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Reflex heating of the skin and telethermography.

Thermography is a very instructive method of showing the interaction between direct changes in heat radiation of the surface of the skin and its relation to reflex processes. Short reflex arcs, such as the axon reflex, and long reflex arcs, such as the flush phenomenon provoked by visceral cutaneous reflexes, can be illustrated by thermography. Reflex changes in skin temperature by cooling the skin of patients with cold contact urticaria with the short-term development of large areas of warmed skin and also reflex changes in cholinergic urticaria after prior heating of such skin areas where the urticarial eruption follows are reflected in the picture of telethermography. Reflex arcs between extremities affect the arterial blood flow and show individual variation, as in the case of the cessation of permanent local vasoconstriction by indirect cooling. The thermographic illustration of convective heat transport to the skin surface can be obstructed by the cooling effect of sweat secretion, e.g., water vapor exudation and by changes in conductive heat transport in abnormal skin.

Adult↗

Forelimb reflexes modulated by tonic neck positions in cats.

The modulation of monosynaptic forelimb reflexes by tonic neck positions was investigated in cats with the head fixed. Lateral flexion of the body in a horizontal plane markedly facilitates reflexes of the deep radial nerve (DR) in the ipsilateral forelimb, while the antagonistic ulnar nerve (ULN) reflexes are strongly inhibited. Opposite effects are seen after contralaternal body movement. Dorsiflexion of the body clearly increases DR-reflexes and exerts a reciprocal although more pronounced inhibition on ULN reflexes. Opposite effects appear after ventriflexion. The reflex modulation starts with head-body displacements of approximately 5 degrees and increases with increasing angles. Furthermore reflex modulation does not depend on the intact cerebrum and cerebellum. The comparison of forelimb and hindlimb reflexes shows a decrease of the neck influences along the spinal cord.

Animals↗

Modulation of hindlimb reflexes by tonic neck positions in cats.

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 antagnostic deep peroneal nerve (DP), occurs following lateral flexion and rotation around the body axis at an angle of 25 degrees. 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 degrees-15 degrees. 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.

Afferent Pathways↗

Genetically associated similarities and differences in the generation of neurons comprising an early developing reflex pathway in mouse spinal cord.

Tritiated thymidine autoradiography has been used to study the generation of lateral motor neurons (LMNs), association interneurons (ANs) and dorsal root ganglion cells (DRGNs) in the spinal cords of genetically diverse strains of mice. The neuronal populations analyzed in this study form an early developing reflex pathway and the ontogeny of this circuit exhibits genetically associated variability. The strains of mice used in this investigation have been shown to differ in the embryonic age at which forelimb reflex movements are first manifest and in the timing of synapse formation within the reflex pathway. A precocious development of these reflex traits occurs in strain C57BL/6J in comparison to embryos of intermediate (CBA/CaJ) and late developing (LP/J) strains. All three inbred strains show the same basic generation sequence for the neuronal populations comprising the forelimb reflex pathway. The generation of LMNs precedes that of ANs, and the generation of ANs, in turn, precedes that of DRGNs. Since this is the same sequence as that observed for the formation of synaptic junctions in all three strains, it is suggested that synaptogenic sequences in reflex circuits may be determined by the generation sequence of the component neuronal populations. Although the strains all exhibit the same basic sequence of neuronal generation, the temporal relationships of the generation of each cell population within this sequence show significant strain dependent variations. C57BL/6J displays a larger temporal separation between the generation of each cell type than LP/J, and CBA/CaJ is intermediate to the other two strains in this respect. The fact that this strain order is identical to that observed for the development of reflex traits suggests that genetically associated differences in the timing of neuronal birthdays within a common generation sequence may have a substantial influence on the timing of synaptogenesis within the reflex pathway.

Animals↗

Role of spinal serotonin1 receptor subtypes in thermally and mechanically elicited nociceptive reflexes.

The ability of 5-HT1A and 5-HT1B agonists to alter a spinal animal's nociceptive threshold was examined using two analgesiometric tests. In the spinal withdrawal reflex test, administration of the selective 5-HT1A agonists ipsapirone, gepirone and PAPP resulted in significant dose-dependent increases in receptive field (RF) area for withdrawal reflexes when compared to predrug baseline values, indicating an increase in nociceptive sensitivity. The average overall percent maximal increase in RF area following administration of 5-HT1A selective compounds was: 80 +/- 16% for the ventroflexion reflex, 90 +/- 6% for the dorsiflexion reflex and 87 +/- 8% for the lateral flexion reflex. Similar to the effects noted with 5-HT1A agonists, administration of 5-HT1B agonists RU24969, mCPP and TFMPP resulted in a hyperalgesic response with an overall percent maximal increase of 43 +/- 6% for the ventroflexion reflex, 51 +/- 6% for the dorsiflexion reflex and 38 +/- 9% for the lateral flexion reflex. In the tail-flick analgesiometric test, administration of the 5-HT1A agonists 8-OH-DPAT and ipsapirone and the 5-HT1B agonists RU24969 and mCPP resulted in a significant dose-dependent increase in tail-flick latencies when compared to predrug baseline values, indicating a decrease in nociceptive sensitivity to noxious thermal stimuli. No differences in magnitude of the effect of the two receptor subtypes were found, indicating that stimulation of either 5-HT1A or 5-HT1B receptors was equipotent in producing the antinociceptive tail-flick response.(ABSTRACT TRUNCATED AT 250 WORDS)

8-Hydroxy-2-(di-n-propylamino)tetralin↗

An experimental psychophysiological approach to human bradycardiac reflexes.

Bradycardic reflexes in man are both of scientific and clinical interest. Using the methods of experimental psychophysiology, control over relevant independent variables permits the study of fine-grained temporal physiologic response topographies, and of psychological factors that may modify the reflex. In addition, information can also be sought through interdisciplinary collaborations with experimental physiologists in order to shed light on the mechanism of the reflexes. These general features of the approach are illustrated by presenting data on two bradycardic reflex preparations: the laboratory dive analog, and the 90-degree negative tilt. The dive-analog studies have shown that a) the dive-reflex proper is a late-occurring bradycardia accompanied by a late-occurring vasoconstriction; and b) for the elicitation of this reflex, both breath-holding and face immersion are necessary. In addition, the physiologic manipulation of temperature affects the reflex in an inverse way over the range of 10 degrees to 40 degrees C, while the sense of control (a psychological variable) attenuates the reflex. The negative-tilt preparation produces a bradycardic response that is ideal as a Pavlovian unconditional response. Some Pavlovian conditioning arrangements, especially an "imaginational" form, do produce significant conditional bradycardic responding, and this has both potential clinical (e.g., biofeedback-related) and theoretical (e.g., S-R vs. S-S accounts of Pavlovian conditioning) applications. The paper ends with a comment on the cognitive paradigm shift in psychology. Although this shift is of importance, it is suggested that it is also important to "remember the response."

Biofeedback, Psychology↗

Effects of changes in hip joint angle on H-reflex excitability in humans.

We examined the amplitude modulation of the soleus (Sol) H-reflex during controlled variations of the hip joint angle in 21 healthy adult human subjects. Hip angle variations were imposed separately, or in combination either with stimulation of the plantar skin or with electrical activation of muscle afferents from the medial gastrocnemius (MG) nerve. We found that with subjects in the supine position, flexion of the hip significantly depressed Sol H-reflex excitability, by as much as 50% of control reflex values (Ho) recorded at 10 degrees of hip flexion. Conversely, significant facilitation of the H-reflex was observed when the hip joint was extended (10 degrees), with amplitudes reaching 200+/-15.3% of Ho. Changes in H-reflex amplitude were also observed during electrical stimulation of either the foot sole or the MG nerve, when stimuli were delivered at different hip angles. Foot sole stimulation resulted in facilitation of the H-reflex with the hip extended while depression of the reflex was recorded with the hip flexed. In contrast, MG nerve stimulation at group-I muscle afferent strength resulted in a significant increase in the Sol H-reflex magnitude with the hip flexed, while during hip extension, no significant effects were observed [corrected]. This study provides evidence for the existence of a spinal mechanism, determined principally by the hip joint angle, which promotes switching between inhibitory and facilitatory pathways during hip flexion and extension. The origins of such a spinal mechanism are discussed.

Adult↗

Characterisation of the quadriceps stretch reflex during the transition from swing to stance phase of human walking.

The main objective of this study was to characterize the stretch reflex response of the human thigh muscles to an unexpected knee flexion at the transition from stance to swing during walking. Eleven healthy subjects walked on a treadmill at their preferred speed. Reliable and constant knee flexions (6-12 degrees amplitude, 230-350 degrees /s velocity, 220 ms duration) were applied during the late swing and early stance phase of human walking by rotating the knee joint with a specifically designed portable stretch apparatus affixed to the left knee. Responses from rectus femoris (RF), vastus lateralis (VL), vastus medialis (VM), biceps femoris (BF), medial hamstrings (MH) and medial gastrocnemius (GM) were recorded via bipolar surface electromyograms (EMG). The onset of the response in the RF, VL and VM, remained stable and independent of the time in the step cycle when the stretch was applied. Across all subjects the response onset (mean +/- SD) occurred at 23+/-1, 24+/-1 and 23+/-1 ms for RF, VL and VM, respectively. The duration of the initial response was 90-110 ms, at which time the EMG signal returned towards baseline levels. Three reflex response windows, labelled the short latency reflex (SLR), the medium latency reflex (MLR) and the late latency reflex response (LLR), were analysed. The medium and late reflex responses of all knee extensors increased significantly ( p=0.008) as the gait cycle progressed from swing to stance. This was not related to the background EMG activity. In contrast, during standing at extensor EMG levels similar to those attained during walking the reflex responses were dependent on background EMG. During walking, LLR amplitudes expressed as a function of the background activity were on average two to three times greater than SLR and MLR reflex amplitudes. Distinct differences in SLR and LLR amplitude were observed for RF, VL and VM but not in the MLR amplitude. This may be related to the different pathways mediating the SLR, MLR and LLR components of the stretch response. As for the knee extensor antagonists, they exhibited a response to the stretch of the quadriceps at latencies short enough to be monosynaptic. This is in agreement with the suggestion by Eccles and Lundberg (1958) that there may be functional excitatory connections between the knee extensors and flexors in mammals.

Adult↗

Stretch reflexes in human abdominal muscles.

Homonymous and heteronymous reflex connections of the abdominal muscles were investigated by the application of a tap to the muscle belly and observation of surface electromyographic responses. Reflex responses of the following abdominal muscles were investigated both ipsilateral and contralateral to the tap: rectus abdominis (RA), external oblique (EO) and internal oblique (IO). 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 also observed in muscles on both ipsilateral and contralateral sides in response to the same stimulus. The latencies of the crossed responses were only marginally longer than responses evoked in the respective ipsilateral muscle. Moreover, the reflexes evoked in the IO muscle from ipsilateral and contralateral IO muscle afferents were of comparable amplitude, as were those reflexes evoked in ipsilateral and contralateral EO and RA muscles when tapping IO. These similarities in the reflex characteristics on the ipsilateral and contralateral sides suggest that abdominal muscle afferents activate similar pathways to muscles on both sides of the body. It follows that if the homonymous stretch reflex of abdominal muscles have a monosynaptic component, then a similar monosynaptic pathway activates synergistic motoneurones, not only ipsilaterally but also contralaterally.

Afferent Pathways↗

Stretch reflex instability compared in three different human muscles.

The possibility of causing instability in the stretch reflex has been examined in three different human muscles: biceps, first dorsal interosseous (FDI) of the hand and digastric. Tremor recorded as fluctuation of isometric force was compared with that occurring during contraction against a spring load. The spring compliance was selected to make the natural frequency of the part in each case appropriate for oscillations in the short latency stretch reflex. A computer model of the whole system was used to predict the frequency at which oscillations should be expected and to estimate the reflex gain required in each case to cause sustained oscillations. Estimates were computed of the autospectra of the force records and of the rectified surface EMG signals and of the coherence functions. Normal subjects showed no evidence of a distinct spectral peak during isometric recording from any of the three muscles. However, in anisometric conditions regular oscillations in force occurred in biceps, but not in FDI or digastric. The oscillations in biceps at 8-9 Hz were accompanied by similar oscillations in the EMG which were highly coherent with the force signal. The results are consistent with the presence of a strong segmental stretch reflex effect in biceps and weak or absent reflex in FDI. Digastric is known to contain no muscle spindles and therefore to lack a stretch reflex. In two subjects who volunteered that they had more tremor than normal, but had no known neurological abnormality, there was a distinct peak in the force spectrum at 8-9 Hz in biceps and FDI in isometric conditions with coherent EMG activity. The peak increased in size in anisometric conditions in biceps but not in FDI. This component appears to be of central rather than of reflex origin. No equivalent component was found in digastric records. The results are discussed in relation to the possible role of the short latency stretch reflex in the genesis of physiological tremor in different muscles.

Adult↗

Exteroceptive reflexes in jaw-closing muscle EMG during rhythmic jaw closing and clenching in man.

Exteroceptive jaw reflexes might play a role in normal functions of the mouth such as mastication. Until now these reflexes have only been studied under isometric conditions. The aim of this study was to compare exteroceptive reflexes in jaw muscle EMG during the closing phase of rhythmic open-close movements and clenching, at the same jaw gape and with similar muscle EMG. Reflexes consisting of successive waves of decreased and increased muscle activity (the Q, R, S and T waves of the post-stimulus electromyographic complex (PSEC)), evoked by light noxious electrical stimulation of the vermillion border of the lower lip, were recorded from the jaw closing muscles of 17 subjects. Differences between the two tasks occurred in two phases of the PSEC: (1) in an early phase, around the R wave, there was significantly less EMG during jaw closing (mean EMG ratio between jaw-closing and clenching 0.71), and (2) in a late phase, around the transition between the S to the T wave, there was significantly more EMG during jaw closing (mean EMG ratio: 1.40). The decrease in EMG activity around the R wave during jaw closing may be due to a change in reflex sensitivity at an interneuron level. The increase in EMG activity around the transition between the S and T waves during jaw closing might, at least in part, be due to a proprioceptive stretch reflex. This reflex is mediated by muscles spindles that are activated by the deceleration of the jaw evoked by the lip stimulus. The finding of inhibitory reflex mechanisms that predominate more during rhythmic jaw movements than during clenching in an early phase of the PSEC might be related to protecting oral tissues from trauma when the jaw is closing with potentially a large muscle force. In contrast, when food is held between the teeth, a possible inhibitory influence of light noxious stimuli is diminished.

Adolescent↗

Subthreshold transcranial magnetic stimulation during the long latency component of the cutaneomotor reflex.

Modulation of ongoing electromyographic (EMG) activity in the small hand muscles can be induced by electrical stimulation of the digital nerves or by stimulation of the skin of the fingers. Several groups have attempted to establish a role for the motor cortex in the generation of the facilitatory component of the cutaneomotor reflex. Our aim was to establish if the facilitatory component of the reflex could be diminished by a procedure known to inhibit the motor cortex, namely, subthreshold transcranial magnetic stimulation. During sustained small contractions of the first dorsal interosseus muscle transcranial magnetic stimuli (TMS), which were subthreshold for the generation of a motor evoked potential, were delivered via a figure-of-eight coil. Inhibition of ongoing EMG was observed in all subjects. In two separate series of trials, TMS was timed so that the resultant inhibition occurred coincident with either the short or long latency stretch reflex or with the initial or later part of the facilitatory component of the cutaneomotor reflex. The short latency stretch reflex is known to involve a largely monosynaptic loop via the spinal cord, whereas the long latency response involves a transcortical loop. The long latency response was reduced in size following subthreshold TMS, whereas the short latency response was unchanged. This provides evidence of the effectiveness of subthreshold TMS in inhibiting a transcortical reflex. When the TMS was timed so that the inhibition occurred coincident with the facilitatory component of the cutaneomotor response, neither the early nor later changes were inhibited. Thus, the pathway of the long-latency cutaneomotor reflex is not similar to the transcortical pathway of the stretch reflex. Either the response does not travel via the cortex or it involves different cortical neurones.

Adult↗

Somatosensory graviception inhibits soleus H-reflex gain in humans during walking.

To investigate the effects of gravity-related somatosensory information on spinal human reflexes, the soleus H-reflex was recorded in ten healthy subjects walking on a treadmill at 2.0 km/h on land and in water. The modulation pattern of the soleus H-reflex was determined in ten different phases of the step cycle. While the subjects were walking in water, the background electromyographic activity (BGA) of the soleus was lower than that on land; on the other hand, the soleus H-reflex amplitude while the subjects were walking in water showed no significant differences throughout the step cycle compared with that while the subjects were on land; the phase-dependent soleus H-reflex modulation pattern was well preserved while walking in water. There was a linear relationship between the BGA and the H-reflex amplitude in each condition; however, the soleus H-reflex gain while walking in water was significantly higher than that on land. These findings suggest that the somatosensory graviception can markedly reduce the spinal reflex excitability. Our findings are discussed in relation to human gait; therefore, further studies are needed to clarify the effect of somatosensory graviception on human neural mechanisms.

Adult↗

Stretch reflex modulation during imposed static and dynamic hip movements in standing humans.

The purpose of this study was to investigate the effects of hip proprioceptors on soleus stretch reflex excitability in standing humans. A custom-made device to stretch the ankle extensors was mounted on the lower leg portion of a gait orthosis and was used to elicit stretch reflex responses while standing. Six subjects with motor complete spinal cord injury (SCI) and six spinal intact subjects were placed in the orthosis, and stretch reflex responses were elicited when static and/or dynamic hip joint angle changes were imposed. We found that static hip extension significantly enhanced the stretch reflex responses as compared to the neutral position and the hip flexion position only in the SCI group. The EMG magnitude induced by hip extension was 142 +/- 16.6% greater than that induced by the neutral position. When the leg was dynamically swung, the reflex responses also changed with the phase of the hip angle in the SCI group; in particular, the reflex amplitude was enhanced with hip extension and in the transition phase from flexion to extension. Although the magnitude of the changes was less than that in the SCI group, a similar type of modulation was found in the normal group. Given the fact that the persons with SCI had lost the neural connection between higher nervous center and the paralyzed lower limb muscles, the mechanism underlying the present results can be attributed to the peripheral afferent input due to the hip angle changes. We concluded that hip mediated afferent input has a significant influence on the excitability modulation of the soleus stretch reflex pathway. Such neural modulation may play a role in the mechanism responsible for the phase-dependent modulation of the stretch reflex while walking.

Adult↗

Comparison of the depression of H-reflexes following previous activation in upper and lower limb muscles in human subjects.

When conditioning-testing (C-T) stimuli are applied to Ia afferents to elicit H-reflexes, the test reflex is abolished immediately following the conditioning reflex. As the C-T interval is increased, the test response slowly begins to recover, taking several hundred milliseconds to attain control values. The time course of this recovery is known as the H-reflex recovery curve. H-reflex recovery curves were compared using surface EMG and single motor unit activities in lower limb soleus and upper limb flexor carpi radialis (FCR) muscles in seven healthy human subjects. Under rest conditions, the recovery of H-reflexes and single motor unit activity was slow for soleus; the recovery was not complete even in 1 s. In comparison, the recovery was very fast for FCR motor units, occurring in 200-300 ms. The effects of rate of stimulation (0.1-10.0 imp/s) were also examined on the magnitude of H-reflex responses. The reflex response declined with increasing rate of stimulation, the decline being slightly greater in soleus than in FCR. When these phenomena were examined with voluntary facilitation of the spinal cord, the time of recovery shortened and the effect of stimulus rate also diminished. Changes with background facilitation were greater in FCR than in soleus. The differences between the two muscles are attributed mainly to differences in presynaptic inhibition in the two spinal segments, and/or to the differences in dynamics of the transmitter release in terminals of Ia afferents synapsing with slow soleus motoneurons and those synapsing with the fast FCR motoneurons.

Adult↗

Context dependency of a limb withdrawal reflex in the caterpillar Manduca sexta.

The proleg withdrawal reflex in the caterpillar Manduca sexta is a robust, well-characterized system for investigating the integration of sensory information with centrally patterned behavior. The reflex is evoked by stimulating mechanosensory hairs--planta hairs--located at the tip of each proleg. We studied the expression of this reflex by combining video recordings and electromyographic recordings from the main retractor muscles of the proleg, the principal and accessory planta retractor muscles. In intact animals, the nature of the response depended on the motor context of the animal. Animals which were standing quietly showed great variability in both the kinematic properties of proleg withdrawal, and the corresponding muscle electrical activity. Animals which were hanging upside down from a wooden dowel exhibited a much faster reflex, with retraction of the proleg occurring slightly faster than in standing animals, but re-extension of the proleg to the substrate being considerably faster. In crawling animals, expression of the reflex depended on the phase of the crawling cycle during which stimulation occurred. The reflex in a given proleg was suppressed during stance phase of that proleg. During swing phase, however, planta hair stimulation evoked proleg withdrawal, resulting in an assistance reflex. In contrast. isolated abdomens showed much less variability in the reflex. A comparison of the relationship between retractor muscle activity and the resulting proleg movement showed significant correlations between both the duration of activity and the number of muscle spikes, and the size of the associated proleg withdrawal. This is a promising system in which to investigate how central neuronal circuits accomplish context-dependency of motor behavior.

Animals↗

Considerations for use of the Hoffmann reflex in exercise studies.

There continues to be great interest in evaluating the adaptive plasticity of the human nervous system in response to exercise training or other interventions. For various reasons, researchers have been interested in estimates of spinal reflex processing in intact human subjects before and after training. A reflex pathway that has been employed in this regard is the Hoffmann (H) reflex. This brief review describes the basic procedure for evoking the H reflex in different muscles. Other sections address methodological issues that affect interpretation of the H reflex. In particular, the role that presynaptic inhibition serves in the modification of the H reflex and how this precludes its use as an unambiguous measure of alpha-motoneuron excitability will be discussed. Applications of the H reflex to study adaptive plasticity in humans is also reviewed, and methodological requirements that should be maintained for accurate interpretation of H reflexes in exercise studies are presented.

Electromyography↗

Changes in reflex excitability following isometric contraction in humans.

Enhancement of muscle stretch following isometric contraction has been thought to occur as a result of inhibitory reflex mechanisms. Experiments with electrical stimulation (H-reflex) have demonstrated maximal H-reflex suppression during force relaxation followed by gradual recovery over the following 20 s. There has been considerable speculation as to whether electrical and mechanical stimulation elicit similar response behaviour. The present study examined postisometric reflex modulation following both stimulation modalities. In ten subjects dorsiflexion stimuli varying in speed and amplitude were applied after 30% and 60% maximal voluntary contraction (MVC). Modulation of the mechanically and electrically evoked responses following isometric plantarflexion was investigated. Reflex responses following both stimulation modalities were depressed during the course of force relaxation. A rather fast recovery was observed in mechanical stimulation. Postisometric response modulation was neither altered by the amount of isometric plantarflexion, nor by the amplitude of the applied stretch stimulus. With increasing velocity of the applied dorsiflexion, however, the shape of the reflex modulation persisted, but the magnitude of the responses was significantly enhanced. In electrical stimulation, however, recovery was delayed. It is suggested that postisometric reflex modulation is due to presynaptic inhibition. Moreover, possible peripheral mechanisms resulting from alpha-gamma-coactivation may also affect the stretch receptor itself because of inherent stiffness properties. The latter possibility particularly would explain the differences between mechanical and electrical stimulus modalities. With respect to practical implications, the very fast recovery (< 400 ms) of the stretch responses to control values strongly contradicts the interpretation that after isometric precontraction, suppression of reflex activity might be used for more efficient stretching of the tendomuscle system.

Adult↗