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Organization of the motoneurons innervating the pelvic muscles of the male rat.

The cytoarchitecture of the motoneuron pool of the male rat was studied at the lumbo-sacral transition area, particularly in L6. In the latter segment a dorso-medial (DM), ventral (V), dorso-lateral (DL), and retrodorso-lateral group (RDL) could be defined. The DL group was associated with a prominent longitudinal dendrite bundle and the CM group with smaller transverse bundles. Moreover, close soma-somatic apposition was found between neurons in these columns. Because L6 gives rise to n. pudendus and contributes to n. ischiadicus, horseradish peroxidase (HRP) was applied to the cut n. ischiadicus and in other experiments injected into the pelvic muscles. Neurons in RDL were labeled following exposure of n. ischiadicus to HRP. Injections in m. levator ani resulted in labeled neurons in the V group, mainly below L6. Injections in m. sphincter urethrae resulted in labeled neurons in the DL group as well as neurons immediately cranial to this column. Musculus ischiocavernosus injections resulted in transport of HRP to neurons in the DL group, primarily in its medial part, and to more cranially located neurons. In addition, some neurons in the V group in L6 were labeled. Following injections in m. bulbocavernosus and m. sphincter ani, labeled neurons were found primarily in the DM group, and to a lesser extent in the V group. Histochemical investigations with staining methods for the localization of acetyl cholinesterase (AChE) and heavy metals (the Timm method) demonstrated that part of the neuropil of DL and of DM were different from the rest of the motoneuron neuropil. In the DL group the area with the diverging staining patterns corresponded to the region of the dendrite bundle. The experimental data indicated and the ultrastructural studies demonstrated that the histochemical differences could be correlated with differences in the composition of the populations of boutons. The comparison of the cytoarchitectural and histochemical data with the results obtained by the aid of the retrograde HRP tracing technique established that mm. sphincter urethrae, ischiocavernosus, bulbocavernosus, and sphincter ani were each innervated by two populations of neurons that were situated in separate areas which had different histochemical properties, and which thus probably have different compositions of their afferent inputs. The duality in the motoneuron pool that innervates the pelvic mucscle might be a reflection of the dual influence on these muscles. As all other striated muscles the pelvic muscles are under voluntary control. However, they are also tightly linked to the function of the pelvic viscera and thus under influence of the autonomic nervous system.

Acetylcholinesterase↗

Propriospinal fibers interconnecting the spinal enlargements in some quadrupedal reptiles.

The cells of origin, course and site of termination of long propriospinal fibers interconnecting the intumescences have been studied with the aid of the horseradish peroxidase technique, as well as with anterograde degeneration techniques, in some quadrupedal reptiles (the lizards Tupinambis nigropunctatus and Varanus exanthematicus, and the turtles Testudo hermanni and Pseudemys scripta elegans). The anterograde degeneration findings suggest that long descending propriospinal fibers from the cervical intumescence are distributed bilaterally to the ventral gray of the lumbar enlargement, including the lateral motoneuron column. Long ascending fibers from the lumbar to the cervical intumescence are distributed, also bilaterally, to the ventromedial part of area VII--VIII. The cells of origin of long descending propriospinal fibers wre found in the medial part of area VII--VIII in the cervical intumescence, particularly contralateral to the injection side. The cells of origin of long ascending propriospinal fibers were found in the lumbar intumescence, also particularly contralateral to the injection side, in the ventromedial part of area VII--VIII. It seems likely that in the reptiles studied the long propriospinal fibers interconnecting the spinal enlargements are in large part organized as crossed connections. The demonstration of long propriospinal fibers in lizards and turtles--i.e. quadrupedal reptiles that move their limbs in a particular diagonal pattern--suggests that these pathways are of great importance for the coordination of forelimb and hindlimb movements.

Afferent Pathways↗

Tectal connections in Python reticulatus.

The origins of the axons terminating in the mesencephalic tectum in Python reticulatus were examined by unilateral tectal injections of horseradish peroxidase. Retrogradely labeled cells were observed bilaterally throughout the spinal cord in all subdivisions of the trigeminal system, with the exception of nucleus principalis, which showed labeled cells only on the ipsilateral side. Labeling of the reticular formation occurred bilaterally in nucleus reticularis inferior magnocellularis, nucleus reticularis lateralis, nucleus reticularis, and the mesencephalic reticular formation. The tectum also receives bilateral projections from the dorsal tegmental field, the nucleus of the lateral lemniscus, and nucleus isthmi, and ipsilateral projections from nucleus profundus mesencephali. A few labeled cells were found ipsilaterally in the locus coeruleus and in nuclei vestibulares ventrolateralis and ventromedialis. In the diencephalon labeled cells were observed ipsilaterally in nucleus ventrolateralis thalami, nucleus ventromedialis thalami, nucleus suprapeduncularis, and in the dorsal and ventral lateral geniculate nuclei. Bilateral labeling was observed in nucleus periventricularis hypothalami. Furthermore, labeling was ipsilaterally present in the ventral telencephalic areas. The tectum in Python reticulatus receives a wide variety of afferent connections which confirm the role of the tectum as an integration center of visual and exteroceptive information.

Afferent Pathways↗

The conduction velocity, number, and diameter of unmyelinated fibers in Remak's nerve.

Remak's nerve in the chicken was examined ultrastructurally and electrophysiologically to determine the characteristics of fibers in the nerve trunk. The ration of unmyelinated fibers to myelinated ones was 111:1. The mean number of unmyelinated fibers was 3555 plus/minus 232 (SEM, n=5) and they had a mean diameter of 0.502 plus/minus 0.034 (SEM) micron. The compound action potential consisted almost entirely of a large diphasic waveform which had a mean peak conduction velocity of 0.62 plus/minus 0.031 (SEM, n=5) m.s-1 at 37 degrees C.

Action Potentials↗

Tactile stimulus predictability modulates activity in a tactile-motor cortical network.

Manipulating objects in the hand requires the continuous transformation of sensory input into appropriate motor behaviour. Using a novel vibrotactile device combined with fMRI, the cortical network associated with tactile sensorimotor transformations was investigated. Continuous tactile stimuli were delivered in a random or predictable pattern to the second digit on the right hand of all subjects. To better distinguish sensory and motor processes, subjects were instructed to make proportionate motor gripping responses with their left hand. A consistent cortical network of activation was revealed that included the supplementary motor, dorsal and ventral premotor, posterior parietal, primary and secondary somatosensory and primary motor cortex. Tracking the unpredictable versus predictable tactile stimulus led to greater delays in motor responses and to increased performance errors. Cortical effects due to stimulus predictability were observed in several components of the network, though it was most evident as increased cortical activation in frontal motor regions during tracking of unpredictable tactile stimuli. In contrast to the proposed hypotheses, primary and secondary somatosensory cortices contralateral to tactile input did not reveal enhanced responses during unpredictable tracking. Facilitation during unpredictable tracking was also observed in primary somatosensory cortex contralateral to motor responses, the receptive site for movement-related afference. The present study provides a novel and controlled approach to investigate the loci associated with tactile-motor processing and to measure the task-specific effect of stimulus predictability on network components.

Adult↗

Response of human jaw muscles to axial stimulation of a molar tooth.

The reflexes of the main jaw-closer muscles (masseter and anterior temporalis) on both sides of the jaw were investigated using surface electromyography to observe reflex activity following mechanical stimulation of the 1st right upper-molar tooth at various forces under a number of levels of jaw-muscle activity. As with analogous studies performed on the incisor, three distinct reflex events were identified in the EMG before the earliest conscious subject reaction: early excitation, inhibition and late excitation. However, contrary to observations found during studies on the incisor, excitation, not inhibition was the primary reflex response. The application of a local anaesthetic block around the stimulated molar showed that the primary agents in eliciting the observed reflexes were not contained within the periodontium of the stimulated tooth. A diminished representation of periodontal mechanoreceptors around the molar teeth and more elaborate root structures, hence a more solid connection to the jaw and consequently less tooth movement, were deemed the likely reason for the distinction between the reflex responses of the incisal and molar regions. In addition to the reflex studies, the minimum reaction time of a number of subjects was determined to permit the distinction of a reflex event and an event that could be a conscious subject reaction. It was found that the reaction time of the temporalis muscles was significantly shorter than those of the masseter, while no significant difference was found between the left and right sides. Overall, the data showed that the presence or absence of background muscle activity and subject variability were the main causes of changes in the reflex response, provided the level of the stimulus was greater than 3 N. The application of local anaesthetic had no impact on the reflexes evoked.

Adolescent↗

Tonic differential supraspinal modulation of PAD and PAH of segmental and ascending intraspinal collaterals of single group I muscle afferents in the cat spinal cord.

We compared in the anesthetized cat the effects of reversible spinalization by cold block on primary afferent depolarization (PAD) and primary afferent hyperpolarization (PAH) elicited in pairs of intraspinal collaterals of single group I afferents from the gastrocnemius nerve, one of the pairs ending in the L3 segment, around the Clarke's column nuclei, and the other in the L6 segment within the intermediate zone. PAD in each collateral was estimated by independent computer-controlled measurement of the intraspinal current required to maintain a constant probability of antidromic firing. The results indicate that the segmental and ascending collaterals of individual afferents are subjected to a tonic PAD of descending origin affecting in a differential manner the excitatory and inhibitory actions of cutaneous and joint afferents on the pathways mediating the PAD of group I fibers. The PAD-mediating networks appear to function as distributed systems whose output will be determined by the balance of the segmental and supraspinal influences received at that moment. It is suggested that the descending differential modulation of PAD enables the intraspinal arborizations of the muscle afferents to function as dynamic systems, in which information transmitted to segmental reflex pathways and to Clarke's column neurons by common sources can be decoupled by sensory and descending inputs, and funneled to specific targets according to the motor tasks to be performed.

Afferent Pathways↗

Responses in sympathetic nerves of the dog evoked by stimulation of somatic nerves.

Responses in thoracic and renal sympathetic nerves evoked by electrical stimulation of cutaneous and muscle nerves in anaesthetized mongrel dogs were observed. Supramaximal stimulation of cutaneous nerves evoked two responses in both thoracic and renal nerves with latencies in the ranges 58--184 msec and 349--733 msec which are referred to as the early and late responses. It was shown that the early and late responses were evoked by group III and group IV afferent fibres respectively. Stimulation of muscle nerves of the forelimb and the hypoglossal nerve evoked smaller early responses which were considered to be due to activation of group III fibres and which had latencies in the range 92--157 msec. Supramaximal stimulation of muscle nerves in the hind limb failed to evoke any responses in approximately two-thirds of preparations and in the remainder only low level inconsistent early responses were observed. No matter how intense the stimuli applied to muscle nerves there were never any responses which could be related to the activation of group IV fibres.

Afferent Pathways↗

Activity of motoneurons during fictitious scratch reflex in the cat.

(1) Intracellular recording of motoneurons of different hindlimb muscles: tibialis anterior (TA), gastrocnemius and soleus (GS), vastus crureus (Vast), posterior biceps and semitendinosus (PBSt), was carried out during the fictitious scratch reflex in decerebrate cats. (2) During the postural stage of the reflex, a depolarizaiton (3.8 mV on average) was observed in TA motoneurons accompanied by tonic discharge. No change of the membrane potential (MP) and no discharge were observed during this stage in GS, Vast and PBSt motoneurons. (3) In the rhythmical stage of the reflex, the MP of TA motoneurons changed only slightly during the 'long' (L) phase of the scratch cycle and remained at approximately the same level as during the postural stage. In this phase, motoneurons discharged at frequencies of 20-100 pps. In the 'short' (S) phase of the scratch cycle a strong repolarization occurred, the MP reached the same level as observed during resting conditons (MP0), and the discharge discontinued. (4) GS motoneurons were gradually depolarized during the second half of the L-phase. The depolarization reached its maximum (5.5 mV on average in relation to the MP0) on average in relation to the MP0) in the S-phase, and several action potentials were generated with intervals of 5-10 msec. Then, at the beginning of the L-phase, the motoneurons were repolarized and the MP reached the level of the MP0. The behavior of Vast motoneurons was essentially similar to that of GS motoneurons. (5) The PBSt motoneurons usually had two peaks of depolarization per cycle--in the S-phase and at the beginning of the L-phase. The maximal depolarization was 3.5 mV (on average). The motoneurons generated action potentials at one or both peaks of depolarization. (6) The possible organization of the central influences upon motoneurons of different muscles during scratching is discussed.

Afferent Pathways↗

Projections from areas 17 and 18 of the visual cortex to the nucleus of the optic tract.

The corticofugal pathway to the nucleus of the optic tract (NOT) in the cat was studied with visual and electrical stimulation in two experimental series. In 10 experiments cells in the NOT were identified, and orthodromic responses evoked by single electric shocks applied through microelectrodes situated at different loci in areas 17 and 18. All but 2 units gave clear responses to shocks applied to either cortical electrode. The mean of response latencies was calculated to be 3.29 ms for area 17 stimulation and 3.04 ms for area 18 stimulation. In 8 further experiments the stimulation and recording sites were reversed; i.e. single shocks through microelectrodes in the NOT were used to elicit antidromic discharges in areas 17 and 18. A third microelectrode was placed in the superior colliculus (SC) at a position in retinotopic register with the units recorded at the NOT stimulation site. Out of 231 cortical units tested, 42 (17%) gave an antidromic response to NOT and/or SC shock. Seventeen units responded both to NOT and SC stimulation, 18 to NOT alone, and 7 to SC alone. The response latencies after NOT shock (mean 2.8 ms +/- 1.5 S.D.) did not differ significantly from those to SC shock (2.9 ms +/- 1.5 S.D.). All cortical cells activated antidromically from NOT and/or SC were located in layer V of areas 17 and 18. These units showed the following response characteristics: they responded well to conventional light bars as well as to large area random dot patterns; they were binocular and showed direction tuning; as compared to NOT cells they were more sharply tuned for stimulus velocity preferring faster movements. The present findings suggest a convergent projection to the NOT from the same type of cortical cells that project to the superior colliculus.

Action Potentials↗

Thalamic bursting mechanism: an inward slow current revealed by membrane hyperpolarization.

Hyperpolarization of the ventrolateral thalamic cell membrane reveals a slow inward current which is not normally observed at the resting membrane potential. The response evoked by depolarizing synaptic potentials or depolarizing current pulses from a hyperpolarized potential consists of a burst of action potentials superimposed upon a slow voltage response, in contrast to the single active response evoked without the background polarization. We propose that such behavior is caused by a slow inward current that is activated at subthreshold potentials and inactivated or masked at resting potential.

Afferent Pathways↗

The intrinsic organization of the vestibular complex: evidence for internuclear connectivity.

The HRP anterograde and retrograde labeling techniques provide evidence for extensive internuclear connectivity within the vestibular complex. Specifically: (1) the superior vestibular nucleus is topographically and reciprocally related to the spinal (spr) and medial vestibular nuclei (mv); (2) the lateral vestibular nucleus (lv) is reciprocally related to the mv, and (3) the lv receives afferent fibers from the spv but does not reciprocate this input.

Afferent Pathways↗

Nipecotic acid: preferential accumulation in the cochlea by GABA uptake systems and selective retrograde transport to brainstem.

[3H]Nipecotic acid was shown to be preferentially accumulated by the same cochlear structures which selectively accumulate [3H]gamma-aminobutyric acid ([3H]-GABA), including the terminals of a subset of olivocochlear neurons. With both amino acids, olivocochlear fibers selectively transported label in a retrograde direction, from cochlea to brainstem. However, only [3H]nipecotic acid produced dense labeling, and labeling of cell bodies in the superior olive, presumably because it is metabolized very slowly. Nipecotic acid appears to provide a selective retrograde tracer, specific to neurons whose terminals exhibit preferential GABA uptake.

Animals↗

Ascending and descending internuclear projections within the trigeminal sensory nuclear complex.

The cells of origin of ascending and descending internuclear pathways in the trigeminal sensory nuclear complex were studied by the method of retrograde transport of horseradish peroxidase in the cat. The cells of origin of the ascending internuclear pathways are distributed in all laminae of the caudal part of the spinal trigeminal nucleus (Vc) except for lamina II and the caudal regions of the pars interpolaris of the spinal trigeminal nucleus (Vi). The cells arising from the Vc project to all rostral trigeminal nuclei except the caudal Vi and dorsal part of the principal trigeminal nucleus (Vpd), and neurons of the caudal Vi project to the dorsomedial (Vo.dm) and rostrodorsomedial (Vo.r) divisions of the spinal trigeminal nucleus and the ventral part of the principal trigeminal nucleus (Vpv), although the main ascending fibers from the Vc arise from laminae III-V and project to the rostral Vi and pars oralis. By contrast, the cells of origin of the descending internuclear pathways are distributed in all trigeminal nuclei, with chain-like connections between the neighboring nuclei, while the caudal regions of the Vi and laminae I-II do not receive any descending projections. The main ascending fibers from the paratrigeminal nucleus (or interstitial nucleus) at the caudal level of the Vi project to the parabrachial nucleus. These findings indicate that the internuclear pathways are differentially organized between the ascending and descending projections, and suggest that the internuclear trigeminal connections have a smaller influence on the trigeminothalamic tract cells in the Vpd, caudal Vi, and lamina I.

Afferent Pathways↗

Reciprocal connections between rostral ventrolateral medulla and inspiration-related medullary areas in the cat.

We investigated connections between the rostral ventrolateral medulla (rVLM) and the two main inspiration-related medullary areas, i.e., the dorsal respiratory group (DRG) and the rostral ventral respiratory group (rVRG) in the cat. Non respiration-related tonically firing units encountered in the rVLM displayed either antidromic or orthodromic responses to DRG or rVRG microstimulation. Some units responded to the stimulation of both regions. We suggest that at least part of rVLM neurons are components of medullary loops operating in the control of breathing.

Afferent Pathways↗

Diagonal ventral forebrain continuum has overlapping telencephalic inputs and brainstem outputs which may represent loci for limbic/autonomic integration.

Growing evidence indicates that three areas within the mammalian basal forebrain share many common features. Based on the similarity of connections and their adjacent spacial proximity, three forebrain nuclei are referred to as a continuum. The components of this diagonal ventral forebrain continuum (DVFC) are the central nucleus of the amygdala, the sublenticular portion of the substantia innominata, and the lateral bed nucleus of the stria terminalis. A primary concern and terminal goal of this study is to determine whether the region of this continuum which projects to the brainstem autonomic nuclei such as the vagal nuclei or the parabrachial nuclei also receives inputs from the basolateral amygdala. The first phase of this study involved determining what autonomic regions receive projections from the basal forebrain. The vagal complex and the parabrachial nuclei were found to receive the densest inputs from the DVFC. The topographic distribution of the respective retrogradely labeled cells and their collateral status is described. The second phase involved looking at afferent inputs from brainstem nuclei. The parabrachial nucleus sends reciprocal projections back to the continuum, which generally overlap the neurons which project back to the brainstem visceral nuclei. The third phase of the study indicated that the cells of the basolateral amygdala contribute a major terminal field which overlaps those cells of the basal forebrain continuum which in turn project to either the nucleus of the solitary tract or the parabrachial nucleus. The possibility that the circuits implied in this study represent the neural circuitry whereby emotional stimuli result in changes in visceral activity is addressed.

Afferent Pathways↗