âAn action, which we ourselves should require experience to enable us to perform, when performed by an animal, more especially by a very young one, without any experience, and when performed by many individuals in the same way, without their knowing for what purpose it is performed, is usually said to be instinctive.â â Charles Darwin
If complex physical structures can evolve, what about complex behaviors? Darwin tackles instinctsâthe inherited behavioral patterns that animals perform without learning. Can behaviors as intricate as the honeybeeâs hexagonal comb or the cuckooâs parasitism be explained by natural selection? Darwin argues yes, and provides some of his most fascinating examples.
Darwin acknowledges the difficulty of defining instinct precisely, but focuses on its key characteristic: instinctive behaviors are inherited, not learned.
An inherited behavioral pattern that animals perform correctly the first time, without instruction or practice. Instincts can be modified by experience but donât depend on it. They are to behavior what complex organs are to anatomy.
The first step is showing that instincts, like physical structures, can vary:
If instincts can vary, and if variations are heritable, then natural selection can act on behavior just as it acts on anatomy.
The European cuckoo lays its eggs in the nests of other birds, leaving foster parents to raise its young. This remarkable behavior troubled naturalistsâhow could such a complex parasitic strategy evolve?
Darwin notes that some birds occasionally lay eggs in othersâ nests. If this accidentally proved advantageousâif foster-raised chicks survived wellâthe behavior could be selected for. Each component of the cuckooâs instincts could evolve incrementally:
Some ant species raid the nests of other species, capturing pupae and raising them as workers. Darwin studied this extensively:
The gradation shows how an extreme instinct could evolve. Early stages involve occasional slave-taking; later stages show increasing dependence until the master species can no longer survive without slaves.
Darwin considered the hexagonal cells of honeybee combs the most wonderful instinct of allâand potentially the most difficult to explain:
Honeybee cells are mathematically perfect hexagonal prisms with pyramidal basesâthe shape that encloses maximum volume with minimum wax. How could âblindâ instinct produce what seems to require advanced geometry?
Darwinâs explanation is a masterpiece of gradualist reasoning:
Darwin shows gradation among bee species:
Darwin showed that if bees simply excavate spherical cells at equal distances, the intersecting walls automatically form the optimal hexagonal pattern. No geometric âknowledgeâ is neededâjust consistent spacing and wall-thinning behavior. The mathematics is a byproduct, not a goal.
Darwin addresses key objections:
How can natural selection act on worker ants and bees, which donât reproduce? Their instincts canât be passed to offspring because they have no offspring.
Selection acts on the reproductive members of the colony. If producing sterile workers with particular instincts benefits the colonyâs reproductive success, those genes spread. The unit of selection is the family or colony, not the individual worker. Darwin anticipates the concept of kin selection developed fully by Hamilton a century later.