Evolution, Science, and Society: a "white paper" on behalf of the field of evolutionary biology [working draft - last updated 4 June 1997] [EXECUTIVE SUMMARY] http://www.rci.rutgers.edu/~ecolevol/execsumm.html Prepared by: Editorial Chair: Douglas J. Futuyma (SUNY-Stony Brook) Organizational Chair: Thomas R. Meagher (Rutgers University) Steering Committee: Michael J. Donoghue (Harvard University) Charles H. Langley (University of California-Davis) Linda Maxson (University of Tennessee) Working Group: Albert F. Bennett (University of California-Irvine) May R. Berenbaum (University of Illinois) H. Jane Brockmann (University of Florida) Marcus W. Feldman (Stanford University) Walter M. Fitch (University of California-Irvine) Laurie R. Godfrey (University of Massachusetts) James Hanken (University of Colorado) David Hillis (University of Texas) David Jablonski (University of Chicago) Mary-Claire King (University of Washington) Carol B. Lynch (University of Colorado) Leslie Real (Indiana University) Margaret A. Riley (Yale University) J. John Sepkoski, Jr. (University of Chicago) Vassiliki Betty Smocovitis (University of Florida) Under the sponsorship of: The A. P. Sloan Foundation The National Science Foundation Endorsed by the following scientific societies: American Society of Naturalists Animal Behavior Society (PENDING) Ecological Society of America (PENDING) Genetics Society of America (PENDING) Paleontological Society (PENDING) Society for Molecular Biology and Evolution (PENDING) Society of Systematic Biologists Society for the Study of Evolution (PENDING) INTRODUCTION The science of evolutionary biology is the study of the history of life and the processes that lead to its unity and diversity. Evolution occupies a central position in the biological sciences. Although sometimes controversial outside of scientific disciplines, evolution has both been fully documented by scientific research and is fundamentally important for understanding biological systems, including ourselves. Moreover, the methods developed to study evolutionary processes and history -- ranging from the molecular to the paleontological - have proved indispensable in other fields. Academic and research institutions in the United States and elsewhere have acknowledged the importance of this field by establishing departments that focus on evolutionary biology. The importance of evolutionary biology is reflected in the rapid growth of journals related to evolution and in the increasing application of evolutionary principles in areas such as human health, agriculture, environmental remediation, behavior, and other aspects of human culture. Public understanding of evolutionary biology has grown through the efforts of museums, zoological parks, and botanical gardens, as well as a profusion of popular books and articles. Fueled by accelerating advances in molecular techniques, information processing, and other technological innovations, the biological sciences are on a steeper learning curve than ever before, and biological knowledge in the next few decades will increasingly affect our lives. Evolutionary biology draws on these technological advances, and in turn contributes to other biological disciplines, ranging from molecular biology to ecology. In order for evolutionary biology to contribute fully to basic biological science and to societal concerns, it is necessary that research and education in critical areas be adequately supported, and that the methods and results of evolutionary research be integrated with other disciplines both within and outside of biology. It is necessary, moreover, that evolutionary research be applied to societal problems, and that the implications of evolutionary research be included in the education of a scientifically informed citizenry. This document is addressed to decision-makers responsible for guiding basic and applied scientific research and for developing educational curricula. It is also addressed to anyone interested in understanding the current and potential accomplishments of evolutionary biology. Its major goals are: * to describe our present understanding of evolution and the major intellectual accomplishments of evolutionary biology; * to identify major questions and challenges in which progress in evolutionary science can be expected in the near future; * to describe past and expected contributions of evolutionary biology both to other sciences and to social needs in areas such as health science, agriculture, and environmental science; and * to suggest ways in which progress can be facilitated in basic research, in applications of evolutionary biology to social needs, and in biological science education. To further these goals, delegates from eight major professional scientific societies in the United States whose subject matter substantially includes evolution have prepared this document. Contributions have also been made by other specialists in various topics. A draft of the document was revised in light of feedback elicited from the community of evolutionary biologists in the United States and by making the draft public at scientific meetings and on the Worldwide Web. Although full agreement cannot be expected on every detail and point of emphasis, the major points and conclusions in the following pages represent a large majority of professional evolutionary biologists. WHAT IS EVOLUTION? Biological evolution consists of change in the hereditary characteristics of groups of organisms over the course of generations. In a long-term perspective, evolution is the descent, with modification, of different lineages from common ancestors. Thus the history of evolution has two major components: branching of lineages and change within lineages. The evidence that all organisms have descended, with modification, from common ancestors is so overwhelming that most biologists consider evolution a fact, not a theory. WHAT IS EVOLUTIONARY BIOLOGY? Evolutionary biology seeks to explain the diversity of life: the variety of organisms and their characteristics, and their changes over time. The two encompassing goals of evolutionary biology are: * to discover the history of life on earth; * to understand the causal processes of evolution. Insights achieved through efforts to meet these goals greatly enhance our understanding of biological systems. As such, evolutionary biologists often work at the interface of multiple sub-disciplines of biology and have developed subject areas such as behavioral evolution, evolutionary developmental biology, evolutionary ecology, evolutionary genetics, evolutionary systematics, and molecular evolution. The subdisciplines of evolutionary biology also have formed direct links with fields such as statistics, economics, geology, anthropology, and psychology. Evolutionary biology offers several important, distinctive perspectives on biology. These include emphasis on the interplay between chance and necessity as conflicting agents in biological dynamics, on variation as an inherent feature of biological systems, and on the importance of biological diversity. Chance and necessity refer to the interplay between random events and predictable, nonrandom agents of change, such as natural selection. This interplay, well understood in theory, has been extensively described by empirical observation as well. Variation is a key concept, since evolutionary change ultimately depends on the differential success of different genetic lineages. Understanding the importance undermined the notion, prevalent until the nineteenth century, that species are fixed and immutable. In our own time, it has disproved theories of fixed racial differences and has shown that all features of organisms, even their fundamental mechanisms of inheritance, are variable and subject to evolutionary change. Finally, the ultimate consequence of variability and evolutionary divergence is biological diversity. Evolutionary biology provides the key to understanding principles governing the origin and extinction of species. HOW IS EVOLUTION STUDIED? Science is a process of understanding. Consequently, any scientific discipline is characterized by the tools it uses to achieve understanding. Evolutionary biology draws on a wide range of methodologies, but there is a series of conceptual approaches that characterize the field itself. Methods for understanding the history of evolution include paleontological observations of the fossil record and categorization and classification of variation among living organisms. Differences and similarities among species, in anatomy, DNA, and other features are analyzed by modern statistical methods of phylogenetic inference. These enable us to estimate the genealogical (phylogenetic) relationships among species and the sequence by which their characteristics evolved. Studies of evolutionary processes employ observation and experimentation, mostly on living organisms. Evolutionary changes are often observed directly and studied experimentally by placing populations of rapidly reproducing organisms in new environments and measuring changes that transpire over the course of generations. Studies of genetic variation within species, including variation in DNA sequences, enable us to characterize mutation, genetic drift, natural selection, and other causes of evolution. Such studies draw both on natural populations of many species and on laboratory populations of model organisms such as fruit flies, bacteria, and Arabidopsis plants, which provide detailed information on the genetic basis of evolutionary change. Many hypotheses are tested by comparing patterns of variation among organisms to the different patterns predicted by contrasting hypotheses or theories. For instance, the "comparative method" tests hypotheses about the adaptive value of various characteristics by contrasting features of species that have adapted to different environments. Finally, evolutionary biology is rich in theory, and sophisticated mathematical models and analytical tools are frequently brought to bear on evolutionary issues. CONTRIBUTIONS OF EVOLUTIONARY BIOLOGY The contributions of evolutionary biology can be viewed on three levels: accomplishments of the field with respect to its own goals, insights gained in other fields of biology and science in general that can be traced to evolutionary biology, and benefits to society at large that can be attributed to evolutionary biology. Achievements in the study of evolution Among many accomplishments in studying the history and processes of evolution, evolutionary biologists have * unequivocally established that all organisms have evolved from common ancestors over the course of more than 3.5 billion years; * successfully developed methods of inferring phylogenetic, or genealogical, relationships among organisms; * documented the tempo and mode of evolution; * described patterns of diversification and extinction in the fossil record; * devised and validated a quantitative theory of fundamental evolutionary processes; * established that populations are highly genetically variable; * directly observed and studied the process of evolution; * clarified the mechanisms by which new species arise; * documented many different forms of natural selection; * developed and tested theories of natural selection that explain the evolution of many puzzling characteristics, such as cooperative behavior and senescence; * elucidated processes of coevolution of interacting species; * made rapid progress in understanding mechanisms, rates, and causes of evolution at the molecular level; * achieved greater understanding of the developmental basis of the evolution of complex characters; * elucidated many aspects of human evolution. Contributions to other biological disciplines Among the biological sciences more broadly, evolution is widely viewed as central to biological understanding. Many biologists in diverse fields regard at least a portion of what they do as evolutionary. Among specific recent accomplishments to which evolutionary biology has contributed are the following: Molecular biology. Evolutionary approaches have contributed insight into the function and structure of molecular processes within cells. Examples include elucidation of the significance of higher-order structure in rRNA, the origins and significance of codon usage bias in gene expression, and the significance of different categories of DNA, including sequences that are expressed as codes for known proteins and sequences that are not expressed directly but that may have other impacts on genome function. Evolutionary research thus points the way to research on fundamental molecular mechanisms. Evolutionary studies have also established the biological importance of phenomena, such as gene duplication, that molecular biologists study in model laboratory organisms. Developmental biology. Similarities among the embryos of species that differ radically as adults were among Darwin's chief sources of evidence for evolution. A resurgence in interaction between developmental biology and evolutionary biology is now under way, in part because of renewed focus on development by evolutionary biologists, and in part because of comparisons among species of the genes that have been identified as playing critical roles in development. For instance, homologous genes in organisms as different as insects and mammals play similar developmental roles in some instances, and surprisingly different roles in other cases. Such studies help to identify the developmental functions of genes, and lead to a deeper understanding of developmental phenomena. Physiology and Morphology. Evolutionary biology has long influenced the study of morphology and physiology in animals and plants, and can make many other contributions that are only now being developed. Some of these contributions can affect human physiology, including related areas such as clinical psychology. The logical perspectives, methods, and comparative data of evolutionary biology can advance our understanding of functional anatomy and physiological mechanisms, and their applications to areas such as medicine, agriculture, and veterinary science. Evolutionary physiology and morphology include comparisons of features among species, experimental studies on the evolution of physiological and morphological characters, and phylogenetic and population genetic analyses of physiological and morphological evolution. Neurobiology and behavior. From its inception, the field of animal behavior has had a strong evolutionary base, for its goals have included understanding the evolutionary origin of behavioral traits and their adaptiveness. The evolutionary study of animal behavior has joined with comparative psychology in several research areas, such as the study of learning and the search for adaptive mechanisms in human cognitive processes. Although neurobiologists recognize that the mechanisms they study are adaptations, they generally do not study behavioral mechanisms in expressly evolutionary terms. There are, however, some notable exceptions. The adaptive perspective of comparative evolutionary biology has led to the discovery of interesting sensory and behavioral mechanisms in a variety of species. Comparative studies, based on understanding the adaptive requirements of different species, thus lead to new understanding of behavioral mechanisms. Contributions to societal needs Finally, evolutionary biology has a long history and potentially a bright future in terms of its ability to address pressing societal needs. The following are areas in which evolutionary biology has already made particularly strong contributions: Human Health and Medicine. Evolutionary biology has been central to understanding human genetic diseases, to tracing the origins and epidemiology of infectious diseases, to describing the evolution of antibiotic resistance in pathogenic microorganisms, and to interpreting normal physiological functions and dietary needs as the consequence of long-term evolution. Methods developed by evolutionary geneticists have played an important role in mapping defective human genes and in genetic counseling. Agriculture and Natural Resources. Principles of plant and animal breeding strongly parallel natural evolutionary mechanisms; and there is a rich history of interplay between evolutionary biology and breeding science. Evolutionary insights have a clear role to play in understanding the ongoing evolution of various pathogens and insect pests, including evolution of resistance to pest-control measures. Methods of evolutionary genetics are used to identify different gene pools of commercially important fish and other organisms, their migration routes, and differences in their physiological and life history characteristics. Evolutionary biology has much to contribute to genetic engineering, both by predicting and analyzing organisms' responses to modification of specific genes and by helping to assess the risks risk of planned introductions of genetically engineered organisms. Finding Useful Natural Resources. Many thousands of natural products are used in medicine, food production and processing, cosmetics, biotechnology, pest control, and industry, but millions of other natural products have yet to be screened or even discovered. Evolutionary principles aid in targeted, rather than random, search, by predicting chemical adaptations to environmental selection pressures, and by identifying organisms related to those that have already yielded useful natural products. Environment and Conservation. Evolutionary insights are of importance to both conservation and management of renewable resources. For example, population genetic tools are frequently used to assess the genetic structure of rare or endangered species as a means of determining appropriate conservation measures. Studies of the genetic composition of wild relatives of crop species can be used to determine potentially useful new genes that could be transferred into cultivated species. Studies of wild plants' adaptations to polluted or degraded soils contribute to reclamation of degraded land. Applications beyond Biology. There have long been healthy interactions between evolutionary biology and other analytical fields, notably statistics and economics. Some of the basic tools in statistics, including analysis of variance and path analysis, were originally developed by evolutionary biologists. Along the same lines, evolutionary algorithms, which employ maximization criteria designed to mimic natural selection in biological systems, are currently showing great potential in computer and systems applications. Understanding Humanity. Evolutionary biology has contributed greatly to our understanding of ourselves, by describing our origins and relationship to other living things and the history and significance of variation within and among different groups of people. Evolutionary anthropologists, psychologists, and biologists have advanced hypotheses (controversial even within these fields) on the biological bases for human culture and behavior. In addition, the evolutionary framework for understanding humanity has had a profound impact on literature, the arts, philosophy, and other areas of the humanities. Conclusions Evolutionary biology plays a critical role in modern biology. It provides causal explanations, based on history and on processes of genetic change and adaptation, for the full sweep of biological phenomena, ranging from the molecular to the ecological. Thus, evolutionary biology allows us not only to determine how and why organisms have become the way they are, but also what processes are acting now to modify or change organisms in the future. The comprehensive conceptual framework, mathematical theory, empirical tradition, and other tools of evolutionary biology can be brought to bear on almost every question in biology. Evolutionary biology is a mature scientific discipline that contributes to both fundamental understanding and human uses of biological systems. FUTURE CONTRIBUTIONS OF EVOLUTIONARY BIOLOGY To an increasing extent, researchers in molecular and developmental biology, physiology, ecology, animal behavior, psychology, anthropology, and other disciplines have adopted the methods, principles, and concepts of evolutionary biology as a framework for research and understanding. Likewise, applied research in forestry, agriculture, fisheries, human genetics, and other areas has increasingly attracted scientists trained in evolutionary biology. Evolutionary biologists have expanded their vision, addressing both basic questions throughout the biological disciplines and problems posed by society's needs. As a result of both the rapid growth of this "evolutionary work force" and of technological advances in areas such as molecular methodology, computing, and information processing, progress in evolutionary biology and related areas is more rapid now than ever before. The twenty-first century promises to be "the age of biology," in which evolutionary biology will unquestionably play a prominent, indispensable role. We are confident that given the necessary support in education and research, the evolutionary disciplines are poised to make ever greater contributions to fundamental and applied knowledge. Basic science In basic science, we stand at the threshold * of more fully documenting biodiversity and describing the relationships among organisms; * of analyzing the history of life in greater depth than ever before; * of more fully discovering and explaining processes of evolution at the molecular level; * of analyzing the molecular and genetic basis of characteristics that vary within and among species; * of understanding how developmental mechanisms evolve and give rise to new phenotypic characteristics; * of elucidating the processes that both cause and constrain adaptations in physiology, endocrinology, and anatomy; * of contributing to a deeper understanding of the adaptive meaning and the mechanisms of behavior; * of developing a predictive theory of coevolution between species, such as pathogens, parasites, and their hosts, and of the impact of coevolution on populations and ecological communities. Applied science In the applied realm, evolutionary biologists are continuing to embrace their social responsibility, and are becoming increasingly aware of ways in which their discipline can help * to understand and combat genetic and infectious disease; * to understand human physiological adaptations to stresses, pathogens, and other causes of ill health; * to understand the origin and dynamics of infectious disease organisms; * to combat the evolution of antibiotic resistance by microbial pathogens; * to improve crops and mitigate damage from pathogens, insects, and weeds; * to provide tools for analyzing human genetic diversity as it applies to health, law, and the understanding of human behavior; * to aid responsible economic use and development of biological resources; * to remedy damage to the environment; * to predict consequences of global and regional environmental change; * to conserve biodiversity, potentially faced with the most massive extinction in the last 65 million years. RECOMMENDATIONS In view of its maturity as a field and its centrality to the biological sciences, we call for the following mechanisms to realize the potential of evolutionary biology we propose: * a cross agency review of funding for evolutionary biology, perhaps conducted by the National Academy of Sciences, to determine where such funding comes from and to make recommendations for its coordination [There is at present no overview of evolutionary biology research at the federal level. Similar analytical overviews have resulted in cross agency funding initiatives for areas such as plant biology and global change; and a cross agency funding initiative for evolutionary biology may be appropriate as well.]; * enhanced support for evolutionary biology through existing or emerging channels, including more support for postdoctoral positions, mid-career opportunities, graduate training grants, and research grants [Indeed, increased funding of evolutionary biology initiatives may be a means by which existing agencies could meet their missions more fully and rationally.]; * that opportunities be identified to capitalize on evolutionary biology as an organizing and integrating principle [Such efforts could facilitate the interdisciplinarity of the funded research. In terms of addressing societal needs, these efforts should include outreach to industry. Specific opportunities could be explored through workshops and plenary sessions at scientific society meetings that make connections among subdisciplines and between academic and industrial research/applications. Examples of appropriate settings for such workshops include the Discussion sessions at the ESA meetings or the Science Innovation panels at the AAAS meetings.]; and * establishment of a National Committee on Evolutionary Biology, an electronically linked virtual committee designed as a rapid response network to provide a conduit for input and assistance from the scientific community into federal agencies, the news media and other entities requiring information from the scientific community [Such a Committee could be developed through a consortium of scientific societies under the leadership of a specific society, such as the American Society of Naturalists or the Society for the Study of Evolution, and its membership could be appointed and structured in much the same way as existing journal editorial boards.]. In view of the many applications of evolutionary research to the biological disciplines and to practical social needs, we urge evolutionary biologists: * to communicate to Federal agencies, and to other institutions that support basic or applied research, the relevance of evolutionary biology to these institutions' missions; * to direct their research not only to basic, but also to applied goals; and * to integrate into their educational activities the relevance of evolutionary biology to societal needs. In view of the relevance of evolution to human endeavors and of the overall importance of scientific literacy, we urge that major efforts be made to strengthen curricula in primary and secondary schools as well as in colleges and universities with respect to modern evolutionary biology. We call for: * support of mid-career training of school teachers in evolutionary biology; * greater emphasis on evolution in the curricula expected of biology majors and available to non majors in colleges and universities; * integration of relevant evolutionary concepts into the post baccalaureate training of all biologists and of professionals in areas such as medicine, law, agronomy, and veterinary medicine; * adequate representation of evolutionary biology, especially of understudied subdisciplines, among faculty positions in colleges and universities; * active contributions from the scientific community toward informing the public about the nature, progress, and implications of evolutionary biology. As physics has shaped the science and technology of the present century, so biology will shape the twenty-first century. 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