Motor unit recruitment for dynamic tasks: current understanding and future directions

Skeletal muscle contains many muscle fibres that are functionally grouped into motor units. For any motor task there are many possible combinations of motor units that could be recruited and it has been proposed that a simple rule, the ‘size principle’, governs the selection of motor units recruited for different contractions. Motor units can be characterised by their different contractile, energetic and fatigue properties and it is important that the selection of motor units recruited for given movements allows units with the appropriate properties to be activated. Here we review what is currently understood about motor unit recruitment patterns, and assess how different recruitment patterns are more or less appropriate for different movement tasks. During natural movements the motor unit recruitment patterns vary (not always holding to the size principle) and it is proposed that motor unit recruitment is likely related to the mechanical function of the muscles. Many factors such as mechanics, sensory feedback, and central control influence recruitment patterns and consequently an integrative approach (rather than reductionist) is required to understand how recruitment is controlled during different movement tasks. Currently, the best way to achieve this is through in vivo studies that relate recruitment to mechanics and behaviour. Various methods for determining motor unit recruitment patterns are discussed, in particular the recent wavelet-analysis approaches that have allowed motor unit recruitment to be assessed during natural movements. Directions for future studies into motor recruitment within and between functional task groups and muscle compartments are suggested.     

Hill muscle model errors during movement are greatest within the physiologically relevant range of motor unit firing rates

This study evaluated the accuracy of Hill-type muscle models during movement. Hill-type models are ubiquitous in biomechanical simulations. They are attractive because of their computational simplicity and close relation to commonly measured experimental variables, but there have been surprisingly few experimental validations of these models during functionally relevant conditions. Our hypothesis was that model errors during movement are largest at the low motor unit firing rates most relevant to normal movement conditions. This hypothesis was evaluated in the cat soleus muscle activated either by electrical stimulation at physiological rates or via the crossed-extension reflex (CXR) thereby obtaining normal patterns of motor unit recruitment and rate modulation. These activation paradigms were applied during continuous movements approximately matched to locomotor length changes. The resulting muscle force was modeled using a common Hill model incorporating independent activation, tetanic length-tension and tetanic force-velocity properties. Errors for this model were greatest for stimulation rates between approximately 10-20Hz. Errors were especially large for muscles activated via the CXR, where most motor units appear to fire within this range. For large muscle excursions, such as those seen during normal locomotion, the errors for naturally activated muscle typically exceeded 50%, supporting our hypothesis and indicating that the Hill model is not appropriate for these conditions. Subsequent analysis suggested that model errors were due to the common Hill model's inability to account for the coupling between muscle activation and force-velocity properties that is most prevalent at the low motor unit firing rates relevant to normal activation.     

The effect of fast and slow motor unit activation on whole-muscle mechanical performance: the size principle may not pose a mechanical paradox

The output of skeletal muscle can be varied by selectively recruiting different motor units. However, our knowledge of muscle function is largely derived from muscle in which all motor units are activated. This discrepancy may limit our understanding of in vivo muscle function. Hence, this study aimed to characterize the mechanical properties of muscle with different motor unit activation. We determined the isometric properties and isotonic force–velocity relationship of rat plantaris muscles in situ with all of the muscle active, 30% of the muscle containing predominately slower motor units active or 20% of the muscle containing predominately faster motor units active. There was a significant effect of active motor unit type on isometric force rise time (p < 0.001) and the force–velocity relationship (p < 0.001). Surprisingly, force rise time was longer and maximum shortening velocity higher when all motor units were active than when either fast or slow motor units were selectively activated. We propose this is due to the greater relative effects of factors such as series compliance and muscle resistance to shortening during sub-maximal contractions. The findings presented here suggest that recruitment according to the size principle, where slow motor units are activated first and faster ones recruited as demand increases, may not pose a mechanical paradox, as has been previously suggested.     

Motor unit activation patterns during concentric wrist flexion in humans with different muscle fibre composition

Muscle activity was recorded from the flexor carpi radialis muscle during static and dynamic-concentric wrist flexion in six subjects, who had exhibited large differences in histochemically identified muscle fibre composition. Motor unit recruitment patterns were identified by sampling 310 motor units and counting firing rates in pulses per second (pps). During concentric wrist flexion at 30% of maximal exercise intensity the mean firing rate was 27 (SD 13) pps. This was around twice the value of 12 (SD 5) pps recorded during sustained static contraction at 30% of maximal voluntary contraction, despite a larger absolute force level during the static contraction. A similar pattern of higher firing rates during dynamic exercise was seen when concentric wrist flexion at 60% of maximal exercise intensity [30 (SD 14) pps] was compared with sustained static contraction at 60% of maximal voluntary contraction [19 (SD 8) pps]. The increase in dynamic exercise intensity was accomplished by recruitment of additional motor units rather than by increasing the firing rate as during static contractions. No difference in mean firing rates was found among subjects with different muscle fibre composition, who had previously exhibited marked differences in metabolic response during corresponding dynamic contractions. It was concluded that during submaximal dynamic contractions motor unit firing rate cannot be deduced from observations during static contractions and that muscle fibre composition may play a minor role. Accepted: 5 May 1998     

A Neuro-Mechanical Model of a Single Leg Joint Highlighting the Basic Physiological Role of Fast and Slow Muscle Fibres of an Insect Muscle System

In legged animals, the muscle system has a dual function: to produce forces and torques necessary to move the limbs in a systematic way, and to maintain the body in a static position. These two functions are performed by the contribution of specialized motor units, i.e. motoneurons driving sets of specialized muscle fibres. With reference to their overall contraction and metabolic properties they are called fast and slow muscle fibres and can be found ubiquitously in skeletal muscles. Both fibre types are active during stepping, but only the slow ones maintain the posture of the body. From these findings, the general hypothesis on a functional segregation between both fibre types and their neuronal control has arisen. Earlier muscle models did not fully take this aspect into account. They either focused on certain aspects of muscular function or were developed to describe specific behaviours only. By contrast, our neuro-mechanical model is more general as it allows functionally to differentiate between static and dynamic aspects of movement control. It does so by including both muscle fibre types and separate motoneuron drives. Our model helps to gain a deeper insight into how the nervous system might combine neuronal control of locomotion and posture. It predicts that (1) positioning the leg at a specific retraction angle in steady state is most likely due to the extent of recruitment of slow muscle fibres and not to the force developed in the individual fibres of the antagonistic muscles; (2) the fast muscle fibres of antagonistic muscles contract alternately during stepping, while co-contraction of the slow muscle fibres takes place during steady state; (3) there are several possible ways of transition between movement and steady state of the leg achieved by varying the time course of recruitment of the fibres in the participating muscles.     

The resilience of the size principle in the organization of motor unit properties in normal and reinnervated adult skeletal muscles

Henneman's size principle relates the input and output properties of motoneurons and their muscle fibers to size and is the basis for size-ordered activation or recruitment of motor units during movement. After nerve injury and surgical repair, the relationship between motoneuron size and the number and size of the muscle fibers that the motoneuron reinnervates is initially lost but returns with time, irrespective of whether the muscles are self- or cross-reinnervated by the regenerated axons. Although the return of the size relationships was initially attributed to the recovery of the cross-sectional area of the reinnervated muscle fibers and their force per fiber, direct enumeration of the innervation ratio and the number of muscle fibers per motoneuron demonstrated that a size-dependent branching of axons accounts for the size relationships in normal muscle, as suggested by Henneman and his colleagues. This same size-dependent branching accounts for the rematching of motoneuron size and muscle unit size in reinnervated muscles. Experiments were carried out to determine whether the daily amount of neuromuscular activation of motor units accounts for the size-dependent organization and reorganization of motor unit properties. The normal size-dependent matching of motoneurons and their muscle units with respect to the numbers of muscle fibers per motoneuron was unaltered by synchronous activation of all of the motor units with the same daily activity. Hence, the restored size relationships and rematching of motoneuron and muscle unit properties after nerve injuries and muscle reinnervation sustain the normal gradation of muscle force during movement by size-ordered recruitment of motor units and the process of rate coding of action potentials. Dynamic modulation of size of muscle fibers and their contractile speed and endurance by neuromuscular activity allows for neuromuscular adaptation in the context of the sustained organization of the neuromuscular system according to the size principle.     

Summation of elementary phonomyograms during isometric twitches in humans

To study its summation principle, the phonomyogram (PMG) from the first interosseus dorsalis muscle was recorded in five subjects during single twitches evoked by electrical stimulation over the motor point. By increasing the current pulse from threshold to maximal intensity, PMG amplitude increased linearly with motor unit recruitment. The twitch amplitude-intensity relationship was also linear. The PMG amplitude was therefore linearly related to the external force. For all these relationships highly significant correlation coefficients were found. These relationships were interpreted as being a consequence of an orderly recruitment, although, contrary to what happens during voluntary contraction, the largest and strongest motor units were recruited before the smallest and weakest ones during axon electrical stimulation. The PMG onset always preceded twitch onsets as indicated by latency measurements [mean 3.2 (SD 1.3) ms versus 11.5 (SD 3.9) ms, respectively]. Moreover, PMG and twitch latencies may have been significantly reduced by recruitment, suggesting that orderly recruitment influenced both PMG amplitude and occurrence. These results were interpreted as being the result of the summation of elementary PMG from every contracting motor unit and the stiffness change of the muscle medium occurring with recruitment. Accepted: 25 August 1997     

Catchlike property in human adductor pollicis muscle

The "catchlike" property is defined as the dramatic force increase in skeletal muscles when a single pulse is added at the onset of a sub-tetanic low-frequency stimulation train. This property has been observed in single motor units, whole animal and human muscles. It is an inherent property of muscle fibres and is not related to an increase in motor unit recruitment. Despite an abundance of observations, its origin remains unclear. The aim of this study was to induce the catchlike property in human adductor pollicis and identify its possible origin. Thumb adduction forces were measured using ulnar nerve electrical stimulation at 10Hz for reference trains (RTs) with one extra pulse 8ms after the first stimulation pulse for the experimental trains (ETs). Tests were performed at two muscle length and three stimulation levels and muscle stiffness and potentiation were quantified for all test conditions. The ETs showed higher forces and greater rates of force increase than the RTs. In addition, force increase was more pronounced at short compared to long muscle length, but no differences were found in force increase for the three stimulation levels. Furthermore, potentiation and stiffness were similar across all experimental conditions. Together, these results suggest that the increase in force associated with the catchlike property is neither caused by an increased proportion of attached cross-bridges nor potentiation of the muscle, but appears to be muscle length dependent and present in both slow and fast motor units.     

Inference of motor unit recruitment order in voluntary and electrically elicited contractions

The relationship between surface myoelectric signal parameters and the level of voluntary or electrically elicited contractions was studied in 32 experiments on the tibialis anterior muscle of 22 healthy human subjects. Contractions were performed at 20 and 80% of the maximum voluntary contraction torque. Two levels of stimulation current were used, yielding, respectively, a maximum M wave and an M wave approximately 30% of the maximum. A four-bar electrode probe was used to detect single- and double-differential signals from which mean and median frequency of the power spectrum and average muscle fiber conduction velocity were estimated. Measurements obtained from voluntary contractions showed a positive correlation between contraction levels and both conduction velocity and spectral parameters. Conduction velocity increased by 21.2 +/- 10.9% when voluntary contraction level increased from 20 to 80% of the maximal value. Spectral parameters increased by similar amounts. Tetanic electrical stimulation was applied to a muscle motor point for 20 s via surface electrodes. Rectangular current pulses with 0.1-ms width and frequencies of 20, 25, 30, 35, and 40 Hz were used. Four types of behavior were observed with increasing stimulation level: 1) the two spectral parameters and conduction velocity both increased with stimulation in 15 experiments, 2) the two spectral parameters decreased and conduction velocity increased in 8 experiments, 3) the two spectral parameters and conduction velocity both decreased in 6 experiments, and 4) the two spectral parameters increased and conduction velocity decreased in 3 experiments. Conduction velocity increased with increasing stimulation current in 72% of the experiments, indicating a recruitment order similar to that of voluntary contractions, although it decreased in the other 28% of the cases, indicating a reverse order of recruitment. Contrary to what is observed in direct stimulation of nerves, motor units are not in general recruited in reverse order of size during electrical stimulation of a muscle motor point. This discrepancy may be the result of geometric factors or a lack of correlation between axonal branch diameter and the diameter of the parent motoneuron axon. Changes of conduction velocity and spectral parameters in opposite directions may be the result of the combined effect of the motor unit recruitment order and of the different tissue filtering function associated with the geometric location of the recruited motor units within the muscle.     

The inverse dynamics problem of neuromuscular control

The myoskeletal inverse dynamics problem and the myocybernetic control inverse problem were investigated with respect to their ill-posedness. The first problem consists of finding from observed experimental motion and reaction force data the resultant muscle moments that generated the observed motion, while the second aims at finding the corresponding neural controls. It is shown that both problems belong to the class of incorrectly posed (ill-posed) problems that, by definition, do not possess unique solutions. To illustrate this point, results of a forward dynamics simulation of a comprehensive neuromusculoskeletal model of the human body are presented. These results demonstrate that fairly chaotic neural control perturbations have very little influence on the resulting motion trajectory, at least in the present example. While a regularization procedure may be applied to solve successfully the myoskeletal inverse dynamics problem, the myocybernetic control inverse problem is unsolvable. The latter fact has the important implication that, based on the somatosensory inputs it receives, the pars intermedia in the cerebellum is not able to control individual motor unit stimulation rates and recruitment patterns but only whole muscles by means of a single compound signal. The latter signal is identified as the “common drive.” Presumably at the spinal level, special neural circuits are used to decompose the common drive signal into motor unit recruitment patterns and stimulation rates that are specific for a given mode of contraction and probably obey certain optimality principles. Received: 26 February 1999 / Accepted in revised form: 11 June 1999     

A teleological explanation of Weber's law and the motor unit size law

It is shown that the Weber-Fechner law. which relates the response of a sensory biosystem to the intensity of the input stimulus, can be derived from a teleological principle of minimum transentropy (maximal noise reduction) provided the relative mean fluctuation (coefficient of variation) of the input intensity can be assumed to be (approximately) constant for all feasible mean input intensities. A law is then deduced from experimental results which quantifies the relationship existing between the relative amount of activated muscle mass and the “size” (which term is clearly defined) of a newly recruited motor unit. This law is found to be formally equivalent to the Weber-Fechner law when applied to motor unit recruitment. It is then shown that, in general, the ratio of the force increment upon recruitment, to the present force output does not obey Weber's law. Finally, it is proved that the “motor unit size law” as derived in this paper implies a fixed sequential order in the recruitment of motor units and that it may be viewed as the realization, by the mammalian neuromuscular system, of a general principle of maximum grading sensitivity.     

Specific contributions of various muscle fibre types to human muscle performance: an in vitro study.

Human skeletal muscle fibres can be divided in five groups: 1, 1-2A, 2A, 2A-2B and 2B, by using myosin heavy chain (MHC) isoforms as molecular markers. This study aimed to define the contribution of each fibre type to the contractile performance of human muscles. Single fibre segments were dissected from bioptic samples of vastus lateralis and chemically skinned. Force-velocity properties, including isometric tension (P0), maximal shortening velocity (Vmax), maximum power output (Wmax) and the velocity at which Wmax is reached (Vopt), were determined at maximum calcium activation. Among these parameters Wmax showed the largest range of variation: about nine times between 2B and slow fibres. Vopt also showed large (about four times) and significant variations between fibre types. Force development at submaximum calcium activation was studied and force-pCa curves were obtained for each fibre type. Calcium sensitivity was greater in 2B than in 2A and in slow fibres. The slope of the force-pCa curve was greater in fast than in slow fibres. At the end of the experiment the MHC isoform composition of each fibre segment was determined by gel electrophoresis. The functional properties of each fibre type are discussed in the light of the motor unit recruitment mechanism to understand their possible physiological role.     

Muscle fibre growth dynamics in diploid and triploid rainbow trout

The effect of triploidy on muscle fibre growth was determined by comparing hyperplasia and hypertrophy of white muscle fibres in all-female, diploid and triploid rainbow trout Oncorhynchus mykiss (100–400 mm total length). Conventional morphometry and protein and DNA concentrations were used to assess muscle fibre hyperplasia and hypertrophy in white muscle samples derived from an anterio-dorsal location. Muscle fibre distributions were significantly different between triploids and diploids in trout <300 mm. The proportion of fibres <20 μm was higher in diploids than in triploids and the proportion of fibres in the 20–40 μm category was higher in triploids than in diploids. This indicates that the hyperplastic fibres of triploids are larger than those of diploids. Larger hyperplastic fibres in triploids are probably due to the combined effect of increased nuclear size in triploids and the relatively high nucleus: cell ratio observed in small muscle fibres. These larger fibres may be less favourable to cellular metabolic exchange because of their smaller surface area to volume ratios, and perhaps account for reduced viability and growth observed in triploids during early life stages. On the other hand, the lack of difference in the distribution of fibres <20 μm between diploids and triploids at larger body size ranges (301–400 mm) imply that triploid trout may have higher rates of new fibre recruitment and growth capacity at these sizes. There was no difference between diploid and triploid trout in the mean size of muscle fibres; however, the number of fibres per unit area was reduced by 10% in triploids. No differences were observed in protein or DNA concentrations in muscle tissues between the two genetic groups. Since triploid nuclei have 1·5 times more DNA than diploid nuclei, this deviation from the expected muscle DNA concentration (1·3–1·4 times more DNA in triploids when the 10% reduction in fibre density is considered) suggests that the number of nuclei per muscle fibre is reduced. In both diploids and triploids, mean fibre size increased with body length while fibre density decreased. Similarly, protein concentration in the muscle tissue increased and DNA concentration declined with increasing body length. Protein/DNA ratio was strongly and positively correlated with fibre size. These results demonstrate that changes in DNA and protein concentrations can be used to assess hyperplasia and hypertrophy in muscle tissues. However, the morphometric procedure provides better insight into muscle fibre growth as it enables the direct visualization and analysis of muscle fibre distribution patterns.     

Simulation of motor unit recruitment and microvascular unit perfusion: spatial considerations

Fuglevand, Andrew J., and Steven S. Segal. Simulationof motor unit recruitment and microvascular unit perfusion: spatial considerations. J. Appl. Physiol.83(4): 1223-1234, 1997.Muscle fiber activity is the principalstimulus for increasing capillary perfusion during exercise. Thecontrol elements of perfusion, i.e., microvascular units (MVUs), supplyclusters of muscle fibers, whereas the control elements of contraction,i.e., motor units, are composed of fibers widely scattered throughoutmuscle. The purpose of this study was to examine how the discordantspatial domains of MVUs and motor units could influence the proportion of open capillaries (designated as perfusion) throughout a muscle crosssection. A computer model simulated the locations of perfused MVUs inresponse to the activation of up to 100 motor units in a muscle with40,000 fibers and a cross-sectional area of 100 mm². The simulation increasedcontraction intensity by progressive recruitment of motor units. Foreach step of motor unit recruitment, the percentage of active fibersand the number of perfused MVUs were determined for several conditions:1) motor unit fibers widely dispersed and motor unit territories randomly located (whichapproximates healthy human muscle),2) regionalized motor unitterritories, 3) reversed recruitmentorder of motor units, 4) denselyclustered motor unit fibers, and 5)increased size but decreased number of motor units. The simulationsindicated that the widespread dispersion of motor unit fibersfacilitates complete capillary (MVU) perfusion of muscle at low levelsof activity. The efficacy by which muscle fiber activity inducedperfusion was reduced 7- to 14-fold under conditions that decreased thedispersion of active fibers, increased the size of motor units, orreversed the sequence of motor unit recruitment. Such conditions aresimilar to those that arise in neuromuscular disorders, with aging, orduring electrical stimulation of muscle, respectively.

     

The effect of activity during early postnatal development on motor unit size

At early stages of neuromuscular development, motor unit territory is expanded, with each muscle fibre being supplied by several axons. During postnatal development, some synapses are eliminated, motor unit size decreases, and the adult distribution of motor unit sizes emerges. This process depends on activity, since it proceeds more rapidly when the nerve is activated and is slower when activity is reduced. Here we studied whether, in addition to influencing the rate of retraction of motor unit territory, activity during the critical period of development affects the final outcome of the distribution of motor unit sizes. The sciatic nerve of 8- to 12-day-old rats was stimulated daily. One week later the tension of the extensor digitorum longus muscle and that of its individual motor units was recorded. The sizes of individual motor units were calculated and compared with those from animals that received no stimulation. The distribution of motor unit sizes from stimulated muscles was not significantly different from those from control muscles. Therefore, we conclude that although activity increases the rate at which motor units attain their adult size, it does not influence the final outcome of motor unit size distribution.     

Morphological Characteristics of Motor Neurons Do Not Determine Their Relative Susceptibility to Degeneration in a Mouse Model of Severe Spinal Muscular Atrophy

Spinal muscular atrophy (SMA) is a leading genetic cause of infant mortality, resulting primarily from the degeneration and loss of lower motor neurons. Studies using mouse models of SMA have revealed widespread heterogeneity in the susceptibility of individual motor neurons to neurodegeneration, but the underlying reasons remain unclear. Data from related motor neuron diseases, such as amyotrophic lateral sclerosis (ALS), suggest that morphological properties of motor neurons may regulate susceptibility: in ALS larger motor units innervating fast-twitch muscles degenerate first. We therefore set out to determine whether intrinsic morphological characteristics of motor neurons influenced their relative vulnerability to SMA. Motor neuron vulnerability was mapped across 10 muscle groups in SMA mice. Neither the position of the muscle in the body, nor the fibre type of the muscle innervated, influenced susceptibility. Morphological properties of vulnerable and disease-resistant motor neurons were then determined from single motor units reconstructed in Thy.1-YFP-H mice. None of the parameters we investigated in healthy young adult mice – including motor unit size, motor unit arbor length, branching patterns, motor endplate size, developmental pruning and numbers of terminal Schwann cells at neuromuscular junctions - correlated with vulnerability. We conclude that morphological characteristics of motor neurons are not a major determinant of disease-susceptibility in SMA, in stark contrast to related forms of motor neuron disease such as ALS. This suggests that subtle molecular differences between motor neurons, or extrinsic factors arising from other cell types, are more likely to determine relative susceptibility in SMA.     

An oscillator theory of motor unit recruitment

The phenomenon of systematic recruitment of motor units with increasing demand load is usually explained by the size principle. Though this principle successfully explains the gain-related aspects of muscle force generation, it does not address the need for desynchronization of motor unit activities in order to produce a smooth tension profile at the level of whole muscle, while individual muscle fibers are "twitching." We propose an oscillator model of motor neurons in which a pool of motor neurons fires a bundle of muscle fibers. Although individual muscle fibers have a complicated tension profile, the tension produced by the entire bundle is regulated and follows a command signal accurately. This is shown to be possible because of uncorrelated activity produced by local inhibitory connections among motor neurons. Connections that produce synchronized oscillations result in uncontrolled contractions of the muscle. These results seem to suggest that while synchronized activity indicates pathology and disease, desynchronized activity is the precondition for normal muscle function. Physiological evidence for the proposed theory of motor unit synchronization is presented.     

Recruitment and maximal diameter of axial muscle fibres in teleosts and their relationship to somatic growth and ultimate size

Growth of white axial muscle fibres of ten species of freshwater teleosts from five families (Cyprinidae, Centrarchidae, Percidae, Salmonidae, Esocidae) possessing widely different growth rates and ultimate sizes have been studied. The dynamics of muscle increase (i.e. increase in fibre numbers and/or diameter) appears to determine the ability for rapid somatic growth and large ultimate size in teleosts. Thus, the largest and fastest growing species (smallmouth bass, lake whitefish, rainbow trout, muskellunge) show evidence of sustained recruitment of muscle fibres to a large size, in contrast to the smaller and slower growing species (bluntnose minnow, longnose dace). Pumpkinseed, bluegill and yellow perch are all intermediate in fibre growth dynamics, growth and ultimate size between the smaller and larger species. Moreover, the ability of teleosts to grow rapidly and attain a large ultimate size is dependent on the body length at which recruitment of new muscle fibres into the growing axial muscle ceases. The regression equation, y =– 0.29 + 2.26 ( x ), showing the relationship (r = 0.95) between ultimate body length, x , and fork length at cessation of recruitment, y , indicated that for these teleosts, recruitment tends to cease when the fork length reaches about 44% of the ultimate body length. Possible mechanisms to account for this relationship are proposed, and the role of the ultimate fibre diameter in posing limits to the ultimate size of the species is discussed.     

Orderly recruitment among motoneurons supplying different muscles.

Virtually all movements involve the recruitment of motor units from multiple muscles. Given the functional diversity of motor units (motoneurons and the muscle fibers they supply), the effective production of specific movements undoubtedly depends upon some principle(s) to organize the ensemble of active motor units. The principle acting to organize the recruitment of motor units within muscles is the size principle, whereby the first motor units to be recruited have the smallest values for axonal conduction velocity and contractile force, and are the slowest to contract and fatigue. Here we consider the possibility that the size principle applies in the recruitment of motor units across muscles, i.e., that regardless of their muscles of origin, active motor units are recruited in rank order, for example, from low to high conduction velocity. The benefits of orderly recruitment across muscles could be similar to the acknowledged advantages of orderly recruitment within muscles. One benefit is that the neural process involved in organizing active motor units would be simplified. In a muscle-based scheme, the size principle would organize only those motor units within individual muscles, leaving the nervous system with the additional task of coordinating the relative activities of motor units from different muscles. By contrast, in an ensemble-based scheme, orderly recruitment of all motor units according to the size principle would automatically coordinate motor units both within and across motor nuclei. Another potential benefit is the provision for movements with smooth trajectory, the result of interleaving the divergent torque contributions made by motor units from muscles that differ in their orientations about joints. Otherwise, if order were restricted within muscles, the torque trajectory of a joint would change unevenly as participating muscles begin contracting at different times and grade activity at different rates. These considerations support speculation that motor units recruited from co-contracting muscles are collectively recruited according to the size principle.     

Organization of skeletal muscle in the urodele Triturus cristatus: muscle fibre types and motor units

Four muscle fibre types are described in the biceps and extensor digitorum communis muscles of the newt forelimb. The histological criteria forming the basis for the distinctions include differential staining with p-phenylenediamine and succinate dehydrogenase histochemistry and electron microscopy. In addition, three distinctive motor unit types are described for the biceps muscle. These are fast units, slow units and intermediate units. The structure of muscle fibre and the physiological characteristics of muscle fibres belonging to each motor unit, have been correlated by using iontophoretic passage of Lucifer yellow into muscle fibres belonging to physiologically characterized motor units and their subsequent histological identification by the succinate dehydrogenase reaction. The three motor unit types correspond to slow muscle fibres, intermediate muscle fibres and two classes of fast muscle fibres.