Modulation of hindlimbreflexes by tonic neck positions in cats

Summary In cats with the head fixed in a stereotactic frame a slight facilitation of the gastrocnemiussoleus (GS) monosynaptic reflexes in the ipsilateral hindlimb, accompanied by an inhibition of the monosynaptic reflex of the antagonistic deep peroneal nerve (DP), occurs following lateral flexion and rotation around the body axis at an angle of 25°. Following dorsiflexion of the body a moderate inhibition of the extensor reflexes (GS) up to 10% and a reciprocal slight excitation in the same range of the monosynaptic reflexes of flexor muscles (DP) is recorded. On the other hand after ventriflexion of the body there is a marked inhibition in the range of 50% of all the monosynaptic extensor and flexor reflexes for all the stimulus intensities used. This inhibition can already be demonstrated in ventriflexion of 10°–15°. All these patterns of reflex modulation are similar in cortically intact, ischemically decorticated and precollicularly decerebrated cats. Furthermore these patterns are not essentially influenced by cerebellectomy, although all reflex amplitudes are reduced. However, the spinal reflexes can not be modulated by these body movements after acute spinalization.Supported by Sonderforschungsbereich Hirnforschung (SFB 70) der Deutschen Forschungsgemeinschaft (DFG)     

Absence of nerve specificity in human cutaneous reflexes during standing

Cutaneous reflexes in lower limb muscles were recorded from healthy human subjects after non-noxious electrical stimulation of superficial peroneal (SP), sural and distal tibial nerves while subjects: (1) made graded voluntary contractions of the ankle and knee extensor and flexor muscles while mimicking late stance or heel strike limb positions; and (2) walked on a treadmill at speeds of 2 and 4 km/h. During standing, net reflexes were predominantly suppressive and graded with background EMG. In contrast, during walking net reflexes were mostly facilitatory and uncorrelated with background EMG. Opposite signs (negative during standing, positive during walking) and significant differences of the reflex ratio (net reflex/background EMG) were seen in most leg muscles. The nerve stimulated did not determine the sign of the net reflex while standing: nerve specificity was absent. We suggest that during standing, where maintenance of posture is of primary importance, there is a reduction of effort that led to increased cutaneous input (i.e., a global suppressive response), while during walking there is a modulation of reflexes which is independent of muscle activation level but closely tied to events occurring in the step cycle.     

Roll tilt reflexes after vestibulospinal tract lesions

Summary The effects of lesions of the vestibulospinal tracts on vestibular reflexes evoked by roll tilt in forelimb and neck extensors were examined in decerebrate cats. Sectioning the medial longitudinal fasciculus, which contains the medial vestibulospinal tract, had no major effect on the phase of the reflex, although some gain was usually lost at high stimulus frequencies. Spinal lesions at C2–C3, both cord hemisections and more restrictive lesions which cut the lateral vestibulospinal tract, produced two major effects on the forelimb. Background EMG activity was usually abolished in the triceps ipsilateral to the lesion, with partial loss of activity in the opposite limb. The tilt reflex response in the ipsilateral limb appeared normal, although it was usually necessary to raise the background excitability of the preparation by administering L-Dopa in order to observe the reflex. In contrast, the response in the contralateral limb showed a phase reversal of 180 deg at low stimulus frequencies, implying that the reflex in intact cats receives a crossed otolith-spinal input. Responses in the neck extensors splenius and biventer, recorded from compartments caudal to the spinal lesion, were relatively unaffected.Partially supported by NASA grant NSG-2380 and PHS grants NS02619 and RR07065Recipient of NIH Postdoctoral Fellowship NS06128Supported in part by NIH Research Service Award 7524     

The dynamic neck-eye reflex in mammals

Summary Stimulation of cervical proprioceptors by torsion of the neck results in movement of the eyes. The pathways of this neck-eye reflex have been identified electrophysiologically, and in individuals with vestibulo-ocular deficits the reflex is often seen to contribute to retinal image stability during head movements. In intact individuals, however, its role in ocular compensation for head movements is questionable. In this and other studies, the reflex eye movements were in the direction opposite the vestibulo-ocular reflex and were, therefore, anticompensatory. In four species of mammal (rat, rabbit, cat, and bush baby — a primate), the reflex was most consistently elicited with an anticompensatory phase; furthermore, when an animal partially stabilizes its head in space (by the vestibulo-collic reflex) during body rotation, the vestibulo-ocular and neck eye reflexes must have opposite polarities if their summation is to be of use to the animal. The neck-eye reflex appears to be absent when the animal actively moves its head; it only appears during the experimental procedure employed to elicit the reflex. An alternative function for the electrophysiologically identified pathway of the neck-eye reflex is suggested.Supported by USPHS Grant NS-13809 and University of Connecticut Research Foundation Grant 35-832     

Hand muscle reflexes following air puff stimulation

Hand muscle reflexes following muscle stretch and electrical nerve stimulation show a typical pattern consisting of short- and long-latency reflexes. The present investigation was designed to test reflexes following pure cutaneous stimulation. Air puffs were delivered to the palmar tip and the nail bed of the first, second and fifth fingers during isotonic contraction of hand muscles. The EMGs from the thenar muscles, the first dorsal interosseous muscle and the hypothenar muscles were recorded. Reflexes were obtained in all muscles, with a typical configuration consisting of a short-latency excitatory component (cutaneous longlatency reflex I, cLLR I) and a second excitatory component (cutaneous long-latency reflex II, cLLR II), with an inhibitory component between them. The size of cLLR II differed depending on the area stimulated and the muscle recorded. We found the largest responses always in the muscle acting on the stimulated finger. The reflex size depended on the strength of air puff stimulation. Allowing small displacements of the fingers led to an additional increase in the size of the reflex. The pattern of reflexes was identical independent of whether the finger tip or the nail bed was stimulated, but the size of the reflexes was smaller following nail bed stimulation. Following blockade of the cutaneous nerve branches of the thumb with local anaesthetics, air puff stimulation of the thumb no longer elicited this reflex pattern. Hence, under our experimental conditions, cutaneous receptors were the only source of afferent input for these reflexes.The results suggest that these cutaneous reflexes are mainly dedicated to controlling the stimulated finger independent of whether the palmar tip or the nail bed is stimulated. A possible physiological function is the adapting of grip force during handling of delicate objects if a perturbation is applied either to the object or the hand.     

Human interlimb reflexes evoked by electrical stimulation of cutaneous nerves innervating the hand and foot

There is some discrepancy over the extent to which reflex pathways from different cutaneous nerves in the hand and foot link the cervical and lumbar spinal cord in neurologically intact humans. The present experiments were designed to determine whether stimulation of a cutaneous nerve in the foot or in the hand evoked reflexes in the non-stimulated limbs (interlimb reflexes). Reflexes were elicited by stimulating (5x1-ms pulses at 300 Hz) the superficial peroneal (SP; innervates the foot dorsum) or superficial radial (SR; innervates the dorsolateral portion of the hand) nerve while subjects (n=10) performed focused contractions of different upper and lower limb muscles. Reflex responses were divided into early (<75 ms), middle (75-120 ms), and late (>120 ms) epochs as determined from averages of 50 sweeps of stimulus-locked electromyographic activity. Significant interlimb reflexes were found at the early latency in 44/106 and 44/103 muscles sampled after SP and SR nerve stimulation, respectively. At the middle latency, significant interlimb reflexes were seen in 89/106 and 87/103 muscles sampled after SP and SR nerve stimulation, respectively. Interlimb reflexes were seen when stimulating at the wrist (i.e. SR nerve) and when stimulating at the ankle (i.e. SP nerve) with an equal probability. The results show that interlimb cutaneous reflexes are widely distributed in humans. The mean latency of the earliest response was quite short and may be mediated by a propriospinal pathway. Functionally, these pathways may provide a substrate for transferring information to coordinate movements between the limb segments.     

How does action resist visual illusion? Uncorrected oculomotor information does not account for accurate pointing in peripersonal space

The amplitudes and signs of cutaneous reflexes are modulated during rhythmic movements of the arms and legs (during walking and arm or leg cycling for instance). This reflex modulation is frequently independent of the background muscle activity and may involve central pattern generator (CPG) circuits. The purpose of the present study was to investigate the nature and degree of coupling between the upper limbs during arm cycling, with regard to the regulation of cutaneous reflexes. Responses to electrical stimulations of the right, superficial radial nerve (five 1 ms pulses, 300 Hz) were recorded bilaterally in six arm muscles of eight participants during arm cycling involving only the limb ipsilateral to the stimulation, only the limb contralateral to the stimulation, and bilateral movement when the limbs were both in-phase and 180° out of phase. The pattern of cutaneous reflex modulation throughout the arm cycle was independent of the functional state of the limb contralateral to the recording site, irrespective of whether recordings were made ipsilateral or contralateral to the stimulation. Furthermore, cutaneous reflexes were significantly (p<0.05) modulated with arm position in only 8% of cases in which the limb containing the responding muscle was either stationary or being moved passively by the experimenter. The results show that there is relatively weak coupling between the arms with regard to the regulation of cutaneous reflexes during rhythmic, cyclical arm movements. This suggests a loose connection between the CPGs for each arm that regulate muscle activity and reflex amplitude during rhythmic movement.     

Comparison of dynamic properties of canal-evoked vestibulospinal reflexes of the neck and forelimb in the decerebrate cat

Summary The responses of neck and forelimb muscles to sinusoidal polarization of the horizontal canal nerve were compared by recording from these muscles simultaneously. Contrary to results on the vestibulocollic reflex, the central phase lag in the vestibulo-forelimb reflex increases with increasing frequencies up to 3 Hz. This demonstrates a difference in the organization of vestibular-driven pathways to neck and forelimb muscles.Supported in part by grant NS 02619 from the National Institutes of HealthRecipient of a fellowship from the China Medical Board. On leave from the Department of Physiology, University of Hong Kong     

Absence of linear correlation between fluctuations in area of simultaneous recorded monosynaptic responses and Hoffmann's reflexes in the rat

In this study we analyze the possible relationship between fluctuations in area of monosynaptic reflex responses (MSR) and Hoffmann's reflex (H reflexes) in the plantar closed loop pathway of the anesthetized rat. These reflexes were evoked by low-frequency stimuli applied to the sciatic nerve or lateral plantar nerve and then concurrently recorded on the distal tibial nerve or lateral plantar nerve, respectively as well as the lateral plantar muscles in the foot of the anesthetized rat. From trial to trial, H reflexes showed higher variability in area than MSR, whether the latter was recorded in the distal tibial nerve (n=8 experiments) or in the lateral plantar nerve (n=5 experiments). No linear correlation was found between changes in area of concurrently evoked MSR and H reflexes (r(MSR-H,n=8)=0.11+/-0.03 and r(MSR-H,n=5)=0.08+/-0.09, respectively). These findings suggest that trial-to-trial fluctuations in area of H reflexes may involve interaction of several sources of variation, among others to MSR variability (due to pre-, and post-synaptic factors influencing the excitability of spinal motoneurons) in combination with those related to peripheral mechanisms, such as trial to trial activation of a different number of muscle fibers, either by the probabilistic transmitter release from neuromuscular junctions, by activation of motor units of variable size or to fluctuations in excitability of muscle fibers.     

Analysis of the forelimb crossed extension reflex in thalamic cats during stepping.

Forelimb crossed extension reflexes were examined in 22 thalamic cats. These reflexes were elicited either by backward passive movement or by repetitive electrical stimulation of cutaneous and joint afferent nerves in the contralateral forelimb. Single stimulation of the superficial radial nerve evoked two types of reflex responses--early (ER) and late (LR)--from the triceps brachii muscle on the contralateral side. The latencies were about 7 and 16-25 ms, corresponding to the propriospinal (PSR) and spino-bulbo-spinal (SBS) reflexes of the ipsilateral flexor, respectively. Repetitive stimulation of the superficial radial nerve evoked the LR but not the ER. The crossed extension reflex and LR were abolished by lesions of the dorsolateral funiculus of the cervical cord on the side opposite to the recording. The tonic EMG activity, crossed extension reflex and LR in the extensor on the side of lesions were abolished by lesions of the ventrolateral funiculus of the cervical cord. During forelimb stepping, the amplitudes of both ER and LR fluctuated depending on the phase of the step cycle. The ER appeared during a narrow period in the early phase of the stance, whereas the LR was observed during a wide period from the middle of the swing to the middle of the stance. Both responses were absent from the middle of the stance to the middle of the swing. These observations suggest that forelimb crossed extension reflexes involve both spinal and supraspinal (SBS) loop mechanisms, and that these are utilized during stepping, with the latter mechanism in particular playing an important part in the extension phase of the forelimb forward movement.     

Relationship of cat vestibular neurons to otolith-spinal reflexes

Summary The dynamics of neurons in the vestibular nuclei of canal-plugged, decerebrate cats were studied in response to lateral (roll) tilt. Forelimb and neck extensor reflexes recorded simultaneously develop a progressive phase lag above 0.1 Hz. Neurons which exhibited a muscle-like phase lag were excited during low frequency stimuli by ipsilateral side-up tilt (beta response). Neurons with alpha responses, excited during side-down tilt, exhibited a constant phase, without a high frequency lag. Vestibulospinal neurons were present in both of these response groups, as were units driven at monosynaptic latencies by electrical stimulation of the ipsilateral labyrinth. The phase-lagging beta responses are appropriate for contributing to the reflexes observed in the ipsilateral neck and contralateral forelimb.Partially supported by NASA grant NSG-2380 and NIH grant NS02619NIH Postdoctoral Fellowship PHS NS06128     

Modulation of cutaneous reflexes by load receptor input during human walking

To investigate the influence of load on the modulation of cutaneous reflexes, evoked by sural nerve stimulation, electromyographic activity in different leg muscles (tibialis anterior, gastrocnemius medialis (GM), biceps femoris, and soleus muscles (SO)) was recorded in healthy humans during treadmill walking with different body loads. Sural nerve stimulation was applied at two times perception threshold during different phases of the step cycle. Reflex amplitudes increased with body unloading and decreased with body loading. The reflex responses were not a simple function of the level of background activity. For example, in GM and SO, the largest reflex responses occurred during walking with body unloading, when background activity was decreased. Hence, stable ground conditions (body loading) yielded smaller reflexes. It is proposed that load receptors are involved in the regulation of cutaneous reflex responses in order to adapt the locomotor pattern to the environmental conditions.     

Conditioned and unconditioned forelimb reflex systems in the cat: involvement of the intermediate cerebellum

 Temporary inactivation of the cerebellar interposed nuclei was used to assess the role of the intermediate cerebellum in the performance of forelimb cutaneo-muscular reflexes in the cat. The following types of reflexive responses were evaluated: the classically conditioned and unconditioned forelimb withdrawal responses and the forelimb tactile placing, hopping and magnet responses. The experiments tested the hypothesis that the intermediate cerebellum is involved in the performance of all the above forelimb reflexes. The forelimb withdrawal reflex was classically conditioned in a newly developed paradigm in which animals were first operantly conditioned to stand on four elevated platforms. Trained animals were microinjected with a γ-aminobutyric acid (GABA) agonist, muscimol, in the interposed nuclei, and the effects of inactivation of the intermediate cerebellar output on the forelimb reflexes were examined. The main findings of these experiments are that unilateral muscimol inactivation of the interposed nuclei in the cat abolished the expression of the classically conditioned limb flexion reflex, suppressed the performance of the unconditioned withdrawal reflex and, in parallel, downregulated the tactile placing, hopping and magnet postural responses in the ipsilateral forelimb. These observations are inconsistent with concepts indicating exclusive involvement of the intermediate cerebellum in the classically conditioned reflexes elicited by aversive stimuli. On the contrary, they support the hypothesis of a more global involvement of this structure in learned and unlearned defensive flexion reflexes and in automatic postural response systems. Received: 29 July 1996 / Accepted: 26 September 1996     

Interaction of tonic neck and vestibular reflexes in the forelimb of the decerebrate cat

Summary Tonic neck reflexes, studied with EMG recording, have similar dynamics in forelimb extensor muscles of acutely labyrinthectomized cats, and in cats with intact labyrinths. The reflex occurs more frequently in the latter and its gain is higher. In intact preparations we evoked vestibular and tonic neck reflexes separately or in combination, at frequencies of 0.05–0.5 Hz. As expected from earlier work, the two reflexes oppose each other and frequently cancel; the two reflexes add linearly.Supported in part by N.I.H. grant NS 02619 and NASA grant NSG 2380     

Spatial coordination by descending vestibular signals

Electromyographic activity of dorsal neck muscles and neck torques was recorded to study vestibulocollic, cervicocollic, and combined reflexes in alert and decerebrate cats during rotations of the whole body, the body except for the head, and the head but not the rest of the body. Cats were rotated about many axes that lay in the frontal, sagittal, and horizontal planes using sinusoidal 0.25-Hz waveforms or sum-of-sinusoid waveforms. Robust electromyographic responses were recorded from six muscles, with response directionality that in most cases did not show strong dependence on the reflex tested or on other factors including exact neck angle, stimulus amplitude from 5° to 60°, and intact versus decerebrate state. Based on the strength of responses to rotations about all the tested axes, neck muscles could be characterized by maximal activation direction vectors representing the axis and direction of rotation in threedimensional space that was most excitatory during reflex responses. Responses to rotations about axes that lay in a coordinate plane were predicted by a cosine function of the angle between the axis under test and the maximally excitatory axis in the plane. All muscles were excited by the nose down phase of pitch rotation and by yaw and roll away from the side on which the muscle lay. Biventer cervicis was best activated by rotations with axes near nose-down pitch, and its axis of maximal activation also had small, approximately equal components of yaw and roll toward the contralateral side. Complexus was best excited by rotations with axes nearest roll, but with large components along all three axes. Occipitoscapularis was best excited by rotations about axes near pitch, but with a moderately large contralateral yaw component and a smaller but significant contralateral roll component. Splenius was best excited by rotations with a large component of contralateral yaw, considerable nose-down pitch, and a smaller component of contralateral roll. Rectus major was best excited by rotations near nose-down pitch, but with a substantial contralateral yaw component and smaller contralateral roll component. Obliquus inferior was best excited by rotations with a large component of contralateral yaw, but with considerable contralateral roll and nose-down pitch components. All muscles responded as though they received convergent input from all three semicircular canals. Vestibulocollic and combined reflex responses in alert cats and vestibulocollic, cervicocollic, and combined responses in decerebrate cats appeared to have the same directionality, as evidenced by insignificant shifts in maximal activation vectors. Cervicocollic responses in alert cats were inconsistent and often absent, but appeared upon decerebration, suggesting that higher centers suppress the cervicocollic reflex in intact animals. Decerebration and partial cerebellectomy had no significant effect on maximal activation directions, although electromyographic response magnitudes increased after each. The results suggest that common circuits or strategies are used by neck stretch and vestibular-neck reflexes. The reflex excitation directions do not match the mechanical actions of the neck muscles but agree fairly well with previously published predictions of a mathematical model of neck motor control.     

Sensitivity of monosynaptic test reflexes to facilitation and inhibition as a function of the test reflex size: a study in man and the cat

Summary In parallel experiments on humans and in the cat it was investigated how the sensitivity of monosynaptic test reflexes to facilitation and inhibition varies as a function of the size of the control test reflex itself. In man the monosynaptic reflex (the Hoffmann reflex) was evoked in either the soleus muscle (by stimulation of the tibial nerve) or the quadriceps muscle (by stimulation of the femoral nerve). In the decerebrate cat monosynaptic reflexes were recorded from the nerves to soleus and medial gastrocnemius muscles; they were evoked by stimulation of the proximal ends of the sectioned L7 and S1 dorsal roots. Various excitatory and inhibitory spinal reflex pathways were used for conditioning the test reflexes (e.g. monosynaptic Ia excitation, disynaptic reciprocal inhibition, cutaneous inhibition, recurrent inhibition, presynaptic inhibition of the Ia fibres mediating the test reflex). It was shown that the additional number of motoneurones recruited in a monosynaptic test reflex by a constant excitatory conditioning stimulus was very much dependent on the size of the test reflex itself. This dependency had the same characteristic pattern whatever the conditioning stimulus. With increasing size of the test reflex the number of additionally recruited motoneurones first increased, then reached a peak (or plateau) and finally decreased. A similar relation was also seen with inhibitory conditioning stimuli. The basic physiological factors responsible for these findings are discussed. Finally, the implications for the interpretation of experiments in man with the H-reflex technique are considered.     

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.     

Bisphenol A attenuates phenylbiguanide-induced cardio-respiratory reflexes in anaesthetized rats

Bisphenol A (BPA), a toxic chemical released from plastics, produces respiratory arrest and hypotension after a latency. The latency was similar to the reflex apnoea induced by the vagal C fibre stimulation. Therefore, the present study was undertaken to examine the effects of chronic and acute exposure to BPA on cardio-respiratory reflexes elicited by phenylbiguanide (PBG). Acute and chronic experiments were performed on adult female rats. In chronic experiments, the animals were ingested with pellets containing BPA (2 μg/kg body weight) or without BPA (time-matched control) for 30 days. Subsequently, the animals were anaesthetized and prepared for recording blood pressure, ECG and respiratory excursions. PBG was injected through jugular vein to evoke reflexes in these animals. In acute experiments, the PBG reflexes were obtained before and after injecting BPA/ethanol. Also vagal afferent activity was recorded in some rats. In time-matched control rats, PBG produced bradycardia, hypotension and tachypnoea over a period of time. The maximal changes were around 50–65%. In BPA treated group, the PBG-induced heart rate and respiratory frequency changes were attenuated significantly. Acute exposure of animals to BPA (35 mg/kg body weight) for 30 min also attenuated the PBG-induced responses significantly. The attenuation of the PBG reflex responses by BPA in acute experiments was associated with decreased vagal afferent activity. The present results indicate that BPA attenuates the protective cardio-respiratory reflexes due to decreased vagal afferent activity.     

Facilitation of soleus H-reflex amplitude evoked by cutaneous nerve stimulation at the wrist is not suppressed by rhythmic arm movement

Neural connections between the cervical and lumbosacral spinal cord may assist in arm and leg coordination during locomotion. Currently the extent to which arm activity can modulate reflex excitability of leg muscles is not fully understood. We showed recently that rhythmic arm movement significantly suppresses soleus H-reflex amplitude probably via modification of presynaptic inhibition of the IA afferent pathway. Further, during walking reflexes evoked in leg muscles by stimulation of a cutaneous nerve at the wrist (superficial radial nerve; SR) are phase and task dependent. However, during walking both the arms and legs are rhythmically active thus it is difficult to identify the locus of such modulation. Here we examined the influence of SR nerve stimulation on transmission through the soleus H-reflex pathway in the leg during static contractions and during rhythmic arm movements. Nerve stimulation was delivered with the right shoulder in flexion or extension. H-reflexes were evoked alone (unconditioned) or with cutaneous conditioning via stimulation of the SR nerve (also delivered alone without H-reflex in separate trials). SR nerve stimulation significantly facilitated H-reflex amplitude during static contractions with the arm extended and countered the suppression of reflex amplitude induced by arm cycling. The results demonstrate that cutaneous feedback from the hand on to the soleus H-reflex pathway in the legs is not suppressed during rhythmic arm movement. This contrasts with the observation that rhythmic arm movement suppresses facilitation of soleus H-reflex when cutaneous nerves innervating the leg are stimulated. In conjunction with other data taken during walking, this suggests that the modulation of transmission through pathways from the SR nerve to the lumbosacral spinal cord is partly determined by rhythmic activity of both the arms and legs.     

Functional organization of vestibular and visual inputs to neck and forelimb motoneurons in the frog.

1. Intracellular responses in neck and forelimb motoneurons to electrical stimulation of the vestibular nerve, the optic tectum, and the optic nerve were studied in frog. 2. Stimulation of the anterior branch of the vestibular nerve typically produced EPSPs, bilaterally, in neck, shoulder (DOR), and forelimb extensor (TRI, RAD) motoneurons, and bilateral IPSPs in forelimb adductor (PED) and flexor (ULN, COR) motoneurons. 3. Latencies of PSPs recorded in neck, shoulder, and proximal extensor motoneurons (TRI) were mostly in the disynaptic range, whereas many of those recorded in distal extensor (RAD) and in adductor and flexor motoneurons involved three synapses. 4. Lesion of the vestibulospinal fibers greatly reduced the vestibular nerve-evoked field potentials in the spinal cord and the occurrence of PSPs in forelimb motoneurons. These results as well as the latency measurements suggest that the pathway linking vestibular nerve and forelimb motoneurons mainly consists of vestibulospinal fibers, though involvement of other structures for production of later PSPs could not be completely ruled out. Hemisection of the brain stem at its most caudal level showed that the pathway to the contralateral motoneurons crosses at the level of brain stem as well as in the spinal cord. 5. Stimulation of the optic tectum produced EPSPs, IPSPs, and a mixture of EPSPs and IPSPs in neck, shoulder, and forelimb motoneurons, bilaterally. Most frequently, a combination of an excitation and inhibition was observed. The pathway from the optic tectum to neck and limb motoneurons is at least dysnaptic in nature. 6. Stimulation of the optic nerve produced IPSPs and a mixture of EPSPs and IPSPs in neck and forelimb motoneurons. Impulses originating from the optic nerve descend as far as to lumbar motoneurons producing EPSP-IPSP sequences bilaterally. 7. Interaction studies suggested that the vestibular and optic pathways to neck and forelimb motoneurons are separate from each other so that the site of integration of vestibular and visual input occurs at the level of motoneurons. 8. Evidence for electronic coupling among forelimb motoneurons and electrical synaptic transmission in th pathway linking vestibular nerve and forelimb motoneurons is presented.