Behavior and modeling of two-dimensional precedence effect in head-unrestrained cats.

The precedence effect (PE) is an auditory illusion that occurs when listeners localize nearly coincident and similar sounds from different spatial locations, such as a direct sound and its echo. It has mostly been studied in humans and animals with immobile heads in the horizontal plane; speaker pairs were often symmetrically located in the frontal hemifield. The present study examined the PE in head-unrestrained cats for a variety of paired-sound conditions along the horizontal, vertical, and diagonal axes. Cats were trained with operant conditioning to direct their gaze to the perceived sound location. Stereotypical PE-like behaviors were observed for speaker pairs placed in azimuth or diagonally in the frontal hemifield as the interstimulus delay was varied. For speaker pairs in the median sagittal plane, no clear PE-like behavior occurred. Interestingly, when speakers were placed diagonally in front of the cat, certain PE-like behavior emerged along the vertical dimension. However, PE-like behavior was not observed when both speakers were located in the left hemifield. A Hodgkin-Huxley model was used to simulate responses of neurons in the medial superior olive (MSO) to sound pairs in azimuth. The novel simulation incorporated a low-threshold potassium current and frequency mismatches to generate internal delays. The model exhibited distinct PE-like behavior, such as summing localization and localization dominance. The simulation indicated that certain encoding of the PE could have occurred before information reaches the inferior colliculus, and MSO neurons with binaural inputs having mismatched characteristic frequencies may play an important role.

Dynamic sound localization in cats.

Sound localization in cats and humans relies on head-centered acoustic cues. Studies have shown that humans are able to localize sounds during rapid head movements that are directed toward the target or other objects of interest. We studied whether cats are able to utilize similar dynamic acoustic cues to localize acoustic targets delivered during rapid eye-head gaze shifts. We trained cats with visual-auditory two-step tasks in which we presented a brief sound burst during saccadic eye-head gaze shifts toward a prior visual target. No consistent or significant differences in accuracy or precision were found between this dynamic task (2-step saccade) and the comparable static task (single saccade when the head is stable) in either horizontal or vertical direction. Cats appear to be able to process dynamic auditory cues and execute complex motor adjustments to accurately localize auditory targets during rapid eye-head gaze shifts.

Simultaneous comparison of two sound localization measures.

Almost all behavioral studies of sound localization have used either an approach-to-target or pointing/orienting task to assess absolute sound localization performance, yet there are very few direct comparisons of these measures. In an approach-to-target task, the subject is trained to walk to a sound source from a fixed location. In an orienting task, finger, head and/or eye movements are monitored while the subject's body is typically constrained. The fact that subjects may also initiate head and eye movements toward the target during the approach-to-target task allows us to measure the accuracy of the initial orienting response and compare it with subsequent target selection. To perform this comparison, we trained cats to localize a broadband noise presented randomly from one of four speakers located ± 30° and ± 60° in azimuth. The cat responded to each sound presentation by walking to and pressing a lever at the perceived location, and a food reward was delivered if the first attempt was correct. In tandem, we recorded initial head and eye orienting movements, via magnetic search coils, immediately following target onset and prior to the walking response. Reducing either stimulus duration or level resulted in a systematic decline in both measurements of localization performance. When the task was easy, localization performance was accurate for both measures. When the task was more difficult, the number of incorrect (i.e., wrong selection) and no-go (i.e., no selection) responses increased. Interestingly, for many of the incorrect trials, there was a dissociation between the orienting response and the target selected, and for many of the no-go trials, the gaze oriented towards the correct target even though the cat did not move to it. This suggests different neural systems governing walking to a target as compared to unconditioned gaze orienting.

Localization of click trains and speech by cats: the negative level effect.

Although localization of sound in elevation is believed to depend on spectral cues, it has been shown with human listeners that the temporal features of sound can also greatly affect localization performance. Of particular interest is a phenomenon known as the negative level effect, which describes the deterioration of localization ability in elevation with increasing sound level and is observed only with impulsive or short-duration sound. The present study uses the gaze positions of domestic cats as measures of perceived locations of sound targets varying in azimuth and elevation. The effects of sound level on localization in terms of accuracy, precision, and response latency were tested for sound with different temporal features, such as a click train, a single click, a continuous sound that had the same frequency spectrum of the click train, and speech segments. In agreement with previous human studies, negative level effects were only observed with click-like stimuli and only in elevation. In fact, localization of speech sounds in elevation benefited significantly when the sound level increased. Our findings indicate that the temporal continuity of a sound can affect the frequency analysis performed by the auditory system, and the variation in the frequency spectrum contained in speech sound does not interfere much with the spectral coding for its location in elevation.

Effects of forward masking on sound localization in cats: basic findings with broadband maskers.

Forward masking is traditionally measured with a detection task in which the addition of a preceding masking sound results in an increased signal-detection threshold. Little is known about the influence of forward masking on localization of free-field sound for human or animal subjects. Here we recorded gaze shifts of two head-unrestrained cats during localization using a search-coil technique. A broadband (BB) noise masker was presented straight ahead. A brief signal could come from 1 of the 17 speaker locations in the frontal hemifield. The signal was either a BB or a band-limited (BL) noise. For BB targets, the presence of the forward masker reduced localization accuracy at almost all target levels (20 to 80 dB SPL) along both horizontal and vertical dimensions. Temporal decay of masking was observed when a 15-ms interstimulus gap was added between the end of the masker and the beginning of the target. A large effect of forward masking was also observed for BL targets with low (0.2-2 kHz) and mid (2-7 kHz) frequencies, indicating that the interaural timing cue is susceptible to forward masking. Except at low sound levels, a small or little effect was observed for high-frequency (7-15 kHz) targets, indicating that the interaural level and the spectral cues in that frequency range remained relatively robust. Our findings suggest that different localization mechanisms can operate independently in a complex listening environment.

Gaze shifts to auditory and visual stimuli in cats.

While much is known about the metrics and kinematics of gaze shifts to visual targets in cats, little is known about gaze shifts to auditory targets. Here, cats were trained to localize auditory and visual targets via gaze shifts. Five properties of gaze shifts to sounds were observed. First, gaze shifts were accomplished primarily by large head movements. Unlike primates, the head movement in cats often preceded eye movement though the relative timing of eye in head and head latencies depended upon the target modality and gaze shift magnitude. Second, gaze shift latencies to auditory targets tended to be shorter than equivalent shifts to visual targets for some conditions. Third, the main sequences relating gaze amplitude to maximum gaze velocity for auditory and visual targets were comparable. However, head movements to auditory and visual targets were less consistent than gaze shifts and tended to undershoot the targets by 30 % for both modalities. Fourth, at the end of gaze movement, the proportion of the gaze shift accomplished by the eye-in-head movement was greater to visual than auditory targets. On the other hand, at the end of head movement, the proportion of the gaze shift accomplished by the head was greater to auditory than visual targets. Finally, gaze shifts to long-duration auditory targets were accurate and precise and were similar to accuracy of gaze shifts to long-duration visual targets. Because the metrics of gaze shifts to visual and auditory targets are nearly equivalent, as well as their accuracy, we conclude that both sensorimotor tasks use primarily the same neural substrates for the execution of movement.

Behavioral and modeling studies of sound localization in cats: effects of stimulus level and duration.

Sound localization accuracy in elevation can be affected by sound spectrum alteration. Correspondingly, any stimulus manipulation that causes a change in the peripheral representation of the spectrum may degrade localization ability in elevation. The present study examined the influence of sound duration and level on localization performance in cats with the head unrestrained. Two cats were trained using operant conditioning to indicate the apparent location of a sound via gaze shift, which was measured with a search-coil technique. Overall, neither sound level nor duration had a notable effect on localization accuracy in azimuth, except at near-threshold levels. In contrast, localization accuracy in elevation improved as sound duration increased, and sound level also had a large effect on localization in elevation. For short-duration noise, the performance peaked at intermediate levels and deteriorated at low and high levels; for long-duration noise, this "negative level effect" at high levels was not observed. Simulations based on an auditory nerve model were used to explain the above observations and to test several hypotheses. Our results indicated that neither the flatness of sound spectrum (before the sound reaches the inner ear) nor the peripheral adaptation influences spectral coding at the periphery for localization in elevation, whereas neural computation that relies on "multiple looks" of the spectral analysis is critical in explaining the effect of sound duration, but not level. The release of negative level effect observed for long-duration sound could not be explained at the periphery and, therefore, is likely a result of processing at higher centers.

The role of spectral composition of sounds on the localization of sound sources by cats.

Sound localization along the azimuthal dimension depends on interaural time and level disparities, whereas localization in elevation depends on broadband power spectra resulting from the filtering properties of the head and pinnae. We trained cats with their heads unrestrained, using operant conditioning to indicate the apparent locations of sounds via gaze shift. Targets consisted of broadband (BB), high-pass (HP), or low-pass (LP) noise, tones from 0.5 to 14 kHz, and 1/6 octave narrow-band (NB) noise with center frequencies ranging from 6 to 16 kHz. For each sound type, localization performance was summarized by the slope of the regression relating actual gaze shift to desired gaze shift. Overall localization accuracy for BB noise was comparable in azimuth and in elevation but was markedly better in azimuth than in elevation for sounds with limited spectra. Gaze shifts to targets in azimuth were most accurate to BB, less accurate for HP, LP, and NB sounds, and considerably less accurate for tones. In elevation, cats were most accurate in localizing BB, somewhat less accurate to HP, and less yet to LP noise (although still with slopes ∼0.60), but they localized NB noise much worse and were unable to localize tones. Deterioration of localization as bandwidth narrows is consistent with the hypothesis that spectral information is critical for sound localization in elevation. For NB noise or tones in elevation, unlike humans, most cats did not have unique responses at different frequencies, and some appeared to respond with a "default" location at all frequencies.

The vestibulo-auricular reflex.

The mammalian orienting response to sounds consists of a gaze shift that can be a combination of head and eye movements. In animals with mobile pinnae, the ears also move. During head movements, vision is stabilized by compensatory rotations of the eyeball within the head because of the vestibulo-ocular reflex (VOR). While studying the gaze shifts made by cats to sounds, a previously uncharacterized compensatory movement was discovered. The pinnae exhibited short-latency, goal-directed movements that reached their target while the head was still moving. The pinnae maintained a fixed position in space by counter-rotating on the head with an equal but opposite velocity to the head movement. We call these compensatory ear movements the vestibulo-auricular reflex (VAR) because they shared many kinematic characteristics with the VOR. Control experiments ruled out efference copy of head position signals and acoustic tracking (audiokinetic) of the source as the cause of the response. The VAR may serve to stabilize the auditory world during head movements.

Sound-localization performance in the cat: the effect of restraining the head.

In oculomotor research, there are two common methods by which the apparent location of visual and/or auditory targets are measured, saccadic eye movements with the head restrained and gaze shifts (combined saccades and head movements) with the head unrestrained. Because cats have a small oculomotor range (approximately +/-25 degrees), head movements are necessary when orienting to targets at the extremes of or outside this range. Here we tested the hypothesis that the accuracy of localizing auditory and visual targets using more ethologically natural head-unrestrained gaze shifts would be superior to head-restrained eye saccades. The effect of stimulus duration on localization accuracy was also investigated. Three cats were trained using operant conditioning with their heads initially restrained to indicate the location of auditory and visual targets via eye position. Long-duration visual targets were localized accurately with little error, but the locations of short-duration visual and both long- and short-duration auditory targets were markedly underestimated. With the head unrestrained, localization accuracy improved substantially for all stimuli and all durations. While the improvement for long-duration stimuli with the head unrestrained might be expected given that dynamic sensory cues were available during the gaze shifts and the lack of a memory component, surprisingly, the improvement was greatest for the auditory and visual stimuli with the shortest durations, where the stimuli were extinguished prior to the onset of the eye or head movement. The underestimation of auditory targets with the head restrained is explained in terms of the unnatural sensorimotor conditions that likely result during head restraint.

Medial pontine hemorrhagic stroke.

BACKGROUND AND PURPOSE: This case report documents a rare opportunity to observe the motor function of an individual for nearly 6 months following a primary pontine hemorrhage in the medial pontine tegmentum of the brain stem. The purpose of this report is to illustrate how knowledge of the location of the hemorrhage, in conjunction with knowledge of brain-stem structure-function relationships, informs physical therapist examination and intervention. CASE DESCRIPTION: RM, a right-handed 81-year-old man with hypertension, had a hemorrhagic brain-stem stroke that severely compromised control of posture and whole-limb movements. Some residual ability to use the right hand and fingers remained, provided the trunk and right upper arm were stabilized. RM had undiminished intellectual abilities and unaltered memory because of sparing of cerebral cortices. RM's cognitive abilities, however, were obscured by severe impairments in interpersonal communication because of extensive damage to cranial nerve structures. Computed tomographic scans verified that the hematoma crossed the midline and was confined to the medial pontine tegmentum. DISCUSSION: We interpret motor deficits resulting from stoke in the medial pontine tegmentum in terms of damage to brain-stem descending motor systems and ascending somatosensory systems. Recognition of cognitive and residual motor abilities following brain-stem stroke can aid in the development of rehabilitation strategies.