Injections of WGA-HRP were made in the rat trigeminal ganglion and C1-3 dorsal root ganglia (DRGs) to study the central projection patterns and their relations to each other. Trigeminal ganglion injections resulted in heavy terminal labeling in all trigeminal sensory nuclei. Prominent labeling was also observed in the solitary tract nucleus and in the medial parts of the dorsal horn at C1-3 levels, but labeling could be followed caudally to the C7 segment. Contralateral trigeminal projections were found in the nucleus caudalis and in the dorsal horn at C1-3 levels. The C1 DRG was found to be inconstant in the rat. When it was present, small amounts of terminal labeling were found in the external cuneate nucleus (ECN) and the central cervical nucleus (CCN). No dorsal horn projections were seen from the C1 DRG. Injections in the C2 DRG resulted in heavy labeling in the ECN, nucleus X, CCN, and dorsal horn, where it was mainly located in lateral areas. Labeling could be followed caudally to the Th 7 segment. C2 DRG projections also appeared in the cuneate nucleus (Cun), in all the trigeminal sensory nuclei, and in the spinal, medial, and lateral vestibular nuclei. A small C2 DRG projection was observed in the ventral cochlear nucleus. C3 DRG injections resulted in heavy labeling in both medial middle and lateral parts of the dorsal horn, in the ECN, and in nucleus X, whereas the labeling in the CCN was somewhat weaker. Smaller projections were seen to trigeminal nuclei, Cun, and the column of Clarke. Comparisons of the central projection fields of trigeminal and upper cervical primary afferents indicated a somatotopic organization but with a certain degree of overlap.

Central distribution of trigeminal and upper cervical primary afferents in the rat studied by anterograde transport of horseradish peroxidase conjugated to wheat germ agglutinin.

Synopsis The term sensorimotor describes all the afferent, efferent, and central integration and processing components involved in maintaining stability in the postural control system through intrinsic motor-control properties. The scope of this paper is to highlight the sensorimotor deficits that can arise from altered cervical afferent input. From a clinical orthopaedic perspective, the peripheral mechanoreceptors are the most important in functional joint stability; but in the cervical region they are also important for postural stability, as well as head and eye movement control. Consequently, conventional musculoskeletal intervention approaches may be sufficient only for patients with neck pain and minimal sensorimotor proprioceptive disturbances. Clinical experience and research indicates that significant sensorimotor cervical proprioceptive disturbances might be an important factor in the maintenance, recurrence, or progression of various symptoms in some patients with neck pain. In these cases, mo...

Sensorimotor Function and Dizziness in Neck Pain: Implications for Assessment and Management

This article sets out to present a fairly comprehensive review of our knowledge about the functions of the receptors that have been found in the extraocular muscles--the six muscles that move each eye of vertebrates in its orbit--of all the animals in which they have been sought, including Man. Since their discovery at the beginning of the 20th century these receptors have, at various times, been credited with important roles in the control of eye movement and the construction of extrapersonal space and have also been denied any function whatsoever. Experiments intended to study the actions of eye muscle receptors and, even more so, opinions (and indeed polemic) derived from these observations have been influenced by the changing fashions and beliefs about the more general question of how limb position and movement is detected by the brain and which signals contribute to those aspects of this that are perceived (kinaesthesis). But the conclusions drawn from studies on the eye have also influenced beliefs about the mechanisms of kinaesthesis and, arguably, this influence has been even larger than that in the converse direction. Experimental evidence accumulated over rather more than a century is set out and discussed. It supports the view that, at the beginning of the 21st century, there are excellent grounds for believing that the receptors in the extraocular muscles are indeed proprioceptors, that is to say that the signals that they send into the brain are used to provide information about the position and movement of the eye in the orbit. It seems that this information is important in the control of eye movements of at least some types, and in the determination by the brain of the direction of gaze and the relationship of the organism to its environment. In addition, signals from these receptors in the eye muscles are seen to be necessary for the development of normal mechanisms of visual analysis in the mammalian visual cortex and for both the development and maintenance of normal visuomotor behaviour. Man is among those vertebrates to whose brains eye muscle proprioceptive signals provide information apparently used in normal sensorimotor functions; these include various aspects of perception, and of the control of eye movement. It is possible that abnormalities of the eye muscle proprioceptors and their signals may play a part in the genesis of some types of human squint (strabismus); conversely studies of patients with squint in the course of their surgical or pharmacological treatment have yielded much interesting evidence about the central actions of the proprioceptive signals from the extraocular muscles. The results of experiments on the eye have played a large part in the historical controversy, now in at least its third century, about the origin of signals that inform the brain about movement of parts of the body. Some of these results, and more of the interpretations of them, now need to be critically re-examined. The re-examination in the light of recent experiments that is presented here does not support many of the conclusions confidently drawn in the past and leads to both new insights and fresh questions about the roles of information from motor signals flowing out of the brain and that from signals from the peripheral receptors flowing into it. There remain many lacunae in our knowledge and filling some of these will, it is contended, be essential to advance our understanding further. It is argued that such understanding of eye muscle proprioception is a necessary part of the understanding of the physiology and pathophysiology of eye movement control and that it is also essential to an account of how organisms, including Man, build and maintain knowledge of their relationship to the external visual world. The eye would seem to provide a uniquely favourable system in which to study the way in which information derived within the brain about motor actions may interact with signals flowing in from peripheral receptors. The review is constructed in relatively independent sections that deal with particular topics. It ends with a fairly brief piece in which the author sets out some personal views about what has been achieved recently and what most immediately needs to be done. It also suggests some lines of study that appear to the author to be important for the future.

The functions of the proprioceptors of the eye muscles.

Forty-six patients suffering from autosomal dominant cerebellar ataxia type I (ADCA I) underwent to a genotype-phenotype correlation analysis by molecular genetic assignment to the spinocerebellar ataxia type 1, 2, or 3 (SCA1, SCA2, SCA3) genetic locus and electro-oculography. Oculomotor deficits that are attributed to dysfunction of cerebellar structures occurred in all three mutations without major differences between the groups. Gaze-evoked nystagmus, however, was not found to be associated with SCA2. Square wave jerks were exclusively observed in SCA3. The gain in vestibulo-ocular reflex was significantly impaired in SCA3 and SCA1. In SCA3 the severity of vestibular impairment increased with CAG repeat length. Severe saccade slowing was a highly characteristic feature of SCA2. In SCA3 saccade velocity was normal to mildly reduced while SCA1 fell into an intermediate range. The present data show that each mutation is associated with a distinct syndrome of oculomotor deficits. Reduced saccade velocity and the absence of both square-wave jerks and gaze-evoked nystagmus allow one SCA2 to be distinguished from SCA3 patients in almost all cases. The eye movement disorder of SCA1 patients, however, overlaps with both SCA2 and SCA3.

Autosomal dominant cerebellar ataxia type I: oculomotor abnormalities in families with SCA1, SCA2, and SCA3.

The role of ocular muscle proprioception in the localization of visual targets has been investigated in normal humans by deviating one eye to create an experimental strabismus. The passively deviated eye was covered and the other eye viewed the target. With a hand-pointing task, targets were systematically mislocalized in the direction of the deviated nonviewing eye. A 4- to 6-degree error resulted when the nonviewing eye was offset 30 degrees from straight ahead. When the eye was deviated, the perceived "straight-ahead" was also displaced, by a similar amount, in the same direction. Since the efferent motor commands to the displaced and to the nondisplaced eyes are presumably identical by the law of equal innervation, the mislocalization of visual objects must be attributed to the change in proprioceptive information issued from the nonviewing, deviated eye. Thus proprioception contributes to the localization of objects in space.

https://science.sciencemag.org/content/sci/249/4964/58.full.pdf

The role of ocular muscle proprioception in visual localization of targets

Brainstem projections of first-order afferent neurons that innervate the suboccipital muscles of the monkey have been determined by using the technique of transganglionic transport of wheat-germ-agglutinin-conjugated horseradish peroxidase (WGA/HRP) and HRP. Neck muscle afferents distribute to several distinct regions located within the caudal brainstem and rostral spinal cord. Terminal labeling was heaviest within the ventral portion of the ipsilateral lateral cuneate nucleus. Muscle afferent terminals also distributed to ventrolateral portions of the pars triangularis division of the cuneate nucleus. Projections were consistent with the known somatotopic (i.e., both place and modality) organization of the cuneate nucleus. Moreover, neck muscle projections to the cuneate nucleus were, in part, coincident with those previously demonstrated for the extraocular muscles (Porter: J. Comp. Neurol. 247:133-143, '86). Sparse terminal projections were noted in the central cervical nucleus. In addition, light terminal labeling was present in group x of the vestibular complex and in an ill-defined region along the lateral margin of the brainstem. Present observations, which provide the first complete description of the central distribution of neck muscle afferent neurons in the primate, may contribute to the known substrate for eye/head coordination.

Daniel P. Edney

Papers

Neck muscle afferent projections to the brainstem of the monkey: implications for the neural control of gaze.