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J C Houk

Publications and source records attributed to J C Houk.

At least 37 records · Page 2Linked to original sources

Motor co-ordinates in primate red nucleus: preferential relation to muscle activation versus kinematic variables.

1. Magnocellular red nucleus (RNm) neurones (n = 158) were recorded from two macaque monkeys during a tracking task using one of six single-degree-of-freedom manipulanda. This task allowed us to study discrete movements about most of the joints of the arm. Single-unit, kinematic and electromyographic (EMG) signals from ten to twenty muscles of the upper limb were collected for approximately 2 min while the monkey used a given manipulandum. Movements about different joints were studied by switching among manipulanda. 2. Cross-correlation functions were calculated between RNm discharge rate and the kinematic variables, position and velocity, and between RNm and each of the EMG signals. Statistically significant cross-correlation peaks were found in 24% of the position correlations, 22% of the velocity correlations and 32% of the EMG correlations. The highest correlations were for EMG, reaching above 0.60. The peak correlation provided an effective means of identifying neurones with strong functional relations to one or more movements and/or muscles. These could then be analysed in detail, on a trial-by-trial basis. 3. The similarity between the dynamics of EMG and velocity signals of many highly practised movements makes it difficult to determine which might be the more likely target of RNm control. Therefore, we sought exceptions to this pattern, in order to distinguish between these two possible modes of control. For example, at the end of a movement, muscles occasionally remained active as velocity approached zero. Small corrective movements were often accompanied by a disproportionately large EMG. During these periods, RNm activity usually followed the time course of one or more of the EMG signals as opposed to the velocity signal. In the majority of cases, RNm responses were bidirectional, less frequently unidirectional and rarely reciprocal. These patterns were similar to the patterns of muscle activity. They did not resemble the velocity signals unless the latter were passed through a rectifier. 4. The results support the hypothesis that the red nucleus generates motor commands in a muscle-based co-ordinate system. Covariation between RNm discharge and velocity may result indirectly from correlations between muscle activation and movement. We discuss how the cerebellar cortex might convert the distributed representation of target position, known to be present in the posterior parietal cortex, directly into dynamic, muscle-based commands in the rubro-cortico-cerebellar limb premotor network.

Animals↗

Three-dimensional reconstruction of the rubrocerebellar premotor network of the turtle.

Neuroanatomical studies have demonstrated that the organization of the reptilian rubrocerebellar limb premotor network is similar to that of mammals. This network is composed of prominent recurrent connections among the red nucleus, lateral cerebellar nucleus and lateral reticular nucleus. In this paper the rubrocerebellar system of the turtle was three-dimensionally reconstructed to permit detailed examination of its anatomical organization. Each nucleus and its major efferent pathway was imaged and reconstructed from separate anatomical cases. Section images were used to draw tissue boundaries, mark cell positions and locate axonal trajectories. For each nucleus, drawings of section images containing labeled cells were stacked in the rostrocaudal direction using anatomical landmarks, and a graphic model of the surface was constructed using the method of triangulation. An ellipsoid of equal concentration was computed for each nucleus to ascertain their three-dimensional boundaries and location within the brainstem. To examine the entire rubrocerebellar network, a template of the turtle brainstem and cerebellum was constructed. The component nuclei of the rubrocerebellar network and their axonal projections were then spatially warped onto the template reconstruction on a section by section basis. The final three-dimensional reconstruction of the turtle rubrocerebellar limb premotor network could be rotated in space, allowing proper visualization of the anatomical details of this system. Furthermore, we were able to mathematically section through the reconstruction to obtain brainstem slices with differing orientations and thickness.

Animals↗

Bistability in cerebellar Purkinje cell dendrites modelled with high-threshold calcium and delayed-rectifier potassium channels.

Phase-plane analysis of the ionic currents underlying dendritic plateau potentials was carried out to study the nonlinear dynamics and steady-state transfer properties of the dendritic tree in cerebellar Purkinje cells. The results of an analysis of the P-type calcium and delayed rectifier potassium channel system are presented in this study. These channels constitute a simple system that can support bistability and plateau potentials. By requiring both the steady-state current-voltage curve and nullclines to mimic basic plateau potential properties, we obtained well-defined ranges of specific conductance that can support bistability. Hysteresis was found to be surprisingly prevalent in this simple ion-channel system. Using the steady-state current voltage relationship, we derive concise, algebraic expressions for the voltage and current thresholds of state transitions as functions of specific conductance. The significance of bistability in this ion-channel system is discussed with respect to the generation of plateau potentials in Purkinje cells dendrites and the role of the cerebellum in motor control.

Action Potentials↗

Synaptic linkages between red nucleus cells and limb muscles during a multi-joint motor task.

The magnocellular red nucleus (RNm) becomes highly active when a monkey reaches to grasp an object. However, the only spike-triggered averaging studies of the RNm to date have been restricted to a simple wrist tracking paradigm and electromyographic (EMG) measurements of muscles of the forearm. We have now measured EMG signals from a large number of muscles throughout the shoulder, arm, forearm, and hand during a variety of tasks, including unconstrained reaching and grasping movements. Relations between these EMG signals and single-unit activity were assessed by on-line spike-triggered averaging and revealed significant post-spike effects among muscles of the shoulder and proximal arm, as well as intrinsic hand muscles. Although there remained a strong bias toward the extensor muscles of the forearm, as has been shown earlier, these results reinforce the importance of the RNm in the control of coordinated, whole-limb reaching movements.

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In vitro classical conditioning of abducens nerve discharge in turtles.

In vitro classical conditioning of abducens nerve activity was performed using an isolated turtle brainstem-cerebellum preparation by direct stimulation of the cranial nerves. Using a delayed training procedure, the in vitro preparation was presented with paired stimuli consisting of a 1 sec train stimulus applied to the auditory nerve (CS), which immediately preceded a single shock US applied to the trigeminal nerve. Conditioned and unconditioned responses were recorded in the ipsilateral abducens nerve. Acquisition exhibited a positive slope of conditioned responding in 60% of the preparations. Application of unpaired stimuli consisting of CS-alone, alternate CS and US, or backward conditioning failed to result in conditioning, or resulted in extinction of CRs. Latencies of CR onset were timed such that they occurred midway through the CS. Activity-dependent uptake of the dye sulforhodamine was used to examine the spatial distribution of neurons labeled during conditioning. These data showed label in the cerebellum and red nucleus during conditioning whereas these regions failed to label during unconditioned responses. Furthermore, the principal abducens nucleus labeled heavily during conditioning. These findings suggest the feasibility of examining classical conditioning in a vertebrate in vitro brainstem-cerebellum preparation. It is postulated that the abducens nerve CR represents a behavioral correlate of a blink-related eye movement. Multiple sites of conditioning are hypothesized, including the cerebellorubral circuitry and brainstem pathways that activate the principal abducens nucleus.

Abducens Nerve↗

Anatomical organization of the limb premotor network in the turtle (Chrysemys picta) revealed by in vitro transport of biocytin and neurobiotin.

The in vitro turtle brainstem-cerebellum preparation has been a valuable tool in the study of central motor programs. In the present study, we investigate the anatomical organization of the turtle rubrocerebellar limb premotor network and its sensory connections in vitro by combining the rapid anterograde and retrograde transport of neurobiotin and biocytin with the extended viability of the isolated turtle brainstem-cerebellum. These compounds retrogradely labeled soma, dendrites, and axons, and orthogradely labeled axons and, to a lesser extent, terminals. The chelonian red nucleus receives a dense input from the contralateral lateral cerebellar nucleus and projects heavily to the contralateral spinal cord. Rubral axons sparsely innervate the lateral cerebellar nucleus and project heavily to the lateral reticular nucleus. Lateral reticular axons heavily innervate the lateral cerebellar nucleus before terminating in the pars lateralis of the cerebellar cortex as mossy fibers. These prominent, recurrent loops among the lateral cerebellar nucleus, red nucleus, and lateral reticular nucleus constitute the turtle rubrocerebellar limb premotor network. Sensory inputs to the red nucleus originate in the contralateral dorsal column nuclei, the principal trigeminal nucleus, and the spinothalamic system. These sites project bilaterally to the lateral reticular nucleus. The lateral cerebellar nucleus receives a contralateral input from the dorsal column nuclei. The red nucleus projects sparsely to the dorsal column nuclei. The red nucleus also receives an ipsilateral descending projection from the suprapeduncular nucleus, located in the diencephalon, and an ascending input from the rostral rhombencephalic reticular formation. An ipsilateral descending pathway originating in the red nucleus is likely to be the rubro-olivary tract.

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Model of topographic map development guided by a transiently expressed repulsion molecule.

The projection from the retina develops into a precise map of the visual world on the surface of the tectum. The search for molecular position cues that mediate map formation has recently yielded a tectal molecule that exerts a repulsion to fibers from the entire temporal half retina. This molecule appears not to function in the generally accepted gradient manner but instead provides only binary position information, and it is only expressed transiently during early development. Here we describe modeling results that compare the efficacy of binary versus graded position cues in topographic map formation; the model also includes an activity dependent process. We find that binary repulsion is more efficient than graded chemoaffinity in the rapid establishment of map polarity, and transient expression of either cue provides sufficient guidance for precise map formation.

Axons↗

Intrinsic and synaptic properties of turtle red nucleus neurons in vitro.

Burst discharges in the red nucleus are correlated with discrete limb movements. Intracellular recordings from red nucleus neurons in the in vitro turtle brainstem-cerebellum was performed to elucidate mechanisms underlying these bursts. Depolarizing intracellular current injection failed to demonstrate endogenous membrane currents that might produce burst discharges, and neurons did not exhibit significant spike frequency adaptation, which is a characteristic of synaptically driven bursts. Responses of red nucleus neurons to synaptic input demonstrated a late, slow depolarizing synaptic potential (slow EPSP) having a latency of 9-12 ms, and a maximal duration of 600 ms. it is concluded that neither intrinsic membrane responses, nor the duration of the slow EPSP, can fully account for the behavior of red nucleus neurons during burst discharge. We hypothesize that activity in the red nucleus is driven by a gradual recruitment of NMDA receptors, and lpr by polysynaptic excitatory pathways.

Animals↗

Receptive fields of single cells from the face zone of the cat rostral dorsal accessory olive.

Natural stimulation was used to map the receptive fields of single cells recorded from the rostral medial portion of the dorsal accessory olive (rDAO) and the subjacent principal olive (PO) of the barbiturate anesthetized cat. Previous reports indicated a somatotopic mapping of the entire contralateral body within the rDAO which included a small face zone and a larger zone with a very precise map for the limbs. While concentrating on the face zone of the rDAO we confirmed the previously reported somatotopy (face: rostral and medial; forelimb: caudal and medial; hindlimb; caudal and lateral; and trunk: rostal and lateral) and found a somatotopy within, and adjacent to, the face zone. At the border between rDAO regions representing forelimb and face, cells with forelimb fields were found to lie dorsally to cells with facial fields. Within the rDAO face region, cells with large facial fields lie dorsally to cells with small facial fields. In both cases, the more ventral cells lie in the ventral lamella of the PO, which suggests a functional as well as physical continuity between rDAO and the ventral lamella of the PO. We therefore conclude that the face zone in the rDAO and the face zone in the PO form one continuous and complete map of the face with an orderly progression of receptive fields. Furthermore, we have found that stimulation of the red nucleus can inhibit rDAO cells with facial receptive fields just as it does cells with receptive fields from the rest of the body.

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Anatomy of the turtle cerebellorubral circuit studied in vitro using neurobiotin and biocytin.

We have combined the rapid anterograde and retrograde transport of neurobiotin and biocytin with the extended viability of the isolated turtle brainstem-cerebellum to conduct in vitro studies of the chelonian cerebellorubral circuit. Tracers were pressure injected in 15-25 nl quantities and the optimal transport time was 16 h. Tissue sections were incubated with avidin-biotin-HRP complex and reacted with DAB. Retrogradely labeled soma, dendrites and axons, and anterogradely labeled axons and to a lesser extent terminals were visible with both tracers. Red nucleus injections resulted in dense retrograde label in the contralateral lateral cerebellar nucleus and a heavily labeled contralateral rubrospinal tract. Cerebellar nucleus injections revealed light retrograde and dense terminal label in the contralateral red nucleus, together with retrograde label in a cell cluster in the ipsilateral ventrolateral medullary reticular formation, an area we identify as the lateral reticular nucleus. Injections into this medullary region resulted in heavy mossy fiber input to the ipsilateral cerebellum and moderate retrograde label in the contralateral red nucleus. These results identify prominent recurrent projections between the lateral cerebellar nucleus, red nucleus and lateral reticular nucleus, in addition to revealing other features of the cerebellorubral circuit.

Animals↗

Distributed motor commands in the limb premotor network.

Neuroanatomical studies have demonstrated extensive interconnections between the motor cortex, red nucleus and cerebellum, forming a premotor network for controlling limb movement. Single-unit studies indicate that command signals for limb movements are distributed broadly throughout this network. Cellular studies have demonstrated multiple recurrent loops in this network, and the presence of excitatory and inhibitory amino acid neurotransmitters. A recent model suggests that movement commands are initiated by sensory inputs to these loops, and that positive feedback, regulated by inhibition from cerebellar Purkinje cells, distributes commands throughout the limb premotor network. This model offers a new framework for exploring relationships between basic neural mechanisms and concepts of motor performance that derive from experimental psychology.

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Correlation of primate red nucleus discharge with muscle activity during free-form arm movements.

1. We recorded from 239 neurons located in the magnocellular division of the red nucleus of four alert macaque monkeys. At the same time, we recorded electromyographic (EMG) signals from as many as twenty electrodes chronically implanted on muscles of the shoulder, arm, forearm and hand. We recorded EMG signals for periods ranging from several months to a year. 2. The monkeys were trained to perform three free-form food retrieval tasks, each of which activated all of the recorded muscles and most of the neurons. The 'prehension' task required simply that the monkey grasp a piece of food from a fixed point in space. The 'barrier' task required the monkey to reach around a small barrier to obtain the food, and the 'Kluver' task required that food be removed from small holes. During the prehension task, we found approximately equal numbers of neurons that were strongly active while the hand was being moved toward the target (70% of units), and while the food was being grasped (60%). Relatively few units were active as the hand was returned to the mouth (15%). 3. Data files of 1-2 min duration were collected while the monkey performed a single behavioural task. Whenever possible, we recorded files for all three tasks from each neuron. For each file we calculated long time-span analog cross-correlations (+/- 1.28 s) between instantaneous neuronal firing rate and each of the full-wave rectified, low-pass filtered EMG signals. We used the peak correlation and the time of the peak as two summary measures of the functional relation between modulation of neuronal activity and EMG. 4. The magnitude of the strongest correlations was between 0.4 and 0.5 (normalized to a perfect correlation of +/- 1.0). Distal muscles were the most frequently correlated, and extensors were more frequently correlated than flexors. For all monkeys, the lags for well correlated muscles were distributed broadly about a uni-modal value near 0 ms. Eighty five per cent of the correlations larger than or equal to 0.25 had peaks between -150 and 200 ms. 5. The activity of each neuron was represented in a muscle co-ordinate system by an n-dimensional 'functional linkage vector', each element of which was the peak correlation with one of n muscles. The vector for any given neuron points in a particular direction in muscle space, depending on the similarity between the activity of the neuron and the activity of each muscle.(ABSTRACT TRUNCATED AT 400 WORDS)

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Output organization of intermediate cerebellum of the monkey.

1. The goal of this study was to investigate the motor organization of monkey nucleus interpositus (NI) and neighboring regions of the lateral nucleus (NL) by correlating discharge of single neurons with active movements. Neurons were surveyed during free-form movements as well as during operation of six devices that required movement about specific forelimb joints. The paradigm allowed us to test the hypothesis that discharge of individual cells relates to movements about individual joints. 2. One hundred sixty-two isolated nuclear neurons from two monkeys were studied. Eighty-three percent showed large increases in discharge (an average of 3 times resting rate for forelimb neurons) during movement of one body part, either forelimb, hindlimb, mouth/face, or eyes. 3. Anterior interpositus contains neurons related to hindlimb movement in anterior regions and neurons related to forelimb movement in posterior regions. A mouth/face-related area exists in the dorsal-posterior regions and is continuous with a mouth/face area in the dorsal regions of NL. Posterior interpositus (NIP) showed no clear separation between forelimb and hindlimb neurons: forelimb neurons were encountered throughout the nucleus, and hindlimb neurons were encountered in the medial-anterior two thirds. A distinct eye movement area exists in lateral, posterior, and ventral regions of NIP. This area borders regions of NL that also contain eye movement-related neurons. 4. Forelimb interpositus neurons discharged strongly during reach and grasp; discharge rates were recorded for 41 neurons during a stereotyped reach and the average depth of modulation was 149 imp/s. Nineteen neurons that modulated during device tracking were also tested during reaching, and the depth of modulation was much greater during reaching. 5. Fifty-nine forelimb neurons were tested with device tracking. Twenty-seven (46%) produced no audible modulation, regardless of the joint being exercised. The remaining 32 neurons modulated during movement on at least one device (mean depth of modulation = 84 imp/s). Comparison of discharge during use of different devices revealed no strong evidence for device-specific discharge. 6. Discharge modulations during device tracking were phasic, preceded movement, and, for a small number of cells, showed consistent parametric relations to duration, amplitude, and velocity of movement. 7. Despite a clear somatotopy within NI and NL, there is no finer mapping based on active movements about individual joints within forelimb regions. Discharge modulation depends on movements involving the whole limb. Progress in understanding the function of intermediate cerebellum depends on determining the variables required to elicit consistent and high modulation of neural discharge.(ABSTRACT TRUNCATED AT 400 WORDS)

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Movement-related inputs to intermediate cerebellum of the monkey.

1. The primary goal of this study was to characterize the information about single-joint forelimb movements supplied to intermediate cerebellar cortex by mossy fibers. Discharge of mossy fibers and Golgi cells was studied while monkeys operated six devices that required movements about specific joints. Additional control experiments in anesthetized cats and monkeys established criteria for identification of mossy fibers and Golgi cells. 2. The control experiments demonstrate that mossy fibers can be distinguished from Purkinje and Golgi cells by the waveshapes of their action potentials. Asynaptic activation from the inferior cerebellar peduncle, in combination with histological localization of recording sites in granular layer or subcortical white matter, verified that mossy fibers produce a variety of waveshapes that are characterized by brief initial phases and relatively small amplitudes. The same waveshapes were observed for the mossy fiber recordings from awake monkeys, and many identified mossy fibers had sensory properties similar to those found in the awake animals. From these combined criteria, we conclude that the recordings in the awake animals were from mossy fibers. Golgi cells, recorded exclusively in the granular layer of cerebellar cortex, were characterized by action potentials of longer duration and larger amplitude as compared with mossy fibers, and none were asynaptically activated from the inferior cerebellar peduncle. 3. Units were isolated while the monkeys made free-form and tracking movements. We studied movement-related discharge of 80 mossy fibers and 12 Golgi cells. Mossy fibers showed high modulations during use of at least one of the six manipulanda and had clear preferences for movement about a specific joint, although they often showed consistent but weaker firing during movement about a neighboring joint. Separation of movements by more than one joint produced a large reduction in discharge: shoulder units never fired well to movements of the finger, and finger units never fired well to movement of the shoulder. 4. The tracking task required maintenance of fixed limb positions (a static phase) as well as movements between these positions (a dynamic phase). Of 80 mossy fibers, 18% had purely tonic discharge patterns, 63% were phasic-tonic, and 20% were purely phasic. Discharge patterns were reciprocal (45%), bidirectional (42%), or unidirectional (13%). 5. Eighty percent of the mossy fibers exhibited tonic discharge that was significantly (P < 0.01) correlated with joint angle (r = 0.65 +/- 0.19, mean +/- SD), and about one third had phasic components that were significantly correlated with movement velocity.(ABSTRACT TRUNCATED AT 400 WORDS)

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Evidence for GABAergic interneurons in the red nucleus of the painted turtle.

Immunocytochemical and electrophysiological evidence supporting the presence of GABAergic interneurons in the turtle red nucleus is presented. Injections of HRP into the spinal cord produced labeling of large neurons in the contralateral red nucleus. The peroxidase-antiperoxidase (PAP) method revealed smaller cells immunoreactive to an antibody against glutamate decarboxylase (GAD), the synthetic enzyme for the inhibitory neurotransmitter GABA, that were interspersed among larger immunonegative neurons. Similar small neurons were densely immunostained by antibodies to GABA-glutaraldehyde conjugates obtained from different sources and applied according to pre-embedding and postembedding protocols. Rubrospinal neurons retrogradely labeled with HRP measured 16 and 27 microns in mean minor and major cell body diameters, while GABA-like immunopositive neurons situated within the red nucleus measured 7 and 13 microns. There was very little overlap in soma size between the two cell populations. Therefore, we suggest that the GAD- and GABA-positive neurons may be local inhibitory interneurons. This notion is further supported by observations of pre-embedding immunostaining for GAD and postembedding immunostaining for GABA showing that the turtle red nucleus is amply innervated by immunoreactive axon terminals. These puncta are closely apposed to cell bodies and dendrites of both immunonegative large neurons and immunopositive small neurons. Moreover, immunogold staining at the electron microscopic level demonstrated that GABA-like immunoreactive axon terminals with pleomorphic synaptic vesicles formed symmetric synapses with cell bodies and dendrites of the two types of red nucleus cells. These ultrastructural features are commonly assumed to indicate inhibitory synapses. A moderately labeled bouton with round vesicles and asymmetric synapses was also observed. In addition, the two types of red nucleus neurons received asymmetric axosomatic and axodendritic synapses with GABA-negative boutons provided with round vesicles, features usually associated with excitatory functions. To obtain electrophysiological evidence for inhibition, intracellular recordings from red nucleus neurons were conducted using an in vitro brainstem-cerebellum preparation from the turtle. Small, spontaneous IPSPs were recorded from 7 out of 14 red nucleus cells studied. These morphological and physiological results provide strong support for concluding that the turtle red nucleus, like its mammalian counterpart, contains GABAergic inhibitory interneurons. While we have not identified the main source of input to these interneurons, in view of the scarce development of the reptilian cerebral cortex, this input is unlikely to come from the motor cortex as it does in mammals.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Spatial overlap of rubrospinal and corticospinal terminals with input to the inferior olive.

Somatosensory responses of cells in the dorsal accessory olive are suppressed following stimulation of the magnocellular red nucleus. Since the magnocellular red nucleus of the cat does not project directly to the dorsal accessory olive, the present experiments were designed to identify indirect pathways that might mediate suppression of olivary responsiveness. Wheat germ agglutinin-horseradish peroxidase was used to compare the location of magnocellular red nucleus terminals with the locations of cells providing input to the rostral dorsal accessory olive. Cells projecting to forelimb rostral dorsal accessory olive can be divided into two main groups: one group comprises a column of large cells located in the ventral caudal cuneate nucleus extending into lamina VI of C1 and C2, and a second group comprises smaller cells located in the ventral rostral cuneate nucleus. Terminations of fibers originating in the magnocellular red nucleus were found to target both groups of cells projecting to the dorsal accessory olive. Therefore, it is possible that the responsiveness of olivary cells is influenced via these terminations. Stimulation of sensorimotor cortex has also been shown to inhibit olivary responsiveness. Terminations from sensorimotor cortex target the same regions of cells that project to the dorsal accessory olive as those of the magnocellular red nucleus, and a similar, perhaps identical, anatomical substrate may serve to modulate olivary sensitivity by the two descending systems.

Animals↗

Sulforhodamine labeling of neural circuits engaged in motor pattern generation in the in vitro turtle brainstem-cerebellum.

A fluorescent molecular probe was used in combination with a novel in vitro preparation to study spatial patterns of neural activity associated with motor pattern generation. The in vitro brainstem-cerebellum preparation takes advantage of the turtle's unusual resistance to anoxia to preserve the entire neural network that connects the cerebellum, red nucleus, and reticular formation. This preparation was bathed in a 0.01% solution of sulforhodamine while it was activated unilaterally by electrical stimulation of the dorsal quadrant of the spinal cord for 1 hr. Sulforhodamine is a small, sulfonated, highly charged fluorescent molecule that is taken up by endocytosis. To examine its distribution in the cerebellum and brainstem, coronal sections were prepared and viewed under epifluorescence illumination. Distinctive spatial patterns of labeling were associated with unilateral electrical stimulation of the in vitro network, suggesting that dye uptake was activity dependent. Blockade of uptake with altered magnesium and calcium concentrations indicated that single spike discharge evoked ortho- or antidromically was insufficient to induce dye uptake. Instead, sulforhodamine staining correlated with the presence of burst discharge that was recorded extracellularly from the red nucleus. Blockade of burst discharge with excitatory amino acid receptor antagonists prevented dye uptake in the red nucleus, the lateral cerebellar nucleus, and other structures that are known to be interconnected by recurrent anatomical pathways. These results suggest that sulforhodamine is internalized by intensely active neurons. The spatial distributions of label support the hypothesis that burst discharges in the turtle red nucleus are mediated by excitatory amino acid neurotransmitters and sustained by recurrent excitation in cerebellorubral synaptic pathways. Positive feedback in these recurrent pathways may provide an important driving force for the generation of motor programs that control limb movements.

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