. (shrink)