Echolocation behavior in big brown bats is not impaired after intense broadband noise exposures.

Echolocating bats emit trains of intense ultrasonic biosonar pulses and listen to weaker echoes returning from objects in their environment. Identification and categorization of echoes are crucial for orientation and prey capture. Bats are social animals and often fly in groups in which they are exposed to their own emissions and to those from other bats, as well as to echoes from multiple surrounding objects. Sound pressure levels in these noisy conditions can exceed 110 dB, with no obvious deleterious effects on echolocation performance. Psychophysical experiments show that big brown bats (Eptesicus fuscus) do not experience temporary threshold shifts after exposure to intense broadband ultrasonic noise, but it is not known if they make fine-scale adjustments in their pulse emissions to compensate for any effects of the noise. We investigated whether big brown bats adapt the number, temporal patterning or relative amplitude of their emitted pulses while flying through an acoustically cluttered corridor after exposure to intense broadband noise (frequency range 10-100 kHz; sound exposure level 152 dB). Under these conditions, four bats made no significant changes in navigation errors or in pulse number, timing and amplitude 20 min, 24 h or 48 h after noise exposure. These data suggest that big brown bats remain able to perform difficult echolocation tasks after exposure to ecologically realistic levels of broadband noise.

The modulation rate transfer function of a harbour porpoise (Phocoena phocoena).

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I-VI), and 1.4 ms (II-IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal's size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 µPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 µs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.

Keeping returns optimal: gain control exerted through sensitivity adjustments in the harbour porpoise auditory system.

Animals that use echolocation (biosonar) listen to acoustic signals with a large range of intensities, because echo levels vary with the fourth power of the animal's distance to the target. In man-made sonar, engineers apply automatic gain control to stabilize the echo energy levels, thereby rendering them independent of distance to the target. Both toothed whales and bats vary the level of their echolocation clicks to compensate for the distance-related energy loss. By monitoring the auditory brainstem response (ABR) during a psychophysical task, we found that a harbour porpoise (Phocoena phocoena), in addition to adjusting the sound level of the outgoing signals up to 5.4 dB, also reduces its ABR threshold by 6 dB when the target distance doubles. This self-induced threshold shift increases the dynamic range of the biosonar system and compensates for half of the variation of energy that is caused by changes in the distance to the target. In combination with an increased source level as a function of target range, this helps the porpoise to maintain a stable echo-evoked ABR amplitude irrespective of target range, and is therefore probably an important tool enabling porpoises to efficiently analyse and classify received echoes.

Stereotypical rapid source level regulation in the harbour porpoise biosonar.

Some odontocetes and bats vary both click intensity and receiver sensitivity during echolocation, depending on target range. It is not known how this so-called automatic gain control is regulated by the animal. The source level of consecutive echolocation clicks from a harbour porpoise was measured with a hydrophone array while the animal detected an aluminium cylinder at 2, 4 or 8 m distance in a go/no-go paradigm. On-axis clicks had source levels of 145-174 dB re 1 µPa peak-to-peak. During target-present trials the click trains reached comparable source levels independent of the range to the target after three clicks. After an additional click, the source level was reduced for the 2 and 4 m trials until it equalled the one-way transmission loss. During target-absent trials, the source level remained high throughout the entire click train. Given typical values of harbour porpoise inter-click intervals, the source level reduction commenced within a few 100 ms from the first click in the click train. This may indicate a sub-cortically regulated source level regulation in the harbour porpoise.

The vocal development of the pale spear-nosed bat is dependent on auditory feedback.

Human vocal development and speech learning require acoustic feedback, and humans who are born deaf do not acquire a normal adult speech capacity. Most other mammals display a largely innate vocal repertoire. Like humans, bats are thought to be one of the few taxa capable of vocal learning as they can acquire new vocalizations by modifying vocalizations according to auditory experiences. We investigated the effect of acoustic deafening on the vocal development of the pale spear-nosed bat. Three juvenile pale spear-nosed bats were deafened, and their vocal development was studied in comparison with an age-matched, hearing control group. The results show that during development the deafened bats increased their vocal activity, and their vocalizations were substantially altered, being much shorter, higher in pitch, and more aperiodic than the vocalizations of the control animals. The pale spear-nosed bat relies on auditory feedback for vocal development and, in the absence of auditory input, species-atypical vocalizations are acquired. This work serves as a basis for further research using the pale spear-nosed bat as a mammalian model for vocal learning, and contributes to comparative studies on hearing impairment across species. This article is part of the theme issue 'Vocal learning in animals and humans'.

Ontogeny of auditory brainstem responses in the bat, Phyllostomus discolor.

Hearing is the primary sensory modality in bats, but its development is poorly studied. For newborns, hearing appears essential in maintaining contact with their mothers and to develop echolocation abilities. Here we measured auditory brainstem responses (ABRs) to clicks and narrowband tone pips covering a large frequency range (5-90 kHz) in juveniles (p7 to p200) and adults of the bat, Phyllostomus discolor. Tone-pip audiograms show that juveniles at p7 are already quite responsive, not only below 20 kHz but up to 90 kHz. Hearing sensitivity increases further until about p14 and is then refined, possibly correlated with growth and differentiation of the animals' outer ears. ABR amplitudes decrease within the first 3-4 months, inversely correlated with the bat weight and forearm length. ABR Wave I latency decreases with increasing stimulation level. ABR duration (measured between Waves I and V) is longer in juveniles and shortens with age which may reflect temporal refinement of auditory brainstem neurons to accommodate the exceptional temporal precision required for effective echolocation. Overall our data show that P. discolor bats have good hearing very early in life. The current method represents a fast and minimally invasive way of characterizing basic hearing in bats.

Sonar beam dynamics in leaf-nosed bats.

Ultrasonic emissions of bats are directional and delimit the echo-acoustic space. Directionality is quantified by the aperture of the sonar beam. Recent work has shown that bats often widen their sonar beam when approaching movable prey or sharpen their sonar beam when navigating through cluttered habitats. Here we report how nose-emitting bats, Phyllostomus discolor, adjust their sonar beam to object distance. First, we show that the height and width of the bats sonar beam, as imprinted on a parabolic 45 channel microphone array, varies even within each animal and this variation is unrelated to changes in call level or spectral content. Second, we show that these animals are able to systematically decrease height and width of their sonar beam while focusing on the approaching object. Thus it appears that sonar beam sharpening is a further, facultative means of reducing search volume, likely to be employed by stationary animals when the object position is close and unambiguous. As only half of our individuals sharpened their beam onto the approaching object we suggest that this strategy is facultative, under voluntary control, and that beam formation is likely mediated by muscular control of the acoustic aperture of the bats' nose leaf.

Estimating colony sizes of emerging bats using acoustic recordings.

The decline of bats demands more widespread monitoring of populations for conservation and management. Current censusing methods are either prone to bias or require costly equipment. Here, we report a new method using passive acoustics to determine bat count census from overall acoustic amplitude of the emerging bat stream. We recorded the video and audio of an emerging colony of Mexican free-tailed bats from two cave locations across multiple nights. Instantaneous bat counts were calculated from the video frames, and the bat stream's acoustic amplitude corresponding to each video frame was determined using three different methods for calculating acoustic intensity. We found a significant link between all three acoustic parameters and bat count, with the highest R (2) of 0.742 linking RMS pressure and bat count. Additionally, the relationship between acoustics and population size at one cave location could accurately predict the population size at another cave location. The data were gathered with low-cost, easy-to-operate equipment, and the data analysis can be easily accomplished using automated scripts or with open-source acoustic software. These results are a potential first step towards creating an acoustic model to estimate bat population at large cave colonies worldwide.