The estimated mechanical advantage of the prosimian ankle joint musculature,and implications for locomotor adaptation

In this study we compared the power arm lengths and mechanical advantages attributed to 12 lower leg muscles across three prosimian species. The origins and insertions of the lower leg muscles in Garnett's galago, the ring‐tailed lemur, and the slow loris were quantified and correlated with positional behaviour. The ankle joint of the galago has a speed‐oriented mechanical system, in contrast to that of the slow loris, which exhibits more power‐oriented mechanics. The lemur ankle joint exhibited intermediate power arm lengths and an intermediate mechanical advantage relative to the other primates. This result suggests that the mechanical differences in the ankle between the galago and the lemur, taxa that exhibit similar locomotory repertoires, reflect a difference in the kinematics and kinetics of leaping (i.e. generalised vs. specialised leapers). In contrast to leaping primates, lorises have developed a more power‐oriented mechanical system as a foot adaptation for positional behaviours such as bridging or cantilevering in their arboreal habitat.     

An allometric analysis of the number of muscle spindles in mammalian skeletal muscles

An allometric analysis of the number of muscle spindles in relation to muscle mass in mammalian (mouse, rat, guinea-pig, cat, human) skeletal muscles is presented. It is shown that the trend to increasing number as muscle mass increases follows an isometric (length) relationship between species, whereas within a species, at least for the only essentially complete sample (human), the number of spindles scales, on average, with the square root rather than the cube root of muscle mass. An attempt is made to reconcile these apparently discrepant relationships. Use of the widely accepted spindle density (number of spindles g(-1) of muscle) as a measure of relative abundance of spindles in different muscles is shown to be grossly misleading. It is replaced with the residuals of the linear regression of ln spindle number against ln muscle mass. Significant differences in relative spindle abundance as measured by residuals were found between regional groups of muscles: the greatest abundance is in axial muscles, including those concerned with head position, whereas the least is in muscles of the shoulder girdle. No differences were found between large and small muscles operating in parallel, or between antigravity and non-antigravity muscles. For proximal vs. distal muscles, spindles were significantly less abundant in the hand than the arm, but there was no difference between the foot and the leg.     

Fibre type composition in the lumbar perivertebral muscles of primates: implications for the evolution of orthogrady in hominoids

The axial musculoskeletal system is important for the static and dynamic control of the body during both locomotor and non‐locomotor behaviour. As a consequence, major evolutionary changes in the positional habits of a species are reflected by morpho‐functional adaptations of the axial system. Because of the remarkable phenotypic plasticity of muscle tissue, a close relationship exists between muscle morphology and function. One way to explore major evolutionary transitions in muscle function is therefore by comparative analysis of fibre type composition. In this study, the three‐dimensional distribution of slow and fast muscle fibres was analysed in the lumbar perivertebral muscles of two lemuriform (mouse lemur, brown lemur) and four hominoid primate species (white‐handed gibbon, orangutan, bonobo, chimpanzee) in order to develop a plausible scenario for the evolution of the contractile properties of the axial muscles in hominoids and to discern possible changes in muscle physiology that were associated with the evolution of orthogrady. Similar to all previously studied quadrupedal mammals, the lemuriform primates in this study exhibited a morpho‐functional dichotomy between deep slow contracting local stabilizer muscles and superficial fast contracting global mobilizers and stabilizers and thus retained the fibre distribution pattern typical for quadrupedal non‐primates. In contrast, the hominoid primates showed no regionalization of the fibre types, similar to previous observations in Homo. We suggest that this homogeneous fibre composition is associated with the high functional versatility of the axial musculature that was brought about by the evolution of orthograde behaviours and reflects the broad range of mechanical demands acting on the trunk in orthograde hominoids. Because orthogrady is a derived character of euhominoids, the uniform fibre type distribution is hypothesized to coincide with the evolution of orthograde behaviours.     

Structure, distribution and innervation of muscle spindles in avian fast and slow skeletal muscle

Muscle spindles in 2 synergistic avian skeletal muscles, the anterior (ALD) and posterior (PLD) latissimus dorsi, were studied by light and electron microscopy to determine whether morphological or quantitative differences existed between these sensory receptors. Differences were found in the density, distribution and location of muscle spindles in the 2 muscles. They also differed with respect to the morphology of their capsules and intracapsular components. The slow ALD possessed muscle spindles which were evenly distributed throughout the muscle, whereas in the fast PLD they were mainly concentrated around the single nerve entry point into the muscle. The muscle spindle index (number of spindles per gram wet muscle weight) in the ALD was more than double that of its fast-twitch PLD counterpart (130.5±2.0 vs 55.4±2.0 respectively, n=6). The number of intrafusal fibres per spindle ranged from 1 to 8 in the ALD and 2 to 9 in the PLD, and their diameters varied from 5.0 to 16.0 μm and 4.5 to 18.5 μm, respectively. Large diameter intrafusal fibres were more frequently encountered in spindles of the PLD. Unique to the ALD was the presence of monofibre muscle spindles (12.7% of total spindles observed in ALD) which contained a solitary intrafusal fibre. In muscle spindles of both the ALD and PLD, sensory nerve endings terminated in a spiral fashion on the intrafusal fibres in their equatorial regions. Motor innervation was restricted to either juxtaequatorial or polar regions of the intrafusal fibres. Outer capsule components were extensive in polar and juxtaequatorial regions of ALD spindles, whereas inner capsule cells of PLD spindles were more numerous in juxtaequatorial and equatorial regions. Overall, muscle spindles of the PLD exhibited greater complexity with respect to the number of intrafusal fibres per spindle, range of intrafusal fibre diameters and development of their inner capsules. It is postulated that the differences in muscle spindle density and structure observed in this study reflect the function of the muscles in which they reside.     

Topography of muscle spindles and Golgi tendon organs in shoulder muscles of "Monodelphis domestica".

The topography of muscle spindles and Golgi tendon organs in the rotator cuff and surrounding shoulder muscles of a small laboratory marsupial (monodelphis domestica) were studied using light microscopy of serial sections. The shoulder joint of monodelphis has a large degree of freedom of movement allowing this animal to use the upper extremities for a wide range of activities like climbing and manipulating food. Thus, similar to the situation in man the shoulder joint is mainly secured by muscles. Silver stained serial paraffin sections were examined under the light microscope and the distribution of muscle spindles and Golgi tendon organs was reconstructed using three-dimensional image processing. In the two animals examined 113 and 131 muscle spindles respectively were found within the 4 rotator cuff muscles. In addition, 76 and 40 Golgi tendon organs respectively were seen at the musculo-tendinous junctions of these muscles preferentially close to the insertion at the humerus head. Also the surrounding shoulder muscles contain both muscle spindles and Golgi tendon organs in large numbers, but the ratio of Golgi tendon organs per muscle spindle appears to be lower. Number and localization of muscle spindles and Golgi tendon organs suggest, that these receptors are important for both reflex control of shoulder muscle tone as well as monitoring of static position and movement in the shoulder joint.     

Comparative histochemical study of prosimian primate hindlimb muscles. II. Populations of fiber types

The populations of fiber types in hindlimb muscles of the tree shrew (Tupaia glis), lesser bushbaby (Galago senegalensis), and the slow loris (Nycticebus coucang) were described and an attempt was made to correlate populations of fiber types and locomotor patterns. Muscle fibers were assigned to one of the following groups: fast-twitch glycolytic (FG), fast-twitch oxidative-glycolytic (FOG), and slow-twitch oxidative (SO). Histochemical techniques for the demonstration of alkaline- and acid-stable ATPases, succinic dehydrogenase, and mitochondrial á-glycerophosphate dehydrogenase were used in the classification of muscle fibers. Results indicated that the FG fiber type is the predominant fiber type in muscles used for jumping, the FOG fiber type is predominant in muscles used for running, and the SO fiber type occurs in high percentages in postural muscles. The SO fiber was also the most common fiber in muscles of the slow loris--a species that exhibits a slow, deliberate, sustained locomotor pattern. Intramuscular regional variations in populations were seen in some larger muscles of the tree shrew, but not in the lesser bushbaby and slow loris. Our results did not support the contentions of others that analogous muscles in different species have similar populations of fiber types.     

Distribution, density, and structure of muscle spindles in the vastus intermedius and the peroneus longus muscles of sheep

Muscle spindles are not always distributed more in postural muscles with many slow-twitch-oxidative (SO) myofibers than in locomotory muscles with few SO myofibers. The purpose of present study was to examine the distribution, density, and structure of muscle spindles in the vastus intermedius muscle: an antigravity muscle and the peroneus longus muscle: a locomotory muscle in the sheep. Muscle spindles were reconstructed from serial sections at 300 microns intervals throughout the muscles. Myofiber types were classified into SO, fast-twitch-oxidative-glycolytic, and fast-twitch-glycolytic myofibers by differences in histochemical reactivity. No significant difference in the density of muscle spindles (DMS) existed between the vastus intermedius (DMS: 5.3) and peroneus longus (DMS: 5.7) muscles. The muscle spindles were distributed more in the distal portion than in the proximal portion of the vastus intermedius muscle. The muscle spindles were distributed in the proximal and middle portion but hardly in the distal portion of the peroneus longus muscle. Muscle spindles were classified into simple, tandem, and compound muscle spindles. Most of the muscle spindles were the simple type. The differences in size of the muscle spindle and numbers of the intrafusal myofibers were not significant between the two muscles. The results show that the density and structure of the muscle spindles do not differ between the postural and locomotory muscles in the sheep.     

Neuromuscular spindles relative to joint movement complexities

Small muscles, such as those producing movements of the digits, are known to contain high densities of muscle spindles compared to larger muscles of the limb. These high densities have been associated with an increased need for proprioceptive feedback during precise manipulative movements. A recent study has indicated that spindle densities are related to muscle size rather than to functional capability. By contrast, the current study examined whether spindle numbers, actual spindle densities and relative spindle densities correlated with the complexity of joint movements in two species with different muscle sizes, human and guinea pig. Published data were used for the histological parameters of each species and measurements were made of movement range in each kinematic degree of freedom for each joint of the human and guinea pig pectoral and pelvic limbs. Muscle weights of newborn humans were 315 times greater than those of guinea pigs but joint movement complexities were almost the same in the two species and actual spindle numbers of human were only 7.3 times those of guinea pig. Joint movement complexity tended to be greatest most proximally on each limb and progressively decreased more distally. Regardless of muscle weight differences, actual spindle numbers and relative spindle densities correlated positively and significantly with joint movement complexity and actual spindle densities correlated negatively in each species. Hence muscle spindle numbers and densities around joints appear to reflect joint functional capability for appropriate proprioceptive feedback to the central nervous system in the control of joint movement.     

The moment arms of the muscles spanning the glenohumeral joint: a systematic review

The moment arm of a muscle represents its leverage or torque‐producing capacity, and is indicative of the role of the muscle in joint actuation. The objective of this study was to undertake a systematic review of the moment arms of the major muscles spanning the glenohumeral joint during abduction, flexion and axial rotation. Moment arm data for the deltoid, pectoralis major, latissimus dorsi, teres major, supraspinatus, infraspinatus, subscapularis and teres minor were reported when measured using the geometric and tendon excursion methods. The anterior and middle sub‐regions of the deltoid had the largest humeral elevator moment arm values of all muscles during coronal‐ and scapular‐plane abduction, as well as during flexion. The pectoralis major, latissimus dorsi and teres major had the largest depressor moment arms, with each of these muscles exhibiting prominent leverage in shoulder adduction, and the latissimus dorsi and teres major also in extension. The rotator cuff muscles had the largest axial rotation moment arms regardless of the axial position of the humerus. The supraspinatus had the most prominent elevator moment arms during early abduction in both the coronal and scapular planes as well as in flexion. This systematic review shows that the rotator cuff muscles function as humeral rotators and weak humeral depressors or elevators, while the three sub‐regions of the deltoid behave as substantial humeral elevators throughout the range of humeral motion. The pectoralis major, latissimus dorsi and teres major are significant shoulder depressors, particularly during abduction. This study provides muscle moment arm data on functionally relevant shoulder movements that are involved in tasks of daily living, including lifting and pushing. The results may be useful in quantifying shoulder muscle function during specific planes of movement, in designing and validating computational models of the shoulder, and in planning surgical procedures such as tendon transfer surgery.     

A comparison of the spindles in two different muscles of the frog

1. The responses of spindles in the iliofibularis muscle of frogs to stretch during either small motor nerve fibre stimulation or the application of suxamethonium were compared.2. All spindles which were excited by small motor nerve fibre stimulation were also excited by suxamethonium, and their responses to these two methods of excitation were very similar. The drug dose was usually 5-10 mug/ml. but smaller and larger doses were effective. Large doses (> 100 mug/ml.) could sometimes lead to a reversible partial block of the spindle response to stretch.3. Suxamethonium also caused a prolonged contraction in extrafusal slow muscle fibres. This contraction was not responsible for the effect on the spindle, because the time course of its action on the muscle tension and on the spindle afferent was different.4. It was concluded that suxamethonium stimulated prolonged contraction in the small intrafusal muscle fibres, which are known to be innervated by the small motor nerve fibres.5. Only about half of the spindles in the iliofibularis muscle were excited by suxamethonium.6. In the sartorius muscle which has no slow extrafusal muscle fibres, no spindles were found to be excited by suxamethonium in the way characteristic of that due to small intrafusal muscle fibre contraction.7. It is concluded that, in frog muscles which have no slow extrafusal fibres, the muscle spindles do not have small intrafusal muscle fibres of the kind found in the iliofibularis muscle.     

Detection of simultaneous movement at two human arm joints

To detect joint movement, the brain relies on sensory signals from muscle spindles that sense the lengthening and shortening of the muscles. For single-joint muscles, the unique relationship between joint angle and muscle length makes this relatively straightforward. However, many muscles cross more than one joint, making their spindle signals potentially ambiguous, particularly when these joints move in opposite directions. We show here that simultaneous movement at adjacent joints sharing biarticular muscles affects the threshold for detecting movements at either joint whereas it has no effect for non-adjacent joints. The angular displacements required for 70% correct detection were determined in 12 subjects for movements imposed on the shoulder, elbow and wrist at angular velocities of 0.25–2 deg s⁻¹. When moved in isolation, detection thresholds at each joint were similar to those reported previously. When movements were imposed on the shoulder and wrist simultaneously, there were no changes in the thresholds for detecting movement at either joint. In contrast, when movements in opposite directions at velocities greater than 0.5 deg s⁻¹ were imposed on the elbow and wrist simultaneously, thresholds increased. At 2 deg s⁻¹, the displacement threshold was approximately doubled. Thresholds decreased when these adjacent joints moved in the same direction. When these joints moved in opposite directions, subjects more frequently perceived incorrect movements in the opposite direction to the actual. We conclude that the brain uses potentially ambiguous signals from biarticular muscles for kinaesthesia and that this limits acuity for detecting joint movement when adjacent joints are moved simultaneously.     

Neural organization of spindles in three hindlimb muscles of the rat

The neuroanatomical organization of the dynamic (bag₁) and static (bag₂ and chain) intrafusal systems was compared by light and electron microscopy of serial sections among 71 poles of muscle spindle in soleus (SOL), extensor digitorum longus (EDL), and lumbrical (LUM) muscles in the rat. Eighty-four percent of 195 fusi-motor (γ) axons to the spindles innervated either the dynamic bag₁ fiber or the static bag₂ and/or chain fibers. Sixteen percent of the γ axons coin-nervated the dynamic and static intrafusal fibers. Some of these nonselective axons were branches of efferents that also gave rise to axons selective to either the dynamic or static types of intrafusal fibers in one or more spindles. Thus activation of individual stem γ efferents might not have a purely dynamic or purely static effect on the integrated afferent outflow from spindles of a hind-limb muscle in the rat. In addition, primary afferents in all muscles had terminations that cross-innervated the dynamic bag₁ and static bag₂ and/or chain intrafusal fibers in individual spindles, an arrangement that may enhance the mixed dynamic/static behavior of afferents when different intrafusal fibers are activated concurrently. Spindles of the slow SOL and fast EDL muscles had similar features, whereas differences were observed in the organization of the proximal (SOL and EDL) and distal (LUM) muscles. Spindles in LUM muscles had fewer static intrafusal fibers, a higher ratio of dynamic to static γ axons, and a higher incidence of skeletofusimotor (β) innervation to intrafusal fibers than spindles in the SOL or EDL muscles. Thus, the relative contribution of dynamic and static systems to muscle afferent outflow may differ among spindles located in different segments of the rat hindlimb. However, the dynamic and static intrafusal systems of spindle were less sharply demarcated in each of the three hindlimb rat muscles than in the cat tenuissimus muscle.     

EMG of scapulohumeral muscles in the chimpanzee during reaching and “arboreal” locomotion

Current views on the function of the deltoid and rotator cuff muscles emphasize their roles in arm-raising as participants in a scapulohumeral force “couple.” The acceptance of such a mechanism is based primarily on a 1944 EMG study of human shoulder muscle action. More recently, it has been suggested that shoulder joint stabilization constitutes a second and equally important function of the cuff musculature, especially in nonhuman primates which habitually use their forelimbs in overhead postural and locomotor activities. Few comparative data exist, however, on the actual recruitment patterns of these muscles in different species. In order to assess the general applicability of a scapulohumeral force couple model, and the functional significance of the differential development of the scapulohumeral musculature among primate species, we have undertaken a detailed study of shoulder muscle activity patterns in nonhuman primates employing telemetered electromyography, which permits examination of unfettered natural behaviors and locomotion. The results of our research on the chimpanzee, Pan troglodytes, on voluntary reaching and two forms of “arboreal” locomotion reveal four ways in which previous perceptions of the function of the scapulohumeral muscles must be revised: (1) the posterior deltoid is completely different in function from the middle and anterior regions of this muscle; (2) the integrity of the glenohumeral joint during suspensory postures is not maintained solely by osseoligamentous structures; (3) the function of teres minor is entirely different from that of the other rotator cuff muscles and is more similar to the posterior deltoid and teres major; and (4) each remaining member of the rotator cuff plays a distinct, and often unique, role during natural behaviors. These results clearly refute the view that the muscles of the rotator cuff act as a single functional unit in any way, and an alternative to the force couple model is proposed.     

Muscle receptors: content of some of the extrinsic and intrinsic muscles of the larynx in the bonnet monkey (Macaca radiata)

The number of muscle receptors and in particular the muscle spindle content of some of the intrinsic and extrinsic laryngeal muscles was evaluated, using bonnet monkeys (Macaca radiata). The technique of Barker and Ip [J. Physiol. (Lond) 169:73-74, 1963] was used to stain the muscle receptors. The results indicate that the intrinsic laryngeal muscles were devoid of muscle spindles. The extrinsic laryngeal muscles that were examined contain muscle spindles, though their density is less than that reported in the spindle-rich muscles like the rotators of the head and some of the intrinsic muscles of the hand (Cooper in: Structure and Function of Muscle, Vol. I, G.H. Bourne, ed, Academic Press, New York and London, 1960).     

The responses of frog muscle spindles and fast and slow muscle fibres to a variety of mechanical inputs

1. The tension in the iliofibularis muscle of frogs was recorded while the muscle was stretched or released. At the same time recordings were made from single spindle afferents in dorsal root filaments. Either large or small motor nerve fibres were stimulated in split ventral root filaments.2. While small motor nerve fibres were stimulated the discharge from muscle spindle afferents was greatly increased by stretching, and greatly reduced by shortening the muscle. This sensitivity to movement was shown even if the movements were small, so that a stretch of 0.2% of the muscle length was sufficient to cause a pronounced increase in the afferent discharge.3. In contrast, during stimulation of the large motor nerve fibres the spindle was much less sensitive to movements with the result that even stretches or releases of the muscle by 1 mm did not cause very large changes in the discharge frequency.4. The tension in slow extrafusal muscle fibres in many ways mirrored the spindle discharge during the stimulation of small motor nerve fibres, for the tension was greatly increased by stretching, even through small distances, and greatly reduced by releasing the muscle. The tension in fast extrafusal muscle fibres was much less changed by such movements, and thus was rather like the spindle discharge during stimulation of large motor nerve fibres.5. As the extrafusal muscle fibres do not directly pull on and excite the spindle afferents, the simplest explanation for the similarities between the muscle tension and the spindle discharge is that the mechanical properties of the intrafusal muscle fibres innervated by the large motor nerve fibres are like those of fast extrafusal muscle fibres, and that the mechanical properties of the small intrafusal fibres are similar to those of slow extrafusal muscle fibres.6. It is shown that the cross-bridge sliding filament mechanism of muscle contraction provides a ready explanation for the differences found between fast and slow muscles, and it is concluded that a most important functional difference between the two sorts of intrafusal muscle fibres is the speed of their contractions, for it is this which determines their contrasting actions on the spindle.7. It was also found that low rates (< 4/sec) of small motor nerve fibre stimulation were often very effective in exciting the spindles. These rates produced rather little extrafusal tension.     

The afferent innervation of two infrahyoid muscles of the cat

1. The afferent innervation of the straplike muscles of the infrahyoid region were investigated in two ways. The morphology of spindles and counts of tendon organs were investigated by the gold chloride technique in ten muscles. Spindle counts were made in forty pairs of thyrohyoid and infrahyoid muscles. De-efferenting of the nerves to these muscles was done in three cats and the calibre spectra of the afferent innervation investigated. These were compared with the total counts of fibres in intact nerves.2. In the thyrohyoid, spindles are frequently absent. No tendon organs were seen. In the large infrahyoid (combined sternohyoid and sternothyroid), spindle counts varied from 0 to 20 and the mean spindle count per gram of muscle was 3.5. A maximum of five tendon organs were seen in the muscle. Both spindle and tendon organ counts are low when compared with a limb muscle of similar weight and size.3. In the infrahyoid muscle complex spindles were about equal in number to simple spindles.4. Counts of spindles in the infrahyoid muscle in families of three or more siblings suggest that some families of kittens tend to have higher spindle counts than other families.5. The afferent innervation of the two muscles varied between 21 and 42% of the total fibre population and the fibre diameter spectrum is in keeping with the low counts of encapsulated endings.     

Difference in muscle spindle structure between pigeon muscles used in aerial and terrestrial locomotion

Muscle spindle density (number of spindes per gram of muscle) of all 29 muscles of the forearm and leg of the domestic pigeon was evaluated by counting receptors in van Gieson-stained serial cross sections. Extra- and intrafusal fiber-type profiles were determined from histochemical preparations. Muscles of the leg had on the average significantly more avian slow-twitch oxidative extrafusal fibers (22.5 vs. 0.8%) and slower contraction times than muscles of the forearm, but fiber-type profiles and gross actions of muscles showed no consistent relation to the relative abundance of receptors. Differences in intrafusal fibertype composition among spindles were sought because of their potential effect on the quality of the afferent discharge. The number of intrafusal fibers per spindle was on the average significantly less (4.57 vs. 5.99) in the muscles of the leg than in those of the forearm; and of spindles with the same number of intrafusal fibers, those in the leg had smaller periaxial spaces. Distribution of intrafusal fiber types identified with the myofibrillar adenosine triphosphatase reaction differed among spindles of varying sizes. An acid- and alkali-labile type occurred most frequently (P = 0.05) in spindles with one to three intrafusal fibers, and an acid-labile and alkali-stable type was most often seen (P = 0.05) in spindles with 4 to 7 intrafusal fibers. The smaller receptors were more abundant in the leg, while the larger ones were about equally distributed between the two extremities. Muscle fibers with dimensions that sometimes approached small extrafusal fibers were present in about 3% of the axial bundles examined, most of them in the forearm. The selective morphological variation of avian muscle spindles may represent the structural basis for qualitatively different afferent discharges that relate to the characteristic types of locomotion served by the two extremities.     

Critical period in muscle spindle regeneration in grafts of developing rat muscles

Extensor digitorum longus (EDL) muscles from rats at various intervals after birth were grafted into EDL muscles of adult recipients. Three to twelve months after the operation, host muscles containing the grafts were removed and examined for the presence of muscle spindles in the graft. The aim of the study was to establish when muscle spindles become capable of regeneration during development. Regenerated muscles grafted during the first week after birth were virtually spindleless. Grafts of muscles transplanted 10 and 15 days postnatally contained only 5–8 muscle spindles on average. In contrast, the regenerated grafts originating from muscles of 24- and 28-day-old rats were spindle-rich as in mature muscle grafts; the number of spindles in the transplanted EDL muscles (25.0±2.3; mean ±SE) attained values comparable to free standard autografts of these muscles in adult animals. Thus, the critical period after grafting, which also involves the loss of a vascular supply, is considerably longer than the critical period for muscle-spindle survival after nerve injury. Fifteen days after birth, when muscle spindles still survive denervation, only a few regenerated spindles were present in the individual muscle regenerates. We assume that the low resistance of immature spindle capsules to ischaemia accounts for their massive degeneration and abortive spindle regeneration in grafts from 10- to 15-day-old rats.     

Discharge of spindle afferents from jaw-closing muscles during chewing in alert monkeys.

The discharge of muscle spindle afferents from monkey spindle afferents from monkey jaw-closing muscles was studied during mastication of natural foods by extracellular recording from the fibers or cell bodies of the tract and mesencephalic nucleus of the fifth nerve. In all, 39 muscle afferents were studied. The spindle associated with 18 of the afferents was positively identified by the afferent's response to gentle, localized palpation of either the temporalis or masseter muscle. Discharge patterns were observed during mastication, and in the majority of cases the qualitative passive response characteristics of the spindle afferent were determined. During steady chewing spindle afferent discharge typically paused briefly during the initial rapid upward part of the chewing cycle. Firing generally began as the jaw slowed its upward movement, and firing rates during the slow grinding portion of the upward movement were within the range of 50-80 spikes/s. All spindles exhibited a brisk discharge during the opening movement, typically within the range of 100-150 spikes/s. One-third of the spindle afferents exhibited a brief, high-frequency burst of firing at the very beginning of the opening movement, presumably as a result of stretch applied to a spindle just previously subjects to fusimotor excitation. Although the results of the study make it clear that spindles in jaw-closing muscles are coactived along with the extrafusal muscle fibers, the fusimotor bias does not seem capable of sustaining discharge in the face of rapid shortening of the muscle. Furthermore, the fact that discharge rate during opening, when the jaw-closing motoneurons are quiescent, is much higher than at any part of the closing cycle, when the motoneurons are active, suggests that the muscle spindles cannot provide the primary excitatory drive to the motoneurons.