Reflex control of ipsilateral and contralateral paraspinal muscles

Homonymous and heteronymous reflex connections of the paraspinal muscles were investigated by the application of a tap to the muscle bellies of the lumbar multifidus and iliocostalis lumborum muscles and observation of surface electromyographic responses in the same muscles on both sides of the trunk. Reflexes were evoked in each of the homonymous muscles with latencies and estimated conduction velocities compatible with being evoked by Ia muscle afferents and having a monosynaptic component. Short latency heteronymous excitatory reflex connections were observed in muscles on the ipsilateral side, whilst reflex responses in the contralateral muscles were inhibitory in response to the same stimulus. The latencies of the crossed responses were on average 9.1 ms longer than the ipsilateral excitatory responses. These results are in contrast to the crossed excitatory responses observed between the abdominal muscles and trapezius muscles on the opposite aspect of the trunk. Such a difference in the reflex pathways between these two groups of trunk muscles compliments the different anatomical arrangement of the muscle groups and suggests a contribution to their commonly observed activation patterns.     

Pattern of monosynaptic heteronymous Ia connections in the human lower limb

Changes in the firing probability of single motor units in response to electrical stimulation of muscle nerves were used to derive the projections of muscle spindle Ia afferents to the motoneurones of various leg and thigh muscles. Discharges of units in soleus, gastrocnemius medialis, peroneus brevis, tibialis anterior, quadriceps, biceps femoris and semitendinosus were investigated after stimulation of inferior soleus, gastrocnemius medialis, superficial peroneal, deep peroneal and femoral nerves. Homonymous facilitation, occurring at the same latency as the H reflex and therefore attributed to monosynaptic Ia EPSPs, was found in virtually all the sampled units. In many motor nuclei an early facilitation was also evoked by heteronymous low-threshold afferents. The heteronymous facilitation was considered to be mediated through a monosynaptic pathway when the difference between the central latencies of heteronymous and homonymous peaks was not more than 0.2 ms. The heteronymous Ia connections were widely distributed. In particular, monosynaptic coupling between muscles operating at different joints appears to be the rule in humans, though it is rare between ankle and knee muscles in the cat and the baboon.     

Neuronal pathways for spinal reflexes activated by group I and group II muscle afferents in the spinal segment (Co1) innervating the tail in the low spinalized cat

 We studied neuronal pathways for spinal reflexes activated by group-I and group-II muscle afferents in the spinal segments innervating the tail in unanesthetized and spinalized (L1) cats. Experiments were performed on 25 adult cats of both sexes. The effects of stimulating nerves innervating six tail muscles on both sides were recorded from tail motoneurons in the first coccygeal spinal segment (Co1) using glass microelectrodes. Stable recordings were obtained from 150 tail motoneurons. Stimulation of group-I muscle afferents (stimulus intensity <1.8 T) often produced EPSPs (82/150) after stimulating nerves innervating neighboring tail muscles. Motoneurons innervating the long-tendoned muscles, M. extensor caudae lateralis and M. flexor caudae longus (ECL and FCL), received heteronymous monosynaptic connections from group-I muscle afferents innervating the ipsilateral tail muscles. The motoneurons innervating segmental muscles, M. extensor caudae medialis and M. flexor caudae brevis (ECM and FCB), received heteronymous monosynaptic connections from group-I muscle afferents innervating tail muscles on both sides. The motoneurons innervating tail muscles originated from the Ossa coxae, M. abductor caudae externus and M. abductor caudae internus (ACE and ACI), received monosynaptic connection from group-I muscle afferents innervating most of the tail muscles on both sides. Crossed disynaptic inhibitory pathways activated by primary muscle afferent inputs were observed in ECM, ACE, FCL, and FCB motoneurons. The effects of group-II afferent inputs were not dependent on the kind of motoneuron, and alternative excitatory and inhibitory pathways were not clearly observed in the tail motoneuron pool. It is suggested that variability of the neuronal pathways from group-I and -II muscle afferents to tail motoneurons corresponds to functional relationships among tail muscles, depending on the tail movements. Received: 9 April 1997 / Accepted: 8 August 1998     

The bilateral reflex control of the trapezius muscle in humans

Abstract. Electromyographic recordings were made from the trapezius muscle in 18 healthy subjects using surface electrodes placed bilaterally. A mechanical tap applied to the insertion of the lower fibres of trapezius evoked a reflex at 11.9 +/- 1.4 ms (mean plus minus SD) in the ipsilateral trapezius muscle. In addition and surprisingly, short latency, facilitatory reflexes were also seen on the contralateral side at a latency of 14.3 +/- 1.7 ms. Electrical stimulation of the afferent nerve to trapezius, the cervical nerve of C3/4, also evoked short latency, facilitatory reflexes from both the ipsilateral and contralateral muscles. Since this nerve carries the Ia fibres from trapezius, this reflex would appear to be the equivalent of the H reflex seen in the soleus muscle. The latency of this reflex was 10.9 +/- 1.4 ms ipsilaterally and 11.9 +/- 1.5 ms contralaterally, the difference being only 1.0 +/- 0.7 ms. Voluntary activity of the ipsilateral trapezius increased the amplitude of both mechanically and electrically evoked ipsilateral and crossed reflexes, and vibration of the ipsilateral trapezius decreased the amplitude of both reflexes. These results together suggest that the earliest parts of both ipsilateral and crossed reflexes are monosynaptic in origin. If the crossed reflex is, in part, due to a common Ia presynaptic input, correlation of bilateral trapezius activity should produce a peak in the cross-correlogram. Indeed, cross-correlation of the bilateral trapezius activity during elevation of the arms produced a short duration peak. Moreover, this was significantly larger when compared to that constructed from activity of the upper and lower fibres of trapezius from one side. These results imply that muscle spindle afferents from the ipsilateral trapezius monosynaptically activate motoneurones contralaterally.     

The actualization of heteronomous monosynaptic connections of IA afferents in the muscle motor centers of the human hand

Reflex excitability of spinal centers of hand muscles was examined in normal subjects and patients with traumatic lesion of forearm nerves. Central and peripheral muscle responses were evoked by stimulation of homonymous and heteronymous nerves. Patients with median and/or ulnar nerve lesion showed the heteronymous abductor pollicis brevis and abductor digit minimi H-responses. The cause of appearance of heteronymous responses is supposed to be actualization of latent heteronymous monosynaptic connections following traumatic lesion and afferent deprivation. It seems obvious that activation of heteronymous pathways corresponds to the process in afferent system known as "revitalization" of median-to-ulnar anastomoses after damage.     

Effects on the fusimotor-muscle spindle system induced by intramuscular injections of hypertonic saline

Intramuscular injection of hypertonic saline (HS) is a procedure widely adopted to experimentally induce deep muscle pain in humans. This study was undertaken to test whether intramuscular injections of HS (5%) influence the activity of primary and secondary muscle spindle afferents (MSAs) from homonymous as well as heteronymous muscles. The experiments were performed on six cats anaesthetised with alpha-chloralose. Usually responses of two to nine MSAs from gastrocnemius medialis (GM) and/or gastrocnemius lateralis (GL) muscles were recorded simultaneously, while HS was injected either into the receptor-bearing muscle (homonymous responses) or into a close (GM/GL) or remote synergistic muscle (posterior biceps, PB, heteronymous responses). The mean rate of discharge and the depth of modulation of the MSA responses to sinusoidal stretching of the receptor-bearing muscle were calculated. Out of the 42 afferents tested (7 from GM and 35 from GL), 38 (90%) exhibited statistically significant responses to injections of HS into homonymous and/or heteronymous muscles. With injections into the homonymous muscle, the average maximal increase in mean rate of discharge was 74% and the average decrease in depth of modulation was --18%. The mean duration of the effects was 2.1 min. The corresponding values for heteronymous injections into a close synergist were 87%, -17% and 2.1 min (GM or GL), and for injections into PB 52%, -11%, and 1.8 min. The majority of the responses (72%) were compatible with reflex action on static fusimotor neurones, whereas 20% of the responses could be attributed to mixed static and dynamic fusimotor action. The remaining 8% of the responses were attributed to inhibition of fusimotor activity. There were no statistically significant differences between the responses following injections into homonymous or heteronymous muscles. Injections of Tyrode's solution did not induce any significant alterations in MSA responses, implying that they were not induced by direct and/or injury effects of the injections. HS-related changes in MSA activity were completely abolished after the nerves to corresponding muscles were cut, confirming the reflex nature of the effects. Thus, intramuscular injections of HS induce reflex changes in MSA activity from both homonymous and heteronymous muscles, most likely via fusimotor reflexes. Predominantly static fusimotor neurones were activated. The possible role of the fusimotor-muscle spindle system in altered motor control during experimentally induced muscle pain is discussed.     

Synaptic connections from large afferents of wrist flexor and extensor muscles to synergistic motoneurones in man

 Short-latency excitatory Ia reflex connections were determined between pairs of human wrist flexor and extensor muscles. Spindle Ia afferents were stimulated by either tendon tap or electrical stimulation. The activity of voluntarily activated single motor units was recorded intramuscularly from pairs of wrist flexor or extensor muscles. Cross-correlation between stimuli and the discharge of the motor units provided a measure of the homonymous or heteronymous excitatory input to a motoneurone. Homonymous motoneurone facilitation was generally stronger than that of the heteronymous motoneurones. The principal wrist flexors, flexor carpi radialis (FCR) and flexor carpi ulnaris (FCU), were tightly connected through a bidirectional short-latency reflex pathway. In contrast, the extensor carpi ulnaris (ECU) and the extensor carpi radialis (ECR) did not have similar connections. ECU motoneurones received no short-latency excitatory Ia input from the ECR. ECR motoneurones did receive excitatory Ia input from ECU Ia afferents; however, its latency was delayed by several milliseconds compared with other heteronymous Ia excitatory effects observed. The wrist and finger extensors were linked through heteronymous Ia excitatory reflexes. The reflex connections observed in humans are largely similar to those observed in the cat, with the exception of heteronymous effects from the ECU to the ECR and from the extensor digitorum communis (EDC) to the ECU, which are present only in humans. The differences in the reflex organization of the wrist flexors versus the extensors probably reflects the importance of grasping. Received: 19 August 1996 / Accepted: 6 March 1997     

Reflex pathways connect receptors in the human lower leg to the erector spinae muscles of the lower back

Reflex pathways connect all four limbs in humans. Presently, we tested the hypothesis that reflexes also link sensory receptors in the lower leg with muscles of the lower back (erector spinae; ES). Taps were applied to the right Achilles’ tendon and electromyographic activity was recorded from the right soleus and bilaterally from ES. Reflexes were compared between sitting and standing and between standing with the eyes open versus closed. Reflexes were evoked bilaterally in ES and consisted of an early latency excitation, a medium latency inhibition, and a longer latency excitation. During sitting but not standing, the early excitation was larger in the ES muscle ipsilateral to the stimulation (iES) than in the contralateral ES (cES). During standing but not sitting, the longer latency excitation in cES was larger than in iES. This response in cES was also larger during standing compared to sitting. Responses were not significantly different between the eyes open and eyes closed conditions. Taps applied to the lateral calcaneus (heel taps) evoked responses in ES that were not significantly different in amplitude or latency than those evoked by tendon taps, despite a 75–94% reduction in the amplitude of the soleus stretch reflex evoked by the heel taps. Electrical stimulation of the sural nerve, a purely cutaneous nerve at the ankle, evoked ES reflexes that were not significantly different in amplitude but had significantly longer latencies than those evoked by the tendon and heel taps. These results support the hypothesis that reflex pathways connect receptors in the lower leg with muscles of the lower back and show that that the amplitude of these reflexes is modulated by task. Responses evoked by stimulation of the sural nerve establish that reflex pathways connect the ES muscles with cutaneous receptors of the foot. In contrast, the large volley in muscle spindle afferents induced by the tendon taps compared to the heel taps did not alter the ES responses, suggesting that the reflex connection between triceps surae muscle spindles and the ES muscles may be relatively weak. These heteronymous reflexes may play a role in stabilizing the trunk for maintaining posture and balance.     

Influence of synaptic identity on single-Ia-afferent connectivity and EPSP amplitude in the adult cat: homonymous versus heteronymous connections.

1. This study makes use of the pattern of synaptic connections between motoneurons and Ia afferents of triceps surae muscles in the cat to test the relative importance of synaptic identity, neuronal size, and neuronal topography as determinants of Ia-afferent connectivity and excitatory postsynaptic potential (EPSP) amplitude. 2. The synaptic actions of single-Ia medial gastrocnemius (MG) afferents were measured by intracellular recording in MG and lateral gastrocnemius (LG) motoneurons. The spike-triggered averaging technique was used to measure EPSPs generated by homonymous or heteronymous Ia afferents and motoneurons, i.e., neurons supplying the same or different muscles, respectively. In agreement with earlier studies, the pooled sample showed that the number of functional connections and the size of EPSPs were both significantly greater for homonymous than for heteronymous neurons. 3. Afferent conduction velocity, motoneuron conduction velocity, rheobase current, and position of the motoneuron relative to the spinal cord afferent entry were all correlated with EPSP amplitude, but the amplitude difference between homonymous and heteronymous connections remained significant after the statistical removal analysis of covariance (ANCOVA) of the contribution of these variables. Stepwise multiple-regression analysis showed that synaptic identity explained the greatest fraction of the variance in EPSP amplitude (9%), with significant but smaller fractions accounted for by rheobase current or motoneuron conduction velocity. 4. In a separate experiment, the monosynaptic affects from both homonymous and heteronymous single-Ia afferents were examined in each of 88 MG or LG motoneurons. The single-Ia afferents used in this portion of the study were sampled from both MG and LG muscles and selected for similar conduction velocities and spinal cord entry points.(ABSTRACT TRUNCATED AT 250 WORDS)     

Reciprocal excitation of muscle antagonists by the primary afferent pathway

Summary Measurements were made from electromyograms, evoked in human soleus and tibialis anterior muscles by tendon taps. Evidence is presented to suggest that in certain motor disorders it is possible to stimulate the stretch reflex afferents from a single muscle and elicit activity both in that muscle and its antagonist at comparable latencies and levels. Responses in both muscles occur at the short latencies of an oligosynaptic or even possibly a monosynaptic pathway.We discuss the proposition that the traditional concept of spinal reciprocal innervation as an inhibitory linking of muscle antagonists must be expanded to include both inhibitory and excitatory connections. Such reciprocal excitatory connections appear pronounced in spastic patients who have suffered perinatal injuries to the immature nervous system.     

Specificity of electrical coupling among neurons innervating forelimb muscles of the adult bullfrog

1. The specificity of electrical connections among sensory fibers and motoneurons in the bullfrog's spinal cord was studied by recording intracellularly from brachial motoneurons. Synaptic potentials evoked by stimulation of individual muscle nerves were recorded in normal and reduced-calcium solutions and after acute section of dorsal or ventral roots. 2. Homonymous motoneurons are electrically coupled. After the dorsal roots were cut to abolish sensory input, short-latency potentials were almost always evoked anti-dromically in a motoneuron by stimulation of its own muscle nerve but rarely by stimulation of nerves innervating other muscles. These potentials differed from chemically mediated synaptic potentials in this preparation; they had a shorter latency and remained after perfusion with reduced-calcium solutions. This evidence suggests that they are mediated electrically. 3. Some motoneurons that innervate functionally equivalent muscles are electrically coupled. Approximately two-thirds of the motoneurons innervating the internal or external heads of the triceps muscles received coupling potentials on stimulation of the other nerve, although never on stimulation of the heteronymous, medial, triceps nerve. 4. The monosynaptic potentials evoked by muscle sensory afferents in motoneurons often have both electrical and chemical components. The electrical component occurred with short delay and persisted in reduced-calcium solutions. The chemical component occurred 1.5-2.0 ms later, at 14 degrees C, and was abolished by reducing calcium in the bathing solution. Muscle sensory afferents make these mixed synapses on homonymous, heteronymous, and other motoneurons.     

Monosynaptic connexions of low threshold muscle afferents with hindlimb motoneurones in the turtle spinal cord

Summary Excitatory postsynaptic potentials (EPSPs) were recorded intracellularly from hindlimb motoneurones of the anaesthetized fresh water turtle. The EPSPs were evoked from low threshold muscle afferents and the amplitudes saturated for stimuli less than two times the nerve threshold. The segmental latencies of these EPSPs, measured from the initial positive peak of the triphasic cord dorsum potential to the onset of the EPSP, ranged from 1.5 to 3.1 ms. The intraspinal conduction time of afferents was estimated by recording afferent volleys in the grey matter along the vertical course of intraspinal afferent fibres. The synaptic delay was estimated by subtracting the latency of the afferent volley at the deepest region of the dorsal horn from the segmental latency of the EPSP (in the range from 1.6 to 2.1 ms) recorded in the same microelectrode track. The average value was 0.99 ms (range: 0.9–1.1 ms), which was close to the known synaptic delay of cold-blooded animals. Therefore, the EPSPs in this range of segmental latencies were regarded as monosynaptic. Taking account of the intraspinal afferent conduction time (0.8 ms on average) and another synaptic delay, the latency for disynaptic transmission would be 2.8 ms or more. Thus, EPSPs having segmental latencies of 1.5–3.1 ms were suggested to be almost all monosynaptic in nature, at least under the present conditions of deep anaesthesia. On the basis of the above criteria for the monosynaptic nature of EPSPs, the pattern of convergence of monosynaptic excitatory inputs from various muscle afferents was investigated. Monosynaptic EPSPs were induced from the homonymous muscle nerve and the nerve innervating the synergist at the same joint. The heteronymous EPSPs were also found between muscles within each group of the anterodorsal musculature and the posteroventral musculature. No monosynaptic connexions were found between anterodorsal and posteroventral muscles except between the muscles innervated by the peroneal and the tibial nerve.     

Stretch and H reflexes in triceps surae are similar during tonic and rhythmic contractions in high decerebrate cats

During locomotion in decerebrate and spinal cats the group Ia afferents from hind leg muscles are depolarized rhythmically. An earlier study concluded that this locomotor-related primary afferent depolarization (PAD) does not contribute to modulation of monosynaptic reflex pathways during locomotion. This finding indicated that the neural network generating the locomotor rhythm, the central pattern generator (CPG), does not presynaptically inhibit monosynaptic reflexes. In this investigation we tested this prediction in decerebrate cats by measuring the magnitude of reflexes evoked in ankle extensor muscles during periods of tonic contractions and during sequences of rhythmic contractions. The latter occurred when the animal was induced to walk on a treadmill. At the similar levels of activity in the soleus muscle there was no significant difference in the magnitude of the soleus H reflex in these two behavioral situations. Similar results were obtained for reflexes evoked by brief stretches of the soleus muscle. We also examined the reflexes evoked by ramp-and-hold stretches during periods of rhythmic and tonic activity of the isolated medial gastrocnemius (MG) muscle. At similar levels of background activity, the reflexes evoked in the MG muscle were the same during rhythmic and tonic contractions. Our failure to observe a reduction in the magnitude of H reflexes and stretch reflexes during rhythmic contractions, compared with reflexes evoked at the same level of background activity during tonic contractions, is consistent with the notion that the CPG for stepping does not presynaptically inhibit monosynaptic reflexes during the extension phase of locomotor activity. Our results indicate that presynaptic inhibition of the monosynaptic reflex associated with normal locomotion in cats or humans arises from sources other than the extensor burst generating system of the central pattern generator.     

Heteronymous reflex connections in human upper limb muscles in response to stretch of forearm muscles

Torque motor produced stretch of upper limb muscles results in two distinct reflex peaks in the electromyographic activity. Whereas the short-latency reflex (SLR) response is mediated largely by the spinal monosynaptic reflex pathway, the longer-latency reflex (LLR) is suggested to involve a transcortical loop. For the SLRs, patterns of heteronymous monosynaptic Ia connections have been well-studied for a large number of muscles in the cat and in humans. For LLRs, information is available for perturbations to proximal joints, although the protocols for most of these studies did not focus on heteronymous connections. The main objective of the present study was to elicit both SLRs and LLRs in wrist flexors and extensors and to examine heteronymous connections from these muscles to elbow flexors (biceps brachii; BiBr) and extensors (triceps brachii; TriBr) and to selected distal muscles, including abductor pollicis longus (APL), first dorsal interosseous (FDI), abductor digiti minimi (ADM), and Thenars. The stretch of wrist flexors produced SLR and LLR peaks in APL, FDI, ADM, Thenars, and BiBr while simultaneously inducing inhibition of wrist extensors and TriBr. When wrist extensors were stretched, SLR and LLR peaks were observed in TriBr, whereas the primary wrist flexors, APL and BiBr, were inhibited; response patterns of FDI, ADM, and Thenars were less consistent. The main conclusions from the observed data are that: 1) as in the cat, afferents from wrist flexors and extensors make heteronymous connections with proximal and distal upper limb muscles; and 2) the strength of heteronymous connections is greater for LLRs than SLRs in the distal muscles, whereas the opposite is true for the proximal muscles. In the majority of observations, SLR and LLR excitatory peaks were observed together. However, on occasion, LLRs were observed without the SLR response in hand muscles when wrist extensors were stretched.     

Corticospinal activation of internal oblique muscles has a strong ipsilateral component and can be lateralised in man

Trunk muscles receive corticospinal innervation ipsilaterally and contralaterally and here we investigate the degree of ipsilateral innervation and any cortical asymmetry in pairs of trunk muscles and proximal and distal limb muscles. Transcranial magnetic stimulation (TMS) was applied to left and right motor cortices in turn and bilateral electromyographic (EMG) recordings were made from internal oblique (IO; lower abdominal), deltoid (D; shoulder) and first dorsal interosseus (1DI; hand) muscles during voluntary contraction in ten healthy subjects. We used a 7-cm figure-of-eight stimulating coil located 2 cm lateral and 2 cm anterior to the vertex over either cortex. Incidence of ipsilateral motor evoked potentials (MEPs) was 85% in IO, 40% in D and 35% in 1DI. Mean (± S.E.M.) ipsilateral MEP latencies were longer (P<0.05; paired t-test) than contralateral MEP latencies (contralateral vs. ipsilateral; IO: 16.1±0.4 ms vs. 19.0±0.5 ms; D: 9.7±0.3 ms vs. 15.1±1.9 ms; 1DI: 18.3±0.6 ms vs. 23.3±1.4 ms), suggesting that ipsilateral MEPs were not a result of interhemispheric current spread. Where data were available, we calculated a ratio (ipsilateral MEP areas/contralateral MEP areas) for a given muscle (IO: n=16; D: n=8; 1DI: n=7 ratios). Mean values for these ratios were 0.70±0.20 (IO), 0.14±0.05 (D) and 0.08±0.02 (1DI), revealing stronger ipsilateral drive to IO. Comparisons of the sizes of these ratios revealed a bias towards one cortex or the other (four subjects right; three subjects left). The predominant cortex showed a mean ratio of 1.21±0.38 compared with 0.26±0.06 in the other cortex (P<0.05). It appears that the corticospinal control of IO has a strong ipsilateral component relative to the limb muscles and also shows hemispheric asymmetry.     

Effects of flexor reflex afferent stimulation on the soleus H reflex in patients with a complete spinal cord lesion: evidence for presynaptic inhibition of Ia transmission

Summary The effects of electrically stimulating the Flexor Reflex Afferent (FRA) on the soleus H reflexes were investigated in 34 paraplegic patients having a clinically complete spinal cord lesion. Conditioning stimuli (5–50 mA) were applied to the ipsilateral or contralateral sural nerve. The conditioning-test interval ranged from 20 to 1000 ms. A late ipsilateral flexor reflex (EMG) was found in all patients. A late contralateral extension reflex was sporadically observed in only 3 patients. The excitability curves usually showed two phases of ipsilateral H reflex inhibition and contralateral H reflex facilitation, one between 50 and 130 ms and the other after over 200 ms. These intervals correspond to early and late flexion reflexes. With high intensity stimulation the early and late ipsilateral inhibition fused. An early low threshold ipsilateral facilitation occured in 9 patients. The contralateral late facilitation was followed by prolonged inhibition in 10 patients. Changes in presynaptic inhibition were assessed by measuring the heteronymous monosynaptic Ia facilitation from quadriceps to soleus. For methodological reasons, it was only possible to investigate the effect of contralateral conditioning volleys which was performed in 5 patients. A significant and regular reduction of the heteronymous Ia facilitation was found in 4 patients. This reduction is taken to indicate that the FRA evokes presynaptic inhibition of Ia transmission to alpha motoneurones. Presynaptic inhibition was also indicated by the enhancement of a vibratory stimulus induced inhibition in 2 subjects. These results are consistent with the hypothesis that the reflex organization in patients with a spinal cord section is similar to that of the acute spinal cat injected with DOPA.     

Synaptic inputs from low threshold afferents of trunk muscles to motoneurons innervating the longissimus lumborum muscle in the spinal cat

(1) We studied the reflex actions of group I and II afferents to longissimus lumborum (Long) motoneurons in the L1–L5 spinal segments from the epaxial muscle, m. Long, and the hypaxial muscle, m. obliquus externus abdominus (OEA). (2) Postsynaptic potentials (PSPs) recorded from 140 Long motoneurons in 30 spinal cats were analyzed. Under the present experimental conditions, the stimulation of Long and OEA nerves at an intensity below 1.5 times threshold (T) activated only group I muscle afferents, while stimulations at 2–5T activated group II muscle afferents as well. (3) The incidence of PSPs was related to the proximity of the spinal segments of the nerves stimulated to the spinal segment of the motoneurons; the shorter the distance the larger the PSPs and higher incidence of PSPs. The Long motoneurons received group I afferent input mainly from the same and adjacent segments, and received group II afferent inputs from a wider range of segments. (4) A short (i.e., less than 1.0 ms) latency of excitatory PSPs (EPSPs) evoked by ipsilateral group I afferents of Long at the same or adjacent segment indicated a monosynaptic connection. In general, the central latencies became longer as the distance between spinal segments of stimulated nerves and motoneurons increased. Major PSP components were produced by polysynaptic neuronal pathways. The spatial facilitation between PSPs evoked by afferents of different nerves (i.e., ipsilateral Long (iLong) and contralateral Long (cLong) of the same segment; iLongs of different segments; and iLong and iOEA of the same segment) indicated that they shared common interneurons. (5) Although iLong and iOEA muscle afferents produced predominantly EPSPs, and contralateral muscle afferents elicited predominantly IPSPs in Long motoneurons at each spinal segment, the patterns of convergence from Long and OEA muscle afferents of different spinal segments and of different sides differed considerably among motoneurons. (6) These findings demonstrated various input patterns of individual motoneurons within the same motoneuron pool, which might reflect the complexity of neuronal control of the back muscles for various trunk movements, including lateral and dorsal bending, rotating, and fixation of the trunk. Electronic Publication     

Exteroceptive Influences on the Lumbar Back Muscle Tone and Reflexes in the Cat

The exteroceptive influences on tonic activity and on stretch reflexes of the longissimus dorsi and multifidus spinae muscles were investigated in decerebrate, spinal and chloralose-anesthetized cats. Adequate skin stimulation was used to map out facilitatory and inhibitory skin areas on the trunk and the extremities. On the trunk facilitatory areas are relatively large and located at the dorsal side while inhibitory areas are confined to the ventrolateral part of the contralateral body half. The facilitatory skin fields are of approximately the same size in decerebrate and spinal cats. On leg skin stimulation facilitation can be evoked from the ipsilateral hind limb while inhibition results from stimulation of the other limbs. Spinal cord transection increased excitatory effects of ipsilateral hind limb stimulation. Reflex responses in the back muscles to applied stretch are described. These reflexes were used as test reflexes in experiments with conditioning stimulation of the peripheral nerves supplying skin areas from which effects on back muscle activity were evoked by adequate stimulation. The conditioning-test experiments and those using adequate stimulation show that the longissimus dorsi and multifidus spinae are activated or facilitated by an ipsilateral stimulus to skin afferents. The extent of the effects induced by stimulation of skin differs in the types of preparation used. These differences may be accounted for by assuming a supraspinal control of the reflex pathways studied.     

Electromyographic study of trunk muscle activity during unresisted twisting posture in various twisting angles

The study investigated muscular activities of trunk twisting during standing posture by observing the surface EMG signal to estimate which muscles were active in graded isometric twisting posture. A laser pointer was used to maintain the twisting angle, while external resistances, which were commonly used to maintain the posture, were avoided. This procedure approached actual working posture in real life. Grading postural effort into various twisting angles provided reliable and natural posture measurements. Erector spinae (ES) at levels of L2 and L5 were selected to represent low back muscles, while external oblique (EO) and internal oblique (IO) were represented as abdominal muscles. Ten male subjects were asked to maintain a trunk twisting posture in various twisting angles for each right and left twisting direction. The EMG of the observed muscles were analyzed by power spectral density, and the activity changes were normalized by procedures of relative presentation and standardization. A Tukey multiple comparison was also applied to compare the activity changes among postural stages. Higher activity in contralateral ES, contralateral EO and ipsilateral IO is shown in the results, however, a significant increase against the initial posture (i.e., no-twisting posture) was found when the twisting angle was more than 30 degrees. Consistency with most past studies was shown in the result for the muscular activity of abdominal muscles, but not in the results for the low back muscles. The result emphasized the consideration that trunk muscle activity under a natural twisting condition was not always the same as that with the twisting under external resistance.     

The lateral reticular nucleus in the cat

The afferent paths from the spinal cord and from trigeminal afferents to the lateral reticular nucleus (LRN) were investigated by intracellular recording from 204 LRN neurones in preparations with a spinal cord lesion at C3 that spared only the ipsilateral ventral quadrant. Stimulation of nerves in the limbs evoked EPSPs and JPSPs in 201 of 204 tested LRN neurones. The strongest input was from the ipsilateral forelimb (iF) which evoked EPSPs in 49% and IPSPs in 73% of the LRN neurones. Each of the other limbs evoked EPSPs in approximately 20% and IPSPs in approximately 25% of the neurones. Stimulation of the ipsilateral trigeminal nerve (iTrig) evoked EPSPs in 32% and IPSPs in 46% of the neurones. The shortest latencies of the EPSPs and IPSPs indicated a disynaptic connection between primary afferents in the iF and iTrig and the LRN. The most direct pathways for excitatory and inhibitory responses from the other limbs were trisynaptic. Stimulation of the ventral part of the ipsilateral funiculus (iVLF) at C3 (C3iVLF) evoked monosynaptic responses in 189 of 201 tested LRN neurones. Monosynaptic EPSPs were recorded in 104 neurones and monosynaptic IPSPs in 126 neurones. Monosynaptic EPSPs and IPSPs were encountered in all parts of the LRN. Stimulation of the iVLF at L1 (L1iVLF) evoked monosynaptic EPSPs and IPSPs in the ventrolateral part of the LRN. The termination areas of excitatory and inhibitory fibres appeared to be the same. LRN neurones without monosynaptic EPSPs or IPSPs from the L1iVLF were located mainly in the dorsal part of the magnocellular division. Stimulation of the dorsal funiculi (DF) at C2 and the ipsilateral trigeminal nerve (iTrig) evoked excitatory and inhibitory responses in the LRN. The shortest latencies of EPSPs and IPSPs indicated disynaptic connections.