RESUMO
The acoustic startle reflex is an oligo-synaptic reflex arc elicited by rapid-onset sounds. Odontocetes evolved a range of specific auditory adaptations to aquatic hearing and echolocation, e.g. the ability to downregulate their auditory sensitivity when emitting clicks. However, it remains unclear whether these adaptations also led to changes of the startle reflex. We investigated reactions to startling sounds in two bottlenose dolphins (Tursiops truncatus) and one false killer whale (Pseudorca crassidens). Animals were exposed to 50â ms, 1/3 octave band noise pulses of varying levels at frequencies of 1, 10, 25 and 32â kHz while positioned in a hoop station. Startle responses were quantified by measuring rapid muscle contractions using a three-dimensional accelerometer attached to the dolphin. Startle magnitude increased exponentially with increasing received levels. Startle thresholds were frequency dependent and ranged from 131â dB at 32â kHz to 153â dB at 1â kHz (re. 1â µPa). Startle thresholds only exceeded masked auditory AEP thresholds of the animals by 47â dB but were â¼82â dB above published behavioural audiograms for these species. We also tested the effect of stimulus rise time on startle magnitude using a broadband noise pulse. Startle responses decreased with increasing rise times from 2 to 100â ms. Models suggested that rise times of 141-220â ms were necessary to completely mitigate startle responses. Our data showed that the startle reflex is conserved in odontocetes and follows similar principles as in terrestrial mammals. These principles should be considered when assessing and mitigating the effects of anthropogenic noise on marine mammals.
Assuntos
Limiar Auditivo , Golfinhos/fisiologia , Reflexo de Sobressalto/fisiologia , Estimulação Acústica/veterinária , Animais , Golfinho Nariz-de-Garrafa/fisiologia , Ecolocação , Feminino , Havaí , MasculinoRESUMO
Toothed whales possess a sophisticated biosonar system by which ultrasonic clicks are projected in a highly directional transmission beam. Beam directivity is an important biosonar characteristic that reduces acoustic clutter and increases the acoustic detection range. This study measured click characteristics and the transmission beam pattern from a small odontocete, the spinner dolphin (Stenella longirostis). A formerly stranded individual was rehabilitated and trained to station underwater in front of a 16-element hydrophone array. On-axis clicks showed a mean duration of 20.1 µs, with mean peak and centroid frequencies of 58 and 64 kHz [standard deviation (s.d.) ±30 and ±12 kHz], respectively. Clicks were projected in an oval, vertically compressed beam, with mean vertical and horizontal beamwidths of 14.5° (s.d. ± 3.9) and 16.3° (s.d. ± 4.6), respectively. Directivity indices ranged from 14.9 to 27.4 dB, with a mean of 21.7 dB, although this likely represents a broader beam than what is normally produced by wild individuals. A click subset with characteristics more similar to those described for wild individuals exhibited a mean directivity index of 23.3 dB. Although one of the broadest transmission beams described for a dolphin, it is similar to other small bodied odontocetes.
Assuntos
Ecolocação/fisiologia , Audição/fisiologia , Ultrassom , Vocalização Animal/fisiologia , Acústica , Animais , Golfinhos , Espectrografia do Som/métodos , StenellaRESUMO
Odontocete marine mammals explore the environment by rapidly producing echolocation signals and receiving the corresponding echoes, which likewise return at very rapid rates. Thus, it is important that the auditory system has a high temporal resolution to effectively process and extract relevant information from click echoes. This study used auditory evoked potential methods to investigate auditory temporal resolution of individuals from four different odontocete species, including a spinner dolphin (Stenella longirostris), pygmy killer whale (Feresa attenuata), long-finned pilot whale (Globicephala melas), and Blainville's beaked whale (Mesoplodon densirostris). Each individual had previously stranded and was undergoing rehabilitation. Auditory Brainstem Responses (ABRs) were elicited via acoustic stimuli consisting of a train of broadband tone pulses presented at rates between 300 and 2000 Hz. Similar to other studied species, modulation rate transfer functions (MRTFs) of the studied individuals followed the shape of a low-pass filter, with the ability to process acoustic stimuli at presentation rates up to and exceeding 1250 Hz. Auditory integration times estimated from the bandwidths of the MRTFs ranged between 250 and 333 µs. The results support the hypothesis that high temporal resolution is conserved throughout the diverse range of odontocete species.
Assuntos
Percepção Auditiva/fisiologia , Golfinhos/fisiologia , Potenciais Evocados Auditivos , Baleias/fisiologia , Animais , Potenciais Evocados Auditivos/fisiologia , Masculino , Processamento de Sinais Assistido por ComputadorRESUMO
Hearing sensitivity change was investigated when a warning sound preceded a loud sound in the false killer whale (Pseudorca crassidens), the bottlenose dolphin (Tursiops truncatus), the beluga whale (Delphinaperus leucas) and the harbor porpoise (Phocoena phocoena). Hearing sensitivity was measured using pip-train test stimuli and auditory evoked potential recording. When the test/warning stimuli preceded a loud sound, hearing thresholds before the loud sound increased relative to the baseline by 13 to 17 dB. Experiments with multiple frequencies of exposure and shift provided evidence of different amounts of hearing change depending on frequency, indicating that the hearing sensation level changes were not likely due to a simple stapedial reflex.
Assuntos
Cetáceos/fisiologia , Potenciais Evocados Auditivos , Audição/fisiologia , Percepção Sonora/fisiologia , Animais , Aprendizagem , SomRESUMO
Hearing sensitivity, during trials in which a warning sound preceding a loud sound, was investigated in two harbor porpoises (Phocoena phocoena). Sensitivity was measured using pip-train test stimuli and auditory evoked potential recording. When a hearing test/warning stimulus, with a frequency of either 45 or 32 kHz, preceded a loud 32 kHz tone with a sound pressure level of 152 dB re 1 µPa root mean square, lasting 2 s yielding an sound exposure level (SEL) of 155 dB re 1 µPa(2)s, pooled hearing thresholds measured just before the loud sound increased relative to baseline thresholds. During two experimental sessions the threshold increased up to 17 dB for the test frequency of 45 kHz and up to 11 dB for the test frequency of 32 kHz. An extinction test revealed very rapid threshold recovery within the first two experimental sessions. The SEL producing the hearing dampening effect was low compared to previous other odontocete hearing change efforts with each individual trial equal to 155 dB re 1 µPa(2) but the cumulative SEL for each subsession may have been as high as 168 dB re 1 µPa(2). Interpretations of conditioned hearing sensation change and possible change due to temporary threshold shifts are considered for the harbor porpoise and discussed in the light of potential mechanisms and echolocation.
Assuntos
Potenciais Evocados Auditivos/fisiologia , Audição/fisiologia , Phocoena/fisiologia , Animais , Audiometria , Limiar Auditivo , Testes Auditivos/métodos , Percepção Sonora/fisiologia , Masculino , SomRESUMO
The hearing sensitivity of a bottlenose dolphin for a warning sound, when the exact time of the arrival of a loud sound could or could not be predicted, was measured. Sensitivity was measured when the time of onset of the loud sound was randomly varied (random-variation sessions) and when the time of onset of the loud sound and the pattern of stimulus levels was constant (fixed-stimulus sessions). The loud sound was kept the same in both of the series. The mean duration and mean range of the levels of the test/warning signal were also kept equal across experimental sessions. Hearing sensitivity was measured using the auditory evoked potential method with rhythmic trains of short pips as test stimuli. With randomly varied warning sounds, thresholds before the loud sound were on average 10.6 dB higher than the baseline thresholds. With fixed warning signals, thresholds were on average 4.4 dB higher than the baseline thresholds. Considering that the loud sounds were identical, the difference between the random-variation and the fixed-stimulus sessions cannot be explained by a direct (unconditioned) influence of sound exposure. Therefore, the data provide reliable evidence for the conditioning nature of the hearing-dampening effect and also demonstrate that hearing sensitivity change also depends on when the animal can expect the loud sound to occur.
Assuntos
Golfinho Nariz-de-Garrafa/fisiologia , Audição/fisiologia , Estimulação Acústica , Animais , Limiar Auditivo , Condicionamento Psicológico , Potenciais Evocados Auditivos , Feminino , Som , Fatores de TempoRESUMO
Ice-dwelling beluga whales are increasingly being exposed to anthropogenic loud sounds. Beluga's hearing sensitivity measured during a warning sound just preceding a loud sound was tested using pip-train stimuli and auditory evoked potential recording. When the test/warning stimulus with a frequency of 32 or 45 kHz preceded the loud sound with a frequency of 32 kHz and a sound pressure level of 153 dB re 1 µPa, 2 s, hearing thresholds before the loud sound increased relative to the baseline. The threshold increased up to 15 dB for the test frequency of 45 kHz and up to 13 dB for the test frequency of 32 kHz. These threshold increases were observed during two sessions of 36 trials each. Extinction tests revealed no change during three experimental sessions followed by a jump-like return to baseline thresholds. The low exposure level producing the hearing-dampening effect (156 dB re 1 µPa(2)s in each trial), and the manner of extinction, may be considered as evidence that the observed hearing threshold increases were a demonstration of conditioned dampening of hearing when the whale anticipated the quick appearance of a loud sound in the same way demonstrated in the false killer whale and bottlenose dolphin.
Assuntos
Limiar Auditivo/fisiologia , Beluga/fisiologia , Encéfalo/fisiologia , Extinção Psicológica/fisiologia , Audição/fisiologia , Aprendizagem/fisiologia , Estimulação Acústica , Animais , Eletroencefalografia , Potenciais Evocados Auditivos/fisiologia , FemininoRESUMO
Understanding the hearing of marine mammals has been a priority to quantify and mitigate the impact of anthropogenic sound on these apex predators. Yet our knowledge of cetacean hearing is still limited to a few dozen species, therefore compromising any attempt to design adaptive management strategies. The use of auditory evoked potentials allows scientists to rapidly and noninvasively obtain the hearing data of species rarely available in captivity. Unfortunately, many practical and ethical reasons still limit the availability of large whales, thus restricting the possibility to effectively ensure that anthropogenic sounds have minimum effects on these species. The example of a recent Blainville's beaked whale (Mesoplodon densirostris) audiogram collected after a stranding indicated, for instance, very specialized hearing between 40 and 50 kHz, which corresponded to the frequency-modulated upsweep signals used by this species during echolocation. The methods used during a stranding event are presented along with the major difficulties that have slowed down the scientific community in measuring the audition of large whales and the potential value in obtaining such results when successful.
Assuntos
Golfinhos/fisiologia , Audição/fisiologia , Baleias/fisiologia , AnimaisRESUMO
Echolocating animals adjust the transmit intensity and receive sensitivity of their sonar in order to regulate the sensation level of their echoes; this process is often termed automatic gain control. Gain control is considered not to be under the animal's cognitive control, but previous investigations studied animals ensonifying targets or hydrophone arrays at predictable distances. To test whether animals maintain gain control at a fixed level in uncertain conditions, we measured changes in signal intensity for a bottlenose dolphin (Tursiops truncatus) detecting a target at three target distances (2.5, 4 and 7 m) in two types of sessions: predictable and unpredictable. Predictable sessions presented the target at a constant distance; unpredictable sessions moved the target randomly between the three target positions. In the predictable sessions the dolphin demonstrated intensity distance compensation, increasing the emitted click intensity as the target distance increased. Additionally, as trials within sessions progressed, the animal adjusted its click intensity even from the first click in a click train, which is consistent with the animal expecting a target at a certain range. In the unpredictable sessions there was no significant difference of intensity with target distance until after the 7th click in a click train. Together, these results demonstrate that the bottlenose dolphin uses learning and expectation for sonar gain control.
Assuntos
Adaptação Fisiológica/fisiologia , Golfinho Nariz-de-Garrafa/fisiologia , Cognição/fisiologia , Acústica , Animais , Ecolocação/fisiologia , Feminino , Aprendizagem/fisiologia , SomRESUMO
Quantifying and understanding the impact of anthropogenic sound on marine mammals has been the focus of many researchers both in laboratory settings as well as in the field. This study presents the audiogram of a sub-adult Blainville's beaked whale that stranded in Hawaii. The hearing measurements were conducted using the non-invasive auditory brainstem response technique. A total of 11 sinusoidally amplitude modulated tones were tested ranging from 5.6 to 160 kHz. The audiogram data indicated that the region of best hearing was found between 40 and 50 kHz with thresholds below 50 dB. This frequency range partially overlaps with the frequency modulated upsweep that Blainville's beaked whales have been reported to use during echolocation. These results match the frequency range obtained from the hearing measurements of a Gervais' beaked whale previously tested using contact acoustic stimulation and emphasize the importance of obtaining rapid hearing measurements on live stranded animals to improve the understanding of poorly known species.