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The Society for Integrative and Comparative Biology
Integrative Biology, an Organismic Biologist's Point of View1
1 Department of Ecology and Evolutionary Biology, University of California, Los Angeles, California 90025-1606
Under conditions of social stability such as existed in an earlier and simpler era when there was little change from one generation to the next, it is reasonable to assume that there was a positive correlation between age and wisdom. Today the validity of that assumption seems tenuous, to say the least. Consider my situation. I am 85 years old. Clearly, I am an old scientist, and I am not sure that I have any scientific wisdom to communicate. My uncertainty is in part related to the dynamics of the scientific enterprise.
Because of the success of the natural sciences and the technology derivative from them, for the past four or five centuries the human species has lived in a state of accelerating technological and cultural change. The rate of change has been particularly high for us scientists. Indeed, we live in a state of continuous intellectual revolution—and that is not an easy situation to which to adjust. As one of the elder members of the community of integrative biologists, I am overwhelmingly aware that during this continuing intellectual revolution, seniority is more likely to be correlated with obsolescence than with wisdom.
However, despite the rapidity of scientific change, science is an intensely historical activity: every investigator stands on the shoulders of those who have preceded him or her. My thoughts may be old-fashioned, perhaps obsolete, and almost certainly they are platitudinous. Nevertheless, they may have some historical interest, because they represent the understanding of a person whose university education and scientific activity spans more than 60% of the Twentieth Century, a period of scientific expansion and intellectual change without precedent in human history.
Most scientists, consciously or subconsciously, operate within a series of assumptions about the nature of their areas of professional concern. I can identify the assumptions that have been the basis of my view of biology. My confidence in them has been sufficient for me to treat them as if they were axioms. In retrospect, it is clear that they have shaped my interpretation of research data and sometimes guided my selection of research problems. Some of these assumptions, which have been axiomatic to me, may be platitudes to you. They reflect the fact that during a lifetime of studying the functional adaptations of organisms I have never been able to decide whether I was a student of natural history, ecology, behavior, or physiology. (In my more manic moments I have even considered myself a student of evolution.) My undergraduate years were the 1930s. The early formative years of my professional life were the late 1940s and 50s—before the DNA revolution.
For better or for worse, here are a few of my personal axioms. I present them to you primarily for historical interest.
- Two of the salient attributes of superior science are originality of conception and generality of application (Bartholomew, 1982
). The example par excellence is Darwinian natural selection. It offers a mechanism so widely applicable that it is almost coexistent with reproduction and so innovative that it shook and continues to shake people's perception of causality, of themselves, and their place in the cosmos. It is the central pillar of my understanding of the biological world.
- Only two processes are known to lead to the evolution of adaptive change, these are natural selection and chance. Natural selection itself is a stochastic process. Chance plays a substantial role both in generating the genetic variability on which natural selection operates and in shaping the ecological arena in which natural selection takes place.
- Natural selection increases fitness, but it produces systems that function no better than they must (Bartholomew, 1986). It yields adequacy of adaptation rather than perfection. The adaptive changes that result from its actions are adjustments only to local and immediate conditions. It is blind to the long-term consequences of the biological changes that it produces. The chances that short-term adaptations will meet long-term environmental changes or some new or future challenge are inevitably slight. Consequently, for a given population or a given species, the long-term probability of extinction is high.
- Despite the high long-term probability of extinction, every organism alive today, including every person reading this paper, is a link in an unbroken chain of parent-offspring relationships that extends back unbroken to the beginning of life on earth. Every living organism is a part of an enormously long success story—each of its direct ancestors has been sufficiently well adapted to its physical and biological environments to allow it to mature and reproduce successfully. Viewed thus, adaptation is not a trivial facet of natural history, but a biological attribute so central as to be inseparable from life itself.
- Diversity of form and diversity of function are among the most salient features of organismic biology. One of the most impressive contrasts in all of biology is the relative uniformity of the physiological machinery at the cellular and molecular levels of integration and the diversity of the adaptive patterns into which this machinery is assembled at the organismic level (Bartholomew, 1987).
- Organism and environment form an inseparable pair. As Claude Bernard pointed out more than a century ago, each can be defined only in terms of the other. Thus, an organism can effectively be visualized as a chemical system whose integrity depends on continuous exchanges with its physical environment, and which has a history continuing back through time in an unbroken series, with each cycle (in most multicellular animals) consisting of a breeding adult, a gamete, a zygote, an embryo, and then another breeding adult. If he has this perspective, a student of organisms is forced to attempt to deal with biological data in an integrated manner. He may even be stimulated to pose research questions that others never ask, particularly if they work rigidly within the constraints of some special field. Physiology comes immediately to my mind because its techniques have been a central to much of my research.
- Each species has evolved a special set of solutions to the general problems that all organisms must face. By the fact of its existence, a species demonstrates that its members are able to carry out adequately a series of general functions, all of which are familiar and most of which are thoroughly identified. These general functions offer a framework within which one can integrate one's view of biology and focus one's research. Such a view helps one to avoid becoming lost in a morass of unstructured detail—even though the ways in which different species perform these functions may differ widely. A few obvious examples will suffice. Organisms must remain functionally integrated. They must obtain materials from their environments, and process and release energy from these materials. They must remain adequately hydrated. If they are vertebrates they must maintain body temperature within the narrow range of approximately 50 degrees (for microorganisms about 100 or so degrees) in a universe where ambient temperatures range from absolute zero to millions of degrees C. They must differentiate and grow, and they must reproduce. By focusing one's questions on one or another of these obligatory and universal capacities, one can ensure that one's research will not be trivial and that it will have some chance of achieving broad general applicability.
- The responses of organisms ignore the categories of scientific specialization. Organisms are functionally indivisible and their activities do not fall neatly into even the broadest of the conventional biological compartments such as physiology, morphology, behavior and genetics. These broad categories, and also the many more restricted areas of specialization into which biologists segregate themselves, do not exist separately in the world of organisms. They are artifacts of convenience and fashion, created by humans and perpetuated by university administrative structures and government bureaucracies, by traditions, and by groups of biologists. It is the intact and functioning organism on which natural selection operates. Organisms are therefore a central element of concern to the biologist who aspires to a broad and integrated understanding of biology.
- An organismic biologist faces inescapable complications because of the nature of the system that he studies. Every organism has an evolutionary history. All organisms face similar problems but various taxa have met these problems in a wide variety of ways. Thus, it is difficult to find general answers to specific questions, particularly when one deals with adaptations. It is more productive and more biologically realistic to expect many different special answers to each general question depending on the taxon and the environment to which it is adjusted.
I shall conclude my homily by talking briefly about scientific creativity and innovation—the qualities that, more than any other, characterize the superior scientist. Creativity can be enhanced by bearing in mind a point that Bernard Cohen, a perceptive student of the history of science, pointed out some years ago. Scientific breakthroughs, from the simplest innovations to sweeping intellectual revolutions, are characterized by the transformation of existing ideas and information. Such a process is analogous to the evolution of adaptations through natural selection. One can sometimes enhance the probability of making an innovative scientific contribution by transferring ideas and questions from one field of specialization to another, or from one taxonomic group to another. This procedure, which can often be done with surprising ease, has been especially important to me and also to many of my students.
This brings me to the final point of my remarks, the relation between creativity and aging, a topic with which I have had substantial experience. Scientific research, until it has gone through the grueling and sometimes painful process of publication, is just play, and play is a characteristic of young vertebrates, particularly young mammals. In some ways, scientific creativity is related to the exuberant behavior of young mammals. Indeed, creativity seems to be a natural characteristic of young humans. If one is fortunate enough to be associated with a university, even as one ages, teaching allows one to contribute to, and vicariously share, in the creativity of youth.
My primary interest has always been research. I produced scientific publications steadily for fifty years, but the scientific relevance of even my best efforts has been at most a decade or two. It is clear that my most lasting contribution to science has been my students and the generations of their students who have followed them.
I will end on a highly personal note. The wisest decision I ever made with regard to science, I made as a child. In the summer of 1932, shortly after my thirteenth birthday, I decided to become a zoologist, because I thought it would be fascinating to visit distant parts of the world and study exotic animals. I was right. It has been.
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ACKNOWLEDGMENTS |
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Of the various professional awards I have received, none has given me more satisfaction than having this SICB lecture series bear my name. I am grateful to my former students, associates and friends, and to my wife Ruth, for the generous financial support that has made this lecture series possible. I am also grateful to Ray Huey and the members of the committees who have selected the awardees and organized today's symposium.
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FOOTNOTES |
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1 From the Symposium Integrative Biology: A Symposium Honoring George A. Bartholomew presented at the Annual Meeting of the Society for Integrative and Comparative Biology, 5–9 January 2004, at New Orleans, Louisiana.
2 Author's New Address, 501 Via Casitas Apt 814, Greenbrae, CA, 94904. E-mail: gbarthol{at}aol.com
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Bartholomew, G. A. 1982. Scientific innovation and creativity: A zoologist's point of view. Amer. Zool, 22:227-235.
Bartholomew, G. A. 1986. The role of natural history in contemporary biology. Bioscience, 36:324-329.[CrossRef]
Bartholomew, G. A. 1987. Interspecific comparisons as a tool for ecological physiologists. In M. E. Feder, A. F. Bennett, W. W. Burggren, and R. B. Huey (eds.), New Directions in Ecological Physiology, pp. 11–35. Cambridge University Press.
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