john hawks weblog

paleoanthropology, genetics and evolution

cooperation

  • Developing the sharing sense

    Mon, 2011-03-21 01:11 -- John Hawks

    Following on after yesterday's post about hunter-gatherer population structure, I ended with the proposal that cooperation may be a "cognitive technology" in the same way suggested for numbers ("Number as cognitive technology").

    The technology perspective attracts me. It seems a productive way to examine the interaction between innate and extrinsic factors leading to human behaviors. We learn about numbers. Without a development of the brain within a cultural setting with widespread counting and training in number use, people don't develop the habits of mind that allow rapid comparison of cardinal values. They can still operate on sets of objects and compare their quantities, but they are missing a shorthand, a symbolic shortcut, that comes with learning and practice. Numerical concepts, invented and repeatedly used by human societies, give learners access to this symbolic method of problem-solving.

    Cooperation and other prosocial behaviors are similar in some respects. Whether you share with another person or not in a particular concept depends on the rules about sharing that you learned as a member of your society. What's interesting is that these rules change with age in various ways. So I went looking in the developmental psychology literature for some data about how kids share. My notes here are just a start -- and I'm pretty sure they're rough to read near the end -- but I found it interesting how the data seem to illuminate the issue of cooperation in the archaeological record.

    Toddlers

    Toddlers can, in some circumstances, exhibit a surprising degree of understanding about the intentions of others. They can also be surprisingly helpful -- that is, they can see when another individual wants something, and can actively help that other person to get it. A paper last fall by Kristen Dunfield and colleagues [1] gives a nice review of this kind of helping behavior in toddlers aged 18 and 24 months.

    Replicating previous work by Warneken and Tomasello (2006, 2007), we found that by 18 months, infants are beginning to identify the situations in which helping behavior is required; that is, they will aid instrumentally by retrieving an item that is out of a person’s reach, thus fulfilling another’s unmet goal. Further, the present study found a similar frequency of helping behavior to Warneken and Tomasello (2006), even though in the current study participants only received one experimental helping trial as opposed to the three trials they received in the previous paradigm. In light of previous studies, helping behavior may also be seen as young as 14 months, though the contexts in which it occurs are less flexible, owing perhaps to an emerging understanding of goal-directed activities (Warneken & Tomasello, 2007), recognition of the means by which certain unmet goals can be fulfilled, and the physical ability to mediate the completion of the goal.

    However, as I well remember from my own toddlers, the "prosocial" characteristics of infants can be temperamental, to say the least. Dunsfield and colleagues considered 18 and 24-month-olds, finding substantial heterogeneity among individuals in the kind of helping or sharing behavior they exhibited.

    While acknowledging the dangers of arguing from a null effect, it is the case that although the majority of the participants engaged in at least some prosocial behavior, there were no correlations between the various prosocial behaviors. Further, the most common pattern of response was to engage in only one type of prosocial behavior (helping or sharing). Although the tendency to engage in prosocial behavior in general tended to increase across our two timepoints, the increase was not the result of systematic development within or between the various subtypes of prosocial behavior. Thus, we have no evidence in the present study for “across the board” prosocial behavior within individuals in these two age groups. With future research that explores the consistency both within and between the multiple specific types of behavior, and that considers enduring behavior over time in a longitudinal manner (Eisenberg et al., 1999), it may be the case that helping, comforting, and sharing do not cluster together within an individual’s repertoire and perhaps should not be grouped together as one general category of unified behavior in infancy.

    A natural question is, what does it take to manage any kind of sharing at all among children this young? By this age most children have experienced thousands of times when an adult or another caregiver has performed the opposite role, giving the child what she cannot reach herself. This long history of positive exemplars for sharing and cooperative behavior nevertheless leaves substantial variation among children in how they actually behave in a similar context.

    The first article by Warneken and Tomasello cited above [2] compared human children with chimpanzee juveniles of a similar age. They showed that the human children did show these prosocial tendencies by 18 months, but that so do chimpanzees -- at least to a certain extent. The chimpanzee juveniles handled the most indexical of the tasks relatively well -- the case where a person is reaching for something but needs help to reach it. Other tasks didn't bring out the cooperative nature in chimpanzee juveniles:

    However, the chimpanzees did not help the human reliably in the other types of tasks—that is, in those involving physical obstacles, wrong results, or wrong means. In a follow-up study, we gave them two additional tasks of these types—designed to make the human's problem especially salient and with more time for a response—and they still did not help in these tasks (14). Presumably, when someone is reaching with an outstretched arm toward an object, the goal is in principle easier to understand and the kind of intervention follows straightforwardly. This could explain why out-of-reach tasks (in contrast to the other scenarios) elicited more helping by children and the only instances of helping by chimpanzees. Children and chimpanzees are both willing to help, but they appear to differ in their ability to interpret the other's need for help in different situations.

    This goes some distance toward explaining what children need to make them potential helpers. They need some way of figuring out the goal of the person who needs help, and they need to have no goal of their own that directly conflicts. Before Warneken and Tomasello's work, chimpanzee juveniles had not shown signs of such prosocial behaviors in other experimental contexts. Those authors attribute the difference to food: Most chimpanzee experiments had involved food treats, attempting to get individuals to share food with each other. The chimpanzee's own desire for the food may directly interfere with the goals of other individuals -- a conflict that is hardly likely to lead to sharing, even in human toddlers.

    There is little sense in calling the chimpanzee behavioral pattern "rudimentary", as psychologists sometimes do. The human pattern here is rudimentary compared to the extent of helping and sharing that occur later in childhood. The human children in this context seem to have an ability to diagnose the intentions of another individual more than do the chimpanzees. They also seem to have more patience for helping, in some sense. Warneken and Tomasello returned to the topic in a 2009 review [3] that puts forward the situation with respect to sharing, helping, and information transfer. They note that human language depends on cooperation in a way that chimpanzee vocalizations do not. It may not be coincidental that language is learned across the same ages as cooperative behaviors.

    Preschool-aged children

    Olson and Spelke [4] reported on a slightly more intricate study with 3.5-year-old children. They assessed sharing behavior in which children had to divide a pool of items among a number of recipients. These potential recipients sometimes included both relatives and strangers. In other instances, the potential recipients varied in terms of whether they had interacted with the children by sharing with them. Olson and Spelke intended to find whether children of this age would engage in direct and indirect reciprocity, and whether they would skew their distribution of the resource toward relatives as opposed to strangers.

    What they found is that kids of this age typically divy things up fairly:

    Children may have distributed resources equally on the four-resource trials for either of two reasons. First, it is possible that children will resort to equal sharing whenever resources are plentiful and will favor family, friends, reciprocators, and generous others only under conditions of scarcity. Such a possibility is consistent with the finding that social conflicts among older children and adults arise primarily when resources are limited ([Jackson, 1993] and [Sherif et al., 1961]). Alternatively, the equality response may be driven by a predisposition to distribute resources in a one-to-one correspondence with recipients whenever such a distribution is possible. That predisposition, in turn, could arise either spontaneously or through the internalization of an explicit rule children are taught by parents and other adults.

    As soon as they can manage matching objects with people, they are parceling out things one to a person. That's obviously an integral part of most children's experience -- everything from passing out parts in a game, to passing out food at dinner. So the behavior itself is highly reinforced if not explicitly taught, and it may well be explicitly taught to most children.

    The children in Olson and Spelke's trials also tended to share more with people who had previously been generous in the past, either directly or indirectly to the child. By rewarding past generosity, the children were fulfilling their end of a reciprocity arrangement. This seems pretty relevant to the dynamics in ancient human groups; if a 3-year-old can manage the basics of reciprocity, it may not have taken much to push people into a stable hunting and gathering economy, which is based on reciprocity.

    School-age children

    Here's what interested me the most. Kids at 3.5 years already get the idea of sharing equally and fairly. So you might think this would be deeply ingrained in older children. But instead what we see is that older children start to reason more and more like adults, which ironically makes them share less evenly. They just get more clever about how to rationalize their choice to be unfair.

    For example, a nice study by Gummerum and colleagues [5] compared students age 9 to age 17 for their performance in the "dictator game."

    The "dictator game" is an experimental model that has been repeatedly employed in adults to study the themes of cooperation and altruism. An individual is given control over how to divide a single sum between herself and another anonymous person. The individual can choose any division down all for himself and zero for the anonymous player.

    Gummerum and colleagues added a twist, making individuals work in groups of three to decide on their offers. The offers then reflected not only the preferences of individuals going into the study but also their moral reasoning with each other after discussing the offers in small groups. This yielded an interesting, almost ethnographic picture of how the children came to make their decisions about appropriate offers.

    They found that the offers made by groups were strongly influenced by the level of moral reasoning employed by group members. When a student who favored a low offer was arguing at a higher level of sophistication, the group was more likely to adopt a low offer. And vice-versa -- when the clever student was arguing for a more equitable offer at a higher level, the group was more likely to give more. Girls gave higher offers than boys in the experiment as well.

    In a game like this, the sharing and reciprocity aspects of prosocial behavior are transformed into moral questions. No punishment befalls students who choose to make low offers in the dictator game; yet there is the consideration of self-regard. And others have heard the arguments that a student makes, affecting her reputation. Moral reasoning is, in other words, public.

    Concluding thoughts

    What I find so interesting about comparing children of different ages, is not about cooperation but instead about how the rules are shifted to higher levels of description. Sharing and reciprocity are quite simple, and children can manage them young, although irregularly. Kids can learn about sharing and helping in a rather unsophisticated way, and their performance reflects very simple expectations. Equal division, turn-taking, and punishment of defectors are all integral parts of early childhood.

    Obviously, any humans living in foraging societies in the recent past have grappled similarly with the moral aspects of cooperation and altruism. But that moral reasoning comes at an age far past when children are taught about the importance of fairness, sharing and helping. The kind of dynamic that concerns many anthropologists -- how do foraging peoples maintain the rules that underlie reciprocity and altruistic behavior -- is simply at a different level than the dynamic that actually inculcates cooperation. Yet with children who learn systematically to help and cooperate, such behaviors have a much higher chance of existing stably, even in small societies. If there is any cognitive invention that a human society would not want to lose, I think some conception of fairness may be it.


    References

  • The chimpanzee males who adopt orphans

    Mon, 2010-02-01 22:59 -- John Hawks

    The value of long-term field studies: Christophe Boesch and colleagues report on adoption in the Taï Forest chimpanzee study population -- where more than 30 years of observations have produced 18 well-defined cases of adoption of orphaned individuals. They considered "adoption" to be the provision of maternal care (e.g., carrying, feeding, food sharing, defense) for more than two months. It's possibly unfortunate terminology, as it leads to headlines like mine. Yet it is really interesting behavior.

    It would be nice to say that these cases represent 18 happy endings, but these adoptions did not increase the probability of survival compared to orphaned individuals who did not receive ongoing care. There were a couple of cases where females breastfed orphaned infants "for many years," but there seem to be several sad stories too.

    Sometimes, the care for the orphaned juveniles was given by males:

    Remarkably, all adult males of the East Group that adopted young orphans went a step further by investing in unweaned small infants and carrying them dorsally during travel for many months (see Figures 3 and 4 of Porthos with Gia) (Table 3). Since, Taï chimpanzees walk about 8 km per day on average, this represents a notable investment. Porthos' adoption of Gia lasted for 17 months, until his death due to Anthrax, and he was seen to carry her even in extremely risky situations, such as during encounters with neighboring communities [26]. Furthermore, some males were seen to share their night nest with their adopted infant (Table 3). Fredy, the 3rd ranking male of the East Group, adopted Victor, the son of Vanessa, who died from Anthrax in late December 2008, and shared his nest with him every night, carried him on his back for all long travels, and shared the Coula nuts he opened from December 2008 to July 2009. For example, on February 17th, Fredy cracked 196 Coula nuts for 2h05mn and shared pieces of 79% of them. This gives a measure of the altruistic investment made in an unrelated infant.

    That sounds pretty amazing. I think it's very relevant to human evolution, as orphaning must have been very common with the high mortality rates of the past.

    The authors propose that adoption is basically a side effect of prosocial behavior in these chimps brought on by leopard predation:

    [T]he resulting high predation pressure exerted by these cats seems to have promoted strong within-group solidarity in the form of care for all injured individuals as well as joint coalition defense against the leopards [16], [26]. Once established, this care for the welfare of others seems to have been generalized to new social contexts, including adoption [26]. Any discussions about the evolution of altruism must include the caveat that dissimilar socio-ecological conditions will lead to important population differences in both chimpanzees and humans and we need to remain very careful before making any claims about species differences.

    Well, if there was such a simple psychological correlation between various kinds of prosocial behavior, it would make life a lot easier for those of us trying to figure out what happened to humans.

    References:

    Boesch C, Bolé C, Eckhardt N, Boesch H. 2010. Altruism in forest chimpanzees: the case of adoption. PLoS ONE 5:e8901. doi:10.1371/journal.pone.0008901

  • Robot genetics

    Sun, 2010-01-31 10:01 -- John Hawks

    Dario Floreano and Laurent Keller describe experiments that combine genetic algorithms and robots. It's a review essay rather than a description of new research, but unlike most descriptions of "evolutionary robotics", it's actually directed toward biologists instead of AI researchers.

    In this essay we will examine key experiments that illustrate how, for example, robots whose genes are translated into simple neural networks can evolve the ability to navigate, escape predators, coadapt brains and body morphologies, and cooperate. We present mostly—but not only—experimental results performed in our laboratory, which satisfy the following criteria. First, the experiments were at least partly carried out with real robots, allowing us to present a video showing the behaviours of the evolved robots. Second, the robot's neural networks had a simple architecture with no synaptic plasticity, no ontogenetic development, and no detailed modelling of ion channels and spike transmission. Third, the genomes were directly mapped into the neural network (i.e., no gene-to-gene interaction, time-dependent dynamics, or ontogenetic plasticity). By limiting our analysis to these studies we are able to highlight the strength of the process of Darwinian selection in comparable simple systems exposed to different environmental conditions.

    Some of the simplest machine learning experiments are basically like those used in behavioral psychology -- put the robots in a maze, make them remember where the food is, that sort of thing. Robots are simpler than rats, so the researchers can reverse-engineer the "evolved" software at the end of a series of experiments to see what worked and why:

    Interestingly, the driving speed of the best-evolved robots was approximately half of the maximum possible speed and did not increase even when the evolutionary experiments were continued for another 100 generations. Additional experiments where the speed was artificially increased revealed that fast-moving robots had high rates of collisions because the 300-ms refresh rate of the sensors did not allow them to detect walls sufficiently in advance at high speed. Thus, the robots evolved to move at intermediate speeds because of their limited neural and sensory abilities.

    Figuring out that particular optimization would drive a team of human programmers crazy. Can you imagine? "Why do they keep running into that wall?!"

    On the other hand, dumb selection took a lot of generations to get to that point. You can't say selection was more efficient. If you had a crew of programming grunts and forced them to sit in a room for 100 robot generations, they'd come up with something.

    It's quite possible that a human would have come up with much better software, by pushing the robots past the limits of mutations on their "genomes". Selection has its own "sensor limitations", it can get stuck in a local optimum, and depends on the mutation structure to explore the landscape.

    It helps if the landscape has some strong correlation structure. That's what came to my mind as I read their account of experiments to make robots cooperate:

    However, when the arena contained both large and small tokens, the behaviour of robots was influenced by the group kin structure. In groups of unrelated robots (i.e., robots whose genomes where not more similar within than between groups), robots invariably specialised in pushing the small objects, which was the most efficient strategy to maximise their own individual fitness them (i.e., large tokens provided an equal direct payoff as a small token but were more difficult to successfully push). By contrast, the presence of related robots within groups allowed the evolution of altruism. When groups were formed of “clonal” robots all having the same genome, individuals primarily pushed the large tokens even though it was costly, in terms of individual fitness, for the robots pushing (Video S6).

    If you wonder how robots have "kin", it's that they share similar (or the same) genomes. The simplicity of the behaviors suggests a functional explanation for kin selection -- for many kinds of tasks, it may simply be easier to cooperate with other individuals who "work" the same way. Different approaches to the same task may clash.

    They describe a similar result for cooperation by information sharing:

    Similar results were obtained in experiments where groups of light-emitting, foraging robots could communicate the position of a food source at a cost to themselves because of the resulting increased competition near food. In these experiments, robots again readily evolved costly communication when they were genetically related, but altruistic communication never evolved in groups of unrelated robots when selection operated at the individual level [38],[39].

    The next logical step for this kind of research is nano-scale: evolutionary robotics on molecular machines. Which is scary. I hope they have the sense not to train them up by eating biological systems...

    There's this old course on the books here, "Human aspects of robotics". I suppose it was taught back in the 80's when robots looked like they would replace all the manufacturing workers. I've often thought that someday it may be revived as with robots as the heroes instead of the villains.

    References:

    Floreano D, Keller L. 2010. Evolution of adaptive behavior in robots by means of Darwinian selection. PLoS Biol 8:e1000292. doi:10.1371/journal.pbio.1000292

  • Evolving swarm bots

    Tue, 2009-10-27 09:48 -- John Hawks

    Robot swarms programmed with genetic algorithms to "evolve" their behavior:

    A more recent 2009 study, again at Lausanne, suggests that swarms of bots don't just evolve cooperative strategies to find food (or avoid poison), they can also evolve the ability to deceive. Bots equipped with artificial neural networks and programmed to find food eventually learn to conceal their visual signals from other robots to keep the food for themselves. “Forget zombies,” a post on Current TV's blog comments about the little bots, “this is the real threat.” (Fortunately, these experimental bots don’t eat brains – at least, not yet.)

    A peeve: I wish people would stop using the word "learn" for this kind of thing. The robots aren't "learning" anything; their genetic algorithms are randomly changed and then subjected to a round of selection. I'm not sure they really qualify as "swarm bots" either, if they're competing instead of cooperating.

    Anyway, the article references my UW colleague Chuck Snowdon's work:

    Communication is very important for social organisms to ensure their ecological success. For example, University of Wisconsin-Madison psychology professor Charles Snowdon offers a perspective on what the early environmental conditions may have been that led to the hominid communicative explosion. His research into the world of nonhuman primates suggests that while apes and monkeys in the Old World tend to be relatively silent creatures, the New World is home to much noisier monkeys such as tararins and marmosets that vocalize more frequently to “show more richness of development and learning in their vocal patterns, and that appear to transmit more information with the sounds they produce than do any of the Old World primates.”

    A key reason, he suggests, is cooperative breeding, which is found in the New World animals to a much greater extent than in the Old World monkeys and apes. New World primates live in circumstances where engaging in rich communicative exchange is advantageous, because parents (and alloparents -- aunts, uncles, and others) engage in cooperative rearing and need to communicate about it. This, Snowdon suggests, may be a critical factor that differentiated our early hominid ancestors from their ape cousins.

    I think monkeys are much more of a threat than bots. Now, if there were swarming monkey bots, that would be different.

  • A Snowdrift game version of hunting

    Fri, 2009-06-05 23:39 -- John Hawks

    I want to run through some examples of how we can apply game theory to consider hunting decisions in human groups. First, I describe a simple Snowdrift model applied to hunting. This is part 2 of a series, part 1 introduces the topic of the Snowdrift game.

    A reader sent along a story after reading the first post:

    In reading your snowdrift blog post, I was reminded of an experiment that does not require game theory to understand. You may have heard of it. Two pigs are in a pen. One is dominant. To get food one of them presses a bar, but the food is dispensed at the other side of the pen. If the subdominant pig presses the bar, it gets no reward, as the dominant pig hogs the food, eating it all. The result is that the dominant pig presses the bar while the subdominant pig waits at the food trough. Then the dominant pig rushes over to the trough to push the subdominant pig aside. Both pigs get fed, but the dominant pig does all the work

    It's a great example of asymmetrical rewards. I'll get to those in the next few posts on this topic, because the asymmetries are very important to understanding dynamics in hunter-gatherers. But first, we have to describe the simple symmetrical case, including the algebra defining the evolutionarily stable equilibrium between the two simple strategies.

    Suppose we have two hunters, who will share whatever game either of them kills. A man may choose on a given day to hunt. By hunting, he suffers a cost c and brings back a large fixed benefit b for each man. The two men may both choose to hunt on the same day, resulting in the same benefit b but a lowered cost c∕2 for each man. The two men decide whether to hunt simultaneously and without conferring — that is, there is no information transfer between them that would affect their decisions.

    Here is the payoff matrix of the game for player 1 (choices on left) given the strategy selected by player 2 (on top):

    hunt no hunt
    hunt b - c∕2 b - c
    no hunt b 0

    Given the existence of the two strategies, “hunt” and “no hunt,” the ESS is the ratio at which the two strategies have equal expected returns. If individuals select a strategy and do not vary, the ESS represents the frequency of these variants in the population. If in contrast, individuals can choose to adopt either strategy, then the ESS also will be the optimal proportion of the two strategies in one individual’s repertoire. The two strategies will yield equal payoffs when the ESS satisfies the following equation, where p represents the proportion of “hunt” and 1 - p the proportion of “no hunt”:

    p(b- c∕2)+ (1- p)(b - c) = pb
    (1)

    …which simplifies to p = 2(b - c)(2b - c). That expression is positive where b > c, and approaches unity where c is very small relative to b. If in contrast b c then the scenario is the Prisoner’s Dilemma, where the only ESS is a pure “no hunt” strategy.

    Let’s also look at a slightly different case. As above, each man’s return from hunting will be b regardless of whether one man or both choose to hunt. But in the payoff matrix below, the cost of hunting is also the same whether one man or both choose to hunt. So there is no reduction in the cost of hunting if both men do it.

    hunt no hunt
    hunt b - c b - c
    no hunt b 0

    Now, in this case, the ESS satisfies the equation:

    p(b- c)+ (1- p)(b - c) = pb
    (2)

    Again, p is the frequency of the “hunt” strategy. This simplifies to p = (b-c)∕b, which again yields the Prisoner’s Dilemma when b < c.

    OK, that’s the simple Snowdrift game model, described in the language of hunting instead of winter car accidents. It is quite simplistic in many ways. We might expect real hunters to have successes and costs that vary as stochastic functions of the environment. A real hunter must decide whether to hunt based not only on the odds his companion will hunt, but also upon some appraisal of the companion’s likelihood of success. Men in hunting societies are not paired up by the buddy system, but instead make their decisions about hunting in the context of a larger group’s activities.

    Maybe most confusing, there are two possible kinds of currency in which benefits and costs may be expressed. A benefit from hunting may be most naturally measured in calories. If we average hunting returns across many episodes, then our result would be mean calories per day, or per hour of effort. Likewise, it might seem natural to discuss costs in terms of calories, as we might consider the cost of locomotion or cost of transport associated with foraging.

    But the only currency that matters to evolution is fitness. We cannot assume that maximizing caloric returns will maximize fitness. Transport and locomotor costs may be minor compared to the mortality risk from predation when foraging far from camp. The caloric benefits from hunting matter more to a starving child than to a satiated adult.

    So the measures of costs and benefits that define the ESS should be expressed in terms of fitness. That’s a problem, because fitness outcomes are a lot harder to measure than caloric returns. To figure out caloric expenditure and returns, you can measure oxygen consumption, work out distances, and weigh meat. To measure fitness, you have to record lifetime reproduction. To assess the relationship between caloric returns and fitness, you need a lifetime of caloric returns.

    So far, hunter-gatherer demographic data and hunting returns are both known from a small number of transverse studies. Longitudinal data on hunter-gatherer demography are limited, and mostly known by retrospective methods — that is, informants share their knowledge about the history of their groups. The fitness effects of a single individual’s hunting effort over time are not known.

    If fitness outcomes are hard for the scientist to measure, they are equally hard for a social actor to predict. Even intelligent actors like humans know little about the effects of their actions upon their future reproduction. Men sometimes do poorly with information directly relevant to fitness, like “Is the child mine?” That’s not to say that men may not follow highly sophisticated strategies to allocate hunting effort. But we should develop explanations that do not assume that a man knows the fitness benefits and costs of his choices.

    Next: Life history and asymmetrical strategies

  • Snowdrift games, cooperation, and "tragedy of the commune"

    Tue, 2009-06-02 23:27 -- John Hawks

    It’s the second day of June, which means it’s a good time to consider snowdrifts. OK, maybe not – but at least we’re far enough from winter now that the thought of snowdrifts out the window isn’t enough to give me a chill.

    The Snowdrift Game is a theoretical model of cooperation within the context of game theory. I gave a short introduction to game theory a couple of years ago, focusing on the games of Chicken and the Prisoner’s Dilemma. There are really only two formal varieties of two-player games involving cooperation or defection in the absence of information transfer. When defection is always the optimal strategy, it’s the Prisoner’s Dilemma. When a mixed strategy of cooperation and defection is optimal, it’s Chicken.

    But there are other names for this game. I’m not sure why, exactly—I suppose it’s because teenage boys in dragsters don’t appeal to everybody. One familiar name is the Hawk-Dove game. An individual can adopt two strategies: either attack and fight for a resource, or share equally and retreat when attacked. In the game, fighting carries a high cost (like wrecking your car into somebody) so a mixed strategy is optimal. When hawks are common, it’s better to be a dove and avoid fighting. When doves are common, it’s better to be a hawk because you always win.

    A third name for this game is Snowdrift. Imagine you’re riding in a car that becomes stuck in a snowdrift. You and a fellow passenger share the same interest: you both want the snowdrift to be removed. But who’s going to get out and shovel? It might seem fair just to get out and shovel the snow together—in other words, to cooperate. But what if the other passenger just sits there and refuses to help? If the cost of shoveling is low compared to the benefit of getting out of the drift, it will be in your interest to shovel by yourself. Sure, the other passenger is a freeloader who shares the benefit undeservedly, but so what? If the cost of shoveling was too high for you to bear, you’d have refused to do it, letting both of you freeze there. That would be the Prisoner’s Dilemma. But if the cost of shoveling is low compared to the costs of doing nothing, then a mixed strategy will be optimal. As long as freeloaders aren’t too common, that strategy will pay off. So a population engaged in the Snowdrift game will come to a mixed proportion of shovelers and freeloaders.

    Doebeli et al. (2004) considered the Snowdrift game as a model for the evolution of cooperation. A mixed strategy of cooperation and defection can emerge under a Snowdrift game system of payoffs, which makes it very different from the Prisoner’s Dilemma. Remember that in the Prisoner’s Dilemma, defection always generates a higher payoff than cooperation, regardless of the opponent’s strategy. So stable cooperation can only evolve under a Prisoner’s Dilemma system of payoffs if some kind of information transfer is possible. One example is the Iterated Prisoner’s Dilemma, in which two players encounter each other repeatedly. In this circumstance, one player can punish defection, leading to conditional strategies — the most famous of which is “tit for tat” — that yield a positive payoff for cooperation. It is worth pointing out that the cumulative payoffs under “tit for tat” or other conditional strategies come to approximate the payoffs of the Snowdrift game. The transfer of information changes one payoff structure into another.

    Here, we have unveiled a different paradox of cooperation, which could be termed the ”tragedy of the commune”: In a cooperative system, in which every individual contributes to a common good and benefits from its own investment, selection does not always generate the evolution of uniform and intermediate investment levels but may instead lead to an asymmetric stable state, in which some individuals make high levels of cooperative investment and others invest little or nothing.

    In practice, it is often difficult to determine the payoffs in social interactions and hence to distinguish prisoner’s dilemma and snowdrift interactions [a phage system marks a rare exception, but interestingly, selection turns the prisoner’s dilemma into a snowdrift game (24)]. Nevertheless, the mere existence of high- and low-investing individuals has often been taken as prima facie evidence that the interaction is governed by a prisoner’s dilemma, with some additional mechanism, such as reciprocity, responsible for the co-existence of altruists and nonaltruists. The tragedy of the commune, however, provides a quite different and, in many ways, simpler explanation for the coexistence of high- and low-investing individuals, which potentially applies to a wide range of cooperative and communal enterprises in biological systems (Doebeli et al. 2004:861–862).

    How is this relevant to paleoanthropology? The last paragraph of the paper suggests one way:

    In behavioral ecology, classical examples of cooperation include collective hunting and territory defense in lions (28), predator inspection in sticklebacks (29), and alarm calls in meerkats (30). In theoretical discussions of these examples, the existence of cooperators providing a common good and defectors exploiting it has been assumed a priori. The tragedy of the commune, however, suggests an evolutionary mechanism for the emergence of distinct behavioral patterns with differing degrees of provisions to the common good. This mechanism may also apply to cultural evolution in human societies, in which large differences in cooperative contributions to communal enterprises could give rise to conflicts on the basis of accepted notions of fairness (Doebeli et al. 2004:862).

    Food sharing in human hunter-gatherers includes many asymmetries. For example, hunters differ greatly in their hunting returns and expenditure of effort. Yet good hunters tolerate the presence of poor hunters and share food with them. As with hunting but extended to both men and women, people invest greatly varying degrees of effort into gathering plant foods, with resulting variation in caloric returns. Some of the variation in investment and success is age-related, some is likely directly environmentally induced, and some may reflect frequency-dependent strategies.

    Over the next few days, I’ll be considering human hunting from the perspective of the Snowdrift game. I’ll start with some very simple deterministic models and then try to make them a bit more relevant by considering the effects of stochastic payoffs and asymmetries among players.

    Next: Defining the Snowdrift game for hunting

    References:

       Doebeli M, Hauert C, Killingback T. 2004. The evolutionary origin of cooperators and defectors. Science 306:859–862. doi:10.1126/science.1101456.

  • Disease, pathogens, and collectivism

    Tue, 2008-04-15 12:03 -- John Hawks

    Sharon Begley in Newsweek reports on a hypothesis about "collectivism" and pathogens:

    The West epitomizes individualistic, do-your-own thing cultures, ones where the rights of the individual equal and often trump those of the group and where differences are valued. East Asian societies exalt the larger society: behavior is constrained by social roles, conformity is prized, outsiders shunned. "The individualist-collectivist split is one of the most powerful differences among cultures," says Nisbett. But the reason a society falls where it does on the individualism-collectivism spectrum has been pretty much a mystery. Now a team of researchers has come up with a surprising explanation: disease-causing microbes. Societies that evolved in places with an abundance of pathogens, they argue, had to adopt behaviors that add up to collectivism, for reasons of sheer preservation. Societies that arose in places with fewer pathogens had the luxury of individualism, which is less effective at limiting the spread of disease but brings with it other social benefits, such as innovation.

    There have been a lot of papers lately trying to match clines (that is, gradients) of phenotypes or genes with current ecological conditions. Climate is the most frequent (often, measured in terms of temperature or rainfall). Pathogens sort of follow the climate gradient, with some exceptions -- sometimes but not always allowing for the fact that malaria is the largest.

    But in some cases, much more important will be the historical dimension. Many, (but not all) people who live in high-pathogen areas today had ancestors who adopted agriculture relatively early, began living in denser concentrations and larger groups, and who therefore experienced in common a range of selective pressures that have nothing to do with pathogens.

    Would it be surprising that early agriculturalists living in emerging villages and cities might have been subject to pressures that enhanced collectivism? Such changes may have been facilitated by genetic changes, but would have also included cultural adaptations. Yet a correlation with pathogens would emerge as a side-effect of the history of agriculturalism, not as a direct cause.

    To my mind, these kinds of historical correlations rooted in ecological change will be a central problem of anthropological genomics. Recent human evolution has been dominated by a few very large changes -- like boulders thrown into a pond that have spread massive ripples through many elements of human genetics, anatomy, and behavior. These changes are not yet complete -- the ripples have not settled down nor reached the shore. For this reason, there will be many correlations that have been induced by the large ecological changes, making bivariate spatial comparisons a poor test of cause.

    I would say that historical correlation is a problem with a number of recent studies of cranial variation. Lately, it has been fashionable to test the hypothesis that cranial variation is adaptive to climate, by looking for spatial correlations between cranial measurements and climatic variables. But this test assumes that the correlations have not emerged as a result of some other cause. That might not be such a bad assumption, if humans had been static within their current geographic range for a long, long time. But humans have been anything but static -- in fact, their dynamism over the last 40,000 years has been the cause of profound changes in human biology. Naturally, things will be correlated with climate, because climate has been correlated with human subsistence and population size changes.

    A better test of adaptation on cranial variables would propose concrete mechanical (or developmental) reasons why a cranial trait helps someone to better survive under a given climatic regime. Finding a correlation in space is not enough. This is why Bergmann's and Allen's rules are more compelling than the more nebulous idea that "facial form" adapts people to their climate.

    Likewise in the case of pathogens -- a correlation between current pathogen load and current behavioral "collectivism" does not suggest a causal relationship between the two. Instead, we would want some kind of functional hypothesis to account for pathogens causing people to change in their attitudes toward cooperating. Concepts like the "luxury of individualism" make little sense. Of course, some people will prefer to behave with individual autonomy. But how would a recurrent epidemic disease stop them?

  • Cooperation, phenotypic vectors, energy

    Wed, 2006-04-26 08:53 -- John Hawks

    Burtsev and Turchin (2006) present the results of simulations of cooperative behavior in self-interested agents. This is a well-established subject, and their contribution is that their strategies are "evolved" from basic behavioral elements within their simulations, instead of being assumed a priori.

    In our model, agents are endowed with a limited set of receptors, a set of elementary actions and a neural net in between. Behavioural strategies are not predetermined; instead, the process of evolution constructs and reconstructs them from elementary actions. Two new strategies of cooperative attack and defence emerge in simulations, as well as the well-known dove, hawk and bourgeois strategies. Our results indicate that cooperative strategies can evolve even under such minimalist assumptions, provided that agents are capable of perceiving heritable external markers of other agents.

    This to me is one of the most interesting aspects of the model: behavioral traits gain random associations with recognizable phenotypes, and individuals shape their behavior according to the phenotypes that they detect around them.

    Each agent has external phenotype that is coded by a vector of integer values (markers). The markers do not influence behaviour but function only as indicators of similarity....All of our simulations were started with an initial population of agents that were unaware of markers (the matrix coefficients connecting input from markers to actions were preset to zero). Thus, the use of markers in a population had to evolve from a blank slate. Because markers and behaviours are not linked (apart from both being inherited from the ancestors), agents can lose cooperative behaviours by mutation while retaining 'in-group' markers. Thus, the structure of the model allows free-riders to arise.

    This "phenotypic association" vector is suggestive. Of course, for real animals it would probably be more effective to recognize the behaviors themselves as signs. But this depends on multiple opportunities -- you have to see somebody else's behavior at least once to judge it. If there were external manifestations associated with behaviors, it would give the opportunity to decide before an interaction what the other individual's likely strategy would be.

    But then, selection would favor mimicry -- free-riders with the phenotypes of cooperators, for example. This force will tend to limit the degree of association between observable traits and behaviors...

    except...

    That observable traits that indicate relatedness will also tend to indicate similarity in cooperator phenotypes. In other words, you can figure that your relatives will tend to act like you, and also tend to look like you. And as a bonus, if you are sharing with a relative, you are increasing your inclusive fitness.

    They find that the evolution of different strategies depends on carrying capacity, and some new strategies emerged. The comparison of these is worth reading, but a bit too long and involved to quote at length. This part is important:

    Our results have important implications for the evolution of territoriality in animals (and private property in humans). With a few exceptions, theorists have paid little attention to the role that cooperation may have in the evolution of territoriality. Our study suggests that cooperative defence of territory can radically change the course of evolution in resource-rich (C > C2) environments. When the amount of resource becomes large enough to support more than one agent, and too large for a single agent to monopolize, solitary bourgeois are replaced by cooperative starlings, provided that agents can recognize in-group members. The starling strategy does not take over completely, however, but coexists with other strategies in a complex dynamical way.

    The "starling" strategy is a mobbing strategy, in which small animals cooperate to attack and drive away large solitary predators.

    One limitation of these kinds of simulations is that they don't include reproductive boundaries. For this reason, they don't really distinguish models of within-species cooperation from between-species mutualism. Different strategies like "hawk" and "dove" might really represent predators and prey species, or they might represent contrasts of competitive behavior within a species.

    So the appearance of stable strategies at any given level of possible complexity might be a constraint on natural communities, but the level of that constraint may not be immediately obvious. To that end, this study has a very large possible set of strategies (more than 101000 combinations), which means it is sampling a richer set of behaviors than most simple game-theoretic models.

    References:

    Burtsev M, Turchin P. 2006. Evolution of cooperative strategies from first principles. Nature 440:1041-1044. DOI link

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