CHAPTER FOUR

What Brings You Here Takes You Away

I SPENT TWO YEARS AFTER I LEFT THE UNIVERSITY OF MICHIGAN engaged in revolutionary politics. Recall the expression “History is like the life of a soldier…”; my work as a revolutionary agitator was much like that. Sometimes I was just passing out leaflets at the Ford River Rouge plant in Dearborn, Michigan, to support a slate of progressives running for union offices. Other times I was sitting in a car with a revolver in my coat to protect the lives of union members who had been threatened by organized crime associated with their union bureaucracy. Still other times I was organizing to help stop Klan terror threats against African Americans who had moved into formerly all-white neighborhoods in the western suburbs of Detroit. During those years I traveled to the United Kingdom to support mine workers striking against Margaret Thatcher’s government for better wages and safer working conditions. On one day I stood arm in arm with miners’ wives, standing up to a charge by mounted police officers attempting to break up their picket lines.¹ On that visit I also spoke at a program addressing racism in the United Kingdom and the United States. After my talk was over, an elderly gentleman in the audience approached me and told me that my lecture was the best he had heard on the subject since that of his close friend Claude McKay in the 1920s.² To say the least, I was humbled.

In the period between the University of Michigan and Wayne State University (WSU), I worked with some of the most intelligent and passionate people whose names you will never know. They worked tirelessly, for little reward, and often risking their lives to combat racism, sexism, antigay bigotry, and class oppression. During my evenings, if I wasn’t at a meeting against some form of injustice, I was reading. I found that immersing myself in works of history, social science, philosophy, and economics was an excellent palliative against hunger. My wife, Sue, and I survived on the various short-term jobs we could find. At one point I was a seasonal worker driving a UPS truck from 5:00 a.m. to 3:00 p.m. and then tutoring mathematics at Wayne County Community College in the evening. Sue worked as a waitress and at other odd jobs. She was not a social activist in the way I was, but she was committed to helping people as she could. Years later she admitted that she often went without food during the day so I would have something to eat when I came back to the apartment. Our first Thanksgiving together included not a turkey but, rather, canned foods that friends had donated to us.

We both spent time as substitute teachers in the Detroit Public Schools system. We experienced Jonathan Kozol’s Savage Inequalities up close and personally.³ We found ourselves using what little money we had to provide lunch money for hungry children. My wife gave away her only winter coat to a child without one. Sometimes we would luck into a long-term assignment to replace a teacher, only to be removed right before the school district would have to give us a contract. The next day we would be called back to the same school and class to resume the same assignment. At one point, I felt that maybe the best I could do with my life would be to go back to school to earn a teaching certificate. However, it became increasingly clear to me that I would never be able to make a lasting change in the public school system because it was not designed to empower children (particularly Black children). Rather, its goal was to indoctrinate them to become pliant workers and docile consumers in American society.⁴

My frustration with the situation in the Detroit Public Schools and our deteriorating economic situation led me to decide to return to school to earn a PhD. It was clear to me that I was never going to get people to really listen to what I said without a credential. I decided my best chance to finish my degree was at Wayne State University (WSU). So I called the biology department and made an appointment with the graduate adviser. I was the biologist who came in from the cold. On the day of my appointment, the snow was blowing, but not quite enough to obstruct my vision as I drove down Woodward Avenue on my way to the campus. My cream-colored Chrysler K-car barely ran on a good day. On that day it was struggling and wasn’t producing any heat. It was the ugliest car to ever roll off the assembly lines of Detroit—certainly not to be compared to Stephen King’s elegant Christine or the 1967 Chevrolet driven by Dean and Sam Winchester in what would become my favorite drama, Supernatural. If I had been a truly merciful person, I would have just put that car out of its misery, but on that day I needed it.

I explained to the WSU biology department graduate adviser, William Moore, that I wanted to apply for admission to the graduate program. He looked at me disapprovingly and said something to the effect of “You understand that this is a very prestigious and rigorous program?” I responded, “Yes, Dr. Moore, I understand, and that is precisely why I want to come here to finish my PhD.” I handed him my unofficial transcripts from Oberlin College and the University of Michigan. He glanced them over and then immediately picked up the phone to dial the office of Leo Luckinbill. (If you recognize the last name, he is the brother of the actor Laurence Luckinbill, who played the part of Spock’s brother in Star Trek V: The Final Frontier.) After a short conversation with Dr. Luckinbill (henceforth Leo in this chapter), I decided to apply to WSU biology. I was admitted, with a solid financial aid package, for the fall of 1985.

In my first year in the WSU graduate school, my head was clearly not fully back in the academy. I wasn’t spending enough time on my classes, and I was virtually a no-show in the laboratory. I spent many of my afternoons hanging out in the gym or in Hart Plaza with the chess hustlers. The most challenging class of that first year was offered by Kazutoshi “Koz” Mayeda, who taught human genetics. I was better prepared than the other first-year graduate students for this course. I already had two years of graduate coursework from Michigan and had done rather well in Julian Adams’s graduate-level population genetics course. Our text for that course was Lewontin’s classic work The Genetic Basis of Evolutionary Change.⁵ None of that mattered with Koz. He was old-school in the most fundamental way. I worked very hard in that course, and I definitely felt he was being harder on me than on any of the other students. I assumed, of course, that this resulted from the same anti-Black racism I had experienced in my interactions with East Asian Americans. Koz gave a monster of a final exam. Nobody left early. However, as we all shuffled out of the classroom, heads down from the ordeal, he pulled me aside. He asked me, “Joe, do you know why I was harder on you than on any of the other students?” I shook my head and said no. His answer shocked me, and it still rings through my psyche to this very day: “Because I know what kind of racism you are going to face in this field, and I wanted to be sure that I did everything in my power to prepare you to face it.” To the say the least, I was shocked. He then told me the story of his family’s internment at the Manzanar War Relocation Center during World War II. I don’t know if his interest in genetics was spurred by Masuo Kodani, a famous Japanese American cytogeneticist who was also at that camp.⁶ Koz would have been thirteen or fourteen years old when his family was interned. It little matters where he picked up his interest in genetics, because that interest spurred him to make sure that a promising graduate student received his full attention. After he finished his story, he asked me for the honor of serving on my dissertation committee. I learned a lot that day about the fallacy of assumptions. Koz clearly saw something in me that I myself didn’t yet see. I am eternally grateful that he did.

The most fateful moment of my life to that point occurred when Leo called me into his office for a conversation. His words were direct and exactly what I needed to hear. He said something to the effect of “Joe, nobody here doubts that you are a really smart guy, but if you don’t get into the lab tomorrow and start working on your thesis, I am going to cut you loose.” His calling me on the carpet forced me to make a decision. Was I really committed to earning a PhD? Or was a failed intellectual all I was to ever be? After I decided to commit to a PhD, I had an additional problem: What exactly was I going to do? I had to design an experiment and get to work on it the next day. PhDs are awarded only to students who demonstrate their capacity to complete a research project, from start to finish. At Michigan I had been an endless font of great and interesting ideas, but I failed at carrying anything through to completion. Imposter syndrome was once again staring me directly in the face, but this time I wasn’t going to be the one to blink.

IN GREEK MYTHOLOGY FATE WAS EMBODIED BY THREE SISTERS (Clotho, life; Lachesis, lifespan; and Atropos, death). As fate would have it, on the snowy day I chose to return to science, I became part of one of the most significant scientific revolutions of the twentieth century concerning the biological basis of their responsibilities. The ancients pretty much uniformly believed that aging, disease, and death entered the world as the result of a curse from the gods. The Greeks blamed Pandora’s curiosity; the Hebrews, Satan’s tempting of Eve to eat the fruit from the tree of knowledge. The Old Testament (Hebrew Bible) recounted conflicting ideas concerning how long an individual might live. In Genesis 5:27 we are told that Methuselah lived 969 years, but the Psalms describe the life of a man as only seventy to eighty years (Psalms 90:10).

As poetic as these ideas are, they give us no mechanistic understanding of why organisms age (or even a clear definition of what aging is). Little progress on these ideas occurred until the late nineteenth century. Darwin didn’t discuss the question of aging, but the codiscoverer of evolution by means of natural selection, Alfred Russel Wallace, did. Wallace suggested that after aged individuals (who were no longer reproducing) had produced a sufficient number of offspring, they would be a detriment to the species because they would be utilizing resources that their progeny needed to survive and reproduce. Natural selection would thus favor mechanisms that would remove them from the population. The problem with this argument is that it can easily be understood as an example of group selection. Group selection is the idea that the group would be favored over the individual. A variety of experiments have shown that this is incorrect in most circumstances. The evolutionary biologist Michael Rose pointed out, however, that this was an incorrect way of understanding Wallace’s idea. Even if one imagined that an organism might have a physiology that would allow immortality (at the cost of sterility), such an organism’s species would still go extinct because of external causes of death (such as predators, forest fires, drowning). Therefore, he reasoned, natural selection should always favor reproduction over an extended life span.⁷ At around the same time Wallace was musing on aging, the German cytogeneticist August Weismann began a more serious examination of the biology of aging.⁸ Weismann’s great revelation was that the pattern of aging and life span in any species could not simply be due to internal biochemical and physiological processes. For Weismann, adaptation to the needs of the species, which was an evolutionary problem, was at the root of the patterns of aging. He intimated that to correctly understand this question, a mathematical theory of natural selection was needed.⁹

The mathematical theory required to solve this problem arrived in the early twentieth century in the form of the neo-Darwinian synthesis (NDS).¹⁰ The mechanism of natural selection was unified with a correct understanding of the mechanisms of inheritance (Mendelian particulate inheritance). This was something Darwin did not have when he wrote On the Origin of Species in 1859. The NDS made possible the correct approach to a number of vexing problems in biological science, including the origin and maintenance of biological variation within species (or the question of what biological races were and how they came about). This problem dominated the second portion of my career. Ronald A. Fisher (who was also a eugenicist) provided a crucial equation that allowed the beginning of the solution to the question of why eukaryotic organisms age. He showed that organismal fitness (evolutionary fitness, not physiological fitness) in species with age structure is the product of their age-specific survival (l_x) and age-specific reproduction (m_x).¹¹ Thus the formula for lifelong evolutionary fitness of an organism with age structure is equal to Σ l_x × m_x across all ages (x). Age-specific survivorship is the probability of living to a certain age. If 100,000 babies are born on a given day, one year later about 99,000 will still be alive. The survivorship probability (l₁) for that year is 99,000 / 100,000, or 0.99. Another important aspect of survivorship probabilities is that they always decrease over time. So imagine that for this same group (cohort), by age fifty 85,000 are still alive, l₅₀ = 85,000 / 100,000 = 0.85. As time goes on, more individuals die, so that at age one hundred, only 1,000 are still alive; l₁₀₀ = 1,000 / 100,000 = 0.01. Because humans cannot reproduce until adolescence, the number of offspring the females could produce at age one is 0. So the age-specific fitness would be calculated as 0.99 × 0.00 = 0.00. Alternatively, at age thirty in the contemporary United States, the average number of children a female has is approximately 2.1, so with l₃₀ about 0.90, the age-specific fitness would be 0.90 × 2.1 = 1.89. Thus, until organisms reproduce, they are potentially evolutionary dead ends, which gives even more support to Wallace’s idea that natural selection will tend to favor reproduction over survival.

Important developments in the evolutionary theory of aging were supplied by the British biologist Peter Medawar in 1952. Medawar’s central idea can be summarized by this passage:

The force of natural selection weakens with increasing age—even in a theoretically immortal population, provided that it is exposed to real hazards of mortality. If a genetical disaster… happens late enough in individual life, its consequences may be completely unimportant.¹²

Medawar’s idea was an extension of Wallace’s original proposal. Medawar is saying that what natural selection is most concerned with is the way an organism operates during its reproductive period. As genetical disasters late in life don’t impair reproduction, genes with these effects will not be removed from any species. In 1957, George C. Williams further improved on Medawar’s ideas. He proposed that there might be genes that were beneficial to an individual early in life but harmful to that same individual late in life (after reproduction was completed). As these genes often affect multiple features of the organism, they would result in an “antagonistic pleiotropy.”¹³ “Pleiotropy” is the term in genetics for a gene that affects multiple traits simultaneously. Finally, a few years later, William D. Hamilton (the same guy I knew at Michigan) formalized the mathematics behind Medawar’s and Williams’s concepts in a paper published in the Journal of Theoretical Biology.¹⁴

These ideas set the stage for Michael Rose’s brilliant experiment.¹⁵ Leo came upon a similar idea at around the same time.¹⁶ Their papers were published one after the other in the same volume of the prestigious journal Evolution in 1984. What Rose (Michael for the rest of this chapter) had realized was that the Medawar/Williams evolutionary theory of aging predicted that the timing of reproduction determined the pattern of senescence. This timing of reproduction could therefore be altered in the laboratory utilizing the tools of experimental evolution. Experimental evolution is the study of the mechanisms of evolution (natural selection, genetic drift) under controlled and replicable conditions either in the laboratory or in field settings.¹⁷ This technique had been used in the discipline since the 1940s.

Michael chose the common fruit fly (Drosophila melanogaster) to test the Medawar/Williams theory. First, this organism fit the requirements of the theory (a separation of germ from somatic tissue). Germ tissue gives rise to an organism’s gametes (eggs or sperm). Somatic tissue (nerves, muscles, bones, etc.) takes care of all the behavioral and physiological requirements for an organism to reproduce. To put it simply, a chicken is just an egg’s way of making the next generation of eggs. This is a key concept. It means that with regard to the fundamental requirements for the evolution of aging, fruit flies and mammals (including humans) are no different. The second benefit of using fruit flies for the experiment was that because they are small and inexpensive to maintain, Michael was able to use population sizes large enough (about 2,000 per population) to avoid any results due to genetic drift (chance fluctuation on gene frequency due to small population size). This also allowed him to replicate his populations fivefold. Again, this guaranteed that results were not just fluke events. Finally, the genetics and the genetic map of Drosophila was already well known at this time. Frédéric Lints and his coworkers had attempted, with no success, to use experimental evolution to alter the pattern of senescence in Drosophila a decade before the Rose and Luckinbill experiments.¹⁸ From his failure he concluded that aging was not under genetic control. However, Lints’s failure resulted not from the fact that genes did not play a role in aging but from the way he constructed his experiments. Specifically, he grew the larvae of the flies he attempted to select for altered life span at a density of ten larvae per vial. For some reason, which we still don’t exactly understand, higher densities (about sixty larvae per vial) are required to stimulate the patterns of gene expression required for the flies to respond to selection for delayed reproduction. Both the Rose and Luckinbill experiments utilized higher numbers of larvae per vial and were able to show clear results of selection acting on the timing of reproduction, resulting in flies evolving longer life spans.

NONSCIENTISTS WOULD PROBABLY BE HORRIFIED BY THE IDEA that some of science’s greatest ideas were arrived at by accident. Michael Rose’s group discovered by accident that his postponed-senescent (longer-lived) fruit flies had superior physiological capacities than the controls, the short-lived fruit flies. According to Michael’s account, he walked into the laboratory one morning and found his laboratory technician in tears. He asked her what was wrong, and she told him she had “killed the flies.” She had forgotten to put the agar plates in their cages over the weekend, and when she opened an incubator she found that the flies had died from starvation and desiccation. Michael panicked and ran over to the incubators. He looked inside, and the flies he saw were fine. But the technician and Michael had opened different incubators. The technician had opened the incubator containing the controls (B stock, short-lived), and Michael had opened the incubator containing the experimental flies (O stock, long-lived). He reasoned that a consequence of the O stock’s postponed aging was a superior physiology, better stress resistance. He and his postdoctoral researcher, Phil Service, got to work right away to validate this, and they published their results in 1985.¹⁹

During the afternoon, evening, and night before Leo’s deadline for kicking me out of the lab, I was feverishly poring through the papers on the evolution and physiology of senescence. I also concluded that postponed senescence must have caused the long-lived flies to attain a dramatically different physiology from that of the short-lived flies. I reasoned that the best place to demonstrate this difference would be in their flight performance. Of course, I had to design an experiment to demonstrate that. It was also critical for me that Phil Service had not examined flight, as I could not earn a PhD for simply repeating another researcher’s experiment. The key, it seemed to me, resided in the fact that all flies have something called the “tarsal reflex”: if you suspend them in the air, they will automatically fly until exhaustion. All I needed to do was devise a method of suspending a very small fly (2 mm wide and 3 mm long) in such a way that its wings still worked. The solution I arrived at was to use a light-test fishing line attached to a Pasteur pipette. The fly was attached to the fishing line with Duco Cement. I used Duco to repair things around my apartment, so I knew it wasn’t a particularly strong glue, but that attribute made it useful for this work. In the morning I went out and bought the fishing line and a tube of Duco. When Leo walked into the laboratory later that afternoon, to his shock, he saw me sitting in front of a line of flies that were suspended from a test-tube rack and flying effortlessly. I had a stopwatch on each fly timing its flight duration. So began the experiment that changed my career forever. When I told Sue about the design of my experiment, she was amazed. It turned out that as a child in her village she used to capture dragonflies and tether them using a small string. The dragonflies were essentially living kites. Furthermore, the English translation of Sue’s name means “long life” or “longevity.” She was given this name because she almost died at birth. So, years later, the man she married was tethering insects to study longevity. You can’t make up stuff like this!

Conducting the flight-duration measurements was both labor- and time-intensive, so I recruited and trained undergraduates to work with me, via independent study. This was a win-win situation: I acted as a research instructor, teaching students about science in the way science was actually achieved, and they provided me with the labor needed to get the experiments completed. I soon found that a disproportionate number of these students were underrepresented minorities (URM) and/or women. This was easy to understand; even though Wayne State had a large enrollment of URM students, there were very few URM faculty or graduate students in biology. Notable exceptions were James “Jim” Jay, one of the first African American microbiologists, and Joe Dunbar (at the Medical School). Both these men provided me with valuable mentorship during my time at WSU and in the early portion of my career. URM and female students populating my research groups would turn out to be a consistent feature of my career. Later on, I would design and direct several NSF and National Institutes of Health (NIH) training programs to address underrepresentation, and I still do to this day. In those early years, I was perfecting a model of tiered mentorship that allowed me to train students, who would in turn train students with less experience than they had. This allowed me to conduct the very large and labor-intensive Drosophila life history experiments that were the primary subject of my work in this period.

VERY EARLY ON IN MY EXPERIMENT IT BECAME CLEAR THAT THE long-lived population had dramatically better flight performance (three to four times better) than the short-lived ones. However, an obvious explanation came to mind: What if the flies had different flight behaviors? What if the long-lived ones flew slower—that is, with a lower wing-beat frequency (WBF)—and were thus more capable of flying longer? Flies (the order Diptera) maintain very high wing-beat frequencies (over two hundred cycles per second). I needed a way to measure instantaneous flight while they were suspended in the air. Leo had a friend in the physics department (a sort of MacGyver-type guy, Angelo Nichols) who cobbled together instruments of all sorts. I talked with Angelo and proposed that we could use a laser setup so that the flies’ wing beats (which are very much like the butterfly stroke human swimmers use) would break the beam. We could simply count the frequency of the wing beats breaking the beam and have the data we needed. First we tried an industrial laser. Well, the flies didn’t respond very well to that setup. I seem to remember a few of them bursting into flame, but I think that was more one of my nightmares than something that actually happened. However, when we finally did get the system to work, we noticed right away that the oscilloscope was not varying from a reading of sixty cycles per second. We immediately realized our error: the oscilloscope was reading the pulse of the laser, and the flies were flying much faster than the laser could register! Angelo realized we needed a better medium, and that’s when he decided to try an infrared beam. The final design of our device is shown in Figure 4.1.²⁰

Figure 4.1. Diagram of an infrared-based wing-beat frequency device. A signal generated by interrupting an infrared beam between emitter and detector diodes passes through wave shaper and counter circuits to online storage in microcomputer memory. To the left is a fly tethered by Duco cement.
Source of diagram on the right: Graves JL, Luckinbill L, and Nichols A, Flight duration and wing beat frequency in long- and short-lived Drosophila melanogaster, Journal of Insect Physiology 34 (1988): 1021–1026.

It was during one of our WBF measurement sessions that I first met Michael Rose. He was in Michigan and had come by our laboratory to talk with Leo about our experiments. When he saw the flight apparatus, he was immediately impressed, both because I had gotten tethered flight to work with Drosophila (Phil Service had tried it in his laboratory and failed), and because I had thought about whether differences in WBF might be involved in differential energy expenditure between the short- and long-lived lines. After Michael left our laboratory, I did not know that he was beginning to hatch a plan to bring me to UC Irvine upon completion of my PhD. The specific mechanism he used was put forward by my old Oberlin colleague, Richard Lenski, who was also a professor at Irvine. A few weeks later I was contacted by Michael and encouraged to apply for the University of California’s President’s Postdoctoral Fellowship program, a program designed to recruit talented URM postdoctoral candidates to the University of California with an eye to eventually moving them into tenure track faculty appointments. The program is still in existence, because despite the academy’s “best” efforts, there has still not been sufficient increase in the numbers of URM scientists taking up faculty appointments at America’s elite universities.

The results of the WBF experiments showed that the long-lived flies actually flew at a slightly faster WBF than the short-lived ones (about 220 cycles per second, compared to about 180 cycles per second). The studies of flight duration and WBF composed Chapter 1 of my doctoral dissertation and were published in 1988 in the Journal of Insect Physiology as my first lead-author, peer-reviewed journal paper. That year, the Annual Drosophila Research Conference was held in Chicago, at the Hilton. I submitted my abstract and it was accepted. I was also nominated for the first Larry Sandler Memorial Lecture. Larry Sandler was a Drosophila biologist who had died the year before. I wasn’t chosen for that honor (Bruce Edgar was). My paper was scheduled as a contributing talk in the section on Methods. If you have ever been in the Hilton, then you have some sense of how large its main ballroom is. I estimated that when our session started, there were more than five hundred people in the room (it was standing-room only). I was sitting in the front row with the other four or five people scheduled to speak in the session. This was my first talk at a national meeting. As a graduate student at Michigan, I had given some talks at the Midwest Population Biology meetings. If there were fifty people at the entire meeting, it was a good year. What I didn’t realize was that the speaker before me was one of the first people to use polymerase chain reaction (PCR) on Drosophila DNA. As soon as his talk was over, the ballroom began to empty, with participants heading off to other concurrent sessions. By the time I reached the podium, there were no more than twenty people in the room. I made an opening joke about whether I had accidentally wandered into a Klan meeting—that got a few chuckles—and gave my flight and WBF talk. I was somewhat disappointed at the low turnout, but the following day I heard groups of people all over the meeting talking about my presentation. Some other graduate students even came up to me and addressed me as “Dr. Graves.” I was amazed, but I admitted to them that I was still completing my dissertation.

In hindsight, Detroit may have been the only city in America that would have allowed me to finish my PhD. The city’s history is deeply entwined with the African American struggle for liberation. On March 12, 1859, John Brown and Frederick Douglass met at William Webb’s house on Congress Street to discuss African American emancipation.²¹ Douglass approved of Brown’s sentiments but disagreed with his plan. A few months after this meeting Brown would attempt to carry out his radical plan for an armed uprising at Harpers Ferry, Virginia. On days when I felt particularly low and my dissertation seemed far out of reach, I would go downtown to the marker commemorating that historical meeting. Detroit is also where the trade union movement first began to make headway with African American workers.²² I knew some of the participants in the Dodge Revolutionary Union Movement (DRUM).²³ Diego Rivera’s frescoes depicting the role of the worker in civilization reside in the Detroit Institute of Arts museum, across the street from the Wayne State campus.²⁴ This was another place I would go to find solace in those torturous days before I completed my dissertation. Finally, Detroit is the home of Motown Records; the Hitsville U.S.A. museum (at 2648 W. Grand Blvd.) and the home Aretha Franklin grew up in (at 7400 LaSalle Blvd.) are both only a couple of miles from the WSU campus. Yes, if there was any city that could have carried me through this ordeal, it was Detroit.

I still had two hurdles to get over before I finished at WSU: my final oral examination and then my dissertation defense. I dreaded my oral examination, as I feared it would provide the faculty with an opportunity to pay me back for my obnoxious behavior during the three years I had been at Wayne. One example is enough to explain why I was so concerned. During my last year at Wayne I was a teaching assistant (TA) for one of the sections of Introductory Biology taught by William Thompson (ornithologist). In one of his lectures he was discussing the causes of insect population cycles. Nothing he said was incorrect, but he was leaving out a lot of important new developments (particularly the possibility that insect numbers were the result of chaotic cycles). I was seated at the top of the lecture hall, but Thompson could clearly see my facial expressions indicating disdain for his comments. Finally, he said out loud, before the entire class, “Well, Mr. Graves, if you think you know this topic better than I do, you are welcome to come down and take over the rest of the lecture!” Maybe he didn’t expect me to take him up on the offer, because he was definitely surprised when I rose from my seat, walked down the aisle to the podium, and took over the lecture. With no prior preparation, I proceeded to give the students a review of the classical ecological theory associated with insect population cycles. I followed that with a discussion of the utility of density-dependent models of population growth with time delay, discrete logistics, and some of my work on competition and dynamical complexity. To my surprise, when the class ended Thompson stopped me on my way out of the lecture hall and asked if he could be on my PhD committee. He obviously wasn’t threatened by my display but, rather, saw an opportunity to help me make the transition to the next level of my career.

With so many examples of that sort of behavior on my part in so many venues, the committee members (the biologists Luckinbill, Moore, Carl Freeman, Mayeda, and Thompson and the statistician Allen Reed) were undoubtedly looking forward to getting me in a closed room for my oral exam. They didn’t disappoint. It felt like being examined by the grand inquisitor Tomás de Torquemada, but without all the blood. The night before the exam I decided to take the Fifth on any questions I felt I could not answer well. By the time the two hours were up, I felt I had taken the Fifth more times than I had actually answered questions. The committee concluded the exam and asked me to leave the room. Waiting in the hall for their decision, I was convinced that I had failed the exam. After a short while they called me back in and told me I had passed. However, they felt that I needed to improve my comprehension of statistical methods and required that in my last semester I enroll in a statistics course. It was a good call on their part, as I had never taken a course in the area and what I knew had been self-taught.

My dissertation defense was scheduled for June 24, 1988. By this time, I had received word that I had been awarded the fellowship at the University of California to work in Rose’s laboratory. All that was required now was to write the dissertation. The personal computer (PC) was just beginning to come into wide use at that time. I had been doing my best to stay out of political activism in the weeks leading up to my defense. However, in the last few weeks before my dissertation defense, I received a call from a friend who worked at a women’s health clinic in a predominantly African American neighborhood. She asked if I would help them provide security against an announced pro-life rally against their clinic. I agreed and called John, a friend I had worked with in my anti-Klan activities when I first came to the city. John, a man of European descent, was a salt-of-the-earth guy—definitely the kind of person you would want in a foxhole with you if things got rough. He made some calls, and we soon had enough people to make sure that patients, doctors, and other clinic employees could be safely escorted through the rally.

Figure 4.2. Celebration photo, June 24, 1988. In the Luckinbill Laboratory after I completed my dissertation. Pictured from left to right: Leo Luckinbill, Sue Graves, and Joseph L. Graves Jr.

We arrived on the scene an hour before the pro-life rally was scheduled to start. The pro-lifers, decked out in “Christian soldier” T-shirts, were already there in force. They moved to block the sidewalk to prevent the people working in the clinic from getting inside. John and I recognized several of the faces in the pro-life group as Klansmen and neo-Nazis we had encountered at previous demonstrations. We asked them, politely and as professionally as we could, to clear a space on the sidewalk to allow people to enter their place of work. Polite didn’t work. When the scuffle started, news trucks were pulling up to cover the demonstration. The resulting film of John and me heavily engaged with the pro-lifers made the evening news. I wasn’t arrested, but the next day I was called in to talk with the chair of the biology department. I was pretty sure I was about to be tossed out of the department. Instead, to my surprise, he congratulated me on my efforts at the clinic. When I left his office, I breathed a huge sigh of relief.

The days leading up to my dissertation defense were a blur. On the afternoon of the event, shortly before the auditorium began to fill, Leo must have been wondering where I was, as he had not seen me in some time. In the days leading up to the defense I had been going without sleep. NoDoz and whatever stimulants I could find had been keeping me awake. In the hours leading up to my talk, Brad Pollack and I were in the graduate office, copying my dissertation. All that remained was to get past “that lady.” Those of you who wrote dissertations in the early days of the word processor know that there was always an administrative assistant in the graduate dean’s office tasked with measuring the margins of each and every page with a ruler. “That lady” finished her measuring and stamped the dissertation as received about thirty minutes before I was scheduled to present. I dashed across campus, ran up four flights of stairs, and burst into our apartment. I jumped in the shower, got dressed, and headed across the parking lot leading to the biological sciences building. By now the auditorium was packed. I don’t remember how the publicity for my dissertation was handled, but there was not a free seat in the auditorium. Leo started his presentation of “the candidate” for the audience with the autobiographical material in my dissertation. He quickly ran out of material and started ad-libbing with anecdotes about me in the lab. I arrived at the top of the lecture hall, and he said something like “And here he is now!” My slide deck must have been preloaded in the projector with all the slides turned upside down and placed opposite to the “this side toward screen” label. I don’t remember loading them myself. When I got to the podium, I was so flustered and mentally exhausted that I forgot the title of my dissertation! So I said to the audience, “I am going to talk to you today about something having to do with life history and aging in Drosophila.” I couldn’t help but notice the eyes of everyone on the committee rolling at that remark. However, I pulled myself together and proceeded to give that audience the best talk I had ever given on my research (Figure 4.3).

THE 1988 ANNUAL MEETING OF THE SOCIETY FOR THE STUDY OF Evolution (SSE) was held in Pacific Grove, California, at the Asilomar Conference Grounds, either shortly before or shortly after my dissertation defense. I flew out to California to attend the meetings and was picked up at San Francisco International Airport by my postdoctoral research adviser, Michael Rose.

Michael had been a child prodigy. He was born in Canada in 1955 and finished his undergraduate degree in 1975 at Queen’s University in Ontario. He completed his PhD work under Brian Charlesworth in 1979 at the University of Sussex in the UK. In 1992 he (with others) was awarded the President’s Prize of the American Society of Naturalists, and in 1997, the Busse Prize of the World Congress of Gerontology. He has published nine books on evolutionary biology (three of them as sole author). His 1991 work, Evolutionary Biology of Aging, caused a seismic shift in the field of gerontology.²⁵ Over the course of his career he has been cited more than twenty thousand times, which places him in the top 2 percent of all scientists. This citation level is consistent with his current title, distinguished professor, within the University of California.

Figure 4.3. Cover of Life magazine, October 1992. The cover article discussed current research about postponing aging and mentioned work by Rose and Graves with regard to extending healthy life span.

I had only talked with Michael for short bits of time during his visit to Detroit. So I felt some apprehension about going to work with someone I didn’t know that well. All I really knew about Michael was that he was brilliant—so brilliant that although we were the same age (thirty-three), he was under review for promotion to full professor at UC Irvine and I was just starting as a postdoctoral researcher. However, when he turned on his car in the parking lot and James Brown came pouring out of the speaker, I knew we were going to be okay.

The first morning at the conference center was like every other professional science meeting I had attended so far. As I sat alone in the cafeteria for breakfast, it was obvious that I was the only African American at the meeting. At one point, I looked up from my breakfast and saw an older gentleman walking toward my table. I was pretty sure that it was John Maynard Smith. I had expected to see great scientists at SSE, but I was shocked when Maynard Smith sat down right next to me. He looked at me and said, “You’re Joe Graves, aren’t you?” I told him yes. What he said next threw me for a loop: “I’ve read some of your papers. They’re pretty good. I hear you are going to work with Michael Rose.” Once again, I told him yes. Then he looked into my eyes with a quasi-serious grin on his face and said, “Well, you’ll have to move him to the left a little bit.” With that he got up and headed off to chat with some other folks. Later that day, or it might have been the next, George C. Williams came up to me and introduced himself. He also was familiar with some of my work. This behavior made it pretty clear to me that I was one of the few (perhaps the only) African American evolutionary biologists they had ever met. It would be a year or two before the title of Black Darwin would be bestowed upon me, but 1988 was now a historic year for the discipline, and more African Americans would follow me into the field in short order.