Information theory analysis of patterns of modulation in the advertisement call of the male bullfrog, Rana catesbeiana.

Male bullfrogs often amplitude modulate the envelopes of the individual notes (croaks) in their multinote advertisement calls. These amplitude modulations change the envelope of the note from smooth and unmodulated to one with varying numbers of modulations. A Markov analysis shows the pattern of change in the envelope to be highly ordered, but not completely so (semi-Markovian). Three simple rules govern the presence or absence of modulations in individual notes. These rules are (1) all calls begin with an unmodulated note; (2) the first note to be modulated will contain only one modulation; and (3) when a change in modulation occurs from one note to the next, it does so with an increase or a decrease of one modulation only. The addition of modulations is correlated with an increase in note duration. Physiologically, the presence of modulations might increase the precision of temporal coding of note periodicities in the central auditory system.