Segmental Afferents Research Articles

Posteroanterior Cervical Transcutaneous Spinal Cord Stimulation: Interactions with Cortical and Peripheral Nerve Stimulation

Transcutaneous spinal cord stimulation (TSCS) has demonstrated potential to beneficially modulate spinal cord motor and autonomic circuitry. We are interested in pairing cervical TSCS with other forms of nervous system stimulation to enhance synaptic plasticity in circuits serving hand function. We use a novel configuration for cervical TSCS in which the anode is placed anteriorly over ~C4–C5 and the cathode posteriorly over ~T2–T4. We measured the effects of single pulses of TSCS paired with single pulses of motor cortex or median nerve stimulation timed to arrive at the cervical spinal cord at varying intervals. In 13 participants with and 15 participants without chronic cervical spinal cord injury, we observed that subthreshold TSCS facilitates hand muscle responses to motor cortex stimulation, with a tendency toward greater facilitation when TSCS is timed to arrive at cervical synapses simultaneously or up to 10 milliseconds after cortical stimulus arrival. Single pulses of subthreshold TSCS had no effect on the amplitudes of median H-reflex responses or F-wave responses. These findings support a model in which TSCS paired with appropriately timed cortical stimulation has the potential to facilitate convergent transmission between descending motor circuits, segmental afferents, and spinal motor neurons serving the hand. Studies with larger numbers of participants and repetitively paired cortical and spinal stimulation are needed.

Assessment of axonal recruitment using model-guided preclinical spinal cord stimulation in the ex vivo adult mouse spinal cord

Spinal cord stimulation (SCS) is used clinically to limit chronic pain, but fundamental questions remain on the identity of axonal populations recruited. We developed an ex vivo adult mouse spinal cord preparation to assess recruitment following delivery of clinically analogous stimuli determined by downscaling a finite element model of clinical SCS. Analogous electric field distributions were generated with 300-µm × 300-µm electrodes positioned 200 µm above the dorsal column (DC) with stimulation between 50 and 200 µA. We compared axonal recruitment using electrodes of comparable size and stimulus amplitudes when contacting the caudal thoracic DC and at 200 or 600 μm above. Antidromic responses recorded distally from the DC, the adjacent Lissauer tract (LT), and in dorsal roots (DRs) were found to be amplitude and site dependent. Responses in the DC included a unique component not seen in DRs, having the lowest SCS recruitment amplitude and fastest conduction velocity. At 200 μm above, mean cathodic SCS recruitment threshold for axons in DRs and LT were 2.6 and 4.4 times higher, respectively, than DC threshold. SCS recruited primary afferents in all (up to 8) caudal segments sampled. Whereas A and C fibers could be recruited at nearby segments, only A fiber recruitment and synaptically mediated dorsal root reflexes were observed in more distant (lumbar) segments. In sum, clinically analogous SCS led to multisegmental recruitment of several somatosensory-encoding axonal populations. Most striking is the possibility that the lowest threshold recruitment of a nonprimary afferent population in the DC are postsynaptic dorsal column tract cells (PSDCs) projecting to gracile nuclei.NEW & NOTEWORTHY Spinal cord stimulation (SCS) is used clinically to control pain. To identify axonal populations recruited, finite element modeling identified scaling parameters to deliver clinically analogous SCS in an ex vivo adult mouse spinal cord preparation. Results showed that SCS first recruited an axonal population in the dorsal column at a threshold severalfold lower than primary afferents. These putative postsynaptic dorsal column tract cells may represent a previously unconsidered population responsible for SCS-induced paresthesias necessary for analgesia.

Sensory Modulation of Locomotor-Like Membrane Oscillations in Hb9-Expressing Interneurons

The central pattern generator can generate locomotor-like rhythmic activity in the spinal cord in the absence of descending and peripheral inputs, but the motor pattern is regulated by feedback from peripheral sensory inputs that adjust motor outputs to external stimuli. To elucidate the possible role of Hb9-expressing interneurons (Hb9 INs) in the locomotor circuitry, we investigated whether their induced oscillatory activity is modulated by low-threshold afferents in the isolated spinal cords of neonatal Hb9:eGFP transgenic mice. Low-intensity stimulation of segmental afferents generated short-latency, monosynaptic excitatory responses in 62% of Hb9 INs. These were associated with longer-latency (approximately 13 ms) excitatory postsynaptic currents that were evoked in all Hb9 INs, probably by slow conducting afferents that synapse directly onto them. Concomitant morphological analysis confirmed that afferent axons with immunoreactive expression of vesicular glutamate transporter-1 and parvalbumin, presumably from primary afferents, contacted somata and dendrites of all Hb9 INs. Most of the putative synaptic contacts were on distal dendrites that extended to an area with profuse afferent projections. We next examined whether low-threshold afferents in upper (flexor-related) and lower (extensor-related) lumbar segments altered the timing of neurochemically induced locomotor-like rhythms in Hb9 INs and motoneurons. Excitation of flexor-related afferents during the flexor phase delayed the onset of subsequent cycles in both Hb9 INs and segmental motoneurons while maintaining the phase relationship between them. The in-phase correlation between voltage oscillations in Hb9 INs and motor bursts also persisted during the two- to threefold increase in cycle period triggered by extensor-related afferents. Our findings that low-threshold, presumably muscle afferents, synapse directly onto these interneurons and perturb their induced locomotor-like membrane oscillations in a pattern that remains phase-locked with motor bursts support the hypothesis that Hb9 INs are part of the sensorimotor circuitry that regulates the pattern of locomotor rhythms in the isolated cord.

Commissural interneurons with input from group I and II muscle afferents in feline lumbar segments: neurotransmitters, projections and target cells

The aim of this study was to analyse neurotransmitter content, projection areas and target cells of commissural interneurons with input from group I and/or II muscle afferents in lumbar segments in the cat. Axonal projections of 15 intracellularly labelled commissural interneurons were reconstructed. Ten interneurons (nine located in laminae VI-VII, one in lamina VIII) were glutamatergic; only one interneuron (located in lamina VIII) was glycinergic. Contralateral terminal projections were found both in motor nuclei and within laminae VI-VIII. In order to identify target cells of commissural interneurons, effects of stimulation of contralateral group I and II muscle afferents were investigated on interneurons within these laminae. Three tests were used: intracellular records from individual interneurons, modulation of probability of activation of extracellularly recorded interneurons and modulation of their actions on motoneurons using disynaptic PSPs evoked in motoneurons as a measure. All these tests revealed much more frequent and/or stronger excitatory actions of contralateral afferents. The results indicate that commissural interneurons with input from contralateral group I and II afferents target premotor interneurons in disynaptic pathways from ipsilateral group I and II afferents and that excitatory disynaptic actions of contralateral afferents on these interneurons are mediated primarily by intermediate zone commissural interneurons. A second group of commissural interneurons activated by reticulospinal neurons, previously described, frequently had similar, but occasionally opposing, actions to the cells described here, thus indicating that these two subpopulations may act on the same premotor interneurons and either mutually enhance or counteract each other's actions.

Activation and inhibition of the micturition reflex by penile afferents in the cat

Coordination of the urinary bladder and the external urethral sphincter is controlled by descending projections from the pons and is also subject to modulation by segmental afferents. We quantified the effects on the micturition reflex of sensory inputs from genital afferents traveling in the penile component of the somatic pudendal nerve by electrical stimulation of the dorsal nerve of the penis (DNP) in alpha-chloralose anesthetized male cats. Depending on the frequency of stimulation (range, 1-40 Hz), activation of penile afferents either inhibited contractions of the bladder and promoted urine storage or activated the bladder and produced micturition. Stimulation of the DNP at 5-10 Hz inhibited distension-evoked contractions and increased the maximum bladder capacity before incontinence. Conversely, stimulation at 33 and 40 Hz augmented distension-evoked contractions. When the bladder was filled above a threshold volume (70% of the volume necessary for distension-evoked contractions), stimulation at 20-40 Hz activated de novo the micturition reflex and elicited detrusor contractions that increased voiding efficiency compared with distension-evoked voiding. Electrical stimulation of the DNP with a cuff electrode or percutaneous wire electrode produced similar results. The ability to evoke detrusor contractions by activation of the DNP was preserved following acute spinal cord transection. These results demonstrate a clear role of genital afferents in modulating the micturition reflex and suggest the DNP as a potential target for functional restoration of bladder control using electrical stimulation.

On organization of a neuronal network in pathways from group II muscle afferents in feline lumbar spinal segments.

The aim of the study was to verify the hypothesis that trisynaptic actions of group II muscle afferents upon motoneurones are, at least in part, mediated by dorsal horn interneurones exciting the same intermediate zone interneurones that are interposed in disynaptic pathways from group II afferents. Population EPSPs (field potentials) and responses of individual interneurones evoked by group II afferents in the dorsal horn and in the intermediate zone were analysed in order to assess the possibility of a causal relationship between them. When direct actions of group II afferents in the intermediate zone were abolished by presynaptic inhibition, distinct later components of field potentials and delayed interneuronal responses were induced at latencies 0.5-1 ms longer than those seen originally. Both the latency and a marked temporal facilitation define these later group II actions as being evoked disynaptically. Under the same conditions, single stimuli activated more than one half of dorsal horn interneurones, and the second and third stimuli activated all of these interneurones. Responses of dorsal horn interneurones preceded disynaptically evoked responses of intermediate zone interneurones. The study indicates that intermediate zone interneurones may be activated by group II afferents both directly and via dorsal horn interneurones and that synaptic actions of group II afferents upon these interneurones, and their subsequent actions upon motoneurones, may be modulated in parallel at the level of intermediate zone and dorsal horn interneurones.

Spinal circuitry of sensorimotor control of locomotion.

During locomotion many segmental hindlimb reflex pathways serve not only to regulate the excitability of local groups of motoneurones, but also to control the basic operation of the central pattern-generating circuitry responsible for locomotion. This is accomplished through a reorganization of reflexes that includes the suppression of reflex pathways operating at rest and the recruitment during locomotion of previously unrecognized types of spinal interneurones. In addition presynaptic inhibition of transmission from segmental afferents serves to regulate the gain of segmental reflexes and may contribute to the selection of particular reflex pathways during locomotion. The fictive locomotion preparation in adult decerebrate cats has proved to be an important tool in understanding reflex pathway reorganization. Further identification of the spinal interneurones involved in locomotor-dependent reflexes will contribute to our understanding not only of reflex pathway organization but also of the organization of the mammalian central pattern generator.

Sacral spinal interneurones and the control of urinary bladder and urethral striated sphincter muscle function.

Normally, during bladder filling (continence) and expulsion (micturition) there is a reciprocity between the pattern of activity in the urinary bladder sacral parasympathetic efferents and the somatic motoneurones innervating the striated external urethral sphincter muscle. The co-ordination of this pattern of reciprocal activity appears to be determined by excitatory and inhibitory actions of a variety of segmental afferents and descending systems with sacral spinal actions. These actions may in part be mediated through lower lumbar and sacral excitatory and inhibitory spinal interneurones. Over the past 30 years, both neuroanatomical and electrophysiological approaches have been used to reveal an ever-increasing richness in the neuronal network in the lower spinal cord related to the bladder and striated external urethral sphincter muscle. The purpose of this review is to present an overview of the identified excitatory and inhibitory spinal interneurones hypothesized to be involved in the central networks controlling the sacral bladder parasympathetic preganglionic neurones and striated urethral sphincter motoneurones during continence and micturition.

Modulation of synaptic transmission from segmental afferents by spontaneous activity of dorsal horn spinal neurones in the cat.

We examined, in the anaesthetised cat, the influence of the neuronal ensembles producing spontaneous negative cord dorsum potentials (nCDPs) on segmental pathways mediating primary afferent depolarisation (PAD) of cutaneous and group I muscle afferents and on Ia monosynaptic activation of spinal motoneurones. The intraspinal distribution of the field potentials associated with the spontaneous nCDPs indicated that the neuronal ensembles involved in the generation of these potentials were located in the dorsal horn of lumbar segments, in the same region of termination of low-threshold cutaneous afferents. During the occurrence of spontaneous nCDPs, transmission from low-threshold cutaneous afferents to second order neurones in laminae III-VI, as well as transmission along pathways mediating PAD of cutaneous and Ib afferents, was facilitated. PAD of Ia afferents was instead inhibited. Monosynaptic reflexes of flexors and extensors were facilitated during the spontaneous nCDPs. The magnitude of the facilitation was proportional to the amplitude of the 'conditioning' spontaneous nCDPs. This led to a high positive correlation between amplitude fluctuations of spontaneous nCDPs and fluctuations of monosynaptic reflexes. Stimulation of low-threshold cutaneous afferents transiently reduced the probability of occurrence of spontaneous nCDPs as well as the fluctuations of monosynaptic reflexes. It is concluded that the spontaneous nCDPs were produced by the activation of a population of dorsal horn neurones that shared the same functional pathways and involved the same set of neurones as those responding monosynaptically to stimulation of large cutaneous afferents. The spontaneous activity of these neurones was probably the main cause of the fluctuations of the monosynaptic reflexes observed under anaesthesia and could provide a dynamic linkage between segmental sensory and motor pathways.

Nociceptive input to spinal interneurones in reflex pathways from group II muscle afferents in cats.

Effects of noxious stimulation of the skin by radiant heat were tested on responses of first order interneurones in reflex pathways from group II muscle afferents in mid-lumbar, lower-lumbar and sacral segments of the spinal cord. In mid- and lower-lumbar segments both background discharges and monosynaptically evoked responses of intermediate zone interneurones were facilitated. Those of mid-lumbar dorsal horn interneurones were also facilitated suggesting that both these interneuronal populations contribute to the facilitation of flexion reflexes by nociceptors. In contrast, the dominating effects of noxious heat on sacral dorsal horn group II interneurones were inhibitory. The effects evoked by selective activation of C fibres, after A-delta fibres had been blocked by TTX, were similar to those obtained before TTX application.

Field potentials generated by group II muscle afferents in the lower-lumbar segments of the feline spinal cord.

The actions of group II muscle afferents projecting to the lower-lumbar (L6 and L7) segments of the cat spinal cord were investigated by recording the cord dorsum and focal synaptic field potentials evoked by electrical stimulation of hindlimb muscle nerves. Cord dorsum potentials recorded over the lower-lumbar segments were generally much smaller than those produced by group II afferents terminating within the midlumbar and sacral segments. Only group II afferents of tibialis posterior produced potentials with an amplitude (mean maximal amplitude 39 microV, n = 7) approaching that of potentials over other segments. Focal synaptic potentials (mean maximal amplitudes 135-200 microV) were evoked by group II afferents of the following muscle nerves, listed in order of effectiveness: quadriceps, tibialis posterior (throughout L6 and L7), gastrocnemius soleus, flexor digitorum longus, posterior biceps-semitendinosus and popliteus (mainly within L7). Field potentials were recorded in the dorsal horn (laminae IV-V) and also more ventrally in a region which included the lateral part of the intermediate zone (lateral to the large group I intermediate field potentials) and often extended into the ventral horn (laminae V-VII). The latencies of the group II potentials are considered compatible with the monosynaptic actions of the fastest conducting group II muscle afferents. The results are compared with morphological evidence on the pattern of termination of group II muscle afferents in the lower-lumbar segments and with previous descriptions of the actions of group II muscle afferents in midlumbar and sacral segments.

Contribution of group III and IV muscle afferents to multisensorial spinal motor control in cats

The contribution of group III and IV muscle afferents to multisensorial segmental reflex pathways was investigated by testing for spatial facilitation between these afferents and non-nociceptive segmental afferents from skin, muscles and joints on postsynaptic potentials (PSPs) in alpha-motoneurones recorded in anaemically decapitated high spinal cats. Group III and IV muscle afferents were activated by intraarterial injection of potassium chloride (320 mM) or bradykinin triacetate (81 microM). Skin, joint and group I-II muscle afferents were stimulated by graded electrical stimulation of various nerves. Conditioning by stimulation of group III and IV muscle afferents spatially facilitated the transmission in segmental reflex pathways from low- to medium-threshold cutaneous and joint afferents as well as from lb and group II muscle afferents. Both excitatory and inhibitory pathways from these afferents were facilitated. Monosynaptic excitation and disynaptic antagonistic inhibition from group Ia afferents remained unaffected. It is concluded that the spatial facilitation observed between group III and IV muscle afferents and the other afferents indicate a convergence from group III and IV muscle afferents and the other afferents on common interneurones in segmental flexor reflex pathways. Under physiological conditions they thus contribute to the multisensorial feedback of the flexor reflex pathways. Pathophysiologically, the observed convergence may aggravate muscle weakness and atrophy of muscles induced by group III and IV muscle afferents.

Effects of monoamines on transmission from group II muscle afferents in sacral segments in the cat.

The effects of one 5-HT1A serotonin agonist (8-OH-DPAT) and of two alpha 2 noradrenaline agonists (tizanidine and B-HT 933) were tested on the transmission between group II muscle afferents and spinal neurons in the sacral segments of the spinal cord in the cat. These agonists have previously been found to depress transmission from group II muscle afferents either in the dorsal horn or in the intermediate zone of midlumbar segments, and this study addressed the question of whether their actions in the sacral segments are similarly selective. The drugs were applied ionophoretically and their effects were tested on field potentials evoked from group II muscle afferents. As judged by changes in the amplitude of the early components of these field potentials, the transmission is effectively depressed by the serotonin agonist (to 56 +/- 26% after 2 min of ionophoresis of 8-OH-DPAT) but not by the noradrenaline agonists (to 97 +/- 12% after 6 min of ionophoresis of B-HT 933 and to 95 +/- 17% after 6 min of ionophoresis of tizanidine). These data suggest that transmission from group II muscle spindle afferents in the sacral segments is under control of serotonin releasing neurons, as in the dorsal horn of midlumbar segments, but leave open the question of the similarities or differences in the mechanisms (pre- and/or postsynaptic) of this control.

Interneurones in pathways from group II muscle afferents in sacral segments of the feline spinal cord.

1. Properties of dorsal horn interneurones that process information from group II muscle afferents in the sacral segments of the spinal cord have been investigated in the cat using both intracellular and extracellular recording. 2. The interneurones were excited by group II muscle afferents and cutaneous afferents but not by group I muscle afferents. They were most effectively excited by group II afferents of the posterior biceps, semitendinosus, triceps surae and quadriceps muscle nerves and by cutaneous afferents running in the cutaneous femoris, pudendal and sural nerves. The earliest synaptic actions were evoked monosynaptically and were very tightly locked to the stimuli. 3. EPSPs evoked monosynaptically by group II muscle afferents and cutaneous afferents of the most effective nerves were often cut short by disynaptic IPSPs. As a consequence of this negative feedback the EPSPs gave rise to single or double spike potentials and only a minority of interneurones responded with repetitive discharges. However, the neurones that did respond repetitively did so at a very high frequency of discharges (0.8-1.2 ms intervals between the first 2-3 spikes). 4. Sacral dorsal horn group II interneurones do not appear to act directly upon motoneurones because: (i) these interneurones are located outside the area within which last order interneurones have previously been found and (ii) the latencies of PSPs evoked in motoneurones by stimulation of the posterior biceps and semitendinosus, cutaneous femoris and pudendal nerves (i.e. the main nerves providing input to sacral interneurones) are compatible with a tri- but not with a disynaptic coupling. Spatial facilitation of EPSPs and IPSPs following synchronous stimulation of group II and cutaneous afferents of these nerves shows, however, that sacral interneurones may induce excitation or inhibition of motoneurones via other interneurones. 5. Comparison of the properties of group II interneurones in the sacral segments with those of previously studied group II interneurones in the midlumbar segments leads to the conclusion that these two populations of neurones are specialized for the processing of information from different muscles and skin areas. In addition, equivalents of only one of the two subpopulations of midlumbar interneurones have been found at the level of the pudendal nucleus: neurones with input from group II but not from group I muscle afferents. Neurones integrating information from group I and II muscle afferents and in direct contact with motoneurones thus seem to be scarce in the sacral segments.(ABSTRACT TRUNCATED AT 400 WORDS)

Ascending tract neurones processing information from group II muscle afferents in sacral segments of the feline spinal cord.

1. Ascending tract neurones located in the dorsal horn of sacral segments of the spinal cord have been investigated by extracellular and intracellular recording in the anaesthetized cat. The aim was to determine whether information from group II afferents that terminate within the sacral segments is conveyed to supraspinal structures and which types of neurones are involved. 2. A considerable proportion of ascending tract neurones found in the dorsal horn in the same segments as the pudendal (Onuf's) motor nucleus were excited by group II muscle afferents. The great majority (93%) of these neurones had axons ascending in ipsilateral funiculi. Spinocervical tract neurones constituted the largest proportion (82%) of such neurones, while very few spinocerebellar tract and propriospinal neurones and no postsynaptic dorsal column neurones were found among them. 3. In addition to activation by group II muscle afferents all of the neurones were strongly excited by cutaneous afferents. The most potent excitation was evoked by afferents of the posterior biceps-semitendinosus and gastrocnemius muscle nerves and by afferents of the cutaneous femoris, sural and pudendal nerves. The latencies of intracellularly recorded excitatory potentials were indicative of a high incidence of monosynaptic coupling between the afferents and ascending tract neurones. 4. The highly effective monosynaptic excitation of spinocervical tract neurones in the sacral segments by group II afferents is in contrast to the weak disynaptically mediated actions of group II afferents on such neurones in the L6-L7 segments but comparable to the actions of group II afferents on ascending tract neurones in the midlumbar segments. 5. Both the patterns of peripheral input and the latencies of synaptic actions in ascending tract neurones were similar to those in interneurones at the same locations (accompanying report). Similar information is therefore likely to be processed by both categories of neurones. 6. The role of sacral spinocervical tract neurones as a system for transmitting information from group II muscle afferents to supraspinal centres and the potential contribution of this system to the perception of limb position are discussed.

Morphology of interneurones in pathways from group II muscle afferents in sacral segments of the cat spinal cord

The morphology of 12 sacral interneurones with peripheral input from group II muscle afferents was analyzed after intracellular injection of horseradish peroxidase (HRP). The neurones were located in Rexed's laminae III-V overlying the pudendal (Onuf's) motor nucleus. The interneurones had medium sized elongated somata and dendrites projecting radially. All of the interneurones were funicular neurones and fell into two categories depending on whether their axons ran within the dorsal part of the lateral funiculus (DLF; n = 7) or within the ventral funiculus, or the ventral part of the lateral funiculus (VF or VLF; n = 4). The latter were located more rostrally. Within the DLF similar proportions of stem axons and secondary axonal branches descended and ascended. Within the VF and VLF all of the axons ascended. Collaterals of axons running in the DLF arborized primarily within the dorsal horn and the intermediate zone; none were found to approach the motor nuclei. In contrast, collaterals of axons running in the VF/VLF arborized in both the intermediate zone and the ventral horn and passed close to the motor nuclei. We conclude that sacral interneurones with group II input are morphologically nonhomogenous and that only those located most rostrally might have direct actions upon motoneurones. Both the axonal projections and the input (from group II but not from group I muscle afferents and from skin afferents) of sacral interneurones indicate that they are homologous to dorsal horn group II interneurones in the midlumbar segments. They appear, however, to form part of more local neuronal networks than their midlumbar counterparts.

A relay for input from group II muscle afferents in sacral segments of the cat spinal cord.

1. A neuronal relay for input from group II afferents of hindlimb muscle nerves has been found in the previously little explored sacral segments of the cat spinal cord. 2. Electrical stimulation of group II muscle afferents of a number of nerves evoked negative potentials on the surface (cord dorsum potentials) and population postsynaptic potentials (field potentials) within the sacral segments. The largest potentials were evoked by stimulation of the posterior biceps-semitendinosus and triceps surae nerves which evoke much smaller potentials in other segments. Group II afferents of other nerves, notably those which have their main relay within the middle lumbar segments, were much less effective. 3. The sites at which cord dorsum and field potentials evoked by group II muscle afferents were recorded varied in relation to the external topography of the L7-S2 spinal segments but were consistent in their location relative to the pudendal motor nucleus (Onuf's nucleus). Potentials evoked by group II afferents of the posterior biceps and semitendinosus nerves peaked at a level corresponding to the rostral half of Onuf's nucleus and potentials evoked by afferents of the gastrocnemius nerves peaked just rostral to this nucleus. The largest field potentials (of 0.5-1.0 mV) were recorded within the dorsal horn. Field potentials in the intermediate zone were much smaller (< 0.3 mV) and were seen less frequently. 4. Evidence was obtained that the dorsal horn field potentials are to a great extent evoked monosynaptically by the fast conducting fraction of group II muscle afferents: (i) they were evoked at short latencies (2.4-2.7 ms from the stimulus; 1.3-1.7 ms from group I components of afferent volleys and 0.5-0.7 ms from group II components of these volleys), (ii) the conduction times of impulses in the fastest conducting fraction of group II afferents, between the sacral segments (where these impulses were induced by intraspinal stimuli) and the peripheral nerves, were only about 0.5 ms shorter than the latencies of field potentials recorded at the site of intraspinal stimulation and evoked by stimulation of the same peripheral nerves and, (iii) the field potentials followed repetitive stimuli without temporal facilitation. 5. Negative cord dorsum and field potentials were also evoked by small stretches of the semitendinosus and triceps surae muscles. Although they were smaller than potentials evoked by electrical stimulation of sensory fibres and appeared at longer latencies, their presence is consistent with a contribution of muscle spindle afferents to the actions of group II muscle afferents within the sacral segments.(ABSTRACT TRUNCATED AT 400 WORDS)

An intracellular study of perineal and hindlimb afferent inputs onto sphincter motoneurons in the decerebrate cat.

The external urethral sphincter (EUS) and external anal sphincter (EAS) are striated muscles that function to maintain urinary and fecal continence respectively. This study examines the short-latency synaptic input from a variety of cutaneous perineal and muscle/cutaneous hindlimb afferents to the motoneurons innervating these muscles. Intracellular recordings from antidromically identified EUS and EAS motoneurons provided records of the postsynaptic potentials (PSPs) produced by electrical stimulation of peripheral afferents in decerebrate or chloralose anesthetized cats. Excitatory postsynaptic potentials (EPSPs) were produced in most EUS and EAS motoneurons by stimulation of ipsilateral and contralateral sensory pudendal (SPud) and superficial perineal (SPeri) cutaneous nerves. The shortest central latencies in the study (1.5 ms) suggest that there are disynaptic excitatory, in addition to tri- and oligosynaptic, connections within these reflex pathways. EPSPs mixed with longer latency inhibitory potentials (E/I PSPs) were observed in both motoneuron populations, but were found more frequently in EAS motoneurons. These E/I PSPs were evoked more often from contralateral afferents than from ipsilateral afferents. Cutaneous nerves innervating the hindlimb had weaker if any synaptic effects on sphincter motoneurons. Stimulation of ipsilateral hindlimb muscle nerves rarely produced PSPs in EUS motoneurons and had weak synaptic actions on EAS motoneurons. In 2 of 22 animals (both decerebrate), large inhibitory potentials predominated over early small EPSPs suggesting that inhibitory pathways from these afferents to sphincter motoneurons can be released under certain circumstances. The relation between the segmental afferents to EUS and EAS motoneurons and the neural circuitry influencing them during micturition and defecation are discussed.

Further evidence for synaptic actions of muscle spindle secondaries in the middle lumbar segments of the cat spinal cord.

1. The aim of this study has been to investigate the receptor origin of postsynaptic actions evoked by group II muscle afferents in mid-lumbar segments of the cat spinal cord. The experiments tested the hypothesis that the afferents involved were the secondary endings of muscle spindles. 2. Spindle afferents were activated by contractions of intrafusal muscle fibres which were induced by electrical stimulation of fusimotor axons in the distal parts of transected ventral roots by one to three stimuli at 150-500 stimuli/s. A separate series of experiments has shown that such stimuli are effective in activating a considerable proportion of muscle spindle secondaries when contractions of extrafusal muscle fibres are eliminated by differential fatigue of these fibres, provided that several fusimotor axons are stimulated simultaneously. 3. Extracellular field potentials were recorded in the dorsal horn, at such locations where synaptic actions were evoked by electrical stimulation of group II but not group Ia muscle spindle or group Ib tendon organ afferents of pretibial flexors. Effects of activation of spindle afferents following stimulation of fusimotor axons were then compared with effects evoked by electrical stimulation of group II afferents of anterior tibial or extensor digitorum longus nerves and by small stretches of these muscles. 4. Distinct field potentials were evoked by stimulation of ventral root fibres at all locations at which field potentials were obtained from group II afferents stimulated electrically. The latencies of these field potentials were in both cases shorter in the dorsal horn than in the ventral horn. 5. The appearance of these field potentials was not related to contractions of extrafusal muscle fibres and was also observed when these contractions were practically eliminated. Furthermore, their threshold and similar dependence on a potentiating effect of two to three stimuli, as found for single secondaries, allow them to be attributed to secondary endings of muscle spindles.

An investigation of local actions of ionophoretically applied DOPA in the spinal cord.

Methyl-L-DOPA (L-beta-3,4-dihydroxy-phenylalanine methyl ester, hydrochloride) was applied ionophoretically to investigate its effects on neurones at various locations in the cat spinal cord. Its actions were tested on monosynaptic field potentials evoked from group I and group II muscle afferents in midlumbar segments. Methyl-L-DOPA has been found to depress field potentials evoked from group II afferents in the ventral horn and in the intermediate zone but not in the dorsal horn, nor field potentials evoked from group I afferents. Its effects were the same as those of systemically applied L-DOPA (L-beta-3,4-dihydroxyphenylalanine), although weaker.