seep across a microscopic gap, and lock onto a receptor in the surface of another neuron, changing its activity and thereby allowing patterns of neural firing to propagate through the brain. One major class of neurotransmitters are the catecholamines, which include dopamine, serotonin, and norepinephrine (also called noradrenaline, and related to the adrenaline that triggers the fight-or-flight response). The catecholamines are used in several motivational and emotional systems of the brain, and their concentration is regulated by proteins that break them down or recycle them. One of those enzymes is monoamine oxidase-A, MAO-A for short, which helps to break down these neurotransmitters, preventing them from building up in the brain. When they do build up, the organism can become hyperreactive to threats and more likely to engage in aggression.
The first sign that MAO-A can affect violence in humans was the discovery of a Dutch family that carried a rare mutation that left half the men without a working version of the gene.152 (The gene is found on the X chromosome, which men have only one of, so if a man’s MAO-A gene is defective, he has no backup copy to compensate.) Over at least five generations, the affected men in the family were prone to aggressive outbursts. One, for example, forced his sisters to disrobe at knifepoint; another tried to run over his boss with his car.
A more common kind of variation is found in the part of the gene that determines how much MAO-A is produced. People with a low-activity version of the gene build up higher levels of dopamine, serotonin, and norepinephrine in the brain. They also are more likely to have symptoms of antisocial personality disorder, to report that they have committed acts of violence, to be convicted of a violent crime, to have amygdalas that react more strongly and an orbital cortex that reacts less strongly to angry and fearful faces, and, in the psychology lab, to force a fellow participant to drink hot sauce if they think he has exploited them.153 Unlike many other genes that affect behavior, the low-activity version of the MAO-A gene seems to be fairly specific to aggression; it does not correlate well with any other personality trait.154
The low-activity version of the MAO-A gene makes people more prone to aggression primarily when they have grown up with stressful experiences, such as having been abused or neglected by their parents or having been held back in school.155 It’s hard to pinpoint the exact stressors that have this effect, because stressful lives are often stressful in many ways at once. In fact, the modulating factor may consist of other genes which are shared with an abusive parent, predisposing both parent and child to aggression, and which may also elicit negative reactions from the people around him.156 But whatever the modifying factor is, it does not turn the effects of the low-activity version of the gene upside down. In all the studies, the gene has an aggregate or main effect in the population that could make it a target of selection. Indeed, Moffitt and Caspi (who first discovered that the effect of the gene depends on stressful experiences) suggest that rather than thinking of the low-activity version of the gene as a contributor to violence, we should think of the high-activity version as an inhibitor of violence: it protects people from overreacting to a stressfilled life. Geneticists have discovered statistical evidence of selection for the MAO-A gene in humans, though the evidence does not single out the low- or high-activity variant; nor does it prove that the gene was selected for its effects on aggression.157
Other genes that affect dopamine have been associated with delinquency as well, including a version of a gene that affects the density of dopamine receptors (DRD2) and a version of a gene for a dopamine transporter (DAT1) that mops up excess dopamine from the synapse and transports it back into the neurons that release it.158 Any of them could be fair game for rapid natural selection.
Genetic tendencies toward or away from violence, then, could have been selected during the historical transitions we have examined. The question is, were they? The mere existence of pathways to evolutionary change does not prove that those pathways were taken. Evolution depends not just on the genetic raw material but on factors such as the demography of a population (including both sheer numbers and the degree to which they have absorbed immigrants from other groups), the roll of the genetic and environmental dice, and the