Functional role of airflow-sensing hairs on the bat wing.

The wing membrane of the big brown bat (Eptesicus fuscus) is covered by a sparse grid of microscopic hairs. We showed previously that various tactile receptors (e.g., lanceolate endings and Merkel cell neurite complexes) are associated with wing-hair follicles. Furthermore, we found that depilation of these hairs decreased the maneuverability of bats in flight. In the present study, we investigated whether somatosensory signals arising from the hairs carry information about airflow parameters. Neural responses to calibrated air puffs on the wing were recorded from primary somatosensory cortex of E. fuscus Single units showed sparse, phasic, and consistently timed spikes that were insensitive to air-puff duration and magnitude. The neurons discriminated airflow from different directions, and a majority responded with highest firing rates to reverse airflow from the trailing toward the leading edge of the dorsal wing. Reverse airflow, caused by vortices, occurs commonly in slowly flying bats. Hence, the present findings suggest that cortical neurons are specialized to monitor reverse airflow, indicating laminar airflow disruption (vorticity) that potentially destabilizes flight and leads to stall. NEW & NOTEWORTHY: Bat wings are adaptive airfoils that enable demanding flight maneuvers. The bat wing is sparsely covered with sensory hairs, and wing-hair removal results in reduced flight maneuverability. Here, we report for the first time single-neuron responses recorded from primary somatosensory cortex to airflow stimulation that varied in amplitude, duration, and direction. The neurons show high sensitivity to the directionality of airflow and might act as stall detectors.

Morphology and deflection properties of bat wing sensory hairs: scanning electron microscopy, laser scanning vibrometry, and mechanics model.

Bat wings are highly adaptive airfoils that enable demanding flight maneuvers, which are performed with astonishing robustness under turbulent conditions, and stability at slow flight velocities. The bat wing is sparsely covered with microscopically small, sensory hairs that are associated with tactile receptors. In a previous study we demonstrated that bat wing hairs are involved in sensing airflow for improved flight maneuverability. Here, we report physical measurements of these hairs and their distribution on the wing surface of the big brown bat, Eptesicus fuscus, based on scanning electron microscopy analyses. The wing hairs are strongly tapered, and are found on both the dorsal and ventral wing surfaces. Laser scanning vibrometry tests of 43 hairs from twelve locations across the wing of the big brown bat revealed that their natural frequencies inversely correlate with length and range from 3.7 to 84.5 kHz. Young's modulus of the average wing hair was calculated at 4.4 GPa, which is comparable with rat whiskers or arthropod airflow-sensing hairs.

Organization of the primary somatosensory cortex and wing representation in the Big Brown Bat, Eptesicus fuscus.

Bats are the only mammals capable of true powered flight. The bat wing exhibits specializations, allowing these animals to perform complicated flight maneuvers like landing upside-down, and hovering. The wing membrane contains various tactile receptors, including hair-associated Merkel receptors that might be involved in stabilizing bat flight. Here, we studied the neuronal representation of the wing membrane in the primary somatosensory cortex (S1) of the anesthetized Big Brown Bat, Eptesicus fuscus, using tactile stimulation with calibrated monofilaments (von Frey hairs) while recording from multi-neuron clusters. We also measured cortical response thresholds to tactile stimulation of the wings.The body surface is mapped topographically across the surface of S1, with the head, foot, and wing being overrepresented. The orientation of the wing representation is rotated compared to the hand representaion of terrestrial mammals, confirming results from other bat species. Although different wing membrane parts derive embryologically from different body parts, including the flank (plagiopatagium), the tactile sensitivity of the entire flight membrane (0.2-1.2 mN) is remarkably close or even higher (dactylopatagium) than the average tactile sensitivity of the human fingertip.

Free-flight encounters between praying mantids (Parasphendale agrionina) and bats (Eptesicus fuscus).

Through staged free-flight encounters between echolocating bats and praying mantids, we examined the effectiveness of two potential predator-evasion behaviors mediated by different sensory modalities: (1) power dive responses triggered by bat echolocation detected by the mantis ultrasound-sensitive auditory system, and (2) ;last-ditch' maneuvers triggered by bat-generated wind detected by the mantis cercal system. Hearing mantids escaped more often than deafened mantids (76% vs 34%, respectively; hearing conveyed 42% advantage). Hearing mantis escape rates decreased when bat attack sequences contained very rapid increases in pulse repetition rates (escape rates <40% for transition slopes >16 p.p.s. 10 ms(-1); escape rates >60% for transition slopes <16 p.p.s. 10 ms(-1)). This suggests that echolocation attack sequences containing very rapid transitions (>16 p.p.s. 10 ms(-1)) could circumvent mantis/insect auditory defenses. However, echolocation attack sequences containing such transitions occurred in only 15% of the trials. Since mantis ultrasound-mediated responses are not 100% effective, cercal-mediated evasive behaviors triggered by bat-generated wind could be beneficial as a backup/secondary system. Although deafened mantids with functioning cerci did not escape more often than deafened mantids with deactivated cerci (35% vs 32%, respectively), bats dropped mantids with functioning cerci twice as frequently as mantids with deactivated cerci. This latter result was not statistically reliable due to small sample sizes, since this study was not designed to fully evaluate this result. It is an interesting observation that warrants further investigation, however, especially since these dropped mantids always survived the encounter.

Auditory sensitivity and frequency selectivity in greater spear-nosed bats suggest specializations for acoustic communication.

We investigated the relationship between auditory sensitivity, frequency selectivity, and the vocal repertoire of greater spear-nosed bats ( Phyllostomus hastatus). P. hastatus commonly emit three types of vocalizations: group-specific foraging calls that range from 6 to 11 kHz, low amplitude echolocation calls that sweep from 80 to 40 kHz, and infant isolation calls from 15 to 100 kHz. To determine if hearing in P. hastatus is differentially sensitive or selective to frequencies in these calls, we determined absolute thresholds and masked thresholds using an operant conditioning procedure. Both absolute and masked thresholds were lowest at 15 kHz, which corresponds with the peak energy of isolation calls. Auditory and masked thresholds were higher at sound frequencies used for group-specific foraging calls and echolocation calls. Isolation calls meet the requirements of individual signatures and facilitate parent-offspring recognition. Many bat species produce isolation calls with peak energy between 10 and 25 kHz, which corresponds with the frequency region of highest sensitivity in those species for which audiogram data are available. These findings suggest that selection for accurate offspring recognition exerts a strong influence on the sensory system of P. hastatus and likely on other species of group-living bats.

Auditory scene analysis by echolocation in bats.

Echolocating bats transmit ultrasonic vocalizations and use information contained in the reflected sounds to analyze the auditory scene. Auditory scene analysis, a phenomenon that applies broadly to all hearing vertebrates, involves the grouping and segregation of sounds to perceptually organize information about auditory objects. The perceptual organization of sound is influenced by the spectral and temporal characteristics of acoustic signals. In the case of the echolocating bat, its active control over the timing, duration, intensity, and bandwidth of sonar transmissions directly impacts its perception of the auditory objects that comprise the scene. Here, data are presented from perceptual experiments, laboratory insect capture studies, and field recordings of sonar behavior of different bat species, to illustrate principles of importance to auditory scene analysis by echolocation in bats. In the perceptual experiments, FM bats (Eptesicus fuscus) learned to discriminate between systematic and random delay sequences in echo playback sets. The results of these experiments demonstrate that the FM bat can assemble information about echo delay changes over time, a requirement for the analysis of a dynamic auditory scene. Laboratory insect capture experiments examined the vocal production patterns of flying E. fuscus taking tethered insects in a large room. In each trial, the bats consistently produced echolocation signal groups with a relatively stable repetition rate (within 5%). Similar temporal patterning of sonar vocalizations was also observed in the field recordings from E. fuscus, thus suggesting the importance of temporal control of vocal production for perceptually guided behavior. It is hypothesized that a stable sonar signal production rate facilitates the perceptual organization of echoes arriving from objects at different directions and distances as the bat flies through a dynamic auditory scene. Field recordings of E. fuscus, Noctilio albiventris, N. leporinus, Pippistrellus pippistrellus, and Cormura brevirostris revealed that spectral adjustments in sonar signals may also be important to permit tracking of echoes in a complex auditory scene.

A computational sensorimotor model of bat echolocation.

A computational sensorimotor model of target capture behavior by the echolocating bat, Eptesicus fuscus, was developed to understand the detection, localization, tracking, and interception of insect prey in a biological sonar system. This model incorporated acoustics, target localization processes, flight aerodynamics, and target capture planning to produce model trajectories replicating those observed in behavioral insect capture trials. Estimates of target range were based on echo delay, azimuth on the relative intensity of the echo at the two ears, and elevation on the spectral pattern of the sonar return in a match/mismatch process. Flapping flight aerodynamics was used to produce realistic model trajectories. Localization in all three spatial dimensions proved necessary to control target tracking and interception for an adequate model of insect capture behavior by echolocating bats. Target capture using maneuvering flight was generally successful when the model's path was controlled by a planning process that made use of an anticipatory internal simulation, while simple homing was successful only for targets directly ahead of the model bat.

Echolocation behavior of big brown bats, Eptesicus fuscus, in the field and the laboratory.

Echolocation signals were recorded from big brown bats, Eptesicus fuscus, flying in the field and the laboratory. In open field areas the interpulse intervals (IPI) of search signals were either around 134 ms or twice that value, 270 ms. At long IPI's the signals were of long duration (14 to 18-20 ms), narrow bandwidth, and low frequency, sweeping down to a minimum frequency (Fmin) of 22-25 kHz. At short IPI's the signals were shorter (6-13 ms), of higher frequency, and broader bandwidth. In wooded areas only short (6-11 ms) relatively broadband search signals were emitted at a higher rate (avg. IPI= 122 ms) with higher Fmin (27-30 kHz). In the laboratory the IPI was even shorter (88 ms), the duration was 3-5 ms, and the Fmin 30- 35 kHz, resembling approach phase signals of field recordings. Excluding terminal phase signals, all signals from all areas showed a negative correlation between signal duration and Fmin, i.e., the shorter the signal, the higher was Fmin. This correlation was reversed in the terminal phase of insect capture sequences, where Fmin decreased with decreasing signal duration. Overall, the signals recorded in the field were longer, with longer IPI's and greater variability in bandwidth than signals recorded in the laboratory.

Vocal control of acoustic information for sonar discriminations by the echolocating bat, Eptesicus fuscus.

This study aimed to determine whether bats using frequency modulated (FM) echolocation signals adapt the features of their vocalizations to the perceptual demands of a particular sonar task. Quantitative measures were obtained from the vocal signals produced by echolocating bats (Eptesicus fuscus) that were trained to perform in two distinct perceptual tasks, echo delay and Doppler-shift discriminations. In both perceptual tasks, the bats learned to discriminate electronically manipulated playback signals of their own echolocation sounds, which simulated echoes from sonar targets. Both tasks utilized a single-channel electronic target simulator and tested the bat's in a two-alternative forced choice procedure. The results of this study demonstrate changes in the features of the FM bats' sonar sounds with echolocation task demands, lending support to the notion that this animal actively controls the echo information that guides its behavior.

Echo-delay resolution in sonar images of the big brown bat, Eptesicus fuscus.

Echolocating big brown bats (Eptesicus fuscus) broadcast ultrasonic frequency-modulated (FM) biosonar sounds (20-100 kHz frequencies; 10-50 microseconds periods) and perceive target range from echo delay. Knowing the acuity for delay resolution is essential to understand how bats process echoes because they perceive target shape and texture from the delay separation of multiple reflections. Bats can separately perceive the delays of two concurrent electronically generated echoes arriving as little as 2 microseconds apart, thus resolving reflecting points as close together as 0.3 mm in range (two-point threshold). This two-point resolution is roughly five times smaller than the shortest periods in the bat's sounds. Because the bat's broadcasts are 2,000-4,500 microseconds long, the echoes themselves overlap and interfere with each other, to merge together into a single sound whose spectrum is shaped by their mutual interference depending on the size of the time separation. To separately perceive the delays of overlapping echoes, the bat has to recover information about their very small delay separation that was transferred into the spectrum when the two echoes interfered with each other, thus explicitly reconstructing the range profile of targets from the echo spectrum. However, the bat's 2-microseconds resolution limit is so short that the available spectral cues are extremely limited. Resolution of delay seems overly sharp just for interception of flying insects, which suggests that the bat's biosonar images are of higher quality to suit a wider variety of orientation tasks, and that biosonar echo processing is correspondingly more sophisticated than has been suspected.

Target flutter rate discrimination by bats using frequency-modulated sonar sounds: behavior and signal processing models.

This study utilized psychophysical data and acoustical measurements of sonar echoes from artificial fluttering targets to develop insights to the information used by FM bats to discriminate the wingbeat rate of flying insects. Fluttering targets were produced by rotating blades that moved towards the bat, and the animal learned to discriminate between two rates of movement, a reference rate (30 or 50 Hz) and a slower, variable rate. Threshold discrimination performance depended on the rotation rate of the reference target, with a difference value of 9 Hz for the reference rate of 30 Hz and 14 Hz for the reference rate of 50 Hz. Control experiments demonstrated that the bats used sonar echoes from the moving targets to perform the discrimination task. Acoustical measurements showed that the moving target produced a Doppler shift in the echo and a concomitant change in the arrival time of each frequency in the linear period FM sweep. The difference in delay between echoes from moving and stationary parts varied linearly with flutter rate and depended on the characteristics of the bat's sonar sounds. Simulations also showed a reduction in average echo bandwidth with increasing flutter rate, which may account for a higher delay discrimination threshold using the 50-Hz reference rate. This work suggests that Doppler-induced changes in echo delays produced by fluttering targets may contribute to the FM bat's perception of flying insect prey.

Spatially selective auditory responses in the superior colliculus of the echolocating bat.

When a bat approaches a target, it continuously modifies its echolocation sounds and relies on incoming echo information to shape the characteristics of its subsequent sonar cries. In addition, acoustic information about the azimuth and elevation of a sonar target elicits orienting movements of the head and pinnae toward the sound source. This requires a common sensorimotor interface, where echo information is used to guide motor behaviors. Using single-unit neurophysiological methods and free-field auditory stimulation, we present data on biologically relevant specializations in the superior colliculus (SC) of the bat for orientation by sonar. In the bat's SC, two classes of spatially tuned neurons are distinguished by their sensitivity to echoes. One population shows facilitated, delay-tuned responses to pairs of sounds, simulating sonar emissions and echoes. Delay tuning, related to encoding target range, may play a role in guiding motor responses in echolocation, because the bat adjusts its emissions with changes in target distance. The delay-facilitated response depends on the direction of stimulation and on the temporal relationship between the simulated emission and echo in the sound pair, suggesting that this class of neurons represents the location of a target in three dimensions. A second population encodes the target in two dimensions, azimuth and elevation, and does not show a facilitated response to echoes delivered from any locus. Encoding of azimuth and elevation may be important for directing head aim, and this class may function in transforming auditory spatial information into signals used to guide acoustic orientation.

The brain-stem auditory-evoked response in the big brown bat (Eptesicus fuscus) to clicks and frequency-modulated sweeps.

Three experiments were performed to evaluate the effects of stimulus level on the brain-stem auditory-evoked response (BAER) in the big brown bat (Eptesicus fuscus), a species that uses frequency-modulated (FM) sonar sounds for echolocation. In experiment 1, the effects of click level on the BAER were investigated. Clicks were presented at levels of 30 to 90 dB pSPL in 10-dB steps. Each animal responded reliably to clicks at levels of 50 dB pSPL and above, showing a BAER containing four peaks in the first 3-4 ms from click onset (waves i-iv). With increasing click level, BAER peak amplitude increased and peak latency decreased. A decrease in the i-iv interval also occurred with increasing click level. In experiment 2, stimuli were 1-ms linear FM sweeps, decreasing in frequency from 100 to 20 kHz. Stimulus levels ranged from 20 to 90 dB pSPL. BAERs to FM sweeps were observed in all animals for levels of 40 dB pSPL and above. These responses were similar to the click-evoked BAER in waveform morphology, with the notable exception of an additional peak observed at the higher levels of FM sweeps. This peak (wave ia) occurred prior to the first wave seen at lower levels (wave ib). As the level of the FM sweep increased, there was a decrease in peak latency and an increase in peak amplitude. Similarity in the magnitude and behavior of the i-iv and ib-iv intervals suggests that wave ib to FM sweeps is the homolog of the wave i response to click stimuli. Experiment 3 tested the hypothesis that wave ia represented activity emanating from more basal cochlear regions than wave ib. FM sweeps (100-20 kHz) were presented at 90 dB pSPL, and broadband noise was raised in level until the BAER was eliminated. This "masked threshold" occurred at 85 dB SPL of noise. At masked threshold, the broadband noise was steeply high-pass filtered at five cutoff frequencies ranging from 20 to 80 kHz. Generally, wave ia was eliminated for masker cutoff frequencies of 56.6 kHz and below, while wave ib was typically observed for masker cutoffs down to 28.3 kHz. The results of these three experiments are compared and contrasted with data from other mammalian BAER studies.

Acoustic information available to bats using frequency-modulated sounds for the perception of insect prey.

Through the present study, the acoustic information available to an echolocating bat that uses brief frequency-modulated (FM) sonar sounds for the pursuit and capture of insect prey has been characterized. Computer-generated sonar pulses were broadcast at tethered insects, and the returning echoes were recorded on analog tape at high speed for off-line analyses. Echoes from stationary and fluttering insects were displayed using time waveform, spectrogram, power spectrum, and cross-correlation representations. The results show echo signatures for the different insect species studied, which change with the angle of incident sound. Sequences of echoes from fluttering insects show irregular changes in sound amplitude and time-frequency structure, reflecting a random temporal relation between the changing wing position and the arrival of incident sound. A set of recordings that controlled the temporal relation between incident sound and insect wing position suggests that information about the spatial profile of a flying insect could be enhanced if the bat were to produce a sequence of sounds that synchronized briefly with the moving target's wing-beat cycle. From this study, it has been proposed that the FM bat receives stroboscopic-like glimpses of fluttering prey whose spatial representation depends on the operation of the bat's sonar receiver.

Acoustic image representation of a point target in the bat Eptesicus fuscus: evidence for sensitivity to echo phase in bat sonar.

Echolocating bats, Eptesicus fuscus, were trained in two distinct behavioral tasks to investigate the images they perceive of a sonar point target. In the first task, bats were trained in a two-alternative forced-choice procedure to detect electronically simulated target echoes at a range of approximately 57 cm. Half of the trials in the detection task contained echoes from a stationary target (simulated by a fixed echo delay) and half contained echoes from a jittering target (simulated by an echo delay alternating between two time values over successive sonar emissions). In the second task, bats were trained in a two-alternative forced-choice procedure to discriminate between electronically simulated stationary and jittering targets, centered about a range of 57 cm. Both target detection and target jitter discrimination performance were assessed as a function of jitter magnitude, with jitter values ranging from 0-60 microseconds (corresponding to a change in distance of 0 to 10.3 mm). In both detection and discrimination tasks, the bat's performance changed cyclically with the magnitude of echo jitter. Specifically, when the phase of the playback echoes was unchanged, performance levels were poorest at 0 and 30 microseconds, and when the phase of the echoes alternated by 180 deg from one to the next, performance levels were poorest at 15 and 40-50 microseconds. The results suggest that Eptesicus is sensitive to the phase reversal of echoes and thus have implications for assessing receiver models of echolocation.

Discrimination of jittered sonar echoes by the echolocating bat, Eptesicus fuscus: the shape of target images in echolocation.

1. Behavioral experiments with jittering echoes examined acoustic images of sonar targets in the echolocating bat, Eptesicus fuscus, along the echo delay or target range axis. Echo phase, amplitude, bandwidth, and signal-to-noise ratio were manipulated to assess the underlying auditory processes for image formation. 2. Fine delay acuity is about 10 ns. Calibration and control procedures indicate that this represents temporal acuity rather than spectral discrimination. Jitter discrimination curves change in phase when the phase of one jittering echo is shifted by 180 degrees relative to the other, showing that echo phase is involved in delay estimation. At an echo detectability index of about 36 dB, fine acuity is 40 ns, which is approximately as predicted for the delay accuracy of an ideal receiver. 3. Compound performance curves for 0 degrees and 180 degrees phase conditions match the crosscorrelation function of the echoes. The locations of both 0 degrees and 180 degrees phase peaks in the performance curves shift along the time axis by an amount that matches neural amplitude-latency trading in Eptesicus, confirming a temporal basis for jitter discrimination.

Convergence of temporal and spectral information into acoustic images of complex sonar targets perceived by the echolocating bat, Eptesicus fuscus.

1. FM echolocating bats (Eptesicus fuscus) were trained to discriminate between a two-component complex target and a one-component simple target simulated by electronically-returned echoes in a series of experiments that explore the composition of the image of the two-component target. In Experiment I, echoes for each target were presented sequentially, and the bats had to compare a stored image of one target with that of the other. The bats made errors when the range of the simple target corresponded to the range of either glint in the complex target, indicating that some trace of the parts of one image interfered with perception of the other image. In Experiment II, echoes were presented simultaneously as well as sequentially, permitting direct masking of echoes from one target to the other. Changes in echo amplitude produced shifts in apparent range whose pattern depended upon the mode of echo presentation. 2. Eptesicus perceives images of complex sonar targets that explicitly represent the location and spacing of discrete glints located at different ranges. The bat perceives the target's structure in terms of its range profile along a psychological range axis using a combination of echo delay and echo spectral representations that together resemble a spectrogram of the FM echoes. The image itself is expressed entirely along a range scale that is defined with reference to echo delay. Spectral information contributes to the image by providing estimates of the range separation of glints, but it is transformed into these estimates. 3. Perceived absolute range is encoded by the timing of neural discharges and is vulnerable to shifts caused by neural amplitude-latency trading, which was estimated at 13 to 18 microseconds per dB from N1 and N4 auditory evoked potentials in Eptesicus. Spectral cues representing the separation of glints within the target are transformed into estimates of delay separations before being incorporated into the image. However, because they are encoded by neural frequency tuning rather than the time-of-occurrence of neural discharges, the perceived range separation of glints in images is not vulnerable to amplitude-latency shifts. 4. The bat perceives an image that is displayed in the domain of time or range. The image receives no evident spectral contribution beyond what is transformed into delay estimates. Although the initial auditory representation of FM echoes is spectrogram-like, the time, frequency, and amplitude dimensions of the spectrogram appear to be compressed into an image that has only time and amplitude dimensions.(ABSTRACT TRUNCATED AT 400 WORDS)

Frequency selectivity of hearing in the green treefrog, Hyla cinerea.

Frequency selectivity of hearing was measured in the green treefrog, Hyla cinerea. A psychophysical technique based on reflex modification was used to obtain masked threshold estimates for pure tones (300-5,400 Hz) presented against two levels of broadband masking noise. A pure tone (S-1) presented 200 ms prior to a reflex-eliciting stimulus (S-2) inhibited the motor reflex response to S-2. The magnitude of this reflex modification effect varied systematically with the sound pressure level (SPL) of S-1, and threshold was defined as the SPL of S-1 at which the reflex modification effect disappeared. Masked thresholds were used to calculate critical ratios, an index of the auditory system's frequency selectivity. The frequency selectivity of the treefrog's hearing is greatest and critical ratios are lowest (22-24 dB) at about 900 and 3,000 Hz, the two spectral regions dominant in the male treefrog's species-specific advertisement call. These results suggest that the treefrog's auditory system may be specialized to reject noise at biologically-relevant frequencies. As in other vertebrates, critical ratios remain constant when background noise level is varied; however, the shape of the treefrog's critical ratio function across frequencies differs from the typical vertebrate function that increases with increasing frequency at a slope of about 3 dB/octave. Instead, the treefrog's critical ratio function resembles its pure tone audiogram. Although the shape of the treefrog's critical ratio function is atypical, the critical ratio values themselves are comparable to those of many other vertebrates in the same frequency range. Critical ratio values here measured behaviorally do not match critical ratio values previously measured physiologically in single eighth nerve fibers.(ABSTRACT TRUNCATED AT 250 WORDS)

Spatial displacement sensitivity of X- and Y-cells in the dorsal lateral geniculate nucleus of the cat.

The sensitivity of X- and Y-cells in the dorsal lateral geniculate nucleus of the cat to small, temporally modulated displacements of grating stimuli was measured at 0.175, 0.25, 0.50, 1.00, and 2.00 c/deg. For every cell, two threshold measures were determined: first, a contrast threshold with a counterphase grating and then a displacement threshold with a grating matched in spatial frequency, but whose contrast was 2.5 times the threshold value. The results showed that displacement thresholds of both X- and Y-cells decreased with increasing spatial frequency. At low spatial frequencies, mean displacement thresholds of X- and Y-cells were similar, but at intermediate spatial frequencies, Y-cell thresholds were lower than X. X-cell displacement thresholds were lower than Y only at the highest spatial frequency tested. Consistent with previous reports, contrast thresholds also varied with spatial frequency for both X- and Y-cells. The local luminance differences produced by the contrast threshold and displacement threshold stimuli for the two classes of cells were compared. Across all spatial frequencies, the change in position of the gratings at displacement threshold produced smaller luminance differences than the counterphase gratings at contrast threshold. This enhanced sensitivity of X- and Y-cells to a local luminance changes produced by grating displacement was related to the high spatial contrast of the grating and not to the displacement per se.

Behavioral audiograms of the bullfrog (Rana catesbeiana) and the green tree frog (Hyla cinerea).

Reflex modification was used in a psychophysical technique to measure absolute auditory sensitivity of two species of anurans. Behavioral audiograms for these animals reveal that the bullfrog can detect sounds from 100 Hz to 3.2 kHz and the green tree frog from 100 Hz to 5 kHz. The shape and the sensitivity of these behavioral audiograms are similar to those of neural evoked-response audiograms of these animals. Absolute auditory sensitivity of anurans is only partially related to the spectral composition of their species-specific vocalizations.