Back to the testicles of sparrows: In a breeding area each little piece of territory has a male sparrow who will defend it against all comers.* Suppose a meddling ornithologist captures one of these territorial males and removes him from the territory. What happens? Other males from adjacent areas—many of them not previously able to defend a territory—move in. Of course they have to threaten and intimidate before they’ll be taken seriously. So the general level of sparrow anxiety rises, both among the newcomers and among unreplaced sparrows in adjacent territories. Political tensions become high. If now we monitor the bloodstreams of the sparrows in the course of their disputes (which from our point of view, of course, seem petty, but to them it’s Quemoy and Matsu), we find that everyone’s testosterone level has risen—the newly introduced males who are trying to establish their territories, and the males of neighboring territories who are now required to do more in the way of defending than has been their recent practice. Something similar is true for many animals.
Those who have more testosterone, by and large, become more aggressive. Those who need more testosterone, by and large, generate it. Testosterone seems to play a vital role as both the cause and the effect of aggression, territoriality, dominance, and the rest of the “boys-will-be-boys” constellation of male behavioral traits. This seems to be true for widely differing species, including monkeys, apes, and humans.
In springtime, stimulated by the increasing day length, the testosterone level in male perching birds and songbirds (such as jays, warblers, and sparrows) goes up; they develop plumage, unveil a scrappy temperament, and begin singing. Males with larger repertoires breed earlier and produce more chicks. The repertoires of the most attractive males range up to dozens of distinguishable songs. Musical variety is the means by which more testosterone is converted into more birds.
When eggs are being laid, the male testosterone level remains high; they’re protecting their mates. Once the females begin incubating the eggs and are uninterested in sexual advances, male testosterone levels fall. Suppose that the females are now given estrogen implants so they remain sexually alluring and receptive, despite their new maternal duties. Then the testosterone levels in the males remain high. As long as the female is sexually available, the male is inclined to be nearby and protective.17
These experiments suggest that an important selective advantage may accrue if a species breaks out of the estrus constraint. Continuous female sexual receptivity keeps the male around for all sorts of useful services. This is just what seems to have happened—maybe through a small adjustment in the DNA code for the internal estrogen clock—in our species.
Testosterone-induced behavior must be subject to limits and constraints. If it were carried to counterproductive lengths, natural selection would quickly readjust the concentration of steroids in the blood. Testosterone poisoning to the point of maladaptation must be very rare. In nectar-eating birds, bats, and insects it’s possible to compare the energy expended in male steroid-driven defense against poachers with the energy that could be extracted from the flowers being guarded.* In fact, territoriality typically turns on only when the energy benefit exceeds the energy cost, only when there are so few delectable flowers to suck that it pays for you to expend the effort to chase away the competition. Nectar-eaters are not rigid territorialists. They won’t fight all comers to protect a wasteland of stones. They make a cost-benefit analysis. Even in a rich garden of nectar-bearing flowers, often no territorial behavior is seen in the morning—because plentiful nectar has been accumulating at night when the birds were asleep. In the morning, there’s enough to go around. Toward noon, when birds from far and wide have been feeding and the resource begins to get scarce, territoriality turns on.18
Wings outstretched, beaks lunging, the locals drive away the intruders. Maybe they feel they’ve been nice guys long enough, but now they’ve had it up to——
Many animals may do it, but at least among rats and mice it’s well-demonstrated: Fear is accompanied by a characteristic odor, a fear pheromone, easily recognized by others.19
Often, as soon as they sense you’re afraid, your friends and relatives run away—useful for them, but not very helpful for you. It may even encourage the rival or predator who has prompted your fear in the first place.