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The Genial Gene: Deconstructing Darwinian Selfishness

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Joan Roughgarden's 2009 book, The Genial Gene, focuses on the present limitations of sexual selection theory. At present, we might all think that such a stance is rubbish. We've seen nature documentaries and been taught in school why such species as the peacock have such great, bright, colorful, and costly displays. In natural selection, the most fit survive, yet having to haul massive tail feathers can leave a bird much more prone to predation. Even more, only the males have the bright displays. Sexual selection teaches us that these bright feathers attract females so that, while costly in one sense, guarantee more mates and those more offspring, who will then themselves have bright massive tail feathers.
Ironically, a 2008 study has found that the tail feathers offer no mating advantage. In this case, the theory is wrong.
Roughgarden cites this study and many more, debunking case after case of what one previously assumed would fall easily under the sexual selection banner. Notable authors in the field will also concede that the definition of sexual selection isn't even clear, and when cases such as the peacock fail, the theory is expanded.
Roughgarden, after showing the difficulty with present theory, offers another approach: social selection. Here, cooperation replaces competition. Birds work as a team to produce chicks. Sperm and egg develop in such a manner as to provide the greatest chance of fertilization. As yet, the theory is untested. However, it has been developed to match existing data and knowledge and easily provides better explanation for natural phenomena than does the current definition of sexual selection.
While I might be skeptical of many of Roughgarden's hypotheses, they are now available, as were Copernicus', Galileo's, and any other groundbreaking scientist's theories and hypotheses. Now, science can go about testing, debunking, or proving social selection.
Whether Roughgarden's theory offers a valid replacement or not, this book is a great read. Not only do you learn about social selection, but you also catch up on sexual selection theory. You learn much more than you'd see in a basic biology class, and you are left with facts and ideas that you can contemplate yourself. The writing is easy to follow, often entertaining, and very informative.

I wrote a longish review of the book in Tikkun, the magazine I work on (here: [...]), but thought I would post some conclusions from that review here to encourage nonscientists to buy the book. I was very impressed by Dawkins's 'The Selfish Gene' when it came out and have been a critical enthusiast for sociobiology ever since: believing in its potential, but troubled by its biases. I never imagined that sexual selection itself might be so riven by a bias towards male/female conflict, rather than cooperation, as to skew the scientific research and interpretation to a point of becoming something closer to pure ideology than science. Roughgarden's book argues this brilliantly, and for the most part very clearly for interested lay people. It was a revelation to me as a non-biologist.

Darwin saw two major forces in evolution: natural selection and sexual selection. Roughgarden proposes replacing the latter with a theory she calls social selection. Here are the conclusions from my review:

At the end of the book she presents a table of twenty-six assumptions or hypotheses in sexual selection theory that are contradicted by equivalents in social selection theory. All of these can potentially be resolved, as to their truth or falsity, by field and experimental research. Roughgarden already presents enough data to show that social selection must be taken very seriously, and will likely prevail. This would be a huge paradigm shift. She finds widespread and deep resistance even to testing the theories.

Let me be clear. Roughgarden's is not an argument against sociobiology, and whatever explanations and descriptions of a universal human nature it can establish. Her beef is only with a vision of sexual and social relations based in selfish, as opposed to genial, genes.

Other books have objected to selfish gene theory because it appears to be an ideology, and one fully in line with modern capitalism and competitive individualism. But this is the first book I have read that attacks the whole ideology of selfish Darwinism on a broad scale from the perspective of purely biological research. As a Christian and transgender woman, Roughgarden has plenty of ideological reasons to oppose selfish gene theory and a worldview based on binary male/female conflict and traditional sex roles. But the beauty of this book is that she has pursued in a strictly scientific manner whatever skepticism her own life experience has taught her to hold about mainstream scientific ideologies. Here she is concerned solely with what is scientifically testable and true.

The very idea that there is truth in these postmodern times, when even physicists seem to have got beyond the hope of it, is a breath of intellectual oxygen. Roughgarden writes:

"As often stated in this book the issue before us is not whether a biological nature predicated on selfishness, deception, and genetic hierarchy is appealing or repugnant compared with a biological nature predicated on teamwork, honesty, and generic equality. The issue is which of these views of biological nature is true."

And when she adds, "I believe I have shown that the overwhelming weight of data and theory reveal that the selfish-gene picture does not truly and accurately describe biological nature," I applaud both her conclusion and her courage in saying it so forcefully, for she is stirring up a storm that will rage for at least a generation.

My reaction on reading the book was, "this is a civilization-changer." Roughgarden is a powerful thinker and writer. I have rarely been as energized and delighted by a book as I was by this one. You could say it's no big deal: our capacity to choose to love each other is experientially real, however biologists choose to explain it. But to the contrary, I think we are in desperate need of significant intellectual support for a post-capitalist, post-patriarchal, post-individualistic social order that prioritizes community, caring, and interdependence. What a gas, what an unexpected bonanza, what a grace, if it turned out that's what biology supports at a genetic level anyway.

This may be the most important book, philosophically speaking, on evolutionary theory in a decade. Not everyone will be persuaded by Roughgarden's critique of sexual selection or her insistence that one must choose between it and her alternative, social selection. Nevertheless, her social selection theory offers an intriguing alternative to the assumption that sex dimorphisms evolve because superior males win choosy, coy females. Instead, organisms choose mates to be good team players in maximizing offspring reared. If Roughgarden is right, males and females evolved as allies, not enemies, and evolutionary theory needs a rethink because competition evolves in a cooperative world, not the other way around.

The book is engagingly written, well organized and provides entry points to the technical literature, making it accessible to scientific specialists and a general audience alike. The book includes sections on cooperation and teamwork, genetic and social systems for sex, and concludes with point by point comparisions of the virtues and limitations of sexual selection and social selection theories.

Social selection theory is a "two tier" theory which holds genes "at arms length" from the behaviors that structure social interactions and the dynamics of evolutionary games between organisms that form teams to pursue reproductive goals. It is particularly valuable for introducing to a wide audience Nash bargaining solutions from game theory. These form part of the cooperative basis for Roughgarden's alternative to Nash competitive equilibrium theory that has supported many of the arguments favoring sexual selection theory.

Roughgarden's Genial Gene offers an engaging, readable account of her social selection theory, and it provides a powerful critique of classical sexual selection theory. The views presented and defended in this book are certainly controversial, but they are clearly articulated and well supported with references to findings in population biology.

Roughgarden has had a distinguished and varied career as a biologist, and this book reflects that variety. The Genial Gene accomplishes several tasks at once. It offers a novel take on the evolution of sex and gender; it suggests a refinement to modeling in behavioral ecology; and it offers a new challenge to the view that selfishness is the route to evolutionary success. The book is also a valuable read for those with an interest in philosophy of science. It illustrates the complex interplay of theory, models and data in evolutionary ecology, and it invites the question of how social norms may relate to scientific findings.

I'm not a biologist, but a one-time physicist (at SLAC many years ago) and software entrepreneur/telecom executive ever since. Being generally curious, I read the Selfish Gene long ago, and was intrigued by the contrasting title of this book.

I'm only about 1/3 of the way through the Genial Gene, but wow! what a lot of interesting ideas! Not being a biologist, I presumed sexual selection was settled theory - on the surface it seems pretty logical. But then you go through the arguments presented here, and clearly the math just doesn't work.

The Genial Gene is an excellent read because
1. It's always fascinating to see science in the making, and clearly this is an area where paradigms are ripe for overturning.
2. It opened my eyes to a whole realm of very interesting problems - not being a specialist in this field, I never realized that biology had such intriguing theoretical issues
3. The writing is clear and approachable for the scientifically-inclined layperson, but it doesn't patronize or gloss over the difficult bits. Unlike, a lot of popularizations, you need to have your mind fully engaged while you read this book.

Darwin believed that there are two kinds of biological selection, natural selection and sexual selection. Natural selection is for traits that confer success the individual's battle with predators and search for prey. Sexual selection is for traits that increase an individual's chance for successful mating. Darwin was particularly struck by the fact that some traits that clearly have fitness costs in the natural selection setting seem to have fitness benefits in the sexual selection setting. For instance, the peacock's gorgeous and abundant plumage is costly to produce and renders the bird vulnerable to predators but desirable to peahens.

The issue of sexual selection was especially important to Darwin because natural selection alone appears unable to explain central characteristics of human beings. For instance, hunter-gatherer tribes rarely have languages in which counting uses numbers beyond one, two, and three, yet humans are capable of prodigious mathematical feats. More generally, of what possible survival value is there in a brain that is extremely costly to maintain, much of whose power is used in singing, dancing, creating art and music, and in other tasks far removed from the daily grind of self-preservation? Lord Russel Wallace chided Darwin at length for maintaining that humans are subject to the same biological laws as other species, and urged him to recognize the intervention of a Divine hand in the creation of our species. Darwin used sexual selection to answer Lord Wallace, saying "He who admits the principle of sexual selection will be led to the remarkable conclusion that the nervous system not only regulates most of the existing function of the body, but has indirectly influenced the progressive development of various bodily structures and of certain mental qualities. Courage, pugnacity, perseverance strength and size of body, weapons of all kinds, musical organs, both vocal and instrumental, bright colours and ornamental appendages, have all be indirectly gained by the one sex or the other, through the exertion of choice, the influence of love and jealousy, and the appreciation of the beautiful in sound, colour or form, and these powers of mind manifestly depend on the development of the brain." Darwin (1879, p. 687.)

Concerning singing and language, Darwin was even more insistent on the importance of sexual selection: "some early progenitor of man probably first used his voice in producing true musical cadences, that is singing, as do some of the gibbon-apes at the present day; and we may conclude from a widely-spread analogy, that this power would have been especially exerted during the courtship of the sexes." Darwin (1879, p. 109)

The mechanism for sexual selection for male traits was outlined by Darwin as follows. A female mutant prefers a certain male ornament, and preferentially mates with males with that ornament. Her female offspring inherit her taste for the ornament, and her male offspring inherit the ornament. Thus this preferential mating scheme is perpetrated in to the next, and all succeeding, generations. Called "runaway selection," this dynamic was analytically specified by the great Ronald Fisher, and there are currently several models of this type in the literature. This is the model of sexual selection, characterized by coy females who instigate intense competition of males for her favors, the males having no role in the sustenance of their offspring and hence of no value except reproductive value, that Roughgarden says is wrong. It its place, Roughgarden proposes a dynamic of "social selection," in which males and females cooperate harmoniously in nurturing and raising their young. Her great example is nesting birds, who work furiously and jointly to tend to their young. Males and females in nesting species both care about the quality of their partners, including not only their ability to compete in reproducing, but also foraging, defending, and provisioning a nest.

Joan Roughgarden is a very famous and accomplished biologist. In recent years, however, she has turned to a virulent form of ideological conflict-aversion that greatly interferes with her ability to get her (important) message across. It is typical of scientists to build on the successes of previous generations (Newton claimed that "if I have seen further, it is by standing on the shoulders of giants"). Roughgarden will have nothing of this. Traditional population biology is incurably infected with patriarchal bias, and must be swept away. The truth for Roughgarden is simply the opposite of the traditional theory; whence the title of the book, in contrast with Richard Dawkins' famous The Selfish Gene. In fact, both conflict and cooperation, as well as selfish and altruistic behavior, are inextricably intertwined aspects of the behavior of many vertebrate species, and the contemporary task is to augment and not simply deny traditional population biology.

Social selection is, as Roughgarden asserts, the more general category of which sexual selection is a subcategory. What is social selection (this is my story---Roughgarden may consider me an enemy, for all I know, but mine is the true story, I assert)? Every sexually reproducing species in which members interact in frequent and routine ways develop standardized patterns of interaction that involved both conflict and cooperation. These patterns are built into their genes and take the form of standardized structures and standardized behaviors. Structural standardization is called "niche-construction," and accounts for the beaver dam, the termite mound, the beehive and so on. See, for instance, Kevin N. Laland, F. John Odling-Smee and Marcus W. Feldman, "Evolutionary Consequences of Niche Construction and Their Implications for Ecology", Proceedings of the National Academy of Sciences 96 (1999):10242-10247; Kevin N. Laland, F. John Odling-Smee and Marcus W. Feldman, "Group Selection: A Niche Construction Perspective", Journal of Consciousness Studies 7,1/2 (2000):221-224; Kevin N. Laland and Marcus W. Feldman, Niche Construction (Princeton: Princeton University Press, 2004); F. John Odling-Smee, Kevin N. Laland and Marcus W. Feldman, Niche Construction: The Neglected Process in Evolution (Princeton: Princeton University Press, 2003). The importance of standardized structures is that these structures then become the environment within which the species evolves. This process is perhaps most dramatic in humans, for whom production techniques and their related cultural forms become the basis for further genetic evolution. This is the so-called gene-culture coevolution. See, for instance, Robin M. Dunbar, "Coevolution of Neocortical Size, Group Size and Language in Humans", Behavioral and Brain Sciences 16,4 (1993):681-735; William H. Durham, Coevolution: Genes, Culture, and Human Diversity (Stanford: Stanford University Press, 1991); Herbert Gintis, "Gene-culture Coevolution and the Nature of Human Sociality", Proceedings of the Royal Society B 366 (2011):878-888; Robert Boyd and Peter J. Richerson, Culture and the Evolutionary Process (Chicago: University of Chicago Press, 1985).

The social behavior side of social selection is much less well developed that the social institutional side, but it includes the typical pattern of male-female mating and care of offspring, as well as the pattern of mate characteristics typical of males and females of that species. Darwin could only think of fitness-reducing characteristics, such as the peacock's feathers, or the attractive female's physical weakness and fragility. But in fact, the characteristics of social selection develop according to the laws of natural selection, just as every other aspect of the constitution of a species. Moreover, it has been recognized for many years that the runaway selection model favored by Darwin and Fisher is not mathematically tenable (see my analysis in Game Theory Evolving (Princeton: Princeton University Press, 2009). The widely preferred theory is the costly signaling theory of Amotz Zahavi and William Hamilton (see Amotz Zahavi, "Mate Selection---A Selection for Handicap", Journal of Theoretical Biology 53 (1975):205-214; Amotz Zahavi and Avishay Zahavi, The Handicap Principle: A Missing Piece of Darwin's Puzzle (New York: Oxford University Press, 1997); William D. Hamilton and M. Zuk, "Heritable True Fitness and Bright Birds: a Role for Parasites?", Science 218 (1982):384-387). Costly signaling theory nicely embraces the dual conflict/cooperation nature of social interaction.

According to the handicap principle, females are indeed choosy, but they choose mates who have "good genes" and hence will contribute to hearty offspring. For instance, a peacock with a heavy parasitic load does not have the resources to produce beautiful plumage. A female does well to choose a mate with light parasitic load, because her offspring will inherit the immunity to pests enjoyed by her mate. To make a long story short, virtually every case of female choice is an instance of costly signaling. Female choice is thus fitness enhancing and evolves by natural selection. Of course, there are times when the cues used by the female as a signal of mate fitness are imperfect signals and can be exploited by males for their own reproductive ends. But in general we can expect mate choice and cooperation in rearing offspring to an intimate aspect of the fitness of a species, and that the strategic interaction of mates will be some pattern of cooperation and competition, as is the case of all cooperative enterprises (see Samuel Bowles and Herbert Gintis, A Cooperative Species: Human Reciprocity and its Evolution (Princeton: Princeton University Press, 2011).

Where does Roughgarden stand on all this? First, she completely rejects costly signaling theory, reciting its failures but not its successes. This is quite disingenuous; Roughgarden's social selection hypothesis is immeasurably strengthened when seen as extending and correcting costly signaling theory. Moreover, costly signaling theory does support some of traditional sexual selection theory, while contradicting other parts of the theory. Similarly, Roughgarden cannot give a cogent explanation of social selection theory because it in fact sometimes supports traditional sexual selection and sometimes does not.

This book is thus very frustrating. There is virtually no chance Roughgarden's message will be accepted by traditional population biologists because they will not get beyond her whole-cloth rejection of classical sexual selection. The layman will see the morality play of the good guys (Roughgarden and allies) against the bad guys (supporters of conflict/competition theories of all types), but they will be confused by the mass of evidence, only half of which supports her position. This book is aching to be rewritten from a balanced perspective.

Several reviews on this page imply that this book advocates a Creationist or Intelligent Design perspective. This is not true. Roughgarden is a well known evolutionary biologist and a strong advocate of Darwin's theory of natural selection (but not his theory of sexual selection). Please don't judge a book by its title/cover!

I finished reading The Genial Gene some weeks ago and found it extremely stimulating. It is about time that we stop considering cooperation in offspring rearing as some kind of epiphenomenon of unsuccessful competition. This is what the whole 'survival of the fittest' crowd does across board without acknowledging that Darwin never coined that phrase. The same orientation applies in competitive game theory.

Worse still, so-called scientists, while paying lip-service to the black-swan model of hypothesis testing, engage in hypocritical greying/whitening of black swans that threaten their cherished theories. This happens even more exaggeratedly in psychology.

Roughgarden is not taking issue with the whole of Darwinian theory, only the theory of sexual selection. She very clearly points out that Nash also developed cooperative game theory that is conveniently excluded by researchers who base their projects on competitive game theory. Roughgarden's work should be taken very seriously by empirical researchers in evolutionary biology and evolutionary psychology as a platform to investigate the relative merits of social selection and sexual selection theory through use of objective measures. Particularly for evolutionary psychology, self-report justications of 'just-so' stories should be abandoned and studies that directly compare the two hypotheses using observable methods should be designed.

To dismiss Roughgarden's work solely on the basis that it challenges cherished orthodoxy is to risk propagating poor science as well as ignoring the heterosexist subtext of sexual selection theory.

I recommend this book for readers interested in the philosophies and theories of evolution. More specially, Joan Roughgarden's theory of social selection in The Genial Gene offers a way to revive the troublesome field of evolutionary psychology for biologists. This work will certainly open discussion on the once blindly accepted sexual selection of Charles Darwin. The Genial Gene's greatest strength is the narrative of evolution, and life, at large that it shares. While this book is largely about animals, I believe an important point of discussion is its potential application to humans, and evolutionary psychology. Roughgarden's writing exemplifies the ideal of popular science writing-- accessible, fun, and well-supported by primary research. Much of the narrative of evolutionary psychology is founded upon sexual selection, a theory that is currently being challenged. Social selection contrasts the traditional sexual selection theory that emphasizes "selfishness, deceit and coercion as naturally inevitable" with one that emphasizes "cooperation". In social selection, the importance is not the mating event, but the rearing of the offspring; social behavior views having offspring reproducing system as opposed to a mating system. As both a psychology and biology major, I found The Genial Gene to be an essential read for all academics as an important starting point for a discussion on diversity in evolution and sexual selection.

Currently, the rhetoric behind many evolutionary psychologists is sexist and misguided, such as Ronald Fisher's runaway theory, which both says that females are choosy but denies them agency to have developed their brains for other reasons than to appreciate the males. Roughgarden's The Genial Gene is transformative because it offers a new and comprehensive view on this rigid theory; her theory of social selection is contemporary and inclusive. If the narrative emphasizes mutual mate choice, like sexual selection, we have to ignore non-traditional genders of strong male and caring female. If the narrative emphasizes the rearing of the offspring, like social selection, we can now include the multiple genders and resulting relationships among them that are evident in our environment today and the past. Social selection allows evolutionary psychology to build a narrative that is not premised on the dimorphism of brain size and distinct, traditional genders that do not exist.

In theory, evolutionary psychology is promising, but in its short history, the sexual selection narrative has corrupted its interpretation of the evolution of the human mind. In order to refocus evolutionary psychology away from sex differences and mate choice based on overgeneralizations, future supporters need to be from a variety of backgrounds, including both more women and more supporters of social selection. Since the power of science ultimately lies in the verifiability and presence of empirical data, it is important to form new testable hypotheses such as through behaviourial dynamics and game theory, like we find in The Genial Gene. A social selection interpretation can then, more accurately, fit the biological evidence. If "evolutionary psychology is dead, but doesn't seem to know it yet," as Paul Ehrlich, a biologist, once remarked, then social selection will be the force that resurrects it wholeheartedly.

I absolutely loved this book. It was extremely enlightening and detailed, providing many legitimate and new arguments against sexual selection. For those looking for an interesting read about evolution with scientific data and concrete examples, this is a must-have. It's also a relatively easy read and isn't convoluted with confusing jargon.

As a Bio major at Stanford, I've never been taught or introduced to anything like this, and it was an incredibly educational text that EVERY scientist, if not person, should read. It's a shame our high school and university curricula don't include any of this because it effect just as many social issues as evolutionary ones.

This book isn't simply about evolution and sexual selection. It addresses the holes in the theories, and proposes new ones. The message is beyond just science, but you'll have to read and find out why.

There's not doubt you'll be the source of many cocktail conversations.