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Neurone firing behaviour is a result of complex interaction between synaptic inputs and cellular intrinsic properties. Intriguing firing behaviour, delayed spiking, was shown in some neurones, in particular, in cat neocortical neurones and rat pyramidal hippocampal neurones. In contrast, the similar spiking mode was not reported for animal spinal motoneurones. In the present study, an attempt was made to look for possible evidence of delayed spiking in human motoneurones firing within the low-frequency, sub-primary range, characteristic for voluntary muscle contractions and postural tasks. Forty-seven firing motor units (MUs) were analyzed in ten experiments on three muscles (the flexor carpi ulnaris, the tibialis anterior, and the abductor pollicis brevis) in four healthy humans. Single MUs were activated by gentle voluntary muscle contractions. MU peri-stimulus time histograms, durations of inter-spike intervals, and motoneurone excitability changes within a target interspike interval were analyzed. It was found that during testing the firing motoneurone excitability by small, transient excitatory Ia afferent volley, depending firstly on volley timing within a target interspike interval and excitatory volley strength, the same motoneurone displayed either the direct short-latency response (the H-reflex) or the delayed response (with prolonged and variable latency). Thus, the findings, for the first time, provide evidence for a possibility of two modes of spiking in firing motoneurones. Methods of the estimation of delayed responses and their possible functional significance are discussed. It is emphasized that, for understanding of this issue, the integration of data from studies on experimental animals and humans is desirable.

This study investigated whether modulation of corticospinal-motoneuronal excitability and/or synaptic transmission of the Ia afferent spinal reflex contributes to decreases in voluntary activation and muscular force after an acute bout of prolonged static muscle stretching. Fifteen men performed five 60-s constant-torque stretches (15-s rest intervals; total duration 5 min) of the plantar flexors on an isokinetic dynamometer and a nonstretching control condition in random order on 2 separate days. Maximum isometric plantar flexor torque and triceps surae muscle electromyographic activity (normalized to M wave; EMG/M) were simultaneously recorded immediately before and after each condition. Motor-evoked potentials (using transcranial magnetic stimulation) and H-reflexes were recorded from soleus during EMG-controlled submaximal contractions (23.4 ± 6.9% EMG maximum). No changes were detected in the control condition. After stretching, however, peak torque (mean ± SD; -14.3 ± 7.0%) and soleus EMG/M (-17.8 ± 6.2%) decreased, and these changes were highly correlated (

Hoffmann (H‐) reflex amplitudes in plantar flexor soleus muscle are modulated by posture, yet dorsiflexor tibialis anterior (TA) H‐reflex parameters have sparingly been studied. The purpose was to investigate modulation of the TA H‐reflex when postural demands are increased from sitting to standing. In this study, data from 18 participants (Age: 25 ± 4 years, Height: 170.9 ± 9.5 cm, Weight: 75.9 ± 17.2 kg) allowed comparison of two experimental conditions involving different postures (i.e. sitting and standing). Maximal amplitude of the TA H‐reflex (H max) as a percent of the maximal M‐wave amplitude (M max) (H max (% M max)) during sitting and standing was compared using ANOVA. Modulation of TA H‐reflex amplitude was found: Eleven participants showed facilitation and seven showed no change of reflex amplitudes. Only participants in the facilitation group showed modulation related to changes in posture (sitting: 8.7 ± 2.9%; standing: 14.8 ± 6.7%, P = 0.005). These data provide evidence of the sensitivity to posture of TA H‐reflexes. As with task‐dependent changes in soleus H‐reflexes, presynaptic regulation of Ia afferent transmission is a possible mechanism. Further investigations into causes of modulation are warranted.

Postactivation depression (PActD) of Ia afferent excitatory postsynaptic potentials (EPSPs) in spinal motoneurons results in a long-lasting depression of the stretch reflex. This phenomenon (PActD) is of clinical interest as it has been shown to be reduced in a number of spastic disorders. Using in vivo intracellular recordings of Ia EPSPs in adult mice, we demonstrate that PActD in adult (100–220 days old) C57BL/6J mice is both qualitatively and quantitatively similar to that which has been observed in larger animals with respect to both the magnitude (with ∼20% depression of EPSPs at 0.5 ms after a train of stimuli) and the time course (returning to almost normal amplitudes by 5 ms after the train). This validates the use of mouse models to study PActD. Changes in such excitatory inputs to spinal motoneurons may have important implications for hyperreflexia and/or glutamate-induced excitotoxicity in the neurodegenerative disease amyotrophic lateral sclerosis (ALS). With the use of the G127X SOD1 mutant mouse, an ALS model with a prolonged asymptomatic phase and fulminant symptom onset, we observed that PActD is significantly reduced at both presymptomatic (16% depression) and symptomatic (17.3% depression) time points compared with aged-matched controls (22.4% depression). The PActD reduction was not markedly altered by symptom onset. Comparing these PActD changes at the EPSP with the known effect of the depression on the monosynaptic reflex, we conclude that this is likely to have a much larger effect on the reflex itself (a 20–40% difference). Nevertheless, it should also be accounted that in aged (580 day old) C57BL/6J mice there was also a reduction in PActD although, aging is not usually associated with spasticity.

Muscle kinematics and kinetics are nonlinearly encoded by proprioceptors, and the changes in muscle length and velocity are integrated into Ia afferent. Besides, proprioceptive signals from multiple muscles are probably mixed in afferent pathways, which all lead to difficulties in proprioceptive recognition for the cerebellum. In this study, the artificial neural networks, whose organizations are biologically based on the spinocerebellar tract and cerebellum, are utilized to decode the proprioceptive signals. Consistent with the controversy of the proprioceptive division in the dorsal spinocerebellar tract, the spinocerebellar tract networks incorporated two distinct inferences, (1) the centralized networks, which mixed Ia, II, and Ib and processed them together; (2) the decentralized networks, which processed Ia, II, and Ib afferents separately. The cerebellar networks were based on the Marr–Albus model to recognize the kinematic states. The networks were trained by a specific movement, and the trained networks were subsequently required to predict kinematic states of six movements. The results demonstrated that the centralized networks, which were more consistent with the physiological findings in recent years, had better recognition accuracy than the decentralized networks, and the networks were still effective even when proprioceptive afferents from multiple muscles were integrated.

The human soleus stretch reflex is comprised of several components with different latencies typically termed the SLR, MLR and in some cases LLR component. The SLR is suggested to be mediated by Gr Ia afferent arising from activation of spindle afferents, the MLR is predominantly due to gr Ib and II feedback while LLR likely contains a cortical component. Here we report data on the changes in both the SLR and MLR following a well-established conditioning protocol. Participants attended 30 sessions where the initial six served as a baseline measure of reflex size and the following 24 sessions were devoted to training the participants to either increase or decrease the SLR component. Following successful up-conditioning of the SLR, the MLR remained unchanged. However the SLR and MLR both decreased with down-conditioning. This suggests that the MLR is not exclusively Gr Ib or II based but has some Ia afferent input.

Background: The muscle spindle is an important sensory organ for proprioceptive information, yet there have been few attempts to use Shannon information theory to quantify the capacity of human muscle spindles to encode sensory input. Methods: Computer simulations linked kinematics, to biomechanics, to six muscle spindle models that generated predictions of firing rate. The predicted firing rates were compared to firing rates of human muscle spindles recorded during a step-tracking (center-out) task to validate their use. The models were then used to predict firing rates during random movements with statistical properties matched to the ergonomics of human wrist movements. The data were analyzed for entropy and mutual information. Results: Three of the six models produced predictions that approximated the firing rate of human spindles during the step-tracking task. For simulated random movements these models predicted mean rates of 16.0 ± 4.1 imp/s (mean ± SD), peak firing rates

Sudden stretch of active muscle typically results in two characteristic electromyographic responses: the short latency M1 and the long latency M2. The M1 response originates from the monosynaptic Ia afferent reflex pathway. The M2 response is less well understood and is likely a compound response to different afferent inputs mediated by spinal and transcortical pathways. In this study the possible contribution of the Ia afferent pathway to the M2 response was investigated. A mechanism was hypothesized in which the M1 response synchronizes the motoneurons, and therewith their refractory periods. Stretch perturbation experiments were performed on the wrist and results were compared with a computational model of a pool of motoneurons receiving tonic and Ia afferent input. The simulations showed the same stretch amplitude, velocity, and duration-dependent characteristics on the M2 as found experimentally. It was concluded that the stretch duration effect of the M2 likely originates from the proposed Ia afferent mediated mechanism.

The central nervous system employs different strategies to execute specific motor tasks. Because afferent feedback during shortening and lengthening muscle contractions differs, the neural strategy underlying these tasks may be quite distinct. Cortical drive may be adjusted or afferent input regulated. The exact mechanisms are not clear. Here, we examine the control of synaptic transmission across the Ia synapse during shortening and lengthening muscle contractions. Subjects were instructed to maintain isolated activity in a single tibialis anterior (TA) motor unit while muscle length was varied from flexion to extension and back. At a fixed interval after a firing of the active motor unit, a single electrical stimulus was applied to the common peroneal nerve to activate Ia afferents from the TA muscle. We investigated the stimulus-induced change in firing probability of 19 individual low-threshold TA motor units during shortening and lengthening contractions. Any change in firing probability depends on both pre- and postsynaptic mechanisms. In this experiment, motoneuron firing rate was similar during both contraction types. There was no difference in the firing probability between shortening and lengthening contractions (0.23 ± 0.03 and 0.20 ± 0.02, respectively). We suggest that there is no contraction type-specific control of Ia input to the motoneurons during shortening and lengthening muscle contractions. Cortical adjustments may have occurred.

Transmission in the corticospinal and Ia pathways to soleus motoneurons was investigated in healthy human subjects during bicycling. Soleus H reflexes and motor evoked potentials (MEPs) after transcranial magnetic stimulation (TMS) were modulated similarly during the crank cycle being large during downstroke [concomitant with soleus background electromyographic (EMG) activity] and small during upstroke. Tibialis anterior MEPs were in contrast large during upstroke and small during downstroke. The soleus H reflexes and MEPs were also recorded during tonic plantarflexion at a similar ankle joint position, corresponding ankle angle, and matched background EMG activity as during the different phases of bicycling. Relative to their size during tonic plantarflexion, the MEPs were found to be facilitated in the early part of downstroke during bicycling, whereas the H reflexes were depressed in the late part of downstroke. The intensity of TMS was decreased below MEP threshold and used to condition the soleus H reflex. At short intervals (conditioning-test intervals of −3 to −1 ms), TMS produced a facilitation of the H reflex that is in all likelihood caused by activation of the fast monosynaptic corticospinal pathway. This facilitation was significantly larger in the early part of downstroke during bicycling than during tonic plantarflexion. This suggests that the increased MEP during downstroke was caused by changes in transmission in the fast monosynaptic corticospinal pathway. To investigate whether the depression of H reflexes in the late part of downstroke was caused by increased presynaptic inhibition of Ia afferents, the soleus H reflex was conditioned by stimulation of the femoral nerve. At a short interval (conditioning-test interval: −7 to −5 ms), the femoral nerve stimulation produced a facilitation of the H reflex that is mediated by the heteronymous monosynaptic Ia pathway from the femoral nerve to soleus motoneurons. Within the initial 0.5 ms after its onset, the size of this facilitation depends on the level of presynaptic inhibition of the Ia afferents, which mediate the facilitation. The size of the facilitation was strongly depressed in the late part of downstroke, compared with the early part of downstroke, suggesting that increased presynaptic inhibition was indeed responsible for the depression of the H reflex. These findings suggest that there is a selectively increased transmission in the fast monosynaptic corticospinal pathway to soleus motoneurons in early downstroke during bicycling. It would seem likely that one cause of this is increased excitability of the involved cortical neurons. The increased presynaptic inhibition of Ia afferents in late downstroke may be of importance for depression of stretch reflex activity before and during upstroke.

In human peripheral nerves, physiological evidence has been presented for a number of biophysical differences between cutaneous afferents and alpha motor axons. The differences in strength-duration properties for cutaneous afferents and motor axons in the median nerve have been attributed to greater expression of a persistent Na(+) conductance (I(Na,P)) on cutaneous afferents. However, it is unclear whether the biophysical properties of human group Ia afferents differ from those of cutaneous afferents. The present studies were undertaken to determine whether the properties of human group Ia afferents can be studied indirectly using 'threshold tracking' to measure the excitability changes in the H reflex, and to determine whether the excitability of group Ia afferents differs from that of cutaneous afferents. The strength-duration properties of the soleus H reflex and soleus motor axons were measured at rest and during sustained voluntary contractions. Similar experiments were performed on the median nerve at the wrist to study the strength-duration properties of cutaneous afferents, alpha motor axons and H reflex of the thenar muscles. In addition, the technique of 'latent addition' was used to determine whether there was a difference in a low-threshold conductance on soleus Ia afferent and motor axons. The present findings indicate that the strength-duration time constant (tau(SD)) for the H reflex is longer than that for alpha motor axons, but similar to that for cutaneous afferents. There were no differences in tau(SD) for the soleus H reflex at rest and during contractions, suggesting that tau(SD) for the H reflex is largely unaffected by changes in synaptic or motoneurone properties. Finally, the difference in latent addition suggests that the longer tau(SD) of the soleus H reflex may indeed be due to greater activity of a persistent Na(+) conductance on Ia afferents than on soleus alpha motor axons.

Mammalian Ia afferents have a tendency to fire action potentials phase-locked to periodic stimuli, when certain static gamma axons are stimulated at constant frequency. This driving is thought to be mediated by chain fibres, which, in contrast to other intrafusal fibres, have fast contraction properties, comparable to type II extrafusal fibres. Intrafusal contractions are mechanically transmitted to the sensory ending, which transduces the movement into a receptor current. The ensuing oscillatory changes in receptor potential which accompany these rapid chain fibre contractions then cause the generation of one spike during each contraction cycle. If stimulation frequencies are high, or the bag2 fibre is coactivated, the afferent may fire at subharmonic frequencies, giving 1:2 or 1:3 driving. Also, very irregular spike intervals can be observed, which gives the impression of an erratic firing response. Changes in the initial muscle length often cause transitions between subharmonic, 1:1 driving, and irregular firing (Boyd, Murphy & Mann, 1985; Banks, 1991).

Recent work has shown that the monosynaptic pathway of the SSR can be operantly conditioned, and that a significant part of the plasticity responsible for the behavioral change resides in the spinal cord. The most likely sites of this activity-driven plasticity are the synapse of the Ia afferent neuron on the motoneuron and/or the motoneuron itself. Because the SSR pathway is the simplest and most accessible stimulus-response pathway in the vertebrate CNS, it may provide a valuable experimental model for elucidating activity-driven CNS changes responsible for learning.

The complexity and inaccessibility of the vertebrate CNS impede the localization and description of memory traces and the definition of the processes that create them. Recent work has shown that the spinal stretch reflex (SSR), which is produced by a monosynaptic twoneuron pathway, can be operantly conditioned, and that memory traces responsible for this behavioral change reside in the spinal cord. The probable locations are the terminal of the Ia afferent neuron on the motoneuron and/or the motoneuron itself. Because it modifies a simple well-defined and accessible pathway, SSR conditioning may be a valuable experimental model for studying vertebrate memory.

1. We have developed a technique to measure the total amount of current from a synaptic input system that reaches the soma of a motoneuron under steady-state conditions. We refer to this quantity as the effective synaptic current (IN) because only that fraction of the synaptic current that actually reaches the soma and initial segment of the cell affects its recruitment threshold and firing frequency. 2. The advantage of this technique for analysis of synaptic inputs in comparison to the standard measurements of synaptic potentials is apparent from Ohm's law. Steady-state synaptic potentials recorded at the soma of a cell are the product of IN and input resistance (RN), which is determined by intrinsic cellular properties such as cell size and membrane resistivity. Measuring IN avoids the confounding effect of RN on the amplitudes of synaptic potentials and thus provides a more direct assessment of the magnitude of a synaptic input. 3. Steady-state synaptic inputs were generated in cat medial gastrocnemius (MG) motoneurons by using tendon vibration to activate homonymous Ia afferents. We found that the magnitude of the Ia effective synaptic current (Ia IN) was not the same in all MG cells. Instead, Ia IN covaried with RN (r = 0.64; P less than 0.001), being about twice as large on average in motoneurons with high RN values as in those with low RN values. Ia IN was also correlated with motoneuron rheobase, afterhyperpolarization duration, and axonal conduction velocity. 4. A comparison of transient Ia EPSPs with steady-state Ia EPSPs (Ia EPSPSS) evoked in the same cells suggested that the effective synaptic current that produces the transient Ia EPSP was also greater in motoneurons with high RN values than in those with low RN values. 5. The factors responsible for the Ia IN-RN covariance are uncertain. However, our finding greater values of Ia IN in high RN motoneurons is consistent with other evidence suggesting that Ia boutons on these motoneurons have a higher probability for neurotransmitter release than those on low RN motoneurons (19). 6. The neural mechanisms underlying orderly recruitment are discussed. The effect of the Ia input is to produce an approximately twofold expansion of the differences in motoneuron recruitment thresholds that are generated by intrinsic cellular properties. It is suggested that the higher efficacy of Ia input in low-threshold motoneurons confers particular importance on this input system in the control of vernier movements (7).

The purpose of this paper is to review research on the reflex action of group II afferent fibers originating in the muscle spindle. Although the reflex actions of the Ia afferent fibers from muscle spindles are well defined, the reflex action of the group II afferent fibers is not clearly understood. Experiments on animals whose spinal cord had been transected gave rise to the idea that group II afferent fibers facilitate flexor muscles and inhibit extensor muscles regardless of the origin of the fibers. This concept has been accepted by neurophysiologists and has even been incorporated into therapeutic exercise procedures by physical therapists. However, accumulating evidence is presented that challenges this classical concept. Physical therapists, when trying to incorporate the reflexes evoked by the group II afferent fibers into treatment, must become better aware of the complex reflex actions of which these fibers are capable.

In cats anesthetized with allobarbitone-urethane, Mn 2+ and Co 2+ (and occasionally La 3+ ) were released extracellularly from micropipettes while recordings were made of monosynaptic excitatory postsynaptic potentials evoked in lumbosacral motoneurones by Ia afferent stimulation. These excitatory postsynaptic potentials consistently showed a marked depression in their rate of rise (by an average of 44%) and an increase in half-amplitude duration (by an average of 50%). There was a less pronounced reduction in peak amplitude and increase in time-to-peak. After applications of Mn 2+ or Co 2+ , presynaptic potentials recorded in motoneurones showed no sign of any depression but the synaptic delay was clearly increased. It is concluded that the monosynaptic excitatory postsynaptic potential evoked by Ia afferents in cat motoneurones is probably mediated by chemical transmission.

1. The individual EPSPs evoked by the action of single Ia fibers from cat triceps surae (MG, LG, SOL) were recorded in homonymous and heteronymous motoneurons innervating these same three muscles. 2. In general, Ia fibers projected to a greater percentage of homonymous than heteronymous motoneurons. One class of Ia afferent evoked EPSPs in virtually all homonymous motoneurons; the other had a substantially lower projection frequency. Possible difficulties introduced by the limited resolution of the averaging technique are discussed. 3. Individual EPSPs were larger on the average if evoked a) in SOL rather than in MG or LG motoneurons, b) by LG rather than by MG or SOL afferent fibers, or c) in homonymous rather than in heteronymous motoneurons. The mean EPSP was larger in homonymous than in heteronymous motoneurons because the largest EPSPs (greater than 150 muV) were found mainly in homonymous motoneurons. 4. Rise times of EPSPs were only slightly shorter in homonymous than in heteronymous motoneurons, suggesting that other factors besides relative location of Ia terminals account for the observed EPSP amplitude differences. Rise times in SOL motoneurons were longer than those in MG or LG. 5. LG afferent fibers tended to produce larger EPSPs in rostral than in caudal LG motoneurons, and MG afferents produced larger EPSPs in caudal than in rostral MG motoneurons. These spatial effects were in accord with the more rostral entry of LG than MG Ia afferents into the spinal cord. The differential projection of SOL afferents to MG and SOL motoneurons which overlap spatially in the spinal cord suggests a species specificity in addition to a location specificity.

The excitability of a firing human motoneurones was tested by Ia afferent excitatory volleys. An H reflex was evoked by stimulation of the tibial nerve while the subjects maintained the background rhythmic firing of single MU of the soleus and of the medialis gastrocnemius during weak voluntary contraction. The mean background firing rate of MUs ranged from 3.6 to 9.4 imp/sec. In order to estimate the ‘effectiveness’ of an afferent volley, PSTHs were constructed. The firing index was used as a measure of motoneurone excitability. It has been shown that the ‘effectiveness’ of an afferent volley decreases with an increase in background firing rate of a MU. Such a dependence was deduced from changes in current motoneurone excitability within an interspike interval. The results indicate that during natural muscle contraction at a low firing rate of a motoneurone, the discharge rate is one of the factors affecting the excitability of a firing motoneurone and, hence, its input-output relations. The method of testing changes in human motoneurone excitability within an interspike interval may prove useful in studies on interaction of volleys arriving at a motoneurone via different excitatory inputs.

Although GABA and piperidine-4-sulphonate depolarize Ia afferent terminations in the lumbar spinal cord by activation of bicuculline-sensitive GABA receptors, no evidence was obtained for abicuculline-sensitive alteration by either GABAmimetic of the excitability of vestibulospinal terminations. This suggests that the terminal arborizations of vestibulospinal fibers, unlike their cell bodies. are devoid of GABA receptors having properties similar to those on either central neuron cell bodies, primary afferent cell bodies or their central terminations.

1. Tonic activation of muscular activity was observed during forced vibration of the gastrocnemius and soleus muscles in the decerebrate cat. Such tonic activity was not observed in the tibialis anterior muscle.2. Intervals of both unitary EMG and motoneuronal spikes were shown to occur according to the principle of integer multiplication of vibratory cyclic time.3. Gradual recruitment of tonic vibration reflex (vibratory facilitation) was found to survive after the cessation of vibratory stimulation (postvibratory facilitation).4. Nonsequential interspikes revealed that gradual tonic recriutment is effected by the spikes with a shorter firing interval.5. Muscle activities during TVR may be effected by PTP with some excitatory frequency associated with that of the applied vibration.6. Postvibratory facilitation did not correlate with the vibratory frequency. The firing of such phases may be determined by the intrinsic repetition rate of the individual motoneuron.

The frequency transfer characteristics of the tonic stretch reflex have been studied using homonymous Ia afferent stimulation as input to the soleus motoneurone. By varying additionally the frequency of inhibitory volleys from either the Ia component of antagonist muscle nerve or the homonymous Ib afferents, the suppressive effect on the excitatory synaptic activity has been investigated for a wide variety of different phase relations.

JANKOWSKA, E., S. LUND and A. LUNDBERG. The effect of DOPA on the spinal cord. 4. Depolarization evoked in the central terminals of contralateral Ia afferent terminals by volleys in the jlexor rejlex afferents. Acta physiol. scand. 1966. 68. 337-341. After injection of DOPA in unanaesthetized spinal cats volleys in the flexor reflex afferents (FRA) evoke a late long lasting contralateral dorsal root potential. Measurements of excitability of contralateral afferent terminals show a corresponding depolarization of Ia, but not of Ib or cutaneous afferent terminals. Results are also presented showing that transmission from the FRA to contralateral Ia afferent terminals can be inhibited from the FRA. In previous papers in this series it has been shown that after an i.v. injection of 1-3,4-dihydroxyphenylalanine (DOPA) volleys in the flexor reflex afferents evoke a late ipsilateral dorsal root potential (DRP) which is caused by a selective depolarization in the central terminals of Ia afferents (Andtn et al. 1966a, b). In the present paper it will be shown that after DOPA volleys in the FRA produce similar effects on the contralateral side. The experimental procedures have been given in the previous papers.

1. The origin of primary afferent depolarization (PAD) of the central terminals of group II afferent fibres of tibialis anterior and extensor digitorum longus muscles has been investigated in the cat. Changes in the excitability of the terminals to intraspinal stimuli, upon application of conditioning stimuli to muscle nerves (quadriceps, sartorius, gracilis, posterior biceps-semitendinosus, anterior biceps-semimembranosus, gastrocnemius-soleus, deep peroneal), cutaneous nerves (sural, superficial peroneal) and the posterior nerve to the knee joint, were used as a measure of PAD. 2. PAD was most readily evoked by conditioning stimuli which were maximal for group II muscle afferents. However, some PAD was also evoked from group I afferents and evidence is presented that group Ia afferents contributed. Afferents of posterior biceps-semitendinosus and sartorius muscles appeared to be most effective. PAD was also evoked by stimulation of cutaneous and joint nerves, often in the same fibres which were affected by group Ia afferents. 3. It is concluded that there are several common sources of PAD of group II and group Ia afferent terminals on the one hand, and group Ib afferent terminals on the other. 4. The properties of PAD of group II afferents are discussed in relation to the problem of how PAD affects transmission from fibres with long terminal branches of small diameter.

When a subject is instructed to resist external load, stretch-induced reflex responses can be observed in the emg of the contracting muscles when the load is abruptly increased. As commonly known, these reflex responses are grouped into segments denoted M1, M2 and M3 according to the nomenclature of Tatton et al. (1975). The long latency components have attracted particular attention; they have been studied in normal as well as in pathologic states, such as in parkinsonism and chorea.

For several years we have studied the input-output relations of muscle spindles and tendon organs using random length perturbations as stimuli. While studying the effect of gamma stimulation on spindle afferent responses, we made the chance observation that although static gamma fibers altered the response of a spindle Ia afferent to random length stimuli, in response to specific length sequences with concurrent gamma stimulation, the receptor produced an unexpectedly large number of spikes in exactly the same temporal relation to the repeated “random” sequence. We report here on the data obtained in one such experiment in which a gamma fiber we had classified as “static” modified the Ia afferent response to muscle stretch.

Publisher Summary This chapter describes the mechanisms of transmitter release at Ia afferent terminations. Fluctuations in the amplitude of a synaptic potential usually indicate that the amount of transmitter released at the nerve terminal is fluctuating. Some details of transmitter release mechanisms can be established from an analysis of these fluctuations. The afferent connection consisted of three boutons on a trunk dendrite. The average excitatory postsynaptic potentials (EPSPs) recorded in the neuron is shown. The standard deviation time course is analyzed. The number of different components in the EPSP is equal to the number of boutons. In two other results of this type, the number of boutons has always been equal to or greater than the number of different components in the EPSP.

The distribution of Ia terminals to alpha-motoneurons has been investigated under conditions where a) the Ia fiber innervates a foreign muscle and b) the motor axon innervates a foreign muscle. This distribution has been assessed by examining which motoneurons develop an EPSP in response to stimulation of a single Ia afferent fiber. Cross-union of the medial gastrocnemius and lateral gastrocnemius-soleus nerves has been performed in 5--8 day old kittens and the effect of this procedure on Ia connections to alpha-motoneurons has been investigated in the adult about one year later. No rearrangement of synaptic connections has been observed under these conditions. The connections from self-unioned Ia afferents and to self-unioned motoneurons are also normal. The individual EPSP's in these motoneurons with regenerated axons have normal rise times. The findings of normal Ia connections as well as normal EPSP rise times in alpha-motoneurons with regenerated motor axons indicates reversibility of the changes seen in axotomized motoneurons.

The spike-triggered averaging technique (26) was used to determine whether the synaptic input from medial gastrocnemius (MG) Ia-afferent fibers to homonymous motoneurons is "topographically weighted" (22) by means of differences in projection frequency, excitatory postsynaptic potential (EPSP) amplitude, or a combination of both factors. Motoneurons were classified as either "same branch" or "other branch," depending on whether a Ia-afferent fiber and motor axon were contained in the same or different intramuscular nerve branches. No difference was found in the projection frequency of Ia-afferents to the same branch and other branch motoneurons (95 versus 94%, respectively). The mean EPSP amplitude was larger in the same branch group of motoneurons (92 +/- 8 (SE) microV; n = V; n = 97) than in the other branch group (77 +/- 7 microV; n = 79). This difference was most striking in high-rheobase (greater than or equal to 10 nA) motoneurons, for which the mean EPSP amplitude in the same branch group was 82 +/- 12 microV (n = 48), whereas that in the other branch group was 52 +/- 5 microV (n = 37). In 60 cases it was possible to compare the EPSPs produced by a same branch afferent and an other branch afferent in the same motoneuron. The same branch afferent produced the larger EPSP in 73% (44/60) of the cases. Moreover, the mean ratio of the same branch to the other branch EPSP amplitudes was 1.7, which was both statistically significant and consistent with analogous results from our preceding study of aggregate EPSPs (22). Mean rise times and half-widths of EPSPs in the same branch group were not significantly different from those in the other branch group. Furthermore, no significant differences in rise times or half-widths between the two groups were evident when motoneurons were segregated according to their rheobase values. This suggests that the segregation of Ia-afferent and motor axons across the intramuscular nerve branches is not reflected in the locations of Ia terminals on the motoneuron somadendritic surface and that other factors must account for observed EPSP amplitude differences. Our data suggest that the topographic weighting of homonymous Ia-afferent input to cat MG motoneurons is mediated by a gradient of EPSP amplitude rather than by a gradient of Ia connectivity and also suggest that the effect is most prominent in high-rheobase motoneurons.

Publisher Summary This chapter discusses the neural mechanisms of voluntary movements and precentral motor area. The separation of motor and sensory areas is artificial. The spinal projections from the areas are important diagnostic characters for distinguishing areas. The relation of the multiple arborizations to different motoneuron species was determined by retrograde labeling of ulnar, median, and radial motoneurons by HRP injected into the nerves. The same corticospinal (CS) axons supplied at least four motoneuron pools, both ulnar and radial, with boutons de passage and boutons terminaux. Most synapses were presumed to be on distal dendrites, which were not filled with horseradish peroxidase (HRP). Some CS axons crossed the midline and arborized bilaterally. The general distribution was coextensive with that of the arborizations of the group Ia afferent axons. It was suggested that the divergence being revealed is in some ways a corollary of the convergence that had been demonstrated and that any repeatable results of cortical stimulation must be because of the existence of certain condensations or concentrations in these otherwise relatively diffuse neuronal networks.

A study has been made of the depression that occurs when a monosynaptic reflex pathway is subjected to repetitive stimulation. Reflex depression has a dual origin. High frequency or early depression is postsynaptic in origin and results from subnormality in the motoneurons. Low frequency, late, or enduring depression is presynaptic in origin. The conditioning volley-test volley technique and the frequency-mean monosynaptic reflex amplitude relation yield similar information concerning reflex depression. Each method has its advantages and for some purposes one or the other of the methods necessarily must be employed. The results of a variety of experiments are consistent with the proposition that reflex depression in the monosynaptic reflex pathway originates by action in the group IA afferent fibers of muscle origin that are responsible for monosynaptic reflex transmission. Depression is present at a frequency of 0.1 per second (6 per minute) and absent at a frequency of 0.05 per second (3 per minute). Thus it is impractical for most purposes to employ repetition rates that satisfy the requirement for designation as "single shock" stimulations. The temporal course of enduring depression has been determined. It is identical with that for a number of other phenomena observable in monosynaptic reflex pathways, which suggests a common origin. The mechanism of low frequency or enduring depression is discussed in the light of this suggestion.

1. Properties of medial gastrocnemius motoneurones in cats were examined with intracellular electrodes 1-77 days after partial transection of the medial gastrocnemius muscle nerve. 2. Three sequential stages of reaction were defined: (i) firstly, conduction velocity of axotomized motoneurones fell, but e.p.s.p. properties and Ia-motoneurone connectivity were normal; (ii) secondly, e.p.s.p.s became characterized by prolonged rise times and half-widths and by diminished amplitudes; (iii) finally, failures of connectivity were also seen, that is, Ia spindle afferent fibres connected with fewer axotomized than normal homonymous motoneurones. 3. The reason for the change of e.p.s.p. properties cannot be stated with certainty: among the factors may be (i) a change of motoneurone membrane properties and morphology; (ii) partial detachment of synaptic boutons before their total disconnexion, and (iii) detachment of Ia afferent branches terminating on the motoneurone soma before those on the dendrites. 4. The previous conclusion that alteration of e.p.s.p. profiles following motoneurone axotomy was due to total disconnexion of those Ia fibres making synapses on the soma and proximal dendrites ("somatic stripping") must be modified to account for the findings that e.p.s.p. changes are seen before connectivity changes.

THE aim of the present investigation has been to give, in cats, excess use to the synapses of some monosynaptic pathways through the spinal cord, keeping other monosynaptic pathways as controls. In an initial aseptic operation the nerves to some muscles of a synergic extensor group in one hind-limb have been severed and capped to prevent regeneration. Since the remaining muscles of such a synergic group have to substitute for the whole group in supporting the weight of the animal, it can be expected that there will be an excess of stress on them; consequently there will be an increased discharge from their stretch receptors along the group Ia afferent fibres, so giving an increased activation of the excitatory synapses on their motoneurones.