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Biomedical subjects

P T Madsen

Publications and source records attributed to P T Madsen.

10 recordsLinked to original sources

Quantitative measures of air-gun pulses recorded on sperm whales (Physeter macrocephalus) using acoustic tags during controlled exposure experiments.

The widespread use of powerful, low-frequency air-gun pulses for seismic seabed exploration has raised concern about their potential negative effects on marine wildlife. Here, we quantify the sound exposure levels recorded on acoustic tags attached to eight sperm whales at ranges between 1.4 and 12.6 km from controlled air-gun array sources operated in the Gulf of Mexico. Due to multipath propagation, the animals were exposed to multiple sound pulses during each firing of the array with received levels of analyzed pulses falling between 131-167 dB re. 1 microPa (pp) [111-147 dB re. 1 microPa (rms) and 100-135 dB re. 1 microPa2 s] after compensation for hearing sensitivity using the M-weighting. Received levels varied widely with range and depth of the exposed animal precluding reliable estimation of exposure zones based on simple geometric spreading laws. When whales were close to the surface, the first arrivals of air-gun pulses contained most energy between 0.3 and 3 kHz, a frequency range well beyond the normal frequencies of interest in seismic exploration. Therefore air-gun arrays can generate significant sound energy at frequencies many octaves higher than the frequencies of interest for seismic exploration, which increases concern of the potential impact on odontocetes with poor low frequency hearing.

Acoustic Stimulation↗

Foraging Blainville's beaked whales (Mesoplodon densirostris) produce distinct click types matched to different phases of echolocation.

Blainville's beaked whales (Mesoplodon densirostris Blainville) echolocate for prey during deep foraging dives. Here we use acoustic tags to demonstrate that these whales, in contrast to other toothed whales studied, produce two distinct types of click sounds during different phases in biosonar-based foraging. Search clicks are emitted during foraging dives with inter-click intervals typically between 0.2 and 0.4 s. They have the distinctive form of an FM upsweep (modulation rate of about 110 kHz ms(-1)) with a -10 dB bandwidth from 26 to 51 kHz and a pulse length of 270 micros, somewhat similar to chirp signals in bats and Cuvier's beaked whales (Ziphius cavirostris Cuvier), but quite different from clicks of other toothed whales studied. In comparison, the buzz clicks, produced in short bursts during the final stage of prey capture, are short (105 micros) transients with no FM structure and a -10 dB bandwidth from 25 to 80 kHz or higher. Buzz clicks have properties similar to clicks reported from large delphinids and hold the potential for higher temporal resolution than the FM clicks. It is suggested that the two click types are adapted to the separate problems of target detection and classification versus capture of low target strength prey in a cluttered acoustic environment.

Animals↗

Marine mammals and noise: problems with root mean square sound pressure levels for transients.

Current mitigation levels for noise transients impinging on marine mammals are specified by rms pressures. The rms measure critically relies upon choosing the size of averaging window for the squared pressures. Derivation of this window is not standardized, which can lead to 2-12 dB differences in rms sound pressure for the same wave forms. rms pressure does not represent the energy of the noise pulse and it does not prevent exposure to high peak pressures. Safety levels for transients should therefore be given by received peak-peak sound pressure and energy flux density instead of rms sound pressure levels.

Journal Article↗

Biosonar performance of foraging beaked whales (Mesoplodon densirostris).

Toothed whales (Cetacea, odontoceti) emit sound pulses to probe their surroundings by active echolocation. Non-invasive, acoustic Dtags were placed on deep-diving Blainville's beaked whales (Mesoplodon densirostris) to record their ultrasonic clicks and the returning echoes from prey items, providing a unique view on how a whale operates its biosonar during foraging in the wild. The process of echolocation during prey capture in this species can be divided into search, approach and terminal phases, as in echolocating bats. The approach phase, defined by the onset of detectable echoes recorded on the tag for click sequences terminated by a buzz, has interclick intervals (ICI) of 300-400 ms. These ICIs are more than a magnitude longer than the decreasing two-way travel time to the targets, showing that ICIs are not given by the two-way-travel times plus a fixed, short lag time. During the approach phase, the received echo energy increases by 10.4(+/-2) dB when the target range is halved, demonstrating that the whales do not employ range-compensating gain control of the transmitter, as has been implicated for some bats and dolphins. The terminal/buzz phase with ICIs of around 10 ms is initiated when one or more targets are within approximately a body length of the whale (2-5 m), so that strong echo returns in the approach phase are traded for rapid updates in the terminal phase. It is suggested that stable ICIs in the search and approach phases facilitate auditory scene analysis in a complex multi-target environment, and that a concomitant low click rate allows the whales to maintain high sound pressure outputs for prey detection and discrimination with a pneumatically driven, bi-modal sound generator.

Acoustics↗

Echolocation clicks of two free-ranging, oceanic delphinids with different food preferences: false killer whales Pseudorca crassidens and Risso's dolphins Grampus griseus.

Toothed whales (Odontoceti, Cetacea) navigate and locate prey by means of active echolocation. Studies on captive animals have accumulated a large body of knowledge concerning the production, reception and processing of sound in odontocete biosonars, but there is little information about the properties and use of biosonar clicks of free-ranging animals in offshore habitats. This study presents the first source parameter estimates of biosonar clicks from two free-ranging oceanic delphinids, the opportunistically foraging Pseudorca crassidens and the cephalopod eating Grampus griseus. Pseudorca produces short duration (30 micro s), broadband (Q=2-3) signals with peak frequencies around 40 kHz, centroid frequencies of 30-70 kHz, and source levels between 201-225 dB re. 1 micro Pa (peak to peak, pp). Grampus also produces short (40 micro s), broadband (Q=2-3) signals with peak frequencies around 50 kHz, centroid frequencies of 60-90 kHz, and source levels between 202 and 222 dB re. 1 micro Pa (pp). On-axis clicks from both species had centroid frequencies in the frequency range of most sensitive hearing, and lower peak frequencies and higher source levels than reported from captive animals. It is demonstrated that sound production in these two free-ranging echolocators is dynamic, and that free-ranging animals may not always employ biosonar signals comparable to the extreme signal properties reported from captive animals in long-range detection tasks. Similarities in source parameters suggest that evolutionary factors other than prey type determine the properties of biosonar signals of the two species. Modelling shows that interspecific detection ranges of prey types differ from 80 to 300 m for Grampus and Pseudorca, respectively.

Animal Communication↗

Sperm whale sound production studied with ultrasound time/depth-recording tags.

Delphinoids (Delphinidae, Odontoceti) produce tonal sounds and clicks by forcing pressurized air past phonic lips in the nasal complex. It has been proposed that homologous, hypertrophied nasal structures in the deep-diving sperm whale (Physeter macrocephalus) (Physeteridae, Odontoceti) are dedicated to the production of clicks. However, air volumes in diving mammals are reduced with increasing ambient pressure, which seems likely to influence pneumatic sound production at depth. To study sperm whale sound production at depth, we attached ultrasound time/depth-recording tags to sperm whales by means of a pole and suction cup. We demonstrate that sperm whale click production in terms of output and frequency content is unaffected by hydrostatic reduction in available air volume down to less than 2% of the initial air volume in the nasal complex. We present evidence suggesting that the sound-generating mechanism has a bimodal function, allowing for the production of clicks suited for biosonar and clicks more suited for communication. Shared click features suggest that sound production in sperm whales is based on the same fundamental biomechanics as in smaller odontocetes and that the nasal complexes are therefore not only anatomically but also functionally homologous in generating the initial sound pulse.

Animals↗

Sperm whale clicks: directionality and source level revisited.

In sperm whales (Physeter catodon L. 1758) the nose is vastly hypertrophied, accounting for about one-third of the length or weight of an adult male. Norris and Harvey [in Animal Orientation and Navigation, NASA SP-262 (1972), pp. 397-417] ascribed a sound-generating function to this organ complex. A sound generator weighing upward of 10 tons and with a cross-section of 1 m is expected to generate high-intensity, directional sounds. This prediction from the Norris and Harvey theory is not supported by published data for sperm whale clicks (source levels of 180 dB re 1 microPa and little, if any, directionality). Either the theory is not borne out or the data is not representative for the capabilities of the sound-generating mechanism. To increase the amount of relevant data, a five-hydrophone array, suspended from three platforms separated by 1 km and linked by radio, was deployed at the slope of the continental shelf off Andenes, Norway, in the summers of 1997 and 1998. With this system, source levels up to 223 dB re 1 microPa peRMS were recorded. Also, source level differences of 35 dB for the same click at different directions were seen, which are interpreted as evidence for high directionality. This implicates sonar as a possible function of the clicks. Thus, previously published properties of sperm whale clicks underestimate the capabilities of the sound generator and therefore cannot falsify the Norris and Harvey theory.

Animal Communication↗

Sperm whales (Physeter catodon L. 1758) do not react to sounds from detonators.

A number of observations show that sperm whales (Physeter catodon L. 1758) react to various man-made pulses with moderate source levels. The behavioral responses are described to vary from silence to fear. Click rates of five submerged male sperm whales were measured during the discharge of eight detonators off Andenes, northern Norway. In addition, the behavioral response of a surfaced specimen was observed. Click rates of the submerged whales and the behavior of the surfaced specimen did not change during the discharges with received sound levels of some 180 dB re 1 microPa peRMS. The apparent lack of response to the discharges could be due to similarity between sperm whale clicks and detonations. Accordingly, it can be speculated that the discharges may have been perceived as isolated clicks from conspecifics.

Animals↗