Cell Types Promoting Goosebumps Form a Niche to Regulate Hair Follicle Stem Cells.

Piloerection (goosebumps) requires concerted actions of the hair follicle, the arrector pili muscle (APM), and the sympathetic nerve, providing a model to study interactions across epithelium, mesenchyme, and nerves. Here, we show that APMs and sympathetic nerves form a dual-component niche to modulate hair follicle stem cell (HFSC) activity. Sympathetic nerves form synapse-like structures with HFSCs and regulate HFSCs through norepinephrine, whereas APMs maintain sympathetic innervation to HFSCs. Without norepinephrine signaling, HFSCs enter deep quiescence by down-regulating the cell cycle and metabolism while up-regulating quiescence regulators Foxp1 and Fgf18. During development, HFSC progeny secretes Sonic Hedgehog (SHH) to direct the formation of this APM-sympathetic nerve niche, which in turn controls hair follicle regeneration in adults. Our results reveal a reciprocal interdependence between a regenerative tissue and its niche at different stages and demonstrate sympathetic nerves can modulate stem cells through synapse-like connections and neurotransmitters to couple tissue production with demands.

The unique axon trajectory of the accessory nerve is determined by intrinsic properties of the neural tube in the avian embryo.

The accessory nerve is a cranial nerve, composed of only motor axons, which control neck muscles. Its axons ascend many segments along the lateral surface of the cervical spinal cord and hindbrain. At the level of the first somite, they pass ventrally through the somitic mesoderm into the periphery. The factors governing the unique root trajectory are unknown. Ablation experiments at the accessory nerve outlet points have shown that somites do not regulate the trajectory of the accessory nerve fibres. Factors from the neural tube that may control the longitudinal pathfinding of the accessory nerve fibres were tested by heterotopic transplantations of an occipital neural tube to the cervical and thoracic level. These transplantations resulted in a typical accessory nerve trajectory in the cervical and thoracic spinal cord. In contrast, cervical neural tube grafts were unable to give rise to the typical accessory nerve root pattern when transplanted to occipital level. Our results show that the formation of the unique axon root pattern of the accessory nerve is an intrinsic property of the neural tube.

Distribution of Neuron Cell Bodies in the Intraspinal Portion of the Spinal Accessory Nerve in Humans.

The spinal accessory nerve is often identified as a purely motor nerve innervating the trapezius and sternocleidomastoid muscles. Although it may contain proprioceptive neurons found in cervical spinal levels C2-C4, limited research has focused on the histology of the spinal accessory nerve. The objective of the present study was to examine the spinal accessory nerve to determine if there are neuronal cell bodies within the spinal accessory nerve in humans. Cervical spinal cords were dissected from eight cadavers that had previously been used for dissection in other body regions. The segmental rootlets were removed to quantify the neuron cell bodies present at each spinal level. Samples were embedded in paraffin; sectioned; stained with hematoxylin and eosin; and examined using a microscope at 4×, 10×, and 40× magnification. Digital photography was used to image the samples. Neuronal cell bodies were found in 100% of the specimens examined, with non-grossly visible ganglia found at spinal levels C1-C4. The C1 spinal level of the spinal accessory nerve had the highest number of neuron cell bodies.

Histologic confirmation of neuronal cell bodies along the spinal accessory nerve.

INTRODUCTION: Most sources conclude that the spinal accessory nerve (SAN) is a purely motor nerve. There are some reports that suggest a sensory component, although the exact nature of such sensory fibers has yet to be elucidated. With such discrepancies in the literature and with well-established pain syndromes of unknown etiology following SAN injury, the authors performed the present study to better clarify this anatomy. MATERIALS AND METHODS: The entire accessory nerve was harvested from 10 adult cadavers. Samples were then submitted for immunohistochemical analyses. RESULTS: Occasional microganglia cells were identified along the SAN in all specimens. These ganglia were most numerous along the intracranial segment of the SAN, but none was found along the cranial rootlets of the accessory nerve. CONCLUSIONS: Neuronal cell bodies were identified along the course of the SAN in human cadavers. Although the function is not certain, such cells have been found in other animals to be nocioceptive in nature. Pending further study, these cells may be found to be involved in enigmatic pain syndromes thought to arise in the sternocleidomastoid and trapezius muscles.

Axotomy alters alternative splicing of choline acetyltransferase in the rat dorsal motor nucleus of the vagus nerve.

Choline acetyltransferase of the peripheral type (pChAT) is a splice variant that lacks exons 6-9 of the common-type ChAT (cChAT); the role of pChAT remains unknown. We investigated the expression of pChAT and cChAT after axotomy to try to elucidate its function. In the dorsal motor nucleus of the vagus nerve (DMNV), nucleus ambiguus (NA), and hypoglossal nucleus (HN) of control rats, we observed neural expression of cChAT but no pChAT-positive neurons. Following nerve transection, we clearly detected pChAT-labeled neurons in the DMNV and weakly labeled neurons in the NA, but pChAT was not seen in the HN. In the DMNV, the mean number of cChAT-positive neurons decreased rapidly to 40.5% of control at 3 days post transection, and to 5.0% of control after 7 days. The number of cChAT-positive neurons then gradually increased and reached a plateau of about 25% of control value at 28 days post transection. pChAT-positive neurons did not appear until 7 days after transection. On the same day, pChAT mRNA was detected in the DMNV neurons by reverse transcription-polymerase chain reaction (RT-PCR) by using laser capture microdissection. The number of pChAT-positive neurons gradually decreased, and only 10% of the cholinergic neurons retained pChAT expression 56 days post transection. Double-immunofluorescence analysis showed that some of the DMNV neurons expressed both cChAT and pChAT upon recovery from axotomy. These results suggest that the expression of pChAT is associated with the regenerative or degenerative processes of motoneurons especially for general visceral efferents.

UNC5C is required for spinal accessory motor neuron development.

In both invertebrates and vertebrates, UNC5 receptors facilitate chemorepulsion away from a Netrin source. Unlike most motor neurons in the embryonic vertebrate spinal cord, spinal accessory motor neuron (SACMN) cell bodies and their axons translocate along a dorsally directed trajectory away from the floor plate/ventral midline and toward the lateral exit point (LEP). We have recently shown that Netrin-1 and DCC are required for the migration of SACMN cell bodies, in vivo. These observations raised the possibility that vertebrate UNC5 proteins mediate the presumed repulsion of SACMN away from the Netrin-rich ventral midline. Here, we show that SACMN are likely to express UNC5A and UNC5C. Whereas SACMN development proceeds normally in UNC5A null mice, many SACMN cell bodies fail to migrate away from the ventral midline and inappropriately cluster in the ventrolateral spinal cord of mouse embryos lacking UNC5C. These results support an important role for UNC5C in SACMN development.

Localization of the spinal nucleus of accessory nerve in rat: a horseradish peroxidase study.

The spinal nucleus of the accessory nerve (SNA) comprises the group of somata (perikarya) of motor neurons that supply the sternocleidomastoid and trapezius muscles. There are many conflicting views regarding the longitudinal extent and topography of the SNA, even in the same species, and these disagreements prompted the present investigation. Thirty Sprague-Dawley rats (15 males, 15 females) were used. The SNA was localized by retrograde axonal transport of horseradish peroxidase. Longitudinally, the SNA was found to be located in the caudal part (caudal 0.9-1.2 mm) of the medulla oblongata, the whole lengths of cervical spinal cord segments C1, C2, C3, C4, C5 and rostral fourth of C6. In the caudal part of the medulla oblongata, the SNA was represented by a group of perikarya of motor neurons lying immediately ventrolateral to the pyramidal fibres that were passing dorsolaterally after their decussation. In the spinal cord, the motor neuronal somata of the SNA were located in the dorsomedial and central columns at C1, in the dorsomedial, central and ventrolateral columns at C2 and in the ventrolateral column only at C3, C4, C5 and rostral quarter of C6. The perikarya of motor neurons supplying the sternocleidomastoid were located in the caudal part (caudal 0.9-1.2 mm) of the medulla oblongata ventrolateral to the pyramidal fibres that were passing dorsolaterally after their decussation. They were also located in the dorsomedial and central columns at C1, in the dorsomedial, central and ventrolateral columns at C2 and only in the ventrolateral column at the rostral three-quarters of C3. The perikarya of motor neurons supplying the trapezius muscle were located in the ventrolateral column only in the caudal three-quarters of C2, the whole lengths of C3, C4 and C5, and in the rostral quarter of C6.

Forced expression of Phox2 homeodomain transcription factors induces a branchio-visceromotor axonal phenotype.

What causes motor neurons to project into the periphery is not well understood. We here show that forced expression of the homeodomain protein Phox2b, shown previously to be necessary and sufficient for branchio-visceromotor neuron development, and of its paralogue Phox2a imposes a branchiomotor-like axonal phenotype in the spinal cord. Many Phox2-transfected neurons, whose axons would normally stay within the confines of the neural tube, now project into the periphery. Once outside the neural tube, a fraction of the ectopic axons join the spinal accessory nerve, a branchiomotor nerve which, as shown here, does not develop in the absence of Phox2b. Explant studies show that the axons of Phox2-transfected neurons need attractive cues to leave the neural tube and that their outgrowth is promoted by tissues, to which branchio-visceromotor fibers normally grow. Hence, Phox2 expression is a key step in determining the peripheral axonal phenotype and thus the decision to stay within the neural tube or to project out of it.

Molecular control of spinal accessory motor neuron/axon development in the mouse spinal cord.

Within the developing vertebrate spinal cord, motor neuron subtypes are distinguished by the settling positions of their cell bodies, patterns of gene expression, and the paths their axons follow to exit the CNS. The inclusive set of cues required to guide a given motor axon subtype from cell body to target has yet to be identified, in any species. This is attributable, in part, to the unavailability of markers that demarcate the complete trajectory followed by a specific class of spinal motor axons. Most spinal motor neurons extend axons out of the CNS through ventral exit points. In contrast, spinal accessory motor neurons (SACMNs) project dorsally directed axons through lateral exit points (LEPs), and these axons assemble into the spinal accessory nerve (SAN). Here we show that an antibody against BEN/ALCAM/SC1/DM-GRASP/MuSC selectively labels mouse SACMNs and can be used to trace the pathfinding of SACMN axons. We use this marker, together with a battery of transcription factor-deficient or guidance cue/receptor-deficient mice to identify molecules required for distinct stages of SACMN development. Specifically, we find that Gli2 is required for the initial extension of axons from SACMN cell bodies, and that netrin-1 and its receptor Dcc are required for the proper dorsal migration of these cells and the dorsally directed extension of SACMN axons toward the LEPs. Furthermore, in the absence of the transcription factor Nkx2.9, SACMN axons fail to exit the CNS. Together, these findings suggest molecular mechanisms that are likely to regulate key steps in SACMN development.

Identification and characterization of a cell surface marker for embryonic rat spinal accessory motor neurons.

The developing mammalian spinal cord contains distinct populations of motor neurons that can be distinguished by their cell body positions, by the expression of specific combinations of regulatory genes, and by the paths that their axons take to exit the central nervous system (CNS). Subclasses of spinal motor neurons are also thought to express specific cell surface proteins that function as receptors which control the guidance of their axons. We identified monoclonal antibody (mAb) SAC1 in a screen aimed at generating markers for specific subsets of neurons/axons in the developing rat spinal cord. During early embryogenesis, mAb SAC1 selectively labels a small subset of Isl1-positive motor neurons located exclusively within cervical segments of the spinal cord. Strikingly, these neurons extend mAb SAC1-positive axons along a dorsally directed trajectory toward the lateral exit points. Consistent with the finding that mAb SAC1 also labels spinal accessory nerves, these observations identify mAb SAC1 as a specific marker of spinal accessory motor neurons/axons. During later stages of embryogenesis, mAb SAC1 is transiently expressed on both dorsally and ventrally projecting spinal motor neurons/axons. Interestingly, mAb SAC1 also labels the notochord and floor plate during most stages of spinal cord development. The mAb SAC1 antigen is a 100-kD glycoprotein that is likely to be the rat homolog of SC1/BEN/DM-GRASP, a homophilic adhesion molecule that mediates axon outgrowth and fasciculation.

Location of the spinal nucleus of the accessory nerve in the human spinal cord.

The segmental extent and topography of the spinal nucleus of the accessory nerve (SNAN) was investigated in the adult human spinal cord. Transverse sections of segments between the lower medulla and C6 were stained with cresyl violet and the motor cell columns identified according to the numerical locations defined by Elliott (1942). The segmental extent and topography of the cervical part of column 2 resembled that previously described for the SNAN of primates.

On the presence of ganglion cells in the intracranial portion of the accessory nerve (XI cranial nerve) in some mammals.

The intracranial tract of the accessory nerve (XI cranial nerve) was studied in some mammals (equines, domestic and wild ruminants, pig, carnivores, rabbit, nutria, guinea pig, hamster, hedgehog). The specimens were embedded in paraffin or paraplast, the sections were stained with cresyl violet, haematoxylin and eosin, or submitted to argentic impregnation. Pseudounipolar ganglion cells were found in all the mammals examined, with the exception of the cat. The number of cells and their variability in the different species and subjects were related. The topography and morphology of the cells were described. This comparative study has demonstrated that the accessory nerve is not a entirely motor nerve, but it is a mixed, motor and sensitive, nerve. Nevertheless, we think further studies are necessary in order to establish the peripheral distribution, the central pathway and the functional role of the pseudounipolar neurons found in the intracranial tract of the accessory nerve.

Functional redundancy and gustatory development in bdnf null mutant mice.

In the mouse nasopalate papilla and in the trenches of the foliate and vallate papillae, taste buds accumulated primarily during the first 2 weeks after birth. Null mutation for brain-derived neurotrophic factor caused extensive death of embryonic taste neurons, with the secondary outcome that most taste buds failed to form. However not all taste neurons died; functional redundancy rescued a variable number. The primary research objective was to identify the likely site of the taste neuron rescue factor that substituted for BDNF. In this quest taste bud abundance served as a useful gauge of taste neuron abundance. The proportion of taste buds that developed was variable and uncorrelated among the nasopalate, vallate, and foliate gustatory papillae within each bdnf null mutant mouse. Thus, in spite of shared IXth nerve innervation, the vallate and foliate papillae independently varied in residual gustatory innervation. This variation rules against the rescue of gustatory neurons by system-wide factors or by factors acting on the IXth ganglion or nerve trunk. Therefore it is likely that surviving BDNF-deprived taste neurons were stochastically rescued by a redundant neurotrophic factor at the level of the local gustatory epithelium. These findings broaden the classic expectation that target tissue supplies only a single neurotrophic factor that can sustain sensory (taste) neurons.

The efferent system of cranial nerve nuclei: a comparative neuromorphological study.

A number of inconsistencies and controversies are inherent in the classification of cranial nerve nuclei based on the concepts of the various head-theories. The assumption of head segmentation, which is common to these theories, serves as the basis for designating the dorsomedial nuclei as the somatomotor column, although they innervate striated muscles of a viscus and a specific sense organ. The ventrolateral nuclei are called the specific visceromotor column; they innervate striated muscles in the branchiogenous area, but many of these muscles insert on skeletal elements. A series of comparative neuromorphological studies investigating the dendritic arborization pattern and axonal trajectory in the frog, lizard, and rat suggests a much more delicate classification in which nine morphologically and functionally different neuron groups can be discerned: 1. The hypoglossal nucleus appears coincidentally with the muscular tongue in amphibia. The spindle-shaped perikaryon, the bipolar dendritic arborization, and the straight ventral trajectory of the axon are characteristic morphological features in all three animal species investigated. 2. The oculomotor, trochlear, and abducens nuclei present a remarkably conservative topography and organization in all vertebrates with a moving eye. With their oval-shaped or polygonal perikarya and radiating dendritic arborization, these neurons distinctly differ from hypoglossal neurons. The ipsilateral axons follow a straight ventral course, the contralateral axons form a dorsal loop before crossing the midline, and the crossing is not consequence of neuron migration to the contralateral side. 3. The accessory abducens nucleus is present in tetrapods except apes and human. The elongated perikaryon and the dorsoventral dendritic orientation distinctly distinguish these neurons from other cranial motoneurons, the nucleus is found in the lateral part of the reticular formation. The neurons differentiate in situ, they do not migrate from the main abducens nucleus. 4. In the submammalian trigeminal and facial nuclei, two basic neuron types can be distinguished on the basis of their morphology. The first type is larger and accumulates in the rostral part of the trigeminal nucleus. This type innervates the jaw closer muscles. The second type is found in the caudal part of the trigeminal nucleus and in the facial nucleus. These neurons innervate the muscular floor of the mouth and the facial contingent supplies the jaw opener muscle. A very characteristic feature in the axonal trajectory is an initial medial course and a hairpin turn, or dorsal loop, at the lateral aspect of the medial longitudinal fasciculus. In addition to the two types of neurons, there is a third type in the frog trigeminal nucleus. This innervates an orbital muscle.(ABSTRACT TRUNCATED AT 400 WORDS)

Prenatal development of the human nucleus ambiguus during the embryonic and early fetal periods.

The ontogenetic development of the nucleus ambiguus was studied in a series of human embryos and fetuses ranging from 3 to 12.5 weeks of menstrual age (4 to 66 mm crown-rump length). They were prepared by Nissl and silver methods. Nucleus ambiguus neuroblasts, whose neurites extend towards and into the IXth and rostral Xth nerve roots, appear in the medial motor column of 4-6-week-old embryos (4.25-11 mm). These cells then migrate laterally (6.5 weeks, 14 mm) to a position near the dorsal motor nucleus of X. At 7 weeks (15 mm), nucleus ambiguus cells begin their migration, which progresses rostrocaudally, into their definitive ventrolateral position. The basic pattern of organization of the nucleus is established in its rostral region at 8 weeks (22.2-24 mm) and extends into its caudal region by 9 weeks (32 mm), when its nearly adult organization is evident. Cells having the characteristics of mature neurons first appear rostrally in the nucleus during the 8.5-9-week period (24.5-32 mm), gradually increase in number, and constitute the entire nucleus at 12.5 weeks (65.5 mm). Definitive neuronal subgroups first appear at 10 weeks (37.5 mm) in the large rostral nuclear region. These features suggest that the human nucleus ambiguus develops along a rostrocaudal temporospatial gradient. Evidence indicates that function of nucleus ambiguus neurons, manifested by fetal reflex swallowing, occurs after the cells migrate into their definitive position, establish the definitive nuclear pattern, and exhibit mature characteristics.

Neurons of origin of the internal ramus of the rabbit accessory nerve: localization in the dorsal nucleus of the vagus nerve and the nucleus retroambigualis.

The neurons of origin of the internal ramus of the rabbit accessory nerve were identified in the dorsal nucleus of the vagus nerve, using bilateral injections of horseradish peroxidase into the inferior vagal ganglion, soft palate, and pharynx, which were preceded by different combinations of the unilateral intracranial severings of the rootlets of the vagus and glossopharyngeal nerves, those of the cranial root of the accessory nerve, and the trunk of its spinal root. The neurons of origin occupied the caudal four-fifths of the dorsal vagal nucleus extending from about 1.0 mm rostral to the obex as far caudally as the second cervical spinal segment, with their number being about half the total number of neurons of the nucleus. Although considerably fewer, they were also located in the nucleus retroambigualis of the caudal half of the first cervical spinal segment and the second segment. Axons of most internal ramus neurons traversed the rootlets of the cranial accessory root. Axons of the few neurons located more caudally than about 1.0 mm caudal to the obex emerged from the upper cervical spinal cord to run along the trunk of the spinal accessory root before finally joining the internal ramus; caudal to the midlevel of the first cervical segment, the dorsal vagal nucleus and the nucleus retroambigualis contained neurons whose axons followed only that course.

Fibers supplying the laryngeal musculature in the cranial root of the rabbit accessory nerve: nucleus of origin, peripheral course, and innervated muscles.

The location of the rabbit laryngeal motoneurons whose axons traverse the cranial root of the accessory nerve was studied with injection of HRP or nuclear yellow into the laryngeal muscles in combination with the intracranial severing of either the rootlets of the vagus nerve or those of the cranial root. The motoneurons were located in the diffuse cell group that forms a subnucleus occupying the caudal two-thirds of the nucleus ambiguus and sending fibers to the inferior laryngeal nerve. The caudal one-third of the diffuse cell group supplying the lateral cricoarytenoid muscle, was occupied only by these motoneurons, whereas in its rostral two-thirds, they were intermingled with motoneurons having axons that traversed the vagal rootlets. The thyroarytenoid and posterior cricoarytenoid motoneurons are present in the rostral two-thirds of the diffuse cell group; axons of the former traversed the rootlets of the cranial root, and of the latter traversed the vagal rootlets. On the other hand, the medial scattered cell group, located in the rostral one-third of the nucleus ambiguus and sending fibers to the cricothyroid muscle via the superior laryngeal nerve, contained only motoneurons with axons traversing the vagal rootlets. The above findings clarified that fibers of the cranial root enter the inferior laryngeal nerve after joining the vagus, and then reach the adductor muscles for the vocal fold, with their neurons of origin in a caudal portion of the nucleus ambiguus. The vagal rootlet fibers, with their neurons of origin situated more rostrally in the nucleus, innervate the tensor and abductor muscles via the superior and inferior laryngeal nerve, respectively.

Horseradish peroxidase study of the localization of motoneurons in the accessory nucleus (XI) of the Japanese toad.

It has been conventionally accepted that the anuran accessory nerve (nXI) only innervates the musculus cucullaris (C). However, we have found that the nXI of the Japanese toad innervated the musculus interscapularis (IS) in addition to the C. The motoneurons innervating these muscles were labelled by the intramuscular injection of horseradish peroxidase (HRP). In the accessory nucleus (XI), the motoneurons were somatotopically organized: the C motoneurons were localized more rostrally, while the IS motoneurons more caudally.

Cobaltic lysine study of the morphology and distribution of the cranial nerve efferent neurons (motoneurons and preganglionic parasympathetic neurons) and rostral spinal motoneurons in the Japanese toad.

The morphology and distribution of the cranial nerve motoneurons (except III, IV, and VI) and rostral spinal motoneurons were systematically studied in the Japanese toad (Bufo japonicus) by retrograde labelling with cobaltic lysine complex. The cobaltic lysine clearly labelled whole neurons, i.e., cell bodies, proximal and distal dendrites, and axons. The branchial motoneurons (V, VII, IX, and X) had similar morphological characteristics and formed a more-or-less continuous cell column through the brainstem. The dendrites could be grouped mainly into the dorsomedial and the ventrolateral dendritic arrays. The dorsomedial dendrites formed a dendritic plexus in the subependymal gray matter, which extended as far peripherally as beneath the ependymal layer. The ventrolateral dendrites formed a broom-like dendritic plexus in the lateral to ventrolateral white matter. They usually extended as far peripherally as the pial surface. The rostrocaudal extent of the dendritic field was also wide and usually exceeded the motor nuclear boundaries. The hypoglossal motoneurons were grouped into the dorsomedial and ventrolateral cell groups, and the latter was considered to be part of the rostral spinal motoneuron column, from their morphology and distribution. The former had well-differentiated dendrites and occupied a more medial position than the branchial motoneurons. Besides the equivalent of the dorsomedial and ventrolateral dendritic arrays of the branchial motoneurons, they had dorsal and commissural dendrites. The accessory motoneurons had morphological characteristics and a distribution pattern similar to those of the rostral spinal motoneurons rather than the branchial motoneurons. The rostral spinal motoneurons had morphological characteristics somewhat different from the branchial motoneurons and the hypoglossal motoneurons (dorsomedial group). Functional implications of the motoneuron morphology are discussed, mainly based on the present results and earlier anatomical and physiological studies of the spinal motoneurons. The present study also revealed the anatomical features of the preganglionic parasympathetic neurons supplying some cranial nerves. These neurons had small somata with less elaborate dendrites and formed an almost continuous cell column that occupied a more dorsal position than the motoneurons of the corresponding nerve. They are thought to be homologous to the salivatory nucleus and the dorsal motor nucleus of the vagus. The basic anatomical organization of the general visceral efferent column seems to be similar throughout vertebrates.

Quantitative analysis of cervical musculature in rats: histochemical composition and motor pool organization. I. Muscles of the spinal accessory complex.

In this paper we characterize the architecture and segmental innervation, histochemical composition, muscle spindle populations, and motor pool organization of rat spinal accessory (SA) muscles: sternomastoid (SM), cleidomastoid (CM), cleidotrapezius (CT), and acromiotrapezius (AT). We also consider whether individual rat neck muscles are supplied by more than one population of motor neurons as they are in turtles and cats and whether in SA muscles motor neuron size scales with target muscle fiber type. SM, CM, and CT are ventral, parallel strap muscles. Each can be divided into grossly visible white and red compartments. AT is a dorsolateral sheetlike muscle that shows no gross compartmentalization. All four muscles are dominated by fast-twitch glycolytic (FG) and fast oxidative glycolytic (FOG) fibers, and FG fibers are significantly more numerous than the FOG type in three out of four muscles. Thus SA muscles in rats appear to be specialized for rapid, phasic head movements. Topographical analyses revealed that there is a striking compartmentalization of fiber types in the ventral muscles that corresponds to the red and white segments seen grossly. Spindles are found only in regions containing slow-twitch oxidative (SO) fibers. Cross-muscle comparisons indicate that there are significant differences between SA muscles in their fiber type composition. The motor pools of SA muscles form a single column from lower medulla to C5. Rostral cells lie dorsomedially in the ventral horn and, at the C1/C2 junction, the column shifts ventrolaterally. Within this column, each motor pool occupies a characteristic rostrocaudal position in the order SM:CM:CT:AT. Thus SM and (in part) CM motor neurons lie more medially than cells supplying the trapezius complex, suggesting that they may be under different patterns of synaptic drive. We saw no evidence that rat SA muscles are supplied by more than one population of motor neurons. Direct comparisons between the soma sizes of motor neurons that supply muscles or parts of muscles with significantly different histochemical compositions indicate that these size differences are in the direction predicted from their histochemical profiles, thus suggesting that in these muscles motor neuron soma size may scale with muscle fiber type.