Mary E. Bates

Bats Use Echo Harmonic Structure to Distinguish Their Targets from Background Clutter

Bats use temporal differences in sonar echoes to suppress interference from background clutter. When echolocating big brown bats fly in complex surroundings, echoes arriving from irrelevant objects (clutter) located to the sides of their sonar beam can mask perception of relevant objects located to the front (targets), causing “blind spots.” Because the second harmonic is beamed more weakly to the sides than the first harmonic, these clutter echoes have a weaker second harmonic. In psychophysical experiments, we found that electronically misaligning first and second harmonics in echoes (to mimic the misalignment of corresponding neural responses to harmonics in clutter echoes) disrupts the bat’s echo-delay perception but also prevents clutter masking. Electronically offsetting harmonics to realign their neural responses restores delay perception but also clutter interference. Thus, bats exploit harmonics to distinguish clutter echoes from target echoes, sacrificing delay acuity to suppress masking.

FM echolocating bats shift frequencies to avoid broadcast–echo ambiguity in clutter

Sonar broadcasts are followed by echoes at different delays from objects at different distances. When broadcasts are emitted rapidly in cluttered surroundings, echo streams from successive broadcasts overlap and cause ambiguity in matching echoes to corresponding broadcasts. To identify reactions to ambiguity in clutter, echolocating bats that emit multiple-harmonic FM sounds were trained to fly into a dense, extended array of obstacles (multiple rows of vertically hanging chains) while the sonar sounds the bat emitted were recorded with a miniature radio microphone carried by the bat. Flight paths were reconstructed from thermal-infrared video recordings. Successive rows of chains extended more than 6 m in depth, so each broadcast was followed by a series of echoes from multiple rows of chains that lasted up to 40 ms. Bats emitted sounds in pairs (“strobe groups”) at short (20–40 ms) interpulse intervals (IPIs) alternating with longer IPIs (>50 ms). For many short IPIs, the stream of echoes from the first broadcast was still arriving when the second broadcast was emitted. This overlap caused ambiguity about matching echoes with broadcasts. Bats shifted frequencies of the first sound in each strobe group upward and the second sound downward by 3–6 kHz. When overlap and ambiguity ceased, frequency shifts ceased also. Frequency differences were small compared with the total broadcast band, which was 75–80 kHz wide, but the harmonic structure of echoes enhances the differences in spectrograms. Bats could use time–frequency comparisons of echoes with broadcasts to assign echoes to the corresponding broadcasts and thus avoid ambiguity.

Jamming avoidance response of big brown bats in target detection

SUMMARY When searching for prey, big brown bats (Eptesicus fuscus) enhance the range of their sonar by concentrating more energy in the nearly constant-frequency (CF) tail portion of their frequency-modulated (FM) sweeps. We hypothesize that this portion of their signals may be vulnerable to interference from conspecifics using the same frequencies in their own emissions. To determine how bats modify their signals when confronted with an interfering stimulus, we compared the echolocation calls of bats when a CF jamming tone was on and off. The bats performed a two-alternative forced-choice detection task in the laboratory that required the use of echolocation. All three bats shifted the tail-end CF component of their emitted frequency bidirectionally away from the CF jamming stimulus only when the jamming frequency was within 2–3 kHz of the preferred baseline frequency of the bat. The duration of their emissions did not differ between the jamming and no-jamming trials. The jamming avoidance response of bats may serve to avoid masking or interference in a narrow range of frequencies important for target detection.

Perception of echo delay is disrupted by small temporal misalignment of echo harmonics in bat sonar

SUMMARY Echolocating big brown bats emit ultrasonic frequency-modulated (FM) biosonar sounds containing two prominent downward-sweeping harmonics (FM1 and FM2) and perceive target distance from echo delay. In naturally occurring echoes, FM1 and FM2 are delayed by the same amount. Even though echoes from targets located off-axis or far away are lowpass filtered, which weakens FM2 relative to FM1, their delays remain the same. We show here that misalignment of FM2 with FM1 by only 2.6 μs is sufficient to significantly disrupt acuity, which then persists for larger misalignments up to 300 μs. However, when FM2 is eliminated entirely rather than just misaligned, acuity is effectively restored. For naturally occurring, lowpass-filtered echoes, neuronal responses to weakened FM2 are retarded relative to FM1 because of amplitude-latency trading, which misaligns the harmonics in the bats internal auditory representations. Electronically delaying FM2 relative to FM1 mimics the retarded neuronal responses for FM2 relative to FM1 caused by amplitude-latency trading. Echoes with either electronically or physiologically misaligned harmonics are not perceived as having a clearly defined delay. This virtual collapse of delay acuity may suppress interference from off-axis or distant clutter through degradation of delay images for clutter in contrast to sharp images for nearer, frontal targets.

Analyzing acoustic interactions in natural bullfrog (Rana catesbeiana) choruses.

Analysis of acoustic interactions between animals in active choruses is complex because of the large numbers of individuals present, their high calling rates, and the considerable numbers of vocalizations that either overlap or show close temporal alternation. The authors describe a methodology for recording chorus activity in bullfrogs (Rana catesbeiana) using multiple, closely spaced acoustic sensors that provide simultaneous estimates of sound direction and sound characteristics. This method provides estimates of location of individual callers, even under conditions of call overlap. This is a useful technique for understanding the complexity of the acoustic scene faced by animals vocalizing in groups.

Effects of filtering of harmonics from biosonar echoes on delay acuity by big brown bats (Eptesicus fuscus)

Big brown bats emit FM biosonar sounds containing two principal harmonics (FM1 approximately 55-22 kHz;FM2 approximately 105-45 kHz). To examine the role of harmonics, they were selectively filtered from stimuli in electronic-echo delay discrimination experiments. Positive stimuli were delayed by 3.16 ms (55 cm simulated target range); negative stimuli were by delayed by 3.96 ms (68 cm). This large 800-micros delay difference (nearly 14 cm) was easily discriminated for echoes containing equal-strength FM1 and FM2. Performance gradually decreased as highpass filters removed progressively larger segments from FM1. For echoes with FM2 alone, performance collapsed to chance, but performance remained good for lowpass echoes containing FM1 alone. Attenuation of FM2 by 3 dB relative to FM1 also decreased performance, but shortening electronic delay of the attenuated FM2 by 48 micros counteracted amplitude-latency trading and restored performance. Bats require the auditory representations of FM1 and FM2 to be in temporal register for high delay acuity. Misalignment of neuronal responses degrades acuity, but outright removal of FM2, leaving only FM1, causes little loss of acuity. Functional asymmetry of harmonics reflects lowpass effects from beaming and atmospheric propagation, which leave FM1 intact. It may cooperate with latency shifts to aid in suppression of clutter.

Spatial location influences vocal interactions in bullfrog choruses

A multiple sensor array was employed to identify the spatial locations of all vocalizing male bullfrogs (Rana catesbeiana) in five natural choruses. Patterns of vocal activity collected with this array were compared with computer simulations of chorus activity. Bullfrogs were not randomly spaced within choruses, but tended to cluster into closely spaced groups of two to five vocalizing males. There were nonrandom, differing patterns of vocal interactions within clusters of closely spaced males and between different clusters. Bullfrogs located within the same cluster tended to overlap or alternate call notes with two or more other males in that cluster. These near-simultaneous calling bouts produced advertisement calls with more pronounced amplitude modulation than occurred in nonoverlapping notes or calls. Bullfrogs located in different clusters more often alternated entire calls or overlapped only small segments of their calls. They also tended to respond sequentially to calls of their farther neighbors compared to their nearer neighbors. Results of computational analyses showed that the observed patterns of vocal interactions were significantly different than expected based on random activity. The use of a multiple sensor array provides a richer view of the dynamics of choruses than available based on single microphone techniques.

Spatial release from simultaneous echo masking in bat sonar

Big brown bats (Eptesicus fuscus) use biosonar to navigate and locate objects in their surroundings. During natural foraging, they often encounter echoes returned by a target of interest located to the front while other, often stronger, clutter echoes are returned from objects, such as vegetation, located to the sides or above. Nevertheless, bats behave as if they do not suffer interference from this clutter. Using a two-choice delay discrimination procedure, bats were tested for the masking effectiveness of clutter echoes on target echoes when the target echoes were delivered from the bats front while clutter echoes were delivered from 90° overhead, a direction of lowpass filtering by the external ears. When clutter echoes are presented from the front at the same delay as target echoes, detection performance declines and clutter masking occurs. When the clutter echoes are presented at the same delay but from overhead, discrimination performance is unaffected and no masking occurs. Thus there is masking release for simultaneous off-axis lowpass clutter compared to masking by simultaneous clutter from the front. The bats performance for simultaneous target and clutter echoes indicates a new role for the mechanism that separates overlapping echoes by decomposing the bats auditory time-frequency representation.

Non‐random patterns of acoustic interactions in chorusing bullfrogs.

Male bullfrogs form choruses to vocally advertise for females and to announce territory occupation to rival males. In dense choruses, calls of individual males may temporally overlap due to the large numbers of vocalizing neighbors. Overlapping calls may be a deliberate communicative strategy, perhaps to form a more salient auditory object in order to effectively guide females to a particular location within the chorus. We used a custom‐written MATLAB program to simulate the proportions of overlapping and nonoverlapping calls in five mock choruses, and then compared model output to empirical data from natural choruses of equal size. The simulation assumed that each bullfrog called independently of his neighbors according to a Poisson process so that overlapping calls occur randomly rather than through cooperation. In four of the five empirical recordings, the number of overlapping calls was significantly smaller than the averages produced by corresponding simulations, indicating that more males vocalized together in bouts than predicted by the random simulation. Empirical and simulated data also differed significantly in the remaining chorus, but in the opposite direction. These data suggest that overlapping calls may confer some communicative advantage. [Work supported by NSF Advance program.]

Echolocation behavior of pairs of flying Eptesicus fuscus recorded with a Telemike microphone.

Four big brown bats (Eptesicus fuscus) were flown singly and in pairs in a room containing a sparse array of vertically hanging plastic chains as obstacles. Each bat carried a lightweight radio telemetry microphone (Telemike) that recorded their emitted echolocation sounds without artifacts from Doppler shifts, directional effects, and atmospheric attenuation. The broadcasts of both bats were also recorded with two stationary ultrasonic microphones located at the far end of the flight room. The echolocation broadcasts of bats flying singly were compared to those emitted when the bats were flown together. The principal change was shifting of harmonic frequencies very slightly (<5 kHz) away from each other and from frequencies used when flying alone. In contrast, the duration of emissions was more stable between single and double bat flights. Changes in ending frequency have been associated with a jamming avoidance response in big brown bats and could indicate attempts to avoid interference while flying wit...