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The cat cervical dorsal root ganglia: general cell-size characteristics and comparative study of neck muscle, neck cutaneous and phrenic afferents.

Sizes of neuronal somata in the cat cervical dorsal root ganglia were determined at different levels (C1-C8). The average value and class distribution of mean cell diameter were analyzed. The ganglia from C1 to C5 could be clearly distinguished from those at levels of brachial plexus afferents (C6-C8) with respect to cell size range, distribution and average. The size distribution, most often limited to 70 microns from C1 to C5, skewed to more than 90 microns from C6 to C8. Cells in the 35-50 microns range of diameter constituted the main portion of the cell population (49-52%) at the C1-C5 levels, whereas from C6 to C8 51-77% of the ganglion cell bodies were more than 50 microns in diameter. The cell size distribution of afferents projecting from C1 to C5 and supplying different muscle or cutaneous targets was studied following retrograde labeling with horseradish peroxidase conjugated to wheatgerm agglutinin. Sizes of cell bodies of biventer cervicis (postural muscle), phrenic (purely respiratory muscle) and cutaneous afferents were all similar. The labelled cell bodies were in the majority (51-64%) less than 35 microns in diameter and ranged towards smaller diameters than counterstained cells in the corresponding ganglia. In spite of similarities in cell size distribution it was estimated from the fiber caliber spectra of the labelled afferents that both unmyelinated and myelinated cutaneous afferents originate from larger cell bodies than muscle afferents in the same diameter range.

Adolescent

Structure of the intraspinal projections of single, identified muscle spindle afferents from neck muscles of the cat.

The morphology and frequency of collaterals originating from single afferents supplying primary endings of muscle spindles in dorsal neck muscles have been examined using intra-axonal injections of HRP. Within the segment in which the afferent entered the spinal cord, one collateral was found for every 3.3 mm of stained axon. In contrast, afferents--one of more segments rostral to the segment in which they entered the spinal cord--had fewer collaterals: One collateral was found for every 6.3 mm of stained axon. The branching structure and terminal distribution of the collaterals were generally similar regardless of the muscle from which the afferent originated and the segment in which the collateral was found. Boutons were found in 2 zones: One of these was located in the intermediate zone, within and around the central cervical nucleus, and the other was found in laminae VIII and IX, including the motoneuron nuclei. The ventral termination zone of collaterals in the same segment as their parent axon entered the spinal cord was larger and had more boutons than the same projection of collaterals whose parent axon entered the spinal cord 1 or 2 segments caudal to the segment in which the collateral was found. These results indicate that afferents supplying primary endings of neck muscle spindles are more likely to contact neurons in the same segment in which the afferent enters the spinal cord than in more rostral segments. However, even within the same segment in which the afferent enters the spinal cord, the projection of neck muscle afferents to the ventral horn is less dense than the corresponding projection of hindlimb muscle spindle afferents in the lumbosacral spinal cord.

Animals

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.

Adaptation, Physiological

EMG activities of neck muscles underlying lateral flexion of the neck during head-turning induced by electrical stimulation of the caudate nucleus in cats.

Patterns of EMG activities of neck muscles underlying the initiation of head-turning, induced by stimulation of the caudate nucleus, were analyzed with special reference to temporal relations between the onset of head-turning and that of changes in EMG activities. These patterns were compared with those associated with the initiation of lateral flexion of the neck which occurred without electrical stimulation of the caudate nucleus in order to examine whether the caudate-induced head-turning was initiated via the same muscular system as that used in non-caudate-induced head movements. Experiments were carried out using 5 awake, unrestrained cats which were trained to stand still with one limb on each of 4 footplates. Trains of stimulating current pulses were applied to several stimulation points in the caudate nucleus while the animal maintained a stable standing posture with its neck extended. Head movements in the horizontal plane and EMGs of 6 neck muscles (splenius, longissimus cervicis, obliquus capitis caudalis, biventer cervicis, complexus and cervical multifidus) were recorded. Patterns of EMG activities of neck muscles around the onset of the caudate-induced head-turning were characterized by an increase in activity of the splenius, the longissimus cervicis and the obliquus capitis caudalis muscles, and by a decrease in activity of the complexus, the biventer cervicis and the cervical multifidus on the side of flexion. It is suggested that an increase in activity of the splenius, the longissimus cervicis and the obliques capitis caudalis muscles was responsible for the initiation of this evoked response. In non-caudate-induced lateral flexion of the neck, patterns of activities of neck muscles were similar to those in caudate-induced head-turning. It is therefore concluded that the caudate-induced head-turning as an evoked behavioral response was initiated through a muscular system similar to that utilized for similar head movements occurring without electrical stimulation of the caudate nucleus, although the pathways involved are thought to be different.

Animals

Projections of extraocular, neck muscle, and retinal afferents to superior colliculus in the cat: their connections to cells of origin of tectospinal tract.

Unit recordings were made in the superior colliculus of cats anesthetized with chloralose and with Pentothal. Electrical stimulation of extraocular muscle afferents and neck muscle afferents excited more units in the superior colliculus than did a variety of moving and stationary visual stimuli. Units responding to neck muscle afferent stimulation fell into three populations; one population firing with a short latency and following stimulus presentation up to 1/s, a second population with a long latency and following stimulus presentation at frequencies lower than 15/min, and a third population exhibiting paired firing. The latencies and firing patterns of the third population combined the characteristics of each of the first two patterns. It is suggested that these characteristics of unit discharges stem from the existence of two pathways from neck muscle afferents to the superior colliculus. The projection is predominantly bilateral. Units responding to neck muscle afferent stimulation are distributed throughout the superior colliculus on the basis of their latencies. Long-latency responses predominate in the superficial layers of the superior colliculus and short-latency responses, while more common in the intermediate and deep layers, predominate in the tegmentum. Extraocular muscle afferent projections to the superior colliculus constitute the single richest projection found in these experiments. While the response patterns and latencies are similar to those of the neck muscle afferents, long-latency responses are the most common and dominate in all collicular regions. Few units in the tegmentum could be excited by extraocular muscle afferents. Both extraocular muscle and neck muscle afferents show considerable convergence with one another and with retinal afferents within the superior colliculus. Cells of origin of the tectospinal tract were identified within the superior colliculus and tegmentum by antidromic excitation from the upper cervical cord. These cells were distributed predominantly within the intermediate and deep layers of the superior colliculus, and sparsely in the superficial layers and tegmentum. Almost 50% of the cells of origin of the tectospinal tract receive a convergent input from extraocular muscle and neck muscle afferents and from the retina. About 30% of the cells were inexcitable to the stimuli employed in these experiments. The significance of these projections is discussed with respect to superior collicular function in the cat and i

Anesthesia, General

[Muscle contraction headache and posture--with special reference to ischemic contraction of the posterior neck muscles].

Cause of muscle contraction headache (MCH) results from sustained contraction of scalp and posterior neck muscles. Recently, we published an effect of posture on the etiology of MCH. According to our data, head bending posture seems to be one of the main causes of sustained contraction of the posterior neck muscles. Wolff presented a hypothesis of ischemic contraction of these muscles as a cause of pain. However, about the blood flow of scalp or posterior neck muscles, only two reports were published so far. These two papers failed to demonstrate a reduction of blood flow in MCH patients, and neglected the effect of posture. The purpose of this report is to examine a change of blood flow of posterior neck muscles with the change of posture. A total of 40 patients with MCH were studied using laser doppler blood flow meter. Needle shaped probe with a diameter of 0.55 mm was inserted 15 mm into the posterior neck muscle. The angle between orbito-meatal line and horizontal plane were measured using a light helmet with goniometer. Surface EMG of the posterior neck muscles was recorded at the same time. In the case of controls who do not experience headache, the amplitude of EMG increases slightly with the bending posture (40 microV with OM line 20 degrees upward from the horizontal plane, 46 microV with OM line horizontal, and 52 microV when 30 degrees downward). In reverse to the increase of the EMG activity, blood flow of the neck muscles decreases (12 ml/100 g/min with OM line 20 degrees upward, 10.8 with OM line horizontal, 7.6 at 10 degrees down, 4.6 at 20 degrees down, and 4.1 at 30 degrees down).(ABSTRACT TRUNCATED AT 250 WORDS)

Adult

Compartmentalization of motor units in the cat neck muscle, biventer cervicis.

1. The neck muscle biventer cervicis is supplied by five separate nerve bundles that originate from segments C2-C5 and enter the muscle at different rostrocaudal levels. We have used the glycogen-depletion method to investigate the distribution of muscle fibers supplied by each nerve bundle and also the extent of motor-unit territories supplied by single motoneurons in the C3 segment. 2. Prolonged intermittent stimulation of each nerve bundle produced glycogen depletion in a compartment of muscle fibers that ran only a fraction of the whole-muscle length. The depleted compartment was separated by tendinous inscriptions from adjacent, serially arranged compartments that were supplied by different nerve bundles. Thus the muscle was divided into five in-series compartments, arranged in the same rostrocaudal sequence as the nerves by which they were supplied. 3. Six fast, glycolytic (FG) and five fast, oxidative-glycolytic (FOG) motor units were depleted by repetitive intracellular stimulation of their antidromically identified motoneurons in the C3 segment. The fibers of each motor unit were confined to a striplike subvolume whose cross-sectional area was only 20-40% of that for the whole compartment in which it was located. Single motor units contained an average of 408 extrafusal fibers (range: 262-582 fibers), and these were distributed with an average density of 20 fibers/mm2 in cross sections through their motor domains. No significant differences were found between the numbers or densities of fibers in FG and FOG motor units. 4. The specialized in-series organization of compartments has functional implications because the forces generated by one compartment of motor units must be transmitted through other in-series compartments of muscle fibers rather than directly onto skeletal attachments. The confined distribution of muscle fibers belonging to a single motor unit suggests that an additional level of organization may exist within individual compartments. The implications of these features for the physiological behavior and neural control of biventer cervicis are discussed.

Animals

Postural effects of neck muscle vibration in man.

Vibration of the dorsal neck muscles in man induces falling reactions, walking and "marche en étoile", probably by muscle afferent activation. It can be used as a reproducible error signal in analyzing the interaction between neck muscle proprioception and vestibular as well as ocular motor systems. These interactions are important for posture and coordinated head-eye movements.

Afferent Pathways

Effect of neck posture on the activation of feline neck muscles during voluntary head turns.

1. To determine whether neck posture affects the usage of neck muscles during a specific motor task, we recorded the electromyographic (EMG) patterns of neck muscles in four cats, which made horizontal, head-turning movements to fixate eccentrically placed targets. In some trials, the cervical column was oriented vertically whereas in other trials, the cervical column was oriented more horizontally. 2. During horizontal head movements, five muscles (obliquus capitis inferior, splenius, levator scapulae, complexus, and biventer cervicis) displayed activation patterns that were consistent from cat to cat and did not change when the cats adopted a different neck posture. Most of these muscles are large, superficial muscles that attach to the skull and span many cervical joints. 3. Posturally dependent patterns of activation were observed in five other neck muscles (semispinalis cervicis, longissimus capitis, levator scapulae ventralis, scalenus anterior, and obliquus capitis superior). Most of these muscles lie deeper and more laterally within the neck musculature and generally span fewer cervical joints than the muscles that displayed invariant patterns of activation. 4. These results suggest that the set of invariantly activated muscles may compose part of a basic motor program that is triggered during head movements in the horizontal plane. This motor program appears to be modified by the selective activation of ancillary muscles, which are recruited in a manner related to the neck posture. The deep positioning of the ancillary muscles may permit them to regulate the mobility of the cervical column and to adjust the net muscular force applied across the neck to the skull. Organizing the motor output in this manner might simplify the task of computing the appropriate patterns of neck-muscle activation.

Animals

[A method for puncturing the deep neck muscles in humans].

The authors developed a method to puncture neck muscles by means of computer tomographic control, enabling the introduction of an electrode into the obliquus capitis inferior muscle, to investigate the particular role of neck muscles in posture and head movement.

Electromyography