"Behavioral science is not for sissies." -Steven Pinker

Thursday, February 18, 2010

Sexual Selection Theory and the Existence of Moral Behavior

Recently I have been re-examining an evolutionary explanation for moral behavior. In particular, I have come across the theory of Sexual Selection and is has changed the way in which I view the evolution of morality. In the following I try to propose that evolution could have shaped our acute moral senses and then look at the implications that having a nuanced theory of morality existence poses to thoughts on self-less behavior and egoism.

There are two major theories for the existence of moral behavior: first there is the claim that it has emerged because of societal influences, and second it has evolved through natural selection via Kin Selection and Reciprocal Altruism. Unfortunately, neither or these explanations capture the full complexity and magnificence of altruistic behavior allowing both to be discarded and morality to remain an unsolved human phenomena. I aim to reclaim morality in the name of sexual selection and show that it has a place in our human nature. Altruism and other moral behaviors are evolved traits though sexual selection, and even though they are evolved and have an adaptive function, genuine altruism void of selfish motivation still exists.

The inability for evolutionary science, both evolutionary biology and psychology, to explain altruism in terms of its adaptive function has opened the door for other disciplines to propose theories as to why altruism cannot be an evolved trait. First, the theory of it being a socially introduced behavior.

Sociologist Barry Schwartz of Swarthmore College presents altruism independent from natural tendencies and evolution, instead centering on societal influences. Schwartz writes that psychology and other social sciences have been plagued by dated theories about human behavior; aspects that we consider to be natural to humans are actually fallacious, in particular, the notion that humans are natural egoists. Schwartz concludes the following about the naturalness of altruism, "However, I suggest that there is nothing natural or inevitable about the pervasiveness of altruism-that large-scale cultural influences that regulate social relations and contribute to establishing the boundaries between self and other can have profound effects on altruism.” 1 This succinct definition of altruism proposes that culture and society shape altruism.

Now that we have examined the abridged explanation of altruism in terms of societal influence, it is time to shift and examine the evolutionary perspective on the matter of morality through natural selection. Biologist P.J. Darlington defines altruism as an action that is potentially reciprocated, potentially profitable to both the giver and receiver, and a net gain lottery balancing itself over time. He goes on to say that the evolution of altruism was extremely slow and imprecise because it emerged as a side-effect of other processes and was countered by competition. 2 The two processes that altruism according to natural selection has emerged from are Kin Selection and Reciprocal Altruism. Kin selection is an organism’s ability to increase it’s own fitness by aiding others who share relatively large amounts of genetic material with them. This is referred to as increasing an organism’s inclusive fitness, that is, the sum of their own reproductive success combined with the fitness and reproductive success of their relatives, kin. Since this behavior increases the likelihood of an individual passing their genes down to the next generation, this behavior was able to evolve. The second major evolutionary process is Reciprocal Altruism, behaving altruistically to non-kin with the expectation of future possible reciprocation. This is also referred to as tit-for-tat strategy, a strategy seen in mating and in economics. Reciprocal Altruism was able to evolve because those individuals who cooperated with others and were able to detect cheaters and not cooperate with them had survival advantages over individuals who did not participate in such a system. 3 Natural selection proposes that general altruism and other charitable actions emerged as a side-effect of these two major processes. 4

Although kin selection and reciprocal altruism can account for much of our moral behavior, there is much more to be desired in an explanation for all moral behavior. Presenting the wide range of morality, from moral leadership to charity, to peacekeeping, to romantic generosity, makes the side-effect hypothesis seem incapable of explaining the vast range of morality seen in humans. This explanation also leaves much to be desired concerning the nature of virtue proposing that all virtue is the side-effect of “nepotism and economic prudence.” 5 Also it cannot see the survival payoffs in what the human mind is best at, humor, story-telling, gossip, art, music, ornate language, imagination, religion, and morality. 6 I will be moving on from the theories of natural selection in explaining morality to the theory of sexual selection from which I will paint a much more elegant portrait of morality.

Sexual selection refers to sexual competition within a species that affects relative rates of reproduction. 7 Plainly put, to survive an organism needs reproduce, and in a sexually reproducing species such as humans, this involves a degree sexual competition among males and females. The implication of sexual competition is that traits and behaviors can provide survival benefits if they are able to help a male or female copulate with another attractive male or female. If a trait or behavior emerges in males that females find attractive, even if the quality serves no survival adaptation, it can be selected for in a population simply because males with this trait have reproduced more than males who do not have this trait making the genes for the trait more prevalent in the next generation. In this way, traits that do not seem adaptive can evolve and become present in a population. In fact, the traits that evolve through sexual selection tend to be traits that are actually the most maladaptive.

During sexual selection, males and females will choose a partner to mate with based on cues that they can perceive in their partner telling them about their fitness. It is crucial for an organism to find a mate that is genetically fit in order to pass down their genes to the next generation. Unfortunately, most organisms do not have the ability to inspect the genes of their partners directly, so, individuals will use indirect cues that have the ability to predict their partner’s fitness. Males and females will look for these traits that can tell them whether or not their partner has good genes. In this way it becomes advantageous for an individual to broadcast the quality of their genes compared to the quality of genes found in other members of the same sex in an attempt to attain the best mate.

It is in this display of genetic quality that males and females separate in their mating strategies. They do this because of the differing physiological stress and time sexual reproduction requires. Long story short, males because of the minimal physical demands required for mating develop elaborate ornamentations and courtship rituals in order to please the choosier female. The female is much choosier because of the much higher physical demands placed upon her during sexual reproduction. This makes her decision on which male to mate with much more critical than the males decision prompting her to choose a mate with caution. This causes an arms race between female preferences and male ornaments to emerge creating and evolutionary treadmill. The choosier females get, the more extreme males will get creating a self-reinforcing cycle; a positive feedback loop. 8

The need to broadcast greater and greater genetic fitness has lead to the development of traits that become more and more costly to the individual. Arbitrary heritable female preferences and male ornamentation interact with one another to produce these grand displays. Female preference is a tricky thing, and its force on the evolution of male ornaments is particularly tricky. Take the following example: in a large group of females one is born with a preference for males with small shoulders and another with a preference for large shoulders. Now it is known that a man’s shoulder size is a good indicator of his fitness, but female preference is not directly driven by what is best. One female prefers males with small shoulders so chooses to mate exclusively with these males and the other who inherited the preference for large shoulders mates with men who have large shoulders. Although neither group has a conscious awareness of it, the female who indiscriminately decided to mate with men with large shoulders will have children who survive better than the female who chose to mate with less fit males.

Over time the inherited preference for large shoulders will become more prevalent in a population of females and make having broad shoulders attractive. When the display for large shoulders becomes attractive to females, males will respond by advertising this trait seeking to mate with the best female. As more and more males become selected for broad shoulders, the male population will continually have males with randomly broad shoulders be favorably selected by females. Over time, this process can rapidly increase the size of male shoulders to levels that may seem useless, but serve the purpose of increasing a male’s chances of mating. 9 Females prefer males with traits that indicate fitness and drive their evolution because previous females who preferred those same traits had offspring who were more energetic and successful surviving more often than children who did not have these qualities. 10 With this, it also important to note that many other mating rules exist for choosing the best mate you can and so forth.

So far we have looked at how sexual selection can allow traits to evolve that have no value other than in their ability to attract the opposite sex, and we have seen how arbitrary female mate choice is, preferences only evolving because of the fitness correlated with their decision resulting in more offspring with that similar preference. So, when it comes to looking at morality through the lens of sexual selection, the same rules apply.

In the words of Geoffrey Miller author of The Mating Mind, “We have the capacity for moral behavior and moral judgments today because our ancestors favored sexual partners who were kind, generous, helpful, and fair.” 11 Traits that males ornament themselves typically involve a large amount of genes. These traits are difficult to maintain because mutations are inevitable, and when something, like the brain, involves an enormous percentage of an organism’s genes, any defects in the organism’s genes will be evident. This makes this trait an accurate indicator of fitness allowing it to be selected by females over and over escalating the size of the trait beyond possible means for the male. The selection of traits that are difficult to maintain is referred to as the Zahavi Handicap Principle, that indicators of fitness will encumber the male and require of him a high level of fitness to survive. An example of this in nature is found in the extravagant plumage of the male peacock. Geoffrey Miller writes, "The peacocks tail is not just a cheap, transient advertisement visible only to peahens. It is heavy, encumbering, hard to grow, hard to preen, and highly visible to predators. Peacocks have to drag it around everywhere they go. Unfit peacocks might be able to grow large tails, but they would not be able to carry them while finding food, or fast enough to escape from predators. Only highly fit peacocks can afford large tails." 12 In the same way that peacocks have to carry an expensive ornament to please the peahens, Zahavi suggests that most sexual ornaments are handicaps. The incredible mental step that we now must take is realizing that the Zahavi Handicap Principle applies not only to peacocks, but directly to the existence of moral behavior and altruism.

Altruism is a behavior that evolved as a fitness indicator continually being selected during mating encounters because of its accuracy in displaying fitness, not a behavior that has emerged because of a particularly conducive current social atmosphere as Schwartz proposed. It fits all of our guidelines for a fitness indicator: its difficult to maintain, it requires a large amount of genetic quality to have the brain capacity to be good at it, and it varies among the population. Altruism operationally defined as agreeableness has been shown to have genetic variability and influence as well. Adoption studies performed by Mednick, Brennan, and Kandal have shown that altruism indeed has a partially genetic basis. Children whose biological parents were criminals are just as likely to show similar characteristics in adulthood regardless of being raised by their biological parents or by their adopted parents. 13 Understanding altruism as a sexually selected trait, it would be an appropriate time to review what this entirely means and implies.

The arbitrary nature of sexual selection has selected altruistic acts to be attractive because of their fitness indicator capability, not because these actions are “good for the species”. Altruism didn’t evolve because it promoted the well-being of the population the actor was in. To reiterate, altruism formed because it was a good fitness indicator. Because of the unpredictable nature of sexually selected traits, from peacock feathers to large brains, the question to why altruism evolved instead of a more malicious behavior equally capable of displaying fitness is an important question. The suspected reason that altruism evolved as a fitness display is because of Equilibrium Selection. Groups could have just as likely preferred behaviors that were not as group beneficial as altruism converging on a different equilibrium (equilibrium in this context is referring to a state where a similar behavior is preferred, such as an equilibrium finding malicious behavior attractive) as long as the behavior was a good fitness indicator. The thing is, group competition can help choose which equilibria will evolve. Individual sexual benefits, not group benefits, always maintain the equilibrium, but the advantages that the group who behaved altruistically had over the group that didn’t could have been enough to allow this behavior to develop. 14 We will soon examine the implications of altruism having evolved in this manner when we examine the nature of moral arguments.

So here we are: can genuine altruism exist. We have now seen how altruism evolved for the gains of the individual, not through survival advantages, but through sexual advantages. The question arises as to whether or not the reality of its existence as a selfish behavior implies anything about the legitimacy of the altruist. Can someone behave genuinely altruistically even though their capacity to behave altruistically, in fact for them to even comprehend their own behavior, evolved as a sexually selected fitness indicator with the purpose of enhancing the individual’s fitness? To answer this question we need to understand the extent to which individuals are aware of the selfish acts that they commit and the importance of evolution in the validity of morality.

Because of the arbitrary direction that sexual selection runs by, moral behavior cannot be seen as either an end-all code of conduct, or as a useless artifact of previous sentiment. Courtship displays are not regulated in their size by an understanding sentient being preventing an organism from dying out due to its own desire to please the opposite sex. On the contrary, there are numerous examples of animals going extinct due to extraneous fitness indicators (A good example of this is the Irish Elk with antlers over six feed wide going extinct). 15 Considering that moral behavior is a fitness indicator in the same way that antlers for the elk were, to say that our current form of morality holds legitimacy over any other possible code of conduct also seems erroneous. Although, this is not to say that our ancestors were selecting individuals with artificial kindness. In fact, our ancestors were looking for morality and altruistic behavior that was genuine. Even though we need to look at our morality with the understanding that it has been sexually selected over many generations, the understanding that what was selected was seen as genuine needs to be considered as well. 16

So our ancestors selected individuals who displayed what they perceived as genuine altruism, does this change the fact that they were selecting it with the subconscious motivation to find the best genes in which to carry on their own genetic material? Perhaps an analogy could be of use. Art, like morality, is a sexually selected trait that carries with it emotional baggage. Although we suspect that it has evolved and flourished because of the high fitness demands associated with its creation, it is important to understand how the artist feels about their work. Geoffrey Miller writes what a conversation with a Satin Bowerbird would be like, a bird whose males spends absurd amounts of time creating enormous huts for females to examine and choose a mate from. He proposes that it would something like this,
I find this implacable urge for self-expression, for playing with color and form for their own sake, quite inexplicable. I cannot remember when I first developed this raging thirst to present richly saturated color-fields with a monumental yet minimalist stage-set... It is a happy coincidence that females sometimes come to my gallery and appreciate my work, but it would be an insult to suggest that I create in order to procreate. We live a post-Freudian, post-modernist era in which crude sexual meta-narratives are no longer credible as explanations of our artistic impulses. 17

Miller suggest that the bowerbird’s motivation lies far from the desire to procreate, but in the desire to create beauty. In my opinion, morality should be viewed in the same way. Many people do altruistic deeds because they desire to do altruistic deeds, we saw this earlier in Schwartz’s research on moral behavior. I would not propose that these individuals are secretly wishing more sexual encounters when they perform kind acts, even though their motivation may have evolved because of altruism’s ability to provide more sex. 18 Further, even though the behavior of altruism cannot create itself because we find pleasure in it, to deny the pleasure that we get would be incorrect. Our motivation to behave altruistically evolved because of the fitness benefits provided to those who behaved this way, but the motivation is separate from this distant goal. 19 We act altruistically because we have an inherent feeling that it is the right thing to do, and although sexual selection invented this motivation, the capacity for humans to exhibit genuine altruism still exists, and is actually a part of our human nature.

Altruism exists because it has been sexually selected for over time due to its ability to indicate fitness, but just because of its inherent function, the human capacity for altruism cannot be overlooked. The argument that because altruism and other moral behaviors have evolved to serve in the best genetic interests of our ancestors they also inherently egotistic is an argument that needs to be continually studied. The precise differences between motivation and action have been haunting philosophers for a long time, take for example David Hume. Hume writes of our capacity to understand the motives behind our behavior that, “Our motive or intention, indeed, frequently concealed from ourselves, when it is mingled and confounded with other motives.” 20 In closing, I would desire to final quote from Geoffrey Miller’s The Mating Mind concerning the legitimacy of morality, “The peacock’s tail is no less beautiful when we understand its sexual function. Nor should the validity of human moral vision be reduced when we understand its origin in sexual choice.” 21 Morality needs to be studied under the lens of evolutionary theory, not in an attempt to diminish its power, far from it. It needs to be studied in order to grasp the power of morality in its evolved and manifested form in humanity; morality being a natural part of our species.

Notes:
1 Barry Schwartz, “Why Altruism Is Impossible ... and Ubiquitous ,” The Social Service Review 67, no. 3 (1993): 314.
2 P.J. Darlington, “Altruism: Its Characteristics and Evolution,” Proceedings of the National Academy of Sciences of the United States of America 75, no. 1 (1978): 385.
3 Robert L. Trivers, “The Evolution of Reciprocal Altruism,” The Quarterly Review of Biology 46, no. 1 (1971): 35-37.
4 Darlington, 385-389.
5 Geoffrey Miller, The Mating Mind: How Sexual Choice Shaped the Evolution of Human Nature (New York: Anchor Books, 2000). 304.
6 Miller, 17-18.
7 Miller, 39.
8 Miller, 56.
9 Miller, 175.
10 Miller, 211.
11 Miller, 282.
12 Miller, 123.
13 Mednick, S. A., Brennan, P., & Kandel, E. “Predispositions to violence,” Aggressive
Behavior 14 (1988): 25-33.
14 Miller, 314-318.
15 Millet, 42.
16 Miller, 325.
17 Miller, 270.
18 Miller, 325.
19 Miller, 296.
20 David Hume, An Enquiry concerning the principles of Morals (Cambridge: Hackett Publishing Company, 1983). 91.
21 Miller, 321.


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