Complex predictive eye pursuit in monkey: a model system for cerebellar studies of skilled movement.

Smooth pursuit eye movements provide a good model system for cerebellar studies of complex motor control in monkeys. First, the pursuit system exhibits predictive control along complex trajectories and this control improves with training. Second, the flocculus/paraflocculus region of the cerebellum appears to generate this control. Lesions impair pursuit and neural activity patterns are closely related to eye motion during complex pursuit. Importantly, neural responses lead eye motion during predictive pursuit and lag eye motion during non-predictable target motions that require visual control. The idea that flocculus/paraflocculus predictive control is non-visual is also supported by a lack of correlation between neural activity and retinal image motion during pursuit. Third, biologically accurate neural network models of the flocculus/paraflocculus allow the exploration and testing of pursuit mechanisms. Our current model can generate predictive control without visual input in a manner that is compatible with the extensive experimental data available for this cerebellar system. Similar types of non-visual cerebellar control are likely to facilitate the wide range of other skilled movements that are observed.

Cerebellar flocculus and ventral paraflocculus Purkinje cell activity during predictive and visually driven pursuit in monkey.

Purkinje cells in the flocculus and ventral paraflocculus were studied in tasks designed to distinguish predictive versus visually guided mechanisms of smooth pursuit. A sum-of-sines task allowed studies of complex predictive pursuit. A perturbation task examined visually driven pursuit during unpredictable right-angle changes in target direction. A gap task examined pursuit that was maintained when the target was turned off. Neural activity patterns were quantified using multi-linear models with sensitivities to the position, velocity, and acceleration of both motor output (eye motion) and visual input (retinal slip). During the sum-of-sines task, neural responses led eye motion by an average of 12 ms, a value larger than the 9-ms transmission delay between flocculus stimulation and eye motion. This suggests that flocculus/paraflocculus neurons drove pursuit along predictable sum-of-sines trajectories. In contrast, neural responses led eye motion by an average of only 2 ms during the perturbation task and by 6 ms during the gap task. These values suggest a follow-up role during tasks more heavily dependent on visual processing. Activity in all three tasks was explained primarily by sensitivities to eye position and velocity. Eye acceleration played a minor role during ongoing pursuit, although its influence on firing rate increased during the high accelerations following unexpected changes in target motion. Retinal slip had a relatively small influence on responses during pursuit. This was particularly true for the sum-of-sines and gap tasks where predictive control eliminated any consistent retinal-slip signals that might have been used to drive the eye. Surprisingly, the influence of retinal slip did not increase appreciably during unpredictable perturbations in target direction that generated large amounts of retinal slip. Thus although visual control signals are needed in varying amounts during the three pursuit tasks, they have been converted to motor control signals by the time they leave the flocculus/paraflocculus system. Individual neurons showed a remarkable constancy in eye-sensitivity direction across tasks that indicated direct links to oculomotor neurons. However, some neurons showed changes in sensitivity magnitude that suggested changes in control strategy for different tasks. Magnitude differences were largest for the perturbation task. We conclude that the flocculus/paraflocculus system plays a major role in driving predictive pursuit. It also processes visually driven control signals that originate in other brain regions after a slight delay.

Cerebellar flocculus and paraflocculus Purkinje cell activity during circular pursuit in monkey.

Responses from 69 Purkinje cells in the flocculus and paraflocculus of two rhesus monkeys were studied during smooth pursuit of targets moving along circular trajectories and compared with responses during sinusoidal pursuit and fixation. A variety of interesting responses was observed during circular pursuit. Although some neurons fired most strongly in a single preferred direction during clockwise (CW) and counterclockwise (CCW) pursuit, others had directional preferences that changed with rotation direction. Some of these neurons showed similar modulation amplitudes during CW and CCW pursuit, whereas other neurons showed a preference for a particular rotation direction. Response specificity also was observed during sinusoidal pursuit. Some neurons showed responses that were much stronger during centrifugal pursuit, others showed a preference for centripetal pursuit, and still others showed responses during both centripetal and centrifugal motion. Differences in preferred response direction were sometimes observed for centripetal versus centrifugal pursuit. CW/CCW and centrifugal/centripetal preferences were not explained by a breakdown in component additivity. That is, modulations in firing rate during pursuit along a circular trajectory equaled the sum of modulations during horizontal and vertical sinusoidal components as well as for diagonal components. Instead all responses were well fit by a model that expressed the instantaneous firing rate of each neuron as a multilinear function of the two-dimensional position and velocity of the eye. This model generalized well to performance at different sinusoidal frequencies. It did somewhat less well for responses during fixation, suggesting some separation in the neural mechanisms of dynamic and static positioning. The model indicates that position sensitivity accounted for approximately 36% of the modulation during circular pursuit, and velocity sensitivity accounted for approximately 64%. When position and velocity sensitivity vectors were aligned, responses were simpler and modulations were similar during CW versus CCW pursuit. In contrast, when these vectors pointed in different directions, response complexity increased. Nonaligned position and velocity influences tended to reinforce during circular pursuit in one direction and to cancel each other during pursuit in the opposite direction. They also tended to produce response differences during centripetal versus centrifugal sinusoidal pursuit. The distinct roles played by position and velocity in shaping Purkinje cell responses are compatible with the control signals required to generate smooth pursuit along circular and other two-dimensional trajectories.

Predictive smooth pursuit of complex two-dimensional trajectories demonstrated by perturbation responses in monkeys.

Two-dimensional sum-of-sines waveforms were pursued by the eye with very small phase delays compared with visual feedback delays estimated in the same monkeys. Processing delays in making smooth corrections averaged 90 msec after infrequent right-angle perturbations from a circular trajectory. These feedback delays were much larger than component phase delays during pursuit that averaged: 10 msec for sinusoids, 3 msec for circles, 20 msec for sum-of-two-sines trajectories, and 19 msec for sum-of-three-sines trajectories. This suggests that predictive control can play a strong role during tracking for a variety of simple and complex target trajectories.

Prediction of complex two-dimensional trajectories by a cerebellar model of smooth pursuit eye movement.

A neural network model based on the anatomy and physiology of the cerebellum is presented that can generate both simple and complex predictive pursuit, while also responding in a feedback mode to visual perturbations from an ongoing trajectory. The model allows the prediction of complex movements by adding two features that are not present in other pursuit models: an array of inputs distributed over a range of physiologically justified delays, and a novel, biologically plausible learning rule that generated changes in synaptic strengths in response to retinal slip errors that arrive after long delays. To directly test the model, its output was compared with the behavior of monkeys tracking the same trajectories. There was a close correspondence between model and monkey performance. Complex target trajectories were created by summing two or three sinusoidal components of different frequencies along horizontal and/or vertical axes. Both the model and the monkeys were able to track these complex sum-of-sines trajectories with small phase delays that averaged 8 and 20 ms in magnitude, respectively. Both the model and the monkeys showed a consistent relationship between the high- and low-frequency components of pursuit: high-frequency components were tracked with small phase lags, whereas low-frequency components were tracked with phase leads. The model was also trained to track targets moving along a circular trajectory with infrequent right-angle perturbations that moved the target along a circle meridian. Before the perturbation, the model tracked the target with very small phase differences that averaged 5 ms. After the perturbation, the model overshot the target while continuing along the expected nonperturbed circular trajectory for 80 ms, before it moved toward the new perturbed trajectory. Monkeys showed similar behaviors with an average phase difference of 3 ms during circular pursuit, followed by a perturbation response after 90 ms. In both cases, the delays required to process visual information were much longer than delays associated with nonperturbed circular and sum-of-sines pursuit. This suggests that both the model and the eye make short-term predictions about future events to compensate for visual feedback delays in receiving information about the direction of a target moving along a changing trajectory. In addition, both the eye and the model can adjust to abrupt changes in target direction on the basis of visual feedback, but do so after significant processing delays.

[Animal experiments for testing a hydroxyapatite ceramic coating under vacuum conditions]. / Tierexperimentelle Testung einer unter Niederdruckbedingungen gespritzten Hydroxylapatitkeramikbeschichtung.

Implants that were coated with hydroxyapatite ceramic (H-A.C.) under atmospheric condition in vivo often showed local areas of delamination. By applying the H-A.C. coating using the vacuum plasma spraying (VPS) technique, the mechanical characteristics of the coating was decidedly improved. We used a standardized rabbit model to compare a coating produced by this new technique with an implant conventionally plasma sprayed under atmospheric condition (APS). Cylindrical implants, 6 x 4 mm in size with a region flattened to a depth of 800 microns, were inserted into distal rabbit femurs underneath the patella. The flattened surfaces were coated with either a 150-micron APS-H-A.C. layer or a 150-micron VPS-H-A.C. layer plasma sprayed on an underlying 50-micron porous titanium layer. The animals were killed after 84 or 365 days. After 84 days histomorphologic evaluation revealed that more than 86% of each surface was covered with mature bone, while the VPS-H-A.C. coating demonstrated an almost two times greater tensile strength than the APS-H-A.C. coating. After 365 days both coatings showed a bony coverage of more than 94%. Again the tensile strength testing revealed much higher values for the VPS-H-A.C. coating. This study demonstrates that after 84 days as well as after 365 days in vivo, the VPS-H-A.C. coating had a significantly greater load capacity than an H-A.C. coating applied under atmospheric condition, and an equal affinity for bone.

Control of remembered reaching sequences in monkey. I. Activity during movement in motor and premotor cortex.

Motor and premotor cortex firing patterns from 307 single neurons were recorded while monkeys made rapid sequences of three reaching movements to remembered target buttons arrayed in two-dimensional space. A primary goal was to study and compare directionally tuned responses for each of three movement periods during 12 movement sequences that uniformly sampled the directional space in front of the monkey. The majority of neurons showed maximal responses during movements in a preferred direction with smaller increases during movements close to the preferred direction. These responses showed a statistically significant regression fit to a cosine function for 72% of the neurons examined. Comparisons among tuning directions computed separately for the first, second, and third movement periods suggested the near constancy of preferred direction across a rapidly executed series of movements even though these movements began at different starting points in space. Although directionally tuned neurons were only broadly tuned for a specific direction of movement, the neuronal ensemble carried accurate directional information. A population vector computed by summing vector contributions from the entire population of tuned neurons predicted movement direction with a mean accuracy of 20 degrees. This population code made consistent predictions for each of the 36 movements that were studied using a single set of population parameters. Most of the remaining neurons (24%) that were not tuned during movement did show significant changes in activity during other aspects of task performance. Some nontuned neurons had nondirectional increases that were sustained during movement, while others showed identical phasic bursts during the three movement periods. These nontuned neurons may control stabilizations of the shoulder, trunk, and forearm during movement, or forearm movements during button pushing.

Control of remembered reaching sequences in monkey. II. Storage and preparation before movement in motor and premotor cortex.

Single-neuron responses in motor and premotor cortex were recorded during a movement-sequence delay task. On each trial the monkey viewed a randomly selected sequence of target lights arrayed in two-dimensional space, remembered the sequence during a delay period, and then generated a coordinated sequence of movements to the remembered targets. Of 307 neurons studied, 25% were tuned specifically for either the first or the second target, but not both. In particular, for neurons tuned during both target presentations, tuned activity related to a particular first target direction were maintained during the presentation of a second target in a different direction. During the delay period, 32% of the neurons were tuned for upcoming movement in a single direction. These delay period responses often reflected activity patterns that first developed during target presentations and may therefore act to maintain target period information during the delay. Neurons with tuned activity during both the delay and movement periods exhibited two patterns: the first exhibited tuned responses during the delay that were correlated with the tuning of first-movement responses, while the second pattern showed delay-period tuning that was better correlated with tuned responses during second movements. This indicates that, before movement, distinct neural populations are correlated with specific movements in a sequence. About half the neurons studied were not directionally tuned during the initiation, target, or delay periods, but did show systematic changes in activity during task performance. Some (34%) were exclusively tuned during movement and appear to be involved in the direct control of movement. Others (17%) showed changes in firing rate from period to period within a trial but showed no directional preference for a particular direction of movement. Population analyses of tuned activity during the target and delay periods indicated that accurate directional information about both first and second movements was available in the neuronal ensemble well before reaching began. These results extend the idea that both motor and premotor cortex play a role in reaching behavior other than the direct control of muscles. While some early neural responses resembled muscle activation patterns involved in maintaining fixed postures before movement, others probably relate to the sensory-to-motor transformations, information storage in short-term memory, and movement preparation required to generate accurate reaching to remembered locations in space.

Predictive smooth pursuit of complex two-dimensional trajectories in monkey: component interactions.

Smooth pursuit eye movements were studied in monkeys tracking target spots that moved two-dimensionally. Complex target trajectories were created by applying either two or three sinusoids to horizontal and vertical axes in various combinations. The chance of observing predictable performance was increased by repeated training on each trajectory. Data analyses were based upon repeated presentations of each trajectory within sessions and on successive days. We wished to determine how accurately monkeys could pursue targets moving along these trajectories and to observe interactions among frequency components. At intermediate frequencies, tracking performance was smooth and consistent during repeated presentations with saccadic corrections that were well integrated with smooth pursuit. The mean gain for eight different sum-of-sines trajectories was 0.83 and the mean magnitude (absolute value) of the phase error was 6 degrees. In light of the long delays that have been associated with the processing of visual information, these values indicate that the monkeys were pursuing predictively. Five factors influenced predictive pursuit performance: (1) there was a decline in performance with increasing frequency; (2) horizontal pursuit was better than vertical pursuit; (3) high-frequency components were tracked with higher gains and phase lags, while lower-frequency components were tracked with lower gains and phase leads; (4) the gain of sinusoidal pursuit was always reduced when a second sinusoid was applied to the same axis or, to a lessor extent, when a second sinusoid of higher frequency was applied to the orthogonal axis; (5) the phase of sinusoidal pursuit shifted from a phase lag to a phase lead when combined with a second sinusoid of higher frequency, but was not affected by the addition of a lower-frequency sinusoid. Findings 1 and 2 confirm, in monkeys, results reported for humans, and 3 extends to monkeys and to two-dimensional pursuit results based upon human subjects. All of these findings demonstrate that complex predictive tracking is controlled by a nonlinear and nonhomogeneous system that uses predictive strategies in concert with feedback control to generate good pursuit.

A possible inflammatory reaction in a lateral neck cyst (branchial cyst) because of odontogenic infection.

We present the case of a woman who suffered from an acutely infected diffuse mass in the right neck. This mass had grown rapidly after difficult extraction of a tooth. Histologic analysis of the excised material revealed a lateral neck cyst with a lymph node that showed signs of an acute inflammation near the cyst. These findings support the theory that a preexisting lateral neck cyst may be "activated" by an intraoral inflammation. The different theories of the origin of lateral neck cysts are presented and discussed in the context of the case description.

A Neural Network Model of Cortical Activity during Reaching.

Abstract A neural network model that produces many of the directional and spatial response properties that have been observed for cortical neurons in monkeys moving toward targets in space is described. These include motor cortex units with broad tuning in a single preferred direction, approximately linear variation in activity for different hold positions, and approximate invariance in preferred direction for different starting points in space. Association cortex units in the model are sometimes irregular and reminiscent of neurons observed in visually responsive brain areas such as the posterior parietal cortex. The model is also compatible with population analyses performed on motor cortical neurons. Across network units, the distribution of preferred directions is uniformly distributed in directional space, and the degree of tuning and response magnitude vary from unit to unit. A population code used to predict accurately the direction of arm movements from a large population of coarsely tuned individual neurons allows predictions using a simulated population of unit responses obtained from the neural network model. This code works for different starting locations in space using the same parameters.

Primate motor cortex and free arm movements to visual targets in three-dimensional space. I. Relations between single cell discharge and direction of movement.

We describe the relations between the neuronal activity in primate motor cortex and the direction of arm movement in three-dimensional (3-D) space. The electrical signs of discharge of 568 cells were recorded while monkeys made movements of equal amplitude from the same starting position to 8 visual targets in a reaction time task. The layout of the targets in 3-D space was such that the direction of the movement ranged over the whole 3-D directional continuum in approximately equal angular intervals. We found that the discharge rate of 475/568 (83.6%) cells varied in an orderly fashion with the direction of movement: discharge rate was highest with movements in a certain direction (the cell's "preferred direction") and decreased progressively with movements in other directions, as a function of the cosine of the angle formed by the direction of the movement and the cell's preferred direction. The preferred directions of different cells were distributed throughout 3-D space. These findings generalize to 3-D space previous results obtained in 2-D space (Georgopoulos et al., 1982) and suggest that the motor cortex is a nodal point in the construction of patterns of output signals specifying the direction of arm movement in extrapersonal space.

Primate motor cortex and free arm movements to visual targets in three-dimensional space. II. Coding of the direction of movement by a neuronal population.

We describe a code by which a population of motor cortical neurons could determine uniquely the direction of reaching movements in three-dimensional space. The population consisted of 475 directionally tuned cells whose functional properties are described in the preceding paper (Schwartz et al., 1988). Each cell discharged at the highest rate with movements in its "preferred direction" and at progressively lower rates with movements in directions away from the preferred one. The neuronal population code assumes that for a particular movement direction each cell makes a vectorial contribution ("votes") with direction in the cell's preferred direction and magnitude proportional to the change in the cell's discharge rate associated with the particular direction of movement. The vector sum of these contributions is the outcome of the population code (the "neuronal population vector") and points in the direction of movement in space well before the movement begins.

Primate motor cortex and free arm movements to visual targets in three-dimensional space. III. Positional gradients and population coding of movement direction from various movement origins.

In one experiment, we studied the relations between the frequency of discharge of 274 single cells in the arm area of the motor cortex of the monkey and the actively maintained position of the hand in space. We found that the frequency of discharge of 63.9% of the cells studied was a multilinear function of the position of the hand in space according to the following equation (multiple linear regression): d = f + fxsx + fysy + fzsz, where d is the discharge rate of a single cell, f, fx, fy, fz are regression coefficients, and sx, sy, sz are the coordinates of the position of the hand. The equation above defines a positional gradient which implies that the frequency of cell discharge will increase at a maximum rate when the position of the hand changes along a certain direction; we call this direction of orientation of the positional gradient, and the rate of change in discharge rate along this orientation, the magnitude of the gradient. The orientations of the positional gradients were distributed throughout three-dimensional (3-D) space and their magnitudes differed among different cells. In a different experiment, we studied the changes in activity of 289 cells in the arm area of the motor cortex when the monkeys made equal-amplitude movements that started from different points in space, were in the same direction, and traveled along parallel trajectories in 3-D space. Four pairs of such movement directions (i.e., a total of 8 movement directions) were studied for every cell, and the changes in cell activity associated with movements within each pair were compared. We found that these changes in cell activity did not differ statistically for 68.4% of the movement pairs studied but did differ for the remaining 31.6%. The data from the whole population of cells studied in this experiment were analyzed using the population vector analysis described in the preceding paper (Georgopoulos et al., 1988). Thus, 8 population vectors were calculated, 1 for each of the 8 movement directions studied. We found that the direction of the population vector was close to the direction of the corresponding movement. These results indicate that the population vector provides unique information concerning the direction of the movement even when the point of origin of the movement varies in 3-D space.

Physiological study of neurons in the dorsal and posteroventral cochlear nucleus of the unanesthetized cat.

The responses of neurons in the posteroventral (PVCN) and dorsal (DCN) cochlear nucleus of the unanesthetized cat were determined for both long and short tones. These results were compared with recent studies in the barbiturate-anesthetized cat conducted in the same laboratory using similar stimuli and analysis programs. Every response pattern (poststimulus time histogram to short tones), which has been observed in previous studies using anesthetized animals, was also observed without anesthetic. The converse was also true: no novel response patterns were observed in the unanesthetized cat. This was also true for interval histogram, response area, isorate curve, and frequency sweep data. Some neurons were difficult to classify into existing descriptions of cochlear nucleus response patterns. For example: primary-like, onset, pauser, and buildup response patterns could also show chopper-like properties; onset-inhibitory, pauser, and buildup neurons appeared to form a response continuum rather than exist as separate response categories; and onset neurons with low characteristic frequencies (CFs) often showed sustained and strongly phase-locked responses below approximately 1,000 Hz. In addition, single neurons often showed more than one response pattern depending on the intensity and frequency of the acoustic stimulus. These ambiguities were also observed under anesthetic. Onset neurons within the PVCN appear to be well suited for the encoding of temporal and intensity information. At low stimulus frequencies they often respond to every cycle of a pure tone stimulus and exhibit the highest degree of phase-locking in the cochlear nucleus. The dynamic ranges associated with many onset neurons can exceed 80 dB compared with the 30- to 40-dB dynamic ranges associated with most other cochlear nucleus neurons. Onset neurons show a similar range of activities in the anesthetized cat. Neurons in the DCN have response properties that are more complex than those seen in the PVCN. Response patterns can change from sustained excitation to complete inhibition and are more often nonmonotonic near CF. DCN neurons can show well-defined tuning in the frequency domain and may be used to encode spectral information, but appear to be poorly suited for encoding temporal or intensity information as they are weakly phase-locked and have relatively small dynamic ranges. When DCN neurons "chop" they usually do so more slowly than do PVCN neurons. DCN neurons recorded in the anesthetized cat behave similarly. The relative frequency of a particular response pattern did vary with anesthetic state.(ABSTRACT TRUNCATED AT 400 WORDS)

Neuronal population coding of movement direction.

Although individual neurons in the arm area of the primate motor cortex are only broadly tuned to a particular direction in three-dimensional space, the animal can very precisely control the movement of its arm. The direction of movement was found to be uniquely predicted by the action of a population of motor cortical neurons. When individual cells were represented as vectors that make weighted contributions along the axis of their preferred direction (according to changes in their activity during the movement under consideration) the resulting vector sum of all cell vectors (population vector) was in a direction congruent with the direction of movement. This population vector can be monitored during various tasks, and similar measures in other neuronal populations could be of heuristic value where there is a neural representation of variables with vectorial attributes.

Topography of binaural organization in primary auditory cortex of the cat: effects of changing interaural intensity.

Responses from neuron clusters were used to derive binaural and aural dominance maps within the 5- to 30-kHz frequency representation of the primary auditory cortical (AI) field in the barbiturate-anesthetized cat. Tone burst stimuli were presented dichotically using a calibrated and sealed acoustic delivery system to parametrically vary interaural intensity difference (IID). Neuron cluster responses were divided into three binaural interaction classes using audiovisual criteria: summation (56%), suppression (25%), and mixed (17%). Neurons in the summation and suppression classes demonstrated a single type of binaural interaction, regardless of intensity manipulations. Neurons in the mixed binaural class demonstrated summation responses when dichotic tonal intensities were near their threshold levels and the IID was small, but suppression responses when the IID was increased. The relative proportions of the three binaural interaction classes changed with distance along the dorsal-to-ventral isofrequency dimension. Nearly equal proportions of each class were observed at the ventral end of field AI, whereas quite different proportions of each class were seen at the dorsal extreme of the field. The average frequency of occurrence of the mixed binaural class increased nearly monotonically with increasing distance from the dorsal end of field AI. The majority of mapped AI loci exhibited a contralateral aural dominance (65%) with equidominance (25%), ipsilateral aural dominance (6%), and predominantly binaural (4%) classes accounting for the remainder. Average topographic distributions of aural dominance suggested that the ventral end of field AI consisted almost exclusively of the contralateral dominance class, whereas more equal proportions of the four classes were observed near the dorsal extreme of the field. The highest average proportions of ipsilateral aural dominance and predominantly binaural classes were found in the dorsal half of field AI. Single neurons, isolated at cortical loci assigned to the mixed binaural class during the mapping of neuron clusters, were shown to demonstrate both summation and suppression responses. Quantitative measurements relating either discharge rate or response latency to changes in the IID appeared to distinguish these cells from other single neurons studied. Typically, the probability of discharge was initially increased and subsequently decreased by progressive changes in IID that increased the intensity of the ipsilateral tone relative to the contralateral tone. The initial changes in IID characteristically shortened the latent period to the binaural response while subsequent increments in IID produced a more comp

Cochlear nucleus, inferior colliculus, and medial geniculate responses during the behavioral detection of threshold-level auditory stimuli in the rabbit.

Rabbits were conditioned to respond behaviorally to auditory stimuli by pairing a white-noise conditioned stimulus (CS) with a corneal airpuff unconditioned stimulus (US). The conditioned response (CR) was movement of the nictitating membrane (NM). After the subjects were responding at better than the 90% correct level, the intensity of the auditory stimulus was reduced to behavioral threshold using a staircase procedure. Simultaneous measurements of neural unit activity and behavioral NM responses were then made in rabbits performing at behavioral threshold. After the experiment was completed neural unit responses during behavioral detection trials were compared to neural responses made during nondetection trials. Neural unit responses to a constant intensity, white-noise stimulus at behavioral threshold were well defined and essentially identical on behavioral detection and nondetection trials in the ventral cochlear nucleus, the ventrolateral division of the central nucleus of the inferior colliculus, and the ventral division of the medial geniculate body. This suggests that an auditory stimulus can be neuronally "detected" without being behaviorally detected, and that the neural "decision" to respond behaviorally is not made in these nuclei. Responses recorded from the dorsomedial division of the central nucleus of the inferior colliculus, the pericentral nucleus of the inferior colliculus, and less commonly in the medial division of the medial geniculate body were also clearly present and nearly identical during the onset of the auditory stimulus, but were sometimes consistently different for detection and nondetection conditions during the latter part of the auditory stimulus. These brain regions appear to receive both auditory and nonauditory inputs, and show responses which are more highly correlated with detection behavior.