Journal of Neurophysiology

Kirsch, R. F.; Rymer, W. Z.

doi: N/Apmid: 3598635

Abstract We have investigated the role of reflex regulation of muscle force in normal human subjects by comparing changes in the stretch-evoked increments in elbow joint flexor electromyogram (EMG) and elbow joint torque before and after fatigue. Elbow flexor muscle fatigue was induced by repetitive voluntary isometric contractions. To assess the appropriateness of the EMG signal as an index of neural excitation of muscle under fatiguing conditions, we examined the time course of recovery of joint torque and EMG power spectrum following fatigue. Fatigue-related changes in the EMG power spectra recovered within 5–10 min after fatiguing exercise was terminated, yet the muscle weakness induced by the exercise lasted greater than 7 h and was substantial in magnitude. The decoupling of torque and EMG recovery allowed us to compare pre- and postfatigue EMG stretch responses without adjusting for differences in EMG spectral content. Torque and EMG responses to stretch were quantified by time-averaging over 250-ms “isometric” and “steady-state” periods, just before and just after a ramp angular stretch of the elbow joint, respectively. The torque increment elicited by stretch was lower following fatigue in seven of eight experiments. However, the average decrease of 20.13 +/- 14.42% in these seven subjects was somewhat smaller than the corresponding average shift in the slope of the isometric EMG-torque relationship of 85.84 +/- 90.29% (n = 8). Furthermore, the stretch-induced EMG increment was larger following fatigue in all eight sessions (average of 56.14 +/- 28.96%, n = 8), with six of the shifts reaching statistical significance for alpha = 0.05. Because the pattern of torque and EMG responses before and after fatigue suggested the presence of an active force regulator, we used a simple model of the neuromuscular system to estimate a loop gain value for each session. When pre- and postfatigue responses were matched by isometric background torque level, an average loop gain value of 7.9 was computed, whereas for responses matched by average prestretch EMG level, the loop gain estimates averaged 2.1. Although our assessment of force regulation was essentially static and derived from the responses to a single type of perturbation, the change in the incremental torque and EMG stretch responses indicates that meaningful neural compensation for fatigue occurred. Moreover, the loop gain estimates derived from these responses are an order of magnitude larger than those previously reported in animal models, suggesting that force regulation may be important in the control of human muscle contraction. Copyright © 1987 the American Physiological Society

Pare, D.; Steriade, M.; Deschenes, M.; Oakson, G.

doi: N/Apmid: 3037038

Abstract This study tested the hypothesis that neurons of thalamic nuclei, which are normally devoid of inputs from the reticular thalamic nucleus, do not display spindle oscillations and related rhythmic spike bursts. This proposal derived from our recent studies indicating that the reticular nucleus is the generator of spindling rhythmicity. We used retrograde tracing methods, intracellular recordings in barbiturized cats, and extracellular recordings of single neurons and field potentials in anteroventral (AV), anteromedial (AM), ventroanterior (VA), ventrolateral (VL), and central lateral (CL) thalamic nuclei in cats with rostral brain stem transections (cerveau isole preparations), before and after administration of barbiturates. The observation that AV and AM nuclei do not receive inputs from the reticular nucleus was confirmed by using injections of horseradish peroxidase conjugated to wheat germ agglutinin confined within the limits of anterior nuclei. Such injections led to massive retrograde labeling in mammillary nuclei and layer VI of the retrosplenial cortex but left free of labeling the neurons of the reticular thalamic nucleus. Intracellular recordings showed that AV-AM neurons discharge tonically in response to a depolarizing current applied at rest, whereas they give rise to a slow spike that underlies a burst of fast action potentials when the membrane is hyperpolarized by 5-12 mV. Despite the fact that they share similar properties with other thalamic neurons, intracellularly recorded AV-AM neurons do not exhibit spindle waves under barbiturate anesthesia, whereas VA-VL, CL, and other thalamocortical neurons that receive afferents from the reticular nucleus commonly display such oscillations. With extracellular recordings performed simultaneously in CL and AV or AM nuclei of the unanesthetized cerveau isole preparation, focal spindle oscillations and related rhythmic high-frequency spike bursts of single CL cells contrasted with absence of spindles and spike bursts in AV or AM neurons. Spindling could be induced in AV-AM nuclei only after administration of barbiturates at doses exceeding 3 mg/kg, and it appeared approximately 35–40 s after the barbiturate effect was detected in the simultaneously recorded CL nucleus. Moreover, the spike bursts that were elicited in AV-AM neurons after barbiturate administration were not temporally related with focal spindles. Since spindle oscillations did not appear intracellularly in AV-AM neurons, the possibility was envisaged that barbiturate-induced spindles were the passive reflection of field potentials actively generated in neighboring thalamic nuclei.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1987 the American Physiological Society

Kajander, K. C.; Giesler, G. J.

doi: N/Apmid: 3598627

Abstract The majority of neurons at the origin of the spinocervical tract are driven by noxious stimulation of their receptive fields. Surprisingly, previous studies have encountered only a small percentage of nociceptive neurons within the terminus of the spinocervical tract, the lateral cervical nucleus (LCN). To determine if previous reports have underestimated the proportion of nociceptive LCN neurons, 129 neurons within the nucleus were physiologically identified and examined in cats prepared using three different methods. Fifty-nine percent of the neurons studied in unanesthetized cats that were decerebrated and spinalized responded either differentially or exclusively to noxious mechanical stimulation of the skin within discrete receptive fields. LCN neurons also gave accelerating responses to increasingly more intense noxious thermal stimuli. LCN neurons are, therefore, capable of coding both the intensity and location of noxious stimuli. Only 6% of LCN neurons responded to noxious cutaneous stimuli in unanesthetized, decerebrated cats in which the spinal cord was intact. Only 4% of LCN neurons in intact urethan-anesthetized cats were driven by noxious stimulation. Several previous studies of the LCN have been performed in cats that were deeply anesthetized with barbiturates. Therefore, the effects of barbiturates on the nociceptive responses of LCN neurons were determined. Subanesthetic doses of intravenously administered barbiturates reduced or eliminated the responses of nociceptive LCN neurons to noxious thermal stimuli in decerebrated and spinalized cats. Responses to innocuous mechanical stimuli by these neurons were not blocked by barbiturates. Nociceptive LCN neurons in decerebrated and spinalized cats were somatotopically organized. Neurons with forelimb receptive fields were located in the ventromedial half of the LCN; neurons with hindlimb receptive fields were located in the dorsolateral half of the nucleus. This report and previous studies of the spinocervical tract suggest that the spinocervicothalamic pathway is capable of playing an important role in nociception. Copyright © 1987 the American Physiological Society

Rutecki, P. A.; Lebeda, F. J.; Johnston, D.

doi: N/Apmid: 3037040

Abstract Using extra- and intracellular recording techniques, we investigated the epileptiform activity induced by low concentrations (5 and 10 microM) of bath-applied 4-aminopyridine (4-AP) in the CA3 subfield of rat hippocampal slices. We also studied the effects of 4-AP on the excitatory and inhibitory synaptic conductance changes in CA3 neurons produced by mossy fiber stimulation. Low concentrations of 4-AP induced spontaneously occurring epileptiform discharges at extracellular potassium concentrations between 1 and 10 mM. In contrast, picrotoxin and bicuculline produced spontaneous epileptiform discharges at extracellular potassium concentrations between 5 and 10 mM. The paroxysmal depolarizing shift (PDS) induced by 4-AP was also investigated. At potentials between -40 and -10 mV, the waveform of the PDS consisted of a depolarizing component enveloped by a hyperpolarizing component. The amplitude of the depolarizing component of the PDS was a monotonic function of the membrane potential, and the mean measured reversal potential was -25.7 mV. Under voltage-clamp conditions, the measured conductance associated with the depolarizing component of the PDS averaged 110 nS, with a reversal potential of -14.1 mV. Application of 5 microM 4-AP produced an increase in the inhibitory synaptic conductance change calculated from currents measured 15 ms following mossy fiber stimulation. The mean value increased from 35.2 to 58.1 nS (P less than 0.05) without a significant change in reversal potential. A concentration of 10 microM 4-AP also produced an increase in this inhibitory synaptic conductance change (from 53.3 to 66.3 nS, P less than 0.05) but caused a significant depolarization of the reversal potential (from -66.5 to -61.6 mV, P less than 0.05). This change in reversal potential may reflect a prolongation of the excitatory synaptic currents produced by 4-AP that contributes to the current measured 15 ms from the stimulus. Following application of either 5 or 10 microM 4-AP, there were no significant changes in the resting potential or input resistance of the neurons studied. Application of 5 microM 4-AP also significantly increased the amplitude of the measured excitatory synaptic conductance change produced by mossy fiber stimulation (from 27.9 to 44.1 nS, P less than 0.05) without producing a change in the reversal potential. In 5 of 21 neurons studied, a long-lasting outward synaptic current was present at holding potentials near rest following mossy fiber stimulation.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1987 the American Physiological Society

Kavanagh, G. L.; Kelly, J. B.

doi: N/Apmid: 3598629

Abstract Ferrets were tested in a semicircular apparatus to determine the effects of auditory cortical lesions on their ability to localize sounds in space. They were trained to initiate trials while facing forward in the apparatus, and sounds were presented from one of two loudspeakers located in the horizontal plane. Minimum audible angles were obtained for three different positions, viz., the left hemifield, with loudspeakers centered around -60 degrees azimuth; the right hemifield, with loudspeakers centered around +60 degrees azimuth; and the midline with loudspeakers centered around 0 degrees azimuth. Animals with large bilateral lesions had severe impairments in localizing a single click in the midline test. Following complete destruction of the auditory cortex performance was only marginally above the level expected by chance even at large angles of speaker separation. Severe impairments were also found in localization of single clicks in both left and right lateral fields. In contrast, bilateral lesions restricted to the primary auditory cortex resulted in minimal impairments in midline localization. The same lesions, however, produced severe impairments in localization of single clicks in both left and right lateral fields. Large unilateral lesions that destroyed auditory cortex in one hemisphere resulted in an inability to localize single clicks in the contralateral hemifield. In contrast, no impairments were found in the midline test or in the ipsilateral hemifield. Unilateral lesions of the primary auditory cortex resulted in severe contralateral field deficits equivalent to those seen following complete unilateral destruction of auditory cortex. No deficits were seen in either the midline or the ipsilateral tests. Copyright © 1987 the American Physiological Society

Miller, A. D.; Tan, L. K.; Suzuki, I.

doi: N/Apmid: 2955084

Abstract The role of ventral respiratory group (VRG) expiratory (E) neurons in the control of abdominal and internal intercostal (expiratory) muscle activity during vomiting was examined in decerebrate cats by recording from these neurons during fictive vomiting in paralyzed animals and comparing abdominal muscle activity during vomiting before and after sectioning the axons of these descending neurons. Fictive vomiting was defined by a series of bursts of coactivation of abdominal and phrenic nerves elicited by either subdiaphragmatic vagus nerve stimulation or emetic drugs. Such coordinated activity would be expected to produce vomiting if the animals were not paralyzed. Data were recorded from 27 VRG E neurons that were antidromically activated from the lower thoracic (T13) or lumbar spinal cord. During fictive vomiting, almost two-thirds of these neurons (17/27) were mainly active in between periods of abdominal and phrenic nerve coactivation, when the internal intercostal motoneurons are known to be active. This group of neurons was termed INT neurons. INT neurons were subdivided according to whether they were active between every burst of phrenic and abdominal nerve coactivation (INTa neurons, n = 10) or only between some bursts (INTb neurons, n = 7). Another one-third of the VRG E neurons had normal or increased levels of activity when the abdominal nerves were active during fictive vomiting (ABD neurons). The one remaining neuron was mainly silent throughout fictive vomiting. ABD neurons were indistinguishable from INT neurons on the basis of their location in the VRG, type of firing pattern (ramp versus step ramp), conduction velocity, or extent of projection in the lumbar cord. However, INTa neurons had a significantly higher discharge rate during respiration than either ABD or INTb neurons. Abdominal muscle EMG and nerve activity were recorded from six unparalyzed cats before and after cutting the axons of VRG E neurons as they cross the midline between C1 and the obex. The lesions abolished or almost eliminated expiratory modulation of abdominal muscle activity. In contrast, the abdominal muscles were always active during vomiting; however, the amplitude of postlesion abdominal activity varied from approximately 70-100% of prelesion values in three cats to 60-70% of normal in a fourth animal to only approximately 20% of prelesion values in two other cats.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1987 the American Physiological Society

Chan, Y. S.; Kasper, J.; Wilson, V. J.

doi: N/Apmid: 2955083

Abstract With the use of floating electrodes we recorded from the C2 dorsal root ganglion of decerebrate cats during sinusoidal and trapezoidal head rotation. Fifty-one spontaneously firing afferents were identified as muscle spindle endings. Some were identified by their excitatory response to injection of succinylcholine, others by the similarity of their behavior to that of endings excited by the drug. Because afferent input to the ganglion was restricted by sectioning most nerve trunks, most spindle endings were presumably located in ventral and ventrolateral perivertebral muscles. The firing of each spindle afferent was modulated most effectively by tilting the head in a specific direction; this direction was termed its response vector. Responses to sine waves and trapezoids were then studied with stimuli oriented as closely as possible to the vertical plane of this vector. Most spindle afferents could be classified in one of two categories. Those with high gain, pronounced nonlinearity, and high dynamic index were called type A. Those classified as type B had low gain, a fairly linear response, and low dynamic index. In response to small (0.5 degrees) stimuli, type A endings had phase leads of approximately 40 degrees at frequencies of sinusoidal stimulation of 0.02-0.1 Hz, increasing to approximately 80 degrees at 4 Hz; with larger (2.5 degrees) stimuli, phase was advanced by an additional 10-20 degrees at all frequencies. Phase of type B responses was less advanced than that of type A responses. Gain slopes of the two types of endings were similar. Bode plots of spindle afferents strongly resembled those of upper cervical neurons whose activity is modulated by head rotation. Each spindle afferent had a response vector whose direction remained stable with time, different frequencies of stimulation, and different stimulus amplitudes. The distribution of response vectors covered approximately 270 degrees, with a gap near nose down pitch. Changing initial head position usually had little effect on the direction of an afferent's response vector or on response dynamics. However, stimulation far from the best plane could transform a type A into a type B response. This raises the possibility that type B receptors could be type A receptors best stimulated by yaw and with only low sensitivity to stimulation in vertical planes. Type A receptors have all the properties of spindle primaries. The identification of type B receptors remains uncertain, because they may include secondary afferents as well as primaries stimulated far from their best three-dimensional vector.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1987 the American Physiological Society

Gulyas, B.; Orban, G. A.; Duysens, J.; Maes, H.

doi: N/Apmid: 3598630

Abstract The suppressive action of a moving textured background on responses to moving bars was investigated in 118 striate neurons, 19 dorsal lateral geniculate neurons, and 5 perigeniculate neurons in paralyzed and anesthetized cats. In standard conditions the background was a two-dimensional (2D) noise pattern, the bar moved at optimal speed, and its contrast level was adjusted to yield 50% of the maximum response. Neuronal responses to the moving bar were suppressed when the background moved at the same speed or faster than the bar. The direction of motion of the bar had little influence. This suppressive effect was equally strong in all three experimental samples. The suppressive effect of the moving background was uniformly distributed among the cortical population, being equally strong in all layers, in all parts of the visual field representation, and for different categories of cortical cells. The suppressive effect of the moving background depended little on the structure of the background or on the speed of the bar. The suppression increased with decreasing contrast of the bar. Many (80%) cortical cells and all geniculate neurons responded to the movement of the 2D noise on its own. Most of these cells responded to isolated features (“grains”) in the pattern rather than to movement of the whole pattern. There was no difference in strength of suppression between cortical neurons responsive and unresponsive to the moving 2D noise. The possible origins of this suppressive influence of moving backgrounds and its significance for the processing of visual scenes, more complicated than a single stimulus, are discussed. Copyright © 1987 the American Physiological Society

Johnson, R. D.; Kitchell, R. L.

doi: N/Apmid: 3598632

Abstract In the dog, we isolated 126 mechanoreceptive afferent fibers of the A-delta myelinated fiber class from the dorsal nerve of the penis using microdissection and extracellular electrophysiological techniques. Receptive fields on the glans penis were stimulated with a computer-controlled mechanostimulator and Peltier effect thermostimulator. The great majority of units were categorized as either rapidly adapting (RA) or slowly adapting (SA) and were located primarily proximally and distally, respectively, on the glans. In comparison to values from other glabrous skin regions in other species, mean displacement and force thresholds of penile mechanoreceptors were high, whereas the mean velocity thresholds were low. SA units, generally poor encoders of static displacement, were distinguishable into two types based on static response firing pattern but were not homologous to either the SA I or SA II mechanoreceptors found in other skin regions. Fifty-five units were given simultaneous mechanical and thermal stimulation. Very few units responded to pure thermal stimulation or increased their discharge frequency to skin cooling. Warm receptive-field temperatures between 35 and 43 degrees C increased mechanical sensitivity, measured by displacement and velocity coding functions, in almost all units tested. We conclude that canine penile mechanoreceptors, capable of encoding a variety of skin movements when the penis is warm, provide the spinal cord with the sensory input necessary to drive the spinal sexual reflexes. However, many appear to be at least physiologically different from mechanoreceptors native to other skin areas. Copyright © 1987 the American Physiological Society

Young, A. A.; Dawson, N. J.

doi: N/Apmid: 3598636

Abstract Thermal clamping of deep-body temperature and 16 fields covering the total truncal skin surface enabled characterization of thermal transmission neurons distributed in a midline medullary location. The total data set comprised 136 neurons from 54 female rats. Relative abundance of neuronal types was 27 to 34 to 75 for cold-responsive, warm-responsive, and thermally unresponsive neurons. Response maxima of thermoresponsive neurons to static thermal stimulation of the total truncal surface were 55 +/- 4 ips (mean +/- SE) at 5 degrees C for cold-responsive neurons and 6.0 +/- 1.6 ips at 35 degrees C for warm-responsive neurons. Dynamic thermal stimulation of the total truncal surface at rates up to +/- 1.6 degrees C/s failed to reveal a clear dynamic thermosensitivity in either cold- or warm-responsive neuronal pools. Instead, the data suggest a preferential passing of the static response relative to the dynamic response. Cutaneous thermal receptive fields were diffuse, occupying most of the truncal surface. Subparts of these fields drove thermoresponsive neurons to variable extents, suggesting convergence from unequally represented multiple cutaneous sources. Noxious stimulation at widely distributed body sites consistently augmented activity in cold-responsive neurons. A thermoregulatory rather than somatesthetic role is proposed for the midline medullary neurons studied here. Copyright © 1987 the American Physiological Society