Biomedical subjects

A K Moschovakis

Publications and source records attributed to A K Moschovakis.

13 recordsLinked to original sources

PubMed1992

The termination pattern and postsynaptic targets of rubrospinal fibers in the rat spinal cord: a light and electron microscopic study.

The spinal course, termination pattern, and postsynaptic targets of the rubrospinal tract, which is known to contribute to the initiation and execution of movements, were studied in the rat at the light and electron microscopic levels by using the anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L) in combination with calbindin-D28k (CaBP), gamma-aminobutyric acid (GABA), and glycine immunocytochemistry. After injections of PHA-L unilaterally into the red nucleus, labelled fibers and terminals were detected at cervical, thoracic, and lumbar segments of the spinal cord. Most of the descending fibers were located in the dorsolateral funiculus contralateral to the injection site, but axons descending ipsilaterally were also revealed. Rubrospinal axon terminals were predominantly found in laminae V-VI and in the dorsal part of lamina VII at all levels and on both sides of the spinal cord, but stained collaterals were also seen in the ventrolateral aspect of Clark's column and in the ventral regions of lamina VII on both sides. The proportion of axonal varicosities revealed on the ipsilateral side varied at different segments and represented 10-28% of the total number of labelled boutons. Most of the labelled boutons were engaged in synaptic contacts with dendrites. Of the 137 rubrospinal boutons investigated, only 2 were found to establish axosomatic synaptic junctions in the lumbar spinal cord contralateral to the PHA-L injection. With the postembedding immunogold method, 80.8% of dendrites establishing synaptic contacts with rubrospinal terminals did not show immunoreactivity for either GABA or glycine, whereas 19.2% of them were immunoreactive for both amino acids. Rubrospinal axons made multiple contacts with CaBP-immunoreactive neurons in laminae V-VI. Synaptic contacts between rubrospinal terminals and CaBP-immunoreactive dendrites were identified at the electron microscopic level, and all CaBP-containing postsynaptic dendrites investigated were negative for both GABA and glycine. The results suggest that rubrospinal terminals establish synaptic contacts with both excitatory and inhibitory interneurons in the rat spinal cord, and a population of excitatory interneurons receiving monosynaptic rubrospinal input is located in laminae V-VI.

Animals↗

PubMed1992

Anatomical and physiological study of interneurons in an oligosynaptic cutaneous reflex pathway in the cat hindlimb.

Morphological and physiological studies were undertaken in the cat lumbosacral spinal cord in order to examine individual last-order excitatory interneurons in the disynaptic pathway between low threshold superficial peroneal (SP) afferents and flexor digitorum longus (FDL) motoneurons. Last-order interneurons projecting to the FDL motor nucleus were visualized with activity-dependent trans-synaptic transport of lectin-conjugated horseradish peroxidase (WGA-HRP). The location of lumbar interneurons that responded at short latencies (less than 3 ms) to electrical stimulation of the SP nerve was further defined using extracellular recording. The projection of individual interneurons to motoneurons in the L6 segment was defined by spike-triggered averaging of population ventral root potentials (VRPs) using the sucrose gap technique. Measurements of the central latencies from the arrival of the SP afferent volley to interneuron firing, and from interneuron spike to the onset of the VRP EPSPs, confirm the existence of a low threshold, disynaptic SP pathway that excited motoneurons. These data provide useful latency constraints for the interpretation of the synaptic length of cutaneous reflex pathways in the cat hindlimb.

Afferent Pathways↗

PubMed1991

The size and dendritic structure of HRP-labeled gamma motoneurons in the cat spinal cord.

We report quantitative data obtained from 60 fully reconstructed dendritic trees belonging to eight gamma-motoneurons (gamma-MNs) and six additional gamma-MNs that were not completely reconstructed. The cells were labeled intracellularly with horseradish peroxidase (HRP). These data are compared to measurements from 79 reconstructed dendrites belonging to seven documented alpha-motoneurons (alpha-MNs), supplemented by a larger sample of alpha-MNs labeled intracellularly or by retrograde transport with HRP. As expected from earlier studies, the soma dimensions and total membrane area of gamma-MNs were smaller than those of alpha-MNs. Although gamma-MN dendrites were, on average, slightly but significantly longer than those of alpha-MNs, the former had, on average, smaller diameter stem dendrites, less membrane area, and less profuse branching, and they tended to branch closer to the soma and to terminate farther from the soma. These differences were evident even when subsets of dendrites with similar stem diameters were compared. Some of the anatomical distinctions suggest that gamma-MNs are qualitatively as well as quantitatively different from alpha-MNs, even though the distributions of many of the morphological variables examined showed no abrupt discontinuities between the two motoneuron groups.

Animals↗

PubMed1991

Differential control of short latency cutaneous excitation in cat FDL motoneurons during fictive locomotion.

Low threshold (less than or equal to 2 x T) cutaneous afferents in the superficial peroneal (SP) and medial plantar (PLNT) nerves both produce short-latency excitatory postsynaptic potentials (EPSPs) in flexor digitorum longus (FDL) alpha-motoneurons, with minimum central latencies (less than or equal to 1.8 ms) that indicate a disynaptic connection. However, SP and PLNT EPSPs in FDL motoneurons are differentially modulated during fictive stepping in decerebrate cats. The early components in SP EPSPs are systematically enhanced during the early flexion phase of fictive stepping (Schmidt et al. 1988) while those in PLNT EPSPs are markedly depressed during flexion. In addition, transmission in the PLNT----FDL pathway is enhanced during occasional step cycles in which the FDL displays firing during the extension phase. This enhancement affects only the trisynaptic components of PLNT EPSPs, is simultaneous with the extension FDL burst, and is not found in SP EPSPs. These results indicate that the SP----FDL and PLNT----FDL pathways are composed of entirely different sets of segmental last-order interneurons, each of which receives sensory information from contiguous, relatively limited regions of skin on the most distal parts of the hindpaw. Possible functional consequences of these interneuronal organizations are discussed.

Animals↗

PubMed1991

Structure of the primate oculomotor burst generator. I. Medium-lead burst neurons with upward on-directions.

1. To investigate the structure of the primate burst generator for vertical saccades, we obtained intra-axonal records from vertical medium-lead burst neurons with upward on-directions (UMLBs) in alert, behaving squirrel monkeys, while monitoring their spontaneous eye movements. After physiological characterization, these UMLBs were injected with horseradish peroxidase. 2. UMLBs (n = 14) had no spontaneous activity and emitted bursts of action potentials that preceded rapid eye movements by approximately 6 ms. Parameters of the burst (duration and number of spikes) were highly correlated with parameters of the rapid eye movement (duration and amplitude of the upward displacement of the eyes). 3. The axons of six UMLBs projected to the oculomotor complex. Their somata (4 were recovered) were all in the rostral interstitial nucleus of the medial longitudinal fasciculus (riMLF). Their axons traveled caudally in the medial longitudinal fasciculus (MLF) and ramified in the interstitial nucleus of Cajal (NIC) before entering the oculomotor nucleus. Five axons terminated bilaterally in the subdivisions innervating the superior rectus and inferior oblique muscles and therefore were presumed to be excitatory. One axon terminated in the ipsilateral inferior rectus and superior oblique subdivisions of the oculomotor complex and was presumed to be inhibitory. 4. Additionally, our data demonstrate that the nucleus of the posterior commissure (nPC) may also contain UMLBs. The axon of one such neuron crossed the midline within the posterior commissure and provided terminal fields to the contralateral nPC, riMLF, NIC, and the mesencephalic reticular formation but not to the oculomotor complex. 5. In conclusion, our data demonstrate that the rostral mesencephalon of the monkey contains neurons that have both the activity and the connections that are necessary either to provide motoneurons innervating extraocular muscles of both eyes with the pulse of activity they display during upward saccades or to inhibit their antagonists. Furthermore, our data demonstrate that some UMLBs are better suited for closing the feedback path of the local feedback loop rather than for providing direct input to extraocular motoneurons.

Action Potentials↗

PubMed1991

Structure of the primate oculomotor burst generator. II. Medium-lead burst neurons with downward on-directions.

1. To investigate the morphology and physiology of vertical medium-lead burst neurons with downward on-directions (DMLBs), we impaled midbrain axons and recorded their discharge patterns in relation to spontaneous saccades of alert, behaving squirrel monkeys. Selected axons were injected with horseradish peroxidase and morphologically characterized. 2. DMLBs emitted bursts of impulses that preceded rapid eye movements by approximately 5 ms. Parameters of the burst (duration and number of spikes) were highly correlated with parameters of the saccadic eye movement (duration and amplitude of the downward displacement of the eyes). 3. Somata of DMLBs were recovered in the rostral interstitial nucleus of the medial longitudinal fasciculus (riMLF, n = 14), and in the interstitial nucleus of Cajal (NIC, n = 2). Fibers originating from riMLF DMLBs projected, usually ipsilaterally, to the NIC as well as in the inferior rectus and the superior oblique subdivisions of the oculomotor complex. The axons of NIC DMLBs projected to the ipsilateral riMLF, NIC, and the mesencephalic reticular formation but not to the oculomotor complex. 4. Our data demonstrate that some DMLBs can provide extraocular motoneurons of both eyes with the pulse of activity they display during downward saccades. In addition, such neurons can supply the NIC with one of the signals that this nucleus is thought to integrate to extract an estimate of the vertical eye position. Finally, our data demonstrate the existence of DMLBs that do not establish direct connections with oculomotoneurons.

Action Potentials↗

PubMed1990

A structural basis for Hering's law: projections to extraocular motoneurons.

Conjugate eye movements are executed through the concurrent activation of several muscles in both eyes. The neural mechanisms that underlie such synergistic muscle activations have been a matter of considerable experimentation and debate. In order to investigate this issue, the projections of a class of primate premotoneuronal cells were studied, namely, the vertical medium-lead burst neurons (VMLBs), which drive vertical rapid eye movements. Axons of upward VMLBs ramify bilaterally within motoneuron pools that supply the superior rectus and inferior oblique muscles of both eyes. Axons of downward VMLBs ramify ipsilaterally in the inferior rectus portion of the oculomotor nucleus and in the trochlear nucleus. Thus, VMLBs can drive vertical motoneuron pools of both eyes during conjugate vertical rapid eye movements; these data support Hering's law.

Action Potentials↗

PubMed1988

Structure-function relationships in the primate superior colliculus. I. Morphological classification of efferent neurons.

1. Neurons in the superior colliculus (SC) of anesthetized paralyzed squirrel monkeys were injected intracellularly with horseradish peroxidase (HRP) to establish a morphological classification of tectal efferent neurons in this species. These neurons were physiologically identified by their antidromic responses following stimulation of the contralateral predorsal bundle or SC. These cells also responded with postsynaptic potentials to stimulation of the ipsilateral substantia nigra and cerebral peduncle and the contralateral tectum. 2. Quantitative light microscopic analysis of the somatodendritic profiles and axonal trajectories of 27 recovered cells revealed the existence of three major groups of tectal efferent neurons: L (n = 7), X (n = 8), and T (n = 12). 3. L neurons are small or medium size cells with relatively elaborate dendritic trees and are located mainly in the superficial layers of the SC. They participate in the ipsilateral descending and dorsal ascending tectofugal bundles. Intrinsic collaterals of L axons deploy a large number of boutons both near the parent cell body and more ventrally within the deeper tectal layers. 4. X neurons are mostly large in size and multipolar in shape with relatively complex dendritic trees. Their cell bodies are situated mainly in the stratum griseum intermedium and occasionally in the stratum opticum. Axons of X neurons participate in the crossed descending and ipsilateral ventral ascending projections of the SC. In addition, the axonal system of about half of the X neurons includes recurrent collaterals. 5. T neurons are located mainly in the ventral stratum opticum and the dorsal stratum griseum intermedium. They have small or medium-sized, trapezoid or ovoid cell bodies and relatively simple radiating or vertical dendritic trees. Their axons usually participate in two of the major tectofugal bundles besides providing a commissural component and recurrent collaterals. 6. Morphological details revealed in the present study support the notion that distinct tectofugal axonal systems originate from efferent neurons of the primate SC that differ both as to their location in the tectum as well as the appearance of their somata and dendritic trees. The resulting morphological classification of tectal efferent cells provides a framework for the analysis of tectal function in terms of populations of identified neurons.

Animals↗

PubMed1988

Structure-function relationships in the primate superior colliculus. II. Morphological identity of presaccadic neurons.

1. Microelectrodes filled with horseradish peroxidase (HRP) were inserted in the superior colliculus (SC) of alert squirrel monkeys. Spontaneous eye movements were monitored in the dark during recording and intraaxonal injection of fibers carrying presaccadic signals. 2. Analysis of the relationship between neuronal activity and saccadic parameters indicates that saccade-related neurons can be functionally classified into: 1) vectorial long-lead burst neurons (n = 31), and 2) directional long-lead burst neurons. 3. Vectorial long-lead burst neurons have little if any spontaneous activity and burst intensely before spontaneous saccades within their movement fields with a latency of approximately 20 ms. Their cell bodies were recovered mostly (4/5) in the stratum opticum of the SC. The mediolateral and anteroposterior location of these tectal long-lead burst neurons (TLLBs) together with their movement fields are consistent with existing descriptions of the motor map of the deeper tectal layers. Due to their somatodendritic morphology and pattern of axonal trajectories, TLLBs belong to the T group of tectal efferent neurons that was described in our companion report. Through its branched axonal system each TLLB can relay a signal coding intended eye displacement to reticular targets of the predorsal bundle (PDB), contralateral tectum, ipsilateral mesencephalic reticular formation (MRF), and rostrally located ipsilateral targets of the SC, besides participating in intratectal information processing. 4. Recovered tectal neurons (n = 4) with activity not related to spontaneous saccades participate in the predorsal and ventral ascending tectofugal bundles as well as the projection to the ipsilateral mesencephalic reticular formation. They do not participate in the commissural projection of the SC and need not have recurrent collaterals. Due to their somatodendritic morphology and pattern of axonal trajectories, these cells belong to the X group of tectal efferent neurons that was described in the preceding paper. 5. Recovered cells of origin of directional long-lead burst fibers recorded in the SC (n = 5) are located in the tectorecipient portion of the MRF and their axonal terminals are entirely contained within the SC. The high-frequency portion of the discharge of these reticulotectal long-lead burst neurons (RTLLBs) precedes most contraversive saccades by approximately 19 ms.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

PubMed1987

Inputs from regularly and irregularly discharging vestibular nerve afferents to secondary neurons in the vestibular nuclei of the squirrel monkey. I. An electrophysiological analysis.

1. The electrical excitability of vestibular nerve afferents is related to their discharge regularity (23). Irregular (I) afferents are more excitable than regular (R) afferents. We explored the possibility that the differences in electrical excitability could be used to determine the profile of monosynaptic inputs from the ipsilateral vestibular nerve (Vi) to secondary neurons of the vestibular nuclei. The growth of monosynaptic Vi excitatory postsynaptic potentials (EPSPs) as shock strength is increased should reflect the kinds of afferent input that a secondary neuron receives. We were particularly interested in seeing if cells in the vestibular nuclei could be distinguished as R or I neurons depending on whether they received predominantly regular or irregular inputs. Barbiturate-anesthetized squirrel monkeys were used. 2. Recordings were made from vestibular nerve afferents. Shock strength was expressed as multiples of T, the value needed to recruit 10% of the afferents or, as determined empirically, to evoke a detectable field potential in the vestibular nuclei. Most I afferents (85/87 = 98%) were recruited below 4 X T, whereas most R afferents (197/212 = 93%) were first activated above 4 X T. The relation between latent period and electrical excitability was flat for units with thresholds in the range 1-4 X T. Latent periods increased for units with higher thresholds, especially those first activated above 8 x T. The threshold differences between I and R afferents are maximal if the shock falls at approximately half the mean interval after a naturally occurring action potential. The same results were obtained by having each unit fire to a maximal (16-32 X T) conditioning shock and then determining the threshold to a test shock presented 4 ms later. The latter stimulus configuration was used to study the Vi monosynaptic inputs to secondary neurons. The test shock was raised by successive doublings from 1 X T to the strength of the conditioning shock (16-32 X T). 3. Intracellular recordings were made from neurons located in the superior vestibular nucleus or the rostral parts of the medical or lateral vestibular nuclei. Amplitudes and latent periods of Vi EPSPs were measured from averages of several repetitions of each stimulus pair. Each EPSP was calculated by subtracting the extracellular from the intracellular averaged response. Of the 122 neurons sampled, 115 were judged to be monosynaptically related to the ipsilateral vestibular nerve because their Vi EPSPs had latent periods in the range of 0.7-1.4 ms.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

PubMed1987

Inputs from regularly and irregularly discharging vestibular nerve afferents to secondary neurons in the vestibular nuclei of the squirrel monkey. II. Correlation with output pathways of secondary neurons.

1. Intracellular recordings were made from secondary neurons in the vestibular nuclei of barbiturate-anesthetized squirrel monkeys. Monosynaptic excitatory postsynaptic potentials (EPSPs) evoked by stimulation of the ipsilateral vestibular nerve (Vi) were measured. An electrophysiological paradigm, described in the preceding paper (26), was used to determine the proportion of irregularly (I) and regularly (R) discharging Vi afferents making direct connections with individual secondary neurons. The results were expressed as a % I index, an estimate for each neuron of the percentage of the total Vi monosynaptic input that was derived from I afferents. The secondary neurons were also classified as I, R, or M cells, depending on whether they received their direct Vi inputs predominantly from I or R afferents or else from a mixture (M) of both kinds of Vi fibers. The neurons were located in the superior vestibular nucleus (SVN) or in the rostral parts of the medical or lateral (LVN) vestibular nuclei. 2. Antidromic activation or reconstruction of axonal trajectories after intrasomatic injection of horseradish peroxidase (HRP) was used to identify three classes of secondary neurons in terms of their output pathways: 1) cerebellar-projecting (Fl) cells innervating the flocculus (n = 26); 2) rostrally projecting (Oc) cells whose axons ascended toward the oculomotor (IIIrd) nucleus (n = 27); and 3) caudally projecting (Sp) cells with axons descending toward the spinal cord (n = 13). Two additional neurons, out of 21 tested, could be antidromically activated both from the level of the IIIrd nucleus and from the spinal cord. 3. The Vi inputs to the various classes of relay neurons differed. As a class, Oc neurons received the most regular inputs. Sp neurons had more irregular inputs. Fl neurons were heterogeneous with similar numbers of R, M, and I neurons. The mean values (+/- SD) of the % I index for the Oc, Fl, and Sp neurons were 34.7 +/- 24.7, 51.9 +/- 30.4, and 61.8 +/- 18.0%, respectively. Only the Oc neurons had a % I index that was similar to the proportion of I afferents (34%) in the vestibular nerve (cf. Ref. 26). 4. The commissural inputs from the contralateral vestibular nerve (Vc) also differed for the three projection classes. Commissural inhibition was most common in Fl cells: 22/25 (88%) of the neurons had Vc inhibitory postsynaptic potentials (IPSPs) and 1/25 (4%) had a Vc EPSP. In contrast, Vc inputs were only observed in approximately half the Oc and Sp neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

PubMed1985

Observations on the somatodendritic morphology and axonal trajectory of intracellularly HRP-labeled efferent neurons located in the deeper layers of the superior colliculus of the cat.

Efferent neurons of the deeper layers of the cat's superior colliculus were stained with horseradish peroxidase (HRP) to demonstrate patterns of somatodendritic morphology and axonal trajectory. A combination of somatodendritic and axonal features of the HRP-labeled cells revealed the existence of three major groups of tectal efferent neurons (X, T, and I). X neurons are mostly large and multipolar and participate in the crossed descending and ipsilateral ventral ascending projections of the superior colliculus. The X group includes multipolar radiating (X1), tufted (X2), large vertical (X3), medium-sized vertical (X4), and medium-sized horizontal (X5) neurons. T neurons participate in one or two of the major tectofugal bundles (medial descending ipsilateral, lateral descending ipsilateral, medial dorsal ascending, crossed descending) besides providing a commissural branch. They also issue recurrent collaterals distributed within a more or less restricted area of the deeper layers. The T group includes medium-sized, trapezoid, radiating (T1) and small or medium-sized, ovoid, vertical (T2) neurons. I neurons participate in the ipsilateral descending projection of the superior colliculus. They are small, triangular or ovoid, sparsely ramified cells that provide long, varicose collaterals irregularly distributed within the deeper layers. The majority of T neurons are located in the ventral stratum opticum or dorsal stratum griseum intermediale; X3 and X5 neurons are situated immediately below in the dorsal stratum griseum intermediale, while X1, X2, X4, and I neurons are indiscriminately distributed within the deeper layers. The polythetic classification presented here provides a conceptual framework for the description of tectal efferent neurons. It is open-ended and can thereby accommodate new cells types as indicated by the disclosure of a small horizontal (A) and a small radiating (unclassified) neuron. Moreover, it does not preclude the construction of alternate taxonomies. A dendro-architectonic classification into four groups [vertical (X3, X4, T2, I), horizontal (X5, A), radiating (X1, T1, I), and tufted (X2)] can be made and would relate to the mode of integration of various tectopetal inputs. A classification based on the dorsoventral location of tectal efferent neurons is also possible and would relate to the dorsoventral distribution of neurons with specific response properties.

Animals↗

PubMed1985

Nigral inhibitory termination on efferent neurons of the superior colliculus: an intracellular horseradish peroxidase study in the cat.

Intracellularly recorded responses of deeper tectal neurons to stimulation of the substantia nigra and the cerebral peduncle were obtained to demonstrate the monosynaptic inhibitory nature of the nigrotectal pathway in the cat. We also employed antidromic stimulation (contralateral predorsal bundle and superior colliculus) and intracellular labeling with HRP to demonstrate which types of tectal efferent neurons are nigrorecipient. The response to nigral stimulation in 61% of the cells studied was a monosynaptic IPSP of short duration. Recovered HRP-labeled nigrorecipient neurons include X1 (large multipolar radiating), X2 (tufted), X4 (medium-size vertical), X5 (medium-size horizontal), T1 (medium-size trapezoid radiating), T2 (small ovoid vertical), I (small sparsely ramified), and A (small horizontal) neurons. Nigrorecipient cells participate in all four of the major efferent axonal systems of the deeper tectal layers: crossed descending (X and T neurons), ipsilateral descending (I and T neurons), ascending (A, X, and T neurons), and commissural (T neurons). EPSPs accompanied by long-lasting hyperpolarizing potentials were recorded from the remaining tectal neurons in response to stimulation of the substantia nigra, cerebral peduncle, and pericruciate cortex. Collision experiments indicate that at least part of the excitatory responses of tectal neurons to nigral and penduncular stimulation are mediated by corticotectal fibers traversing the cerebral peduncle and the substantia nigra. Excitatory effects of nigral, peduncular, and cortical stimulation were disclosed in X neurons including the non-nigrorecipient large vertical neurons of the X3 subgroup. Cortical excitatory and nigral inhibitory inputs converge only on X neurons (X1, X2, X4, X5). In this case, nigrally evoked IPSPs were preceded by a brief EPSP. Collectively, these results demonstrate the inhibitory termination of the nigrotectal pathway on a wide variety of deeper tectal efferent neurons. Such findings imply the versatility of the nigral involvement in tectal mechanisms of gaze control. We suggest that the substantia nigra pars reticulata contacts tectal neurons differing as to their response properties and shapes the signal carried by all the major tectofugal bundles.

Animals↗