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Emotions are hardwired by genes we share and presumably share with other primates and indeed other mammals, as Darwin himself noticed. In us, of course, they get harnessed together with our highly developed theory-of-mind ability and with norms adaptive in our environments. They motivate enforcement of the norms they get paired up with, on others and on ourselves. Some of these norms solve design problems common to humans in all the environments we inhabit. These are parts of the moral core we all share. Others will not be part of core morality but will be locally restricted to the different ecologies that different groups inhabit. Some examples will illustrate how this works.

Anthropologists have uncovered how different ecologies select for differences in local moral norms. For example, there will be quite different norms in pastoral versus agricultural cultures. Shepherds and herders need to solve the problem of asserting ownership over flocks that may spread far beyond their immediate supervision. Among pastoral people, the norm of going to any lengths to punish rustling will be highly adaptive. So there will be strong selection for emotions of anger, revenge, disdain that accompany the norms prohibiting theft of stock animals. Where norms against rustling are not linked with these emotions, the motivation to act on them is reduced. And that lowers the fitness of the pastoralists who endorse the norms but don’t have the right emotions. Again, the process is the same as the one that linked sex and pleasure. Nature was selecting among variations for fitness. Of course, in the emotion/norm combination, the linkage is culturally established and transmitted.

By contrast with pastoralists, sedentary farmers can keep an eye on their plows, even when others borrow them without asking. Among cultivators, there will be selection for norms that allow a certain amount of informal borrowing without permission. The emotions that go along with their norms will be different or weaker than among sheep and cattle herders.

The differences in which behaviors are fitness enhancing dictate which norms persist. Variation and selection harness preexisting emotions to enforce them. Not much genetic hardwiring is required for the Darwinian mechanism that finds and enforces moral norms, sometimes globally and sometimes locally. The emotions are bodily feelings and they are hardwired, subject to the usual Darwinian range of variation. But which norms they get harnessed to depends on what is fitness enhancing in an environment. Different environments make different packages of norm and emotion fitness enhancing. Some packages are fitness enhancing in all the environments we have occupied. These are the ones that form our moral core. They are the ones packaged with the same emotions universally, or nearly so. All societies and cultures have words for anger, disdain, shame, and guilt, and in each, they name roughly the same set of bodily feelings. In each culture, they are linked to common norms—the ones in core morality. They are also linked to different norms—ones that reflect local culture and local environmental filtration.

If you think about how, for example, shame and guilt work, you will see that they are emotions practically designed to solve nature’s problem of getting us to do the right thing, to be nice, at least for the sake of our genes’ futures.

In brief, shame is the unpleasant feeling we get when others catch us not abiding by the norms almost everyone (including us) embraces. Guilt is the unpleasant feeling that overcomes us when we aren’t caught. They are both strong enough by themselves to solve the problem nature created for itself when it made us prone to immediate gratification and resistant to long-term benefits.

Long ago in human cultural evolution, parents began drumming tit-for-tat norms into kids already prone to niceness. Shame at being caught free-riding on others’ niceness and shame at being taken for a sucker by others’ free-riding is easy