Biomechanical organization of single motor units in two multi-tendoned muscles of the cat distal forelimb

Summary In anesthetized cats single motor units (MUs) of the extensor carpi ulnaris (ECU) and extensor digitorum communis (EDC) muscles were selectively activated by stimulation of cervical ventral root filaments. The distribution of force developed by single MUs at the four distal tendons of the EDC muscle and at three portions of the distal tendon of the ECU muscle was analysed. In general, single MUs of both muscles distributed force over all tendons in a unimodal pattern, with the maximal force levels generated at one specific tendon which was termed the best-tendon. Distributions of force were quantitatively described by a parameter representing the mean direction of force output (output-index) and a further one representing the dispersion of force over the distal tendons (divergence). Generally, these parameters and the best-tendon remained stable when a MU was stimulated at different frequencies, but varied from MU to MU. Despite the general stability of the force distribution, slight systematic changes were regularly found in EDC MUs, when they developed a higher amount of force due to a higher frequency of stimulation: the relative amount of force at the best-tendon increased; e.g. the MUs got more selective for the best-tendon. These changes were partly due to overcoming mechanical cross-coupling between neighbouring compartments of the EDC muscle. Such changes of force distribution were only found in a part of the ECU MUs; other ECU MUs did not change their force distribution at all or became less selective for the best-tendon. The phenomenon that MUs of multi-tendoned muscles distribute their force output to the distal tendons in specific patterns is probably due to mechanical partitioning of the parent muscles: the localization of spatial territories of MUs within different anatomical muscle compartments should correspond to the best-tendon. Complex mechanisms allowing passive transmission of force from limited territories along the transverse axis of both muscles must be assumed in order to explain why most MUs act on all tendons and why force distributions change with increasing stimulus frequency. In addition, specific relations between unit type and force distributions were found within both muscles. Fatigue-resistant EDC MUs have broader force distributions than fatigue-sensitive EDC MUs and slow ECU MUs were found to act predominantly on the most ulnar part of the distal tendon. These biomechanical properties of MUs are discussed as supporting the specific functions of the respective muscles.     

Relations among motor unit types,generated forces and muscle length in single motor units of anaesthetized cat peroneus longus muscle

The active length-tension curves of identified single motor units (MUs) belonging to peroneus longus muscle (PL) of anaesthetized adult cats were obtained by eliciting isometric single twitches and tetani. The recorded responses were evaluated by measuring the peak tension amplitude and the tension-time area at muscle lengths extending throughout the physiological length range of the muscle (mean 5.5 mm, standard deviation ±0.8). The muscle lengths at which each tested MU developed its maximal twitch (L _tw) and tetanic (L _te) tensions were determined and compared with the muscle length (L _o) at which the stimulation of all the -axons, innervating PL and contained in L7 ventral root, developed their maximal twitch tension. The mean of single MU L _tw values was at L _o+1.08±1.1 mm. Slow MUs showed the longest values of L _tw(L _o+1.6±1.0 mm). Single MUs stimulated at tetanic frequencies presented their L _te at values shorter than L _o (L _o–2.8±1.7 mm). Slow MUs had the shortest L _te (L _o–3.4±1.5 mm). For all the units L _te was shorter than L _tw. L _tw and L _te were, respectively, negatively and positively correlated with the developed tension. Optimal length values also appeared to be related to the MU types. The possibility is discussed that the muscle and tendon compliances and the high non-linearities to the applied forces are the main factors which can determine the differences among L _o, L _tw and L _te values. The relationships between MU type and optimal length values are suggested to be, at least partly, an epiphenomenon due to the different contraction strengths of the various MU types. However, the heterogeneous distribution of the MU types is brought into account to explain the dependence of L _tw and L _te values on MU type.     

Physiological properties of the motor units of the wrist extensor muscles in man

Summary The physiological properties of 355 motor units (MUs) recorded in the extensor carpi radialis muscles were studied in 34 healthy human subjects during isometric contractions. MU selective twitches were educed from the whole muscle force using the spike-triggered averaging method. The twitch contraction times and twitch forces were measured. From these data it was attempted to estimate the distribution of fast and slow MUs in the muscles studied. MU recruitment thresholds were systematically measured during stereotyped slow ramp contractions (force increase=0.25 N·s^-1). Degrees of correlation between contraction times, twitch forces and recruitment thresholds were pair analysed by computing simple regression curves and correlation coefficients. The degrees of correlation were compared between 245 MUs recorded in 34 subjects and 66 MUs recorded in a single subject. Analysis of the instantaneous discharge frequency of 132 MUs showed the existence of a remarkable degree of correlation (correlation coefficient, r=-0.75) between the frequency rise times (discharge onset to maximal frequency) and the MU twitch contraction times; i.e., the frequency rise times increase when the twitch contraction times decrease. The possibility that muscle contraction may be differentially modulated on the basis of this discharge property of the MUs is discussed. The results are compared to previous data and the limitations of the spike-triggered averaging method applied to long muscles in man are extensively discussed.     

Contractile properties of rat soleus motor units following 14 days of hindlimb unloading

The purpose of this study was to compare the isometric contractile properties of rat soleus motor units after 14 days of hindlimb unloading (HU) to those under control conditions. The motor units (MU) were classified using two mechanical criteria: the presence or not of a sag during unfused tetani and the value of the twitch time-to-peak (TTP). Under control conditions, the soleus muscle was composed of 85% of slow-type (sag −, TTP > 20 ms) and 15% of fast-type (sag +, TTP < 20 ms) units. Following HU, these two populations were still present and results showed: (1) large decreases in their maximal tetanic tensions (of −67% and −60% for slow- and fast-type, respectively), and (2) changes in their relative proportions, i.e. a decrease in the percentage of slow-type units and a twofold increase in the percentage of fast-type units were observed. These latter changes might be the consequence of a complete transformation of slow-towards fast-type units. A third population appeared in the HU solei, 26% of the samples, combining the presence of a sag and speed-related properties between those of slow- and fast-type units. These slow-intermediate units might come from slow units partially transformed into a faster type during HU. Thus the present study showed that unloading conditions induced a reorganisation of the soleus motor unit profile. The complete or partial transformation of the motor units could be related to the changes in the electromyographical activity of the unloaded soleus. Received: 30 June 1995/Received after revision: 12 January 1996/Accepted: 22 January 1996     

No graded responses of finger muscles to TMS during motor imagery of isometric finger forces

Previous studies of motor imagery have shown that the same neural correlates for actual movement are selectively activated during motor imagery of the same movement. However, little is known about motor imagery of isometric force. The aim of the present study was to investigate the neural correlates involved in motor imagery of isometric finger forces. Ten subjects were instructed to produce a finger flexion or extension force ranging from 10% to 60% of maximal isometric force and to mentally reproduce the force after an eight-second delay period. Transcranial magnetic stimulation (TMS) was applied over the hand motor area during imagining the force. We measured the amplitude of motor evoked potentials (MEPs) from the flexor digitorum superfialis (FDS) and the extensor digitorum communis (EDC) muscles and TMS-induced forces from the proximal phalanxes. The results showed that, as compared to the rest condition, the MEP amplitude was greater in the FDS during imagining flexion forces, whereas it was greater in the EDC during imagining extension forces. MEP amplitudes were similar for motor imagery of graded flexion or extension forces. Also, TMS produced flexion forces during imagining flexion forces, whereas it produced extension forces during imagining extension forces. There was no change in the amplitude of TMS-induced forces across graded motor imagery task. These results support the notion that the same neural correlates for actual movement could be selectively activated during motor imagery of the same movement, but demonstrated that the magnitude of isometric force could not be mentally simulated.     

Contractile properties of cat motor units enlarged by motoneurone sprouting

Summary The fast-twitch flexor digitorum longus (FDL) muscle was partially denervated by a unilateral section of the L₇ ventral root. After approximately 100 days the isometric, force-velocity and histochemical properties of single motor units from the partially denervated muscle were determined. The diameter of the nerve axons supplying the muscle were also examined. The extrapolated maximum speed of sarcomere shortening (V_max) of the enlarged motor units was significantly less than that of the control whole muscle V_max. It is concluded that the motoneurone is capable of expanding its territory and therefore its tension generating capability without any apparent change in axonal diameter. In addition, the intrinsic properties of the muscle fibres are altered.     

Relations between identified tendon organs and motor units in the medial gastrocnemius muscle of the cat

Summary In one medial gastrocnemius muscle of each of several cats, the response was recorded of a single tendon organ to the contraction of a single motor unit which strongly excited the receptor. The motor unit was depleted of its glycogen and the depleted muscle fibres identified in PAS-stained transverse sections. The site of maximum tendon organ sensitivity was marked and the tendon organ identified in the same sections. Five pairs of tendon organs and motor units were studied completely. Each tendon organ was found to have one or two (mean 1.6) depleted muscle fibres attached to it, included in the bundle of fibres attached to the end (mean no. 14.4) and side (mean no. 5.6) of the tendon organ. A correlation was found between tendon organ discharge rate and the tension calculated from cross-sectional area measurements of the depleted muscle fibres attached to the tendon organ, with variation between individual pairs of tendon organs and motor units. One estimate of the average sensitivity of the sample was 28 imp/s/mN. A nearly linear discharge rate vs. tension relation was found for single tendon organ and motor unit pairs when tension was graded during a series of fatiguing contractions. Under these conditions the sensitivity, measured as the slope of the relation between discharge rate and motor unit tension recorded at the common tendon, varied between 0.11 and 0.30 imp/s/mN for 6 pairs.     

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.     

Mechanical properties of single motor units in the rabbit masseter muscle as a function of jaw position

Positions and contractile properties of rabbit masseter motor units were investigated at different jaw gapes. Twitch responses were measured at gapes ranging from dental occlusion (0 degree) to maximum opening (21 degrees), in steps of 3 degrees. The twitches were elicited by stimulating motoneurons extracellularly in the trigeminal motor nucleus. The units appeared to produce a large variety of force vectors. On average motor units in the deep parts of the masseter produced considerably less twitch force (average: 25-30 mN) than those in the superficial parts (average: 45-50 mN) and anteriorly located motor units were slower than posteriorly located units. With an increase of jaw angle, twitches became slower, reflected by an increase (30%) of the twitch contraction time. Most motor units had a parabolic-like active jaw angle-force relationship. A large variation in the shape of the curves was found. The average optimum jaw angle was reached at 12 degrees jaw opening. In general, force output was relatively low (20-60% of maximum force) at occlusion and relatively high (60-100% of maximum force) at maximal jaw opening. Anteriorly and posteriorly located motor units differed significantly in their angle-force curves. Anteriorly located motor units produced less relative force at occlusion, showed a steeper increase of force with an increase of jaw angle, reached maximum force at larger jaw angles and produced larger forces at maximum jaw opening. The larger force changes in the more anterior units are probably related to their longer distance from the axis of jaw rotation. The large variability of motor unit properties and angle-force curves suggests that a fine gradation of both force magnitude and direction is possible within the masseter and that the angle-force curve of the whole muscle or of whole muscle parts is broader than that of individual motor units. This broadening may be considered as a mechanism to sustain active muscle force throughout a large movement range.     

Distribution of physiological types of motor units in the cat peroneus tertius muscle

Summary Motor units of the cat peroneus tertius muscle were systematically analyzed using the criteria established by Burke et al. (1973). On the basis of their speed of contraction and resistance to fatigue, 121 (97%) of 125 motor units examined in ten adult cats could be classified as belonging to one of four types: fast-fatiguable (FF), fast-resistant (FR), fast-intermediate (FI), and slow-resistant (S).Peroneus tertius was found to contain 30% FF motor units, 9% FI units, 39% FR units, and 22% S units. Contraction times of fast motor units (FF, FR, and FI) ranged from 15 to 27 ms and those of S units from 26 to 42 ms. The mean tetanic tensions were 37 g for FF units, 29 g for FI units, 7.5 g for FR units, and 1.1 g for S units.Fast motor units displayed considerable post-tetanic potentiation of twitch tension. Under similar conditions of stimulation, FF units appeared able to potentiate more and faster than FR units.Supported in part by the Fondation pour la Recherche Médicale Française     

Effects of activation rate on contractile properties of rabbit masseter motor units

Force-frequency curves of rabbit masseter motor units ( n=20) were determined, in order to study the capacity of these motor units for rate gradation and to establish the relationship between twitch contraction time (TCT) and the shape of the curves. Motor unit force responses were elicited by stimulating motoneurons in the trigeminal motor nucleus extracellularly. A sequence of pulse trains with increasing frequency rates was followed by trains with decreasing frequency rates. All motor units were classified as fast (F) units. The ascending force-frequency curves showed a distinct sigmoid appearance; the descending curves were shifted toward lower stimulation rates. The position and shape of the force-frequency curves related significantly to the TCT. The curves of slower units were located at lower frequencies and had a larger inclination. In addition, slower units had a lower fusion frequency and a larger twitch-tetanus ratio. Hence, slower units started to fuse and reached maximum force at lower stimulation rates than faster units and needed a smaller change in simulation frequency to achieve the same relative force. It can be concluded that the capacity for rate gradation differs between rabbit masseter motor units and that the TCT is a determinant for the position and shape of the force-frequency curves.     

In situ measurement of the innervation ratio of motor units in human muscles

Summary Volume conduction measurements were carried out on the brachial biceps, tibialis anterior and deltoid, in normal human subjects. Attenuation constant K and time constant t of the muscle tissue transfer function were measured, and the average electrode uptake area calculated for the three muscles. The average number of muscle fibres in the motor unit, i.e., the innervation ratio, was calculated from the electrode uptake area, data on the motor unit territory, and measurements of fibre density. The innervation ratios for the brachial biceps, tibialis anterior and deltoid were 209, 329, and 239 fibres, respectively. It was found that the anatomical scatter of fibres belonging to the same motor unit was smaller in brachial biceps than in tibialis anterior, whereas the electrophysiological fibre density was higher in tibialis anterior. The implications of these findings for the interpretation of normal and abnormal electromyographic findings are discussed.     

Non-invasive method to detect motor unit contractile properties and conduction velocity in human vastus lateralis muscle

The contractile properties and conduction velocity of motor units are estimated by using surface array electrodes during voluntary isometric contractions of the human vastus lateralis muscle. The subjects develop and maintain sufficient force to steadily discharge a given motor unit, assisted by visual feedback from an oscilloscope. The torque curve developed around the knee joint is triggered by an individual motor unit and averaged. 31 motor units in five subects are studied. The twitch tension detected ranges from 3 to 27 m Nm with a mean of 12·3 m Nm. The threshold force ranges from 1·88 to 10·12 Nm with a mean of 5·48 Nm, which is 3% of the maximal voluntary contraction. The rise time ranges from 56 to 106 ms with a mean of 83 ms. The mean value of conduction velocity is 4·64 m s⁻¹. The twitch tension is positively correlated to the threshold force (r=0·839, p<0.01), but has no relation to the other parameters. It is concluded that the use of non-invasive surface array electrodes provides the contractile properties of motor units and muscle fibre conduction velocity during weak contractions.     

Differential impairments in reaching and grasping produced by local inactivation within the forelimb representation of the motor cortex in the cat

This study analyzed changes in the performance of a reaching task and its adaptive modification produced by reversible inactivation of three sites within the forelimb representation of the motor cortex (MCx, area 4) in five cats by microinjections of muscimol. Two sites were located in the lateral MCx, rostral (RL-MCx) and caudal (CL-MCx) to the end of the cruciate sulcus, where intracortical microstimulation (ICMS) produced contraction of the most distal muscles. The third site was located more medially, in the anterior sigmoid gyrus (RM-MCx) where ICMS primarily produced contraction of more proximal muscles. The task required the animals to reach into a horizontal target well, located in front of them at one of three possible heights, to grasp and retrieve a small piece of food. The height of the reach was primarily achieved by elbow flexion. Grasping consisted primarily of digit flexion, and food retrieval consisted of forearm supination and shoulder extension. In some blocks of trials, an obstacle was placed in the path of the limb to assess the animal's ability to adaptively adjust the kinematic characteristics of their response trajectory. In normal animals, contact with the bar on the first trial triggered a corrective response at short latency that allowed the paw to circumvent the bar. On all subsequent trials, the trajectory was adapted to prevent contact with the obstacle, with a safety margin of about 1 cm. Inactivation at all sites produced a slowing of movement, a protracted and extended forelimb posture, and increased variability of initial limb position. In addition, inactivation of RL-MCx immediately produced systematic reaching errors, consisting of hypermetric movements, as well as impaired grasping and food retrieval. The degree of hypermetria was similar for all target heights and was not associated with alterations in trajectory control. During inactivation, animals did not compensate for the hypermetria by reducing paw path elevation, suggesting a defect in kinematic planning or in adaptive control. This was confirmed by finding that trajectory adaptation to avoid bar contact was impaired during RL-MCx inactivation. The short latency corrective response, triggered by contact of the limb with the obstacle was, however, preserved. Inactivation of CL-MCx did not impair aiming, grasping, or adaptation immediately after injection. However, impairments occurred after about 1 h postinjection, and at that time mimicked the effects of RL-MCx inactivation. This delay suggests that the drug was acting indirectly on the RL-MCx. Inactivation of RM-MCx did not impair the control of distal muscles, but the reaches became hypometric. The hypometria was greater for higher targets, suggesting that it resulted from weakness. Our results suggest that both rostral regions of the forelimb area of MCx play a more important role in the planning and execution of the prehension response than the caudal portion. We hypothesize that (1) the slowing of movement, forelimb postural changes, hypometria, and grasping and food retrieval impairments are due to defective control of muscles represented locally at each site in MCx and that (2) aiming and adaptation defects, which are produced only by RL-MCx inactivation, result from disruption of integrative mechanisms underlying sensorimotor transformations that normally assure movement accuracy.     

Task-related coding of stimulus and response in cat motor cortex

Summary In a previous study in the cat, we have reported that motor cortex neurons discharging before the initiation of an aimed forearm response (lead cells) are better timed to movement of a display (stimulus) than to the response. The present study was done to distinguish the coding of stimulus and response features in the discharge patterns of such early activity in motor cortex. Single neurons were recorded in the arm area of motor cortex in three cats performing the same pair of responses (forearm flexion and extension) but to display movements in either of the two directions by changing display polarity. The modulation of lead cell activity was contingent on the occurrence of the learned motor response and timed to the stimulus in all conditions. The majority of lead cells (88%, n = 50) fell into one of two distinct classes. In one class of neurons, force-direction (56%, n = 32), activity was contingent on a single direction of forelimb response (flexion or extension) and was thus independent of the direction of the display stimulus. The only muscles whose patterns matched the activity of this class of response-related neurons were forelimb flexors and extensors. In these neurons, the onset of modulation was timed to one or the other of the two stimuli according to the stimulus direction which elicited the appropriate response. Thus, the display-related input to these neurons varied according to the response required. In the second class of neurons, stimulus-direction (32%, n = 18), modulation was associated with a specific stimulus direction rather than the response direction. The pattern of activity of these neurons was similar to the pattern of EMG signals of shoulder and neck muscles during the different task conditions. The contraction of proximal and axial muscles corresponded to a second response elicited by the stimulus, namely attempts at head rotation towards the moving display and was independent of the conditioned forelimb response in both time of onset and direction. To test the possibility that stimulus-direction neurons participated in the control of head rotation we trained two of the animals to also produce isometric changes in neck torque in the direction of the moving display without making the forelimb response. The activity of stimulus-direction neurons was similarly modulated during performance of the neck task. By contrast, force-direction neurons examined during the neck task were either unmodulated or discharged after the neck response. These data suggest that force-direction neurons participate in response initiation and that their activity is triggered by stimuli specific for the task. The reorganization of the inputs to motor cortex is likely to result from gating mechanisms associated with behavioral set. Such neural gates could provide for the efficient transfer of any member of an array of behaviorally relevant stimuli to restricted sectors of the somatotopically organized motor areas.     

Different effects of physical training on the morphology of motor nerve terminals in the rat extensor digitorum longus and soleus muscles

Summary The morphology of nerve terminals in the rat extensor digitorum longus and soleus muscles was studied with light microscopy in 13-week-old male animals after 6 weeks of treadmill running and compared with data from untrained controls. The terminals were stained with methylene blue. Physical training tended to increase the area and length of the nerve terminals in relation to the corresponding muscle fiber diameter, and to reduce the density of nerve terminal varicosities, but significant differences between the trained group and the control group were obtained only in the extensor digitorum longus muscle. The different degrees of effect on the nerve terminals in the two muscles may be due to different abilities to respond to the training, but may also be due to differences in work load caused by the training. The effect of training on extensor digitorum longus junctions may reflect some transformation from fast to slow morphological characteristics.     

Automatic postural responses in the cat: responses of proximal and distal hindlimb muscles to drop of support from a single hind- or forelimb

Summary Cats respond to drop of the support from beneath a single limb with the diagonal stance response (Coulmance et al. 1979). They load the limbs on the diagonal opposite to the one containing the dropped limb and unload the third supporting limb in the diagonal containing the dropped limb. Characteristic biomechanical delays in limb motion and in vertical force changes imposed upon the limbs are observed. These delays range from 30 to 45 ms, depending upon the location of the dropped limb. This study describes the kinematics of the diagonal stance response and the activation of selected agonist-antagonist muscle pairs acting on the joints of the hindlimb during the response. Proximal and distal hindlimb muscles respond to perturbations in groups that are appropriate to the vertical forces imposed upon the limb. When the hindlimb containing the recording electrodes is loaded by drop of the contralateral hindlimb or the ipsilateral forelimb medium latency (25–45 ms) EMG responses occur in the extensors. This response serves to stiffen the limb against the increased vertical force of loading. A similar response is observed when the hindlimb is reloaded after being dropped. In this case, however, short latency responses precede the medium latency responses in muscles that are passively stretched by the limb drop. When drop of the diagonal forelimb unloads the hindlimb containing the electrodes, medium latency responses are observed in the distal hindlimb flexors, which indicates that the unloading is evoked in part by active lifting of the limb. In most cases, the medium latency responses precede or are coincident with the changes in force imposed on the limb, suggesting that the observed responses are centrally programmed.     

Integration in descending motor pathways controlling the forelimb in the cat

Summary With intracellular recording from forelimb motoneurones the spatial facilitation technique has been used to investigate interaction between descending pathways and forelimb afferents.As previously shown for the hindlimb, pyramidal volleys effectively facilitate interneuronal transmission in reflex pathways from different primary afferents. Evidence is presented suggesting disynaptic excitation from corticospinal fibres of interneurones in the reciprocal Ia inhibitory pathway. Interneurones of other reflex pathways from group I muscle afferents receive monosynaptic pyramidal excitation. During pyramidal facilitation volleys in cutaneous afferents may evoke PSPs in motoneurones after a central delay of 1.3 ms suggesting that the minimal linkage is disynaptic.Information regarding convergence on the neurones intercalated in the disynaptic cortico-motoneuronal pathway was obtained by investigating the effect from primary afferents and from other descending pathways on the disynaptic pyramidal EPSPs. Volleys in cutaneous and group I muscle afferents facilitate transmission in the disynaptic cortico-motoneuronal pathway with a time course showing oligosynaptic (probably monosynaptic) action on the intercalated neurone. Rubrospinal volleys likewise effectively facilitate disynaptic cortico-motoneuronal transmission with a time course showing monosynaptic action on the intercalated neurone. Spatial facilitation experiments involving three tests revealed that those intercalated neurones which receive convergent monosynaptic excitation from corticospinal and rubrospinal fibres are excited also from cutaneous forelimb afferents.Disynaptic cortico-motoneuronal transmission was also monosynaptically facilitated by stimuli in the dorsal mesencephalic tegmentum probably activating tectospinal fibres. Disynaptic, presumed tectospinal EPSPs were facilitated from cutaneous forelimb afferents.The convergence onto the neurones intercalated in the disynaptic excitatory cortico-motoneuronal pathway suggests that these neurones integrate the activity in different descending pathways and primary forelimb afferents.Supported by the Deutsche ForschungsgemeinschaftIBRO/UNESCO Fellow     

Changes in rat plantaris motor unit profiles with advanced age

The physiological characteristics of single motor units in rat plantaris muscles were determined in situ, for young adult (3 months) and very old (30–34 months) Fischer 344 rats. Old muscles generated 43% less tetanic force (P₀) per gram. Motor units classified as “slow”, using criteria of fatigue resistance and “sag” during unfused tetani, had a mean P₀ which was 255% of that in young muscles, while fast motor units were similar in P₀ in the two groups. Estimates were made of motor unit numbers using whole muscle and mean motor unit P₀ values. The typical young plantaris contained 48 units, of which 5–6 were slow, while old plantaris contained 29 units, of which 11 were slow. In spite of this large increase in slow motor unit presence (increased mean motor unit P₀, plus increased number) in old muscles, a comparatively modest (72%) increase occurred in the muscle cross-section occupied by histochemically demonstrated slow fibres. During senescence, there occurs a loss in muscle tetanic force capability which is accompanied by a loss of motor units and a reorganization of the remaining motor unit profile. An increase in slow motor unit number and size with advancing age can evidently occur without concomitant histochemical changes. Motor units do not “dedifferentiate”, but maintain their physiological distinctiveness into very old age.