Cutaneous Inputs Research Articles

All-Source Basal Vitamin D Inputs Are Greater Than Previously Thought and Cutaneous Inputs Are Smaller

AbstractThe magnitude of vitamin D inputs in individuals not taking supplements is unknown; however, there is a great deal of information on quantitative response to varying supplement doses. We reanalyzed individual 25-hydroxyvitamin D [25(OH)D] concentration data from 8 studies involving cholecalciferol supplementation (total sample size = 3000). We extrapolated individual study dose-response curves to zero concentration values for serum 25(OH)D by using both linear and curvilinear approaches and measured seasonal oscillation in the serum 25(OH)D concentration. The total basal input (food plus solar) was calculated to range from a low of 778 IU/d in patients with end-stage renal disease to a high of 2667 IU/d in healthy Caucasian adults. Consistent with expectations, obese individuals had lower baseline, unsupplemented 25(OH)D concentrations and a smaller response to supplements. Similarly, African Americans had both lower baseline concentrations and lower calculated basal, all-source inputs. Seasonal oscillation in 4 studies ranged from 5.20 to 11.4 nmol/L, reflecting a mean cutaneous synthesis of cholecalciferol ranging from 209 to 651 IU/d at the summer peak. We conclude that: 1) all-source, basal vitamin D inputs are approximately an order of magnitude higher than can be explained by traditional food sources; 2) cutaneous, solar input in these cohorts accounts for only 10–25% of unsupplemented input at the summer peak; and 3) the remainder must come from undocumented food sources, possibly in part as preformed 25(OH)D.

Posterior hypothalamic modulation of light-evoked trigeminal neural activity and lacrimation

Enhanced light sensitivity is a common feature of many neuro-ophthalmic conditions and some chronic headaches. Previously we reported that the bright light-evoked increases in trigeminal brainstem neural activity and lacrimation depended on a neurovascular link within the eye (Okamoto et al., 2012). However, the supraspinal pathways necessary for these light-evoked responses are not well defined. To assess the contribution of the posterior hypothalamic area (PH), a brain region closely associated with control of autonomic outflow, we injected bicuculline methiodide (BMI), a GABAa receptor antagonist, into the PH and determined its effect on the encoding properties of ocular neurons at the ventrolateral trigeminal interpolaris/caudalis transition (Vi/Vc) and caudalis/upper cervical cord junction (Vc/C1) regions and on reflex lacrimation in male rats under isoflurane anesthesia. BMI markedly reduced light-evoked (>80%) responses of Vi/Vc and Vc/C1 neurons at 10min with partial recovery by 50min after injection. BMI also reduced (>35%) the convergent cutaneous receptive field area of Vi/Vc and Vc/C1 ocular neurons indicating that both intra-ocular and periorbital cutaneous inputs were affected by changes in PH outflow. Light-evoked lacrimation was reduced by >35% at 10min after BMI, while resting mean arterial pressure increased promptly and remained elevated (>20mmHg) throughout the 50-min post-injection period. These results suggested that PH stimulation, acting in part through increased sympathetic activity, significantly inhibited light- and facial skin-evoked activity of ocular neurons at the Vi/Vc and Vc/C1 region. These data provide further support for the hypothesis that autonomic outflow plays a critical role in mediating light-evoked trigeminal brainstem neural activity and reflex lacrimation.

Circuits for Grasping: Spinal dI3 Interneurons Mediate Cutaneous Control of Motor Behavior

Accurate motor performance depends on the integration in spinal microcircuits of sensory feedback information. Hand grasp is a skilled motor behavior known to require cutaneous sensory feedback, but spinal microcircuits that process and relay this feedback to the motor system have not been defined. We sought to define classes of spinal interneurons involved in the cutaneous control of hand grasp in mice and to show that dI3 interneurons, a class of dorsal spinal interneurons marked by the expression of Isl1, convey input from low threshold cutaneous afferents to motoneurons. Mice in which the output of dI3 interneurons has been inactivated exhibit deficits in motor tasks that rely on cutaneous afferent input. Most strikingly, the ability to maintain grip strength in response to increasing load is lost following genetic silencing of dI3 interneuron output. Thus, spinal microcircuits that integrate cutaneous feedback crucial for paw grip rely on the intermediary role of dI3 interneurons.

Amplification of interlimb reflexes evoked by stimulating the hand simultaneously with conditioning from the foot during locomotion

BackgroundWidespread interlimb reflexes evoked in leg muscles by cutaneous stimulation of the hand are phase-modulated and behaviorally relevant to produce functional changes in ankle trajectory during walking. These reflexes are complementary to the segmental responses evoked by stimulation at the ankle. Despite differences in the expression of reflex amplitude based upon site of nerve stimulation, there are some common features as well, suggesting the possibility of shared interneuronal pathways. Currently little is known about integration or shared reflex systems from interlimb cutaneous networks during human locomotion. Here we investigated convergent reflex effects following cutaneous stimulation of the hand and foot during arm and leg cycling (AL) by using spatial facilitation. Participants performed AL cycling and static activation of the target muscle knee extensor vastus lateralis (VL) in 3 different randomly ordered nerve stimulation conditions: 1) superficial radial nerve (SR; input from hand); 2) superficial peroneal nerve (SP; input from foot); and, 3) combined stimulation (SR + SP). Stimuli were applied around the onset of rhythmic EMG bursts in VL corresponding to the onset of the power or leg extension phase.ResultsDuring AL cycling, small inhibitory (~80 ms) and large facilitatory reflexes (~100 ~ 150 ms) were seen in VL. The amplitudes of the facilitatory responses with SR + SP stimulation were significantly larger than those for SP or SR stimulation alone. The facilitation was also significantly larger than the simple mathematical summation of amplitudes from SP and SR trials. This indicates extra facilitation beyond what would be accounted for by serial neuronal processing and was not observed during static activation.ConclusionsWe conclude that AL cycling activates shared interneurons in convergent reflex pathways from cutaneous inputs innervating the hand and leg. This enhanced activity has functional implications for corrective responses during locomotion and for translation to rehabilitation after neurotrauma.

Activity‐dependent development of tactile and nociceptive spinal cord circuits

Developing brain circuits are shaped by postnatal sensory experience, but little is known about this process at the level of the spinal cord. Here we review the mechanisms by which cutaneous sensory input drives the maturation of spinal sensory circuits. Newborn animals are highly sensitive to tactile input and dorsal horn circuits are dominated by low threshold A fiber inputs. We show that this arises from the absence of the functional, targeted glycinergic inhibition of tactile activity that emerges only in the second week of life. Selective block of afferent C fibers in postnatal week 2 delays the maturation of glycinergic inhibition and maintains dorsal horn circuits in a neonatal state. We propose that in the newborn strong tactile A fiber input facilitates activity-dependent synaptic strengthening in the dorsal horn, but that this ends with the arrival of nociceptive C fiber spinal input that drives the maturation of targeted glycinergic inhibition.

Corticokinematic coherence during active and passive finger movements

Corticokinematic coherence (CKC) refers to coupling between magnetoencephalographic (MEG) brain activity and hand kinematics. For voluntary hand movements, CKC originates mainly from the primary sensorimotor (SM1) cortex.To learn about the relative motor and sensory contributions to CKC, we recorded CKC from 15 healthy subjects during active and passive right index-finger movements. The fingertip was either touching or not touching table, resulting in active-touch, active-no-touch, passive-touch, and passive-no-touch conditions. The kinematics of the index-finger was measured with a 3-axis accelerometer. Beamformer analysis was used to locate brain activations for the movements; somatosensory-evoked fields (SEFs) elicited by pneumatic tactile stimulation of the index finger served as a functional landmark for cutaneous input.All active and passive movements resulted in statistically significant CKC at the movement frequency (F0) and its first harmonic (F1). The main CKC sources at F0 and F1 were in the contralateral SM1 cortex with no spatial differences between conditions, and distinct from the SEF sources. At F1, the coherence was by two thirds stronger for passive than active movements, with no difference between touch vs. no-touch conditions.Our results suggest that the CKC occurring during repetitive finger movements is mainly driven by somatosensory, primarily proprioceptive, afferent input to the SM1 cortex, with negligible effect of cutaneous input.

Change in EMG with skin friction at different frequencies during elbow flexion

Modulation of muscle activation in superficial and deeper regions may be induced by tactile stimulation. The purpose of this study was to examine changes in muscle activation with skin friction. Subjects performed an isometric elbow flexion at 30% maximal voluntary cotraction (MVC) with skin friction at different frequencies (0.5–2.7 Hz). Surface electromyography (S-EMG) and intramuscular EMG were obtained from the elbow flexor muscles (BBS: short head of biceps brachii, BBL: long head of biceps brachii, BRA: brachialis). S-EMG activity decreased at a higher frequency of 2.7 Hz and increased linearly with an increase in skin friction frequency (0.5–2.7 Hz) in BBS. A decrease in high-threshold motor unit (HT-MU) firing rate in superficial regions and an increase in low-threshold motor unit (LT-MU) firing rate in deeper regions were observed with skin friction (2.7 Hz) in BBS. The actions of inhibitory interneurons may be influenced by cutaneous afferent input with skin friction. Muscle activation of BBS depended on the intensity of the stimulus. Skin friction over BBS results in an inhibitory response in superficial regions of BBS, most likely due to the increase in firing rate of low-threshold cutaneous mechanoreceptors.

When Standing on a Moving Support, Cutaneous Inputs Provide Sufficient Information to Plan the Anticipatory Postural Adjustments for Gait Initiation

Gait initiation is preceded by initial postural adjustments whose goal is to set up the condition required for the execution of the focal stepping movement. For instance, the step is preceded by a shift of the body’s center of mass towards the stance foot unloading the stepping leg. This displacement is produced by exerting forces on the ground (i.e., thrust) while the body is still motionless. The purpose of this study was to identify whether the mere cutaneous inputs from the feet soles evoked by a lateral translation of the support could be used to scale the initial postural adjustments. Participants stood with their eyes closed on a force platform that could be moved laterally with a low acceleration (between 0.14 m/s2 and 0.30 m/s2) to reach a constant velocity of 0.02 m/s. This translation resulted in a change in the somatosensory cues from the feet soles without modifying vestibular inputs. Participants were instructed to produce a step with the right foot as soon as they felt the platform start to move (on either side) or heard an auditory cue. In the latter case, the platform stayed stationary. We found that the thrust duration was lengthened when the platform moved towards the supporting foot. In this condition, the cutaneous stimulation provided information related to a body shift towards the stepping leg. This increased thrust duration likely helped overcoming the non-functional body shift perceived towards the stepping leg. This result highlights the accuracy with which the actual standing position can be determined from foot sole cutaneous cues in the absence of visual and vestibular or proprioceptive inputs.

Cutaneous and periodontal inputs to the cerebellum of the naked mole-rat (Heterocephalus glaber)

The naked mole-rat (Heterocephalus glaber) is a small fossorial rodent with specialized dentition that is reflected by the large cortical area dedicated to representation of the prominent incisors. Due to naked mole-rats’ behavioral reliance on the incisors for digging and for manipulating objects, as well as their ability to move the lower incisors independently, we hypothesized that expanded somatosensory representations of the incisors would be present within the cerebellum in order to accommodate a greater degree of proprioceptive, cutaneous, and periodontal input. Multiunit electrophysiological recordings targeting the ansiform lobule were used to investigate tactile inputs from receptive fields on the entire body with a focus on the incisors. Similar to other rodents, a fractured somatotopy appeared to be present with discrete representations of the same receptive fields repeated within each folium of the cerebellum. These findings confirm the presence of somatosensory inputs to a large area of the naked mole-rat cerebellum with particularly extensive representations of the lower incisors and mystacial vibrissae. We speculate that these extensive inputs facilitate processing of tactile cues as part of a sensorimotor integration network that optimizes how sensory stimuli are acquired through active exploration and in turn adjusts motor outputs (such as independent movement of the lower incisors). These results highlight the diverse sensory specializations and corresponding brain organizational schemes that have evolved in different mammals to facilitate exploration of and interaction with their environment.

Subthreshold Vestibular Reflex Effects in Seated Humans Can Contribute to Soleus Activation When Combined With Cutaneous Inputs

The integration of vestibular and somatosensory information for the control of lower limb musculature remains elusive. To determine whether a subthreshold vestibular input influences the cutaneous evoked response, the isometric EMG activity in the posturally inactive soleus muscles of 13 healthy, seated subjects was collected. Vestibular afferents were activated using galvanic vestibular stimulation (GVS; 1.8-2.5mA, 500ms), while percutaneous electrical stimulation was delivered to the distal tibial nerve (11ms train of 3 × 1.0 ms pulses, 200Hz) to activate foot sole skin afferents. GVS elicited responses in soleus both independently and when combined with cutaneous stimulation. The responses to the combined sensory input showed an interaction between the two sensory modalities to influence muscle activation. Of note is the presence of significant muscle modulation in the combined condition, where subthreshold vestibular inputs altered the outcome of the cutaneous reflex response. This finding has implications for individuals with sensory deficiency. In the case of an absent or deficient sensory modality, balance protective reflexes to maintain postural equilibrium may be enhanced with targeted sensory augmentation.

Ankle joint movements are encoded by both cutaneous and muscle afferents in humans

We analyzed the cutaneous encoding of two-dimensional movements by investigating the coding of movement velocity for differently oriented straight-line movements and the coding of complex trajectories describing cursive letters. The cutaneous feedback was then compared with that of the underlying muscle afferents previously recorded during the same "writing-like" movements. The unitary activity of 43 type II cutaneous afferents was recorded in the common peroneal nerve in healthy subjects during imposed ankle movements. These movements consisted first of ramp-and-hold movements imposed at two different and close velocities in seven directions and secondly of "writing-like" movements. In both cases, the responses were analyzed using the neuronal population vector model. The results show that movement velocity encoding depended on the direction of the ongoing movement. Discriminating between two velocities therefore involved processing the activity of afferent populations located in the various skin areas surrounding the moving joint, as shown by the statistically significant difference observed in the amplitude of the sum vectors. Secondly, "writing-like" movements induced cutaneous neuronal patterns of activity, which were reproducible and specific to each trajectory. Lastly, the "cutaneous neuronal trajectories," built by adding the sum vectors tip-to-tail, nearly matched both the movement trajectories and the "muscle neuronal trajectories," built from previously recorded muscle afferents. It was concluded that type II cutaneous and the underlying muscle afferents show similar encoding properties of two-dimensional movement parameters. This similarity is discussed in relation to a central gating process that would for instance increase the gain of cutaneous inputs when muscle information is altered by the fusimotor drive.

Capture of kinesthesis by a competing cutaneous input

In four experiments, blindfolded participants were presented with pairs of stimuli simultaneously, one to each index finger. Participants moved one index finger, which was presented with cutaneous and/or kinesthetic stimuli, and this movement caused a raised line to move underneath the other, stationary index finger in a yoked manner. The stimuli were 180º rotations of each other (e.g., < and >), and thus when a < was traced with the moving finger, it caused a > to be felt at the stationary finger. When asked to report the experience, participants predominantly reported the cutaneous stimulus, seemingly being ignorant of the kinesthetic stimulus. This appears to be an intrahaptic capture phenomenon, which is of interest because it suggests that conflict between intrahaptic sensory stimuli can go unnoticed; sometimes we are unaware of how we moved, and sometimes we do not know what we touched. The results are interpreted in light of optimal integration, perceptual suppression, reafference suppression, and inattentional blindness.

Cutaneous inputs from the back abolish locomotor-like activity and reduce spastic-like activity in the adult cat following complete spinal cord injury

Spasticity is a condition that can include increased muscle tone, clonus, spasms, and hyperreflexia. In this study, we report the effect of manually stimulating the dorsal lumbosacral skin on spontaneous locomotor-like activity and on a variety of reflex responses in 5 decerebrate chronic spinal cats treated with clonidine. Cats were spinalized 1month before the terminal experiment. Stretch reflexes were evoked by stretching the left triceps surae muscles. Crossed reflexes were elicited by electrically stimulating the right tibial or superficial peroneal nerves. Wind-up of reflex responses was evoked by electrically stimulating the left tibial or superficial peroneal nerves. We found that pinching the skin of the back abolished spontaneous locomotor-like activity. We also found that back pinch abolished the rhythmic activity observed during reflex testing without eliminating the reflex responses. Some of the rhythmic episodes of activity observed during reflex testing were consistent with clonus with an oscillation frequency greater than 3Hz. Pinching the skin of the back effectively abolished rhythmic activity occurring spontaneously or evoked during reflex testing, irrespective of oscillation frequency. The results are consistent with the hypothesis that locomotion and clonus are produced by common central pattern-generators. Stimulating the skin of the back could prove helpful in managing undesired rhythmic activity in spinal cord-injured humans.

The upright posture improves plantar stepping and alters responses to serotonergic drugs in spinal rats

Recent studies on the restoration of locomotion after spinal cord injury have employed robotic means of positioning rats above a treadmill such that the animals are held in an upright posture and engage in bipedal locomotor activity. However, the impact of the upright posture alone, which alters hindlimb loading, an important variable in locomotor control, has not been examined. Here we compared the locomotor capabilities of chronic spinal rats when placed in the horizontal and upright postures. Hindlimb locomotor movements induced by exteroceptive stimulation (tail pinching) were monitored with video and EMG recordings. We found that the upright posture alone significantly improved plantar stepping. Locomotor trials using anaesthesia of the paws and air stepping demonstrated that the cutaneous receptors of the paws are responsible for the improved plantar stepping observed when the animals are placed in the upright posture.We also tested the effectiveness of serotonergic drugs that facilitate locomotor activity in spinal rats in both the horizontal and upright postures. Quipazine and (±)-8-hydroxy-2-(dipropylamino)tetralin hydrobromide (8-OH-DPAT) improved locomotion in the horizontal posture but in the upright posture either interfered with or had no effect on plantar walking. Combined treatment with quipazine and 8-OH-DPAT at lower doses dramatically improved locomotor activity in both postures and mitigated the need to activate the locomotor CPG with exteroceptive stimulation. Our results suggest that afferent input from the paw facilitates the spinal CPG for locomotion. These potent effects of afferent input from the paw should be taken into account when interpreting the results obtained with rats in an upright posture and when designing interventions for restoration of locomotion after spinal cord injury.

Are all spinal segments equal: intrinsic membrane properties of superficial dorsal horn neurons in the developing and mature mouse spinal cord

Neurons in the superficial dorsal horn (SDH; laminae I-II) of the spinal cord process nociceptive information from skin, muscle, joints and viscera. Most of what we know about the intrinsic properties of SDH neurons comes from studies in lumbar segments of the cord even though clinical evidence suggests nociceptive signals from viscera and head and neck tissues are processed differently. This ‘lumbar-centric' view of spinal pain processing mechanisms also applies to developing SDH neurons. Here we ask whether the intrinsic membrane properties of SDH neurons differ across spinal cord segments in both the developing and mature spinal cord. Whole cell recordings were made from SDH neurons in slices of upper cervical (C2-4), thoracic (T8-10) and lumbar (L3-5) segments in neonatal (P0-5) and adult (P24-45) mice. Neuronal input resistance (R(IN)), resting membrane potential, AP amplitude, half-width and AHP amplitude were similar across spinal cord regions in both neonates and adults (∼100 neurons for each region and age). In contrast, these intrinsic membrane properties differed dramatically between neonates and adults. Five types of AP discharge were observed during depolarizing current injection. In neonates, single spiking dominated (∼40%) and the proportions of each discharge category did not differ across spinal regions. In adults, initial bursting dominated in each spinal region, but was significantly more prevalent in rostral segments (49% of neurons in C2-4 vs. 29% in L3-5). During development the dominant AP discharge pattern changed from single spiking to initial bursting. The rapid A-type potassium current (I(Ar)) dominated in neonates and adults, but its prevalence decreased (∼80% vs. ∼50% of neurons) in all regions during development. I(Ar) steady state inactivation and activation also changed in upper cervical and lumbar regions during development. Together, our data show the intrinsic properties of SDH neurons are generally conserved in the three spinal cord regions examined in both neonate and adult mice. We propose the conserved intrinsic membrane properties of SDH neurons along the length of the spinal cord cannot explain the marked differences in pain experienced in the limbs, viscera, and head and neck.

Changes in correlation between spontaneous activity of dorsal horn neurones lead to differential recruitment of inhibitory pathways in the cat spinal cord

Simultaneous recordings of cord dorsum potentials along the lumbo-sacral spinal cord of the anaesthetized cat revealed the occurrence of spontaneous synchronous negative (n) and negative-positive (np) cord dorsum potentials (CDPs). The npCDPs, unlike the nCDPs, appeared preferentially associated with spontaneous negative dorsal root potentials (DRPs) resulting from primary afferent depolarization. Spontaneous npCDPs recorded in preparations with intact neuroaxis or after spinalization often showed a higher correlation than the nCDPs recorded from the same pair of segments. The acute section of the sural and superficial peroneal nerves further increased the correlation between paired sets of npCDPs and reduced the correlation between the nCDPs recorded from the same pair of segments. It is concluded that the spontaneous nCDPs and npCDPs are produced by the activation of interconnected sets of dorsal horn neurones located in Rexed's laminae III–IV and bilaterally distributed along the lumbo-sacral spinal cord. Under conditions of low synchronization in the activity of this network of neurones there would be a preferential activation of the intermediate nucleus interneurones mediating Ib non-reciprocal postsynaptic inhibition. Increased synchronization in the spontaneous activity of this ensemble of dorsal horn neurones would recruit the interneurones mediating primary afferent depolarization and presynaptic inhibition and, at the same time, reduce the activation of pathways mediating Ib postsynaptic inhibition. Central control of the synchronization in the spontaneous activity of dorsal horn neurones and its modulation by cutaneous inputs is envisaged as an effective mechanism for the selection of alternative inhibitory pathways during the execution of specific motor or sensory tasks.

Perceptual constancy of texture roughness in the tactile system.

Our tactual perception of roughness is independent of the manner in which we touch the surface. A brick surface feels rough no matter how slowly or how rapidly we move our fingers, despite the fluctuating sensory inputs that are transmitted to the finger. Current theories of roughness perception rely solely on inputs from the cutaneous afferents, which are highly affected by scan velocity and force. The question then is: how is roughness constancy achieved? To this end, we characterized the subject's perceived roughness in six scanning conditions. These included two modes of touch: direct touch, where the finger is in contact with the surface, and indirect touch, where the surface is scanned with a hand-held probe; and three scanning modes: active (moving the hand across a stationary surface), passive (moving the surface across a stationary hand), and pseudo-passive (subject's hand is moved by the experimenter across a stationary surface). Here, we show that roughness constancy is preserved during active but not passive scanning, indicating that the hand movement is necessary for roughness constancy in both direct and indirect touch. Roughness constancy is also preserved during pseudo-passive scanning, which stresses the importance of proprioceptive input. The results show that cutaneous input provides the signals necessary for roughness perception and that proprioceptive input resulting from hand movement-rather than a motor efference copy-is necessary to achieve roughness constancy. These findings have important implications in providing realistic sensory feedback for prosthetic-hand users.

Cutaneous afferent input does not modulate motor intracortical inhibition in ageing men

Afferent input has been shown to be a powerful modulator of cortical inhibition. Such modulation is likely to be important for the control of ongoing movement, but may also play a role in facilitating neuroplastic reorganisation. Human motor control and neuroplasticity both decline with ageing, whereas the efficacy of short-interval intracortical inhibition (SICI) appears not to. We examined if ageing alters the efficacy of afferent modulation of SICI. Previously, electrical cutaneous stimulation of a finger has been shown to reduce SICI in the motor cortices of young adults. Paired-pulse transcranial magnetic stimulation was used to assess SICI in the cortical representation of the first dorsal interosseous muscle. SICI was assessed separately under two conditions: with and without prior afferent input from electrical cutaneous stimulation of the index finger. Fifteen 'young' (20.1 ± 2.1 years) and 15 'old' male humans (65.5 ± 3.9 years) were studied. SICI did not differ when young and old males were compared. However, when preceded by electrical cutaneous finger stimulation, SICI was reduced in young men but not old men. Reflex testing indicated preservation of the afferent volley to the cortex. These findings suggest that a contributing factor in the decline of motor function, and possibly neuroplasticity, with ageing is loss of SICI modulation, probably due to altered cortical sensorimotor integration of afferent input.

Enhanced propriospinal excitation from hand muscles to wrist flexors during reach-to-grasp in humans

In humans, propriospinal neurons located at midcervical levels receive peripheral and corticospinal inputs and probably participate in the control of grip tasks, but their role in reaching movements, as observed in cats and primates, is still an open question. The effect of ulnar nerve stimulation on flexor carpi radialis (FCR) motor evoked potential (MEP) was tested during reaching tasks and tonic wrist flexion. Significant MEP facilitation was observed at the end of reach during reach-to-grasp but not during grasp, reach-to-point, or tonic contractions. MEP facilitation occurred at a longer interstimulus interval than expected for convergence of corticospinal and afferent volleys at motoneuron level and was not paralleled by a change in the H-reflex. These findings suggest convergence of the two volleys at propriospinal level. Ulnar-induced MEP facilitation was observed when conditioning stimuli were at 0.75 motor response threshold (MT), but not 1 MT. This favors an increased excitability of propriospinal neurons rather than depression of their feedback inhibition, as has been observed during tonic power grip tasks. It is suggested that the ulnar-induced facilitation of FCR MEP during reach may be due to descending activation of propriospinal neurons, assisting the early recruitment of large motoneurons for rapid movement. Because the feedback inhibitory control is still open, this excitation can be truncated by cutaneous inputs from the palmar side of the hand during grasp, thus assisting movement termination. It is concluded that the feedforward activation of propriospinal neurons and their feedback control may be involved in the internal model, motor planning, and online adjustments for reach-to-grasp movements in humans.

Persistence of locomotor-related interlimb reflex networks during walking after stroke

ObjectiveCutaneous nerve stimulation evokes coordinated and phase-modulated reflex output widely distributed to muscles of all four limbs during walking. Accessibility to this distributed network after stroke offers insight into the pathological changes and suggests utility for therapeutic applications. Here we examined muscles in both the more (MA) and less affected (LA) legs evoked by stimulation at the ankle and wrist during walking in chronic (>6months post CVA) stroke. MethodsStroke and control participants walked on a treadmill with a harness support system. Reflexes were evoked with trains of electrical stimuli delivered separately to the cutaneous superficial peroneal (SP; at the ankle) and superficial radial (SR; at the wrist) nerves. Background locomotor and reflex EMG were phase-averaged across the gait cycle and analyzed off line. ResultsLocomotor background muscle activation patterns were altered bilaterally in stroke, as compared with control. Phase-dependent modulation of interlimb cutaneous reflexes was found in both stroke and control subjects with stimulation of each nerve, but responses were blunted in stroke. Reflex reversal in tibialis anterior (TA) at heel strike with SP nerve stimulation was present in both groups. Notably, SR nerve stimulation produced facilitation during the swing-to-stance transition in the TA and suppression of MG in the MA leg during stance. ConclusionsInterlimb cutaneous inputs may access coordinated reflex pathways in the MA limb during walking after stroke. Importantly activation in these pathways could provoke responses to counter foot drop during swing phase of walking. Additionally, our data support the perspective that there is no “unaffected” side after stroke and that caution should be used when interpreting the LA side as “control” after stroke. SignificanceThe presence of functionally-relevant interlimb cutaneous reflexes in the MA leg presents a substrate that may be strengthened by rehabilitation.