Caudal Zone Research Articles

Identification of the Purkinje cell/climbing fiber zone and its target neurons responsible for eye-movement control by the cerebellar flocculus

We identified 3 Purkinje cell/climbing fiber zones in the cat cerebellar flocculus. The zones were perpendicular to the long axes of the crooked floccular folia, forming the crooked zones. Each zone was different in axonal projection areas of its target neurons. From the neuronal networks it is theoretically expected that activity changes of a particular zone control eye movement in a particular plane: (1) the rostral and caudal zones on one side control movement in the anterior canal plane oh the side of the activity changes and those on both sides control movement in all vertical planes from sagittal to transverse planes; and (2) the middle zone controls movement in the horizontal plane by reciprocal activity changes on both sides. The zone-specific climbing fiber input to a particular zone may contribute to activity changes of the zone in response to mossy fiber input spreading across several zones. Electrical stimulation of each zone evoked the same pattern of eye movement as that theoretically expected from the neuronal networks. This is the first indication that there are indeed functional differences between the Purkinje cell zones in the cerebellum. Our findings support Oscarsson's proposal that each Purkinje cell/climbing fiber zone plus its target neurons may be an operational unit for control of a given motor function.

Operational unit responsible for plane-specific control of eye movement by cerebellar flocculus in cat

1. Main findings in our previous studies are as follows: 1) there are three Purkinje cell zones running perpendicular to the long axis of the folia in the cat flocculus, 2) the caudal zone controls activity of the superior rectus (SR) and inferior oblique (IO) extraocular muscles via the y-group and oculomotor nucleus (OMN) neurons, and 3) the middle zone controls activity of the lateral (LR) and medial rectus (MR) muscles via the medial vestibular (MV) and abducens nucleus (ABN) neurons. In the present study, the neuronal pathways from the remaining rostral zone were investigated in the anesthetized cat. 2. Target neurons of rostral zone inhibition in the superior vestibular nucleus (SV) were identified by observing cessation of spontaneous discharges after rostral zone stimulation. Efferent projections were studied by the use of systematic microstimulation techniques. Unitary responses to stimulation of the eighth nerves were also investigated. 3. There are two types of the target neurons: 1) those, being located in the central and dorsal parts of the SV, project to the trochlear and oculomotor nuclei innervating superior oblique and inferior rectus muscles via the ipsilateral medial longitudinal fasciculus (MLF); and 2) those, being located along the dorsal border of the SV, project to the contralateral oculomotor nucleus innervating superior rectus and inferior oblique muscles via the extra-MLF route. 4. Both types receive monosynaptic anterior canal nerve input but not posterior canal nerve input. Some neurons receive polysynaptic excitatory input from the contralateral eighth nerve, although commissural inhibition was never observed. 5. From neuronal connections of the rostral and caudal zones and action of the extraocular muscles, it was expected that 1) activity changes of Purkinje cells in the rostral and/or caudal zones on one side resulted in conjugate eye movement in the plane of the anterior canal on the side of the activity changes, 2) cooperative increased activity on both sides resulted in conjugate downward eye movement, and 3) increased activity on one side and decreased activity on the other side resulted in conjugate rotatory eye movement. The rostral and caudal zones may be responsible for eye-movement control in the sagittal plane by cooperative activity changes on both sides and in the transverse plane by reciprocal activity changes on both sides. For eye-movement control in the anterior canal plane, Purkinje cell activity on one side would be sufficient to produce the required movement. In a functional sense, we call the rostral and caudal zones, the vertical-plane zones.(ABSTRACT TRUNCATED AT 400 WORDS)

Type X collagen synthesis by cultured chondrocytes derived from the permanent cartilaginous region of chick embryo sternum

In the developing chick embryo sternum, type X collagen is synthesized by chondrocytes from the cephalic region (presumptive mineralization zone) but not by chondrocytes from the caudal region (permanent cartilaginous zone) (Gibson et al., 1984, J. Cell Biol. 99, 208–216). To distinguish between two possibilities, the presence of a nonpermissive microenvironment in the permanent cartilage or the intrinsic inability of caudal chondrocytes to become hypertrophic, type X-producing cells, we have isolated chondrocytes from the caudal third of stage 44 chick embryo sterna and grown them in suspension on agarose-coated dishes. We have found that in these conditions chondrocytes from the caudal zone differentiate to hypertrophic chondrocytes and synthesize large amount of type X collagen, as revealed by the electrophoretic pattern of labeled proteins made in vitro and by slot blot analysis of mRNAs with specific cDNA probes.

Target neurons of floccular caudal zone inhibition in Y-group nucleus of vestibular nuclear complex.

Extracellular unit spikes were recorded in and around the Y-group nucleus in the anesthetized cat. Target (T) neurons of floccular caudal zone inhibition were identified by observing cessation of their spontaneous discharges following stimulation of the floccular caudal zone. The axonal trajectories of the T neurons to the rostral brain stem were studied by observing the antidromic responses of single neurons during systematic tracking with a stimulating microelectrode in the brain stem. The axons of the T neurons pass through a region closely ventral to the lateral part of the brachium conjunctivum (BC), continue rostrally in a region between the BC and the lateral lemniscus, arch medially around the rostral part of the nucleus reticularis tegmenti pontis, cross the midline, continue to the contralateral side by about 1.5 mm lateral from the midline, arch rostrally, run in the central tegmental field on the contralateral side, arch dorsomedially around the caudal pole of the red nucleus, and enter the contralateral oculomotor nucleus (OMN) from the ventrolateral side. In the caudal half of the contralateral OMN, the axons of the T neurons branch out and terminate. The T neurons were exclusively located in the dorsal subdivision of the Y-group nucleus (DY), whereas some were in the medial part of the subnucleus lateralis parvocellularis (SLP, Ref. 12) of the lateral cerebellar nucleus. T neurons were not found in the ventral subdivision of the Y-group nucleus (VY). Differences in neuronal connections between the DY and VY neurons were investigated by observing responses of single neurons to stimulation of the contralateral OMN, the ipsilateral floccular caudal zone, the ipsilateral eighth nerve (i8N), and the contralateral eighth nerve (c8N). Most neurons in the DY and the adjacent medial part of the SLP, receiving inhibitory inputs from the ipsilateral flocculus (exclusively from the caudal zone), project to the contralateral OMN, and about one-half of these neurons receive polysynaptic inputs from the i8N and the c8N. On the other hand, most neurons in the VY receive monosynaptic inputs from the i8N, and some of these neurons project to the ipsilateral flocculus. The neuronal tract via the ventral part of the pontine tegmentum demonstrated in the present experiments is distinct from the classically established vestibulooculomotor tracts via the BC, the medial longitudinal fasciculus, or the ascending tract of Deiters. We call this tract the 'crossing ventral tegmental tract'. Previously, we reported that electrical stimulation of the caudal zone elicited conjugate downward eye movement.(ABSTRACT TRUNCATED AT 400 WORDS)

Morphology and distribution of cystic cavities in the normal murine thymus.

The normal murine thymus was examined by light- and electron microscopy to determine the distribution and morphology of extracellular cystic cavities. Most cavities were confined to the cranial half of each gland, situated at the junction between cortex and medulla. They varied in size and shape, and gave rise to narrow channels that coursed to the capsular surface of the gland. Large cavities could be divided into three zones. A short cranial zone exhibited gland-like features, consisting of cells lining a clear lumen. A central zone was lined by a diverse population of cells. Some possessed secretory granules, while others exhibited an apical ciliated border. Lining cells inter-digitated with each other and were joined laterally by intercellular junctions. The lumen of the central zone contained lymphocytes and macrophages in an amorphous extracellular matrix. The caudal zone of each cavity had an attenuated and incomplete cellular lining, communicating directly with the surrounding thymic parenchyma. Thymic cavities may represent the initial part of the efferent lymphatic system of the gland, beginning in the tissue spaces at the corticomedullary junction. Selected cells could then enter and interact with the luminal contents in the central zone of the cavity. Ciliated cells may then propel lymphocytes and secretions into the narrow channels radiating from the uppermost part of the chamber, leaving a cell-free lumen in this region. These cavities may function in sequestering lymphocytes, macrophages and thymic secretions before their exit from the gland.

Functional organization of thalamic projections to the motor cortex. An anatomical and electrophysiological study in the rat

In rats, horseradish peroxidase crystals were injected in motor cortical foci functionally identified by means of the motor effects evoked by electrical stimulations. The location in the thalamus of the neurons linked to different motor cortical foci was studied. Thalamic neurons were retrogradely labeled in both “motor” (ventralis lateralis and ventralis medialis) and “non-motor” nuclei: centralis lateralis, lateralis posterior, mediodorsalis and posterior thalamic nuclear group, as well as the ventrobasal complex. The ventrobasal complex was labeled after horseradish peroxidase injections in hindlimb and trunk motor areas. The ascending projections toward the motor cortex from both “motor” and “non-motor” thalamic nuclei are organized more precisely and more elaborately than previously reported. The motor cortical afferents from the nucleus ventralis lateralis are organized in three planes, rostrocaudally, dorsoventrally and mediolaterally. An inverted relation exists in the rostrocaudal plane between the nucleus ventralis lateralis and the motor cortex: the caudal motor cortex region (hindlimb) receives fiber inputs from the rostral region of the nucleus ventralis lateralis, whereas the caudal zone of the nucleus ventralis lateralis projects to the rostral motor cortex region (forelimb and vibrissae). A dorsoventral organization has also been observed in the rostral region of the nucleus ventralis lateralis: the ventral aspect is the source of fibers directed to the distal hindlimb region, whereas fibers originating from the dorsal aspect are directed to the proximal hindlimb area. A mediolateral relationship exists between medial and lateral sides of the nucleus ventralis lateralis and, respectively, proximal and distal forelimb cortical areas. There is some overlap between the various nuclear regions thus delineated. Four functional zones were found in the lateral half of the nucleus ventralis medialis and were classified according to their projection to the motor cortex; these are involved in motor control of the proximal and distal forelimb, vibrissae and ocular movements. The projection is topographically organized according to both an inverted rostrocaudal and a direct dorsoventral-mediolateral arrangement. Caudally dorsal and ventral nuclear parts project to rostromedial (vibrissae) and rostrolateral (distal forelimb) regions of the motor cortex, respectively. More rostral nuclear zones project to more caudal (proximal forelimb, eye) cortical regions. There is little overlap between these four nuclear subdivisions. The nucleus centralis lateralis projects to vibrissae and proximal, as well as distal, forelimb areas. The nucleus mediodorsalis was labeled after horseradish peroxidase injections in the vibrissae motor area. Fibers arising from the lateralis posterior and posterior group reach, predominantly, the hindlimb and forelimb motor areas, respectively. The somatotopic organization of the cerebello-thalamo-cortical pathway in the rat was examined by matching the topographical organization of the cerebellothalamic pathway, as obtained in a former study 1 to the results of the present study. Comparisons were attempted with data from other species, particularly the monkey.

Tracing of single fibers of the nervus terminalis in the goldfish brain

Central projections of the nervus terminalis (n.t.) in the goldfish were investigated using cobalt- and horseradish peroxidase-tracing techniques. Single n.t. fibers were identified after unilateral application of cobalt chloride-lysine to the rostral olfactory bulb. The central course and branching patterns of individual n.t. fibers were studied in serial sections. Eight types of n.t. fibers are differentiated according to pathways and projection patterns. Projection areas of the n.t. include the contralateral olfactory bulb, the ipsilateral periventricular preoptic nucleus, both retinae, the caudal zone of the periventricular hypothalamus bilaterally, and the rostral optic tectum bilaterally. N.t. fibers cross to contralateral targets in the anterior commissure, the optic chiasma, the horizontal commissure, the posterior commissure, and possibly the habenular commissure. We propose criteria that differentiate central n.t. fibers from those of the classical secondary olfactory projections. Branching patterns of eight n.t. fiber types are described. Mesencephalic projections of the n.t. and of secondary olfactory fibers are compared and discussed with regard to prior reports on the olfactory system of teleosts. Further fiber types for which the association with the n.t. could not be established with certainty were traced to the torus longitudinalis, the torus semicircularis, and to the superior reticular nucleus on the ipsilateral side.

The neuronal pathway from the flocculus to the oculomotor nucleus: An electrophysiological study of group y nucleus in cats

In 17 cats anesthetized by Ketalar, neuronal activities were recorded extracellularly by a tungsten microelectrode placed in the group y nucleus of the vestibular nuclei. Forty-seven of 139 neurons tested were activated antidromically by stimulation of the superior rectus and inferior oblique subdivisions of the oculomotor nucleus. Twenty-five of these 47 neurons ceased their spontaneous discharges after stimulation of the caudal zone of the flocculus. In addition, 18 out of these 25 units responded polysynaptically to stimulation of the vestibular nerve.

Peripheral and central oculomotor organization in the goldfish, Carassius auratus.

Peripheral and central oculomotor organization was studied in the goldfish. The sizes of the extraocular muscles were quantified by counting the fibers contained in a given muscle and by area measurements of the cross-sectional surfaces. All the muscles were of approximately similar size. Kinematics were determined by electrical stimulation of a given muscle. The macroscopic appearance and kinematics of the muscles had the characteristics of other lateral-eyed animals (e.g., rabbit). Locations of extraocular motor neurons were found by retrograde transport of horseradish peroxidase (HRP) following injections into individual extraocular muscles. The eye muscles were innervated by four ipsilateral (lateral rectus, medial rectus, inferior oblique, inferior rectus) and two contralateral (superior rectus, superior oblique) motor neuron pools. The oculomotor nucleus was found in the midbrain, at the level of the caudal zone of the inferior lobe of the hypothalamus. Inferior rectus motor neurons were located rostrally in the oculomotor nucleus, whereas medial rectus, superior rectus, and inferior oblique motor neurons were intermingled in its more caudal portions. All labelled cells were located dorsally and medially to the medial longitudinal fasciculus (MLF) in close proximity to either the floor of the ventricle or the midline region. Occasionally, motor neurons were interspersed within the fiber bundles of the MLF or the exiting fibers of the oculomotor nerve. The trochlear nucleus, containing superior oblique motor neurons, was found in the immediate lateral and caudal neighborhood of the oculomotor nucleus, where its rostral border overlapped with the caudal border of the latter. The abducens nucleus, containing lateral rectus motor neurons, was located in the posterior brainstem in the neighborhood of the vestibular nuclear complex. This nucleus was divided into a rostral and a caudal portion. The axons of ipsilaterally projecting motor neurons headed toward their respective nerve roots via the shortest possible route, as did the axons of superior rectus motor neurons, which crossed the midline without detour to enter the contralateral oculomotor nerve. In contrast, trochlear motor neuron axons arched around the dorsal aspect of the ventricle through the cerebellar commissure to reach the contralateral trochlear nerve. The morphology of individual motor neurons was visualized by intrasomatic injection of HRP. Cell somata had oblong shapes, and their large dendrites were oriented laterally and ventrally. The axons did not collateralize within the midbrain region or the oculomotor nerve as far as they could be traced.(ABSTRACT TRUNCATED AT 400 WORDS)

Contrasting properties of neurons in two parts of the primary motor cortex of the awake rat

Features of neuronal activity in two subdivisions of primary motor cortex (MI) were recorded in awake rats. Neurons in the caudal part of MI, which overlaps part of the somatic sensory cortex, discharge with brief bursts in conjunction with isometric bar pressing with the forelimb. Cells in this caudal region are activated by cutaneous stimuli. In the rostral part of MI, neurons discharge prior to and during forelimb force changes, begin to discharge earlier than in the caudal zone, and have non-cutaneous or unidentifiable receptive fields. These results suggest separate motor control functions for rostral and caudal parts of rat MI.

小脳片葉の入出力と機能局在

The cerebellar flocculus of the cat be divided into three zones (rostral, middle and caudal) anatomicaly and physiologically. The rostral zone receives climbing fiber input from the rostral part of the dorsal cap and the ventrolateral outgrowth, and sends output to the central part of the superior vestibular nucleus. The middle zone receives climbing fiber input from the caudal part of the dorsal cap and sends output to the medial vestibular nucleus. The caudal zone receives climbing fiber input from the ventrolateral outgrowth and the rostral part of the dorsal cap, and projects to the Y-group and the medial part of the subnucleus lateralis parvocellularis of the lateral cerebellar nucleus, Activation of the rostral, middle and caudal zones produces an upward, an ipsilateral horizontal and a downward eye movement, respectively.

Regional differences in the inductive activity of the mesenchyme of the embryonic mouse urogenital sinus

AbstractEmbryonic urogenital sinuses (UGS) of 16‐day‐old mice were divided into two or three zones (cranial half and caudal half; or cranial third, intermediate, and caudal thirds). Following tryptic digestion, these zones were separated into their mesenchymal (UGM) and epithelial (UGE) components. The cranial and caudal zones of the UGM were recombined separately with the different zones of the UGE or with adult mouse urinary bladder epithelium (BLE). Following four weeks of growth in syngeneic male hosts, tissue recombinants of cranial UGM + caudal UGE contained numerous prostatic ducts and urethral glands. Conversely, recombinants composed of caudal UGM + cranial UGE developed into fibromuscular tissue covered with urethral epithelium and did not contain prostate, but occasionally contained urethral glands. Urethral glands were usually present in grafts of cranial UGM + cranial UGE or in grafts of the intact intermediate third of the UGS. In heterotypic recombinants of the UGM + adult BLE, prostatic glands were induced only when cranial UGM was utilized. Urethral glands were not observed in tissue recombinants prepared with adult BLE. These data suggest that regional differences in mesenchymal inductive ability and epithelial responsiveness play a role in the harmonious morphogenesis of the male lower genitourinary tract.

Forebrain projections to the hypothalamic locomotor region in cats

Descending projections from the cortex and basal ganglia to the hypothalamic locomotor region were studied in cats by the retrograde axonal transport of horse-radish peroxidase method. Neurons forming direct projections to the physiologically identified hypothalamic locomotor region are diffusely scattered over different gyri, but predominantly in areas 4 and 6 of the motor cortex, and also in the entopeduncular nucleus. Powerful cortico- and pallido-hypothalamic projections mainly do not reach as far as the most caudal zones of the hypothalamus, where the region whose electrical stimulation evokes locomotion lies.

Two types of eye movements during electrical stimulation of the rostral and caudal parts of the medial frontal "oculomotor" area in cats

The character of eye movements during electrical stimulation of the medial wall of the brain beneath the cruciate sulcus and of the inferior wall of the cruciate sulcus itself in the frontal cortex was investigated in waking cats. Stimulation of this part of the brain evoked two types of eye movements: unidirectional concomitant saccades, whose direction and amplitude were independent of the original position of the eyes in the orbits, and saccades into the central position (so-called centering saccades). Unidirectional saccades appeared in response to stimulation of the caudal part of the investigated zone, centering saccades in response to stimulation of its rostral part. Analysis of the directions and amplitudes of unidirectional saccades suggested the retinotopic organization of the caudal zone. Systematic changes in the magnitude and direction of vertical saccades during stimulation of the deep parts of the cruciate sulcus indicated previsely the projection of the vertical meridian of the retina. Reappearance of vertical saccades evoked by stimulation of certain parts of the medial wall of the brain suggests the existence of more than one retinotopically organized zone in this region. Inconstancy of stereotaxic coordinates of the oculomotor area, studied in different animals, was noted.

Inhibition of jaw opening reflexes by stimulation of the central gray matter and raphe nuclei in cats

The effect of stimulation of the mesencephalic central gray matter and raphe nuclei on jaw opening reflexes evoked by excitation of high-threshold (dental pulp) and low-threshold (A-alpha) fibers of the infraorbital nerve afferents was studied in cats anesthetized with chloralose and pentobarbital. The jaw opening reflex evoked by stimulation of the dental pulp was shown to be effectively suppressed by conditioning stimulation of the central gray matter and raphe nuclei. The reflex evoked by stimulation of low-threshold infraorbital nerve afferents also was depressed (but less deeply and for a shorter period than the reflex evoked by stimulation of the dental pulp) during stimulation of the raphe nuclei and caudal zone of the central gray matter, but was unchanged after stimulation of the points located in the rostral zone of the central gray matter. Application of single stimuli or bursts of five stimuli with a frequency of 100 Hz had no effect on the reflexes studied. Short-term stimulation with a burst of 10–20 stimuli with a following frequency of 200–400 Hz led to inhibition of the reflexes, which lasted 450–1000 msec. Long-term stimulation of the central gray matter and raphe nuclei for 30 sec with a frequency of 50 Hz caused inhibition of jaw opening reflexes evoked by stimulation of both high- and low-threshold afferents for 60 min. Impulses from the central gray matter and raphe nuclei thus have a mainly inhibitory action on the jaw opening reflex evoked by stimulation of high-threshold afferents, but they act less effectively on the reflex evoked by stimulation of low-thres-hold afferents. The duration of inhibition depends essentially on the parameters of stimulation.

Neuronal pathway from floccular caudal zone contributing to vertical eye movements in cats—role of group y nucleus of vestibular nuclei

In ketamine-anesthetized cats, electric microstimulation of the group y 1 nucleus of the vestibular nuclei evoked a slow and smooth upward eye movement. Destruction of the group y nucleus eliminated a slow and smooth downward eye movement evoked by stimulation of the caudal zone 11 of the flocculus. These data support the interpretation that Purkinje cell activity in the caudal zone of the flocculus can evoke vertical eye movements by inhibiting the activity of neurons of the group y nucleus.

Differences in the neurons that project from the dorsal column nuclei to the diencephalon, pretectum, and tectum in the cat.

The dorsal column nuclei (DCN) project to a number of targets in the nervous system besides the ventroposterolateral nucleus (VPL) of the thalamus. Recent evidence obtained using double-labeling techniques indicates that DCN's diencephalic-projecting neurons differ in their location and morphology from those that project to some of its other targets, such as the cerebellum and tectum. The purpose of the present study was to characterize anatomically the DCN neurons that project another of DCN's targets, the pretectum, and to determine if any of these neurons have collateral projections to the tectum or diencephalon. The projections were studied using two double-labeling methods. One method made use of either tritiated inactivated horseradish peroxidase ([3H]apoHRP) or tritiated N-acetyl wheatgerm agglutinin ([3H]WGA) as a marker and HRP or WGA conjugated to HRP. The other method made use of the dyes Fast Blue and Nuclear Yellow. In each cat, one marker was injected into the DCN-recipient portions of the pretectum, tectum, or diencephalon, and the other marker was injected into another of these three targets. Neurons labeled by pretectal or tectal injections were of all sizes, fusiform and multipolar in shape, and similarly located. They were scattered through the rostral zone of DCN, but were distributed at the periphery of and at the junction between the gracile and cuneate nuclei in DCN's middle and caudal zones. In contrast to the pretectal- and tectal-labeled neurons, neurons labeled by diencephalic injections were round and large. They were found throughout the DCN complex, but were concentrated in DCN's middle and caudal zones. When both the pretectum and diencephalon were injected in the same cat, the two groups of neurons occupied similar locations in the rostral zone, but were distinct in the middle and caudal zones, with the pretectal-projecting neurons surrounding the clusters of diencephalic-projecting neurons. Very few neurons were double-labeled. These results demonstrate that the projections to the pretectum, tectum, and diencephalon originate from different populations of neurons within specific domains in DCN. When these results are compared with the results of electrophysiological and other anatomical studies, it appears that the pretectal- and tectal-projecting neurons may be part of a previously unrecognized system originating in DCN.(ABSTRACT TRUNCATED AT 400 WORDS)

眼球運動に関連したネコ小脳片葉の機能的局在

The cat's cerebellar flocculus can be anatomically divided into three microzones on the basis of differences of their efferent projection sites. We investigated the eye moverents evoked by electric stimulation of each microzone of the flocculus in ketamine-anesthetized cats.An upward eye movement was evoked by stimu-lation of the rostral zone, an ipsilateral horizontal eye movement by stimulation of the middle zone, and a downward eye movement by stimulation of the caudal zone. Stimulation of the flocculus might activate Purkinje cells which inhibit the vestibular nucleus neurons and might cause eye movement.An upward eye movement was evoked by electric microstimulation of group y of the vestibular nuclei. Destruction of group y eliminated the downward eye movement evoked by stimulation of the caudal zone of the flocculus. Purkinje cells in the caudal zone might inhibit group y neurons and might contribute to vertical eye movements in cats.

Functional localization in the three floccular zones related to eye movement control in the cat

Anatomically, the cat's cerebellar flocculus can be divided into 3 zones on the basis of differences in their efferent projection sites 13,14. The functional differences of these 3 zones in relation to eye movement control were investigated by observing the eye movements evoked by electric stimulation of each zone of the flocculus in ketamine-anesthetized cats. Stimulation of the flocculus elicited a slow eye movement. The direction of the slow eye movement was mapped. A downward eye movement was evoked by stimulation of the caudal zone. An ipsilateral horizontal eye movement was induced from the middle zone. An upward eye movement was elicited from the rostral zone. When prolonged stimulation was applied to the flocculus, the slow eye movement was followed by nystagmus in the opposite direction. This nystagmus persisted for many seconds after cessation of stimulation (afternystagmus). Nystagmus and afternystagmus could not be elicited in deeply anesthetized cats. Possibilities as to how the stimulation leads to various eye movements are discussed.

Binaural interaction in low-frequency neurons in inferior colliculus of the cat. I. Effects of long interaural delays, intensity, and repetition rate on interaural delay function.

Detailed, quantitative studies were made of the interaural phase sensitivity of 197 neurons with low best frequency in the inferior colliculus (IC) of the barbiturate-anesthetized cat. We analyzed the responses of single cells to interaural delays in which tone bursts were delivered to the two ears via sealed earphones and the onset of the tone to one ear with respect to the other was varied. For most (80%) cells the discharge rate is a cyclic function of interaural delay at a period corresponding to that of the stimulating frequency. The cyclic nature of the interaural delay curve indicates that these cells are sensitive to the interaural phase difference. These cells are distributed throughout the low-frequency zone of the IC, but they are less numerous in the medial and caudal zones. Cells with a wide variety of response patterns will exhibit interaural phase sensitivities at stimulating frequencies up to 3,100 Hz, although above 2,500 Hz the number of such cells decrease markedly. Using dichotic stimuli we could study the cell's sensitivity to the onset delay and interaural phase independently. The large majority of IC cells respond only to changes in interaural phase, with no sensitivity to the onset delay. However, a small number (7%) of cells exhibit a sensitivity to the onset delay as well as to the interaural phase disparity, and most of these cells show an onset response. The effects of changing the stimulus intensity equally to both ears or of changing the interaural intensity difference on the mean interaural phase were studied. While some neurons are not affected by level changes, others exhibit systematic phase shifts for both average and interaural intensity variations, and there is a continuous distribution of sensitivities between these extremes. A few cells also showed systematic changes in the shape of the interaural delay curves as a function of interaural intensity difference, especially at very long delays. These shifts can be interpreted as a form of time-intensity trading. A few cells demonstrated orderly changes in the interaural delay curve as the repetition rate of the stimulus was varied. Some of these changes are consonant with an inhibitory effect that occurs at stimulus offset. The responses of the neurons show a strong bias for stimuli that would originate from he contralateral sound field; 77% of the responses display mean interaural phase angles that are less than 0.5 of a cycle, which are delays to the ipsilateral tone.(ABSTRACT TRUNCATED AT 400 WORDS)