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C E Carr

Publications and source records attributed to C E Carr.

At least 37 records · Page 2Linked to original sources

Chick cochlear efferent neurons are not immunoreactive for calcitonin gene-related peptide.

The fluorescent retrograde tracer, rhodamine dextran amine, was unilaterally injected into the cochlear duct of anesthetized chicks. Retrogradely labeled cells were found bilaterally in the brainstem ventromedial to the superior olivary nucleus and to the ventral facial nucleus, and in the caudal pontine reticular formation between the dorsal facial nucleus and the abducens nerve root. Tissue sections containing retrogradely labeled cells were incubated in an antiserum to calcitonin gene-related peptide (CGRP) and a fluorescently labeled secondary antiserum. No double-labeled neurons were found, suggesting that chick cochlear efferent neurons do not contain CGRP or that levels of CGRP in these cells may be too low to be detected immunohistochemically.

Animals↗

Enkephalin-like immunoreactivity in the chick brainstem: possible relation to the cochlear efferent system.

Mammalian lateral olivocochlear (LOC) neurons that are immunoreactive for choline acetyltransferase (ChAT) are also immunoreactive for enkephalin (Enk). To determine whether cochlear efferent neurons in birds might also contain Enk-like immunoreactivity (Enk-LI), we studied the auditory brainstem of the domestic chicken using antisera to ChAT, leucine-enkephalin (L-Enk) and methionine-enkephalin (M-Enk). Enk-LI terminals are found around, but not within, the superior olivary nucleus (SO) and the nucleus of the lateral lemniscus, pars intermedia (LLi). A moderate concentration of Enk-LI terminals is found ventromedial to the ventral facial nucleus (VIIv) where the ventrolateral group of ChAT-I cochlear efferent neurons is located. After colchicine injections into the lateral ventricle, a population of intensely stained Enk-LI perikarya was found in the nucleus of the lateral lemniscus, pars ventralis (LLv) with scattered cells in the LLi and the nucleus subceruleus ventralis (SCv). The distribution of Enk-LI and ChAT-I somata, however, never overlapped, even in adjacent sections. Thus, in the chick, Enk-LI perikarya are not distributed in areas where cochlear efferent neurons are found. Instead, a dense concentration of Enk-I terminals can be found in areas containing ChAT-I cochlear efferent neurons. The source of these enkephalinergic terminals may be a population of Enk-LI cells in the LLv.

Animals↗

Choline acetyltransferase-immunoreactive cochlear efferent neurons in the chick auditory brainstem.

Cholinergic neurons in the chick auditory brainstem were studied with the aid of an antiserum to choline acetyltransferase (ChAT), the biosynthetic enzyme for acetylcholine. ChAT-immunoreactive (ChAT-I) neurons were found in a ventrolateral and a dorsomedial cell group. The ventrolateral group is a rostrocaudally directed column of cells that surround the superior olive (SO), are ventromedial to the ventral facial nucleus (VIIv), and are lateral to the nucleus pontis lateralis (PL) as far rostrally as the nucleus subceruleus ventralis. Cells in the dorsomedial group were found in the pontine reticular formation medial to the dorsal facial nucleus and lateral to the abducens nerve root. Occasionally, small ChAT-I cells were found in the crossed dorsal cochlear tract and in the medial vestibular nucleus near the dorsal border of the caudal nucleus magnocellularis (NM). No ChAT-I neurons or fibers were observed in NM, nucleus angularis, nucleus laminaris, in the nuclei of the lateral lemniscus, or in the nucleus mesencephalicus lateralis pars dorsalis. To determine which cholinergic neurons project to the cochlea, a double-labeling technique was used combining ChAT-I and the retrograde transport of biotinylated dextran amine (BDA) from the inner ear. Double-labeled cells were found bilaterally in both the ventrolateral and dorsomedial cell groups, with the exception of large ChAT-I cells dorsal to the SO, which do not appear to project to the cochlea. Cholinergic cells that project to the cochlea were classified into three morphological groups: multipolar, elongate, and round-to-oval. Both the ventrolateral and the dorsomedial cell groups appear to have a mixture of these different cell types. The average somal area of cholinergic cochlear efferents was 246 microns 2. Only about 70% of the cochlear efferent neurons, however, are cholinergic.

Animals↗

An axon with a myelinated initial segment in the bird auditory system.

In the bird auditory brainstem, nucleus laminaris neurons compute interaural time differences by comparing inputs from two ears. Laminaris neurons in barn owls have short dendrites and axons with myelinated initial segments. We examined laminaris cells in chickens with a range of dendritic lengths to determine if neurons with long dendrites had normal axons. Neurons with short dendrites had myelinated initial segments while neurons with long dendrites had normal initial segments. Thus, the myelinated initial segment may serve to move the spike-initiation zone away from the somatodendritic compartment that integrates inputs from the two ears.

Animals↗

Organization of the nucleus magnocellularis and the nucleus laminaris in the barn owl: encoding and measuring interaural time differences.

The circuit from the cochlear nucleus magnocellularis to the nucleus laminaris supports the encoding and measurement of interaural time differences in the auditory brainstem. Specializations for the encoding of temporal information include the few and/or short dendrites and thick axons of the magnocellular and laminaris neurons, and the high degree of convergence in the circuit. Magnocellular cells have large cell bodies covered with somatic spines. The cells have few dendrites, and the number of dendrites decreases from low to high best frequency regions of the nucleus. Magnocellular neurons receive both auditory nerve terminals and GABAergic terminals with symmetric synapses and terminals filled with pleomorphic vesicles. The axonal projections of magnocellular neurons to the nucleus laminaris form maps of interaural time difference. About 100 magnocellular afferents from each side converge on each laminaris neuron, and the terminals from each side do not occupy separate domains on the cell. These terminals form punctate asymmetric synapses on both the dendrites and the cell bodies of laminaris neurons. Laminaris neurons also receive GABAergic terminals which form symmetric synapses. Laminaris neurons have oval cell bodies covered with very short dendrites. The cells in the low best frequency region of the nucleus laminaris have longer dendrites.

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Central projections of auditory nerve fibers in the barn owl.

The central projections of the auditory nerve were examined in the barn owl. Each auditory nerve fiber enters the brain and divides to terminate in both the cochlear nucleus angularis and the cochlear nucleus magnocellularis. This division parallels a functional division into intensity and time coding in the auditory system. The lateral branch of the auditory nerve innervates the nucleus angularis and gives rise to a major and a minor terminal field. The terminals range in size and shape from small boutons to large irregular boutons with thorn-like appendages. The medial branch of the auditory nerve conveys phase information to the cells of the nucleus magnocellularis via large axosomatic endings or end bulbs of Held. Each medial branch divides to form 3-6 end bulbs along the rostrocaudal orientation of a single tonotopic band, and each magnocellular neuron receives 1-4 end bulbs. The end bulb envelops the postsynaptic cell body and forms large numbers of synapses. The auditory nerve profiles contain round clear vesicles and form punctate asymmetric synapses on both somatic spines and the cell body.

Animals↗

Calbindin-like immunoreactivity in the central auditory system of the mustached bat, Pteronotus parnelli.

With the aid of a polyclonal antibody specific for Calbindin D-28k, we studied the distribution of this calcium-binding protein in the central auditory system of the mustached bat, Pteronotus parnelli. Components of the cochlear nucleus (CN) that were calbindin-positive (cabp(+] included the root of the auditory nerve, multipolar and globular bushy cells in the anteroventral CN, multipolar and octopus cells in the posteroventral CN, and small and medium-size cells in the dorsal CN. Not stained were spherical bushy cells of the anteroventral CN and pyramidal/fusiform cells in the dorsal CN. In the superior olivary complex, labeled cells were found in the lateral and medial nuclei of the trapezoid body, the ventral and ventromedial periolivary nuclei, and the anterolateral periolivary nucleus. No cellular labeling was seen in the lateral superior olive. In the medial superior olive, only marginal cells were cabp(+). Labeled fibers could be seen surrounding the gosts of unlabeled cells in both the latter nuclei. Most cells in the intermediate nucleus and the columnar division of the ventral nucleus of the lateral lemniscus were cabp(+). However, the dorsal nucleus was cabp(-). A group of cabp(+) cells was also seen in the paralemniscal zone. The inferior colliculus had a relatively low density of cabp(+) cells. Labeled cells were more common in the caudal half of the central nucleus, and in the external nucleus and dorsal cortex. In the auditory thalamus, nearly every cell in the medial geniculate body was cabp(+), but those in the suprageniculate nucleus and in the posterior group did not stain. Small cells in the intermediate layer and giant cells in the deep layers of the superior colliculus were densely cabp(+). In the pons, cabp(+) cells and neuropil could be seen in the medial and lateral pontine nuclei (pontine gray). In conclusion, calbindin-like immunoreactivity was found in most of the brainstem auditory system, as well as in regions associated with acoustic orientation or control of vocalization. However, except for a minority of cells of the medial superior olive, it is conspicuously absent from the nuclei receiving binaural input below the level of the inferior colliculus.

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A circuit for detection of interaural time differences in the brain stem of the barn owl.

Detection of interaural time differences underlies azimuthal sound localization in the barn owl Tyto alba. Axons of the cochlear nucleus magnocellularis, and their targets in the binaural nucleus laminaris, form the circuit responsible for encoding these interaural time differences. The nucleus laminaris receives bilateral inputs from the cochlear nucleus magnocellularis such that axons from the ipsilateral cochlear nucleus enter the nucleus laminaris dorsally, while contralateral axons enter from the ventral side. This interdigitating projection to the nucleus laminaris is tonotopic, and the afferents are both sharply tuned and matched in frequency to the neighboring afferents. Recordings of phase-locked spikes in the afferents show an orderly change in the arrival time of the spikes as a function of distance from the point of their entry into the nucleus laminaris. The same range of conduction time (160 mu sec) was found over the 700-mu m depth of the nucleus laminaris for all frequencies examined (4-7.5 kHz) and corresponds to the range of interaural time differences available to the barn owl. The estimated conduction velocity in the axons is low (3-5 m/sec) and may be regulated by short internodal distances (60 mu m) within the nucleus laminaris. Neurons of the nucleus laminaris have large somata and very short dendrites. These cells are frequency selective and phase-lock to both monaural and binaural stimuli. The arrival time of phase-locked spikes in many of these neurons differs between the ipsilateral and contralateral inputs. When this disparity is nullified by imposition of an appropriate interaural time difference, the neurons respond maximally. The number of spikes elicited in response to a favorable interaural time difference is roughly double that elicited by a monaural stimulus. Spike counts for unfavorable interaural time differences fall well below monaural response levels. These findings indicate that the magnocellular afferents work as delay lines, and the laminaris neurons work as co-incidence detectors. The orderly distribution of conduction times, the predictability of favorable interaural time differences from monaural phase responses, and the pattern of the anatomical projection from the nucleus laminaris to the central nucleus of the inferior colliculus suggest that interaural time differences and their phase equivalents are mapped in each frequency band along the dorsoventral axis of the nucleus laminaris.

Acoustic Stimulation↗

Distribution of GABAergic neurons and terminals in the auditory system of the barn owl.

Antisera to GAD (glutamic acid decarboxylase) and GABA were used to determine the distribution of GABAergic cells and terminals in the brainstem and midbrain auditory nuclei of the barn owl. The owl processes time and intensity components of the auditory signal in separate pathways, and each pathway has a distinctive pattern of GAD- and GABA-like immunoreactivity. In the time pathway, all the cells of the cochlear nucleus magnocellularis and nucleus laminaris receive perisomatic GABAergic terminals, and small numbers of GABAergic neurons surround both nuclei. The ventral nucleus of the lateral lemniscus (anterior division) contains both immunoreactive terminals and some GABAergic neurons. In the intensity pathway, dense immunoreactive terminals are distributed throughout the cochlear nucleus angularis, which also contains a small number of GABAergic neurons. The superior olive contains two GABAergic cell types and immunoreactive terminals distributed throughout the neuropil. All the neurons of the nucleus of the lateral lemniscus (ventral part) appear to be GABAergic, and this nucleus also contains a moderate number of immunoreactive terminals. Immunoreactive terminals are distributed throughout the neuropil of the ventral nucleus of the lateral lemniscus (posterior division), whereas multipolar and small fusiform GABAergic neurons predominate in the dorsal regions of the nucleus. The time and intensity pathways combine in the inferior colliculus. The central nucleus of the inferior colliculus contains a larger number of fusiform and stellate GABAergic neurons and a dense plexus of immunoreactive terminals, whereas the external nucleus contains slightly fewer immunoreactive cells and terminals. The superficial nucleus contains dense, fine immunoreactive terminals and a small number of GABAergic neurons.

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Axonal delay lines for time measurement in the owl's brainstem.

Interaural time difference is an important cue for sound localization. In the barn owl (Tyto alba) neuronal sensitivity to this disparity originates in the brainstem nucleus laminaris. Afferents from the ipsilateral and contralateral magnocellular cochlear nuclei enter the nucleus laminaris through its dorsal and ventral surfaces, respectively, and interdigitate in the nucleus. Intracellular recordings from these afferents show orderly changes in conduction delay with depth in the nucleus. These changes are comparable to the range of interaural time differences available to the owl. Thus, these afferent axons act as delay lines and provide anatomical and physiological bases for a neuronal map of interaural time differences in the nucleus laminaris.

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Calcium binding protein-like immunoreactivity labels the terminal field of nucleus laminaris of the barn owl.

Nucleus laminaris (NL) is the site at which the timing of sounds arriving in the 2 ears is compared in the auditory system of the barn owl. Earlier studies have reported vitamin D-dependent calcium binding protein (CaBP)-like immunoreactivity in the somata of NL. We report here that CaBP-like immunoreactivity stains the terminal field of NL. The specific CaBP immunoreactivity is localized to a dense plexus of fibers that have bouton-like swellings, usually around unstained somata. This type of immunoreactivity is found in a restricted portion of the central nucleus of the inferior colliculus (ICc), in the anterior division of the ventral lateral lemniscal complex (VLVA), and in the superior olivary nucleus (SO), all of which have been shown by anterograde transport of 3H-proline to be innervated by NL. The immunoreactivity is absent from the posterior division of ventral lateral lemniscal complex and from the region that surrounds the portion of ICc innervated by NL. A restricted lesion in NL results in a localized deficit in immunoreactivity in those regions of ICc and VLVA that are known to be innervated by the lesioned area of NL. In adjacent sections processed by the Fink-Heimer method, degenerating axons are present in the region of the deficit in immunoreactivity.

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Time coding in electric fish and barn owls.

A comparison of the coding and processing of temporally ordered information in electric fish and barn owls demonstrates a number of similar design features including different time and intensity channels and distinct morphological characteristics. Similar mechanisms, such as the presence of delay lines, underlie the sensitivity of both animals to small temporal disparities. Differences between the two systems may reflect the different substrates upon which evolution acted, and the different requirements of the two systems.

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A time-comparison circuit in the electric fish midbrain. II. Functional morphology.

The weakly electric fish Eigenmannia is able to detect temporal disparities as small as 400 nsec between two signals from different parts of the body surface (Carr et al., 1986). The elements of this time-comparison circuit have been identified by EM reconstruction of its component cells. Information about the timing of the zero-crossing of signals on each area of the body surface is coded in phase-coder receptors, a subset of tuberous electroreceptors. Electroreceptors on the body surface are innervated by primary afferents with their central termination on the spherical cells of the medullary electrosensory lateral line lobe. These cells project to lamina VI of the midbrain torus, a structure similar to the inferior colliculus. Afferents entering lamina VI form a very restricted terminal arbor in which they synapse on the three cell types of this lamina. Each afferent makes gap-junction synapses on one or two giant cell somata and morphologically mixed synapses on the distal dendrites of two types of small cell. The afferent terminals thus encode the timing of the electric signal on a local patch of the body surface, forming a somatotopic map of the body surface in lamina VI. The giant cells are adendritic and their axonal arbor is such as to distribute timing information originating from one part of the body surface throughout lamina VI, so that each region of lamina VI receives information about the timing of zero-crossings from the entire body surface from giant cells, as well as information from a local portion of the body surface from the afferent terminals. The giant cells terminate exclusively on the cell bodies of the small cells of lamina VI, shown to be sensitive to small temporal disparities by Heiligenberg and Rose (1985). Thus, each small cell receives a single synapse on its soma from a giant cell that conveys phase-coding information from some portion of the body surface and receives local phase-coding input onto its dendrites from spherical cell afferents. The sensitivity of the small cells to temporal disparities appears to be conferred by their segregation of inputs from two different parts of the body surface onto dendrites and soma, respectively. We propose that the dendritic input acts as a delay line, and the small cell fires maximally when the inputs from the dendrites and the giant cell input onto the soma coincide.

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A time-comparison circuit in the electric fish midbrain. I. Behavior and physiology.

Behavioral experiments show that the weakly electric fish, Eigenmannia, detects differences in timing as small as 400 nsec between electric signals from different parts of its body surface. The neural basis of this remarkable temporal resolution was investigated by recording from elements of the phase-coding system, a chain of electrotonically connected neurons devoted to the processing of temporal information. Each element of this system fires a single action potential for every cycle of the electric signal (either the fish's own electric organ discharge or a sinusoidal signal of similar frequency). For phase-coding primary afferents and midbrain neurons, the temporal resolution was determined by measuring each unit's capacity to lock its spike to a particular phase of the stimulus cycle. The jitter of a neuron's response (measured as the standard deviation of the timing of the spikes with respect to the stimulus) decreases from the level of the primary afferent (mean = 30 microsec) to the midbrain torus (mean = 11 microsec); these results can be correlated with morphological measures of convergence. The temporal resolution of single neurons is still inferior to that displayed at the behavioral level.

Action Potentials↗

A Golgi study of the cell types of the dorsal torus semicircularis of the electric fish Eigenmannia: functional and morphological diversity in the midbrain.

The dorsal torus semicircularis (torus) of the gymnotiform fish Eigenmannia was examined in Golgi-impregnated material. These results were correlated with those of a previous HRP study which used retrograde labelling techniques to identify the efferent cell types of the torus (Carr et al., '81, J. Comp Neurol. 203:649-670). The torus is a laminated midbrain nucleus of the electrosensory system. It receives somatotopically ordered electrosensory input from the medulla and caudal lobe of the cerebellum, proprioceptive input from the descending nucleus of the trigeminal nerve, and input from the optic tectum. The torus projects to the nucleus praeeminentialis, the optic tectum, nucleus electro-sensorius, parts of the central posterior thalamus, the pretectum, the lateral mesencephalic reticular formation (LMRA), the reticular formation, and the inferior olive. The torus has 12 laminae and 48 cell types by Golgi criteria. There are three major orientations to the dendritic fields of the toral neurons: purely horizontal neurons with dendrites confined to a single lamina, multipolar neurons whose dendrites often do not respect laminar boundaries, and vertical cells with dendrites that travel in the vertical bundles of dendrites and axons which pierce the torus at regular intervals. There are four major groups of vertically oriented neurons. The first has a predominantly horizontal dendritic tree with one or two vertical dendrites which connect the cell to a distant lamina. The second consists of "U"-shaped neurons with a horizontal arbor and two major dendrites which ascend in adjacent vertical bundles. The third group is made up of bilaminar neurons which receive input from two vertically separated dendritic arbors, and the fourth group is purely vertical in orientation. A group of four tegmental cell types in the LMRA also send their dendrites into the efferent tracts of the torus, and into lamina IX. The torus is similar in complexity and number of cell types to the mammalian inferior colliculus. The large number of cell types in these midbrain sensory nuclei, compared to the number of afferent inputs (seven or more for the torus) is notable and may reflect the parcellation of function associated with the parallel processing of these inputs.

Afferent Pathways↗

Peripheral organization and central projections of the electrosensory nerves in gymnotiform fish.

The electrosensory system of weakly electric gymnotiform fish is described from the receptor distribution on the body surface to the termination of the primary afferents in the posterior lateral line lobe (PLLL). There are two types of electroreceptor(ampullary and tuberous) and a single type of lateral line mechanoreceptor (neuromast). Receptor counts in Apteronotus albifrons show that (1) neuromasts are distributed as in other teleosts; (2) ampullary receptors number 151 on one side of the head and 208 on one side of the body; (3) tuberous receptors were estimated to number 3,000-3,500 on one side of the head and 3,500-5,000 on one side of the body. The distribution of each receptor type is described. Each receptor is innervated by a single primary afferent. Electrosensory afferents have myelinated cell bodies in the ganglion of the anterior lateral line nerve (ALLN). The distribution of these ganglion cell diameters is strongly bimodal in Apteronotus and Eigenmannia: The smaller-diameter cells may be those which innervate ampullary electroreceptors, the larger-diameter tuberous electroreceptors. Transganglionic HRP transport techniques were used to determine the first-order connections of the anterior lateral line nerve in six species of gymnotiform fish. Small branches of the ALLN were labeled so as to determine the somatotopic organization in the PLLL. The PLLL is divided into four segments from medial to lateral, termed medial, centromedial, centrolateral, and lateral segments (Heiligenberg and Dye, '81). Representations of the head are found rostrally in each zone, and the trunk is mapped caudally in each zone. Thus there are four body maps in the PLLL. The medial segment receives ampullary input (Heiligenberg and Dye, '82) and maps the dorsoventral body axis mediolaterally, as does the tuberous centrolateral segment. The tuberous centromedial and lateral segments map the dorsoventral axis lateromedially. Thus the medial and centromedial segments meet belly to belly, the centromedial and centrolateral segments meet back to back, and the centrolateral and lateral segments meet belly to belly. Adjacent electrosensory maps within the PLLL are therefore always mirror images.

Afferent Pathways↗

Efferent projections of the posterior lateral line lobe in gymnotiform fish.

The posterior lateral line lobe (PLLL) of gymnotoid fish has efferent projections to two midbrain regions: the nucleus praeeminentialis dorsalis (n.P.d.) and the torus semicircularis dorsalis (T.Sd.). Both ipsilateral and contralateral connections are present; the n.P. d. receives nearly equal input from both sides while the T.Sd. receives a stronger contralateral input. The PLLL projection to n.P.d. merely maps medial PLLL to ventral n.P.d. and lateral PLLL to dorsal n.P.d., thus preserving the separate topography and relative orientation of the four electrosensory maps found in the PLLL. Only PLLL pyramidal cells (basilar and nonbasilar pyramids) contribute to this projection. The four PLLL electrosensory maps converge onto T.Sd. so that they map the dorsal body surface onto medial T.Sd. and the ventral body surface onto lateral T.Sd. Pyramidal cells, spherical cells, and multipolar cells contribute to this projection. A small commissural connection links homologous segments of the PLLL; these fibers arise from polymorphic cells of the PLLL.

Animals↗