Quantitative inter-segmental and inter-laminar comparison of corticospinal projections from the forelimb area of the primary motor cortex of macaque monkeys

Corticospinal projections from the forelimb area of the primary motor cortex to the C2-Th2 spinal cord segments were quantitatively analyzed using the high resolution anterograde tracer, biotinylated dextran amine (BDA), in rhesus monkeys (n=5). The majority of descending axons were located in the contralateral dorsolateral funiculus (DLF) (85-98%), but a minor portion was observed in the ipsilateral DLF (1-12%) and ventromedial funiculus (VMF) (1-7%). In the gray matter, axon collaterals and terminal buttons were found mainly in the contralateral laminae VI-VII and IX and ipsilateral lamina VIII. The majority of projections to the contralateral gray matter originated from the contralateral DLF, but a minority originated from the ipsilateral DLF. Axons from the ipsilateral DLF were not found to project collaterals on the ipsilateral side, but directly entered the contralateral side after crossing the midline. On the other hand, projections to the ipsilateral lamina VIII were from the ipsilateral VMF, and commissural axons were from the contralateral DLF. Terminal buttons in the motoneuron pool in the contralateral lamina IX were found mainly at the C7-Th1 spinal cord segments, whereas the projections to the contralateral laminae VI-VII, ipsilateral lamina VIII, and commissural axons were also found in more rostral segments, abundantly at the C4-C8 segments, 1-3 segments rostral to the motoneuronal projections. These results suggest that cortical control of contralateral forelimb motoneurons accompanies regulation of interneuronal systems in the contralateral laminae VI-VII and the ipsilateral lamina VIII located a few segments rostral to the motoneurons.     

Spinal commissural neurons mediating vestibular input to neck motoneurons in the cat upper cervical spinal cord.

Spinal commissural neurons (CNs) activated di- or trisynaptically by stimulation of ipsilateral vestibular afferents were stained with intraaxonal injection of horseradish peroxidase in the cat upper cervical spinal cord. Stem axons of CNs in lamina VIII or VII, after crossing the midline, had ascending and/or descending main branches that gave off multiple axon collaterals to laminae IX and VIII over a few cervical segments. Terminal boutons appeared to make contact with proximal dendrites and somata of retrogradely-labelled neck motoneurons. Therefore, these CNs were regarded as mediating vestibular afferent input to contralateral neck motoneurons trisynaptically at the shortest.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary Collateralization and termination of single C3-C4 propriospinal neurones (PNs) have been studied in the C6-Th1 segments of the cat using two methods: threshold mapping for antidromic activation of C3-C4 PNs and intra-axonal injection of horseradish peroxidase. Low threshold points for antidromic activation of C3-C4 PNs were found in the region of different motor nuclei in lamina IX both at one level and at different segmental levels, in all parts of lamina VII, in the lateral part of lamina VI and in the dorsal and ventral parts of lamina VIII. Collaterals were found from C6 to Th1. A marked decrease of conduction velocity of the stem axon occurred in the caudal region of termination, while it was almost constant in the rostral region of termination. HRP was injected iontophoretically in C6-Th1 into stem axons of neurones, which were activated antidromically from the ventral part of the lateral funiculus in C5/C6, from the lateral reticular nucleus (LRN) and monosynaptically from the corticospinal fibres (stimulated in the contralateral pyramid) which were transected in C5/C6. Reconstruction of successfully stained stem axons, revealed collaterals with terminals on presumed motoneurones in different parts of lamina IX and on interneurones in laminae IV–VIII. These findings confirm previous results which showed monosynaptic projections from C3-C4 PNs to forelimb motoneurones and Ia inhibitory interneurones. With respect to termination in the region of the motoneurones in lamina IX and in the region of Ia inhibitory interneurones in lamina VII, three patterns were found: 1) termination mainly in lamina IX (n=1) 2) termination in laminae IX and VII (n=15) and 3) termination mainly in lamina VII (n=2). However, in some cases the same stem axon gave off collaterals which terminated either on motoneurones in lamina IX or on presumed Ia inhibitory interneurones in lamina VII. Furthermore, when the stem axons had collaterals which terminated in different motor nuclei only some of these collaterals had additional terminations on presumed Ia inhibitory interneurones. This result suggest that C3-C4 PNs do not follow a strict Ia pattern of reciprocal innervation. It is tentatively proposed that the difference of innervation may be related to the type of multi-joint movement, such as target-reaching with the forelimb, which has been shown to be controlled by the C3-C4 PNs. Termination in laminae VI, VIII and different parts of lamina VII indicates that C3-C4 PNs also project to other types of neurones than motoneurones and Ia inhibitory interneurones. Injection of wheat germ agglutinated horseradish peroxidase (WGA-HRP) laterally in laminae VI-VII in C3 and C4 caused anterograde labelling of axonal bundles from neurones in these segments. Labelled axons were found mainly in the lateral funiculus with the highest density in the ventral part. These axons could be traced throughout the forelimb segments and also to the LRN.     

Retrograde labeling of migrating spinal motoneurons in bullfrog larvae

Horseradish peroxidase (HRP) was applied to the ventral root or developing limb bud of bullfrog tadpoles. In animals younger than Stage XII/XIII [9] ventral root application consistently labeled neurons located between the lateral motor column (LMC) and the neuroepithelium. Only occasional motoneurons outside the LMC were labeled in animals older than Stage XIV/XV. Limb bud application of HRP in any-stage tadpole did not label ectopic LMC motoneurons. These results indicate that LMC motoneurons have sent their axons out the ventral root before completion of migration but that these axons may not as yet have reached the limb bud.     

Permanent central synaptic disconnection of proprioceptors after nerve injury and regeneration. I. Loss of VGLUT1/IA synapses on motoneurons

Motor and sensory proprioceptive axons reinnervate muscles after peripheral nerve transections followed by microsurgical reattachment; nevertheless, motor coordination remains abnormal and stretch reflexes absent. We analyzed the possibility that permanent losses of central IA afferent synapses, as a consequence of peripheral nerve injury, are responsible for this deficit. VGLUT1 was used as a marker of proprioceptive synapses on rat motoneurons. After nerve injuries synapses are stripped from motoneurons, but while other excitatory and inhibitory inputs eventually recover, VGLUT1 synapses are permanently lost on the cell body (75-95% synaptic losses) and on the proximal 100 μm of dendrite (50% loss). Lost VGLUT1 synapses did not recover, even many months after muscle reinnervation. Interestingly, VGLUT1 density in more distal dendrites did not change. To investigate whether losses are due to VGLUT1 downregulation in injured IA afferents or to complete synaptic disassembly and regression of IA ventral projections, we studied the central trajectories and synaptic varicosities of axon collaterals from control and regenerated afferents with IA-like responses to stretch that were intracellularly filled with neurobiotin. VGLUT1 was present in all synaptic varicosities, identified with the synaptic marker SV2, of control and regenerated afferents. However, regenerated afferents lacked axon collaterals and synapses in lamina IX. In conjunction with the companion electrophysiological study [Bullinger KL, Nardelli P, Pinter MJ, Alvarez FJ, Cope TC. J Neurophysiol (August 10, 2011). doi:10.1152/jn.01097.2010], we conclude that peripheral nerve injuries cause a permanent retraction of IA afferent synaptic varicosities from lamina IX and disconnection with motoneurons that is not recovered after peripheral regeneration and reinnervation of muscle by sensory and motor axons.     

Spinal cord localization of the motoneurons innervating the sacrococcygeus dorsi lateralis muscle and their noradrenergic nerve terminals in rats

We examined in the present study the spinal cord localization of motoneurons innervating the caudal portion of the sacrococcygeus dorsi lateralis (SCDL) muscle and their noradrenergic nerve terminals in Sprague-Dawley rats, using horseradish peroxidase (HRP) and dopamine-β-hydroxylase (DBH) double-labeling techniques. Retrogradely HRP-labeled motoneurons innervating the caudal part of the SCDL muscle were located ipsilaterally in the ventromedial aspect of the ventral horn (lamina IX) in spinal segments of S₂–S₄. These cells were polygonal in shape, with an average soma diameter of 37.0 ± 1.1 μm (mean ± S.E.M., n = 95) and amounted to 33.6 ± 5.7 (n = 7) in the horizontal plane. Of note was the presence of abundant DBH-positive nerve terminals arborizing on the soma and dendrites of HRP-labeled motoneurons. These results provided anatomical evidence to further support our previous findings that the coerulospinal noradrenergic neurotransmission is involved in the mediation of fentanyl-induced muscular rigidity.     

Arborization of axons in oculomotor nucleus identified by vestibular stimulation and intra-axonal injection of horseradish peroxidase

Summary Axons in the medial rectus (MR) subdivisions of the oculomotor nucleus were identified by horizontal rotation and by electrical stimulation of the vestibular nerves and abducens nuclei. Three types of axons (vestibular type I and II and abducens interneurons) were then injected intra-axonally with horseradish peroxidase (HRP). Each injected axon was reconstructed under the microscope in the frontal and horizontal planes and terminal arborization and boutons contacting with MR motoneurons were studied. The MR motoneurons were identified by retrograde uptake of HRP, HRP being injected in the MR muscle prior to the intra-axonal experiment.The main types of horizontal canal-related axons were as follows: (1) ATD-unilateral termination axons: Most type I axons were of this type. Axons ascended in ascending tract of Deiters (ATD) to the oculomotor nucleus and terminated in ipsilateral MR area. (2) ATD-bilateral termination axons: Very few secondary canal responsive axons were in this group. Axons ascended in ATD to the oculomotor nucleus and terminated in MR motoneuron areas bilaterally and in the Edinger-Westphal nucleus. (3) MLF-bilateral termination axons: Most type II neurons were in this group. Axons went up in the contralateral MLF and into both oculomotor nuclei. Their branches distributed to several motoneuron areas but only infrequently to the MR area; and to the Edinger-Westphal nucleus. (4) AB interneuron axons: Axons ascended in the MLF contralateral to cells of origin and terminated in the contralateral MR motoneuron area.Supported by USPHS Grant No. 06658     

An HRP study of location of the rabbit palatopharyngeal motoneurons and peripheral course of their axons

The location of the rabbit palatopharyngeal motoneurons and the peripheral course of their axons were investigated with intramuscular injection of HRP either in the normal animal or in conjunction with intracranial severing of the vagal rootlets. Labeled palatopharyngeal motoneurons were ipsilaterally located within a subdivision of the nucleus ambiguus which is formed by a compact arrangement of the smallest neurons of the nucleus and situated in the rostral of the nucleus. We named that subdivision the compact cell group (CoG). The labeled motoneurons occupied the caudal half of the CoG at a level from about 500 to 1,900 microns rostral to the obex. Labeling was completely abolished by severing the vagal rootlets, indicating that the axons of the palatopharyngeal motoneurons traversed the vagal rootlets.     

Motoneuronal and motor axonal innervation in the rat hindlimb: A comparative study using horseradish peroxidase

In order to validate the horseradish peroxidase (HRP) technique as a quantitative method for assessing neuronal pools, we compared counts of labeled motoneurons to numbers of corresponding motor axons. Rat spinal motoneurons were labeled by immersing in HRP either the L4 radicular nerve or the normal or deafferented nerve to the tibialis anterior muscle (TA). The technique slightly under-valued the L4 motoneuronal population estimated by counting ventral root motor fibres, while in deafferented TA, counts of labeled neurons closely reflected numbers of motor axons. Measurements of neuron and nerve fibre diameters revealed good correlations of alpha and gamma motor estimates at spinal cord and nerve levels. Values were considered representative because of the approximation obtained in HRP-estimates between control and deafferented animals, indicating no significant damage from the surgical deafferentation.     

An electron microscopic examination of the corticospinal projection to the cervical spinal cord in the rat: lack of evidence for cortico-motoneuronal synapses

We investigated whether direct, cortico-motoneuronal connections are present in the rat, using both light microscopic and electron microscopic techniques. Corticospinal fibres were labelled using the anterograde tracer, biotinylated dextran-amine (BDA), which was injected into forelimb sensorimotor cortex. Motoneurons were retrogradely labelled after injection of cholera toxin subunit B (CTB) into forelimb muscles, contralateral to the injected hemisphere. Terminals of peripheral afferent fibres, which were also labelled by CTB, were easily distinguishable from, and much larger than, BDA-labelled corticospinal terminals. At the light microscope level, corticospinal terminals were found in all laminae contralateral to the injection site, most extensively in laminae VI and VII of cervical segments C5–C8. Although labelling in the ventral horn (lamina IX) was present, it was extremely sparse. A total of 47 corticospinal synapses were studied at the electron microscope level; most of these were in lamina VII and the majority (35/47; 74%) made axo-dendritic contacts with asymmetrical synapses; one made an axo-somatic synapse, and in the remaining 11 cases no postsynaptic structure could be identified. All corticospinal terminals contained spherical boutons. Serial sectioning of eight BDA-labelled corticospinal boutons in lamina IX revealed that most (seven out of eight) did not make synaptic contacts with any neuronal structure, and none made any contact with adjacent dendrites of CTB-labelled motoneurons. Thus these results provide no positive ultrastructural evidence for direct cortico-motoneuronal synaptic connections within lamina IX between corticospinal axon boutons and the proximal dendrites of forelimb motoneurons. The results confirm other lines of evidence suggesting that such connections are not present in the rat. Electronic Publication     

Synaptic organization of dorsal root projections to lumbar motoneurons in the clawed toad (Xenopus laevis)

Summary Synaptic connexions between dorsal root primary afferents and lumbar motoneurons have been investigated in the isolated spinal cord of the clawed toad. The study of monosynaptic actions evoked in motoneurons by 9th or 10th dorsal root stimulation or by impulses in single primary afferents provided evidence for both electrical and chemical junctional transmission at the sensory-motor synapses. The anterograde filling of the 9th and 10th dorsal roots with horseradish peroxidase (HRP) shows that afferents do project to the motoneuron field of the segments IX and X. Some of the fibres not only reach the dorsally located motoneurons, but also cross the lateral motor column (LMC) and terminate in the marginal zone of ventral horn gray matter. The projections of the 9th and 10th dorsal root fibres are most numerous in the caudal part of segment X. Simultaneous HRP labeling of single motoneurons and the whole 10th dorsal root has revealed that afferent fibres make contacts not only on the distal dendrites of the motor cells, but also on the proximal ones. This latter finding is in a good agreement with the electrophysiological data.Dr. Shiriaev died September 11, 1984     

Axonal trajectory of single group Ia and Ib fibers in the cat spinal cord

Intracellular staining with HRP of physiologically identified group Ia and Ib afferent fibers in the adult cat lumbosacral cord revealed that group Ia and Ib fibers take a similar course in the dorsal funiculus, but the collaterals emerging from them show a different topographical distribution and a different mode of branching in the gray matter. Ia collaterals terminate in laminae VI, VII, IX, and sometimes VIII, whereas Ib collaterals terminate only in lamina VI, or both VI and IX. In lamina IX, two large motor-type neurons received terminations of both Ia and Ib fibers at the same time.     

Retrograde HRP study of neurons in the cervical enlargement projecting to the cerebellum in the cat

Summary After cerebellar HRP injections in kittens labeled neurons were found in laminae V–VIII in the cervical enlargement. Most of the labeled neurons were localized in two groups, one in laminae V–VI, the other centrally in lamina VII. Labeled neurons were also observed in the medial part of lamina VII of C5 and T1 and a few in lamina VIII. Neurons in the cervical enlargement seem to terminate largely in cerebellar lobules IV–V of the anterior lobe. Some neurons in laminae V–VI terminated in the ipsilateral paramedian lobule. Neurons in laminae V–VI and central lamina VII of C5–T1 had uncrossed axons. Neurons in medial lamina VII of C5, in lamina VIII and neighbouring parts of lamina VII of C6–T1 had crossed axons. The ramifications of proximal dendrites and axons of the labeled neurons are described using the tetramethylbenzidine (TMB) method for HRP histochemistry. The neurons in the various laminae differed in their characteristic morphology. In conclusion, the findings of Matsushita et al. (1979) concerning the localization and axonal course of cerebellar projecting neurons in the cervical enlargement are confirmed. In addition new data concerning the morphology of the labeled neurons are presented.     

Central distribution of efferent and afferent components of the pudendal nerve in rat

Summary Central distribution of efferent and afferent components of the pudendal nerve was examined in the rat by the horseradish peroxidase (HRP) method after HRP application to the central cut end of the pudendal nerve. The pudendal motoneurons were located in the dorsolateral, dorsomedial and lateral groups at L5 and L6. Each of the dorsolateral and dorsomedial groups constituted a slender longitudinal cell column. Pudendal motoneurons in the lateral group were scattered at L5, rostrodorsally to the dorsolateral group. The neurons in the dorsolateral and lateral groups were labelled with HRP applied to the nerve branch innervating the ischiocavernosus and sphincter urethrae muscles. The neurons in the dorsomedial group were labelled with HRP applied to the branch supplying the sphincter ani externus and bulbospongiosus muscles. Some dendrites of pudendal motoneurons in the dorsomedial group extended to the contralateral dorsomedial group. These crossing dendrites were observed not only in male rats but also in female. The average number of the pudendal motoneurons in the dorsolateral and dorsomedial groups were larger in male rats than in female. A few neurons of the intermediolateral nucleus at upper L6 were also labelled with HRP applied to the dorsalis penis (clitoridis) nerve. Axon terminals of the pudendal nerve were distributed, bilaterally with an ipsilateral predominance, to the gracile nucleus, as well as to the dorsal horn and dorsal commissural gray from L4 to S2. A few labelled axons were seen in the intermediolateral nucleus at L6 and S1. Axon terminals from the dorsalis penis nerve were distributed more medially in the dorsal horn than those from the perinealis nerve.     

Location of the motoneurons innervating the transverse mandibular muscle in the guinea pig.

Motoneurons supplying the transverse mandibular muscle (TMM) in the guinea pig have been traced by injecting wheat germ agglutinin-horseradish peroxidase (WGA-HRP) in the TMM, and after applying HRP to the mylohyoid nerve. The TMM is bilaterally innervated by 22-36 motoneurons in each trigeminal motor nucleus, forming a column located ventrolaterally along the entire length of the superficial masseter motoneuron group. The axons are incorporated to the mylohyoid nerve. The location, the axon pathways in the brainstem and the pattern of the dentritic tree suggest that in the guinea pig the TMM motoneurons are involved in the masticatory movements in coordination with other jaw-closing muscles.     

The morphology of hair follicle afferent fibre collaterals in the spinal cord of the cat

1. The enzyme horseradish peroxidase (HRP) was injected into single axons that innervated hair follicle receptors to study the morphology of their collaterals in the dorsal horn of the cord. The axons were impaled near the dorsal root entrance zone in the lumbosacral spinal cord of anaesthetized cats and HRP injected by passing current through the intra-axonal micro-electrode. The morphology was revealed by subsequent histochemistry.2. Thirteen hair-follicle afferent fibres were stained including six that innervated tylotrichs (type T hair follicle afferent units) and one that innervated guard hairs (type G unit). The remaining six axons were not classified according to hair type, but, on the basis of their axonal conduction velocities, would have been either type G or T.3. Eleven axons could be traced back into the dorsal roots. Eight of these, upon entering the cord, turned and ran towards the brain. They did not divide into rostral and caudal branches. Three of the eleven did divide and gave rise to both rostral and caudal branches.4. Sixty-three collaterals were given off the thirteen stained axons. All well-filled collaterals had a strikingly similar morphology. They descended through laminae I-III of the dorsal horn into the deeper parts of lamina IV or into lamina V, before turning and ascending back into superficial lamina IV and lamina III where they branched profusely to give rise to their terminal arborizations. Terminal boutons, most commonly of the ;en passant' type, were numerous in lamina III, but were also seen in the dorsal part of lamina IV and in ventral lamina II. None were observed in dorsal lamina II or near the junction of the grey and white matter (lamina I) or in lamina V.5. The terminal arborizations of collaterals from a single hair follicle afferent fibre were in line with one another in the longitudinal axis of the cord. In the better-stained preparations the terminal arborizations of adjacent collaterals from a single axon formed a continuous longitudinal column through the dorsal horn. There was a gradual shift of the column of arborizations from lateral to medial as the more rostral collaterals were given off.6. The hair-follicle afferent fibre collaterals are now identified as the ;collaterales grosses et profondes de la substance de Rolando' of Ramón y Cajal (1909) which give rise to the ;flame-shaped arbors' of Scheibel & Scheibel (1968).7. The importance of the longitudinal organization of the terminal arborizations for an understanding of the topographical properties of dorsal horn neurones is discussed.     

Disynaptic tectal and pyramidal excitation of hindlimb motoneurons mediated by pontine reticulospinal neurons in the cat

Summary 1 The pathway mediating disynaptic tectal and pyramidal excitation of hindlimb motoneurons was analyzed in cats anesthetized with chloralose or pentobarbital. Stimulation of the contralateral tectofugal fibers induced EPSPs in flexor and extensor hindlimb motoneurons (118/171). EPSP latencies, measured from the monosynaptically evoked descending volley, were 0.8 ms or less in 34 of the 118 motoneurons, suggesting disynaptic linkage from the tectum. The latencies tended to be shorter in motoneurons innervating proximal muscles than in those innervating distal muscles. 2. Stimulation of the cerebral peduncle induced EPSPs only in a small proportion of motoneurons (7/32). But the peduncular stimulation exhibited a marked facilitatory effect on the tectal EPSPs in most of the tested motoneurons (23/27), showing convergence of tectal and peduncular inputs onto relay cells. 3. In animals whose pyramid was transected, the tectal EPSPs were still facilitated by peduncular stimulation in 45 of 48 tested motoneurons. The time course of facilitation indicated convergence of tectofugal and corticofugal fibers onto brainstem relay neurons. 4. Projection of single neurons in the nucleus reticularis pontis caudalis (NRPC) to the gray matter of the hindlimb segments was examined by mapping thresholds of antidromic activation. Twelve of 13 tested neurons were excited by contralateral tectal stimulation at short latencies, probably monosynaptically. Four of them were found to project to lamina IX. Two of the 3 tested neurons projecting to lamina IX were found to receive excitatory input from the cerebral peduncle. 5. Stimulation of NRPC induced monosynaptic EPSPs in hindlimb motoneurons. In 19 motoneurons, the NRPC-induced monosynaptic EPSPs were facilitated by a conditioning tectal shock. This indicated that the tectal stimulus lowered thresholds of direct activation of cell bodies of premotor NRPC neurons. The time course of the facilitation indicated that the NRPC neurons received monosynaptic tectal excitation. The results provide strong evidence that NRPC neurons are involved in mediating disynaptic tectal excitation of hindlimb motoneurons.     

Increased expression of phosphatase and tensin homolog in reactive astrogliosis following intracerebroventricular kainic acid injection in mouse hippocampus

New techniques were applied for maintaining viable motoneurons in rat cervical spinal cord slices to study electrical and morphological properties from postnatal day (PD) 2-49. Lucifer Yellow injections showed nine to 12, or more, viable motoneurons/slice at PD2, reduced to two to three in lamina IX by PD9. At PD2 and from PD14 onward healthy motoneurons were electrically similar to those of adults. Motoneurons exhibited variable electrical properties and morphology around PD5. They were sensitive to kainate and AMPA at all ages. The sensitivity to N-methyl-D-aspartate (NMDA) was significant at PD2, less at PD9 and virtually absent at PD14. Our observations suggest that NMDA receptors play a role in regulation of motoneuron survival in the early postnatal period, but are lost from adult motoneurons.     

Activity-dependent development of cortical axon terminations in the spinal cord and brain stem

 Corticospinal (CS) axon terminations in several species are widespread early in development but are subsequently refined into a spatially more restricted distribution. We studied the role of neural activity in sensorimotor cortex in shaping postnatal development of CS terminations in cats. We continuously infused muscimol unilaterally into sensorimotor cortex to silence neurons during the postnatal CS refinement period (weeks 3–7). Using anterograde transport of WGA-HRP, we examined the laterality of terminations from the muscimol-infused (i.e., silenced) and active sides in the spinal cord, as well as in the cuneate nucleus and red nucleus. We found that CS terminations from the muscimol-infused cortex were very sparse and limited to the contralateral side, while those from the active cortex maintained an immature bilateral topography. Controls (saline infusion, noninfusion) had dense, predominantly contralateral, CS terminations. There was a substantial decrease in the spinal gray matter area occupied by terminations from the side receiving the blockade and a concomitant increase in the area occupied by ipsilateral terminations from the active cortex. Optical density measurements of HRP reaction product from the active cortex in muscimol-infused animals showed substantial increases over controls in the ratio of ipsilateral to contralateral CS terminations for all laminae examined (IV–V, VI, VII). Our findings suggest that ipsilateral dorsal horn terminations reflect new axon growth during the refinement period because they are not present there earlier in development. Those in the ventral horn are present earlier in development and thus could reflect maintenance of transient terminations. Increased ipsilateral terminations from active cortex were due to recrossing of CS axons in lamina X and not to an increase in labeled CS axons in the ipsilateral white matter. Examination of brain stem terminations suggested that, between postnatal weeks 3 and 7, development of corticocuneate terminations also is activity-dependent but that development of corticorubral terminations is not. Activity-dependent CS development is a plausible mechanism by which early motor experiences could shape the anatomical and functional organization of the motor systems during a critical postnatal period. Received: 4 August 1998 / Accepted: 2 September 1998     

Location of the motoneurons supplying the rabbit pharyngeal constrictor muscles and the peripheral course of their axons: a study using the retrograde HRP or fluorescent labeling technique

The location of the motoneurons supplying the rabbit pharyngeal constrictor muscles (the superior and the middle constrictors, and the thyropharyngeus and the cricopharyngeus; the last two collectively compose the inferior constrictor) was investigated with intramuscular injection of HRP or the fluorescent tracer nuclear yellow into the individual muscles. Moreover, the peripheral course of their axons was investigated by injection of HRP into all of the pharyngeal constrictors in conjunction with intracranial severing of either the vagus or the glossopharyngeal nerves. The pharyngeal constrictor motoneurons were ipsilaterally located within a subdivision of the nucleus ambiguus which is formed by a compact arrangement of the smallest neurons of the nucleus and situated in the rostral half of the nucleus. We named that subdivision the compact cell group (CoG). Axons of the pharyngeal constrictor motoneurons traversed the vagal rootlets. The rostrocaudal extent of the pharyngeal constrictor motoneurons covered almost the entire length of CoG at a level from about 500 to 2,900 microns rostral to the obex, with their number being most numerous in the middle one-third level of the CoG. Although the motoneurons of the superior constrictor, those of the middle constrictor, and those of the thyropharyngeous and the cricopharyngeus overlapped considerably in location, they tended to be arranged rostrocaudally in that order. At the middle one-third level of the CoG, where the CoG is subdivided into dorsomedial and ventrolateral subgroups of neurons, the superior and the middle constrictor motoneurons were confined to the medial portion of the dorsomedial subgroup, while the inferior constrictor motoneurons were distributed throughout its entirety.