Ib Afferents Research Articles

Modelling of impulse activity of afferent fibers of antagonist muscles during transcutaneous electrical stimulation of the spinal cord during walking

The article describes the results of studies on the impulse activity of various groups of afferent fibers and EMG patterns of lower leg antagonist muscles when walking without, during and after transcutaneous electrical stimulation of the dorsal roots of the lower thoracic spinal cord of a person. Using a mathematical model based on the prediction of the triggering of muscle spindles, variability in the manifestation of impulse activity of various afferents tibialis anterior muscle (TA) and gastrocnemius medialis muscle (GM) when walking under different experimental conditions is shown. It was found that walking on a movable treadmill tape in the absence of spinal cord stimulation was accompanied by strong impulse activity of afferents I (Ia and Ib) and II groups GM, increased excitability of its motorneuron pool and weakening of afferent activity and excitability of TA. On the contrary, electrical stimulation of the spinal cord during walking caused strong impulsive activity of group II TA afferents and moderate — GM, while the activity of Ia fibers TA and GM decreased to moderate impulsivity, Ib afferents of the same muscles had the weakest activity, and the excitability of the GM motorneuron pool was greater than TA. During the postactivation period, walking was accompanied by increased impulses of afferent fibers of group Ib and II GM, weakening of afferent flows of Ib TA and Ia afferents GM, but along with this, afferent signals of group Ia and II to the motorneuron nucleus TA decreased to moderate impulses, and excitability of the motorneuron pool GM was higher than TA. The supposed reflex mechanisms of locomotion regulation are discussed on the basis of well-known phenomena associated with the interaction of various afferent inputs to the spinal cord neuronal apparatus in the system of lower leg antagonist muscles.

Postsynaptic potentials of soleus motor neurons produced by transspinal stimulation: a human single-motor unit study.

Transspinal (or transcutaneous spinal cord) stimulation is a noninvasive, cost-effective, easily applied method with great potential as a therapeutic modality for recovering somatic and nonsomatic functions in upper motor neuron disorders. However, how transspinal stimulation affects motor neuron depolarization is poorly understood, limiting the development of effective transspinal stimulation protocols for rehabilitation. In this study, we characterized the responses of soleus α motor neurons to single-pulse transspinal stimulation using single-motor unit (SMU) discharges as a proxy given the 1:1 discharge activation between the motor neuron and the motor unit. Peristimulus time histogram, peristimulus frequencygram, and surface electromyography (sEMG) were used to characterize the postsynaptic potentials of soleus motor neurons. Transspinal stimulation produced short-latency excitatory postsynaptic potentials (EPSPs) followed by two distinct phases of inhibitory postsynaptic potentials (IPSPs) in most soleus motor neurons and only IPSPs in others. Transspinal stimulation generated double discharges at short interspike intervals in a few motor units. The short-latency EPSPs were likely mediated by muscle spindle group Ia and II afferents, and the IPSPs via excitation of group Ib afferents and recurrent collaterals of motor neurons leading to activation of diverse spinal inhibitory interneuronal circuits. Further studies are warranted to understand better how transspinal stimulation affects depolarization of α motor neurons over multiple spinal segments. This knowledge will be seminal for developing effective transspinal stimulation protocols in upper motor neuron lesions.NEW & NOTEWORTHY Transspinal stimulation produces distinct actions on soleus motor neurons: an early short-latency excitation followed by two inhibitions or only inhibition and doublets. These results show how transspinal stimulation affects depolarization of soleus α motor neurons in healthy humans.

Mechanosensory encoding in ex vivo muscle-nerve preparations.

Our objective was to evaluate an ex vivo muscle-nerve preparation used to study mechanosensory signalling by low threshold mechanosensory receptors (LTMRs). Specifically, we aimed to assess how well the ex vivo preparation represents in vivo firing behaviours of the three major LTMR subtypes of muscle primary sensory afferents, namely type Ia and II muscle spindle (MS) afferents and type Ib tendon organ afferents. Using published procedures for ex vivo study of LTMRs in mouse hindlimb muscles, we replicated earlier reports on afferent firing in response to conventional stretch paradigms applied to non-contracting, that is passive, muscle. Relative to in vivo studies, stretch-evoked firing for confirmed MS afferents in the ex vivo preparation was markedly reduced in firing rate and deficient in encoding dynamic features of muscle stretch. These deficiencies precluded conventional means of discriminating type Ia and II afferents. Muscle afferents, including confirmed Ib afferents were often indistinguishable based on their similar firing responses to the same physiologically relevant stretch paradigms. These observations raise uncertainty about conclusions drawn from earlier ex vivo studies that either attribute findings to specific afferent types or suggest an absence of treatment effects on dynamic firing. However, we found that replacing the recording solution with bicarbonate buffer resulted in afferent firing rates and profiles more like those seen in vivo. Improving representation of the distinctive sensory encoding properties in ex vivo muscle-nerve preparations will promote accuracy in assigning molecular markers and mechanisms to heterogeneous types of muscle mechanosensory neurons.

The acute effects of periodic and noisy tendon vibration on wrist muscle stretch responses

Mechanical muscle tendon vibration activates multiple sensory receptors in the muscle and tendon. In particular, tendon vibration tends to activate the Ia afferents the strongest, but also will activate group II and Ib afferents. This activation can cause three main effects in the central nervous system: proprioceptive illusions, tonic vibration reflexes, and suppression of the stretch response. Noisy tendon vibration has been used to assess the frequency characteristics of proprioceptive reflexes and, interestingly there appeared to be no evidence for proprioceptive illusions or tonic vibration reflexes during standing [9]. However, it remains unknown if noisy vibration induces a suppression of the muscle stretch response. Therefore, the purpose of this study was to investigate the effects of noisy and periodic tendon vibration on the stretch response in the flexor carpi radialis muscle (FCR). We examined FCR stretch responses with and without periodic (20 and 100 Hz) and noisy (∼10–100 Hz) tendon vibration. We additionally had participants perform the task under the instruction set to either not respond to the perturbation or to respond as fast as possible. The key finding from this study was that both periodic and noisy vibration resulted in a reduced stretch response amplitude. Additionally, it was found that a participant’s intent to respond did not modulate the amount of suppression observed. The findings from this study provide a more detailed understanding of the effects of tendon vibration on the muscle stretch response.

Proprioceptive Recognition with Artificial Neural Networks Based on Organizations of Spinocerebellar Tract and Cerebellum.

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.

Neuromodulation by combined sensory and motor stimulation in the peripheral nerve: tendon organ afferent activity

Objective. Neuromuscular stimulation is a therapeutic approach to treat impairments such as stroke or pain, although the resulting inputs to the nervous system due to the stimulation are not well quantified. Stimulation activates both afferents and efferents, and the resulting neural activity is a mix of the effects of both: the changes in afferent activity due to efferent physiological actions plus the alterations due to afferent stimulation. This study quantitatively describes the resulting Golgi tendon organ Ib afferent activity in response to mixed afferent and efferent stimulation. Approach. Neural and stimulated action potentials interact by means of collision, neural resetting, and refractory block. We simulated the action potential patterns of Ib afferents in the human first dorsal interosseous during constant voluntary, stimulated, and combined contractions, varying both stimulation rate and location, and both with and without simultaneous Ib stimulation. Main results. Efferent stimulation affects the firing rate versus force relationships of individual tendon organs, but the effects on the population mean are very small. In contrast, afferent stimulation changes the firing rate versus force relationship of individual afferents by increasing the firing rate, decreasing the force resolution, broadening the distribution of action potential rates, and temporally phase locking a portion of the action potentials. Force resolution is retained in the population mean. The effects of afferent stimulation change with the stimulation rate and location, and with receptor firing rates. Significance. This is the first quantitative description of the changes in afferent feedback during combined efferent and afferent nerve stimulation. The small effects of efferent stimulation on the population response implies that tendon organs could provide accurate force feedback during stimulated contractions. The effects of afferent stimulation can be altered by choice of stimulus rate and site and are generalizable to other afferents.

Descending Inputs to Spinal Circuits Facilitating and Inhibiting Human Wrist Flexors.

Recently we reported in humans that electrical stimulation of the wrist extensor muscle extensor carpi radialis (ECR) could facilitate or suppress the H reflex elicited in flexor carpi radialis (FCR), for inter-stimulus intervals (ISIs) of 30 ms or 70 ms, respectively. The facilitation at 30 ms may be produced by both flexor afferents and extensor Ib afferents acting on a spinal circuit; the origin of the suppression at 70 ms is less certain. In this study, we investigated possible descending inputs to these systems. We used magnetic stimulation of the contralateral primary motor cortex, and click sound stimulation, to activate the corticospinal and the reticulospinal tracts respectively, and measured the effects on the H reflex conditioned by ECR stimulation. Corticospinal inputs reduced both the 30 ms facilitation and 70 ms suppression, indicating corticospinal inhibition of both circuits. By contrast, we failed to show any effect of clicks, either on the H reflex or on its modulation by ECR stimulation. This suggests that click-activated reticulospinal inputs to these circuits may be weak or absent.

Convergent Spinal Circuits Facilitating Human Wrist Flexors.

Noninvasive assessment of spinal circuitry in humans is limited, especially for Ib pathways in the upper limb. We developed a protocol in which we evoke the H-reflex in flexor carpi radialis (FCR) by median nerve stimulation and condition it with electrical stimulation above motor threshold over the extensor carpi radialis (ECR) muscle belly. Eighteen healthy adults (8 male, 10 female) took part in the study. There was a clear reflex facilitation at a 30 ms interstimulus interval (ISI) and suppression at a 70 ms ISI, which was highly consistent across subjects. We investigated the following two hypotheses of the possible source of the facilitation: (1) ECR Ib afferents from Golgi tendon organs, activated by the twitch following ECR stimulation; and (2) FCR afferents, from spindles and/or Golgi tendon organs, activated by the wrist extension movement that follows ECR stimulation. Several human and monkey experiments indicated a role for both of these sets of afferents. Our results provide evidence for a spinal circuit in which flexor motoneurons receive convergent excitatory input from flexor afferents as well as from extensor Ib afferents; this circuit can be straightforwardly assessed noninvasively in humans.SIGNIFICANCE STATEMENT Here we described a novel spinal circuit, which is easy to assess noninvasively in humans. Understanding this circuit in more detail could be beneficial for the design of clinical tests in neurological conditions.

Cardinal features of involuntary force variability can arise from the closed-loop control of viscoelastic afferented muscles.

Involuntary force variability below 15 Hz arises from, and is influenced by, many factors including descending neural drive, proprioceptive feedback, and mechanical properties of muscles and tendons. However, their potential interactions that give rise to the well-structured spectrum of involuntary force variability are not well understood due to a lack of experimental techniques. Here, we investigated the generation, modulation, and interactions among different sources of force variability using a physiologically-grounded closed-loop simulation of an afferented muscle model. The closed-loop simulation included a musculotendon model, muscle spindle, Golgi tendon organ (GTO), and a tracking controller which enabled target-guided force tracking. We demonstrate that closed-loop control of an afferented musculotendon suffices to replicate and explain surprisingly many cardinal features of involuntary force variability. Specifically, we present 1) a potential origin of low-frequency force variability associated with co-modulation of motor unit firing rates (i.e.,‘common drive’), 2) an in-depth characterization of how proprioceptive feedback pathways suffice to generate 5-12 Hz physiological tremor, and 3) evidence that modulation of those feedback pathways (i.e., presynaptic inhibition of Ia and Ib afferents, and spindle sensitivity via fusimotor drive) influence the full spectrum of force variability. These results highlight the previously underestimated importance of closed-loop neuromechanical interactions in explaining involuntary force variability during voluntary ‘isometric’ force control. Furthermore, these results provide the basis for a unifying theory that relates spinal circuitry to various manifestations of altered involuntary force variability in fatigue, aging and neurological disease.

P238 Homonymous Ib inhibition during remote muscle contraction

Objective Conflicting theoretical models exist regarding the mechanism related to the ability of the Jendrassek maneuvre to reinforce reflex parameters. Our objective was to investigate if eventually inhibition of the spinal Ib motoneurone during remote muscle contraction would contribute to H-reflex enhancement. Method Soleus H-reflex from 26 healthy subjects was evoked on both sides by stimulating the tibial nerve at rest and during bilateral vigorous hadgrip. This procedure was repeated using paired stimulation at interstimulus intervals ranging from 2 ms to 15 ms. Mean conditioned H-wave amplitude expressed as a percentage of its unconditioned value (H-ratio) evoked during handgrip was compared with the H-ratio obtained at the same interstimulus interval at rest. Results The size of H-reflex evoked during handgrip was significantly higher then of that evoked at rest. The H-ratio value plotted against the interstimulus interval showed both at rest and during handgrip a similar decrease that was maximal at 5 ms and lasted less than 10 ms. Discussion The size of the conditioned H-reflex is determined by the balance between test Ia excitation and Ib inhibition elicited by the conditioning stimulus. According to our results the interneurones intercalated in pathways from Ib afferents retain their inhibitory effect on the alpha motoneurone during remote muscle contraction. Conclusion H-reflex enhancement during Jendrassek maneuvre is not due to inhibition of Ib interneurones. Significance Further studies are needed in order to test the role of other spinal circuits or descending drives that produce subthreshold excitation of alpha motoneurones by remote muscle contraction.

Muscle proprioceptors in adult rat: mechanosensory signaling and synapse distribution in spinal cord.

The characteristic signaling and intraspinal projections of muscle proprioceptors best described in the cat are often generalized across mammalian species. However, species-dependent adaptations within this system seem necessary to accommodate asymmetric scaling of length, velocity, and force information required by the physics of movement. In the present study we report mechanosensory responses and intraspinal destinations of three classes of muscle proprioceptors. Proprioceptors from triceps surae muscles in adult female Wistar rats anesthetized with isoflurane were physiologically classified as muscle spindle group Ia or II or as tendon organ group Ib afferents, studied for their firing responses to passive-muscle stretch, and in some cases labeled and imaged for axon projections and varicosities in spinal segments. Afferent projections and the laminar distributions of provisional synapses in rats closely resembled those found in the cat. Afferent signaling of muscle kinematics was also similar to reports in the cat, but rat Ib afferents fired robustly during passive-muscle stretch and Ia afferents displayed an exaggerated dynamic response, even after locomotor scaling was accounted for. These differences in mechanosensory signaling by muscle proprioceptors may represent adaptations for movement control in different animal species.NEW & NOTEWORTHY Muscle sensory neurons signal information necessary for controlling limb movements. The information encoded and transmitted by muscle proprioceptors to networks in the spinal cord is known in detail only for the cat, but differences in size and behavior of other species challenge the presumed generalizability. This report presents the first findings detailing specializations in mechanosensory signaling and intraspinal targets for functionally identified subtypes of muscle proprioceptors in the rat.

Analysis of Proprioceptive Sensory Innervation of the Mouse Soleus: A Whole-Mount Muscle Approach.

Muscle proprioceptive afferents provide feedback critical for successful execution of motor tasks via specialized mechanoreceptors housed within skeletal muscles: muscle spindles, supplied by group Ia and group II afferents, and Golgi tendon organs, supplied by group Ib afferents. The morphology of these proprioceptors and their associated afferents has been studied extensively in the cat soleus, and to a lesser degree, in the rat; however, quantitative analyses of proprioceptive innervation in the mouse soleus are comparatively limited. The present study employed genetically-encoded fluorescent reporting systems to label and analyze muscle spindles, Golgi tendon organs, and the proprioceptive sensory neuron subpopulations supplying them within the intact mouse soleus muscle using high magnification confocal microscopy. Total proprioceptive receptors numbered 11.3 ± 0.4 and 5.2 ± 0.2 for muscle spindles and Golgi tendon organs, respectively, and these receptor counts varied independently (n = 27 muscles). Analogous to findings in the rat, muscle spindles analyzed were most frequently supplied by two proprioceptive afferents, and in the majority of instances, both were classified as primary endings using established morphological criteria. Secondary endings were most frequently observed when spindle associated afferents totaled three or more. The mean diameter of primary and secondary afferent axons differed significantly, but the distributions overlap more than previously observed in cat and rat studies.

Are Human Digit Muscles Devoid of Recurrent Inhibition?

Are Human Digit Muscles Devoid of Recurrent Inhibition?

Degradation of mouse locomotor pattern in the absence of proprioceptive sensory feedback.

Mammalian locomotor programs are thought to be directed by the actions of spinal interneuron circuits collectively referred to as "central pattern generators." The contribution of proprioceptive sensory feedback to the coordination of locomotor activity remains less clear. We have analyzed changes in mouse locomotor pattern under conditions in which proprioceptive feedback is attenuated genetically and biomechanically. We find that locomotor pattern degrades upon elimination of proprioceptive feedback from muscle spindles and Golgi tendon organs. The degradation of locomotor pattern is manifest as the loss of interjoint coordination and alternation of flexor and extensor muscles. Group Ia/II sensory feedback from muscle spindles has a predominant influence in patterning the activity of flexor muscles, whereas the redundant activities of group Ia/II and group Ib afferents appear to determine the pattern of extensor muscle firing. These findings establish a role for proprioceptive feedback in the control of fundamental aspects of mammalian locomotor behavior.

Simulating spinal border cells and cerebellar granule cells under locomotion--a case study of spinocerebellar information processing.

The spinocerebellar systems are essential for the brain in the performance of coordinated movements, but our knowledge about the spinocerebellar interactions is very limited. Recently, several crucial pieces of information have been acquired for the spinal border cell (SBC) component of the ventral spinocerebellar tract (VSCT), as well as the effects of SBC mossy fiber activation in granule cells of the cerebellar cortex. SBCs receive monosynaptic input from the reticulospinal tract (RST), which is an important driving system under locomotion, and disynaptic inhibition from Ib muscle afferents. The patterns of activity of RST neurons and Ib afferents under locomotion are known. The activity of VSCT neurons under fictive locomotion, i.e. without sensory feedback, is also known, but there is little information on how these neurons behave under actual locomotion and for cerebellar granule cells receiving SBC input this is completely unknown. But the available information makes it possible to simulate the interactions between the spinal and cerebellar neuronal circuitries with a relatively large set of biological constraints. Using a model of the various neuronal elements and the network they compose, we simulated the modulation of the SBCs and their target granule cells under locomotion and hence generated testable predictions of their general pattern of modulation under this condition. This particular system offers a unique opportunity to simulate these interactions with a limited number of assumptions, which helps making the model biologically plausible. Similar principles of information processing may be expected to apply to all spinocerebellar systems.

Control of ankle extensor muscle activity in walking cats

Our objective was to gain insight into the relative importance of feedforward control and different proprioceptive feedback pathways to ongoing ankle extensor activity during walking in the conscious cat. We asked whether the modulation of stance phase muscle activity is due primarily to proprioceptive feedback and whether the same proprioceptive gains and feedforward commands can automatically generate the muscle activity required for changes in walking slope. To test these hypotheses, we analyzed previously collected muscle activity and mechanics data from cats with an isolated medial gastrocnemius muscle walking along a sloped pegway. Models of proprioceptor dynamics predicted afferent activity from the measured muscle mechanics. We modeled muscle activity as the weighted sum of the activity predicted from the different proprioceptive pathways and a simple model of central drive. We determined the unknown model parameters using optimization procedures that minimized the error between the predicted and measured muscle activity. We found that the modulation of muscle activity within the stance phase and across walking slopes is indeed well described by neural control that employs constant central drive and constant proprioceptive feedback gains. Furthermore, it is force feedback from Ib afferents that is primarily responsible for modulating muscle activity; group II afferent feedback makes a small contribution to tonic activity, and Ia afferent feedback makes no contribution. Force feedback combined with tonic central drive appears to provide a simple control mechanism for automatically compensating for changes in terrain without requiring different commands from the brain or even modification of central nervous system gains.

Positive afferent feedback to the human soleus muscle during quiet standing

In this study we investigated the mechanisms responsible for soleus muscle contraction during quiet standing. Subjects stood on a platform that was randomly moved forward or downward or rotated around the ankle. Downward perturbation caused a short-latency drop in averaged rectified soleus electromyography (SOL EMG). SOL drop increased monotonically with downward acceleration amplitude. Ischemia above the knee abolished or diminished this drop. Ischemia above the ankle had no diminishing effect. Vibration of the Achilles tendon had a diminishing effect on the amplitude of SOL responses. The short-latency drop in SOL observed for downward perturbation might be due to a decrease in positive afferent feedback due to the sudden decrease in body weight. This implies the existence of an ongoing afferent feedback loop toward the SOL motoneuron pool from force-sensitive receptors. Both Ia and Ib afferents probably play a role in the responses observed.

Functional subdivision of feline spinal interneurons in reflex pathways from group Ib and II muscle afferents; an update

A first step towards understanding the operation of a neural network is identification of the populations of neurons that contribute to it. Our aim here is to reassess the basis for subdivision of adult mammalian spinal interneurons that mediate reflex actions from tendon organs (group Ib afferents) and muscle spindle secondary endings (group II afferents) into separate populations. Re-examining the existing experimental data, we find no compelling reasons to consider intermediate zone interneurons with input from group Ib afferents to be distinct from those co-excited by group II afferents. Similar patterns of distributed input have been found in subpopulations that project ipsilaterally, contralaterally or bilaterally, and in both excitatory and inhibitory interneurons; differences in input from group I and II afferents to individual interneurons showed intra- rather than inter-population variation. Patterns of reflex actions evoked from group Ib and II afferents and task-dependent changes in these actions, e.g. during locomotion, may likewise be compatible with mediation by premotor interneurons integrating information from both group I and II afferents. Pathological changes after injuries of the central nervous system in humans and the lineage of different subclasses of embryonic interneurons may therefore be analyzed without need to consider subdivision of adult intermediate zone interneurons into subpopulations with group Ib or group II input. We propose renaming these neurons 'group I/II interneurons'.

Collateral Actions of Premotor Interneurons on Ventral Spinocerebellar Tract Neurons in the Cat

Strong evidence that premotor interneurons provide ventral spinocerebellar tract (VSCT) neurons with feedback information on their actions on motoneurons was previously found for Ia inhibitory interneurons and Renshaw cells, while indications for similar actions of other premotor interneurons were weaker and indirect. Therefore the aim of the present study was to reexamine this possibility with respect to interneurons relaying actions of group Ib afferents from tendon organs and group II afferents from muscle spindles. In all, 133 VSCT neurons in the L3-L5 segments (including 41 spinal border neurons) were recorded from intracellularly in deeply anesthetized cats to verify that stimuli applied in motor nuclei evoked monosynaptic inhibitory postsynaptic potentials (IPSPs) attributable to stimulation of axon collaterals of premotor interneurons. IPSPs were found in over two thirds of the investigated neurons. When intraspinal stimuli were preceded by stimuli applied to a muscle nerve at critical intervals, IPSPs evoked from motor nuclei were considerably reduced, indicating a collision of nerve volleys in axons of interneurons activated by group I and group II afferents. In individual VSCT neurons monosynaptic IPSPs were evoked from both biceps-semitendinosus and gastrocnemius-soleus motor nuclei, in parallel with disynaptic IPSPs from group Ib and group II as well as group Ia afferents. These observations indicate that individual VSCT neurons may monitor the degree of inhibition of both flexor and extensor motoneurons by premotor interneurons in inhibitory pathways from group Ib and group II afferents to motoneurons. They may thus be providing the cerebellum with feedback information on actions of these premotor interneurons on motoneurons.

Interneuronal Activity in Reflex Pathways from Group II Muscle Afferents Is Monitored by Dorsal Spinocerebellar Tract Neurons in the Cat

The main aim of the study was to investigate whether group II muscle afferents contribute to the inhibition of dorsal spinocerebellar tract (DSCT) neurons and thereby modulate information provided by these neurons in the cat. In intracellular recordings, we found disynaptic IPSPs from group II afferents in the majority of DSCT neurons, most often in parallel with IPSPs evoked from group I afferents. In an attempt to identify interneurons that mediate these IPSPs, the second aim of the study, laminas IV-VII in midlumbar segments were searched for interneurons antidromically activated by stimuli applied within Clarke's column. Such interneurons were found in regions in which focal field potentials were evoked by group I and II afferents, or ventral to them, and most were coexcited by these afferents. The input to these interneurons and their location indicate that they belonged to the previously identified population of premotor interneurons in disynaptic pathways between group I and II afferents and hindlimb motoneurons. The study leads thus to the conclusion that inhibitory actions of group II afferents on DSCT neurons are collateral to actions on motoneurons and that DSCT neurons monitor inhibitory actions of group II afferents on motoneurons as closely as they monitor actions of group I afferents. The results also indicate that interneurons mediating disynaptic reflex actions from tendon organ (group Ib) afferents and those mediating disynaptic actions from secondary muscle spindle (group II) afferents to motoneurons may be parts and parcel of the same interneuronal population rather than constitute distinct interneuronal populations.