Given a range of possible song models during isolated rearing, a chick

selects an example from its own species and memorizes it. If it hears only
songs of other species, the mature song is the unmodified innate song (Fig.
49.1). Thus there is an innate bias in the initial learning. Where this bias has
been studied, it appears to depend on acoustic sign stimuli (i.e., species-
specific “syllables”). Chicks are able to extract syllables of their species
embedded in foreign songs, or scored in a way never found in their species
(e.g., a syllable that is repeated at an accelerating rate presented to a species
that sings syllables at a constant rate, or vice versa will be extracted and
used in the species-typical manner).
Practice is essential in the normal development of birdsong, and part of
this practice occurs in a babbling phase known as subsong, which begins at
a species-typical age. A bird deafened after its sensitive period, but before it
begins producing notes in preparation for singing (subsong), is unable to
produce even an innate song (Fig. 49.1). During the earliest parts of
subsong, birds try out a number of notes. These notes are typical of the
species, but most are absent from the song they eventually sing. The
learning process may involve producing each member of an innate
repertoire of notes, listening to them, checking to see if they match any
element in the memorized song, discarding the unnecessary ones, and
rearranging, scoring, and modifying the others to produce a reasonable copy
of the original song heard during the sensitive period.
There is some flexibility in song development. For example, when the
chick has heard two very different specimens of its own species’ song, it
will often incorporate elements from each. Similarly, when the chick has
been exposed to the sight of a singing conspecific and simultaneously the
sound of a heterospecific song, it may pick out elements of the abnormal
song and adapt them as best it can into its own species-specific
organization.
Birdsong, therefore, depends on two processes that involve an
interaction between innate capacities and learning: imprinting the song in
memory and then learning to perform it. This system is flexible, but only
within clear limits.

Communication Systems in Primates