Gain control in the sonar of odontocetes

The site and physiologic mechanism(s) responsible for the generation of odontocete biosonar signals have eluded investigators for decades. To address these issues we subjected postmortem toothed whale heads to interrogation using medical imaging techniques. Most of the 40 specimens (from 19 species) were examined using x-ray computed tomography (CT) and/or magnetic resonance imaging (MR). Interpretation of scan images was aided by subsequent dissection of the specimens or, in one case, by cryosectioning. In all specimens we described a similar tissue complex and identified it as the hypothetical biosonar signal generator. This complex includes a small pair of fatty bursae embedded in a pair of connective tissue lips, a cartilaginous blade, a stout ligament, and an array of soft tissue air sacs. Comparing and contrasting the morphologic patterns of nasal structures across species representing every extant odontocete superfamily reveals probable homologous relationships, which suggests that all toothed whales may be making their biosonar signals by a similar mechanism.

In bats and technological sonars, the gain of the receiver is progressively increased with time after the transmission of a signal to compensate for acoustic propagation loss. The current understanding of dolphin echolocation indicates that automatic gain control is not a part of their sonar system. In order to test this understanding, we have performed field measurements of free-ranging echolocating dolphins. Here we show that dolphins do possess an automatic gain control mechanism, but that it is implemented in the transmission phase rather than the receiving phase of a sonar cycle. We find that the amplitude of the dolphins' echolocation signals are highly range dependent; this amplitude increases with increasing target range, R, in a 20 log(R) fashion to compensate for propagation loss. If the echolocation target is a fish school with many sound scatterers, the echoes from the school will remain nearly constant with range as the dolphin closes in on it. This characteristic has the same effect as time-varying gain in bats and technological sonar when considered from a sonar system perspective.

The echolocation transmission beam pattern of a false killer whale (Pseudorca crassidens) was measured in the vertical and horizontal planes. A vertical array of seven broadband miniature hydrophones was used to measure the beam pattern in the vertical plane and a horizontal array of the same hydrophones was used in the horizontal plane. The measurements were performed in the open waters of Kaneohe Bay, Oahu, Hawaii, while the whale performed a target discrimination task. Four types of signals, characterized by their frequency spectra, were measured. Type-1 signals had a single low-frequency peak at 40 +/- 9 kHz and a low-amplitude shoulder at high frequencies. Type-2 signals had a bimodal frequency characteristic with a primary peak at 46 +/- 7 kHz and a secondary peak at 88 +/- 13 kHz. Type-3 signals were also bimodal but with a primary peak at 100 +/- 7 kHz and a secondary peak at 49 +/- 9 kHz. Type-4 signals had a single high-frequency peak at 104 +/- 7 kHz. The center frequency of the signals were found to be linearly correlated to the peak-to-peak source level, increasing with increasing source level. The major axis of the vertical beam was directed slightly downward between 0 and -5 degrees, in contrast to the +5 to 10 degrees for Tursiops and Delphinapterus. The beam in the horizontal plane was directed forward between 0 degrees and -5 degrees.(ABSTRACT TRUNCATED AT 250 WORDS)