Echolocation Sounds Research Articles

The development of vocalizations in Pipistrellus pipistrellus (Chiroptera: Vespertilionidae) during post‐natal growth and the maintenance of individual vocal signatures

The development of vocalizations during post‐natal growth in the microchiropteran bat Pipistrellus pipistrellus is described. Vocalizations served as precursors of echolocation sounds and as isolation calls used to attract mothers. Although calls judged to be echolocation precursors tended to be of short duration and possessed high rates of frequency modulation when compared with isolation calls, a wide spectrum of intermediate calls existed making classification diffuse. Changes in the frequencies and durations of calls during growth are described. Multiple discriminant analysis showed considerable inter‐individual variation in isolation calls, while the calls of any individual were remarkably consistent in their structure at both six days and 15 days of age. The relative importance of acoustic call parameters in contributing towards inter‐individual variation changed between six days and 15 days. Vocal signatures of youngsters were probably important in allowing a mother to identify her own offspring in a creche thus preventing misdirected maternal care in this species, though vocal signatures of infants changed during development. Echolocation calls of flying juveniles were compared with calls from their mothers.

Echolocation in the notch-eared bat, Myotis emarginatus

In Myotis emarginatus, the patterns of echolocation sounds vary with different foraging habitats: In commuting flights the echolocation sounds are linearly frequency modulated sweeps that start at about 100 kHz, terminate at 40 kHz, and have a duration of 1–3 ms. They consist of a loud first harmonic. The second and third harmonics are at least 15 dB fainter than the first one and often undetectable. A distinctly different type of sound is emitted when the bats search for flying insects in open spaces. The sounds are reduced in bandwidth and elongated by a constant frequency component that follows the initial frequency modulated part. Typically, sounds start at about 94 kHz and terminate in a constant frequency component at about 40–45 kHz. The average duration of the constant frequency tail is 2.8 ms; this approximately doubles the length of the pulse, with the longest recorded sound lasting 7.2 ms. When bats are foraging near and within foliage, and gleaning prey from foliage, echolocation sounds are brief (average 1 ms) frequency modulated pulses with a broad bandwidth. The pulses start at about 105 kHz and sweep down to 25 kHz. During gleaning within a building, the frequency range of the sounds is shifted to higher frequencies and extends from 124 to 52 kHz. When the bats forage for aireal insects in a confined area that creates echo-clutter, they emit sounds similar to those used during gleaning within buildings except that sound durations are extended to about 1.8 ms. In each foraging area, the echolocation sounds emitted during the search for and approach to prey are similar in structure. Sound and pause durations are reduced in the approach phase. Irrespective of foraging style and habitat, immediately before capture the bat emits a rapid and stereotyped sequence of 2-10 echolocation pulses (final buzz). These pulses are brief (0.2–0.5 ms), frequency modulated sounds with a reduced bandwidth. The sounds start at 45 kHz and sweep down to 35–20 kHz. The repetition rate is increased up to 200 pulses/s.

The structure of echolocation sounds used by the big brown bat Eptesicus fuscus: Some consequences for echo processing

Analysis of the sonar emissions produced by five big brown bats (Eptesicus fuscus) during detection and range-discrimination tests show that each bat has a distinctive, “personal” emission. This may help bats distinguish their own emissions from those of others and thereby prevent jamming. Emissions usually contain three harmonics and are strongly frequency swept. Modeling of the sweep shape showed it to be more nearly linear in log time than hyperbolic (linear period modulation), which is optimal for Doppler tolerance. However, logarithmic time and linear period modulation are similar in their Doppler properties, and so bats may suffer little penalty for using a slightly nonoptimal sweep shape. We suggest that bats use logarithmic time modulation rather than linear period modulation in order to simplify signal processing during approach to prey, as this type of modulation stabilizes the filtering of echoes resulting from the bat’s automatic gain control. Likewise, the use of several harmonics in the emission may simplify signal processing of Doppler-shifted echoes by reducing the number of matched filters needed for reception.

Discrimination of wingbeat motion by bats, correlated with echolocation sound pattern

Bats of the species Rhinolophus rouxi, Hipposideros lankadiva and Eptesicus fuscus were trained to discriminate between two simultaneously presented artificial insect wingbeat targets moving at different wingbeat rates. During the discrimination trials, R. rouxi, H. lankadiva and E. fuscus emitted long-CF/FM, short-CF/FM and FM echolocation sounds respectively. R. rouxi, H. lankadiva and E. fuscus were able to discriminate a difference in wingbeat rate of 2.7 Hz, 9.2 Hz and 15.8 Hz, respectively, between two simultaneously presented targets at an absolute wingbeat rate of 60 Hz, using a criterion of 75% correct responses. The performance of the different bat species is correlated with the echolocation signal design used by each species, particularly with the presence and relative duration of a narrowband component preceding a broadband FM component. These results provide behavioral evidence supporting the hypothesis that bats that use CF/FM echolocation sounds have adaptations for the perception of insect wingbeat motion and that long-CF/FM species are more specialized for this task than short CF/FM species.

Audition in vampire bats, Desmodus rotundus

1. Within the tonotopic organization of the inferior colliculus two frequency ranges are well represented: a frequency range within that of the echolocation signals from 50 to 100 kHz, and a frequency band below that of the echolocation sounds, from 10 to 35 kHz. The frequency range between these two bands, from about 40 to 50 kHz is distinctly underrepresented (Fig. 3B). 2. Units with BFs in the lower frequency range (10–25 kHz) were most sensitive with thresholds of -5 to -11 dB SPL, and units with BFs within the frequency range of the echolocation signals had minimal thresholds around 0 dB SPL (Fig. 1). 3. In the medial part of the rostral inferior colliculus units were encountered which preferentially or exclusively responded to noise stimuli. — Seven neurons were found which were only excited by human breathing noises and not by pure tones, frequency modulated signals or various noise bands. These neurons were considered as a subspeciality of the larger sample of noise-sensitive neurons. — The maximal auditory sensitivity in the frequency range below that of echolocation, and the conspicuous existence of noise and breathing-noise sensitive units in the inferior colliculus are discussed in context with the foraging behavior of vampire bats.

Auditory adaptations for prey capture in echolocating bats

Auditory adaptations for prey capture in echolocating bats

A gating mechanism for sound pattern recognition is correlated with the temporal structure of echolocation sounds in the rufous horseshoe bat

The rufous horseshoe bat, Rhinolophus rouxi, was trained to discriminate differences in target distance. Loud free running artificial pulses, simulating the bat's natural long-CF/FM echolocation sounds, interfered with the ability of the bat to discriminate target distance. Interference occurred when the duration of the CF component of the CF/FM artificial pulse was between 2 and 70 ms. A brief (2.0 ms) CF signal 2–68 ms before an isolated FM signal was as effective as a continuous CF component of the same duration. When coupled with the bat's own emissions, a 2 ms FM sweep alone was effective in interfering when it came 42 to 69 ms after the onset of the bat's pulse. The coupled FM artificial pulses did not interfere when they began during the bat's own emissions.

Discrimination of fluttering targets by the FM-batPipistrellus stenopterus?

Five bats of the speciesPipistrellus stenopterus were trained in a two-alternative forced-choice procedure to discriminate between two fluttering targets. The positive target simulated an insect with a 50 Hz wingbeat rate. The negative target was varied between 0 and 48 Hz. The bats were able to discriminate a target with 41 Hz from a target with 50 Hz with 75% correct choices. In the discrimination task, they typically emitted echolocation calls of 2–4 ms duration sweeping from 60 kHz to 30 kHz. The duty cycle (i.e. fraction of time filled with echolocation sounds) increased when the targets fluttered, but was always lower than 3%. The performance ofP. stenopterus in discriminating fluttering targets is comparable to that of bats emitting longer sounds with constant-frequency (CF) components and a higher duty cycle. The FM-sounds ofP. stenopterus are short compared with the period of the fluttering targets, and therefore make it difficult for the animal to measure the time interval between two acoustic glints. Other cues may be prominent, such as the frequency modulation by Doppler shifts from the moving blades.

Foraging behaviour and echolocation in the rufous horseshoe bat (Rhinolophus rouxi) of Sri Lanka

In October 1984 foraging areas and foraging behaviour of the rufous horseshoe bat, Rhinolophus rouxi, were studied around a nursery colony on the hill slopes of Sri Lanka. The bats only foraged in dense forest and were not found in open woodlands (Fig. 1). This strongly supports the hypothesis that detection of fluttering prey is by pure tone echolocation within or close to echo-cluttering foliage. During a first activity period after sunset for about 30–60 min, the bats mainly caught insects on the wing. This was followed by a period of inactivity for another 60–120 min. Thereafter the bats resumed foraging throughout the night. They mainly alighted on specific twigs and foraged in flycatcher style. Individual bats maintained individual foraging areas of about 20x20 m. They stayed in this area throughout the night and returned to the same area on subsequent nights. Within this area the bats generally alighted on twigs at the same spots. Foraging areas were not defended against intruders. The bats echolocated throughout the night at an average repetition rate of 9.6±1.4 sounds/s. While hanging on twigs they scanned the surrounding area for flying prey by turning their bodies continuously around their legs. On average they performed one brief catching flight every 2 min and immediately returned to one of their favourite vantage points. Echolocation sounds may consist of up to three parts, a brief initial frequency-modulated (FM) component, a long constant frequency (CF) part lasting for about 40–50 ms, and a final FM part again (Fig. 4b, c). Adult males and females emitted pure tone frequencies in separate bands, the males from 73.5–77 kHz and the females from 76.5–79 kHz (Fig. 5). During scanning for prey from vantage points, the bats mostly emitted pure tones without any FM component (Fig. 4a). The last few pure tones emitted before take-off were prolonged to about 60 ms duration. The final FM part was therefore not an obligatory component of the echolocation signals in horseshoe bats. During flight and especially during emergence from the cave, most sounds consisted of a pure tone and loud initial and final FM sweeps. We therefore suggest that the initial FM part might also be relevant for echolocation. From our observations we conclude that the FM components are especially important during obstacle avoidance. In most sounds emitted in the field a fainter first harmonic was present. It was usually up to 30 dB fainter than the second harmonic, but in some instances it was as loud or even distinctly louder than the second one (Fig. 6a). Even within one sound the intensity relationship between the two harmonics may be reversed. We therefore suggest that the first harmonic is an integral part of the signal and relevant for information analysis in echolocation.

Influence of echolocation pulse rate on Doppler shift compensation control system in the greater horseshoe bat

1. Doppler shift compensation (DSC) behavior of the greater horseshoe bat,Rhinolophus ferrumequinum, was compared during spontaneous emission of echolocation sounds and during electrically elicited vocalization. 2. At low to moderate intensities of electrical stimulation in regions near the midbrain periaqueductal grey, the Doppler shift compensation performance showed the same characteristics as during spontaneous emission. This was also true for artificially increased emission rates (Fig. 1). 3. Electrically induced high sound emission rates improved the dynamic response properties of the Doppler shift compensation control system (Figs. 2, 3, 4). 4. At deep caudal stimulation sites in the midbrain electrical stimulation at moderate to high intensities led either to reversible suppression of DSC, or to reduction and distortion of the compensation (Fig. 5).

Accuracy of distance measurement in the bat Eptesicus fuscus: theoretical aspects and computer simulations.

Behavioral experiments of Simmons [J. Acoust. Soc. Am. 54, 157-173 (1973) and Science 204, 1336-1338 (1979)] on the ranging accuracy in the bat Eptesicus fuscus have led to far-reaching postulates on the existence of optimal and phase-conserving processing mechanisms in the bat. In this paper, the results of computer simulations of these experiments are presented. Two receiver types are investigated: the fully coherent cross-correlation receiver and the cross-correlation receiver with envelope processing (semicoherent). It is shown that Simmons' experiments cannot be treated as a simple estimation of distance, but require at least two (range difference experiment; see Simmons, 1973) or four (range jitter experiment; see Simmons, 1979) echolocation sounds for one decision. The performance of the bat in both experiments is much worse than predicted for a coherent and a semicoherent receiver type. The bat's accuracy in Simmons' range difference experiment is at least 18 dB worse than predicted for an optimal receiver. The results of the jitter experiment cannot be interpreted in a simple way as proof that bats are able to evaluate phase information as in a fully coherent cross-correlation receiver.

Control of echolocation pulses by neurons of the nucleus ambiguus in the rufous horseshoe bat, Rhinolophus rouxi. I. Single unit recordings in the ventral motor nucleus of the laryngeal nerves in spontaneously vocalizing bats.

The vocal motor control of the larynx was studied with single unit recordings from the efferent motor nucleus (nucleus ambiguus) in the CF-FM-bat Rhinolophus rouxi, spontaneously emitting echolocation sounds. The experiments were performed in a stereotaxic apparatus that allowed differentiation of activities in the recorded nucleus depending on the electrode position (Fig. 1). Echolocation calls and respiration activity were monitored simultaneously, thus it was possible to compare the time course of the motor control activity during respiration with and without concurrent vocalization. Unit discharges were classified as laryngeal motoneuron activity according to their correlation with the time course (onset and end) of echolocation calls and their discharge rate as: Pre-off-tonic, pre-off-phasic, off-pauser, off-tonic, on-chopper, on-tonic, prior-tonic and inhibitory (Fig. 4). The on-chopper and on-tonic discharge patterns were assigned to the motor activity of the lateral cricoarytenoid muscle and the off-pauser and off-tonic discharge patterns to the motor activity of the posterior cricoarytenoid muscle controlling the time course of vocal pulses. Motoneuron activities recorded under the condition of systematically shifted frequencies in the emitted echolocation calls were investigated in Doppler-shift compensating bats responding to electronically simulated echoes. Of all neurons classified as motor control, only units of the pre-off-tonic discharge type (cricothyroid muscle) changed their activity with frequency shifts in the vocalized pulses; they showed a positive linear correlation with the emitted sound frequency (Fig. 6). In addition, single unit activities in strict synchronization to vocalization were recorded, that by their low discharge rate were not valid as motor control, and were considered to represent activities of interneurons or internuclear neurons connecting the nucleus ambiguus with other vocalization- and respiration-centers (Fig. 3c). Electric lesions in the brain stem and iontophoretically applied horseradish peroxidase (HRP) served as references for localization and morphological identification of the recording sites in cell stained brain slices.

The detection of phantom targets in noise by serotine bats; negative evidence for the coherent receiver.

Using a target simulator three serotine bats, Eptesicus serotinus, were trained to judge whether a phantom target was present or absent. The echolocation sounds emitted by the bats during the detection were intercepted by a microphone, amplified and returned by a loudspeaker as an artificial echo, with a delay of 3.2 ms and a sound level determined by the overall gain and cry amplitude. The cry level of each pulse was measured and the echo level received by the bat was calculated. The target was presented in 50% of the trials and the gain adjusted using conventional up/down procedures. Under these conditions between 40 and 48 dB peSPL were required for 50% detection (Figs. 2, 3). In a subsequent experiment the phantom target was masked with white noise (No) with a spectrum level of -113 dB re. 1 Pa X Hz-1/2. The thresholds were increased by 7-14 dB. Energy density (S) of a single pulse was measured and used to estimate S/No, which ranged from 36-49 dB at threshold. Theoretically the coherent receiver model predicts the ratio between hits and false alarms observed for the bats at a S/No of ca. 1-2 dB. Since the bats require 40-50 dB higher S/No (Fig. 3), this is taken as negative evidence for coherent reception (cross correlation). Furthermore, a strong sensitivity to clutter was found since there seemed to exist a fixed relationship between thresholds and clutter level.

Frequency tracking and Doppler shift compensation in response to an artificial CF/FM echolocation sound in the CF/FM bat,Noctilio albiventris

Bats of the speciesNoctilio albiventris emit short-constant frequency/frequency modulated (short-CF/FM) pulses with a CF component frequency at about 75 kHz. Bats sitting on a stationary platform were trained to discriminate target distance by means of echolocation. Loud, free-running artificial pulses, simulating the bat's natural CF/FM echolocation sounds or with systematic modifications in the frequency of the sounds, were presented to the bats during the discrimination trials. When the CF component of the artificial CF/FM sound was between 72 and 77 kHz, the bats shifted the frequency of the CF component of their own echolocation sounds toward that of the artificial pulse, tracking the frequency of the artificial CF component.

Echolocation sounds and hearing of the greater Japanese horseshoe bat (Rhinolophus ferrumequinum nippon)

The relationship between the orientation sounds and hearing sensitivity in the greater Japanese horseshoe bat,Rhinolophus ferrumequinum nippon was studied.

Automatic gain control in the bat's sonar receiver and the neuroethology of echolocation

The sensitivity of the echolocating bat, Eptesicus fuscus, to sonar echoes at different time delays after sonar emissions was measured in a two-choice echo detection experiment. Since echo delay is perceptually equivalent to target range, the experiment effectively measured sensitivity to targets at different ranges. The bat's threshold for detecting sonar echoes at a short delay of only 1.0 msec after emissions (corresponding to a range of 17 cm) was 36 dB SPL (peak to peak), but the threshold decreased to 8 dB SPL at a longer delay of 6.4 msec (a range of 1.1 m). Prior research has shown that, at even longer delays (corresponding to ranges of 3 to 5 m), the bat's threshold is in the region of 0 dB SPL. Contractions of the bat's middle ear muscles synchronized with the production of echolocation sounds cause a transient loss in hearing sensitivity which appears to account for the observed echo detection threshold shifts. The bat's echo detection thresholds increase by approximately 11 dB for each reduction in target range by a factor of 2 over the span from 17 cm to 1.1 m. As range shortens, the amplitude of echoes from small targets also increases, by 12 dB for each 2-fold reduction in range. Thus, when approaching a target, the bat compensates for changes in echo strength as target range shortens by changing its hearing threshold. Since this compensation appears to occur in the middle ear, the bat regulates echoes reaching the cochlea to a stable amplitude during its approach to a target such as a flying insect. In addition to this automatic gain control linked to target range, the bat aims its head to track a target's position during approach, thus stabilizing echo amplitude even if the target's direction changes. We hypothesize that the bat's directional emissions, directional hearing, middle ear muscle contractions, and head aim response collectively create a three-dimensional spatial tracking filter which the bat locks onto targets to stabilize echo amplitudes during interception of prey. We further hypothesize that this regulation, which cancels echo amplitude changes caused by the target's changing spatial position, leaves the bat free to observe echo amplitude changes caused by the target's own actions, such as insect wing beats. Elimination of spatially dependent echo amplitude changes removes the cause of potentially troublesome changes in neural response latency and keeps stimulation from echoes in the "tip" region of auditory nerve fiber tuning curves.(ABSTRACT TRUNCATED AT 400 WORDS)

Echolocation and hearing in the mouse-tailed bat,Rhinopoma hardwickei: acoustic evolution of echolocation in bats

1. Mouse-tailed bats (Rhinopoma hardwickei) use short-duration, multiple-harmonic, constant-frequency (CF) or slightly frequency-modulated (FM) signals for echolocation of flying insect prey. The frequency components are 18–20 kHz (first harmonic), 36–40 kHz (second harmonic), 56–60 kHz (third harmonic), and 75–80 kHz (fourth harmonic). The second harmonic is the strongest component, with the third harmonic 2 to 10 dB weaker, and the first and fourth harmonics about 10 to 20 dB weaker than the second. 2. The bat's hearing, as indicated by N4 auditory evoked potentials, is moderately sharply tuned to the second harmonic, broadly sensitive to the first harmonic and to lower frequencies, and moderately sensitive to the third and fourth harmonics. The degree of tuning is sufficient to indicate some specialized function for the second harmonic, perhaps in the task of detecting targets at maximum range, since this is the lowest of the strong harmonics and least affected by atmospheric attenuation. These bats probably do not perform the Doppler compensation response even though their hearing is tuned. 3. The pattern of emission of sonar sounds (repetition rate, duration) during interception of prey is similar to that observed in most other species of bats. The signals are emitted at repetition rates of 10–20/s during the search stage, 20–40/s during the approach stage, and about 100/s during the terminal stage of the pursuit process. The sonar sounds emitted during the search stage are 6–10 ms long, during the approach stage they are 2–4 ms long, and during the terminal stage they are less than 1 ms long. 4. The principal acoustic dimension of echolocation sounds that relates to perception of targets is signal bandwidth, whichRhinopoma hardwickei manipulates throughout the pursuit process by shortening the duration of its sonar sounds and also by slightly broadening the FM sweeps in its terminal-stage sounds. The onset-time of the long, search-stage sounds and the longer, early approach-stage sounds is more abrupt than for the shorter, late approach-stage and the short terminal-stage sounds. This apparently deliberate transient beginning broadens the signal's bandwidth at the onset relative to the narrower bandwidth prevailing for the rest of the CF harmonic structure which follows. In the shorter sounds, which are emitted during the late approach stage and terminal stage, the onset is more gradual, but the duration (the envelope) and the offset-time are now short enough that the bandwidth of the signal as a whole is increased. The bandwidth is manipulated primarily by these changes in signal envelope and secondarily by the increased FM sweep-width in the terminal stage. 5. Except for duration these signals are relatively inflexible and suggestive of a primitive kind of echolocation in which only one dimension is changed to achieve qualities which most other species of bats obtain by changing a variety of signal dimensions simultaneously. The abrupt signal onsets may be an indication that multiple-harmonic CF and FM echolocation sounds evolved from click-like echolocation sounds of the type emitted byRousettus and some terrestrial mammals.

Audiograms of a South Indian bat community

1. Audiograms are recorded from one non-echolocating and nine echolocating sympatrically living bat species of South India. These species areCynopterus sphinx (non-echolocating),Tadarida aegyptiaca, Taphozous melanopogon, T. kachhensis, Rhinopoma hardwickei, Pipistrellus dormeri, P. mimus, Hipposideros speoris, H. bicolor andMegaderma lyra. 2. InRhinopoma hardwickei a highly sensitive frequency range was found which is narrowly tuned to the frequency band of the bat's CF-echolocation call (32–35 kHz, Fig. 3). In hipposiderids a ‘filter’ narrowly tuned to the frequency of the CF-part of the CF-FM echolocation sounds (137.5 kHz inH. speoris and 151.5 kHz inH. bicolor, Fig. 5) could be recorded from deeper parts of IC. 3. In the echolocating species the best frequency of the audiograms closely matched with that frequency range in the echolocation calls containing most energy. 4. In bat species foraging flying prey best frequencies of audiograms and height of preferred foraging areas are inversely related, i.e. bat species hunting high above canopy have lower best frequencies than those foraging close to or within canopy (Fig. 6). 5. A hypothesis is forwarded explaining how fluttering target detection by constant frequency echolocation might have evolved from long distance echolocation by pure tone signals.

Tonotopical organization and pure tone response characteristics of single units in the auditory cortex of the Greater Horseshoe Bat

Tonotopical organization and frequency representation in the auditory cortex of Greater Horseshoe Bats was studied using multi-unit recordings. The auditory responsive cortical area can be divided into a primary and a secondary region on the basis of response characteristics forming a core/belt structure. In the primary area units with best frequencies in the range of echolocation signals are strongly overrepresented (Figs. 6–8). There are two separate large areas concerned with the processing of the two components of the echolocation signals. In one area frequencies between the individual resting frequency and about 2 kHz above are represented, which normally occur in the constant frequency (CF) part of the echoes (CF-area), in a second one best frequencies between resting frequency and about 8 kHz below are found (FM-area). In the CF-area tonotopical organization differs from the usual mammalian scheme of dorso-ventral isofrequency slabs. Here isofrequency contours are arranged in a semicircular pattern. The representation of the cochlear partition (cochleotopic organization) was calculated. In the inferior colliculus and auditory cortex there is a disproportionate representation of the basilar membrane. This finding is in contradiction to the current opinion that frequency representation in the auditory system of Horseshoe Bats is only determined by the mechanical tuning properties of the basilar membrane. Response characteristics for single units were studied using pure tone stimuli. Most units showed transient responses. In 25% of units response characteristics depended on the combination of frequency and sound pressure level used. Frequency selectivity of units with best frequencies in the range of echolocation sounds is very high. Q-10dB values of up to 400 were found in a small frequency band just above resting frequency.

Auditory representation in the cerebellum of the CF-FM bat, Pteronotus parnellii parnellii

Auditory representation in the cerebellum of the mustache bat, Pteronotus parnellii parnellii, was studied with microelectrode mapping technique. This study reveals that a large area of the bat's cerebellum contains units responding to acoustic stimuli. A study of the frequency representation and frequency tuning of the recorded units show that auditory specialization for processing of species-specific echolocation sounds also exists in its cerebellum. There is a large number of units with best frequencies between 60 and 64 kHz. These units have extremely sharp threshold curves with a Q10-dB value as high as 394. These auditory units are organized into a fractured pattern of small patches within which units are tonotopically organized. The units also appear to be columnarly assembled according to their best frequencies. Although different in sensory modality, such a finding is highly comparable to the fractured somatotopy in the tactile areas of cerebellar hemispheres [Shambes et al. Brain Behav. Evol. 15, 94–140 (1975)] [Work supported by NSF 80-07348 and USPH 1-K04-NS00433 to P. Jen.]