We are now in evolution's third century. The nineteenth century brought the view that the world is the result of processes of change. But a common concept does not always lead to common conclusions. Nowhere is this more ironically seen than in Highgate Cemetery in London. Two of the notables interred at Highgate are the social thinkers Karl Marx and Herbert Spencer. Marx and Spencer might both be proud to be at Highgate, but they lie in locations they could hardly find restful. Indeed, these locations seem like somebody's idea of a morbid joke: Very distant in life, they are eternally close in death, facing each other across a narrow path (Fig. 1). Strange bedfellows. Karl Marx (1818–1882, left top) and Herbert Spencer (1820–1903, left bottom) buried across from each other (right) in London's Highgate Cemetery. Photos by the author; map from Highgate Cemetery. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] I discovered this irony while walking through the cemetery on a gray English day. As I was contemplating these great thinkers, a long-haired young American in an old army jacket came strolling along. He stopped and asked another tourist to take a picture of him standing at the side of Marx's memorial and, as he did that, to everyone's surprise, he burst into a full-throated rendition of the “The Internationale.” As he sang out “This is the final struggle!” Spencer must have been struggling in his grave, because in real life Marx and Spencer hardly saw eye to eye. Ever since Isaac Newton, there has been a belief in Western science that nature follows universal laws. To Darwin, Spencer, and Marx the law was evolution. Spencer argued that a single law of evolution extended from Darwin's ideas about organisms to the nature of society. He attempted to connect the physical laws of energy and thermodynamics to organisms and, via the role of energy in mental states, to society.1 In 1851, before Darwin's Origin of Species, he was echoing Malthus about the futility of protecting the weak: The “decrees” of nature's “far-seeing benevolence” are “pitiless in the working out of good” via the “harsh fatalities” among the inferior members of society, all in the “interests of universal humanity” if we but have “the nerve to look this matter fairly in the face.”2 Darwin developed his law of natural selection to explain organic evolution, but in Descent of Man3 he praised Spencer and suggested how behaviors such as cooperation would have been favored by natural selection as human cultures morphed from primitive bands into modern civilization.3 Marx was interested in this same social progression. He and Darwin had a brief correspondence. Marx called the Origin of Species “a book which contains the basis of natural history for our views” and, in 1873, Darwin wrote of Marx's Das Kapital that though their ideas differed they “both earnestly desire the extension of Knowledge, and that this is in the long run sure to add to the happiness of Mankind.”4 But ultimately Marx's evolutionary law was about resolution against rather than acquiescence to inequity. Spencer had predictably negative views of socialism1 and referred to it only briefly and unflatteringly in his Principles of Sociology.5:596 Nonetheless, although Marx's and Spencer's political ideologies were poles apart, their theories, along with Darwin's, shared the central idea that progress resulted from conflict and contrast. The internal complexities of society included social classes competing for resources much as biological evolution was based on competition among biological unequals within species. The idea that society was evolving toward some final state was widespread at the time.6 Both Marx and Spencer held that traits acquired by society were inherited in a Lamarckian way, learned by society in the march of progress. For Marx, the internal contentions in class society would soon resolve into a classless communist society. For Spencer, too, “Civilization is the last stage” of nature's harsh process toward a societal equilibrium of “ultimate perfection” that would embody and be enabled by inequalities based on merit and a kind of anarchic utopia of individual freedom.2 But to Spencer, this would happen by purging rather than resolving. Despite these common elements, we usually see an unbridgeable crevasse between Darwin's views and those of Marx and Spencer and the other social dreamers. Darwin's lasting idea is that natural selection screens variation that arises randomly with respect to function. Though Darwin toyed with Lamarckian inheritance in his theory of pangenesis,7 Darwinian evolution was ever-divergent, because environments would always change and new variation would always arise, occasioning new adaptation. That does not converge on any endpoint, as the presence today of monads and men amply shows.8 Yet, even Darwin and his supporters, including Alfred Wallace and Thomas Huxley, came to believe in a kind of final and even imminent utopia. Thanks to modern society, humans had evolved the cultural ability to stop blind natural selection and control life to our plans and desires, a cultural Lamarckism. The notion that we can control our own destiny by improving our biological stock in ways that we, not nature, choose was embodied in the eugenics movement and exists today in proponents of personalized genetic engineering. The literature of the time shows how pervasive evolutionary ideas were in Victorian intellectual society.9 In Anthony Trollope's political novel The Prime Minister,10 the PM describes his philosophy: You have to strive “to improve the condition of the people” and reduce the distance between rich and the “lower man,” but any final success is “so distant that we need not even think of it as possible.” Equality is a dream because “Men's intellects are at present so various that we cannot even realise the idea of equality.”10:127 Among contemporary authors, Trollope's friend George Eliot was probably the most savvy about science. Evolutionary concepts are found in several of her novels.9 She even had a platonic love affair with Spencer himself. It did not work out. She wrote that he was an internal iceberg who rebuffed her expressions of love,11, 12 while he claimed that she was masculine and too ugly.11-13 Soon after being dumped by Spencer, Eliot took up with another leading intellectual, George Henry Lewes. They were among Darwin's earliest supporters in the Victorian intelligentsia11 and were friends with Trollope, though his discussions with them seem to have been “the closest he ever came to scientific thought.”6, 9 Eliot and Lewes were both buried in Highgate cemetery, right around the path from Marx and Spencer's eternal standoff (Fig. 2). Since Spencer attended both their funerals,11 he had probably trod, hat in hand, on his own future resting place. Even stranger bedfellows. George Eliot (left) and her lover George Lewes (center) in their locations in Highgate (right). Photos by the author; map from Highgate Cemetery. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] During evolution's second century, the twentieth, anthropology developed a long legacy of thinking of both cultural and biological change in evolutionary terms. If nature is law-like and anthropology is a single science, and if humans are biological organisms whose existence has always depended on culture, shouldn't Spencer be right that the same process must explain our joint biological and cultural evolution? Spencer's specific ideas did not stand the test of time, but were followed by continued discussion of evolutionary concepts in anthropology. At least two obvious ways were considered in which the same universal evolution could sensibly connect human biology and culture.14, 15 First would be a functional explanation based on our shared biology. As celebrity chef Wolfgang Puck would put it, what we do is “Eat, love, live!” Society must be structured to meet these needs, and there has been a large body of theory to dissect cultures in that context. Spencer himself discussed the evolution of the “social organism,” a forerunner of ethnology's structuralism and functionalism. More interesting and, of course, long debated among ethnologists, psychologists, and other social scientists is whether, other than being trivially Puckish about human biology, cultural evolution even follows its own laws. Can we explain or predict the forms of society without deeper reference to human biology? Marx thought so, and heavily borrowed anthropological ideas from the American anthropologist Lewis Morgan. The possibility that there might be a single cultural law was supported by the independent presence of many shared aspects of society, such as different kinds of kinship-based social organization, division of labor, status hierarchies, and so on. That seemed law-like, and since similar structures were found globally, human beings appeared to be a constant relative to cultural variation, implying that cultural facts must be understood strictly in terms of cultural facts. That point was made by several important nineteenth- and early twentieth-century evolutionary ethnologists.14 But if they were right, it would disconnect the two kinds of evolution rather than uniting them. Another way there could be a single evolutionary anthropology law is if biological and cultural variation control each other. There are classic examples of the way in which natural selection connects culture and biology. Genetic changes that lead some adults to be able to digest milk co-evolved with the cultural trait of dairying. Settled agriculture enabled our exposure to malaria in Africa, leading to a Darwinian adaptation of genetic resistance such as the sickle cell mutation. We have genetically lost most pheromone- and estrus-based reproductive characteristics, and have evolved cultural bases for choosing mates. And as we teach in introductory anthropology classes, there are thumbs and tools, language, and group behaviors. However, these specific co-adaptations hardly seem like the basis for any universally shared evolutionary law. By the latter decades of the twentieth century, Marx's and Spencer's and others' attempts at a general scientific theory of cultural evolution had clearly failed to reach the kind of consensus that Darwin's has achieved in biology. The idea just petered out, or perhaps its claims even engendered a generation of anti-science reaction in anthropology. Nonetheless, there were renewed attempts to unite cultural and biological concepts by considering culture in terms of gene-analogs called memes, to which one could apply a kind of Darwinian evolutionary theory.14, 16 Memes are rather difficult to identify precisely. They do not evolve in the same way that genes do; for example, they are transmitted among contemporaries, not just inherited from parents. Whatever their merits, since memes do not map directly onto genes, they do not automatically provide the basis for a single theory of evolution. So many social scientists are now turning to genes themselves. As a result of burgeoning genetic research since I first commented on this general subject in these pages,14 we now know of many specific genes that, when mutated, affect complex social behaviors ranging from language to various measures of intelligence, as well as sex, gender, and beyond. Similar findings will be pouring forth in the coming years. They are hungrily picked up by the popular news media, where they are essentially touted as explaining complex social behavior; that is, how people really are, as distinct from our self-illusions. Among the more prominent recent examples are statistical associations between specific variants in neurotransmitter genes and behavior. These include, for example, associations between the dopamine receptor DRD2 and compulsive or risk-taking behavior, including gambling; DRD4 and novelty-seeking; and the SLC6A4 serotonin transporter and anxiety-related responses. There are recent studies of DRD2 variation that claim to predict people's level of partisanship and political participation.17-19 And did you know that your marital fidelity is also written in your genome?20 The supporting evidence is variable. Moreover, many of the claims are controversial, in part due to a tendency to claim stronger or more predictive effects than actually seems to be the case. For example, even follow-up studies that confirm the general behavior of association may find much weaker effects or the effects may be replicable in some but not all studies or may involve different aspects of behavior. The SLC6A4 regulatory variant is classically claimed to confer poor stress response, for example, but recent results show that the same variant can be protective against stress if you have high socioeconomic status (D. Notterman, personal communication). Most of these studies have been done within industrial nations, so that the effect of the same genotype in very different cultures can not be specified a priori even if the genotype exists there. But despite these issues, proponents believe that this kind of evolutionary and behavioral social, psychological, political, and economic genetic study constitutes a new science of “human nature.”19, 21 Such a union of genetics and behavior would have pleased Spencer, though not Marx, for whom social inequities were learned rather than inherent. There is, of course, great resistance to these studies. Some see them as assaults on the sense of freedom of personal action. There also is concern that such work could lead to societal abuse of individuals by pigeon-holing them based on weak statistical results from studies of particular genes in particular contexts. However, these are personal or political objections, not genetic ones, and are not pertinent to whether such results can ultimately lead, as many believe they will, to a unified science of evolutionary anthropology. Basically every human trait involves the activity of many genes. These will all vary to some extent in any population. That is why, with regard to most traits, relatives resemble each other more than they do the average person in the population. For funding reasons, genetics research to date has concentrated on disease, and has almost always found at least some similarity of risk among close relatives. Several genes have usually been identified in which variation convincingly contributes to that risk. Nobody thinks disease is not a legitimate target of genetics research. But this large body of genetics research clearly implies that other traits, including behavior, even culturally sensitive behavior, must also be affected by genetic variation. That view is being confirmed by the kinds of studies referred to above. However, this raises an important question that is directly relevant to Marx and Spencer: Is the current turn to genetics going to provide a universal theory of human biocultural life or is it mainly a hijack of social science by geneticists? The answer is not obvious, as an illustration from evolution's first century may help to show. Besides writing very entertaining novels, Anthony Trollope was fond of playing whist at his London club (Fig. 3).22 For gentlemen “well stricken in years,” a daily whist session “makes easy the passage to the grave.” Some are more suited than others to the excitement of bidding and betting (DRD2, variants?), working out feints during play (DRD4?), or holding their cards close to their chests. Some will be winners and some, sadly, will not. Some of them will take their losses harder than others will (victims of SLC6A4?). Of whist and waistcoats: Trollope and his friends play whist at their London club. From ref23. In what ways can this variation explain whist, London clubs, strategy-books, glasses of sherry, or waistcoats? None of these cultural traits are built into our genomes, yet each can be seen to have an evolutionary history. Whist, for example, apparently goes back a few centuries in France. The players' skills and gambling verve may be variable for genetic or other reasons, but as far as Darwinian selection goes, they are all past reproductive age, so their winnings do not translate into genetic evolution. Whist, played by elderly gentlemen with varying skills and temperaments, evolved into bridge which, shockingly, is played even by women in the same population. So where are the genes for that? There is a parallel in biology. Darwinian theory provides mechanisms for the way that variation arises and becomes more common, but does not account for what specific variations will arise or what they will do. Brains and thumbs (to evaluate and hold whist hands, respectively) are presumably the result of biological evolution, but can't be specifically predicted by population genetics theory, and evolved before there was any whist to play. They have to be explained in their own terms, which are related to the historical value of environmental risk-assessment and, perhaps, tool use. Those explanations are not specifically genetic, even if the traits clearly involve genetic change, and it is interesting to understand how genes bring them about. But they don't explain whist or waistcoats. Unless one thinks we can actually construct the evolution of cultural traits step-by-step in terms of specific genetic variants over past millennia, we will not be much helped by the current interest in behavioral genetics. Genetic variation changes far too slowly to account for cultural traits and their variation. So even if we may be able to use genetics to predict who will be the best whist player, genes will not explain whist, which is too many levels of organization removed from the individual contributors' effects. Even if it is true that a mutation increases one's chances of managing stress badly, gambling, or voting Republican, this does not explain our financial system, good or bad investment, the nature of gambling in our society, or the two-party system. It means that a fraction of the variation in these behaviors, in today's culture, is affected by genetic variation. The only thing that is new is that some of the genes can be identified. Observing the effects of genetic variants today does not lead to automatic evolutionary understanding. An adventurous, risk-taking whist player will lose his waistcoat if he is not quite intelligent or is too emotional, or nearsighted, or credulous, or has a grumbly gut after eating a glutinous pudding, traits that are also affected by genetic variation. Evolution and whist play out only in the net effects of these and countless other factors, plus luck, so that to concentrate on the individual genetic traits is a mistaken way to view human evolution, biological or cultural. Again, this does not mean that culture has no bearing on our biology or vice versa. If there is any profound anthropological truth, it is that our biology and culture have evolved together from the beginning. Just as some people are genetically more susceptible to heart disease and need medication, some people may be more susceptible to heavy gambling or voting Republican, and perhaps they, too, need therapy. But to understand such traits, or whist, in the twenty-first century, evolution's third century, some of the anthropologists in evolution's first century were right: Culture has to be understood in cultural, not genetic, terms. Unfortunately, as the failure of Marx and Spencer shows, the challenge to explain culture in cultural terms is not new. If we care to solve the important central problems of social science, we need better cultural anthropology. We cannot expect genetics to provide the solution.24 Whether elements (or elememes) of existing theories will coalesce in some new form, only time will tell. We can hope that a return to evolutionary social anthropology will make the attempt. But even that would still leave us with the question of whether there must exist at least some kind of meaningful theory if we can but discover it, that applies with more than superficial similarities to both culture and biology. Humans will eventually go extinct and our evolution will cease. The evolution of culture may be responsible for our final state, but Marx and Spencer, who came to opposing views, cannot both be right about what that state will be. Despite what they or even Darwin said, nothing in science suggests that evolution has reached or is aiming toward any particular resolution, unless it be that our demise will restore some ecological imbalances we have caused, to the benefit of the world's species other than cows, corn, and chickens. Still, the arrangements of destiny are interesting even if they are unpredictable. In life, Marx and Spencer never interacted, so we can not know how they might have tried to reconcile their antipodal laws of nature in light of modern genetics. In death, their Highgate destiny is to be locked in an eternal glare over the unresolved questions about their ideas. But perhaps capitalism has resolved their inherent contradictions because, more than a century later (Fig. 4), all over Britain, Marks and Spencer together provide a great place to shop for answers. Marks and Spencer: Together, capitalism style. Oxford and Aylesbury (top right), UK branch. Photos courtesy of R. K. Weiss. [Color figure can be viewed in the online issue, which is available at wileyonlinelibrary.com.] I welcome comments on this column: [email protected] Anne Buchanan and I have a blog that deals with relevant subjects at EcoDevoEvo. blogspot.com. I thank Anne and John Fleagle for critically reading this manuscript, and Mark Francis and Frank Elwell for information on Spencer and Marx. This column is written with financial assistance from funds provided to Penn State Evan Pugh professors.