Electromyographic studies of neck muscles in the intact cat

Summary Natural head movements in alert, unrestrained cats were studied using video-filming, videofluoroscopy and electromyographic (EMG) recording methods. In each cat, up to sixteen neck muscles or neck-muscle compartments were implanted with recording electrodes. Patterns of muscle recruitment were examined during systematically-selected behavioral epochs in which the cat held a range of stationary postures, and when it performed volitional and exploratory behaviors such as flexion-extension or turning, grooming, eating, or headshaking. Patterns of muscular activity were interpreted with reference to simultaneous video images of head and neck movements. In separate, videofluoroscopic analysis, flexion-extension movements were examined to gain insight into the underlying movements of the skull and cervical vertebrae. These and other movements were found commonly to depend upon changes in joint angles between lower as well as upper cervical joints. Stationary postures in which the neck was held vertically were consistently associated with tonic EMG activity in only two long dorsal muscles, biventer cervicis and occipitoscapularis. Less consistent activity was also present in dorsal intervertebral muscles crossing lower cervical joints. When the neck was held horizontally, the long dorsal muscles increased their EMG activity and moderate activity was also recorded in deeper intervertebral and suboccipital muscles. When flexion-extension occurred around upper cervical joints, greatest activity was recorded in rectus capitis posterior and complexus, but when it involved the lower cervical joints, large changes in EMG activity could also be detected in biventer cervicis, occipitoscapularis, and the intervertebral muscles crossing lower cervical joints. During specialized, sagittal-plane movements such as grooming, well-defined patterns of synergy could be recognized that varied according to the degree of involvement of upper and lower cervical joint-sets. Movements in the horizontal plane were associated with EMG activity in a largely different subset of neck muscles including splenius, longissimus capitis and obliquus capitis inferior. The levels of EMG activity during flexion-extension or turning movements were much lower than those observed during other more vigorous behaviors, such as head shaking. Some neck muscles, such as clavotrapezius and sternomastoideus, could only be recruited during forceful or ballistic head movements. Results showed that the patterns of muscular activation were linked not only to the speed and trajectory of the movements of the skull, but also to the kinematics of the motion occurring across different parts of the cervical column.     

Coexistence of peptide immunoreactivity in sensory neurons of the cat

The coexistence of the neuropeptides substance P, cholecystokinin, somatostatin and vasoactive intestinal polypeptide in cat sensory neurons has been examined using peroxidase-anti-peroxidase immunocytochemistry. Attempts were also made to locate cells containing bombesin, neurotensin, [Met]enkephalin and [Leu]enkephalin but no immunoreactivity was found when antisera to these peptides was used. Cells in the dorsal root ganglia were studied by cutting 5 microns serial wax sections or 15 microns cryostat sections. Coexistence was established by applying the antiserum to each peptide to serially adjacent 5 microns sections and establishing the presence of peptide-like immunoreactivity in each of 4 different sections through a single cell. Results showed that the distribution and combinations of coexistence of these neuropeptides in the cat is extremely complex; three and sometimes all four antisera showing immunoreactivity with a single cell. About 21% of all ganglion cells contained some immunoreactivity but there were certainly some small cells which did not contain any immunoreactivity. The coexistence of these peptides differed markedly from that previously reported in the rat suggesting that interspecific differences in the neuropeptide content of cells might be much greater than they are for classical neurotransmitters. The results are discussed in relation to the possible role of neuropeptides and the regulation of their production by sensory neurons.     

Torque vectors of neck muscles in the cat

Summary Anatomical texts describe the neck musculature without measurements of muscle locations or quantitative estimates of pulling actions (torques). This study is based on measurements in stereotaxic coordinates of cat neck muscle origins and insertions, and neck intervertebral rotation axes. Torque vectors in three dimensions were calculated for 14 pairs of dorsal and ventral muscles that insert on the skull or first cervical vertebra. Predicted torque vectors were in general agreement with qualitative statements in the literature. Biventer cervicis and the rectus capitis major, medius, and minor muscles act mainly to raise the head, and longus capitis acts almost exclusively to lower the head. Longissimus capitis, sternomastoid, and cleidomastoid act mainly to roll the head. Complexus acts about equally to raise the head and roll it. Splenius and occipitoscapularis have torque in all three coordinate directions. Torques were altered by changing the pitch of the head with respect to the neck. The calculated neck muscle torques did not correspond to previously reported directions of neck muscle excitation during the vestibulocollic reflex. The neck musculature appears to be a complex, multidimensional system that presents interesting problems in motor control.     

Distribution of neuropeptides in the limbic system of the rat: the hippocampus

The distribution of several neuropeptides (vasoactive intestinal polypeptide, cholecystokinin octapeptide, substance P, neurotensin, methionine-enkephalin and somatostatin) in the hippocampal formation has been studied with immunocytochemical techniques. Numerous vasoactive intestinal polypeptide, cholecystokinin-octapeptide and somatostatin-positive cell bodies were found within the hippocampus and subiculum. Neurotensin-positive cell bodies were found within the subiculum, but no substance P or methionine-enkephalin-containing cell bodies were seen in either hippocampus proper or subiculum. Vasoactive intestinal polypeptide and cholecystokinin-octapeptide-positive cell bodies were predominantly located in the stratum moleculare and stratum radiatum of CA 1-2 regions and dentate gyrus, whilst somatostatin-containing cell bodies were found mainly in the stratum oriens. Nerve fibres containing each of the six peptides were found within the hippocampus. Large numbers of vasoactive intestinal polypeptide, cholecystokinin-octapeptide and somatostatin fibres innervated the pyramidal and granule cell layers, with smaller numbers in the stratum radiatum and fewer still in the stratum moleculare and stratum oriens. Other than a moderately dense neurotensin-positive fibre plexus in the dorsal subiculum, fewer neurotensin, substance P and methionine-enkephalin fibres were present. However, when present, these three peptides had a distribution restricted to a region close to the pyramidal layer in the CA 2/3 region and to the stratum moleculare of the CA 1 region. Peptide-containing fibres were identified entering or leaving the hippocampus in three ways, via (i) the fornix (all six peptides), (ii) the dorsal subiculum (neurotensin-positive fibres projecting to the cingulate cortex: somatostatin, vasoactive intestinal polypeptide, and cholecystokinin-octapeptide present in fibres running between the dorsal subiculum and occipito-parietal cortex) and (iii) the ventral subiculum (vasoactive intestinal polypeptide, cholecystokinin-octapeptide and somatostatin in fibres running between entorhinal cortex and hippocampus, and all six peptides in fibres crossing the amygdalo-hippocampal border). These findings indicate a major distinction between those peptides (vasoactive intestinal polypeptide, cholecystokinin-octapeptide, somatostatin, neurotensin) which are found in cell bodies intrinsic to the hippocampal formation and those peptides (substance P, methionine-enkephalin) which are found only in hippocampal afferents.(ABSTRACT TRUNCATED AT 400 WORDS)     

Auditory corticocortical interconnections in the cat: evidence for parallel and hierarchical arrangement of the auditory cortical areas

Summary The origin and laminar arrangement of the homolateral and callosal projections to the anterior (AAF), primary (AI), posterior (PAF) and secondary (AII) auditory cortical areas were studied in the cat by means of electrophysiological recording and WGA-HRP tracing techniques. The transcallosal projections to AAF, AI, PAF and AII were principally homotypic since the major source of input was their corresponding area in the contralateral cortex. Heterotypic transcallosal projections to AAF and AI were seen, originating from the contralateral AI and AAF, respectively. PAF received heterotypic commissural projections from the opposite ventroposterior auditory cortical field (VPAF). Heterotypic callosal inputs to AII were rare, originating from AAF and AI. The neurons of origin of the transcallosal connections were located mainly in layers II and III (70–92%), and less frequently in deep layers (V and VI, 8–30%). Single unit recordings provided evidence that both homotypic and heterotypic transcallosal projections connect corresponding frequency regions of the two hemispheres. The regional distribution of the anterogradely labeled terminals indicated that the homotypic and heterotypic auditory transcallosal projections are reciprocal. The present data suggest that the transcallosal auditory interconnections are segregated in 3 major parallel components (AAF-AI, PAF-VPAF and AII), maintaining a segregation between parallel functional channels already established for the thalamocortical auditory interconnections. For the intrahemispheric connections, the analysis of the retrograde tracing data revealed that AAF and AI receive projections from the homolateral cortical areas PAF, VPAF and AII, whose neurons of origin were located mainly in their deep (V and VI) cortical layers. The reciprocal interconnections between the homolateral AAF and AI did not show a preferential laminar arrangement since the neurons of origin were distributed almost evenly in both superficial (II and III) and deep (V and VI) cortical layers. On the contrary, PAF received inputs from the homolateral cortical fields AAF, AI, AII and VPAF, originating predominantly from their superficial (II and III) layers. The homolateral projections reaching AII originated mainly from the superficial layers of AAF and AI, but from the deep layers of VPAF and PAF. The laminar distribution of anterogradely labeled terminal fields, when they were dense enough for a confident identification, was systematically related to the laminar arrangement of neurons of origin of the reciprocal projection: a projection originating from deep layers was associated with a reciprocal projection terminating mainly in layer IV, whereas a projection originating from superficial layers was associated with a reciprocal projection terminating predominantly outside layer IV. This laminar distribution indicates that the homolateral auditory cortical interconnections have a feed-forward/feed-back organization, corresponding to a hierarchical arrangement of the auditory cortical areas, according to criteria previously established in the visual system of primates. The principal auditory cortical areas could be ranked into 4 distinct hierarchical levels. The tonotopically organized areas AAF and AI represent the lowest level. The second level corresponds to the non-tonotopically organized area AII. Higher, the tonotopically organized areas VPAF and PAF occupy the third and fourth hierarchical levels, respectively.Abbreviations AAF anterior auditory cortical area - AI primary auditory cortical area - AII secondary auditory cortical area - BF best frequency - C cerebral cortex - CA caudate nucleus - CL claustrum - D dorsal nucleus of the dorsal division of the MGB - ea anterior ectosylvian sulcus - ep posterior ectosylviansulcus - IC internal capsule - LGN lateral geniculate nucleus - LV pars lateralis of the ventral division of the MGB - LVe lateral ventricule - M pars magnocellularis of the medial division of the MGB - MGB medial geniculate body - MGBv ventral division of the MGB - OT optic tract - OV pars ovoidea of the ventral division of the MGB - PAF posterior auditory cortical area - PH parahippocampal cortex - PO lateral part of the posterior group of thalamic nuclei - PU putamen - RE reticular complex of thalamus - rs rhinal sulcus - SG suprageniculate nucleus of the dorsal division of the MGB - ss suprasylvian sulcus - TMB tetrametylbenzidine - VBX ventrobasal complex - VLa ventrolateral complex - VL ventro-lateral nucleus of the ventral division of the MGB - WGA-HRP wheat germ agglutinin conjugated to horse-radish peroxidase - WM white matter - VPAF ventro-posterior auditory cortical area     

Synaptology and sources of vasoactive intestinal polypeptide and substance p containing axons of the cat celiac ganglion. An experimental electron microscopic immunohistochemical study

We have examined the possible origin of vasoactive intestinal polypeptide (VIP) and substance P-containing axons and the synapses formed by these axons in the celiac ganglion of the cat, by means of ultrastructural immunohistochemistry combined with various surgical lesions.Axons containing VIP as well as those immunopositive for substance P, formed axo-dendritic and axosomatic synapses with principal ganglion cells. After transaction of the superior mesenteric nerve numerous degenerated axonal profiles could be found in the celiac ganglion. Some of these contained VIP immunoreactivity. By contrast in alternate sections stained with substance P antibody only intact axons were labelled.Bilateral vagotomy resulted in the appearance of degenerating axonal profiles in the celiac ganglion, some of which could be stained with substance P antiserum but not with VIP antiserum. Following removal of dorsal root ganglia (Th₆-Th₁₂) from both sides, a large number of degenerated axons were found, many of which were immunopositive for substance P but not for VIP.We conclude that the peripheral input to the celiac ganglion contains VIP fibers which form synapses with principal ganglion cells. Substance P-containing fibers reach the celiac ganglion via the vagal nerve as well as from the dorsal root ganglia, and they form synapses with the principal ganglion cells. It is proposed that the VIP and the substance P-containing axons are involved in two different peripheral reflex loops.     

Further studies on the fiber connections of the central cervical nucleus in the cat

Summary The course and cerebellar termination of the axons of the cells in the central cervical nucleus (CCN) was studied in five cats after injections of wheat germ agglutinin-horseradish peroxidase conjugate (WGA-HRP) into the C1-4 segments involving the CCN. In two cats a hemisection was performed ipsilateral to and above the injections in order to prevent transport of WGA-HRP from cerebellar-projecting neurons in lamina VI, just dorsal to the CCN. Labeled axons were found in the brain stem contralateral to the injections just lateral to the spinal trigeminal tract and the vestibular nuclei, and in the reticular formation. Some fibers terminated in the vestibular nuclei (especially group x), the reticular formation, and the inferior olive. The axons entered the cerebellum via the superior cerebellar peduncle. In the cerebellum mossy fiber terminals were found bilaterally in the deep vermal parts of lobules I–VIII. Only in cases without lesions were terminals found in the paramedian lobule ipsilateral to the injection, suggesting that neurons in lamina VI, but not in the CCN, project to the paramedian lobule. In the brain stem retrogradely labeled neurons (possible afferents to the CCN) were found in the vestibular (lateral, inferior and medial) nuclei, the reticular formation, and the trigeminal (spinal and mesencephalic) nuclei. The cerebellar distribution of mossy fiber terminals suggests that spinocerebellar fibers from lower parts of the spinal cord passing through the injection area do not transport WGA-HRP to their terminals.     

Horizontal eye position-related activity in neck muscles of the alert cat

Summary The activity of neck muscles was recorded in the alert, head-fixed cat together with the horizontal and vertical components of eye movements. Electromyographic activity of obliquus capitis cranialis and caudalis, and longissimus capitis, is closely related to horizontal eye position in the orbit both during spontaneous eye movements and vestibular nystagmus. The activity of splenius also shows this relationship but the coupling is less tight, probably because of the postural function of this muscle.Supported by INSERM PRC grant no. 50-7723     

The vestibular nuclei of the cat receive a primary afferent projection from receptors in extraocular muscles

Summary The vestibular nuclei of adult cat were injected with retrogradely transported tracers, WGA-HRP or fluorescent Diamidino-Yellow. Labelled cells were found in the caudal half of the ipsilateral mesencephalic trigeminal nucleus, in the area where ganglionic cells of the sensory receptors in the extraocular muscles have previously been described. Double labelling experiments were carried out with Diamidino Yellow injected in vestibular nuclei and Fast Blue in extraocular muscles. Some cells in the mesencephalic trigeminal nucleus were found to contain both tracers, providing evidence that vestibular neurons do receive direct afferent signals from extraocular muscles. Therefore, this anatomical demonstration suggests a direct feed-back control between the extraocular muscle receptors and the vestibular nuclei.     

Effect of substance P on CCK- or VIP-induced choleresis in anesthetized dogs

10 anesthetized dogs were provided with acute common bile duct fistulas and the gallbladder was excluded. Hepatic bile output and biliary content of sodium, potassium and amylase were studied. 6 caval infusions were administered of CCK, 0.3 Ivy U-kg^-1 min^-1, with a superimposed infusion of SP, 20 ng kg^-1 min^-1. 7 caval infusions were given of VIP, 50 ng-kg^-1 min^-1, with a superimposed infusion of SP, 20 ng·kg^-1 min^-1. CCK increased bile output and biliary content of sodium, potassium and amylase by 78–110%. The corresponding increase induced by VIP was 55–85%. Biliary pH was not influenced. SP abolished the effects of both CCK and VIP. It is suggested that all peptides studied influenced canalicular bile secretion by changing the electrolyte excretion.     

大鼠终纹床核卵圆核及其邻近区域的传出纤维联系——WGA-HRP顺行追踪法研究

本实验选用150～260g的雄性Sprague-Dawley大鼠13只,把WGA-HRP/HRP混合水溶液加压注入一侧终纹床核群前外侧区的卵圆核区域,冰冻切片,TMB法呈色后,在中枢看到顺行标记终末最密集的部位是:下丘脑后部外侧区、中央杏仁核、中脑中央灰质、臂旁核、三叉神经中脑核、蓝斑;比较多的部位是视前区、下丘脑室周区、弓状核、丘脑中线核群、内侧纽核、腹侧背盖核、脚桥背盖核、中脑网状结构、中缝背核以及迷走神经复合体;在线形中缝核、中央上核、腹侧背盖区、黑质,以及延髓中介核,也看到少量标记终末。本工作对卵圆核的传出纤维联系,进行了较全面的观察。     

Organization of cingulo-ponto-cerebellar connections in the cat

Summary This study deals with three different aspects of the organization of connections from the cingulate gyrus to the cerebellum. (1) With the use of wheat germ agglutinin-horseradish peroxidase as a retrograde tracer, the distribution of cingulate neurons projecting to the pontine nuclei was studied. Retrogradely labeled cells were found in layer 5 in all parts of the cingulate gyrus. Average densities of cingulopontine cells were similar in the different cytoarchitectonic subdivisions, although some density gradients were observed. The projection was found to be remarkably strong. Average densities of corticopontine cells in the cingulate gyrus ranged from 500–700 cells per mm² cortical surface, and the total number of neurons was in the range of 75000–105000 (n=4). (2) A topographical organization of terminal fields of fibers originating in different parts of the cingulate gyrus was demonstrated with the combined use of anterograde degeneration and anterograde transport of wheat germ agglutinin-horseradish peroxidase. Terminal fibers originating in different zones of the cingulate gyrus were distributed in a patchy mosaic within a narrow band along the ventromedial aspect of the pontine nuclei. (3) We confirm, with the combined use of lesions in the cingulate gyrus and injections of wheat germ agglutinin-horseradish peroxidase in the ventral paraflocculus, that there is considerable overlap between terminal fibers originating in the cingulate gyrus, and cells retrogradely labeled from the ventral paraflocculus. The role of the ventral paraflocculus as a receiver of limbic input is discussed.This paper is dedicated to Professor Fred Walberg on the occasion of his 70th birthday     

Physiological properties of muscle spindles in dorsal neck muscles of the cat.

1. Single-fiber recording was used to examine the properties of 107 spindle endings in cat biventer cervicis (BC) and complexus (CM) muscles. Responses of receptors were examined following muscle contraction and ramp and hold stretch. Twenty-two endings in splenius (SP) were also examined, but their responses could not be quantitated because the anatomy of SP prevented the application of appropriate stretches. 2. Conduction velocitites of spindle afferents ranged from 13 to 90 m/s. Endings with primary response patterns usually had faster conduction velocities than secondary endings, but there was overlap in the conduction velocity ranges of the two subgroups. 3. Most neck spindle afferents could be classified as either primary or secondary by a constellation of physiological criteria including dynamic response pattern, dynamic index, and variability of resting discharge frequency. However, 22 of 107 endings from BC and CM had responses with characteristics intermediate between primary and secondary responses. The possible sources of these characteristics are discussed. 4. Despite the similarity in properties between spindles of different neck muscles, the length sensitivities of CM spindles were high compared to those of BC spindles. CM spindles showed length-related modulation of firing frequency over a more restricted range of initial muscle lengths than did BC spindles. 5. Eight Golgi tendon organs (GTO) were identified by their characteristics responses. Conduction velocities obtained for five GTO afferent nerves ranged from 50 to 67 m/s. Recordings were also made from receptros in deep muscles surrounding the vertebrae. These receptors had properties characteristic of muscle spindles.     

Efferent fibers from the motor cortex terminate bilaterally in the thalamus of rats and cats

Summary The anterograde transport of lectin-conjugated horseradish peroxidase (WGA-HRP) was here employed in order to visualize crossed cortico-thalamic efferents of the motor cortex in rats and cats. After WGA-HRP cortical injections in the rat retrogradely labeled cells were observed in the ipsilateral thalamus, and heavy anterograde labeling was observed both in the ipsi-and contralateral thalamus. The contralateral anterograde labeling was less intense than the ipsilateral one and it was distributed in the anterior intralaminar structures, in the parafascicular nucleus, in the ventromedial, ventrolateral and ventrobasal nuclei and in the posterior complex, symmetrically to the labeling observed on the ipsilateral side. Further experiments were made in the rat in order to ascertain that the bilateral anterograde labeling in the thalamus derived unilaterally from the cortex. To this purpose, kainic acid was injected unilaterally either into the frontal cortex or into the thalamus, and WGA-HRP was later injected on the same side in the frontal cortex. Moreover, WGA-HRP was injected into the frontal cortex after splitting of the corpus callosum. The results obtained in these experiments confirmed that cortical neurons projected bilaterally upon the thalamus. Further, these experiments indicated that at least the majority of the contralateral fronto-thalamic fibers crossed the midline in the thalamic massa intermedia. WGA-HRP injections into the pericruciate cortex in the cat confirmed the presence of anterogradely labeled terminals in the contralateral anterior and posterior intralaminar, ventral anterior, ventromedial and ventrolateral nuclei. The labeling was in all cases heavier in the intralaminar nuclei than in the other structures, but it was less intense than that observed in the rat. The labeling in the contralateral principal nuclei was heavier in one case in which the injection area diffused into the proreal cortex than in the other cats. Thus, the present results indicate that frontal efferents terminate more densely in the rat's than in the cat's contralateral thalamus. In the latter species frontal fibers are mainly distributed upon the contralateral intralaminar nuclei, whereas in the rat frontal efferents are also heavily distributed upon the ventral nuclear complex.     

大鼠屏状核的传入联系

采用WGA－HRP和CB－HRP法，追踪了16只大鼠屏状核的传入纤维联系，结果表明大脑皮质的躯体感觉区，视皮质及扣带皮质有细胞发出纤维投射到屏状核，后脑腹侧核，未定带，中缝背核及脑脚周核投射到屏状核，下后脑外侧核，视前大细胞核，斜角带核水平支和蓝斑青少量纤维投射到屏状核。     

Distribution of neuropeptides in the limbic system of the rat: the bed nucleus of the stria terminalis, septum and preoptic area

The distribution of the neuropeptides vasoactive intestinal polypeptide, cholecystokinin octapeptide, substance P, neurotensin, methionine-enkephalin and somatostatin has been mapped immunocytochemically in the bed nucleus of the stria terminalis, one of the major sites of termination for efferent projections from the amygdala. Immunoreactive fibres and terminals were distributed more or less topographically and largely in accordance with the previously described localization of peptide-containing cell bodies in the amygdala and the amygdaloid projection fields in the bed nucleus as described by neuroanatomical techniques. Thus, vasoactive intestinal polypeptide, which was found in some of the lateral amygdaloid nuclei, had a substantial projection to the lateral bed nucleus. The lateral bed nucleus also contained cholecystokinin-octapeptide, substance P, neurotensin and methionine-enkephalin immunoreactivity which probably derived from the central amygdaloid nucleus, whilst cholecystokinin-octapeptide, and especially substance P-containing fibres, were found in the medial bed nucleus and probably arise from cells in the medial amygdala. Reciprocal amygdalopetal projections were suggested by the presence of substance P- and somatostatin-containing cell bodies in the mediodorsal bed nucleus and vasoactive intestinal polypeptide cells in the lateral bed nucleus, but somatostatin otherwise had a widespread distribution. Numerous local peptidergic connections were also noted both within the bed nucleus and between it and adjacent structures, especially the preoptic area, hypothalamus and the basal ganglia. These data provide further evidence that neuropeptides play a major role in the connectivity of the limbic system and show that the bed nucleus of the stria terminalis is an important relay station, particularly between amygdaloid efferents and other forebrain areas.     

Electromyographic investigations of the intercostal muscles in experimental poliomyelitis

Summary A study was made of the electrical activity of the intercostal muscles during the development of experimental poliomyelitis.Action potentials were recorded with animals breathing naturally and also during action of specific respiratory stimulants — CO₂ (3 and 10%) and corconium (1 and 10 mg/kg). In the incubation period of the disease the electromyographic picture of the intercostal muscles, was normal, both during natural respiration and during the action of respiratory stimulants. With further progress of the disease there was a rise of the excitation of the intercostal muscle centers and of their regular reaction in response to weak and strong stimuli. The results obtained could be explained by the capacity of the corresponding nerve centers to become rapidly inhibited, the latter state presenting features of a parabiotic character.(Presented by Active Member AMN SSSR S. V. Anichkov) Translated from Byulleten' Éksperimental'noi Biologii i Meditsiny, Vol. 57, No. 3, pp. 49–53, March 1964