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J W Schnupp

Publications and source records attributed to J W Schnupp.

7 recordsLinked to original sources

Effects of altering spectral cues in infancy on horizontal and vertical sound localization by adult ferrets.

We investigated the behavioral consequences of removing the pinna and concha of the external ear bilaterally in infancy on the sound localization ability of adult ferrets. Altering spectral cues in this manner has previously been shown to disrupt the development of the neural representation of auditory space in the superior colliculus. Using broadband noise stimuli, we tested pinnae-removed ferrets and normal ferrets in three sound localization tasks. In each case, we found that both groups of animals performed significantly better when longer duration noise bursts were used. In a relative localization task, we measured the acuity with which the ferrets could discriminate between two speakers in the horizontal plane. The speakers were placed symmetrically either around the anterior midline or around a position 45 degrees lateral to the midline. In this task, the pinnae-removed ferrets achieved very similar scores to the normal ferrets. By contrast, in another relative localization task that measured localization ability in the midsagittal plane, pinnae-removed ferrets performed less well than normals. In an absolute localization task, 12 speakers were spaced at 30 degrees intervals in the horizontal plane at the level of the ferrets' ears. Overall, the pinnae-removed ferrets also performed poorly in this task compared with normal ferrets: they made significantly fewer correct responses, larger localization errors and more front-back errors. Both normal and pinnae-removed animals showed an improvement in performance with practice, although the pattern of improvement differed for each group. The largest improvements in localization accuracy were achieved by the pinnae-removed ferrets, particularly at the frontal positions, and their performance eventually approached that of the normal animals. Nevertheless, some intergroup differences were still present. In particular, the pinnae-removed ferrets continued to make significantly more front-back errors than the normals. These deficits can be attributed to differences in the spectral localization cues available to the animals. Acoustical measurements showed that, compared with normal animals, the head-related transfer functions in the horizontal plane were largely ambiguous around the interaural axis and also contained fewer location-dependent features in the midsagittal plane.

Acoustic Stimulation↗

Signals from the superficial layers of the superior colliculus enable the development of the auditory space map in the deeper layers.

We have examined whether the superficial layers of the superior colliculus (SC) provide the source of visual signals that guide the development of the auditory space map in the deeper layers. Anatomical tracing experiments with fluorescent microspheres revealed that a retinotopic map is present in the newborn ferret SC. Aspiration of the caudal region of the superficial layers of the right SC on postnatal day 0 did not cause a reorganization of this projection. Consequently, recordings made when the animals were mature showed that visual units in the remaining superficial layers in rostral SC had receptive fields that spanned a restricted region of anterior space. Auditory units recorded beneath the remaining superficial layers were tuned to corresponding anterior locations. Both the superficial layer visual map and the deeper layer auditory map were normal in the left, unoperated SC. The majority of auditory units recorded throughout the deeper layers ventral to the superficial layer lesion were also tuned to single sound directions. In this region of the SC, however, we observed much greater scatter in the distribution of preferred sound directions and a significant increase in the proportion of units with spatially ambiguous responses. The auditory representation was degraded, although many of these units were also visually responsive. Equivalent lesions of the superficial layers made in adult ferrets did not alter the topographic order in the auditory representation, suggesting that visual activity in these layers may be involved in aligning the different sensory maps in the developing SC.

Age Factors↗

Sensory neuroscience: visualizing the auditory cortex.

Recent studies suggest that the auditory cortex may use sensory processing strategies analogous to those already established for the visual cortex. Nevertheless, fundamental differences in the way the visual and auditory worlds are structured have to be borne in mind.

Auditory Cortex↗

Altered spectral localization cues disrupt the development of the auditory space map in the superior colliculus of the ferret.

Spectral localization cues provided by the outer ear are utilized in the construction of the auditory space map in the superior colliculus (SC). The role of the outer ear in the development of this map was examined by recording from the SC of anesthetized, adult ferrets in which the pinna and concha had been removed in infancy. The acoustical consequences of this procedure were assessed by recording outer ear impulse responses via a probe-tube microphone implanted in the wall of the ear canal. Both monaural and binaural spectral cues normally show a number of asymmetric features within the horizontal plane, which allow azimuthal locations on either side of the interaural axis to be discriminated. These features were eliminated or altered by chronic pinnectomy. The responses of auditory units in the SC to noise bursts presented in the free field were examined at sound levels of approximately 10 and 25 dB above unit threshold. After bilateral pinnectomy, the representation of auditory space was severely degraded at both sound levels. In contrast to normal ferrets, many units had bilobed azimuthal response profiles, indicating that they were unable to resolve sound locations on either side of the interaural axis. There was also much less order in the distribution of best azimuths or elevations of those units that were tuned to a single direction. Some units were tuned to locations that extended much further into the hemifield ipsilateral to the recording side than the normal range of best azimuths. Unilateral removal of the outer ear, which disrupts the monaural spectral cues for one side only, had a much smaller effect on the development of the auditory representation. At supra- and near-threshold sound levels, the representation of sound azimuth in the SC on both sides of the brain was less scattered than that found after bilateral pinna removal. Nevertheless, units with bilobed responses, broader tuning, and inappropriate best azimuths were observed in both the left and right SC of ferrets in which the left pinna and concha had been removed in infancy. These data illustrate that the localization cues provided by the outer ear play a critical role in the development of the auditory space map in the SC. In contrast to other manipulations of either auditory or visual inputs, the map does not appear to adapt to the changes in spectral cues brought about by pinna removal, suggesting that residual binaural cues are, by themselves, insufficient for its normal maturation.

Acoustic Stimulation↗

Coding for auditory space in the nucleus of the brachium of the inferior colliculus in the ferret.

Coding for auditory space in the nucleus of the brachium of the inferior colliculus in the ferret. J. Neurophysiol. 78: 2717-2731, 1997. The nucleus of the brachium of the inferior colliculus (BIN) projects topographically to the deeper layers of the superior colliculus (SC), which contain a two-dimensional map of auditory space. In this study, we have used broadband stimuli presented in the free field to investigate how auditory space is represented in the BIN of the ferret. Response latencies and temporal firing patterns were comparable with those in the SC, and both properties showed some variation with stimulus location. We obtained spatial response profiles at two sound levels (5-15 and 25-35 dB above unit threshold). A large proportion of azimuth profiles (41% in the suprathreshold condition, 80% in the near-threshold condition) presented a single peak, indicating that they were tuned to single regions in space. For some of these units, the preferred speaker position varied considerably with sound level. The remaining units showed predominantly either broad "hemifield" or spatially ambiguous "bilobed" response profiles. At suprathreshold sound levels, the preferred azimuths of the tuned cells were ordered topographically along the rostrocaudal axis of the BIN, although this representation is considerably more scattered than that in the SC. In contrast to the SC, we observed no systematic variation in the distribution of near-threshold best azimuths, which were instead concentrated around the interaural axis in the contralateral hemifield. The azimuth tuning of individual units in the BIN was generally broader at both sound levels than that in the SC. Many units also were tuned for the elevation of the sound source (48% for supra-, 77% for near-threshold stimulation), but there was no evidence for topographic order in the distribution of preferred elevations within the BIN. These results suggest that the BIN sends inputs to the SC that are already selective for sound azimuth and elevation and that show some degree of topographic order for sound azimuth. These inputs then presumably are sharpened and their topography refined by a mechanism that is likely to involve convergence of other inputs and activity-dependent fine tuning of terminal connections, to result in a precise two-dimensional map of auditory space in the SC.

Acoustic Stimulation↗

The development of topographically-aligned maps of visual and auditory space in the superior colliculus.

The role of the superior colliculus in attending and orienting to sensory stimuli is facilitated by the presence within this midbrain nucleus of superimposed maps of different sensory modalities. We have studied the steps involved in the development of topographically-aligned maps of visual and auditory space in the ferret superior colliculus. Injections of fluorescent beads into the superficial layers showed that the projection from the contralateral retina displays topographic order on the day of birth (PO). Recordings made from these layers at the time of eye opening, approximately 1 month later, revealed the presence of an adult-like map of visual space. In contrast, the auditory space map in the deeper layers emerged gradually over a much longer period of postnatal life. In adult ferrets in which one eye had been deviated laterally just before eye opening, the auditory spatial tuning of single units recorded in the contralateral superior colliculus was shifted by a corresponding amount, so that the registration of the visual and auditory maps was maintained. Chronic application of the NMDA-receptor antagonist MK801 disrupted the normal development of the auditory space map, but had no effect on the visual map in either juvenile or adult animals, or on the auditory map once it had matured. These findings indicate that visual cues may play an instructive role, possibly via a Hebbian mechanism of synaptic plasticity, in the development of appropriately tuned auditory responses, thereby ensuring that the neural representations of both modalities share the same coordinates. Changes observed in the auditory representation following partial lesions of the superficial layers at PO suggest that these layers may provide the source of the visual signals responsible for experience-induced plasticity in auditory spatial tuning.

Animals↗

NMDA-receptor antagonists disrupt the formation of the auditory space map in the mammalian superior colliculus.

In the ferret (Mustela putorius) the map of auditory space in the deeper layers of the superior colliculus (SC) matures over a period of several postnatal weeks, a process known to be guided by both visual and auditory experience. The auditory responses are initially very broadly tuned, and gradually become more selective for specific sound locations that coincide with the visual receptive fields recorded in the same region of the SC. To investigate the possible involvement of NMDA-type glutamate receptors in the postnatal development of this auditory representation, we have reared ferrets in which 400 microns thick sheets of the slow-release polymer Elvax, containing the NMDA receptor antagonists MK801 or APV, were placed on the dorsal surface of the SC. The Elvax was implanted on postnatal day (P) 25-27, just before the onset of hearing, and removed 5-6 weeks later, just prior to recording from the SC on around P61-70. In vitro measurements with Elvax containing 3H-MK801 revealed that the amount of drug released declined sharply over the first 10 d and then stabilized at a fairly constant rate for the following 5 weeks. These in vitro data were found to parallel the in vivo release of MK801 from implanted Elvax slices. Diffusion of MK801 from the implant was measured and significant levels were found within 800 microns of the SC surface, suggesting that the action of MK801 was restricted to the superficial and intermediate layers of the nucleus. Extracellular recordings were made from visual and auditory units in the SC in response to free-field stimulation. The visual responses of units recorded in the superficial layers appeared to be unaffected by either of the drug treatments, and formed a normal, adult-like map of visual azimuth along the rostrocaudal axis of the SC in all animals. Most of the auditory single-unit responses recorded at this age in normal, unoperated controls were spatially tuned and topographically organized, although the map of sound azimuth was less precise than that in adult ferrets. Data from age-matched control animals that had been reared with drug-free Elvax implants were not statistically different from the unoperated juvenile ferrets. However, in animals reared with APV or MK801 Elvax implants, there was an increase in the relative numbers of auditory units that were ambiguously tuned to two or more locations.(ABSTRACT TRUNCATED AT 400 WORDS)

2-Amino-5-phosphonovalerate↗