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indignation at cheating, and methods to detect cheating. Many common emotions can be understood as being built around the expectation of reciprocity and the negative reaction when it is made to fail. If we like a person, we are willing to exchange favors with them. We are angry at those who fail to return favors. We seek punishment for those who take advantage of us. We feel guilty if we fail to return a favor, and shame if publicly exposed. If we believe someone is genuinely sorry about a failure to reciprocate, we trust them. But if we detect they are simulating contrition, we mistrust them.194 The instinct for reciprocity, and the cheater-detection apparatus that accompanies it, seem to be the basis for a fundamental human practice, that of trade with neighboring groups. Long distance trade is one of the characteristic behaviors of the human societies that emerged in the Upper Paleolithic age starting some 50,000 years ago. Tribal societies developed trading systems of considerable sophistication. The Yit Yoront, a foraging society of northern Australia, lived until recently in the Stone Age. One of their most necessary possessions, used in everything from hunting to wood-gathering, were hafted stone axes. But they lived on an alluvial coast and the nearest stone quarry was four hundred miles away. How did they acquire their polished stone axes? They made a product much in demand with their neighbors to the south, spears tipped with the barbs of stingrays. The spears were traded inland, through a long line of trading partners, being exchanged at each stage for a varying number of stone axes. The spear/axe exchange rate was sufficient at each trading post to push stone axes northward and pull barb-tipped spears southward.195

Trade is a foundation of economic activity because it gives the parties to a transaction a strong incentive to specialize in making the items that the other finds valuable. But trade depends on trust, on the decision to treat a total stranger as if he were a member of the family. Humans are the only species to have developed such a degree of social trust that they are willing to let vital tasks be performed by individuals who are not part of the family. This set of behaviors, built around reciprocity, fair exchange and the detection of cheaters, has provided the foundation for the most sophisticated urban civilizations, including those of the present day.

Reciprocity, and an ability to calculate the costs and benefits of cooperation, underpin our social life, writes the economist Paul Seabright, "making it reasonable for us to treat strangers as though they were honorary relatives or friends." It is remarkable that this behavior evolved at a time when primitive warfare was at its most intense and people had every reason to regard strangers with deep suspicion. Strangers can still be dangerous, yet in the right circumstances we habitually trust them. "The knowledge that most people can be trusted much of the time to play their part in the complex web of social cooperation has had dramatic effects on the psychology of our everyday life," Seabright says, making it possible "to step nonchalantly out of the front door of a suburban house and disappear into a city of ten million strangers."196 Without this innate willingness to trust strangers, human societies would still consist of family units a few score strong, and cities and great economies would have had no foundation for existence.

How might this greater level of trust have arisen? Two hormones, known as oxytocin and vasopressin, are emerging as central players in modulating certain social behaviors in the mammalian brain. The hormones are generated in the pituitary gland at the base of the brain and have effects both on the body and in the brain. Oxytocin induces both labor in childbirth and the production of milk. Its effects on the mind, at least in experimental animals, have the general property of promoting affiliative or trusting behavior, lowering the natural resistance that animals have to the close proximity of others. So what does oxytocin do in people? Researchers at the University of Zurich have found that it substantially increases the level of trust. Oxytocin, they say, "specifically affects an individual's willingness to accept social risks arising through interpersonal interactions." The findings emerged from giving subjects a sniff of oxytocin before playing a game that tested 197

trusting behavior.

If the biological basis of trusting behavior is mediated in this manner, the degree of trust could easily be ratcheted up or down in the course of human evolution by genetic changes that either increased individuals' natural production of the hormone or enhanced the brain's response to it. Thus hunter-gatherers might have a genetically lower response to oxytocin while city-dwellers would have evolved a greater sensitivity. Whatever the exact mechanism, it is easy to see how greater levels of trust might have evolved at various stages in human evolution, given that there is a biological basis for the behavior. Trust is an essential part of the social glue that binds people together in cooperative associations. But it increases the vulnerability to which all social groups are exposed, that of being taken advantage of by freeloaders. Freeloaders seize the benefits of social living without contributing to the costs. They are immensely threatening to a social group because they diminish the benefits of sociality for others and, if their behavior goes unpunished, they may bring about the society's dissolution.

Human societies long ago devised an antidote to the freeloader problem. This freeloader defense system, a major organizing principle of every society, has assumed so many other duties that its original role has been lost sight of. It is religion.

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