THE FIRST TASK OF MY POSTDOCTORAL WORK WAS TO PROVE to Michael Rose that flight was a reliable characteristic for demonstrating physiological alterations that resulted from selection for postponed senescence. Michael had actually seen me measuring the wing beats of tethered flies in Leo’s lab. Still, his group, particularly Phil Service, had not been able to do so with their flies. The difference between the postponed-aging flies in the two laboratories was important. Quite frankly, Michael’s experimental protocol was better. He had created five replicated populations of long-lived flies (called O stocks) that had been maintained with effective population sizes of more than two thousand throughout their evolution. His O stocks also lived in plexiglass cages during their adult lives. In contrast, the Luckinbill long-lived stocks had only two replicates and lived in milk bottles at effective population sizes of about three hundred flies.
To my great surprise, the first time I tried to measure flight duration in Rose flies, I couldn’t do it either. Though one might think this represented a great scientific setback, in actuality it was an opportunity. Experiments conducted using the same protocols on the same materials should give the same results. I needed to figure out what was different between the protocols I had used to get Leo’s flies to fly and the ones I was attempting to use with Michael’s. With a little bit of thought the important difference in the protocols became obvious. I had anesthetized Leo’s flies with diethyl ether, and Michael used carbon dioxide (CO2) to knock out his flies. The problem with using CO2 is that it has a lingering effect on the fly’s blood pH (making it more acidic), and this in turn incapacitates its flight enzymes for a longer period of time. As soon as I obtained some ether, I was able to repeat in the Rose stocks the results on flight duration that I had achieved in the Luckinbill stocks. In addition, since the Rose flies were fivefold replicated, the statistical differences between the short-lived and long-lived lines were much easier to demonstrate. In showing that flies that lived longer also had superior physiological performance, we postulated that treatments that could postpone aging in reality would increase individuals’ health span (that is, the length of time an individual is healthy). Our first paper on this result received a lot of attention, eventually even getting coverage in such popular magazines as Discover, The Economist, Life, and Scientific American. The work was also the primary subject of the Black Issues in Higher Education profile article “The Evolutionary Genius of Dr. Joseph L. Graves Jr.,” published in January 1991. Eventually the idea that scientists at UC Irvine were attempting to prolong the life of fruit flies came to the attention of the scriptwriters for NBC’s The Tonight Show and showed up in one of Jay Leno’s opening monologues, which I’m paraphrasing: “I hear scientists in Irvine are trying to extend the life of fruit flies.… Aren’t we trying to get rid of those things?” Leno was referring to the Mediterranean fruit fly outbreak that had caused great damage to California citrus produce.
Our results spurred me to pursue more deeply the connections between postponed aging, physiology, and energy reserves. I began this work as a postdoc and completed it as an assistant professor. The paper in which I examined the relationship between aging, flight, desiccation resistance, and glycogen content was published in the prestigious journal Physiological Zoology (now called Physiological and Biochemical Zoology) in 1992.1 The publication of this paper led me to be recognized as one of the leading young researchers in the nascent field of evolutionary physiology. That same year I would speak at a symposium devoted to evolutionary physiology at the annual meeting of the SSE.
Despite the success of my experimental work as a postdoc, there was no guarantee that I would be hired in a faculty position at a top-tier research institution. The University of California’s President’s Postdoctoral Fellowship was designed to help locate and recruit talented early-career URM and female scientists with the goal of eventually hiring them into the UC system as faculty members, but this goal was not well known to or necessarily supported by the STEM departments that had to do the hiring. In my case, my first application to be hired as a faculty member at UC Irvine was mysteriously “lost.” When the applications came into the department, mine was not among them. Michael called me up, livid, almost screaming over the phone, “Joe, how could you have not gotten your application in on time?” I told him that in fact I had turned it in on time. An internal departmental investigation determined that my application had been received on time and misplaced, though it didn’t explain how such a thing could have happened. Whatever the reason, my supporters in the department were really upset and insisted that the next time the competition was held, my application be handled correctly.
During the following year I was interviewing for faculty positions across the country at top-tier institutions (both public and private). What followed was an amazing litany of excuses for not offering me an appointment. In one case, I had a verbal offer from a dean. We had discussed my salary, and I had been shown my research space. A week later, this same dean called me up, almost incapable of saying the words, to tell me that he had to rescind his verbal offer. He told me he didn’t believe these things happened anymore. So at the end of my interview tour, I had one employment offer from a midtier institution and an upcoming job seminar at Irvine, where the department was ambivalent about having me join the faculty.
The ambivalence was related to several issues. First was a long-standing tradition of not hiring postdocs as faculty at the same institution where they did their postdoctoral research. Second was the claim that I was simply not qualified to be on the faculty. Third were questions about how acceptable I was to some of the white faculty, one of whom actually approached Michael and asked him if he could convince me to “not act so Black.” Going into the job seminar, I knew that there were eight people solidly in my corner, six people solidly opposed to having me on the faculty, and at least ten people in the department who simply didn’t know me or my work.
Candidates for faculty positions are expected to present a seminar explaining their research. As I prepared for the seminar, I was well aware who the leader of the opposition to my appointment was. I prepared for my talk by examining every possible objection to my research methods and conclusions. When I was done, the opposition leader’s hand was the first to go up in the question-and-answer period. I am sure he expected me to avoid him; instead, he was the first person I called on. I thanked him for his question and demonstrated that each of his criticisms of my work and conclusions was invalid. When my answer concluded, he simply sat down, resigned that he had failed to show I was unqualified to be a faculty member. The final vote included only three persons opposed to my appointment, which meant I had won over all the undecided individuals and three of those who had been opposed to me from the start.
When I joined the faculty at Irvine there were only a handful of African American faculty in STEM areas, and there were probably twice as many Chicanos (Mexican Americans) as African Americans. The most senior and influential Chicano faculty member in my department was Eloy Rodriguez. Eloy met with me very soon after my appointment and began to bring me into the programs designed to increase the participation of URM students in science. One of the best decisions we made in that regard was to get both of us assigned to teach the Introductory Biology course (the third member of the team of instructors for this course was National Academy of Sciences member Francisco Ayala). The idea was to expose the students in the introductory course to people who were among the most prestigious and talented scientists and instructors on the campus and who were also URMs. At this time it was well-known that the attrition of URM students from the STEM pipeline began in their first courses at the university.
My first lecture in the course was momentous. I was wearing a Public Enemy T-shirt, complete with their crosshairs logo on the back. There were over two hundred students in the lecture hall. The lecture was organized around Dobzhansky’s dictum “Nothing in biology makes sense except in the light of evolution”; it is the means to understand the diversity of animal life. At its conclusion, dozens of students waited to speak to me. I could see in the back a tall, somewhat shy-looking student patiently waiting for all the students who were just trying to get noticed by the instructor to finish. I will never forget what he said to me: “Dr. Graves, please tell me what I need to do to become a scientist just like you.” I told him to come by my office and we could talk about getting him involved in research. Luciano Matzkin went on to earn his PhD in Drosophila evolutionary genetics and is now a professor at the University of Arizona. He began working with me as a freshman and stayed with my research group until I left UC Irvine.
Luciano was just one of many undergraduates who joined my research group. At its height, it numbered over fifty undergraduates. I directed the senior undergraduate thesis of at least twenty students who went on to professional careers in science and medicine. By 1993 I was the lead principal investigator on a project funded by the National Institute on Aging attempting to repeat the Rose postponed-aging experiment in different species of fruit flies (e.g., Drosophila simulans, D. pseudoobscura, the cactophilic species D. arizonae, and D. virilis). The goal of that project was to determine whether experimental evolution would also alter the pattern of aging in other species of flies. We proposed that that must be the case, but a hypothesis and actual evidence supporting it are not the same thing. The expansion of the work in my laboratory was made possible by the dedication of my undergraduate research students. These experiments showed that evolution for postponed senescence did occur in other species of fruit flies, and they also let us examine some of the underlying genetics allowing this to occur.2
In 1992 UC Irvine was awarded one of the first National Science Foundation Alliances for Minority Participation awards, the California Alliance for Minority Participation (CAMP). These would later be renamed the Louis Stokes Alliances for Minority Participation, which are now featured all over the country. Eloy was one of the principal investigators for our CAMP program. It has long been known that the majority of URM students in post-K–12 education are found in the nation’s community colleges, where they complete the first two years of their undergraduate degree. Transfer academies are intended to ease their transition to four-year campuses. Eloy asked me to direct the CAMP transfer academy to bring those students to campus in the summer before they entered their junior year to acclimate them to what academic work would be like at UC Irvine and to prepare them to navigate a potentially hostile campus environment. My experience with the CAMP transfer academy would structure all the programs I would subsequently design and direct to provide URM students training and guidance at historically white institutions (HWIs).
Of all my student-related accomplishments at Irvine, one of the most important resulted from a conversation I had with a member of our graduate admissions committee. This faculty member came to me with an application to our graduate program that the committee was considering. The applicant was an African American, and if he was admitted he would be the first in the program’s history. At this time admission to our program was quite competitive. Most people in the know ranked our department and the one at the University of Chicago as the two strongest in the nation, perhaps the world. Upon reviewing the application my response was simple: “You should admit this student; he has a better record than I did when I was at the same stage of my career.” They agreed, and in the fall class I had the opportunity to meet Paul Turner. It is not an understatement to say that that fall class in our department was amazing, but Paul performed as well as any of those admitted. He decided to join the laboratory of my colleague Richard Lenski, and when Richard moved to Michigan State, Paul went with him. By my calculation Paul is the third African American to have earned a PhD in evolutionary biology. Eventually Paul became an assistant professor in ecology and evolution at Yale University, where he earned the rank of full professor and was department chair for a while. In 2019 he was awarded the Rachel Carson Endowed Chair in Ecology and Evolutionary Biology at Yale University. In that same year he was elected to both the American Academy of Arts and Sciences and the National Academy of Sciences. I am absolutely sure that if Paul had not been admitted to our program, he would have gone somewhere else and done something just as fantastic. However, I am proud of my role in bringing him into the field.
The moniker “Black Darwin” was bestowed on me by African American undergraduates at UC Irvine because of my work championing evolutionary thinking in the curriculum, developing and directing programs to benefit URM students, and providing research mentorship for students across racially subordinated groups at the college. This work was part of the reason I was elected a fellow of the council of the American Association for the Advancement of Science (AAAS) in biological sciences in 1994. The entire citation read, “For original contributions to the evolutionary theory of aging through an examination of its population genetic and physiological basis and for the successful mentoring of minority graduate and undergraduate students in the biological sciences.” This is one of the highest honors any scientist can earn in America (short of election to the National Academy of Sciences).
ESTABLISHING THAT SEVERAL EVOLUTIONARILY SEPARATED SPECIES of fruit flies all responded to selection for postponed senescence was important. However, it is a long way from demonstrating that the same is true in mammals, because their physiology is quite different from that of insects. I remember having to explain that to some European molecular biologists at an international meeting on the biology of aging held at the Beckman Center (West Coast headquarters of the National Academy of Sciences) next to the UC Irvine campus. During my talk I was explaining that flies from the Rose laboratory that had been selected for increased starvation resistance had also increased their fat reserves and also displayed longer life spans compared to the control populations. I was interrupted by an audience member who stated that this simply wasn’t possible, as it was a well-known fact that increased fat deposits resulted in lowering life span because of atherosclerosis (hardening of the arteries). Doing my best to hide my frustration, I explained as patiently as I could that fruit flies do not have that problem because insects have open circulatory systems (thus, no arteries to clog). This kind of response was par for the course during this period of my career. Too often, the audience saw a “Black man” in front of them rather than a talented scientist. It got to the point in many of these conferences that Michael began to repeat the things I said during the discussion because he could see that few people were listening to me. Once he said the same thing I had (often without changing the wording at all), the comment took on new significance to the white males in the audience.
The comment about the open and closed circulatory systems got me to think more deeply about how much the research we were conducting in flies could be applied to understanding aging-related phenomena in mammals (particularly humans). The evolutionary theory holds that aging results from the fact that early in the history of life, organisms developed a dichotomy between cells that were primarily involved with maintenance of the body (somatic cells) and those involved in reproduction (germ cells). Insects and mammals share this feature from an ancient common ancestor (about 531.5 million years ago), so the population-genetic mechanisms associated with senescence (mutation accumulation and antagonistic pleiotropy) operate in both groups. We had already established this theoretically, but it was important to work out the details of how it operated physiologically if any successful antiaging treatments could be devised. At this point I recognized that the negative genetic correlation between longevity and reproductive effort produced by selection was paralleled, in some cases, by some environmental manipulations. Specifically, it had been shown that life span could be increased by dietary (caloric) restriction in mammals, including primates. I reasoned that the increase in life span was explained as an incidental consequence of lower reproductive effort. There were good reasons for this response to have evolved in nature, because animals often faced food shortages. Thus, reducing reproductive output could have been an adaptation that enhanced fitness in some species that faced uncertain food supplies. This condition probably evolved independently in a wide variety of animal groups. My idea was supported by evidence from gerontological work on life-span extension, by reproductive physiologists concerned with factors that affect fertility, and by life-history studies in various groups of mammals.3
I also recognized the practical difficulties of implementing caloric restriction strategies as a way to increase human life span. First, how long a life span is extended is directly related to how many calories are cut from the diet. Maximum impacts had been shown in some primate species with 50 percent caloric restriction (which also completely shut down their reproduction). Less caloric restriction resulted in less life-span extension. In addition, there is also the problem of when to introduce caloric restriction. Certainly one would not want to cut calories for adolescents, but the later in life the caloric restriction is implemented, the less impact it can have. And further, the issue is also complicated by the fact that altering human behavior, particularly with regard to our eating habits, has been shown to be particularly difficult. This is why most diets fail. The difficulty lies in the fact that we evolved to seek high-calorie foods, and the centers in our brains associated with various reward systems (food, sex, alcohol, other drugs) are all interrelated.
Still, what caloric restriction research tells us is that individuals who have the means and mental fortitude to control how much and what kind of food they eat can significantly extend their life and health span. Obviously, for the vast majority of people in the world (including the poor of the United States) who experience food deserts, caloric restriction will never be of much significance. For these people, our task is to provide them with a steady supply of nutritious food.
DESPITE MY ACADEMIC SUCCESS AT IRVINE, I WAS STILL WRESTLING with my personal demons. The greatest of these was imposter syndrome. On the one hand, I had established that as “Black Darwin” I could hold my own scientifically with some of the greatest evolutionary thinkers on the planet, people like Francisco Ayala, Michael Rose, Larry Mueller, Steve Frank, Richard Lenski, and others. My scholarly work was also gaining respect among the few African American professors on campus (e.g., Joe White, psychology; Anthony James, molecular biology). It was at this time in my career that I began to think more seriously about how my knowledge and skills as an evolutionary biologist might be turned to addressing the issues that were contributing to the oppression of Black people.
However, I was still a person unmoored from his Black working-class origins and his professional life, which was so completely submerged in European American culture and perspectives. That contradiction came out in bad ways, eroding my personal relationships. In many ways the pressure of my position in the academy was twisting me into someone extremely self-centered—someone who, in hindsight, I really don’t like. I vacillated between actions that appeared supremely arrogant and actions that displayed an utter lack of self-confidence. It was the struggle between, on the one hand, the values instilled in me by a mother who worked in rich people’s houses during the day, pulled a factory shift at night, and was always in church on Sunday and, on the other hand, my inflated vision of myself as an “evolutionary genius” that would shape the next stage of my career.