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W B Warr

Publications and source records attributed to W B Warr.

17 recordsLinked to original sources

Changes in spontaneous otoacoustic emissions produced by acoustic stimulation of the contralateral ear.

Spontaneous otoacoustic emissions (SOAEs) were measured in human ear canals before, during and after presentation of tonal stimuli to the contralateral ear. Stimuli were presented in 1/8 octave steps from 2 octaves below to 1 octave above the SOAE frequency at sound levels below the observed contralateral acoustic reflex threshold. For certain conditions there was an abrupt upward frequency shift at stimulus onset. For a fixed level the effect was frequency selective; the maximum frequency shift was obtained with tones approximately 1/2 octave below the SOAE. SOAE amplitude usually decreased but in some cases increased or remained unchanged. When amplitude changes were observed, the maximum shifts were observed for tones at or near the SOAE frequency. Changes in SOAEs were not observed for stimulus levels below 60 dB SPL. The effect is believed to be mediated by medial efferent neurons of the uncrossed olivocochlear bundle which arise in the medial region of the superior olivary complex and terminate on outer hair cells (OHCs). These results support those models which attribute SOAE generation to OHCs, and are indicative of an efferent influence on cochlear mechanics. A simple model is presented that proposes that efferent activity alters the tuning of the emission generator by causing changes in OHC membrane conductance.

Acoustic Stimulation

Descending projections from the superior olivary complex to the cochlear nucleus of the cat.

Subdivisions of the cochlear nuclear complex give rise to a number of discrete projections to certain cell groups of the superior olivary complex and also received substantial descending projections from the periolivary nuclei. In the present study, we sought to determine by means of retrograde transport of horseradish peroxidase (HRP), and anterograde transport of radiolabeled protein, if the periolivary nuclei give rise to discrete projections to the various subdivisions of the cochlear nuclear complex. Following medium to large injections of HRP into the cochlear nucleus, irrespective of location, labeled cells were found in all periolivary nuclei bilaterally. In every case more than 40% of the labeled cells were found in the lateral nucleus of the trapezoid body on the same side and the ventral nucleus of the trapezoid body of both sides. Other periolivary nuclei contributing more than 5% of the total number of cells in individual cases were the contralateral lateral nucleus of the trapezoid body and the ipsilateral anterolateral and dorsal periolivary nuclei. Injections of tritiated leucine into periolivary nuclei gave rise to axonal labeling to the trapezoid body and the dorsal acoustic stria, usually bilaterally, and to terminal labeling that was widely distributed within the cochlear nuclear complex. In several cases with small injections, particularly in the lateral nucleus of the trapezoid body, the projections from the periolivary nuclei to the anteroventral and dorsal cochlear nuclei connected areas described as having similar best-frequency representation. The autoradiographic data corroborated the main results from the HRP experiments and provided additional information permitting these conclusions: the projections from the periolivary nuclei to the cochlear nuclear complex are organized tonotopically, at least in part; each periolivary nucleus (and perhaps individual cells), projects widely throughout the cochlear nuclear complex; the pattern of termination of projections from different periolivary nuclei to a given region of the cochlear nuclear complex are similar, as seen in autoradiograms, and the lateral and dorsal periolivary nuclei project mainly ipsilaterally, while the medial periolivary nuclei project bilaterally with a contralateral bias. The magnitude of these projections and their widespread distribution within the cochlear nuclear complex would suggest an important role for the descending projections in the normal functioning of the cochlear nucleus.

Acetylcholinesterase

The projections of principal cells of the medial nucleus of the trapezoid body in the cat.

Previous studies suggest that the principal cells of the medial nucleus of the trapezoid body (MNTB) give rise to the projection from MNTB to the lateral superior olivary nucleus (LSO) of the same side, where they mediate rapid inhibitory effects of contralateral sound stimulation. In the present study, we explored certain morphological features of this connection as well as several other projections of the MNTB by using anterograde and retrograde axonal tracing methods. Following injections of tritiated leucine into MNTB, labeled axons reached LSO by passing ventral to, dorsal to, and through the medial superior olivary nucleus, and gave rise to labeling around the somata and proximal dendrites of LSO fusiform cells. As measured in autoradiograms of 2 micron plastic sections, these axons had a modal diameter of 5-6 micron. Terminal labeling, tentatively attributed to principal cell axons, was also seen in the ventral nucleus of the lateral lemniscus (VNLL) and the dorsomedial and ventromedial periolivary nuclei. HRP injections into the LSO and the VNLL showed that the principal cell projected to both of these nuclei and revealed a topographic arrangement of the projection to the LSO which is consistent with tonotopic maps determined electrophysiologically. Control HRP injections demonstrated that other minor projections of the MNTB arose from minor cell populations in this nucleus. The findings provide a morphological correlate of certain physiological findings and suggest a wider role for the MNTB in the ascending auditory system than previously has been supposed.

Animals

Topographic organization of the olivocochlear projections from the lateral and medial zones of the superior olivary complex.

By anterograde tracing using autoradiography, we have found topographic organizations in the projections of both medial and lateral olivocochlear (OC) systems in the cat. Lateral-zone injections show an ipsilateral cochleotopic projection pattern with more medial injections projecting more basally in the cochlea. In the contralateral cochlea, in contrast, the projections from all of the lateral-zone injections were predominantly to the apex. However, detailed analysis suggests the possibility that the contralateral lateral-zone projections may have the same cochleotopic organization as the ipsilateral projections but with a heavy bias toward the apex. Medial-zone injections show a pattern in which more dorsal regions project more basally in both cochleas. The ipsilateral projections of lateral OC neurons appear to connect regions with similar best frequencies but the projections of medial OC neurons do not. Summation of data from all of the injections in each zone indicates that lateral OC projections are relatively evenly distributed throughout the ipsilateral cochlea but are predominantly to the apex in the contralateral cochlea. Medial OC projections are predominantly to the middle and basal parts of the cochlea on both sides with contralateral projections somewhat more basal than ipsilateral projections.

Animals

Differential olivocochlear projections from lateral versus medial zones of the superior olivary complex.

An anterograde tracer (35S-methionine) was injected unilaterally in the superior olivary complex (SOC) at regions previously demonstrated by retrograde labeling to contain olivocochlear (OC) cell bodies. Quantitative analysis of cochlear autoradiographs from these cats demonstrates that there are two OC systems. The lateral OC system has cell bodies lateral to the medial superior olivary nucleus (MSO) and projects to the inner hair cell (IHC) region bilaterally (mostly ipsilaterally). The medial OC system has cell bodies medial, ventral, and anterior to the MSO and projects to the outer hair cell (OHC) region bilaterally (mostly contralaterally). A single medial OC neuron innervates many small patches of OHCs with substantial gaps between the patches. Medial OC neurons also appear to project to the IHC region to a small extent. A review of the literature with the medial-lateral division of OC efferents in mind reveals many differences between these two systems. In particular, lateral OC axons are unmyelinated and innervate the dendrites of radial afferent fibers under IHCs, whereas medial OC axons are myelinated and directly innervate OHCs. Although both systems appear to be cholinergic, the lateral OC system also shows met-enkephalin-like immunoreactivity. The synapses of the medial OC system are formed in development before those of the lateral OC system and they degenerate more slowly after the OC axons are cut. The many differences between these two OC systems suggest that they are functionally separate systems.

Animals

The dual origins of the olivocochlear bundle in the albino rat.

Recent studies of the origins and terminations of the olivocochlear bundle (OCB) in the cat provide evidence that separate efferent systems differentially innervate the two types of hair cells in the organ of Corti. To begin to test the generality of these separate olivocochlear systems, the cells of origin of the OCB were determined in the albino rat by using axonal transport of horseradish peroxidase. Our findings are that, as in the cat, two classes of olivocochlear (OC) neurons project to the cochlea. These neurons could be dichotomized according to their location in the superior olivary complex (lateral or medial), their size (small or large), and their preferred side of projection (ipsilateral or contralateral). All labeled OC neurons also exhibited a positive reaction for acetylcholinesterase. In the rat, however, lateral and medial OC neurons are each restricted to a single nucleus, and, furthermore, the lateral OC neurons project only ipsilaterally. The rat also has significantly fewer mean totals of both lateral (240 vs. 710) and medial (240 vs. 520) OC neurons than the cat. The present methods also demonstrated the course of axons of the OCB and axon collaterals entering the cochlear nuclear complex. Vestibular efferent neurons were also labeled bilaterally medial and lateral to the facial genu. Our results suggest that the general organizational plan of OC neurons in the rat may offer advantages over the situation in the cat for studies of connectional neuroanatomy, neurophysiology, and behavioral functions of the two olivocochlear systems.

Animals

Distribution and light microscopic features of granule cells in the cochlear nuclei of cat, rat, and mouse.

In the present study the cytology and the topography of the cochlear granule cell domain (a comprehensive term introduced here for all granule cell-containing regions of the cochlear nuclear complex) have been studied light microscopically in Nissl, Bielschowsky, and Golgi-Del Rio-Hortega material of cats, rats, and mice; in Golig rapid material of 0-14-day-old kittens; and in sections of 6-week-old kittens following HRP injections in the superficial dorsal cochlear nucleus (DCN). The domain has been parcellated in seven subdivisions which, in spite of some species' differences, are easily identifiable in all of the included animals. The cochlear granule cells are considered as a particular class of neuron, which is slightly different from, but nevertheless principally similar to the cerebellar granule cells in both shape and mode of neuronal connections. The digitiform terminals of the cochlear granule cells differentiate after the first two weeks of extrauterine life. In several respects these cells show larger variation among species than do the cerebellar granules, the similarity between the two classes of granule cells being most conspicuous in the rodent. The silver, Golgi rapid, and HRP material suggest that all, or at least the majority, of the granule cell axons project to the molecular layer of the DCN, forming parallel fibers similar to those of the cerebellar cortex. Also, the cochlear parallel fibers traverse the spiny apical dendrites of principal neurons (the pyramidal cells) and the smoother dendrites of molecular layer stellate cells.

Animals

Origins of axons in the cat's acoustic striae determined by injection of horseradish peroxidase into severed tracts.

Origins and terminations of fibers of the dorsal and intermediate acoustic striae were studied by surgically severing these tracts and injecting HRP into the incision. This procedure results in filling the severed axons with HRP. Filled axons were traced to cell groups of origin and to some terminations of the acoustic striae. HRP-labeled terminals were found in the cochlear nuclei as well as in periolivary cell groups. Filling of cells with HRP RANged from being complete, resulting in Golgi-like images, to being barely detectable. Labeled cells were abundant in the dorsal and posteroventral cochlear nucleus adjacent to the injection as well as scattered throughout the periolivary cell groups of both sides, being highest in concentration around the ipsilateral lateral superior olive. On the side contralateral to the injection, labeled cells were found along the medial border of the dorsal cochlear nucleus, in the interstitial nucleus of the stria of Held, and sparsely throughout the ventral cochlear nucleus. The distribution of labeled cells was similar following HRP injections of the dorsal cochlear nucleus, except that these injections revealed additional descending projections from the inferior colliculi and from the ventral nucleus of the trapezoid body of both sides. These additional projections were interpreted as entering the CN by a ventral route. Findings of this study are in accord with physiological recordings made from fibers of the acoustic striae.

Afferent Pathways

Olivocochlear and vestibular efferent neurons of the feline brain stem: their location, morphology and number determined by retrograde axonal transport and acetylcholinesterase histochemistry.

Anterograde degeneration studies have shown that the cochlear and vestibular receptor organs receive an efferent innervation from neurons in the brain stem. This pathway may provide a mechanism by which the CNS could modulate its own afferent input. The neurons which provide this innervation have so far escaped positive identification with methods which depend on retrograde cell changes after axotomy. In the present study, horseradish peroxidase (HRP) was injected into the labryinths of kittens and after allowing 24 hours for the retrograde axonal transport of this tracer, its presence in neurons of the brain stem was demonstrated histochemically. Because there is evidence that the efferent innervation of the labyrinth is cholinergic, acetylcholinesterase (AChE) was also demonstrated histochemically in the same or in adjacent tissue sections. Neurons labelled with HRP were found bilaterally in most periolivary cell groups of the superior olivary complex (cochlear efferents) and in the parvocellular reticular nucleus lateral to the abducens nucleus (vestibular efferents). Counts of labelled neurons yielded estimated totals of 1,700-1,800 cochlear and 400-500 vestibular efferent neurons. Approximately 60% of the neurons in each total were located on the side ipsilateral to the injection. The distribution of HRP-labelled neurons was virtually identical to that of AChE-positive neurons found in adjacent sections, and in those regions with predominantly ipsilateral or contralateral projections, there was an approximate correspondence in number of HRP- and AChE-positive neurons. In tissue sections processed successively for demonstration of HRP and AChE, virtually all HRP-labelled neurons were found to be AChE-positive. These findings suggest that a number of current conceptions regarding labyrinthine efferent systems may need revision.

Acetylcholinesterase

Efferent components of the auditory system.

The origins and terminations of the olivocochlear bundle, which provides an efferent innervation to the organ of Corti, are described on the basis of experiments using axonal transport of tracer substances and light microscopy in the cat. The cells of origin were labeled by the retrograde tracer horseradish peroxidase which was injected unilaterally into the cochlea. Labeled cells in the superior olivary complex could be dichotomized according to their location (lateral or medial), their size (small or large), and their preferred side of projection (uncrossed or crossed). All labeled olivocochlear neurons exhibited a positive reaction for acetylcholinesterase. To determine the cochlear projections of the neurons, injections of a radioactive amino acid were made into either the lateral or medial olivocochlear cell group. After allowing time for synthesis and axonal transport of radio-labeled protein to reach synaptic endings in the cochleas, the tissue sections of these specimens were processed for autoradiography. The results indicate that lateral olivocochlear neurons project to the regional beneath the inner hair cells of both sides, whereas medial olivocochlear neurons project to the region beneath the outer hair cells of both sides. These findings are in substantial accord with previous experimental work but suggest that the organ of Corti receives a dual efferent innervation which is organized according to the location and morphology of its cells of origin. Accordingly, it is proposed that the two efferent components of the cochlear innervation described here be referred to as the lateral and medial olivocochlear systems, replacing the current designations of crossed and uncrossed olivocochlear bundles, the latter which are demonstrably heterogeneous in their origins and terminations and, probably, also in their functions.

Animals