Add to bookbag
Authors : Virginia J. Vitzthum, Alan G. Fix, Amy Livingstone
Title: The Four-Field Anthropology and Multi-faceted Life of Frank B. Livingstone
Publication Info: Ann Arbor, MI: MPublishing, University of Michigan Library
2006
Availability:

This work is protected by copyright and may be linked to without seeking permission. Permission must be received for subsequent distribution in print or electronically. Please contact mpub-help@umich.edu for more information.

Source: The Four-Field Anthropology and Multi-faceted Life of Frank B. Livingstone
Virginia J. Vitzthum, Alan G. Fix, Amy Livingstone


vol. 16, no. 1, 2006
Issue title: Retrospectives: Works and Lives of Michigan Anthropologists
URL: http://hdl.handle.net/2027/spo.0522508.0016.107
PDF: Link to full PDF [303kb ]

The Four-Field Anthropology and Multi-Faceted Life of Frank B. Livingstone

Virginia J. Vitzthum

Anthropology Department, Indiana University, Bloomington

Alan G. Fix

Department of Anthropology, University of California, Riverside

Amy Livingstone

History Department, Wittenberg University, Springfield, OH

Most graduate students only dream of writing a dissertation that really makes a difference; Frank Livingstone actually did it! Based on J. B. S. Haldane’s 1949 suggestion that the lethal anemia caused by a double-dose of the hemoglobin S (sickle cell) gene could be balanced by a protective advantage against malaria in heterozygous individuals, and Allison’s 1954 demonstration of such an advantage, Livingstone set out to study sickling in West Africa for his doctoral research. One of his advisors at the University of Michigan was James V. Neel, the geneticist who had elucidated the inheritance of the sickle cell gene.

Frank designed his research to examine the effects of sickling on child growth. Imagine his shock, as he collected blood samples and tested for the presence of S alleles, to find that there were none in that population! Frank’s whole project depended on examining the relationship between hemoglobin S and growth. No S, no relationship, no dissertation.

He was sampling individuals from Liberia, an area not previously characterized for sickling. However, the region was highly malarious (holoendemic) and therefore, according to the malaria hypothesis, the sickle cell gene should have high selective value and be present in high frequency. After ruling out the obvious possibility of some failure of his microscopic identification of sickle cell, he was left with a quandary. Was the malaria hypothesis wrong? Unlikely, since it was strongly supported by several lines of evidence. Or was something different about Liberia? Frank figured it out and produced a dissertation demonstrating the power of an anthropological approach to understanding human variation and evolution. This approach continued to yield numerous insights throughout his career.

Published in 1958 in American Anthropologist as “Anthropological Implications of Sickle Cell Gene Distribution in West Africa,” Frank’s solution became an almost instant classic. It has been reprinted many times and continues to inform research on malaria and human polymorphisms. The logic was simple—once it was explained—but required linking biology (genetics, evolutionary theory, and disease ecology) with anthropology (archaeology, linguistics, subsistence regimes, and settlement practices).

The key question was why the S gene was absent from Liberia, even though malaria was holoendemic. The answer lies in the difference in timing between the introductions of malaria and the sickle cell gene. Transmission of malaria to humans depends on the disease-causing protozoan parasite (Plasmodium falciparum in this case) and the vector (members of the Anopheles gambiae species complex) that transmits the parasite as it feeds on human blood. Since mosquitoes and plasmodia rapidly colonize favorable habitats, malaria can spread quickly. Genes, on the other hand, even when as advantageous as the S gene, increase in frequency only very slowly. During each human generation, roughly 27 years, strong natural selection for heterozygotes gradually adds to the number of S genes in the population. Depending on the intensity of the selection, it may take hundreds or thousands of years to reach an equilibrium frequency of S, in which loss of the allele through the deaths of those who are homozygous for the sickle gene is balanced by its retention due to protection against lethal malaria in heterozygotes.

What if malaria was a recent arrival in Liberia, not yet allowing sufficient time for the build-up of the sickle gene? If so, then it needed to be determined why malaria was recent in Liberia, as opposed to many other areas in Africa where both malaria and sickle cell were well established. Frank’s solution was to show that West African Anopheles vectors of malaria require stagnant, sun-lit pools of water in order to breed. These conditions are not found in dense forests where streams are shaded and usually fast-moving. If Liberia had been forested until recently, then mosquito transmission would not have taken place, and malaria would not have been present. Frank then showed that, indeed, several linguistic groups heavily represented in his Liberian population were relatively late adopters of agriculture. Previously living as hunter-gatherers in uncut forests, these populations would have been malaria free, and hemoglobin S would have had no selective advantage. However, with the adoption of agriculture, the forest was cleared and compacted soils were created where rain could pool in the sun. These farmers, now living at higher density in clearings near their fields, had provided an optimum environment, including an attractive host, for mosquitoes and plasmodia.

Having established these themes in his dissertation, Frank followed them throughout his long career. He became a recognized authority on malaria and human genetic polymorphism, championed the role of natural selection in understanding the distribution of human biological variation, emphasized the role of culture and environment in causing genetic change, and argued against extreme forms of sociobiology that tried to put genes in the causal role. One of his most influential papers, “On the non-existence of human races,” also followed from this basic logic. If human variation is primarily a response to environmental selective conditions, and selection differs for different genetic loci, we should not expect to see neatly patterned human racial isolates. Rather, we should expect what is, in fact, the case: geographic gradients (clines) of gene frequencies arrayed across human populations.

In addition to these foci, Frank had an enlarged curiosity that ranged from the mathematics of kinship to linguistic abilities in the australopithecines. The rich and broad intellectual adventure that was Frank Livingstone cannot be encapsulated by these few pages. Explore Frank’s work for yourself—even the ideas that some would say are passé (for example, the debates among lumpers v. splitters or selectionists v. neutralists). Have a beer (or two or three!) while poring over the “Frankfest” festschrift volume (Vitzthum and O’Rourke 2003) to get a greater appreciation of Frank’s lasting contributions to anthropology. In the meantime, we will strive to give you a small measure of this kind and gentle man, this rigorous imaginative delighted scholar, this helluva hockey player...

Beginnings

In his presentation at the Frankfest in 2002, Theodore (Ted) Steegmann (Buffalo, SUNY) carried the audience back nearly 50 years to an America bursting with optimism and overconfidence, enthusiastically embracing the promises of science and technology. The atom had been cracked, all mysteries seemed penetrable, all problems eventually solvable. The U.S. National Science Foundation was created in 1950 to fund the basic research needed to achieve such goals, and both public and private American educational institutions flourished as never before (Steegmann 2003).

The University of Michigan was no small beneficiary of these developments, and physical anthropology rose on the wave. Under the direction of James Spuhler, the physical anthropology laboratory shared the university’s former maternity hospital with the Human Genetics Department (administratively within the Medical School) directed by James Neel and Jack Schull. Ted described how the exciting intellectual atmosphere fostered by the proximity of these colleagues and their students was furthered by exchanges with those cultural anthropologists—Leslie White, Elman Service, and Marshall Sahlins—who “promoted a structural-functional view that emphasized cultural evolution and societies as adaptive systems” (Steegmann 2003).

From this perhaps unique combination of approaches and expertise emerged a biocultural perspective on human evolution that rejected typological thinking in favor of evolutionary genetics and rigorous quantitative testing. Frank contributed mightily to this effort at the University of Michigan for more than 40 years. How did he come to be there?

Frank Brown Livingstone was born to Margery Brown Livingstone and Guy P. Livingstone on December 8, 1928, in Winchester, MA, an origin that accented his speech even after half a century in the Midwest. His life-long passions for fly-fishing and ice hockey can be traced to ample opportunities for both on the New England waters of his boyhood. Little could deter Frank from pursuing these joys. He first traveled to Liberia for his dissertation fieldwork on a passenger boat with plenty of room for “research equipment.” But on his second trip, this time by air, he had to stuff the fishing gear in his pockets so as to make the airline’s weight limit—one wonders what was left behind in favor of the tackle!

In 1946, Frank graduated from Winchester High School and entered Harvard because his “mom thought it was a good school.” Although his Bachelors degree was in mathematics (1950), it was an anthropology course taken during his junior year at Harvard that sparked Frank’s lifelong intellectual passion. (Harvard’s anthropology department would come to offer him a job in the 1960s. Frank turned them down, repeating with great glee to his family the assessment by that department when he had sought graduate training there: “He wasn’t Harvard material!”).

After two years in the U.S. Army at the Chemical Center in Fort Dix, NJ, Frank began graduate studies in anthropology at the University of Michigan “because it was the cheapest.” To supplement the GI Bill’s educational benefits, which barely covered tuition and expenses, Schull and Neel hired Frank as a human calculator at $1/hour (once upon a time, “calculator” referred only to a person, and ‘electronic’ was the necessary modifier).

In the fall of 1953, Frank and Ronald Nishiyama, a U of M medical student, struck up a friendship that was to last for nearly 52 years—despite the fact that they shared an apartment for two years while students! As luck would have it, that apartment was in the same building as a young mover and shaker in the Democratic Party. Carol Ludington and Frank married on August 13, 1960. Their daughter, Amy, was born the following summer.

Frank’s choice of a dissertation topic was born of a driving intellectual curiosity. In his own words: “Figuring out what caused hemoglobin variation was the state of the science, the most cutting edge issue of the time. Figuring that out would be a major advancement.” Frank was awarded master’s (1955) and doctoral (1957) degrees in anthropology by the University of Michigan, and he joined its anthropology faculty in 1958. In 1962, Spuhler and Livingstone were joined by a third physical anthropologist, Ernst Goldschmidt, destined to become another life-long member of the extended Livingstone clan.

Only a few students come to study at Michigan’s Anthropology Department because of a specific faculty member. But I (AGF) knew exactly why I was willing to put up with Ann Arbor winters. My advice to undergraduates looking for a graduate program is to find an article they wish they had written and to go study with that person. That exactly describes my own choice. I went to the University of Michigan in 1964 because Livingstone’s study of the anthropology of sickle cell was the kind of work I wanted to do. When I arrived in Ann Arbor, I went to see Frank. Sporting a bachelor’s in anthropology, I thought I was ready to start doing research. Frank informed me that I would, instead, be taking courses, and sent me off to Jim Spuhler for advising. Thus began a process of both learning but also unlearning a lot of what I thought I knew. Although I have never quite produced my own equivalent of the “Anthropological implications of sickle cell gene distribution in West Africa,” Frank’s model of the productive interplay of genetics, ecology, and culture has always been at the center of my thinking.

Most everyone who studied anthropology at Michigan from the 1960s through the 1990s seems to have a “first time I saw Frank” story, and it usually involves beer. For me (VJV), it was early September 1977, late afternoon. The new graduate student orientation had just ended, and the faculty had invited us for refreshments on the inner patio of the Student Union. Round metal tables with big sun umbrellas and rusted folding chairs contrasted strangely with the red brick and carved stone of the stately building. The sun was warm, the place was packed, and I wasn’t sure I recognized anyone. And then I heard Frank, in a broad Boston-tinged drawl—“Mimi’s gawl-darned parrot swooped down on my head!”—gesticulating the flight of the bird while pouring a pitcher and passing glasses—“and then it shit on me! Right there! Right on my head!”—everyone’s laughing, and somehow Frank didn’t slop a drop of beer and he didn’t miss a beat. “And what I thought...Really, I was sitting there at Loring’s table, and what I thought was, What’s in this stew?” We laughed so hard we cried; it wasn’t just the story, it was the way of telling, the gusto, the mirth. Of course, as many know, Mimi Brace was an inspired chef; Frank just couldn’t resist stretching a tale for a good laugh.

In Ann Arbor, beginnings for new faculty can be tough, as Lisa Sattenspiel (University of Missouri) learned. Fortunately for many, the Livingstones shared their hospitality as easily as their laughter.

My first post–Ph.D. job was as a temporary faculty member at Michigan from 1984–1987 to replace Frank while he was department chair...Had I known what I was stepping into, I’m not sure I would have had the gumption to take it on. But ignorance is bliss, so in the summer of 1984 I set off cross-country with my husband, two-year-old son, two cats, and all our worldly belongings. On our first day in Ann Arbor we experienced, like many others I’m sure, Carol and Frank’s extreme generosity. Although we found an apartment pretty quickly, it was not going to be available for two to three weeks. But our truck had to be returned in two to three days. Frank stepped in to help us out in our time of dire need, offering us his basement to store all our goods and then helping us transport everything to our new apartment once it became available.

Scientist

In this section, Alan Fix, Frank’s first graduate student,explores Livingstone’s research career, both documenting the themes and exploring some of the apparent excursions.

Distribution of Abnormal Hemoglobin Variants Several of Frank’s earliest papers documented the distribution of various hemoglobin variants in West Africa, based on his collaboration with James Neel and other human geneticists (Hiernaux et al. 1956; Livingstone et al. 1956; Livingstone 1958b, 1960c; Olesen 1959; Sutton et al. 1959). The geographic pattern of genetic variants is critical to understanding the evolutionary causes for their occurrence, and Frank initially contributed data based on his field research. As he turned to modeling these patterns, he saw the need for a compendium of frequency data and provided it in his 1967 book, Abnormal Hemoglobins in Human Populations. He updated this work first in 1973 as Data on the Abnormal Hemoglobins..., published by the U of M Museum of Anthropology in their Technical Reports series, and again in 1985 as Frequencies of the Hemoglobin Variants..., for Oxford University Press. Although these are primarily lists of gene frequencies, the first book, in particular, contains a long discussion of the population genetics of red cell variants and a review of the malaria hypothesis for each of the major regions of the world.

Explaining the Distribution of Hemoglobin Variants Using population genetics theory to explain the geographic distribution of these genes played a central role in Frank’s work. Clearly the malaria hypothesis was the core idea guiding his research. From his 1957 paper on “Sickling and Malaria” through his dissertation, and in numerous subsequent publications (1960a, 1961, 1964a, 1964b, 1969b, 1971, 1983, 1989a, 1989b), he showed how to balance the human hemoglobin polymorphisms. Even outside anthropology he was recognized as an expert in this field, being invited to contribute “Malaria and Human Polymorphisms” (1971) to the Annual Review of Genetics as well as to contribute to several discussions and reviews of hemoglobin variants (e.g., 1983).

Sickle cell hemoglobin in humans is probably the best-known example of a “balanced polymorphism.” When two opposed selective factors balance each other, equilibrium gene frequencies are attained. This equilibrium depends entirely on the magnitude of selection. Thus, if s1 measures selection against the hemoglobin AA “normal” individual due to increased susceptibility to malaria, and s3 represents selection against the SS homozygotes due to anemia’s lethality, then qS = s1 /(s1+ s3) where qS is the equilibrium frequency of the sickle cell allele. So, for instance, in several areas of Africa prior to medical intervention, sickle cell anemia was usually a lethal disease; few individuals survived to pass on their genes (s3 = 1.0). At the same time, the protection against holoendemic malaria in heterozygotes meant that the selective differential against homozygotes was often 25% (s1 = 0.25). Thus, the expected equilibrium frequency (qS) was 0.2, a commonly observed value in many African populations. Explaining the frequency of sickle cell in any particular environment would entail assessing the long-term selective impact of malaria—except in those areas where insufficient time had elapsed for selection to increase the allele to equilibrium frequency. It was exactly this situation that Livingstone identified in Liberia and in some other parts of West Africa.

Understanding the process whereby the sickle cell gene (Hb*S) and the other malaria-related blood polymorphisms such as Hb*C, Hb*E, thalassemia, glucose-6-phosphate dehydrogenase deficiency, and ovalocytosis spread across the tropical malarious regions of the world became a major research interest for Frank. For natural selection to favor the sickle cell gene, it first must be present in the population. The origin of the allele, as for all primary genetic variants, was by mutation, a substitution of one DNA base for another. But such point mutations are exceedingly rare, and Hb*S may have originated only once, or perhaps as many as four times, in human history. Since it is now widely spread not only in Africa but also in parts of Arabia, India, etc., the mutation(s) must have diffused via gene flow.

As early as 1960, Frank was describing “the wave of advance of an advantageous gene.” He revisited this model in many papers extending nearly to the end of his career. Beginning in 1969, he employed computer simulation to model the process, first using a now extinct language, Michigan Algorithm Decoder (MAD), then FORTRAN. I have recently reviewed this work for the Livingstone Festschrift, and this can be consulted for a more detailed history of Frank’s simulation work (Fix 2003).

In the manner of Darwin himself, Livingstone saw natural selection as evolution’s primary force, the principle that explained the most salient features of human variation. Few would now disagree regarding sickle cell and some other hemoglobin variants. But the use of purported biological “markers” to reconstruct population history has also been a long-standing tradition in physical anthropology. On this view, genetic variation arises from random mutations that are neutral in their effects on survival and reproduction. The presence of particular alleles in a population reflects only random genetic drift and/or gene flow from other populations. Perhaps the best-known example of this genetic taxonomy is the “big book” of Cavalli-Sforza and colleagues, The History and Geography of Human Genes (1994). It shows branching tree diagrams said to represent genetic affinities of the world’s populations. However, such phylogenetic trees presume that shared genetic characters are due only to common ancestry. But natural selection can and does increase or decrease gene frequencies, regardless of descent. For example, whatever the closeness of their ancestry, two populations living in very different malarial environments might differ greatly in sickle-cell frequencies.

Disregarding selection’s role, Cavalli-Sforza and his collaborators have used blood group and HLA (Human Leucocyte Antigen) gene frequency data to track ostensible genetic history. This approach simply “baffled” Frank (Livingstone 1995, personal email communication to author). There was just too much evidence and theory for natural selection’s role in determining variation in these genetic systems to treat their alleles as passive markers of population movement and affinity (Livingstone 1991).

Frank also rejected racial classification and for much the same reason (Livingstone 1962a). Some populations of Africans, Arabians, and South Asians—ostensibly representing three different “races”—nonetheless share high frequencies of the sickle cell gene. According to the racial hypothesis, this can’t be because they share racial affinity, for that has already been ruled out of court. The evident similarities in Hb*S frequency must, therefore, be caused by adaptation to a common (malarial) environment, not descent. This was Frank’s very point. By the same token, these populations, like all others, are indeed connected by networks of gene flow, and this too refutes the racial hypothesis. The bottom line is that various gene loci show different, often clinal, patterns of variation in response to different selective regimes.

Natural Selection and Disease The malaria hypothesis provided the classic textbook case for the important role of disease as a selective agent. However, Haldane’s suggestions regarding malaria and sickle cell were also part of a broader argument (Haldane 1949). This related to changes in human ecology brought about by the adoption of settled agriculture to an increasing impact of disease on human evolution. Frank’s thesis had beautifully demonstrated the details of the Hb*S/ malaria/agriculture links. But beginning as early as 1960 in “Natural Selection, Disease, and Ongoing Human Evolution, as Illustrated by the ABO Blood Groups” he also championed natural selection via disease as the primary cause of variation in other genetic systems, especially the blood groups (Livingstone 1960b). Although the statistical demonstration of selective differentials had not been very successful, as he pointed out in a review of selection and the “standard genetic markers,” there were nonetheless good reasons to believe that selection has shaped variation among human populations (Livingstone 1980b). Mechanisms such as frequency-dependent selection, or temporally variable selection, could account for blood groups’ polymorphisms (Livingstone 1978, 1980b).

Frequency dependent selection is the result of predators targeting the most common prey, thereby allowing rare variants to increase in frequency, whereupon they, in turn, become the targeted prey. Because advantage is based on frequency, variants oscillate from rare to common over time. In the context of disease selection, parasites adapt to the antigens or other cell attributes of the host. Any modification from the “normal” condition might be less attractive or beneficial to the invading parasite. Thus “abnormal” or rare alleles would protect the host. As these initially rare genes increased in frequency, the disease organisms would adapt to the new host antigens, advantaging alternative rare variants.

Loci with alleles subject to temporally variable selection should also be polymorphic. Again, the statistical evidence for such selection is skimpy, but the theory is compelling. If, for instance, a human blood antigen is similar to the antigenic structure of a disease organism, say smallpox, then it would be difficult for the human host to mount a strong immune response to the parasite; the antibody would also be directed at the host’s own red blood cells. The cyclical nature of epidemic diseases would result in periodic selection against the antigen. Other epidemic diseases, occurring at different times, might select against other alleles, thus maintaining polymorphism at the locus.

While these arguments may seem excessively theoretical, they are bolstered by a number of empirical demonstrations of selection for blood group alleles. One example is the Duffy negative phenotype, a case that Frank reviewed (1984). In parts of West Africa, almost the entire population lacks both the a and b antigens of the Duffy blood group system, whereas they are polymorphic in many human populations. These antigens, residing on the surface of the red blood cell, are the receptors that allow the Plasmodium vivax species of malaria to recognize its host tissue (Miller et al. 1975). Lacking the antigens, Duffy negative individuals are essentially completely resistant to this form of malaria.

It finally comes down to a matter of emphasis. Livingstone argued that selection could not be ignored. Human variation is not simply the random drift of alleles but, rather, has been profoundly shaped by natural selection. Studies lumping dozens of loci to reconstruct population affinities and history both compound and conflate the evolutionary forces of selection, drift, and gene flow. Even where many loci are neutral, if several are not, estimates of historical population “affinity” will be contaminated. Frank’s selectionist “voice-in-the-wilderness” has been a powerful reminder of this fact, particularly now that one of the pioneers of mtDNA studies, Douglas Wallace, has provided evidence that some haplotypes of this widely utilized, ostensibly “neutral” genome may in fact show selective differentials (Ruiz-Pesini et al. 2004). This should not be surprising, since the mitochondrion is the energy-producing organelle of the cell. Wallace and his colleagues point out that some types are more efficient energy producers while others may generate more heat. In a cold climate, the heat-producing haplotypes ought to be favored. It turns out that one of these, haplotype A, is indeed prevalent in the cold northern portions of North America. Now since the entire mitochondrial genome behaves as a single linked locus, such selection would “carry along” all mtDNA loci. Thus natural selection may, in fact, be impacting the distribution of this universally employed “neutral marker” system, with the result that false population histories will be inferred.

The “Other” Forces: Founder Effect and Gene Flow Population genetic theory identifies four forces of evolution: mutation, natural selection, genetic drift, and gene flow. Although Frank emphasized selection as the most important determinant of human genetic variation, he often pointed out that in many contexts the other forces played equally crucial roles. A salient case was the so-called Brandywine isolate, a small population of African descent living in a historically non-malarial environment in southern Maryland, yet possessing a relatively high frequency of Hb*S (0.1). Livingstone pointed out that the population originated in a few families in the early 18th century, and had remained highly endogamous since then (1969a). These are exactly the conditions promoting founder effect, one form of genetic drift. He then showed by simulation that the anomalously high sickle cell frequency would indeed be likely. Later he asked the question, “How many deleterious genes in polymorphic frequencies would be expected in a large population that resulted from rapid growth from a few founders?” (Livingstone 1980a:97). Using simulation, he found that such a population history could lead to a surprising number of lethal genes reaching relatively high frequency.

Livingstone’s extensive simulation experiments on the spread of hemoglobin variants, although based on the adaptive advantage of these alleles, led him to the conclusion that long-range migration, or gene flow, was a critical factor. The wave of advance of an advantageous gene clearly depends on gene flow; the allele will not increase in a population until it is has been introduced either through mutation—an extremely rare event—or gene flow from a neighboring population. Interestingly, Frank found that gene flow between closely neighboring populations was not sufficient to allow rapid diffusion of these alleles (e.g., 1969b, 1989a). However, a small amount of genetic exchange between widely separated populations brought his models into conformity with the time line for malarial selection. “[A] minimal amount of long range gene flow makes a great difference” (Livingstone 1995, personal communication). Sewall Wright, one of the founders of population genetics, pointed out in 1931 that evolution is the result of a shifting balance of forces. Although Livingstone was often characterized, even by himself, as a “pan-selectionist,” he clearly recognized the importance of those other forces.

Thus, whether the important determinant of genetic variation is genetic drift or the systemic pressures of selection, migration, and perhaps mutation depends to a considerable extent on the populational level being examined. Among the small isolates of the human species, genetic drift (including founder effect) may well be most important, but this does not imply that the major genetic differences within this species are also due to drift. [Livingstone 1973b:66]

No Races; Only Clines Perhaps even more influential within anthropology than his sickle cell work is Livingstone’s (1962a) article, “On the Nonexistence of Human Races.” Although the topic of race continues to be of central concern to anthropologists, most no longer defend a biological race concept for humans. That this is so is due in no small part to Frank’s widely reprinted paper. His rejection of the race concept in biology stemmed from his view that natural selection has determined the major patterns of human genetic variation. Since selection affects each locus independently, different genetic characters are not correlated in terms of their geographic distributions. There are no neat packages of concordant phenotypes. The cline for the sickle cell gene will be determined by malarial intensities, a pattern that need not match that of any other gene loci; each will be subject to different selective pressures, and so will also manifest different selective gradients. Thus, if there were such a thing as a “sickle cell race,” it would not be the same as one defined on the basis of ABO blood groups, skin color, etc.

Anti-Sociobiology It might seem that Livingstone, as a strong selectionist, was inconsistent in not being a champion of sociobiology, the application of natural selection theory to social behavior. However, his position followed logically from his background and previous research. As a graduate student at Michigan, Frank was influenced by the culturological approach of Leslie White, who vigorously argued for the primacy of cultural causes of human social behavior. Another influence was James Spuhler, the founding member of anthropological genetics at the University of Michigan. The title of Spuhler’s well-known edited volume, The Evolution of Man’s Capacity for Culture (1959), nicely summarizes his view of the roles of biology and culture. In Frank’s words:

There are obviously biological aspects of human behavior, and of course every human behavior has a biological basis in the sense that it is dependent on the biological abilities of the human species; but the causes or determinants of human behavior are very different from these generalities. [1980c:308]

Livingstone’s (1958a) own research stressed the critical role of cultural change in determining the selective environment for the sickle cell gene. It was the adoption of agriculture that brought about the series of ecological changes favoring mosquito vectors of malaria; in turn these caused the selective advantage for the Hb*S heterozygote. Human environments are partly shaped by culture. Cultural changes occur much more rapidly than genetic ones. Thus cultural behavior “has priority as a determinant of genetic change” (Livingstone 1980c:325).

Frank’s approach to cultural and genetic causes of behavior can also be seen in his treatment of the incest taboo (1969c). As he said, “There is an increasing consensus—not including me—that the deleterious genetic effects of close inbreeding are the cause of the supposed universality of this taboo in human societies” (1980c:316). He points out that the actual data regarding the effects of close inbreeding in human populations are quite equivocal; moreover, many animal societies do inbreed regularly. Using a simulation model, he demonstrated that inbreeding could actually lower the frequency of deleterious genes (1969c). His general point, again, is that mating behavior is a component of population structure, part of the culture of a population, determined by ecological and social factors.

Kinship Math Livingstone’s interest in cultural anthropology went beyond the role of culture as a selective environment for human genes. Social anthropology of the late 1950s and 1960s was concerned—some would say obsessed—with the analysis of marriage systems, particularly forms of cousin marriage, e.g., prescriptive patrilateral cross-cousin marriage. With an undergraduate degree in mathematics, Frank was well equipped to model these formal systems, and he published several articles from early in his career (1959a, 1959b, 1965, 1969d). These papers form part of a continuing literature on the formal analysis of kinship (Kronenfeld 2001). His early advocacy of graph theory as a tool for understanding cultural patterns also continues in modern work (Hage and Harary 1996).

Impact of Livingstone’s Research Frank Livingstone’s impact outside of anthropology came primarily from his central role in the sickle cell/malaria hypothesis. It should be remembered that sickle cell has been called “the leading paradigm for heterosis” (balanced polymorphism) in humans (Vogel and Motulsky 1997). His numerous studies of the population dynamics of the variant hemoglobins, and his compilations of data on population variation of the malaria related polymorphisms, have secured his enduring reputation in human genetics.

For anthropology, Frank’s research represents a sterling example of the value of the four-field approach. Though his seminal paper was titled the “Anthropological Implications of Sickle Cell Gene Distribution...” in another sense it was about the implications of anthropology as a whole for studying human genetic variation. This work demonstrated why a unified anthropology spanning the breadth of the subdisciplines was crucial to human biology. Natural selection may be the driving force, but population structure and history provide the selective environment and the demographic conditions for evolution. Human populations are structured by culture, e.g., by mating preferences and patterns of gene exchange between groups; and human population history is reflected in more than genes. So anthropological genetics is about much more than DNA. Indeed, as Ken Weiss (2003) points out, the incredible growth in the GSP (Gross Scientific Product) of contemporary human genetics, with the characterization of “the” human genome as well as large catalogs of human gene differences, may not have been matched by growth in understanding of human evolution. As Ken says, “The data we have are voluminous, but the trade of technology for intellectual breadth has had a price...those data need to be understood in terms of realistic models of hunter–gatherer or primate behavior and population structure” (Weiss 2003:628).

With the “molecular revolution,” neutral theory in genetics began to replace the balanced selection hypothesis as the cause of extensive variation. Thus Livingstone’s view that natural selection was paramount has been partly superseded by one emphasizing genetic drift. Perhaps “neutral” genes thus serve as “markers” of population history (Darwin’s “descent”), rather than as indicators of environmental selective histories (Darwin’s “modifications”). But the pendulum swings, and today’s dogma becomes tomorrow’s antiquated error. In the early days of biological anthropology, ostensibly neutral markers (either morphological or metrical) were used to reconstruct human racial history. Today neutral gene markers are used to construct gene trees supposedly reflecting population histories. But one of Livingstone’s enduring legacies is that history and culture also shape gene distributions. Therefore it is a mistake to try to read history from genes alone.

Mentor

During his long career at Michigan, Frank mentored many dozens of graduate students and taught several thousand undergraduates, personally counseling hundreds of them. His door was open to all, for debate or advice, without concern or preference for rank. Frank’s pedagogical approach didn’t differ much by context—office, bar, living room, lecture hall—he challenged you, he expected you to give back measure for measure. Frank wasn’t the least bit interested in having students regurgitate whatever it was he could have said during an hour-long monologue. The teaching, the learning, for us, for him, lay in the exchanges amongst Frank and whomever happened to be within earshot. And if one were of a more timid nature, “Well, Goddamnit, you’d better speak up for yourself, because nobody else is going to do it for you!”

The fact is, no matter where or with whom he happened to be, as Lisa Sattenspiel recalls, Frank just couldn’t not teach...

Early in the fall of my first year (1984), Frank and Carol held a departmental picnic at their house. At one point, Frank asked me about my research. I described how I had developed a mathematical and computer simulation model to study the spread of hepatitis A in Albuquerque, New Mexico, day care centers. He listened, and then gruffly confronted me with, “And what does that have to do with anthropology?” Initially taken aback by the vehemence of his question, I did my best to answer it—all the while feeling I was falling flat on my face in my first academic encounter with one of the senior authorities in my field. I will never know what kind of impression I really made, but it doesn’t matter. Frank’s brusque and deceptively innocent question made me instantly realize the importance of being able to explain to anyone why what you do fits within the framework you have chosen for it. To me, in that very brief encounter, Frank expressed the essence of being a great teacher: to stimulate others to always examine, justify, and support their ideas about the topic they are studying.

Ted Steegmann (2003) described graduate student training in physical anthropology during his years (1958–1964) in Ann Arbor: In addition to the coursework,

One of my greatest learning experiences came as a teaching fellow for James Spuhler’s Biology, Society, and Culture course, a piece of astonishing intellectual exposition on the biobehavioral sciences, then in their infancy. As an assistant to Ernst Goldschmidt, I came to understand how difficult it is to teach introduction to physical anthropology, and had a chance to study with a master of the art.

Frank later took over Jim’s class, teaching it regularly through the 1990s. Ernst would continue to teach introductory physical anthropology to ever-larger classes until 1987. A quick calculation finds that at least 10,000 students learned about genetics, human evolution, human adaptation, and the myth of human races from Ernst. Dozens of graduate student teaching assistants went on to adapt and teach these courses in other educational institutions, forming lineages that stretch back through Frank and Ernst to Jim Spuhler and forward to thousands of students.

At the Frankfest, Bob Eckhardt (Pennsylvania State) whimsically suggested that no matter what the course’s title, Livingstone’s subject matter was abnormal hemoglobins—eliciting a hearty laugh from Frank’s former students. Then drawing from class notes he still has decades later, Bob sketched the essential cohesiveness of Frank’s classes, which were, of course, always about variation and evolution. Holly Smith (University of Michigan) also still remembers the impact of Frank’s classes:

Although I entered graduate school to study paleontology, Frank’s graduate courses were my favorites. I took every one he offered. Frank was a knowledgeable and engaging lecturer. He shaped my understanding of evolution with population genetics. I have never forgotten his examples showing how even the slightest selection inevitably rolls through populations unless the effective breeding size is tiny. Specify a few parameters, and Frank was off and rolling.

Frank didn’t see “the academy” as something apart from “real life.” Science in general, and evolutionary anthropology in particular, could and should be used to gain new understandings of everyday problems experienced by real people, and to expose fallacious arguments. Reading at the Frankfest from a 1972 issue of Time magazine “dedicated to the wonders of the human genome,” Ken Weiss (Pennsylvania State) gave a good feel for Frank’s ability to “point out less-than-critical thinking in such articles, [and then] relate them to contemporary knowledge in evolution and genetics” (Weiss 2003).

Expecting that the struggle to understand the material would do more good than any of his explanations, Frank directed students to the original literature (author and title, and sometimes even library stack location!). He had little use for examinations and was disdainful of “prelims,” the obligatory entrance exams into the doctoral program. By the time I (VJV) took these, the physical anthropology prelims had grown to three 8-hour days of written essays crammed into one week. “What’s the point?!” Frank fumed. “You’ll never do anything like this again for the rest of your life!” He was right, again. Holly Smith recalls that even in standard courses, Frank “wasn’t much for essays. I’ll never forget one question: ‘The human species is presently undergoing the process of speciation—True or False.’ Every once in a while I still ponder the ins and outs of an essay one might write on that question, but Frank didn’t need to read it.”

Ted Steegmann (2003) reminds us that Frank’s lasting influence on students and anthropology began early in his career:

[Livingstone] offered an evolutionary analytical model so original that it has been hard to find parallels since then.... His work on hemoglobin polymorphisms was interesting as molecular biology, but made its eternal mark on the field by explaining how linguistic, archeological, and cultural patterns in West Africa shaped the pattern of a major biological trait. It is one of the great triumphs of anthropology as an integrated field. Second, he caused us to think about the distribution of human variation as a series of often-incongruent clines. Nothing did a better job of rescuing us from ancient and entrenched typological thinking—concepts of human variation that were alive and well in the 1950s. Finally, Frank opened our minds to the infinite usefulness of statistics as a logical system and key research tool.

Frank’s pedagogical legacy—Be critical! Be rigorous!—was evident in the Frankfest papers: The single species hypothesis may not be dead (Hunt 2003), human babies may not all pass through a crawling stage (Tracer 2002), the Black Death may not have been bubonic plague (Wood et al. 2003), balancing selection may not explain HLA-A polymorphism (Williams 2003), genetically based medical treatments may not be superior to environmental and behavioral interventions (Vitzthum 2003). Karen Rosenberg (University of Delaware) echoes this legacy:

[I] remember his brilliance, kindness and indeed his quirkiness. Willing to question everything no matter how much people believed it, he was the person that one could always rely on to tell everyone that the emperor has no clothes and that everything that we thought was true was in fact nonsense (of course, his vocabulary would have been a little more colorful). I think of him walking across campus, dog at his side, on his way to the computing center to pick up output, irreverent as ever.

I (VJV) share with others the sense that being Frank’s student was a privilege. But as one of his very few women students, my appreciation for Frank’s tutelage is even richer. That there were so few women graduate students in physical anthropology speaks to the culture, the times, and long-standing prejudices. But to Frank, it wasn’t the chromosomes you carried that mattered, but what you thought about chromosomes—and genes and variation and evolution. Frank disliked intellectual laziness. He expected from me, as he did from everyone, that I should disagree with him...and I had better do it well! Frank made that clear to me the night I asked him to be my advisor, in the crowded noisy kitchen of a small faculty house on the Old West Side. Waiting for a break in the boisterous debate Frank was having with some grad students, I grabbed my opportunity in a lull afforded by the handing out of more cold beer. Frank’s reply: “Awh, you’re a girrll—you’re probably afraid of numbers!” At which I laughed right in his face, followed by a few thoughts of my own about who’s afraid of what. He signed the papers naming me as his student the next week.

Although it’s fair to say that Frank could be intimidating, was often gruff, and always expected one to stand up for one’s self and one’s ideas, he wasn’t a bully. Frank never failed to support his students when the need arose. He didn’t run a factory, and he didn’t create carbon copies of himself. In Frank’s own words, “If you can’t take 95% of what I’ve done and show me why it’s wrong, and take the other 5% and make it better, then what the hell are you doing here?!”

Colleague

Frank didn’t rein in his contrarian ways just because he happened to be talking with colleagues of equal or greater rank, as illustrated by a popular story retold by many through the years: In a faculty meeting, Frank made a proposal met by silent rejection, and then only seconded by someone out of politeness. Given the opening, Frank went on at length, laying out many arguments in favor of his proposal. He was extremely persuasive, and when the count was finally taken, everyone had voted for it—except Frank. When asked why, he exclaimed, “I changed my mind!”

Turnover was low among the Michigan physical anthropology faculty, and through the decades, shared intellectual perspectives and lasting friendships intertwined. Frank Livingstone and Loring Brace, who joined U of M in 1967, were the foundation of what would come to be known as the “Michigan School” in physical anthropology, emphasizing the efficacy of natural selection, the pivotal role of culture in human evolution and adaptations, and the unity of the human species. This unity applied both to much of the fossil record (“the single species hypothesis”) and to the relations among contemporary populations (the race–defeating recognition of clinal variation). A story, told pretty much the same by both Frank and Loring, presaged their combined future scientific achievements. In 1961, at the 30th Annual Meeting of the American Association of Physical Anthropologists in Columbus, Ohio, Brace and Livingstone were on the program back-to-back, in alphabetical order. Loring presented the heart of a paper that he would subsequently publish in Current Anthropology, “The fate of the ‘classic’ Neanderthals...” As Frank approached the podium, he clearly looked very agitated and blurted, “He...He...He just gave my paper!” Defying the audience to leave or even yawn, Frank continued, “Well, you’re just going to have to listen to it anyway!” and began to read.

As Loring explains, Frank is the reason he joined the University of Michigan:

When Governor Ronald Reagan fired Clark Kerr, the President of the University of California, with the words, “The State of California has no business subsidizing intellectual curiosity,” it was perfectly clear to me that I had no future as a faculty member of...that University in Santa Barbara. I was in negotiations with a good half dozen places, but I chose to come to Ann Arbor specifically because Frank was here. My outlook had been heading in the same direction as his for some years, but when he wrote, “There are no races, there are only clines,” he said succinctly what I had only done with less clarity. In my job talk, I noted that “you have no fossil man” at Michigan. The double entendre was deliberate. Ultimately I was hired to cover the fossil record, but of course I was always interested in what that record led up to and why. Frank, on his part, worked from an understanding of contemporary human variation but was always interested in where it had come from and why. After I had been here for a couple of years, we decided to switch courses. He took my fossil course, and I taught his course, “Problems of Race,” which I have been teaching ever since.

One of the things that has interested me has been if there is no biological entity that corresponds to “race,” how come everybody believes in it as somehow self-evident? The second half of the course I inherited from Frank has dealt at some length with that question, and I have at long last gotten that whole matter into print in “Race” Is a Four-Letter Word: The Genesis of the Concept (2005). I have mentioned Frank prominently in the text. I salute him with admiration and gratitude for what he has done for our field over an awesome and productive career.

Honors

Upon his retirement in 1998, Frank B. Livingstone was named Professor Emeritus of Biological Anthropology at the University of Michigan. Accompanying four decades of service as a faculty member, including a three-year stint as Department Chair, Frank had advised the Human Genetics Section of the World Health Organization and was a Research Associate of the Liberian Institute of Tropical Medicine.

In recognition of his groundbreaking research on sickle cell anemia, the Southern Christian Leadership Conference bestowed upon Frank its highest honor, the Martin Luther King Award. In 1999, the American Association of Physical Anthropologists presented Frank with its most prestigious tribute, the Charles R. Darwin Award for Lifetime Achievement.

As mentioned earlier, Frank’s exemplary accomplishments in both research and teaching were heralded at a Festschrift at the April 2002 meeting of the American Association of Physical Anthropologists in Buffalo, New York. Its proceedings were summarized in Evolutionary Anthropology (Vitzthum and Hunt 2003) and published in a special issue of Human Biology (Vitzthum and O’Rourke 2003). Ample evidence of Frank’s lasting intellectual contributions to the field can be found in journals, citation indices, and the warm appreciation of an audience overflowing the Frankfest hall.

Father

The View from the Family Room: Reflections of an Anthropologist’s Daughter

When asked what their fathers do for a living, most children are able to answer easily: “My Dad’s a fireman, doctor, teacher, salesman...” But being an anthropologist’s offspring makes the response to this simple question a bit complicated. Answering with, “My Dad’s an anthropologist,” can elicit responses such as, “An anthro-what?” or “That’s nice, dear; we’re Episcopalian.”

As a youngster, I was often unsure of what it was my father actually did for a living. I knew he taught at the university, and I understood that the piles of printouts and IBM cards scattered all over our family room had something to do with that. But the exact connection between the two was mysterious. Over time I came to realize that his work had to do with diseases, for when Frank was awarded the Martin Luther King Award by the Southern Christian Leadership Conference for his research on malaria, his colleagues gave him a shirt that said “Mr. Sickle Cell.” I didn’t know what a sickle cell was, but I knew that sickle cells made people sick. Over the years, figuring out what my father did, as well as discovering for myself his profound impact on the anthropology community, turned out to be quite an adventure.

Anthropology was more than a career to my father; it was a part of our family life. I remember “potsherd alerts.” We would be driving down the road, only to encounter a freshly plowed field. We’d pull off the road and my parents would leap from the car to search the field for unknown treasures of the remote past. While I, too, would tromp through the dirt, my enthusiasm was dampened when the lovely piece of Blue Willow porcelain that I had found was dismissed as unimportant. I was miffed since my “pottery shirt” was much prettier than anything my parents had uncovered.

The first time Frank brought home an assortment of fossil casts—australopithecines, Homo erectus, Neanderthals—I was fascinated by the skulls lined up on our coffee table. “These are your ancestors,” my father proclaimed. Later that afternoon, I could be found guiding my playmates through the collection, repeating that these were “my ancestors.” Being a literalist, I gave them the names of those I knew to be dead: Aunt Alice and Grandma Fenner.

When I was fairly young, my father began holding evening seminars at our home. Nearly every Monday night of my childhood, dinner had to be on the table by 6:30, the dishes done and put away by 7:00, so my father could begin preparations—which generally meant popping lots of corn and being sure the drinks were plentiful and cold. By 7:30 the carloads of students would begin to arrive. I can remember lying in my bed and hearing the conversations. Sometimes I would hear my father’s voice but most often it would be those of the students, developing their own ideas about genetics and evolution.

In addition to these seminars, our front door virtually revolved with graduate students and colleagues stopping by for a beer and, if they were lucky, an invitation to stay for one of my mother’s fabulous dinners. As a kid, I wasn’t quite sure what these conversations were about. Over time, however, I came to realize that students came to talk with Frank about their own ideas and research. Many a doctoral dissertation was devised over a beer from the ever-present cooler in our family room. Plans were made for research trips; encouragement was given for students to develop their own ideas and to challenge the accepted paradigms.

Life advice was also given, peppered with frequent (and sometimes salty) interjections from my mother. The 1960s were a turbulent era, perhaps felt most acutely by the youth of that period. Lively discussions at the Livingstones covered many issues—politics, the draft, the war—but the message from Frank and Carol was invariably the same: be true to what you believe and who you are, no matter how difficult that might be.

Newly arrived assistant professors also made the trip to Rock Creek Drive, often to discuss “ten-year”—“Why was everyone so worried about getting ten years? Weren’t they old enough already?” my young mind wondered—and how to go about getting it. Colleagues’ visits also revolved around departmental issues and the intricacies of anthropology. These tended to be lively conversations, punctuated with heated debates and the occasional expletive. As an adult academic, I now realize exchanges that often appeared casual were really Frank’s mentoring style—the shaping and support of young anthropologists while never forgetting that they were also people.

The rigors of seminars, mentoring, administrating, and research were balanced by enthusiastic socializing amongst departmental members. My parents hosted many a party, and it was always an adventure having anthropologists in the house. I recall Napoleon Chagnon trying to teach Yanomamo to the guests, and once I had to rescue my pet guinea pig from Roberto Frisancho when he threatened to cook him in the toaster oven—suggesting that since people in Peru ate guinea pigs that my pet should also be on the party table. He was only kidding, of course, but it certainly made an impression on a ten-year-old.

My favorite anthropology get-together was the annual picnic. Esteemed full professors, younger faculty, and graduate students—all accompanied by families and pets—gathered early in the fall to get to know each other and for general conviviality. There was usually a roasted pig or lamb, and plenty of refreshments. Barriers of rank faded as assistant professors tossed Frisbees or footballs with senior colleagues (Frank would be among the first to volunteer for any sporting event), and students helped professors tap the beer keg. These picnics were always great fun, and like the parties at my parents’ home, they fostered community, collegiality, and affection among the anthropologists.

It wasn’t until I entered college that I became consciously aware of the influence of the department at the University of Michigan on the discipline of anthropology. Nor had I previously appreciated the full significance of my own father’s contributions to evolution and population genetics. At Michigan State I took several classes in anthropology. In my texts I would frequently encounter references to the contributions of people who had passed time in our family room: Leslie White, James Griffin, Eric Wolf, Vern Carroll, Sherry Ortner, Richard Ford, Jeff Parsons, Loring Brace, and others. It was particularly startling to realize that the man whom I knew as “Uncle Leslie,” who literally bounced me on his knee when I was a baby and always sent me a Valentine Day’s card, was one of the founding fathers of modern anthropology. Sometimes an unaware friend would jokingly inquire if the Frank Livingstone cited in these texts was any relation. Imagine the surprise of learning he was my father: the same character who showed up on weekends, smoking his cigar and asking whether we could have beer in our room. Most of my friends came from non-academic backgrounds. It was impossible for them to imagine their parents’ work being the subject of their coursework. For me, it was startling but also a tremendous source of pride because Frank’s work shaped the way my peers looked at the world and understood its workings.

Not surprisingly, my father played an important role in my growth as a scholar. No less than with his own students, Frank frequently played devil’s advocate, challenging my research assumptions. While sometimes aggravating, Frank’s persistence inevitably helped me to clarify my own thinking and approaches. In particular, he “encouraged” me to use statistics to support my points more effectively. Several years ago, Frank had the immense pleasure of watching me eat crow upon my admission that I planned to employ statistical analyses in my study of aristocratic family life in the Middle Ages. My own experiences made me aware of what a wonderful mentor he was to his own students. Frank never sugar- coated his responses or criticism. Upon reflection, and from discussions with his students, I realized Frank wasn’t being unkind. He viewed his students, and me, as his intellectual equals and expected us to be able to hold up our own end of the bargain.

During my own studies, my father was also struggling with a new challenge: being department chair. Nothing in our academic training prepares us for the role of administrating a department. No longer operating in isolation, the concerns and needs of numerous people must be carefully balanced in things as mundane as scheduling classes and as tricky as promotion and compensation. Frank suddenly had to mediate disagreements that he previously could have ignored. Now the well-being of all graduate students—and their treatment by faculty—came within his purview. These are dangerous waters, and Frank encountered his share of turbulence. He didn’t make many friends among the higher administration for his sometimes unpopular or impolitic positions. Sad to say, relationships with some colleagues were also strained. Perhaps to mend bridges, gatherings continued to be held at Chez Livingstone, a testament to the bonds among departmental colleagues. As I am now chair of my own department, I understand more fully the difficulties Frank experienced. I find myself thinking back on his experiences and try to use them to help me chair my own department and interface with the administration.

After his official retirement from the university, Frank continued to walk between home and campus regularly to meet colleagues and advise undergraduates. His own contributions to anthropology were acknowledged with the bestowing of several honors. In one venue or another, someone would realize I am the daughter of THE Frank Livingstone and, more often than not, an invitation would be extended for him to speak. Sadly, his declining health prevented these engagements in his last years. While the impact of my father’s reputation has been something to which I’ve grown accustomed, I have come to realize just how unusual and special it was to come of age in the anthropology department at Michigan in the 1960s and 1970s. It was certainly an adventure knowing and interacting with its faculty as well as the many graduate students who have gone on to acclaim in their own right. I am forever grateful that I had this opportunity. I also feel a tremendous sense of gratitude, for it was at the cocktail parties and anthropology picnics that I was first exposed to the life of the mind and the excitement of ideas. The mark they left was deep for I have chosen academe as my life’s work. To all the anthropologists, past and present, but most particularly my father, I would like to extend my profound thanks. It has been one amazing journey.

Partner

Any balanced retrospective of Frank’s life should rightly give equal time to Carol Livingstone. Married for nearly 45 years, Carol and Frank were partners and collaborators in “his” academic life as well as in their family life. Our inability to do justice here to Carol’s contributions reflects the enormity of such a task—and the fact that Carol is still fully able to come after us and exact retribution should we be so foolish as to write anything even remotely soft-headed, sentimental, or vainglorious! If it can be said that Frank lacked pretension, it can more so be said that Carol is its antithesis. And so we will limit ourselves to a few factual observations.

Not an evening’s seminar, not a reception for some honored speaker, not a party, not a spring barbeque, not a consultation between Frank and anyone could have happened at Rock Creek Drive without Carol. Frank took care of the drinks and the dishwashing (every night!); Carol took care of the food (her lasagna is legendary!). Every guest, drop-in, dog, cat, raccoon, and student left their home warmed by their hospitality; most returned again and again (including the family of raccoons!). Carol’s accomplishments extend far beyond these ministrations. She is a first-rate carpenter (and has some pretty hairy stories about male counterparts’ lack of acceptance), wood carver, and furniture finisher, and a dedicated politico in the Michigan Democratic Party. She was the campaign manager for a re-election of G. Mennon “Soapy” Williams, Michigan’s popular six-term Governor, and a delegate to the 1960 Democratic convention. Carol was Frank’s equal in every way and then some. And he knew it. Frank was lucky to have her. And so are we.

Carol and Frank’s partnership was brought to a close with his death at the age of 76 on March 21, 2005, in Springfield, Ohio, from the complications of Parkinson’s disease. Frank’s bones lie not far from those of Leslie White, near the arboretum through which he and Irving-the-dog would run daily, no matter the weather. Carol now lives in Springfield, Ohio, near their daughter Amy, son-in-law Gordon Thompson (Frank’s perennial golfing, fishing, and hockey “bud”), and grandsons Samuel and William. Ernst can be found today tending the house on Rock Creek Drive that welcomed so many, so warmly, so often.

Farewells

“The golden years are here at last;
The golden years can kiss my ass!”

With these words, Carol opened a celebration of Frank’s life, attended by some 200 of Frank’s family, friends, colleagues, and former students at the Michigan League on April 16, 2005. The Vandenberg Room—dark polished wood, tall leaded windows brimming with spring sunlight—was both elegant and welcoming. A Dixieland band matched its music to the spirit of the man to be honored as guests circled round posterboards spilling with photos and the exhibited tools of Frank’s second trade—a hockey stick and helmet—and its rewards, an imposing trophy (one of many).

In turn, each family member and friend delighted us with their favorite memories of Frank:

“His great love as a boy was fishing—we called him ‘Frog’ till he had a Ph.D.”

“The TV remote remained an enigma.”

“His all-purpose fix-it approach: hit it...then hit it harder!”

“Even as a grad student, he would complete the New York Times crossword in less than an hour.”

“Anxious for the year abroad I was about to begin, I wished it were over; my father’s reply, ‘Don’t wish your life away—live it!’”

Frank certainly had taken his own advice! We laughed with each speaker and, remembering a story or two of our own, smiled to ourselves with the recollection. The band struck up again, glasses filled with beer and wine, the buffet table sagged with shrimp and other Frank favorites. More stories exchanged, more catching up amongst folks long out of touch, more beer. It could easily have been another grand Livingstone party on Rock Creek Drive!

“I had one kid, one wife, one house, one job—I got all the important things right the first time!”

–Frank B. Livingstone

References cited

Allison, A. C.1954. Protection Afforded by Sickle-Cell Trait against Subtertian Malarial Infection. British Medical Journal 1:290–294.

Boren, T., P. Falk, K. A. Roth, G. Larson, and S. Normark. 1993. Attachment of Helicobacter pylori to Human Gastric Epithelium Mediated by Blood Group Antigens. Science 262:1892–1895.

Brace, C. Loring. 2005. “Race” Is a Four-Letter Word: The Genesis of the Concept. Oxford: Oxford University Press.

Cavalli-Sforza, L. L., P. Menozzi, and A. Piazza. 1994. The History and Geography of Human Genes. Princeton: Princeton University Press.

Fisher, R. A. 1937. The Wave of Advance of an Advantageous Allele. Annals of Eugenics 7:355–369.

Fix, A. G. 2003. Simulating Hemoglobin History. Human Biology 75:607–618.

Hage, P., and F. Harary. 1996. Island Networks: Communication, Kinship, and Classification Structures in Oceania. Cambridge: Cambridge University Press.

Haldane, J. B. S. 1949 . Disease and Evolution. Ricerca Scientifica (Suppl.) 19:3–10.

Hiernaux, J., J. Linhard, F. B. Livingstone, J. V. Neel, A. Robinson, and W. W. Zuelzer. 1956. Data on the Occurrence of Hemoglobin C and Other Abnormal Hemoglobins in Some African Populations. American Journal of Human Genetics 8:138–150.

Hunt, K. D. 2003. The Single Species Hypothesis: Truly Dead and Pushing Up Bushes, or Still Twitching and Ripe for Resuscitation. Human Biology 75:485–502.

Kronenfeld, D. B. 2001. Introduction: The Uses of Formal Analysis Re Cognitive and Social Issues. Anthropological Theory 1:147–172.

Livingstone, F. B., M. J. Miller, J. V. Neel, A. R. Robinson, and W. W. Zuelzer. 1956. Two “Fast” Hemoglobin Components in Liberian Blood Samples. Blood 11:902–906.

Livingstone, F. B. 1957. Sickling and Malaria. British Medical Journal 1:762–763.

———. 1958a. Anthropological Implications of Sickle Cell Gene Distribution in West Africa. American Anthropologist 60:533–562.

———. 1958b. The Distribution of the Sickle Cell Gene in Liberia. American Journal of Human Genetics 10:33–41.

———. 1959a. A Further Analysis of Purum Social Structure. American Anthropologist 61:1084–1087.

———. 1959b. A Formal Analysis of Prescriptive Marriage Systems among the Australian Aborigines. Southwestern Journal of Anthropology 15:361–372.

———. 1960a. The Wave of Advance of an Advantageous Gene: the Sickle Cell Gene in Liberia. Human Biology 32:197–202.

———. 1960b. Natural Selection, Disease, and Ongoing Human Evolution. Human Biology 32:17–27.

———. 1960c. The Distribution of Several Blood Group Genes in Liberia, the Ivory Coast, and Upper Volta. American Journal of Physical Anthropology 18:161–178. With H. Gershowitz, J. V. Neel, W. W. Zuelzer, and M. D. Solomon.

———. 1961. Balancing the Hemoglobin Polymorphisms. Human Biology 33:205–219.

———. 1962a. On the Non-Existence of Human Races. Current Anthropology 3:279–281.

———. 1962b. Reconstructing Man’s Pliocene Pongid Ancestor. American Anthropologist 64:301–305

———. 1964a. The Distribution of the Abnormal Hemoglobin Genes and their Significance for Human Evolution. Evolution 16:685–699.

———. 1964b. Aspects of the Population Dynamics of the Abnormal Hemoglobin and Glucose-6-Phosphate Dehydrogenase Deficiency Genes. American Journal of Human Genetics 16:435–450.

———. 1965. Mathematical Models of Marriage Systems. Man 65:146.

———. 1967. Abnormal Hemoglobins in Human Populations. Chicago: Aldine.

———. 1969a. The Founder Effect and Deleterious Genes. American Journal of Physical Anthropology 30:55–60.

———. 1969b. Gene Frequency Clines of the ß-hemoglobin Locus in Various Human Populations and Their Simulation by Models Involving Differential Selection. Human Biology 41:223–236.

———. 1969c. Genetics, Ecology, and the Origins of Incest and Exogamy. Current Anthropology 10:45–61.

———. 1969d. The Application of Structural Models to Marriage Systems in Anthropology. In Game Theory in the Behavioural Sciences. I. H. Buchler and H. G. Nutini, eds. Pp. 235–251. Pittsburgh: University of Pittsburgh Press.

———. 1971. Malaria and Human Polymorphisms. Annual Review of Genetics 5:33–64.

———. 1973a. Data on the Abnormal Hemoglobins and Glucose-6-Phosphate Dehydrogenase Deficiency in Human Populations 1967–1973. Technical Reports No. 3. Ann Arbor: Museum of Anthropology, University of Michigan.

———. 1973b. Gene Frequency Differences in Human Populations: Some Problems of Analysis and Interpretation. In Methods and Theories of Anthropological Genetics. M. H. Crawford and P. Workman, eds. Albuquerque: University of New Mexico Press.

———. 1976. Hemoglobin History in West Africa. Human Biology 48:487–500.

———. 1978. Frequency-Dependent Selection and the ABO blood groups. In Evolutionary Models and Studies in Human Diversity. R. J. Meir, C. M. Otten, and F. Abel-Hameed, eds. Pp. 127–139. New York: Mouton.

———. 1980a. Natural Selection and Random Variation in Human Evolution. In Current Developments in Anthropological Genetics, Volume 1: Theory and Methods. J. H. Mielke and M. H. Crawford, eds. New York: Plenum.

———. 1980b. Natural Selection and the Origins and Maintenance of Standard Genetic Marker Systems. Yearbook of Physical Anthropology 23:25–42.

———. 1980c. Cultural Causes of Genetic Change. In Sociobiology: Beyond Nature/Nurture? G. W. Barlow and J. Silverberg, eds. Pp. 307–330. Boulder: Westview Press.

———. 1983. The Malaria Hypothesis. In Distribution and Evolution of Hemoglobin and Globin Loci. J. E. Bowman, ed. New York: Elsevier.

———. 1984. The Duffy Blood Groups, Vivax Malaria, and Malaria Selection in Human Populations: A Review. Human Biology 56:413–425.

———. 1985. Frequencies of Hemoglobin Variants: Thalassemia, the Glucose-6-Phosphate Dehydrogenase Deficiency, G6PD Variants, and Ovalocytosis in Human Populations. New York: Oxford University Press.

———. 1989a. Simulation of the Diffusion of the ß -globin variants in the Old World. Human Biology 61:297–309.

———. 1989b. Who Gave Whom Hemoglobin S: Use of Restriction Site Haplotype Variation. American Journal of Human Biology 1:289–302.

———. 1989c. Update to “The Wave of Advance of an Advantageous Gene: The Sickle Cell Gene in Liberia.” Human Biology 61:831–834.

———. 1991. Phylogenies and the Forces of Evolution. American Journal of Human Biology 1:83–89.

———. 1993. The Roles of Mutation, Selection, Gene Flow, and Gene Conversion in Determining the Geographical Distributions of the Beta-S Globin Haplotypes (abstract). American Journal of Physical Anthropology, Supplement 16:135.

Miller, L. H., S. J. Mason, J. A. Dvorak, M. H. McGinniss, and I. K. Rothman. 1975. Erythrocyte Receptors for (Plasmodium knowlesi) Malaria: Duffy Blood Group Determinants. Science 189:561–563.

Olesen, E. B., K. Olesen, F. B. Livingstone, F. Cohen, W. W. Zuelzer, A. R. Robinson, and J. V. Neel. 1959. Thalassaemia in Liberia. British Medical Journal 1:1485–1487.

Ruiz-Pesini, E., D. Mishmar, M. Brandon, V. Procaccio, and D. C. Wallace. 2004. Effects of Purifying and Adaptive Selection on Regional Variation in Human mtDNA. Science 303:223–226.

Spuhler, J. 1959. The Evolution of Man’s Capacity for Culture. Detroit: Wayne State University Press.

Steegmann, Jr., T. 2003. Anthropology at the University of Michigan in the 1950s. Human Biology 75:619–624.

Sutton, H. E., J. V. Neel, F. B. Livingstone, G. Binson, P. Kunstadter, and L. E. Trombley. 1959. The Frequencies of Haptoglobin Types in Five Populations. Annals of Human Genetics 23:175–183.

Tracer, D. 2002. Did the Australopithecines Crawl? (abstract). American Journal of Physical Anthropology, Supplement 34:156–157.

Vitzthum, V. J. 2003. A Number No Greater Than the Sum of Its Parts: The Use and Abuse of Heritability. Human Biology 75:539–558.

Vitzthum, V. J., and K. D. Hunt. 2003. AAPA Honors Frank B. Livingstone. Evolutionary Anthropology 12:161–163.

Vitzthum, V. J., and D. O’Rourke, eds. 2003. Special Issue in Honor of Frank B. Livingstone: Papers Presented at the 2002 Annual Meeting of the American Association of Physical Anthropologists, Buffalo, New York, April 10–13, 2002. Theme issue, Human Biology 75(4).

Vogel, F., and A. G. Motulsky. 1997. Human Genetics: Problems and Approaches. Berlin: Springer.

Weiss, K. M. 2003. THINK! Being a Student of Frank B. Livingstone. Human Biology 75:625–628.

Williams, R. C. 2003. The Mind of Primitive Anthropologists: Hemoglobin and HLA, Patterns of Molecular Evolution. Human Biology 75:577–584.

Wood, J. W., R. J. Ferrell, and S. N. Dewitte-Aviña. 2003. The Temporal Dynamics of the 14th Century Black Death. Human Biology 75:427–448.

Wright, S. 1931. Evolution in Mendelian Populations. Genetics 16:97–159.