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1 Introduction,
2 What Is Group Selection?,
3 Group Selection in the 1970s,
4 Career Beginnings and Science after the Thesis,
5 Experimental Studies of Population Heritability,
6 Population Ecology and Population Heritability,
7 The Evolution of Sociality,
8 Calibrating the Laboratory to Nature,
9 Experimental Studies of Wright's Shifting Balance Theory,
10 Beyond the Shifting Balancing Theory,
Acknowledgments,
Reference List,
Index,
Introduction
The central question guiding my research throughout my career has been this: How is the process of adaptation different if the members of a population live clustered in small groups instead of being homogenously distributed like grass on a lawn? The field is called "evolution in subdivided populations" or "adaptation in metapopulations." It has led me to investigate a diverse array of topics, including group selection, family selection, kin selection, and sexual selection, as well as speciation genetics, maternal and paternal genetic effects, and host-symbiont coevolution. In my lab, my students and I have approached these topics using a combination of theoretical, field, and laboratory studies and a diversity of living systems ranging from our laboratory model of flour beetles in the genus Tribolium to other animals, plants, and microbes. Through the generosity of the National Science Foundation Opportunities for Promoting Understanding through Synthesis (OPUS) Program, the Sabbatical Scholars Program at the National Evolutionary Synthesis Center, and the Sabbatical Leave Program at Indiana University, I have had the opportunity to write a conceptual, historical synthesis of the findings from these studies and the relationship between adaptation in metapopulations and broader questions in evolutionary genetics.
This first chapter is an overview of and introduction to concepts considered in greater depth in later chapters. I first provide a bit of personal background about my family and my schooling, since both influenced my career and shaped my research interests.
The organization of later chapters follows the flow from questions and concepts to laboratory experiments and to field studies. The majority of my mathematical models developed from the physical activity of moving flour beetles within and among families (kin selection) or within and among populations (group selection) and the synthetic activity of analyzing data. The dates on some theoretical publications precede those of the experimental works that inspired them, but only because the design, execution, and analysis of experiments is much slower (and generally more tedious) than working out the results of mathematical models.
New data are included in later chapters; data from experiments that went unpublished, owing to interruptions from teaching, administrative tasks, or lapses in funding, as well as to the dispersion of young collaborators away from my lab to career opportunities elsewhere. Published or not, results from these studies became part of lab lore and influenced our thinking and the planning of subsequent research.
Personal Background
In 1949, I was born in Evanston, Illinois, the oldest of eight children, into a fundamentalist, Irish Catholic family. I spent most of my waking hours in the swamps near our home in Westport, Connecticut, collecting frogs, toads, turtles, snakes, and butterflies at Lee's Pond, Willow Brook Cemetery, and the banks of the Saugatuck River. My parents encouraged our interest in nature by allowing us to keep anything we could catch and my dad built elaborate circus wagons, wheeled cages, for moving our menagerie in and out of the garage. My childhood ambition was to become not a scientist, but rather curator of reptiles at the Bronx Zoo, inspired by Marlin Perkins and his show, The Wild Kingdom.
Although neither of my parents had completed college, my mother emphasized our education above all else. (Long after we left home, my mom returned to college and completed her degree at St. Joseph's University of Pennsylvania in 1983 at the age of 61.) And, although she did not believe in evolution herself, she introduced us to it by reading us "dinosaur books" at bedtime. Each summer, we took family trips to Yale's Peabody Museum, where we spent our allowances on rubber dinosaurs and the army men to fight them.
In 1963, my father was appointed director of research for the Triangle Broadcast Center and we moved from Connecticut to Drexel Hill, Pennsylvania, a Catholic enclave near Philadelphia. I attended St. Joseph's Preparatory High School and my endless hours in the swamps were replaced by 5–6 hours of homework each night; a test every week in every subject; and daily quizzes in Latin and math. The Prep faculty had two of my best teachers, Mr. Earl Hart in honors mathematics and Stephen A. Garber, S. J., in honors chemistry. In 1967, I entered another Jesuit institution, Boston College, on scholarship. College was easy compared to high school and a year of advanced placement credit allowed me the freedom to experiment with majors in chemistry, English, sociology, mathematics, and biology. I spent my senior year writing plays, oil painting, and learning Boolean algebra as a "Scholar of the College" before graduating in 1971 with a double major in mathematics and biology. Somewhat unfocused, I applied to law school as well as to graduate schools in anthropology, linguistics, and evolution and ecology. With guidance from my biophysics professor, Dr. Donald J. Plocke, S. J., I applied to and was accepted by the Department of Theoretical Biology at the University of Chicago, where I was supported by a four-year National Institutes of Health (NIH) Graduate Training Fellowship (1971–1975). It was amazing to get paid to go to school, especially to a doctoral program uniquely suited to my combined interests in biology and math. The Chicago faculty applied mathematical models to everything from development to neuroscience to biological clocks. My background in natural history caused me to gravitate toward ecology and evolution. My dissertation was coadvised by Drs. Thomas Park, founder of the field of laboratory ecology and soon to be professor emeritus, and Montgomery Slatkin, a biomathematician and beginning assistant professor. Both were challenging and supportive mentors who guided my earliest ventures into group selection and the evolutionary genetics of metapopulations (chapters 3 and 4).
Interactions and Context
In graduate school, I learned for the first time that Darwinian evolution required variation, replication, and heredity and that any system whose units had those properties could evolve (Lewontin 1970b; later, Maynard Smith 1976) and that selection could operate simultaneously at more than one level and in more than one direction. This made it particularly difficult to determine a priori whether selection at one level was more efficient in producing evolutionary change than selection acting at another. Quantifying the relative efficacy of multilevel selection required an understanding of selection and heredity at each level. The goal of my research was to develop methods for this kind of multilevel quantification and to test the relative efficacy of the levels of...
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