Phrenic motor unit recruitment during ventilatory and non-ventilatory behaviors

Phrenic motoneurons are located in the cervical spinal cord and innervate the diaphragm muscle, the main inspiratory muscle in mammals. Similar to other skeletal muscles, phrenic motoneurons and diaphragm muscle fibers form motor units which are the final element of neuromotor control. In addition to their role in sustaining ventilation, phrenic motor units are active in other non-ventilatory behaviors important for airway clearance such as coughing or sneezing. Diaphragm muscle fibers comprise all fiber types and are commonly classified based on expression of contractile proteins including myosin heavy chain isoforms. Although there are differences in contractile and fatigue properties across motor units, there is a matching of properties for the motor neuron and muscle fibers within a motor unit. Motor units are generally recruited in order such that fatigue-resistant motor units are recruited earlier and more often than more fatigable motor units. Thus, in sustaining ventilation, fatigue-resistant motor units are likely required. Based on a series of studies in cats, hamsters and rats, an orderly model of motor unit recruitment was proposed that takes into consideration the maximum forces generated by single type-identified diaphragm muscle fibers as well as the proportion of the different motor unit types. Using this model, eupnea can be accomplished by activation of only slow-twitch diaphragm motor units and only a subset of fast-twitch, fatigue-resistant units. Activation of fast-twitch fatigable motor units only becomes necessary when accomplishing tasks that require greater force generation by the diaphragm muscle, e.g., sneezing and coughing.     

Morphological and histochemical organization of the flexor carpi radialis muscle in the cat

Most studies concerning the structure and function of skeletal muscle have utilized the hind limb of the experimental animal. However, it has been shown that the number of behavioral tasks performed by the cat's forelimb is greater than that of the hind limb. In addition, the forelimb muscles exhibit a functional complexity not observed in hind-limb musculature. The purpose of this study was to investigate the distribution of fast-twitch and slow-twitch muscle fibers and muscle spindles in the flexor carpi radialis muscle (FCR) and to correlate the distributional patterns in these structures with muscle tendon architecture and muscle function. It was found that the FCR, a wrist flexor, contains 37% slow-twitch fibers and 63% fast-twitch fibers. However, the slow-twitch fibers were concentrated in the deep region located between the tendons of origin and insertion, while the fast-twitch-glycolytic fibers were concentrated more peripherally. Muscle spindles were associated with the slow-twitch region and were never found in the region containing high concentrations of fast-twitch-glycolytic fibers. Fast-twitch-oxidative-glycolytic fibers were uniformly distributed throughout the muscle. It is proposed that the association of muscle spindles with slow-twitch fibers and the differential distribution of muscle fibers into slow-twitch and fast-twitch regions might allow these regions to function independently of one another when called upon to perform complex behavioral tasks.     

Changes of skeletal muscle proteases activities during a chronic low-frequency stimulation period

Modifications of muscular contractile patterns by chronic low-frequency stimulation induce structural, physiological, and biochemical transformations in fast-twitch fibers that cause them to act like slow-twitch muscle. During this transformation many changes in protein pattern appear and the proteolytic system may be involved in those changes. The activities of cathepsin L, B, H, D, the level of cystatin, as well as the calpain activity in rabbit fast-twitch muscle have been compared with those of slow-twitch muscle. The results show that fast-twitch muscle has lower cathepsin activities and higher calpain activities than slow-twitch muscle. Chronic low-frequency stimulation was applied for 24 days to fast-twitch muscles and changes in proteases and protease inhibitors (cystatin and calpastatin) were studied. After 7-14 days of stimulation, lysosomal cathepsin L, B, and D and cytoplasm calpain and proteosome activities increased several-fold. Involvement of the phagocyte cells in the protein fiber turnover was minimal. Although the turnover of contractile proteins during muscle electrostimulation takes place in synchrony with changes in the muscle proteolytic system, the stimulation period used did not attain the total transformation from fast- to slow-twitch muscle proteolytic pattern.     

Heterogeneity of motor units activating single Golgi tendon organs in cat leg muscles

Summary The aim of this study was to investigate whether an individual Golgi tendon organ can signal the contraction of motor units with different physiological properties. The axonal conduction velocity and tetanic tension of motor units were examined in four muscles of the cat leg (peroneus brevis, peroneus longus, tibialis anterior and soleus). The motor units which were found to activate a given tendon organ had contractile properties dispersed over the same range as those of the whole muscle population. The proportion of tendon organ-activating motor units found in the studied samples suggests that altogether, the Golgi tendon organs of a muscle monitor the contraction of every motor unit in this muscle.     

Enlarged motor units resulting from partial denervation of cat hindlimb muscles

1. It was the aim of this study to determine the extent to which a mammalian motoneuron can sprout in a partially denervated muscle, which motor unit types are involved in sprouting, and whether polyneuronal innervation exists between sprouted units. 2. The fast-twitch flexor digitorum longus (FDL) and slow-twitch soleus were partially denervated by unilateral section of the L7 ventral root in 12-wk-old kittens. After approximately 100 days single motor units were isolated, and their isometric contractile characteristics were determined. FDL units were also tested for their resistance to fatigue and categorized as fast-twitch, fatiguing fibers (FF), fast-twitch, fatigue-resistant fibers (FR), and slow-twitch, fatigue-resistant fibers (S). The presence of polyneuronal innervation was investigated between pairs of like and unlike units. 3. The extent of the original denervation was variable and was estimated from the distribution of motor axons innervating the muscle via the L7 and S1 (soleus) or L6 and L7 (FDL) ventral roots on the contralateral side. In soleus, denervations ranged from 75 to 98%; in FDL, 60 to 97% (denervations less than 60% were not investigated). In general, motor-unit force increased in proportion to the extent of the denervation. 4. Within soleus, unit force increased to over 2 N, which was about 16 times greater than the average for a normal muscle (133 mN). However, most units increased in force to between five and 12 times normal. 5. Within FDL, the force development of type S units was unaffected by partial denervation. Type FF units increased by up to 11 times (4.3 N) compared with normal FF units (395 mN) with most increasing between two and four times. FR units exhibited the greatest relative increase in force [up to 19 times (4.3 N) compared with normal (225 mN)]. Most units were two to seven times the normal. 6. A few FDL units were glycogen depleted, the muscles frozen, and cross sections prepared for histochemical analysis. This indicated that the largest units contained approximately 5,000 fibers, and there was little fiber hypertrophy. In the extensively denervated soleus muscle, large numbers of small, presumably denervated fibers were observed. The innervation ratio of several large units was determined indirectly using mean fiber area. This gave estimates of 3,000-4,000 fibers for the largest units. Again, fiber hypertrophy contributed little to the increase in unit force. It was concluded that the increased force of units in both muscles was largely attributable to terminal and axonal sprouting of the intact motor axons. 7. No evidence for polyneuronal innervation was found in either FDL or soleus muscle.(ABSTRACT TRUNCATED AT 400 WORDS)     

Myosin heavy chain expression in developing rat intrafusal muscle fibers

The immunocytochemical expression of several isoforms of myosin heavy chains (MHC) was determined in developing intrafusal and extrafusal fibers of the soleus muscle of prenatal and postnatal rats. At the onset of spindle assembly, both bag2 intrafusal myotubes and primary extrafusal myotubes bound a slow-twitch MHC antibody, whereas the bag1 and chain myotubes expressed a fast-twitch MHC isoform identical to that expressed by secondary extrafusal myotubes. Subsequently, developing intrafusal fibers began to express unique myosin isoforms, and ceased to express some of the myosin isoforms present initially. The initial similarity in MHC composition of intrafusal and extrafusal fibers suggests that these two kinds of mammalian muscle cell originate from a common pool of bipotential myotubes. Differences in MHC expression by intrafusal and extrafusal fibers in adult muscles might result from the effect of sensory neurons on the developing intrafusal myotubes.     

Effects of streptozotocin-induced diabetes on leg muscle contractile properties and motor neuron morphology in rats

Skeletal muscle fiber subtypes are differentially sensitive to diabetes-related pathology; For example, fast-twitch muscles exhibit severe decreases in contraction force while slow-twitch muscles demonstrate prolonged half-relaxation time. However, such alterations have only been examined after a relatively short period following diabetes onset, with no information available regarding muscle damage caused by longer disease periods (>20 weeks). This study examined alterations in the contractile properties of the medial gastrocnemius (fast-twitch) and soleus (slow-twitch) muscles, as well as morphological changes in their motor neurons 12 and 22 weeks after diabetes onset. Adult male Wistar rats were divided into diabetic (12- or 22-week post-streptozotocin injection) and age-matched control groups. Electrically evoked maximum twitch and tetanic tension were recorded from leg muscles. Additionally, motor neuron number and cell body size were examined. At 12 weeks after diabetes onset, decreases in twitch force were observed predominantly in medial gastrocnemius muscles, while soleus muscles exhibited prolonged half-relaxation time. However, these differences became ambiguous at 22 weeks, with decreased twitch force and prolonged half-relaxation time observed in both muscles. On the other hand, reduction in soleus motor neurons was observed 12 weeks after diabetes onset, while medial gastrocnemius motor neurons were diminished at 22 weeks. These data indicate that experimental diabetes induces differential damage to medial gastrocnemius and soleus muscles as well as motor neurons. These diabetes-induced differences may partly underlie the differential deficits observed in gastrocnemius and soleus.     

A comparative study of muscle spindles in slow and fast neonatal muscles of normal and dystrophic mice

Muscle spindles from the slow-twitch soleus and the fast-twitch extensor digitorum longus (EDL) muscles of genetically dystrophic mice of the dy^2J/dy^2J strain were compared with age-matched normal animals at neonatal ages of 1–3 weeks according to histochemical, quantitative, and ultrastructural parameters. Intrafusal fibers in both the soleus and EDL exhibited similar regional differences in myosin ATPase activity, and conformed to those noted previously in various adult species. In distal polar regions, all nuclear bag fibers resembled extrafusal fibers of the type 1 variety, whereas in capsular zones they could be divided into two subtypes. Nuclear chain fibers possessed a staining pattern similar to type 2 extrafusal fibers, and in contrast to the bag fibers they exhibited no regional variations. These features were consistently observed in both the normal and dystrophic muscles at all ages. Spindles varied only slightly in their number and distribution in the two types of muscle, and their location followed the neurovascular branching pattern in each. Irrespective of age or genotype, spindles in the soleus were more homogeneously dispersed, but those in the EDL were concentrated along the dorsal aspect of the muscle. No significant differences were noted in the total number of spindles between normal and dystrophic muscles. In addition, no dramatic differences were observed in the muscle spindle index for soleus and EDL. The first obvious disease-related changes were noted in extrafusal fibers of the soleus of 3-week-old mice, and spindles were often located close to these areas of fiber degeneration. Despite alterations in the surrounding tissue, however, spindles appeared morphologically unaltered in dystrophy. These observations indicate that intrafusal fibers of spindles in neonatal mice appear enzymatically and histologically unaffected in incipient stages of progressive muscular dystrophy.     

Quantitative ultrastructure of histochemically identified avian skeletal muscle fiber types

A cryostat retrieval method and combined adenosine triphosphatase (ATPase) and acetylcholinesterase (AChase) method were used to study the ultrastructure and innervation of histochemically identified skeletal muscle fibers in different pigeon muscles. The Z-line structure and volume percentage sarcotubular system were analyzed from different muscles selected for their composition by fiber type. Histochemically, three main fiber types were investigated: slow tonic fibers with a moderate ATPase activity after preincubation at acid or alkaline pH; fast-twitch fibers that had high activity after alkaline treatment and low activity after acid preincubation; and a type considered to be slow-twitch that had low activity after alkaline, and high after acid preincubation. Both the slow tonic and slow-twitch fibers had multiple, en grappe innervation, while the fast-twitch fibers had robust, single end plates. The Z-line of the fast-twitch and slow-twitch fibers had a regular square lattice pattern, in contrast to the granular, nonlattice structure of the slow tonic Z-line. The volume percentage sarcotubular system of the slow-twitch fibers was intermediate between and significantly different from that of the fast-twitch and slow tonic fibers. These correlative analyses suggest that the avian muscles contain not only the fast-twitch and slow tonic fibers previously known, but also a slow-twitch fiber that appears to be intermediate between the tonic and the mammalian slow-twitch fiber type. Based on the abundance of the sarcotubular system, this fiber type appears to be fast-contracting and -relaxing, in spite of being multiply innervated.     

Effects of diabetes on protein synthesis in fast- and slow-twitch rat skeletal muscle

The effects of acute (2-day) and long-term (7-day) diabetes on rates of protein synthesis, peptide-chain initiation, and levels of RNA were examined in rat skeletal muscles that are known to have differing proportions of the three fiber types: fast-twitch white, fast-twitch red, and slow-twitch red. Short-term diabetes resulted in a 15% reduction in the level of RNA in all the muscles studied and an impairment in peptide-chain initiation in muscles with mixed fast-twitch fibers. In contrast, the soleus, a skeletal muscle with high proportions of slow-twitch red fibers, showed little impairment in initiation. When the muscles were perfused as a part of the hemicorpus preparation, addition of insulin to the medium caused a rapid reversal of the block in initiation in mixed fast-twitch muscles but had no effect in the soleus. The possible role of fatty acids in accounting for these differences is discussed. Long-term diabetes caused no further reduction in RNA, but resulted in the development of an additional impairment to protein synthesis that also affected the soleus and that was not corrected by perfusion with insulin. The defect resulting from long-term diabetes may involve elongation or termination reactions.     

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.     

Non-neural and neural expression of myosin heavy chains by regenerated intrafusal fibers of rats.

Expression of myosin heavy chain (MHC) isoforms was studied in rat soleus (SOL) and extensor digitorum longus (EDL) muscles which regenerated in the presence or absence of innervation. Frozen sections of two 5 day denervated SOL and EDL grafts, two 40 day denervated SOL and EDL grafts, and two reinnervated 40 day SOL and EDL grafts were processed for demonstration of motor endplates, sensory endings, myosin adenosine triphosphatase (mATPase) and for expression of 4 MHCs. No qualitative differences in MHC expression were noted between 5 day or 40 day denervated grafts of the SOL and EDL muscles. All regenerated intrafusal and extrafusal myotubes or myofibers reacted to antibodies against neonatal and fast-twitch MHCs, but not to antibodies against slow-twitch and slow-tonic MHCs in these grafts. These data indicate that MHCs expressed by regenerated intrafusal myotubes do not parallel those expressed by myotubes which give rise to the three types of intrafusal fibers during development and that MHC expression by regenerated intrafusal myotubes parallels that of regenerated extrafusal myotubes prior to innervation. However, some regenerated intrafusal fibers in 40 day nerve-intact grafts bound antibodies to slow-twitch and slow-tonic MHCs, thus expressions of these two MHCs are nerve-dependent in regenerated muscle spindles.     

Ensemble discharge from Golgi tendon organs of cat peroneus tertius muscle

1. The responses of individual tendon organs of the cat peroneus tertius muscle to motor-unit contractions were recorded in anesthetized cats during experiments in which all the Ib-afferent fibers from the muscle had been prepared for recording in dorsal root filaments. This was possible because the cat peroneus tertius only contains a relatively small complement of approximately 10 tendon organs. 2. Motor units of different physiological types were tested for their effects on the whole population of tendon organs in the muscle. Effects of unfused tetanic contractions were tested under both isometric and anisometric conditions. Each motor unit activated at least one tendon organ, and each tendon organ was activated by at least one motor unit. Individual slow-type units were found to act on a single or two receptors, whereas a fast-type unit could activate up to six tendon organs. 3. In one experiment, the effects of 8 motor units on 10 tendon organs were examined. One fast-twitch, fatigue resistant (FR)-type unit acted on six tendon organs, of which four were also activated by another FR unit. The contraction of each unit, on its own, elicited a range of individual responses, from weak to strong. The discharge frequencies of individual responses displayed no clear relation with the strength of contraction, nor did they accurately represent the shape of force profiles. But when all the discharges were pooled, a fairly good correspondence appeared between variations of contractile force and variations of averaged discharge frequencies.(ABSTRACT TRUNCATED AT 250 WORDS)     

Histochemical profiles of fibers in the rat tibialis anterior muscle during early postnatal development

The enzyme activities of intra- and extrafusal fibers in the tibialis anterior muscle of rats during postnatal development have been investigated. Muscle fibers 1 day after birth showed a uniform reaction for adenosine triphosphatase (ATPase), succinate dehydrogenase (SDH), and alpha-glycerophosphate dehydrogenase (alpha-GPD) activities. Fast-twitch (F) and slow-twitch (S) fibers with ATPase activity were found at 9 and 11 days. Thereafter, the type shift of muscle fibers from S to F was observed in the deep and middle portions. Fast-twitch oxidative glycolytic (FOG), fast-twitch glycolytic (FG), and slow-twitch oxidative (SO) fibers with ATPase, SDH, and alpha-GPD activities were found at 15 (the superficial portion) and 17 days (the deep and middle portions). The histochemical differentiation of intrafusal muscle fibers (7 and 9 days) was found earlier than that of extrafusal muscle fibers.     

Changes in properties of the medial gastrocnemius motor units in aging rats

1. The properties of motor units were investigated in the medial gastrocnemius (MG) of old rats [27.5 +/- 1.6 (SD) mo old, n = 18]. Individual motor units were functionally isolated by ventral root fiber splitting and grading stimulus intensity. The muscle-unit portion of the motor unit was identified by the glycogen depletion method. The physiological properties of 77 motor units in 6 animals and the histological results of 7 slow-twitch (type S) muscle units were compared with data from motor units in the same muscle of middle-aged rats (12.8 +/- 1.6 mo old, n = 33). 2. The motor units were classified into four types of categories [FF (fast-twitch motor units with a fatigue index less than or equal to 0.5), FI (fast-twitch motor units with a fatigue index greater than 0.5 but less than 0.75), FR (fast-twitch motor units with a fatigue index greater than or equal to 0.75), S (slow-twitch motor units with a fatigue index greater than 0.75)] using the same criteria (i.e., presence or absence of the "sag" property and fatigability) used for middle-aged rats. No significant difference in the relative distributions of these unit types was detected, although the MG muscle in old rats exhibited a relatively high proportion of type S units and fewer type FR units. 3. The mean tetanic tensions for type FF + FI and FR units were significantly smaller than those in the middle-aged rats. On the other hand, type S motor units produced more tension than in the middle-aged rats. 4. The conduction velocity of motor axons was considerably slower in any unit type of old motor units, and the most marked change was found in type FR units. 5. The general morphological features of the old rat MG were fiber-type grouping, disseminated atrophic or angulated fibers, a decrease in the total number of muscle fibers, and an increase in the number of type I muscle fibers. The major distribution patterns of fibers of different types were the same as those in the middle-aged MG. 6. Seven type S units that produced large tetanic tension were depleted of glycogen in the muscle-unit portions. These units had a large innervation ratio compared with those in the middle-aged rats, whereas the mean cross-sectional area of muscle fibers and the calculated specific tension remained unaltered.(ABSTRACT TRUNCATED AT 400 WORDS)     

Postnatal expression of myosin heavy chains in muscle spindles of the rat

Summary The immunocytochemical expression of two myosin isoforms in intrafusal muscle fibers was examined in soleus muscles of neonatal (zero to six days postpartum) and adult rats. Monoclonal antibodies specific for myosin heavy chains of the slow-tonic anterior latissimus dorsi (ALD58) and fast-twitch pectoralis (MF30) muscles of the chicken were used. In adults ALD58 bound to the intracapsular regions of bag₁ and bag₂ fibers and MF30 bound to the intracapsular regions of bag₂ and chain fibers. The extracapsular regions of intrafusal fibers and all extrafusal fibers did not react to ALD58 or MF30. Bag₁ and bag₂ fibers of neonatal rats expressed immature myosin patterns but chain fibers did not. The adult pattern of immunoreactivity of intrafusal fibers developed by the fourth postnatal day, when the patterns of motor but not sensory innervation in the spindle are still immature. Data suggest that the expression and maintenance of the specific anti-myosin immunoreactivity of intrafusal fibers during postnatal development of rat spindles is dependent upon sensory but not motor innervation. Moreover, afferents might regulate the gene expression responsible for synthesis of myosins isoforms specific to intrafusal fibers only in those myonuclei located within the capsule, but not in the myonuclei in extracapsular regions of intrafusal fibers.     

Effects of an anabolic steroid and sprint training on selected histochemical and morphological observations in rat skeletal muscle types

Summary The effects on selected histochemical and morphological parameters of anabolic steroid administration and of high-intensity sprint running, separately, and in combination, were studied in young adult male rats. Dianabol (methandrostenolone) 1 mg/day for 8 weeks had no significant effects on phosphorylase or glycogen staining intensities and on fiber area in skeletal muscles of either trained or sedentary animals. The program of sprint training resulted in significantly decreased intensities of phosphorylase in all ten regions of the gastrocnemius, plantaris, and soleus muscles that were studied. Glycogen localization was significantly increased with training in five regions of the gastrocnemius and plantaris muscles which contained predominantly fast-twitch fibers. No changes in fiber area occurred with the training program. We conclude from these results that (a) normal androgen levels in young, healthy male animals are sufficiently high so that the intake of large doses of anabolic steroid does not result in the stimulation of glycogen metabolism or hypertrophy of skeletal muscle; (b) the changes induced by high-intensity, short-duration sprint training suggest that the existing glycolytic capacity of muscle is adequate to supply the muscles energy needs even during the stress of very strenuous exercise, and that more fast-twitch fibers were recruited by the exercise regimen than slow-twitch fibers.     

Origin of intrafusal fibers from a subset of primary myotubes in the rat

S46, a monoclonal antibody (mAb) specific for the SM-1 and SM-2 isoforms of avian slow myosin heavy chains (MHC), was used to study the earliest stages of development of intrafusal fibers in muscle spindles of the rat hindlimb. Spindles formed only in the regions of fetal muscles that contained primary myotubes reactive to mAb S46, such as the axial region of the tibialis anterior muscle. The first intrafusal fiber to form, the nuclear bag₂ fiber, originated from within the population of S46-reactive primary myotubes. Binding of mAb S46 by myotubes giving rise to the bag₂ fibers preceded the appearance of encapsulated spindles in the muscles by electron microscopy. However, reactivity to S46 intensified in the myotubes transforming into bag₂ fibers after the innervation of the fibers by afferents, and dissipated in myotubes differentiating into slow-twitch (type I) extrafusal fibers. Thus, afferents may enhance intrafusal expression of the MHC isoform reactive to mAb S46. The pattern of S46 binding to nuclear bag and chain intrafusal fibers in both developing and adult spindles was the same as that reported for the mAb ALD19, suggesting that both antibodies bind to the same MHC isoform. This isoform is probably a developmental form of slow myosin, because it was transiently expressed during the development of type I extrafusal fibers. The origin of bag₂ intrafusal and type I extrafusal fibers from a bipotential subpopulation of primary myotubes reactive to mAb S46 correlates with the location of muscle spindles in the slow regions of muscles in adult rat hindlimbs.     

Frequency of tendon organ discharges elicited by the contraction of motor units in cat leg muscles.

1. The responses elicited in individual tendon organs by the contraction of single motor units were studied in peroneus longus, peroneus brevis, tibialis anterior and soleus muscles. 2. No simple relation was found between the discharge frequency of a tendon organ and the tension produced in the muscle tendon by the contraction of individual motor units. 3. The sensitivity of a given tendon organ to contractile tension was not the same for each of the motor units which elicited its discharge. There was no correlation between the sensitivity of the receptor and the strength of the motor units. 4. Upon repetitive stimulation of a tendon-organ-activating motor unit at increasing rates, the frequency of the receptor sustained discharge reached a maximal value for rates of stimulation eliciting submaximal tetanic tension. Higher rates only produced an increase in the dynamic component of the tendon organ response. 5. These observations show that the contractile tension sensed by a tendon organ is not a simple fraction of the tension which appears at the muscle tendon. They might be accounted for as consequences of the fine structure of tendon organs and of variations in the number of muscle fibres contributed by different motor units to the bundle inserted on each receptor. The location of most tendon organs at musculo-aponeurotic junctions rather than in the tendon proper, could also be responsible for some of the observed discrepancies.