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Much of the repertoire of muscle function performed in everyday life involves isotonic dynamic movements, either with or without an additional load, yet most studies of single motor units measure isometric forces. To assess the effects of muscle load on the contractile response, we measured the contractile properties of single motor units supplying the toe extensors, assessed by intraneural microstimulation of single human motor axons, in isotonic, loaded isotonic, and isometric conditions. Tungsten microelectrodes were inserted into the common peroneal nerve, and single motor axons (n = 10) supplying the long toe extensors were electrically stimulated through the microelectrode. Displacement was measured from the distal phalanx of the toe with either an angular displacement transducer for the unloaded (i.e., no additional load) and loaded (addition of a 4-g mass) isotonic conditions or a force transducer for the isometric conditions. Mean twitch profiles were measured at 1 Hz for all conditions: rise time, fall time, and duration were shortest for the unloaded isotonic conditions and longest for the isometric conditions. Peak displacements were lower in the loaded than unloaded isotonic conditions, and the half-maximal response in the loaded condition was achieved at lower frequencies than in the unloaded isotonic condition. We have shown that the contractile responses of single motor units supplying the human toe extensors are influenced by how they are measured: twitches are much slower when measured in loaded than unloaded isotonic conditions and slowest when measured in isometric conditions., (Copyright © 2015 the American Physiological Society.)

During locomotion, stepping over an obstacle under visual guidance is crucial to continuous safe walking. Studies of the role of the central nervous system in stepping movements have focused on cerebral cortical areas such as the primary motor cortex and posterior parietal cortex. There is speculation that the lateral cerebellum, which has strong anatomical connections with the cerebral cortex, also plays a key role in stepping movements over an obstacle, although this function of the lateral cerebellum has not yet been elucidated. Here we investigated the role of the lateral cerebellum during obstacle avoidance locomotion in rats with a lateral cerebellar lesion. A unilateral lesion in the lateral cerebellum did not affect limb movements during overground locomotion. Importantly, however, the lesioned animals showed overshooting of the toe trajectory specific to the leading forelimb ipsilateral to the lesion when stepping over an obstacle, and the peak toe position, in which the toe is maximally raised during stepping, shifted away from the upper edge of the obstacle. Recordings of EMG activity from elbow flexor and extensor muscles suggested that the overshooting toe trajectory in the ipsilateral leading forelimb possibly resulted from sustained elbow flexion and delayed elbow extension following prolonged activity of the biceps brachii. These results suggest that the lateral cerebellum specifically contributes to generating appropriate toe trajectories in the ipsilateral leading forelimb and to controlling related muscle activities in stepping over an obstacle, especially when accurate control of the distal extremity is achieved under visual guidance.

We investigate whether imagery of voluntary movements of different body parts activates somatotopical sections of the human motor cortices. We used functional magnetic resonance imaging to detect the cortical activity when 7 healthy subjects imagine performing repetitive (0.5-Hz) flexion/extension movements of the right fingers or right toes, or horizontal movements of the tongue. We also collected functional images when the subjects actually executed these movements and used these data to define somatotopical representations in the motor areas. In this study, we relate the functional activation maps to cytoarchitectural population maps of areas 4a, 4p, and 6 in the same standard anatomical space. The important novel findings are 1). that imagery of hand movements specifically activates the hand sections of the contralateral primary motor cortex (area 4a) and the contralateral dorsal premotor cortex (area 6) and a hand representation located in the caudal cingulate motor area and the most ventral part of the supplementary motor area; 2). that when imagining making foot movements, the foot zones of the posterior part of the contralateral supplementary motor area (area 6) and the contralateral primary motor cortex (area 4a) are active; and 3). that imagery of tongue movements activates the tongue region of the primary motor cortex and the premotor cortex bilaterally (areas 4a, 4p, and 6). These results demonstrate that imagery of action engages the somatotopically organized sections of the primary motor cortex in a systematic manner as well as activating some body-part-specific representations in the nonprimary motor areas. Thus the content of the mental motor image, in this case the body part, is reflected in the pattern of motor cortical activation.

To investigate the specificity of fusimotor (gamma) drive during locomotion, gamma-efferents were recorded from the flexor digitorum longus (FDL) and flexor hallucis longus (FHL) nerves in a decerebrate cat preparation. These nerves innervate hindlimb muscles that differ in some aspects of their mechanical action. For both FHL and FDL two stereotyped patterns of gamma activity were distinguished. Tonic units fired throughout the step cycle and had less modulation, but higher minimum rates, than phasic units, which were mainly recruited with ankle extensor [soleus (SOL)] electromyogram (EMG) activity. Differences in the relative timing of these patterns were apparent. In FHL the activity of phasic and most tonic neurons peaked after EMG onset. With FDL, tonic units generally reached maximum rate before, while phasic units peaked after, the beginning of EMG activity. During locomotion FHL and FDL alpha activity were rhythmically recruited with SOL. However, consistent with previous reports, FHL and FDL differed in their patterns of alpha activity. FHL was stereotyped while FDL was variable. Both FHL and FDL had activity related to ankle extensor EMG, but only FDL exhibited a peak around the end of this phase. No corresponding gamma activity was observed in FDL. In conclusion, 1) FHL and FDL received tonic and phasic fusimotor drive; 2) there was no alpha/gamma linkage for the late FDL alpha burst; 3) phasic gamma-efferents in both muscles received similar inputs, linked to plantar flexor alpha activity; and 4) tonic gamma-efferents differed, to the extent that they were modulated at all. The FHL units peaked with the plantar flexor alphas. The FDL neurons generally peaked before alpha activity even began.

This study investigates the effects of parkinsonism and dopamine replacement therapy (levodopa) on centrally initiated postural activity preceding rising onto the toes. The electromyographic (EMG) and force magnitude, scaling, sequencing, and postural stabilization were compared when rising-to-toes under two conditions, slow/low versus fast/high, for parkinsonian patients and elderly control subjects. Parkinsonian subjects were tested after withholding their levodopa medication for 12-16 h and again 1 h after taking their medication when parkinsonian signs were diminished. Parkinsonian subjects showed reduced magnitudes and delayed timing of the postural and voluntary components of the rise-to-toes task, as if they had difficulty turning off the postural, tibialis anterior (TIB) component and initiating the voluntary, gastrocnemius (GAS) component. Dopamine improved the relative timing, as well as the magnitude of both postural and voluntary components of rise-to-toes. Although the magnitude of dorsiflexion torque was smaller for parkinsonian subjects ON and OFF than for healthy elderly controls, the parkinsonian subjects showed intact scaling of the magnitude of postural activity. Parkinsonian subjects do not perform the rise-to-toes task like normal subjects who are instructed to rise slowly; the relative timing of TIB and GAS activation was different even at comparable speeds of performance. Parkinsonian subjects, both ON and OFF, exhibited greater risk of falling than elderly control subjects when rising to toes. This increased risk of falling was reflected in a smaller safety margin between the peak center of mass (CoM) and peak center of pressure (CoP) during the task. The magnitude of mean postural dorsiflexion torque in the rise-to-toes task was highly correlated with a clinical rating scale of gait and balance, suggesting that force control is a critical factor influencing postural control in patients with Parkinson's disease.

The postsynaptic potentials elicited in peroneal motoneurons by either mechanical stimulation of cutaneous areas innervated by the superficial peroneal nerve (SP) or repetitive electrical stimulation of SP were compared in anesthetized cats. After denervation of the foot sparing only the territory of SP terminal branches, reproducible mechanical stimulations were applied by pressure on the plantar surface of the toes via a plastic disk attached to a servo-length device, causing a mild compression of toes. This stimulus evoked small but consistent postsynaptic potentials in every peroneal motoneuron. Weak stimuli elicited only excitatory postsynaptic potentials (EPSPs), whereas increase in stimulation strength allowed distinction of three patterns of response. In about one half of the sample, mechanical stimulation or trains of 20/s electric pulses at strengths up to six times the threshold of the most excitable fibers in the nerve evoked only EPSPs. Responses to electrical stimulation appeared with 3-7 ms central latencies, suggesting oligosynaptic pathways. In another, smaller fraction of the sample, inhibitory postsynaptic potentials (IPSPs) appeared with an increase of stimulation strength, and the last fraction showed a mixed pattern of excitation and inhibition. In 24 of 32 motoneurons where electrical and mechanical effects could be compared, the responses were similar, and in 6 others, they changed from pure excitation on mechanical stimulation to mixed on electrical stimulation. With both kinds of stimulation, stronger stimulations were required to evoke inhibitory postsynaptic potentials (IPSPs), which appeared at longer central latencies than EPSPs, indicating longer interneuronal pathways. The similarity of responses to mechanical and electrical stimulation in a majority of peroneal motoneurons suggests that the effects of commonly used electrical stimulation are good predictors of the responses of peroneal motoneurons to natural skin stimulation. The different types of responses to cutaneous afferents from SP territory reflect a complex connectivity allowing modulations of cutaneous reflex responses in various postures and gaits.

There is a need for a hand-heating system that will keep the hands warm during cold exposure without hampering finger dexterity. The purpose of this study was to examine the effects of torso heating on the vasodilative responses and comfort levels of cooled extremities during a 3-h exposure to -15 degreesC air. Subjects were insulated, but their upper extremities were left exposed to the cold ambient air. The effect of heating the torso [torso-heating test (THT)] on hand comfort was compared with a control condition in which no torso heating was applied, but Arctic mitts were worn [control test (CT)]. The results indicate that mean finger temperature, mean finger blood flow, mean toe temperature, mean body skin temperature, body thermal comfort, mean finger thermal comfort, and rate of body heat storage were all significantly (P < 0.05) higher on average (n = 6) during THT. Mean body heat flow was significantly (P < 0.05) lower during THT. There were no significant differences (P >/= 0.05) in rectal temperature between CT and THT. Mean unheated body skin temperature and mean unheated body heat flow (both of which did not include the torso area in the calculation of mean body skin temperature and mean body heat flow) were also calculated. There were no significant differences (P >/= 0.05) in mean unheated body skin temperature and mean unheated body heat flow between CT and THT. It is concluded that the application of heat to the torso can maintain finger and toe comfort for an extended period of time during cold exposure.

1. Single motor axons innervating human toe extensor muscles were selectively stimulated through a tungsten microelectrode inserted percutaneously into the peroneal nerve. Twitch and tetanic forces were measured from a strain gauge over the proximal phalanx of the toe generating the greatest force. Twitch data were obtained from 19 single motor units in nine subjects: 8 motor units supplied extensor hallucis longus (EHL), 5 motor units supplied extensor digitorum longus (EDL), and 6 motor units supplied extensor digitorum brevis (EDB). Unpotentiated twitch forces ranged from 6.3 to 78.1 mN (20.0 +/- 4.0 mN, mean +/- SE), with the distribution highly skewed toward small forces. Twitch contraction and half-relaxation times were 74.8 +/- 3.9 and 78.6 +/- 6.0 ms, respectively. Compared with motor units in human thenar muscles, those in human toe extensor muscles were stronger but slower. However, as in thenar motor units, twitch force and contraction time were not related. 2. Force-frequency relationships were determined for 13 units (5 EDL, 5 EHL, 3 EDB) by stimulating each unit with short trains (1.0-5.0 s) of constant frequency (2-100 Hz). Peak force was related to stimulus frequency in a sigmoid fashion. The steep region of the curve extended from 5.5 +/- 0.7 (SE) Hz to 16.3 +/- 1.1 Hz for all units, and the stimulus frequency required to generate half-maximal force (9.6 +/- 0.6 Hz) was close to the center of the steep range. This frequency, which was inversely related to twitch contraction time, was lower than the frequency required to develop half-maximal force of human thenar motor units (12 +/- 4 Hz, mean +/- SD). The slopes of the regression lines relating force to frequency, computed over the steep range for each unit, were also lower for the toe extensors (3.7 +/- 0.7 mN/Hz) than for the thenar muscles (6 +/- 1 mN/Hz). 3. Maximal tetanic forces ranged from 29.9 to 188.1 mN (89.0 +/- 16.5 mN, mean +/- SE), and were generated at stimulus frequencies from 15 to 100 Hz (median 50 Hz). The stimulation frequency required for fused tetani (absence of noticeable force fluctuation) was generally less than that required for maximum tetanic force. The mean twitch-tetanus ratio, calculated for unpotentiated twitches, was 0.22 +/- 0.02 (range 0.15-0.41). This ratio was higher than for human thenar motor units (0.14 +/- 0.06, mean +/- SE). After twitch potentiation of 10 units, the mean twitch-tetanus ratio increased to 0.28 +/- 0.04. 4. The effects of preceding each stimulus train with a short interstimulus interval (10 ms) on force production at each frequency were examined in nine motor units. Peak forces at the onset of each contraction were higher when such an "initial doublet" preceded stimulus trains of < or = 20 Hz, but the mean force at the end of each stimulus train was not significantly affected at any frequency. 5. Eight units were stimulated with a train that increased in frequency continuously from 2 to 80 Hz, and then decreased symmetrically. This pattern resulted in peak forces that were higher on the descending limb of the stimulus train, the force-frequency relationship tracing a hysteresis loop. Hysteresis was exhibited because damping in the neuromuscular system causes the mechanical output of muscle to lag behind neural input. Thus, in non-steady-state conditions (as in most forms of natural activity), somewhat higher firing rates may be required to attain a particular level of force; once attained, force output will be transiently unresponsive to diminution of firing rate. 6. We conclude that there are differences in the contractile properties of single motor units in human toe extensor muscles (involved in posture and locomotion) and thenar muscles (involved in prehension and manipulation). Twitch-tetanus ratios were greater for motor units in the toe extensors, and this property accounted for the lower force sensitivity of these units to increases in frequency. (ABSTRACT TRUNCATED)

1. Single neurons in the ventroposterior lateral thalamic nucleus were studied in 10 anesthetized raccoons, 4 of which had undergone amputation of the fourth digit 4-5 mo before recording. Neurons with receptive fields on the glabrous skin of a forepaw digit were examined in response to electrical stimulation of the "on-focus" digit that contained the neuron's receptive field and stimulation of an adjacent, "off-focus" digit. 2. In normal raccoons all neurons responded to on-focus stimulation with an excitation at a short latency (mean 13 ms), whereas only 63% of the neurons responded to off-focus digit stimulation. The off-focus responses had a longer latency (mean 27.2 ms) and a higher threshold than the on-focus responses (800 and 452 microA, respectively). Only 3 of 32 neurons tested with off-focus stimulation had both a latency and a threshold within the range of on-focus values. Inhibition following the excitation was seen in the majority of neurons with both types of stimulation. 3. In the raccoons with digit removal, the region of the thalamus that had lost its major peripheral input (the "deafferented" region) was distinguished from the normal third and fifth digit regions on the basis of the sequence of neuronal receptive fields within a penetration and receptive field size as described previously. 4. Almost all of the neurons in the deafferented region (91%) were excited by stimulation of one or both adjacent digits. The average latency for these responses was shorter (15.3 ms) and the threshold was lower than was the case with off-focus stimulation in control animals. These values were not significantly different from the responses to on-focus stimulation in the animals with digit amputation. 5. These results confirm that reorganization of sensory pathways can be observed at the thalamic level. In addition to the changes in the somatotopic map that have been shown previously with the use of mechanical stimuli, the present paper demonstrates an improvement in several quantitative measures of single-unit responses. Many of these changes suggest that this reorganization could be explained by an increased effectiveness of preexisting, weak connections from the off-focus digits; however, the increase in the proportion of neurons responding to stimulation of adjacent digits may indicate that sprouting of new connections also occurs.

1. The somatotopic representation of the human primary motor cortex was examined noninvasively with estimates of cerebral blood flow (CBF) obtained with positron emission tomography. Twelve normal subjects performed a motor tracking task with the arm, first finger, tongue, and great toe commensurate with the bolus injection of radioactive H215O. Images of the relative percent increase of blood flow, compared with control studies, demonstrated reproducible foci of CBF increases in the motor cortex in every subject. Each motor task could be localized to a predictable site on a coronal section containing the precentral gyrus. 2. In reproducibility experiments of repeated measures, it was determined that two foci of activation in the primary motor cortex could be discriminated with a 95% confidence if they were separated by 5.4 mm. 3. In five subjects with matched magnetic resonance imaging studies, the sites of activation were variable with respect to surface anatomy and could be found at the depth of sulci or the surface of gyri. The findings were similar to previously reported electrophysiological studies using direct cortical stimulation. 4. The method may be applied to the in vivo functional mapping of the primary motor cortex in patients with cerebral disorders.

1. The effect of transection and ligation of the digital nerves of either one (toe 3) or two (toe 3 and toe 4) hindpaw digits, in the first postnatal week, on the tactile receptive fields (RFs) of spinocervical tract (SCT) neurons was studied in adult, alpha-chloralose-anesthetized cats. Immediately before recording, the digital nerves of the corresponding digit(s) of the opposite, intact hindpaw were transected, and the neonatally lesioned digital nerves were recut proximal to the transection neuroma. 2. In the medial part of the dorsal horn at the L6-L7 level, the digits of the hindlimb are represented in the RFs of SCT cells in a precise axial sequence from the most medial digit (toe 2) rostrally to the most lateral digit (toe 5) caudally. Acute denervation of one or two digits in the adult produced an area in the ipsilateral dorsal horn in which SCT cells lacked any RFs. When acute denervation was restricted to a single digit, the unresponsive region of dorsal horn was approximately 3 mm in length, and when two digits were denervated the unresponsive zone was approximately 6 mm long. Because the representation of the toes of the left hindpaw is a mirror image of that of the right, the rostrocaudal extent and position of the region of unresponsive SCT cells was used to assess the location of the borders of the chronically deprived region on the opposite side of the cord. 3. In all cats examined after neonatal denervation of toe 3, most (89%) of the SCT cells sampled within the chronically deprived toe 3 representation had RFs. These RFs were either on toe 2 (44%) or toe 4 (18%), and a large proportion of cells (38%) had multiple RFs with components on both toe 2 and toe 4. In most cases the cells fired briskly to displacement of hairs or light touch of the skin within these RFs. SCT cells with a RF on toe 2 and/or toe 4 were found throughout the whole 3-mm length of the chronically deprived toe 3 region, but cells with a RF on toe 2 were more commonly found than cells with a RF on toe 4 at axial distances greater than or equal to 1.5 mm from the boundary of the normal representations of the respective digit. 4. After chronic, neonatal denervation of both toe 3 and toe 4, 59% of SCT cells sampled overall had RFs, but there was a large degree of interanimal variation in the proportion of unresponsive neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

1. "Unmasking" of weak synaptic connections has been suggested as a mechanism for the early changes in cortical topographic maps that follow alterations of sensory activity. For such a mechanism to operate, convergent sensory inputs must already exist in the normal cortex. 2. We tested for topographic and cross-modality convergence in primary somatosensory cortex of raccoon. The representation of glabrous skin of forepaw digits was chosen because, even though it is dominated by inputs from the glabrous skin of a single digit, it nevertheless comes to respond to stimulation of other digits when, e.g., a digit is removed. 3. Intracellular recordings were made from 109 neurons in the representation of glabrous skin of digit 4. Neurons were tested for somatosensory inputs with electrical and natural stimulation of digits. 4. Excitatory postsynaptic potentials (EPSPs) were evoked in 100% of the neurons (109/109) by electrical stimulation of glabrous skin of digit 4, and in 79% (31 of 39) by vibrotactile stimulation. 5. Glabrous skin of digit 4 was not the sole source of somatosensory inputs. A minority of neurons generated EPSPs after electrical stimulation of hairy skin of digit 4 (10 of 98 neurons, 10%). Electrical stimulation of digits 3 or 5 evoked EPSPs in 22 of 103 neurons (21%). Natural stimulation (vibrotactile or hair bending) was also effective in most of these latter cases (digit 3, 6/7; digit 5, 9/10). 6. Intracortical microstimulation of the "heterogeneous zone" was used to test for corticocortical connections to neurons in the glabrous zone.(ABSTRACT TRUNCATED AT 250 WORDS)