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In four squirrel monkeys ( Saimiri sciureus), the tracer biotin dextranamine (BDA) was injected into the ventrolateral pons at a site at which injection of the glutamate antagonist kynurenic acid blocked vocalization electrically elicited from the periaqueductal gray (PAG). Anterograde projections could be traced into all cranial motor and sensory nuclei involved in phonation, that is, the nucleus ambiguus, facial, hypoglossal and trigeminal motor nuclei, the motorneuron column in the ventral gray substance innervating the extrinsic laryngeal muscles, the nucleus retroambiguus, solitary tract and spinal trigeminal nuclei. Projections were also found into a number of auditory nuclei, namely the nucleus cochlearis-complex, superior olive, ventral and dorsal nuclei of the lateral lemniscus and inferior colliculus. Furthermore, there were projections into the reticular formation of the lateral and dorsocaudal medulla and lateral pons, into nucleus gracilis, inferior and medial vestibular nuclei, lateral reticular nucleus, ventral raphe, pontine gray, superior colliculus, PAG and mediodorsal thalamic nucleus. Injection of the tracer wheat germ agglutinin-conjugated horseradish peroxidase into the ventrolateral pontine vocalization-blocking area in one animal yielded retrograde labeling throughout the PAG. Injection of BDA into a vocalization-eliciting site of the PAG in another animal yielded projections into the ventrolateral pontine vocalization-blocking area. It is concluded that the ventral paralemniscal area in the ventrolateral pons represents a relay station of the descending periaqueductal vocalization-controlling pathway. [ABSTRACT FROM AUTHOR]

The aim of this study was to determine if the angular vestibulo-ocular reflex (VOR) in response to pitch, roll, left anterior-right posterior (LARP), and right anterior-left posterior (RALP) head rotations exhibited the same linear and nonlinear characteristics as those found in the horizontal VOR. Three-dimensional eye movements were recorded with the scleral search coil technique. The VOR in response to rotations in five planes (horizontal, vertical, torsional, LARP, and RALP) was studied in three squirrel monkeys. The latency of the VOR evoked by steps of acceleration in darkness (3,000°/s² reaching a velocity of 150°/s) was 5.8±1.7 ms and was the same in response to head rotations in all five planes of rotation. The gain of the reflex during the acceleration was 36.7 ±15.4% greater than that measured at the plateau of head velocity. Polynomial fits to the trajectory of the response show that eye velocity is proportional to the cube of head velocity in all five planes of rotation. For sinusoidal rotations of 0.5-15 Hz with a peak velocity of 20°/s, the VOR gain did not change with frequency (0.74±0.06, 0.74 ±0.07, 0.37±0.05, 0.69±0.06, and 0.64±0.06, for yaw, pitch, roll, LARP, and RALP respectively). The VOR gain increased with head velocity for sinusoidal rotations at frequencies ≥4 Hz. For rotational frequencies ≥4 Hz, we show that the vertical, torsional, LARP, and RALP VORs have the same linear and nonlinear characteristics as the horizontal VOR. In addition, we show that the gain, phase and axis of eye rotation during LARP and RALP head rotations can be predicted once the pitch and roll responses are characterized. [ABSTRACT FROM AUTHOR]

The contribution of neck proprioceptive signals to signal processing in the vestibular nucleus was studied by recording responses of secondary horizontal canal-related neurons to neck rotation in the squirrel monkey. Responses evoked by passive neck rotation while the head was held stationary in space were compared with responses evoked by passive whole body rotation and by forced rotation of the head on the trunk. Most neurons (76%; 45/59) were sensitive to neck rotation. The nature and strength of neck proprioceptive inputs varied and usually combined linearly with vestibular inputs. In most cases (94%), the direction of the neck proprioceptive input was "antagonistic" or "reciprocal" with respect to vestibular sensitivity and, consequently, reduced the vestibular response during head-on-trunk rotation. Different types of vestibular neurons received different types of proprioceptive input. Neurons whose firing behavior was related to eye position (position-vestibular-pause neurons and position-vestibular neurons) were often sensitive to the position of the head with respect to the trunk. The sensitivity to head position was usually in the same direction as the neuron's eye position sensitivity. Non-eye-movement related neurons and eye-head-velocity neurons exhibited the strongest sensitivity to passive neck rotation and had signals that were best related to neck velocity. The results suggest that neck proprioceptive inputs play an important role in shaping the output of the primate vestibular nucleus and its contribution to posture, gaze and perception. [ABSTRACT FROM AUTHOR]

Eye position and angular velocity were measured in squirrel monkeys ( Saimiri sciureus) by means of the electromagnetic scleral search-coil technique. Horizontal sigma-optokinetic nystagmus (sigma-OKN) was elicited by a stationary, stroboscopically illuminated, periodic, vertical-stripe pattern lining a vertical cylinder. The relationship between the mean slow-phase eye angular velocity, Ve, of sigma-OKN and the product of pattern period, Ps, and flash frequency, fs, was determined. When Ve approximated k·Ps·fs (deg·s–1) and k was an integer ≥l, the sigma-paradigm was fulfilled. Sigma-OKN could be evoked in different ”modes”, whereby k approximated 1, 2,... n. The sigma-OKN properties of squirrel monkeys were similar to those measured for sigma-OKN in the ”stare” mode in man, with the exception of a long-lasting optokinetic afternystagmus (sigma-OKAN) appearing in the monkey. A considerable amount of temporal variability in flash sequence intervals (”temporal noise”), causing retinal error signals that interfered with the sigma-paradigm, was accepted by the visuo-motor system without interruption of sigma-OKN. This observation was explained by the operation of a short memory device for perception of visual motion. The internal gain, gi, which relates the retinal ”error” displacement velocity, Vr, and Ve depended, in turn, on Vr according to a function resembling the known relationship between neuronal activity of NOT (nucleus of the optic tract) nerve cells and Vr. This observation may be taken as direct proof that sigma-OKN can be explained by a centrally preprogrammed relationship between the retinal velocity, Vr, and the OKN slow-phase eye velocity, Ve. It is stipulated that the sum of Vr and efference copy signals generated in cortical or subcortical gaze centers is the essential component controlling the perceived velocity of the sigma-movement, whereby a short-term integrator plays a role for squirrel monkey sigma-OKN. When the flash frequency, fs, was modulated periodically according to a sinewave or ”triangular” function at a rate below 0.5 cycles·s–1, Ve was found to respond with a corresponding modulation, provided the modulation amplitude did not exceed 50% of the mean flash rate. When the latter occurred, nonlinear responses could be observed. A similar response was found when the speed of ”real” optokinetic stimuli was varied sinusoidally. Under these experimental conditions, however, the amplitude of the Ve variation yielded up to 1.0 approximately linear responses. [ABSTRACT FROM AUTHOR]

Natural head movements include angular and linear components of motion. Two classes of vestibulo-ocular reflex (VOR), mediated by the semicircular canals and otoliths (the angular and linear VOR, or AVOR and LVOR, respectively), compensate for head movements and help maintain binocular fixation on targets in space. In this study, AVOR/LVOR interactions were quantified during complex head motion over a broad range of fixation distances at a fixed stimulus frequency of 4.0 Hz. Binocular eye movements were recorded (search-coil technique) in squirrel monkeys while fixation distance (assessed by vergence) was varied using brief presentations of earth-fixed targets at various distances. Stimuli consisted of rotations around an earth-vertical axis and therefore always activated the AVOR. Horizontal and vertical AVORs were assessed when the head was centered over the axis of rotation and oriented upright (UP) and right-side-down (RD), respectively. AVOR gains increased slightly with increasing vergence in darkness, as expected given the small anterior position of the eyes in the head. Combined AVOR/LVOR responses were recorded when subjects were displaced eccentrically from the rotation axis. Eccentric rotations activated the AVOR just as when the head was centered, but added a translational stimulus which generated an LVOR component in response to interaural (IA) or dorsoventral (DV) tangential accelerations, depending on whether the head was UP or RD, respectively. When the head was eccentric and facing nose-out, the AVOR and LVOR produced ocular responses in the same plane and direction (coplanar and synergistic), and response magnitudes increased with increasing vergence. With the head facing nose-in, AVOR and LVOR response components were oppositely directed (coplanar and antagonistic). The AVOR dominated the response when fixation distance was far, and phase was compensatory for head rotation. As fixation distance decreased toward the rotation axis, responses declined to near zero, and when fixation distance approached even closer, the LVOR component dominated and response phase inverted. The same pattern was observed for both horizontal (head UP) and vertical (head RD) responses. The LVOR was recorded directly by rotating subjects eccentrically but in the nose-up (NU) orientation. The AVOR then generated torsional responses to head roll, coexistent with either horizontal or vertical LVOR responses to tangential acceleration when the subject was oriented head-out or right-side-out, respectively. Only the LVOR response components were modulated by vergence. A vectorial analysis of AVOR, LVOR, and combined responses supports the conclusion that AVOR and LVOR response components combine linearly during complex head motion. [ABSTRACT FROM AUTHOR]

The discharge of neurons in the vestibular nuclei was recorded in alert squirrel monkeys while they were being sinusoidally rotated at 2 Hz. Type I position-vestibular-pause (PVP I) and vestibular-only (V I) neurons, as well as a smaller number of other type I and type II eye-plus-vestibular neurons were studied. Many of the neurons were monosynaptically related to the ipsilateral vestibular nerve. Eye-position and vestibular components of the rotation response were separated by multiple regression. Anodal currents, simultaneously delivered to both ears, were used to eliminate the head-rotation signals of irregularly discharging ( I) vestibular-nerve afferents, presumably without affecting the corresponding signals of regularly discharging ( R) afferents. R and I inputs to individual central neurons were determined by comparing rotation responses with and without the anodal currents. The bilateral currents, while reducing the background discharge of all types of neurons, did not affect the mean vestibular gain or phase calculated from a population of PVP I neurons or from a mixed population consisting of all type I units. From this result, it is concluded that I inputs are canceled at the level of secondary neurons. The cancellation may explain why the ablating currents do not affect the gain and phase of the vestibulo-ocular reflex. While cancellation was nearly perfect on a population basis, it was less so in individual neurons. For some neurons, the ablating currents decreased vestibular gain, while for other neurons the vestibular gain was increased. The former neurons are interpreted as receiving a net excitatory (I-EXC) I input, the latter neurons, a net inhibitory (I-INH) input. When compared with the corresponding R inputs, the I inputs were usually small and phase advanced. Phase advances were larger for I-EXC than for I-INH inputs. The sign and magnitude of the I inputs were unrelated to other discharge properties of individual neurons, including discharge regularity and the phase of vestibular responses measured in the absence of the ablating currents. Unilateral currents were used to assess the efficacy of ipsilateral and contralateral pathways. Ipsilateral pathways were responsible for almost all of the effects seen with bilateral currents. The results suggest that the vestibular signals carried by central neurons, even by those neurons receiving a monosynaptic vestibular-nerve input, are modified by polysynaptic pathways. [ABSTRACT FROM AUTHOR]

The vestibulo-ocular reflex (VOR) was studied in three squirrel monkeys subjected to rotations with the head either centered over, or displaced eccentrically from, the axis of rotation. This was done for several different head orientations relative to gravity in order to determine how canal-mediated angular (aVOR) and otolithmediated linear (lVOR) components of the VOR are combined to generate eye movement responses in three-dimensional space. The aVOR was stimulated in isolation by rotating the head about the axis of rotation in the upright (UP), right-side down (RD), or nose-up (NU) orientations. Horizontal and vertical aVOR responses were compensatory for head rotation over the frequency range 0.25-4.0 Hz, with mean gains near 0.9. The horizontal aVOR was relatively constant across the frequency range, while vertical aVOR gains increased with increasing stimulation frequency. In the NU orientation, compensatory torsional aVOR responses were of relatively low gain (0.54) compared with horizontal and vertical responses, and gains remained constant over the frequency range. When the head was displaced eccentrically, rotation provided the same angular stimuli but added linear stimulus components, due to the centripetal and tangential accelerations acting on the head. By manipulating the orientation of the head relative to gravity and relative to the axis of rotation, the lVOR response could be combined with, or isolated from, the aVOR response. Eccentric rotation in the UP and RD orientations generated aVOR and lVOR responses which acted in the same head plane. Horizontal aVOR-lVOR interactions were recorded when the head was in the UP orientation and facing toward ('nose-in') or away from ('nose-out') the rotation axis. Similarly, vertical responses were recorded with the head RD and in the nose-out or nose-in positions. For both horizontal and vertical responses, gains were dependent on both the frequency of stimulation and the directions and relative amplitudes of the angular and linear motion components. When subjects were positioned nose-out, the angular and linear stimuli produced synergistic interactions, with the lVOR driving the eyes in the same direction as the aVOR. Gains increased with increasing frequency, consistent with an addition of broad-band aVOR and high-pass lVOR components. When subjects were nose-in, angular and linear stimuli generated eye movements in opposing directions, and gains declined with increasing frequency, consistent with a subtraction of the lVOR from the aVOR. This response pattern was identical for horizontal and vertical eye movements. aVOR and lVOR interactions were also assessed when the two components acted in orthogonal response planes. By rotating the monkeys into the NU orientation, the aVOR acted primarily in the roll plane, generating torsional ocular responses, while the translational (lVOR) component generated horizontal or vertical ocular responses, depending on whether the head was oriented such that linear accelerations acted along the interaural or dorsoventral axes, respectively. Horizontal and vertical lVOR responses were negligible at 0.25 Hz and increased dramatically with increasing frequency. Comparison of the combined responses (UP and RD orientations) with the isolated aVOR (head-centered) and lVOR (NU orientation) responses, indicates that these VOR components sum in a linear fashion during complex head motion. [ABSTRACT FROM AUTHOR]

The neural response to amplitude-modulated sinus sounds (AM sound) was investigated in the auditory cortex and insula of the awake squirrel monkey. It was found that 78.1% of all acoustically driven neurons encoded the envelope of the AM sound; the remaining 21.9% displayed simple On, On/Off or Off responses at the beginning or the end of the stimulus sound. Those neurons with AM coding were able to encode the AM sound frequency in two different ways: (1) the spikes followed the amplitude modulation envelopes in a phaselocked manner; (2) the spike rate changed significantly with changing modulation frequencies. As reported in other species, the modulation transfer functions for rate showed higher modulation frequencies than the phaselocked response. Both AM codings exhibited a filter characteristic for AM sound. Whereas 46.6% of all neurons had the same filter characteristic for both the spike discharge and the phase-locked response, the remaining neurons displayed combinations of different filter types. The discharge pattern of a neuron to simple tone or noise bursts suggests the behaviour of this neuron when AM sound is used as the stimulus. Neurons with strong onset responses to tone/noise bursts tended to have higher phase-locked AM responses than neurons with weak onset responses. The spike rate maxima for AM sound showed no relation to the tone/noise burst discharge patterns. Varying modulation depth was encoded by the neuron's ability to follow the envelope cycles and not by the non-phase-locked spike rate frequency. The organization of the squirrel monkey's auditory cortex has previously been established by an anatomical study. We have added two new fields using physiological parameters. All fields investigated showed a clear functional separation for time-critical information processing. The best temporal resolution was shown by the primary auditory field (AI), the first-temporal field (T1) and the parainsular au ditory field (Pi). The neural data in these fields and the amplitude modulation frequency range of squirrel monkey calls suggest a similar correlation between vocalization and perception as in human psychophysical data for speech and hearing sensation. The anterior fields in particular failed to follow the AM envelopes. For the first time in a primate, the insula was tested with different sound parameters ranging from simple tone bursts to AM sound. It is suggested that this cortical region plays a role in time-critical aspects of acoustic information processing. The observed best frequencies covered the same spectrum as AI. As in the auditory fields, most neurons in the insula encoded AM sound with different filter types. The high proportion of neurons unable to encode AM sound (40.6%) and the low mean best modulation frequency (9.9 Hz) do not support a prominent role of the insula in temporal information processing. [ABSTRACT FROM AUTHOR]

In 11 squirrel monkeys (Saimiri sciureus), the brain stem was systematically explored with electrical brain stimulation for sites affecting the acoustic structure of ongoing vocalization. Vocalization was elicited by electrical stimulation of different brain structures. A severe deterioration of the acoustical structure of vocalization was obtained during stimulation of the caudoventral part of the periaqueductal grey, lateral parabrachial area, corticobulbar tract, nucl. ambiguus and surrounding reticular formation, facial nucleus, hypoglossal nucleus, solitary tract nucleus and along the fibres crossing the midline at the level of the hypoglossal nucleus. It is suggested that these structures are part of, or at least have direct access to, the motor coordination mechanism of phonation. Complete inhibition of phonation was obtained from the raphe and raphe-near reticular formation. [ABSTRACT FROM AUTHOR]

The brain stems of 17 squirrel monkeys ( Saimiri sciureus) were systematically explored for vocalization-related single-unit activity during calls electrically elicited from the periaqueductal grey. Of 12,280 cells tested, 1151 fired in relation to vocalization. Of these, 587 reacted to external acoustic stimuli and started firing after vocalization onset. As most of these cells were located in classical auditory relay structures, they probably represent auditory neurones reacting indirectly to self-produced vocalization due to auditory feedback. Seven cells reacted to acoustic stimuli but fired in advance of self-produced vocalization. These cells were locoated in the pericentral inferior colliculus, dorsal nucleus of the lateral lemniscus, dorsomedial to the ventral nucleus of the lateral lemniscus and immediately lateral to the central grey. They are probably engaged in tuning the auditory system to process self-generated sounds differently from external sounds. 261 neurones reacted to nonphonatory oral movements (chewing, swallowing) and started firing after vocalization onset. These neurones were widely distributed within the brain stem, with the highest density in the spinal trigeminal nucleus and medially adjacent reticular formation. The majority of these cells seem to react to proprioceptive and tactile stimuli generated by phonatory and nonphonatory oral activities. Some of them may exert motor control on muscles that come into play at later stages of phonation. 57 neurones reacted to nonphonatory oral movements but fired in advance of vocalization onset. These neurones were located mainly in the trigeminal motor nucleus, nucl. ambiguus, reticular formation around these nuclei, parabrachial region and lateral vestibular nucleus. Their role in motor control seems to be related to specific muscles rather than specific functions. 100 of the vocalization-related cells showed a correlation with respiration. Expiration-related cells were found in and around the rostral nucl. ambiguus and in the reticular formation dorsal to the facial nucleus. Inspiration-related cells were located in the rostral and caudal nucl. ambiguus regions, ventrolateral solitary tract nucleus and the lateral reticular formation below the trigeminal motor nucleus. Most of these cells probably represent premotor neurones of respiratory muscles and laryngeal motoneurones of the cricothyroid and posterior cricoarytenoid muscles. Finally, a last group of cells was found that was unresponsive to chewing and swallowing movements, quiet breathing and acoustic stimuli, but changed activity during vocalization. 38 of them became active before vocalization and cricothyroid activity, and 101 afterward. Both types were completely intermingled and scattered widely in the brain stem, including the nucl. ambiguus region, solitary tract nucleus, nucl. reticularis parvocellularis and gigantocellularis, parabrachial region, pericentral colliculus inferior, vestibular complex, periventricular grey and laterally adjacent tegmentum. Some of these cells may be related to vocalization in a more specific way. [ABSTRACT FROM AUTHOR]

Squirrel monkeys were trained to cancel their vestibulo-ocular reflex (VOR) by fixating a visual target that was head stationary during passive vestibular stimulation. The monkeys were seated on a vestibular turntable, and their heads were restrained. A small visual target (0.2°) was projected from the vestibular turntable onto a tangent screen. The monkeys' ability to suppress their VOR by fixating a head stationary target while the turntable was moving was compared to their ability to pursue the target when it was moved in the same manner. Squirrel monkeys were better able to suppress their VOR when the turntable was moved at high velocities than they were able to pursue targets that were moving at high velocities. The gaze velocity gain during VOR cancellation began to decrease when the head velocity was above 80°/s, and was greater than 0.6 when the head velocity was above 150°/s. However, gaze velocity gain during smooth pursuit decreased significantly when the target velocity was greater than 60°/s, and was less than 0.4 when the target velocity was 150°/s or more. The latency of VOR suppression was significantly shorter than the latency of smooth pursuit while the monkey was cancelling its VOR. When an unpredictable step change in head acceleration was generated while the monkey was cancelling its VOR, the VOR evoked by the head acceleration step began to be suppressed shortly after the initiation of the step (≈ 30 ms). On the other hand, the latency of the smooth pursuit eye movement elicited when the visual target was accelerated in the same manner during VOR cancellation was ≈ 100 ms. The comparison between these two results suggests that the monkeys did not use visual information related to target motion to suppress their VOR at an early latency. The monkeys' ability to suppress the VOR evoked by an unexpected change in head acceleration depended on the size of the head acceleration step. The VOR evoked by unexpected step changes in head acceleration was progressively less suppressed at an early latency as the size of the acceleration step increased, and was not suppressed at an early latency when the step change in head acceleration was greater than 500°/s. During smooth pursuit eye movements, unexpected step changes in head acceleration evoked a VOR that was suppressed at an early latency (≈ 50 ms) if the head movement was in the same direction as the ongoing smooth pursuit eye movement. The amount of early VOR suppression increased as the pursuit eye velocity increased. We conclude that squirrel monkeys utilize a fast, non-visual mechanism for cancelling their VOR while they are fixating a visual target and their head is moving. This non-visual mechanism appears to be turned on when the head is moving and the monkey is fixating a head stationary target. The mechanism probably utilizes a voluntarily gated vestibular signal to cancel the signals in VOR pathways at the level of the extraocular motorneurons. Although the VOR cancellation mechanism is not capable of completely suppressing the VOR evoked by large unexpected changes in head acceleration, we suggest that it is capable of suppressing the VOR generated by most voluntary head movements during combined eye and head gaze pursuit and that the function of this gated VOR cancellation system is to extend the range and accuracy of eye-head tracking movements. [ABSTRACT FROM AUTHOR]

Recordings were made from secondary vestibular axons in the medial longitudinal fasciculus (MLF) of barbiturate-anesthetized squirrel monkeys. Antidromic stimulation techniques were used to identify the axons as belonging to one of three classes of neurons: vestibulo-oculo-collic (VOC) neurons project both to the extraocular motor nuclei and to the spinal cord; vestibulo-ocular (VO) neurons do not have a spinal projection; and vestibulocollic (VC) neurons do not have an oculomotor projection. Galvanic stimulation was used to show that axons of all three classes received excitatory inputs from one labyrinth and inhibitory inputs from the other. VOC axons were confined to the MLF contralateral to the labyrinth from which they were excited. They made up more than half of the vestibular axons descending in the contralateral medial vestibulospinal tract (MVST), but less than one-quarter of those ascending in the contralateral MLF to the level of the oculomotor nucleus. Spinal projections were restricted to cervical segments with about half of the axons reaching segment C6. Conduction velocities, measured for Co-projecting axons, were similar for VOC and VC axons and were typically 25-50 m/s. Unlike the situation in the rabbit (Akaike et al. 1973) and cat (Akaike 1983), none of the MVST axons had conduction velocities > 75 m/s. The morphology of VOC neurons was studied by injection of horseradish peroxidase (HRP) into 60 physiologically identified axons in the MLF. Since individual axons were only stained for short distances, it was not possible to ascertain their complete branching patterns. Labeled fibers could be traced to an origin in and around the ventral lateral vestibular nucleus. This localization was confirmed by comparing the distributions within the vestibular nuclei of neurons retrogradely labeled from the upper cervical spinal cord (this study) and from the oculomotor nucleus (McCrea et al. 1987a; Highstein and McCrea 1988). VOC axons reached the contralateral MLF at the level of the abducens nucleus and immediately divided into an ascending and a descending, usually thicker, branch. Seven VOC axons could be traced to the extraocular motor nuclei; three terminated in the medial aspect of the oculomotor nucleus bilaterally and four terminated in the medial aspect of the contralateral abducens nucleus. The former axons may be part of a crossed, excitatory anterior-canal pathway; the latter, part of a similar horizontal-canal pathway. There were no terminations in the trochlear nucleus even though 12 labeled fibers passed close to it. VOC axons projected to several brainstem nuclei, including the contralateral interstitial nucleus of Cajal, cell groups in the region of the medial longitudinal fasciculus rostral to the abducens nucleus, the nucleus prepositus, the nucleus raphe obscuris, Roller's nucleus, and the paramedian medullary reticular formation. Virtually all of the above connections, except for the bilateral projection to the oculomotor nucleus, were contralateral to the cells of origin. The results in the squirrel monkey are compared with previous studies of VOC neurons in the cat (Isu and Yokota 1983; Uchino and Hirai 1984; Isu et al. 1988). In both species, VOC neurons make up a large proportion of contralaterally projecting MVST fibers. On the other hand, such dual-projecting neurons may provide a considerably smaller fraction of the secondary vestibular axons reaching the oculomotor nucleus in the monkey than they do in the cat. [ABSTRACT FROM AUTHOR]

We studied the influence of static head position on the horizontal nystagmus produced by caloric, rotational and optokinetic stimulation in alert squirrel monkeys. Caloric nystagmus is stronger for nose up (NU) than for nose down (ND) pitches; so, for example, slow-phase eye velocity is four times larger in supine than in prone positions. A similarly directed asymmetry occurs in the horizontal vestibulo-ocular (HVOR) responses to longduration, constant angular-head accelerations, but not to midband (0.1 Hz) sinusoidal head rotations. Consistent with a first-order model of the HVOR, the low-frequency or acceleration gain of the reflex (G) is equal to the product of the midband velocity gain (G) and a time constant (T). G is proportional to the cosine of the angle between the horizontal-canal plane and the plane of rotation, from which it is concluded that signals from the horizontal, but not from the vertical canals contribute to the HVOR. T can be as much as twice as large in NU than in ND positions. G is proportional to T and it, too, shows a NU-ND asymmetry. The time constant of optokinetic afternystagmus (T) was also studied. Since T and T are modified in similar ways by static tilts, it is concluded that head position affects the time constants by way of velocity-storage mechanisms. Evidence is presented that the position-dependent modification of velocity storage is otolith-mediated. The results are used to analyze the mechanisms of caloric nystagmus. The caloric response consists of a convective component (C), as originally envisioned by Bárány (1906), and a nonconvective component (N). C accounts for 75% of the caloric response in the conventional supine testing position. Both components can be affected by the position-dependent modification of T or, equivalently, of G. It has been suggested that two mechanisms might contribute to N: 1) a direct thermal effect on hair cells or afferents; or 2) a thermal expansion of labyrinthine fluids that results in a cupular displacement. Both theoretical and experimental evidence indicates that only the first of these mechanisms could result in the steady-state caloric response that is observed in the absence of convection (e.g., in spaceflight and after canal plugging) and that contributes to the prone-supine asymmetry seen in caloric testing. [ABSTRACT FROM AUTHOR]

The responses of single and multi units in the medial geniculate body of the squirrel monkey ( Saimiri sciureus) to modulation frequency, modulation depth and changes in absolute intensity of sinusoidally amplitude modulated (AM) sounds were studied. Both spike-frequency and spike rate modulation were used as a measure for neuronal response. Spike rate modulation was derived from FFT (Fast-Fourier-Transformation) analysis of the PSTHs. In all cases ( n = 133) spike rate modulation was shown to be dependent on the stimulus modulation frequency: Most neurons responded best to one modulation frequency, i.e., they showed a modulation transfer function with bandpass characteristic; only a few displayed a low pass or multiple peaked transfer characteristic. The majority of the neurons responded best in a range from 4 to 64 Hz, with a peak at 32 Hz and a median at 16 Hz. Such modulation frequencies are common in parts of the species vocal repertoire. [ABSTRACT FROM AUTHOR]

Pure tone bursts elicited in primary auditory cortex (AI) cells of the awake squirrel monkey a wide range of response patterns which consisted of one or more excitatory or inhibitory temporal response components. In almost 60% of these cells, response patterns were frequency and/or intensity dependent. Response components such as early and late onset excitation, offset excitation and on-off excitation; as well as tonic excitation or inhibition often varied independently with changes in these stimulus parameters. Individual cells were therefore considered as multiple bandpass filters, and each discrete response component was analyzed separately for its tuning properties. A correlation between best frequencies of the various excitatory components (BEF), and between BEFs and best frequencies of inhibitory components (BIF), in cells which responded with more than one discrete response component, disclosed a significantly higher correlation between BEF/BIF pairs compared with BEF/BEF pairs, presumably reflecting certain 'lateral inhibition like' processes. Applying Q factor, and √Hf-√Lf bandwidth at 10 dB above threshold, as measures of the 'sharpness' of response areas, revealed that approximately 65% of all response areas could be defined as 'narrow' by either one of these 2 measures, with no distinction, in that regard, between excitatory and inhibitory components. The average response bandwidths of the narrowly and the broadly tuned components, at 10 dB above threshold, were 0.4±0.18 and 1.42±0.68 octaves respectively. A comparison with the medial geniculate body (MGB) of the squirrel monkey, applying the √Hf-√Lf measure of sharpness of tuning, showed a significantly higher proportion of narrow response areas in the AI. 'Narrow' response areas in both these regions were equally narrow, whereas the 'broad' response areas of MGB cells were significantly broader. These results suggest a sharpening of response areas throughout the geniculo-cortical transformation. [ABSTRACT FROM AUTHOR]

In 10 squirrel monkeys (Saimiri sciureus) uni- or bilateral lesions were placed in the nucl. solitarius, parabrachial nuclei, nucl. ventralis posterior medialis thalami or face area of primary sensory cortex. The effects of these lesions on vocalization were compared with those after transection of the internal branch of the superior laryngeal nerve. It was found that neither the cortical nor thalamic or parabrachial lesions changed the acoustic structure of vocalization. In contrast, destruction of the nucl. solitarius, like transection of the internal branch of the superior laryngeal nerve, affected vocalization severely. It is concluded that the production of species-specific vocalization depends upon a di- or, possibly, tri-synaptic laryngeal reflex control from tactile and proprioceptive laryngeal mechanoreceptors via nucl. solitarius and, possibly, lateral medullary reticular formation to nucl. ambiguus. [ABSTRACT FROM AUTHOR]

In 39 squirrel monkeys (Saimiri sciureus), the effects of various brain lesions on vocalizations elicited from the precallosal cingulate gyrus were tested. It was found that lesions abolishing the 'cingular vocalization' completely can be traced from the stimulation site continuously down to the laryngeal motoneurons in the nucleus ambiguus. The pathway thus determined (Fig. 4) travels from the precallosal cingulate gyrus through the frontal white matter and enters the internal capsule from a dorsolateral position. The pathway then follows this structure in a medio-caudal direction down to the caudal diencephalon. Here, the effective lesions leave the corticospinal tract and ascend dorsally into the periaqueductal grey. The pathway follows this structure to its end where it sweeps lateral through the parabrachial area and then descends through the lateral pons and ventrolateral medulla to the nucleus ambiguus. In nine of the animals, in addition, the effects of bilateral anterior cingular lesions on vocalizations elicited in other brain areas were tested. It was found that the only vocalization-eliciting area which becomes ineffective after destruction of the anterior cingulate gyrus is the postero-medial orbital cortex. [ABSTRACT FROM AUTHOR]

Eleven squirrel monkeys (Saimiri sciureus) were trained to discriminate species-specific calls from non-species-specific complex sounds in a go, no-go procedure with social contact as positive reinforcement. The task required that the animals not only responded to a particular call but that this response should be generalized to any squirrel monkey call, whether or not it had been presented previously in training. After having reached a performance level of 75% correct responses in three consecutive sessions, seven animals received bilateral lesions of the auditory cortex; the other four animals served as controls. It was found that small lesions within the superior temporal gyrus did not interfere with the discrimination task. Lesions destroying about three quarters of the auditory cortex led to loss of retention; during retraining the animals did not reach criterion, but performed significantly above chance. These animals were able, however, to master a simplified version of the task where one species-specific call had to be discriminated from one non-species-specific sound. Animals with almost total ablation of the auditory cortex were capable of mastering neither the generalized task nor the simplified version. From these results, together with those of the literature, it is concluded 1) that recognition of complex sounds is not possible after complete auditory cortex ablation, probably because of interference with gestalt-formation processing, and 2) that species-specific calls are processed in the auditory system in the same way as other complex sounds. [ABSTRACT FROM AUTHOR]

The projections of four different sub-areas within the anterior limbic cortex, all yielding vocalization when electrically stimulated, were compared in six squirrel monkeys by the autoradiographic tracing technique. Areas of convergence of the projections from all four vocalization loci were the cortex within the anterior cingulate sulcus, a zone following the inferior thalamic peduncle from the central amygdaloid nucleus through the substantia innominata into the midline thalamus, a second zone following the periventricular fibre system from the anterior diencephalon to the caudal midbrain and dorsolateral pontine tegmentum and, finally, the tail of the caudate nucleus. Except for the latter, all of these brain structures produce vocalization when electrically stimulated. The call types elicitable from these projection areas are sometimes different from those elicitable from the anterior limbic cortex. It is hypothesized that the anterior limbic cortex controls vocalization directly, independently of the specific motivational state underlying it. [ABSTRACT FROM AUTHOR]

The projections from the cortical vocal fold area were studied in five squirrel monkeys (Saimiri sciureus) with the aid of the autoradiographie tracing technique. The location of the cortical vocal fold area was determined by exploring the exposed frontal cortex with roving electrodes while examining the larynx for vocal fold adduction. The following projections were found: To the orbital cortex (area 11), dorsomedial frontal cortex (areas 6 and 8), Broca's area (area 44), lower fronto-parietal cortex (areas 6, 4, 3 and 1), fronto-parietal operculum (area 50), insula (areas 14 and 13), caudatum, putamen, claustrum nucl. reticularis th., nucl. ventralis anterior, nucl. ventralis lateralis, nucl. ventralis posteromedialis, nucl. centralis inferior, nucl. centralis lateralis, nucl. medialis dorsalis, nucl. pulvinaris medialis, griseum pontis, nucl. parabrachialis medialis and lateralis, nucl. tr. spinalis n. trigemini and nucl. tr. solitarii. A comparison of this projection system with a previous mapping study for vocalization (Jürgens and Ploog, 1970) revealed that there are two areas yielding vocalization when electrically stimulated which receive direct projections from the cortical larynx area, namely, the cortex around the anterior sulcus cinguli and the parabrachial nuclei at the pons-midbrain transition. The possible relevance of these structures for vocalization is discussed. [ABSTRACT FROM AUTHOR]

In 38 squirrel monkeys 251 vocalization-producing electrode positions were tested for their positive and negative reinforcing properties. Two groups of vocalization-producing brain areas could be distinguished: One group in which the electrically elicited vocalization was independent of the accompanying reinforcement effect, and a second group in which vocalization and reinforcement effect were correlated. The first group included the anterior cingulate gyrus, the adjacent supplementary motor area, gyrus rectus, ventromedial edge of the capsula interna, caudal periaqueductal gray and adjacent parabrachial region. The second group consisted of the caudatum, septum, substantia innominata, amygdala, inferior thalamic peduncle, stria terminalis, midline thalamus, ventral and periventricular hypothalamus, substantia nigra, rostral periaqueductal gray, dorsolateral midbrain tegmentum and lateral medulla. It is hypothesized that the first group contains predominantly or exclusively 'primary' vocalization subtrates; the second group is thought to be composed mainly of structures whose stimulation yields vocalization secondarily due to stimulus induced motivational changes. [ABSTRACT FROM AUTHOR]

The responses of single units in the superior temporal gyrus of awake squirrel monkeys to tone pips, noise, clicks, and frequency-modulated sounds were recorded with extracellular microelectrodes. A majority of the units responded to acoustic stimulation, pure tone pips being the most effective in terms of the percentage of units responding (71%). No simple catalogue of response types could be elicited. Units varied in terms of the combinations of stimuli to which they were responsive, the frequency range over which pure tones were effective, and the temporal pattern of discharge to different frequencies and different types of stimuli. A substantial majority of units gave precise timed responses to the onset or offset of the stimuli, while a few introduced long delays between the stimulus and the response. With regard to the area studied, acoustic units could be found between stereotaxic coordinates A3 and A10, both on the lateral surface of the hemisphere and in the superior temporal plane, as well as in the caudal insular region. No precise tonotopic organization could be discerned. [ABSTRACT FROM AUTHOR]

Action potentials of single auditory cortical neurons of the squirrel monkey were recorded in a chronic, unanesthetized preparation. The responsiveness of units was tested with various types of simple and complex acoustic stimuli in a free field situation. As simple auditory stimuli, bursts of pure tones, clicks, and white noise were utilized. Species-specific vocalizations served as complex, biologically significant stimuli. The data are based on 48 neurons which showed a discrete response to speciesspecific vocalizations. In 63% the response to calls could be predicted from the units' responses to simple stimuli. Thirty-seven percent of the neurons were classified as unpredictable with respect to their responsiveness to vocalizations. The response of most units was restricted to call stimuli which showed similarities in their frequency-time characteristics. About 7% of the 116 units responding to calls were classified as 'selective responders' because they were not excited by any other stimulus tested. It was not possible to single out the acoustic features to which these units responded. [ABSTRACT FROM AUTHOR]

The technique of telestimulation was employed in studying the social behavior of squirrel monkeys (Saimiri sciureus). Within a group of five monkeys an animal is electrically stimulated. Its stimulus response and the consecutively occurring reactions of the non-stimulated animals caused by the stimulus response form a sequence of actions. Using stimulus parameters which remained constant, the stimulation of the same electrode in the same animal was repeated automatically. The sequences of actions accumulated thereby were then analysed mathematically on the basis of probability theory. The frequency distributions, the transition matrices and the courses of the frequencies of the actions over the sequences provided the basis for a classification of the behavior units. The courses of the values of entropy and conditional entropy provided summarized information about the sequential process of the animals' behavior. The study of the structure of the sequences was approached by appropriate tests. The applicability of the mathematical methods employed to the material at hand is discussed. [ABSTRACT FROM AUTHOR]

Social signals of squirrel monkeys (Saimiri sciureus) were elicited by remote-controlled brain stimulation. The method employed permits a study of signal-response interdependencies in varying social situations which can be chosen at the experimenter's will. The function of a signal and thereby its information content are largely determined by the response of the recipients. The analysis of signal function was attempted under five conditions of social interactions: (1) reactions of group members to attack behavior of the stimulated animal, (2) reactions to dominance gestures of the stimulated animal, (3) reactions to signals that are indiscernible to the human observer, (4) reactions that are unusual or abnormal, and (5) reactions to signals sent over greater or lesser distances. Several aspects of the signal proved to be decisive for the prediction of behavior: (1) the signal intensity, (2) the space between animals, and (3) the context. The control of these variables by means of radio stimulation permits further quantitative studies of signal characteristics and their effects on the modification of social behavior. [ABSTRACT FROM AUTHOR]

Detailed histochemical observations have been made on the various components of the spinal cord. The gray matter showed much greater enzymatic activity than the white matter. The neuropil and substantia gelatinosa showed mild G6PD, AChE and BChE activity; moderate SDH, LDH, SE and AC activity; and strong AP, ATPase, and AMPase activity. The pia-arachnoid showed moderate to strong activity of dehydrogenases and phosphatases. The cytoplasm of the neurons and neuronal processes gave moderate to very strong reaction for most of the enzymes except BChE. The areas of synaptic endings show the presence of oxidative enzymes, phosphorylases, ATPase and AMPase. It has been interesting to note that phosphorylases, which have been mostly observed in the cytoplasm, nucleoli and synaptic areas of various neurons and in the neuropil, also show intensely strong reactions in the nuclei of ependymal cells lining the central canal as well as in nuclei of some glial cells. [ABSTRACT FROM AUTHOR]

The distribution of succinic dehydrogenase (SDA) and cytochrome oxidase (Cy. O) in serial sections of the cervical region of the spinal cord and the medulla oblongata, arranged caudo-cranially, has been described. The motor cranial nerve nuclei exhibit strong SDA and Cy. O activity in the neurons and neuropil. The nuclei gracilis, cuneatus, olivaris inferior, cochlearis and vestibularis likewise show strong enzyme activity. Nucleus intercalatus and nucleus tractus solitarius, however, show weak and moderate enzyme activity respectively. The lateral part of formatio reticularis myelencephali shows less SDA and Cy. O compared to the medial part, which shows some accumulation of these enzymes in the neuropil. The neuropil of the molecular layer of cerebellar cortex and the perikarya and dendrites of the Purkinje cells show strong SDA and Cy. O activity. The granular layer exhibits stronger SDA and Cy. O in the synaptic glomeruli. The cerebellar nuclei possess stronger enzyme activity in the neurons and dendritic branches, compared to mild activity in the neuropil. [ABSTRACT FROM AUTHOR]

The distribution of succinic dehydrogenase (SDA) and cytochrome oxidase (Cy. O) has been investigated in a series of sections through the pons and mesencephalon of the squirrel monkey brain. The localization of the two enzymes is very similar in the various regions and shows only slight differences. The epiphysis, however, shows moderately strong SDA and very mild Cy. O activity. Particularly strong SDA and Cy. O activity has been observed in the cell bodies of the various cranial nerve nuclei, nucleus colliculi inferioris, colliculi superioris, nuclei griseum pontis, reticularis tegmenti pontis, lemnisci lateralis pars dorsalis, geniculatum laterale and mediale, and pulvinaris. The enzyme content of the neurons and cell bodies is generally stronger compared to the neuropil which often occurs in smooth, loose, compact and reticulated forms. Any special relationship between the neurons and neuropil with regard to their enzyme content has, however, not been observed. The cranial nerves, and fibers of the brachium conjunctivum, corpus callosum, and fornix show very mild enzyme activity except those of the trapezoid complex which show moderate enzyme activity. [ABSTRACT FROM AUTHOR]

Incremental brightness thresholds (DI) were psychophysically determined at several background illumination intensities for three squirrel monkeys. Gross asymmetrical electrodes were then chronically implanted in the lateral geniculate nucleus of the same animals, and activity was recorded in stimulus conditions identical to behavioral testing. Overall activity, recorded through an integrating voltmeter, showed 1. a tendency to decrease as steady background illumination increased, and 2. an abrupt transient increase at both onset and offset to DI test flashes, directly proportional to test flash intensity. Background illumination, in proportion to its intensity, depressed response to a superimposed test flash. Test flashes below intensity DI at the various levels of background illumination produced no measurable response. The quantity DI was shown to be a function of the depressive or inhibitory effect of background illumination on the capacity of the system to respond to transient stimulation. A secondary determinant of DI appeared to be the amount of variability in ongoing neural activity upon which the DI flash is imposed. The author is indebted to the supervisor of her dissertation, Dr. L. R. Pinneo, for introducing her to the recording technique and for his help towards the completion of this work. [ABSTRACT FROM AUTHOR]