Impulse patterns of Ia afferents and Ia-activated DSCT neurons during sinusoidal muscle stretch in cats

Summary The cat gastrocnemius muscles of one hind leg were stretched sinusoidally with amplitudes between 10 m and 2.5 mm and frequencies between 1 and 30 Hz. The stretch response of deefferented muscle spindle afferents and of Ia-activated cells within Clarke's column were investigated by means of extra-cellular recordings of action potentials. The responses to 20–50 cycles were displayed in impulse patterns (raster diagrams) of the responding action potentials. The impulse patterns of Ia afferents exhibited a high degree of phase-locking (regularity) on the stretch cycle of amplitudes of about 50 m at 3 Hz and all higher amplitudes or frequencies. At comparable stretch parameters the regularity in Ia afferents was 4–6 times larger than in Ia-activated DSCT neurons. The regularity in the DSCT patterns increased with an increase in stretch frequency. The impulse patterns of DSCT cells exhibited a high negative correlation between successive interspike intervals (–0.4 to –0.6) at low stretch frequencies (<3 Hz), which decreased with an increase in stretch frequency.     

Physiological properties of primary sensory neurons appropriately and inappropriately innervating skeletal muscle in adult rats.

1. We have studied the physiology of primary sensory neurons innervating rat hindlimb muscle in the following: 1) normal control animals; 2) animals in which the gastrocnemius nerve (Gn) had regenerated to its original muscle target; and 3) animals in which the cutaneous sural nerve (Sn) had regenerated to a foreign target, muscle. 2. Single-unit recordings were made from 115 afferents in normal, intact animals. They had conduction velocities of 0.8-67.2 m/s, which were distributed with peaks at approximately 1.25, 17.5, and 47.5 m/s. Of the myelinated fibers, 88% had low-threshold mechanosensitive receptive fields and responded to ramp-and-hold stretches of the muscle. The large majority of these fibers (85%) gave slowly adapting responses to ramp-and-hold stretches or direct muscle probing. Stretch-sensitive afferents could be divided (on the basis of their responses to active muscle contraction) into in-parallel or in-series receptors (presumed muscle spindles and Golgi tendon organs, respectively). The in-parallel receptors outnumbered the in-series receptors by approximately 3:2. The 12% of fibers that were insensitive to stretches of the muscle in the physiological range could be divided into roughly equal groups of totally insensitive fibers and high-threshold fibers, which required excessive stretching of the muscle. 3. In the animals with regrown Gn, 94 single fibers with conduction velocities ranging from 11 to 60.6 m/s were studied. The myelinated conduction velocity distribution exhibited only one peak, at approximately 37.5 m/s. Only 67% of the afferents were stretch sensitive (vs. 88% in normal animals), and only about two-thirds of these (vs. 85% in normal animals) gave slowly adapting responses to ramp-and-hold stretches or muscle probing. The incidence of in-series receptors was also increased among regenerated gastrocnemius afferents. The 33% of fibers that were stretch insensitive were mostly unresponsive to even extreme forms of mechanical stimuli. This group presumably represents afferents that failed to make appropriate endings. 4. In the animals with Sn directed to muscle, 460 single afferents were recorded. Their conduction velocities ranged from 0.7 to 67.9 m/s, and the distribution exhibited only a single peak for myelinated fibers at approximately 22.5 m/s, significantly lower than for intact or regrown Gn. Only 41% of the myelinated fibers were stretch sensitive. Nearly all of these (98%) were rapidly adapting to ramp-and-hold stretches or muscle probing, in marked contrast to the other groups. Also, unlike other groups, nearly all stretch-sensitive afferents appeared to be in-series.(ABSTRACT TRUNCATED AT 400 WORDS)     

Responses of cortical neurons (areas 3a and 4) to ramp stretch of hindlimb muscles in the baboon.

1. A study was made of the response of single cortical units in areas 3a and 4 to electrical stimulation of hindlimb muscle nerves and to ramp stretch of hindlimb muscles in baboons anesthetized with chloralose.2. Stimulation of hindlimb muscle nerves revealed a group I projection primarily to area 3a but with some input into adjacent area. 4. A major group II projection was found in area 4 adjacent to area 3a. A small number of area 3a neurons receive convergence from both group I and group II muscle afferents.3a. On the basis of their response pattern to ramp stretch, units were classified into one of six categories and their cytoarchitectonic location was determined. Units in area 3a had hynamic sensitivities equivalent to that of the primary spindle afferents. Although the discharge of some area 3a neurons also reflected differences in muscle length, most area 3a neurons had low position sensitivities. One unit type in area 3a did not respond to maintained muscle stretch and signaled only velocity of stretch.4. Units in area 4 had position sensitivities equivalent to that of primary and secondary spindle afferents. Although the discharge of some area 4 units reflected different velocities of muscle stretch, these units had dynamic sensitivities similar to those of secondary spindle afferents rather than those of primary afferents. One type of unit in area 4 had no dynamic component to muscle stretch and signaled only muscle length.5. The results demonstrate that there is a transfer of dynamic and position sensitivity from spindle afferents to cortical neurons. Furthermore, data processing has occurred because some units respond only to the steady-state length of muscle, while other units encode only the dynamic phase of stretch. This behavior is different from the responses to ramp stretch of either group I or group II muscle afferents in the baboon.6. The results demonstrate that single units in cerebral cortex can encode the information transmitted to the central nervous system by muscle spindle afferents. The purpose for which this information is used remains undetermined.     

Angular tuning and velocity sensitivity in different neuron classes within layer 4 of rat barrel cortex

Local circuitry within layer IV whisker-related barrels is preferentially sensitive to thalamic population firing synchrony, and neurons respond most vigorously to stimuli, such as high-velocity whisker deflections, that evoke it. Field potential recordings suggest that thalamic barreloid neurons having similar angular preferences fire synchronously. To examine whether angular tuning of cortical neurons might also be affected by thalamic firing synchrony, we characterized responses of layer IV units to whisker deflections that varied in angular direction and velocity. Barrel regular-spike units (RSUs) became more tuned for deflection angle with slower whisker movements. Deflection amplitude had no affect. Barrel fast-spike units (FSUs) were poorly tuned for deflection angle, and their responses remained constant with different deflection velocity. The dependence of angular tuning on deflection velocity among barrel RSUs appears to reflect the same underlying response dynamics that determine their velocity sensitivity and receptive field focus. Unexpectedly, septal RSUs and FSUs are largely similar to their barrel counterparts despite available evidence suggesting that they receive different afferent inputs and are embedded within different local circuits.     

Response of primate joint afferent neurons to mechanical stimulation of knee joint.

1. One hundred thirty-eight knee joint afferents from posterior articular nerve (PAN), in primates, were recorded in dorsal root filaments. Responses of afferents were studied in relation to both passive manipulations of the knee and active contractions of quadriceps, semimembranosus, and gastrocnemius muscles. 2. When the knee was passively rotated, most neurons discharged only when extreme angular displacements were achieved. Response of neurons responding to passive extensions was linearly related to the torque applied to the knee. With maintained extensions, discharge in extension neurons adapted slowly. Some of the time constants of adaptation were similar to those for simultaneously recorded torque relaxation. 3. Contractions of quadriceps, semimembranosus, or gastrocnemius muscles could activate many neurons in the absence of changes in joint angle. For quadriceps-activated neurons, rather high torques (mean = 2,450 g with cm) were required. 4. The results support the hypothesis that joint afferents function as capsullar stretch receptors, responding to those mechanical events which result in loading of the capsule.     

Responses of cat dorsal spino-cerebellar tract neurons to sinusoidal stretching of the gastrocnemius muscle

A defined class of cells within the nucleus dorsalis (Clarke's column) receives excitatory input from Ia afferents of mainly one muscle. Action potentials were recorded from axons of these cells (DSCT neurons) which are excited by Ia afferents of the gastrocnemius muscles. We investigated the response to sinusoidal muscle stretch over a wide range of amplitudes (10 m–4 mm) and frequencies (0.1–130 Hz) in the deefferented preparation. The dynamic stretch was superimposed on a moderate static muscle stretch to ensure that the muscle was not slack during the phase of release. The response up to 10 Hz was displayed as PST histograms (cycle histograms) and a sinewave of stretch frequency was fitted to the PST histograms to define amplitude and phase of a response sinewave.At a constant frequency of about 3Hz, the relation between stretch amplitude and response amplitude could well be described by decelerating intensity functions: the hyperbolic or tanh log function and a modified power function (exponent 0.48±0.12). The phase lead of the response sinewave increased with increasing stretch amplitudes of up to 0.5 mm and then decreased.At constant stretch amplitudes of 0.5–2.0 mm the frequency response was investigated. In relation to stretch frequencies between 0.1 and 1 Hz an increase in the response amplitude of 4.4dB was observed and an increase for 13dB/decade between 3 and 10 Hz. At 0.1 Hz the phase of the response sinewave was 48° in advance and increased to a maximum lead of 89° at 6–8Hz. Above 10Hz the positions of the responding action potentials with respect to the stretch cycle were used to define a phase, which was in advance up to 60 Hz but decreased and changed to a phase lag at higher frequencies.If in PST histograms no periods of silence occurred during the phase of stretch release, the mean discharge rate was found to be independent of the sinusoidal stretching. If the pauses were present the mean rate increased with increasing stretch frequencies or amplitudes.     

Responses of cat peroneus brevis muscle spindle afferents during recovery from nerve-crush injury

The responses of regenerated muscle spindle afferents to ramp-and-hold stretch of the peroneus brevis muscle in the cat were recorded at periods from 26 to 140 days after crushing the common peroneal nerve. During the early stages of recovery a number of abnormally responding afferents were observed. The most marked abnormality was the absence or rapid failure of firing during the held phase of the stretch. The proportion of abnormal afferents became less as recovery progressed. Electrical stimulation of isolated static and dynamic gamma-axons increased the firing rates of the afferents during the ramp-and-hold stretch such that a gamma static axon would restore the response of an abnormal afferent to the held phase of the stretch. The regenerated afferents have been classified according to the degree of abnormality displayed. These abnormalities can be accounted for by assuming a subtractive reduction in the firing frequency of the regenerated afferents. This is attributed to an increase in the pacemaker threshold.     

Response properties of vibrissa units in rat SI somatosensory neocortex

1. Glass microelectrodes were used to record extracellular responses from 308 SI cortical neurons to deflections of the contralateral vibrissae in 21 unanesthetized, paralyzed rats. Controlled deflections of individual hairs were produced by means of an electromechanical stimulator. Fast green dye marks were made to aid histological reconstructions of electrode tracks. 2. TS) were observed throughout layers II--VI; "fast" cortical spikes (FS) were less frequently encountered and largely restricted to layer IV. Although both types of potentials had similar negative-positive waveforms. FSs were distinguished from RSs by their comparatively rapid time course, about half that of RSs. RS units (RSU) discharged spontaneously at rates of less than 1--15/s, whereas FS units (FSU) displayed rates of 15--50/s. The amplitudes of FSs, which were generally smaller than those of RSs, often decreased during high-frequency discharges. 3. With sinusoidal oscillations of a vibrissa FSUs responded more reliably and over a broader range of frequencies (3 to at least 40 Hz) than did RSUs , particularly in layer IV. In addition, FSUs typically responded to whisker deflections over a range of 360 degrees, whereas many RSUs in layer IV displayed sharp spatial-tuning charcteristics, responding over a restricted range of less than 90 degrees. 4. Of all units, 58% responded preferentially to stimulus transients (vibrissal movements), 32% displayed sustained responses to stimulus stead-states (fixed vibrissal displacement). For the remaining 10% of units the appropriate stimulus could not be specified; these units were particularly common in layer V. 5. Computation of quantitative stimulus-response relations showed that many units increased their rate of discharge with increasing stimulus intensities. 6. Most units were directionally selective, responding preferentially to deflections of a whisker in one or more of four quadrants. 7. In radial penetrations through the cortex there was a columnar patern so that units were activated at least by the same (i.e., the "principal") whisker. In a number of cases these could be directly correlated with the barrels. Of all units, 55% responded to deflections of single vibrissa only, the remaining 45% to 2--12 adjacently situated vibrissae. For most multiple-whiser units the responses elicited by deflection of any one hair activating the neuron was qualitatively similar to those elicited by deflection of any other hair activating it. The principal whisker of the penetration was typically associated with the most vigorous responses. 8. In layer IV, 85% of neurons responded to deflections of one hair only, the remaining 15% to two or more hairs. In layers II and III 39% of units were activated by more than one vibrissa; in layers V and VI multiple-whisker units predominated (64%). Whisker configurations in the deep layers were larger than those in other layers. 9...     

Physiological evidence for caudal brainstem projections of jaw muscle spindle afferents

Retrograde transport and intra-axonal labeling studies provide convincing evidence that jaw-muscle spindle afferents project to the caudal medulla by way of Probst’s tract. However, functional properties of this caudal projection are not well understood. Extracellular recordings were made in cats at the level of the subnucleus interpolaris (Vi) to identify single units that showed consistent responses to ramp-and-hold stretches of the jaw. In this report, we present data from 20 central units with properties indicating that they received input from trigeminal muscle spindle afferents. All units were activated by gentle palpation of jaw muscles, and none had superficial receptive fields. Two groups of neurons could be defined based on their responses to passive jaw movements. One group (n=12) showed an obvious dynamic response (i.e., a higher level of activity at the onset of stretch than during the hold period). Activity was maintained during the hold phase, and the units stopped firing (unloaded) for a brief period upon jaw closure. The other group (n=8) lacked a dynamic response. Instead, they showed an increase in firing with onset of stretch that was maintained during the hold phase. Thirteen units, which were tested with more than three different jaw stretch speeds and/or amplitudes, were further characterized by analyzing dynamic index (DI) and mean firing rate (MFR) during each phase of the ramp-and-hold movement as well as interspike interval (ISI) variability. All but one unit with a dynamic response showed a speed-sensitivity. In all cases, the MFR was a more sensitive indicator of changes in jaw speed than DI. Neurons in the other group (5/5 tested) showed a high position-sensitivity, i.e., their firing rates varied as a function of amplitude of jaw opening. The percent change in ISI variability for all neurons ranged from 37–84%. The response characteristics of these central neurons were compared to known physiological properties of muscle spindle afferents. The results provided compelling evidence for jaw-muscle-spindle afferent projection onto these neurons. Reconstruction of recording sites showed that medial Vi, and the adjacent reticular formation, are likely recipients for the caudal projections from jaw-muscle-spindle afferents. We suggest that muscle spindle input to this region is well suited for influencing the coordination of motor behavior during feeding and for the integration and processing of kinesthetic information. Received: 22 December 1998 / Accepted: 7 May 1999     

Tendon organs of cat medial gastrocnemius: responses to active and passive forces as a function of muscle length.

The responses of 13 Golgi tendon organs to graded force development of 29 motor units in medial gastrocnemius of the cat have been studied in five experiments. Of the 13 tendon organs, 11 were responsive to passive stretch within the physiological range of muscle length and 5 were "spontaneously" active at very short lengths where no passive tension could be recorded. The relationship between passive force and the firing rates of the various afferents ranged from a linear one to a power relation (Y = Axb + c) with b, a widely varying exponent. Results support the general conclusion that although many Ib afferents respond to passive force within the physiological range of muscle stretch, this form of stimulus is not a particularly effective one. The statis responses of Golgi tendon organs to active force development produced by single motor units was studied at different muscle lengths. In all cases the apparent sensitivity (change in firing rate per active force developed) decreased as muscle length approached Lo. The static responses of Golgi tendon organs to force developed by single motor units were also studied during fatiguing contractions. The data suggest a sigmoid relationship between force developed at the tendon and the Ib response. The collective response of all 13 tendon organs to active and passive forces at different muscle lengths was also examined. This analysis offered further support for the viewpoint that active motor unit contractions provide themost significant excitatory input to tendon organs and that changes in passive force during muscle stretch have comparatively little effect on the collective tendon organ response. The interaction between active and passive force inputs to the Golgi tendon organs is discussed in relation to the complicated nature of the relationship between forces measured at the tendon and those acting within the receptor capsule. When these complications were taken into account it was possible to explain the differences in responsiveness of a given tendon organ to active contraction of several motor units and to passive force in terms of a single force-firing rate curve for the receptor. It is concluded that changes in the force of contraction of single motor units result in relatively small changes in Ib afferent firing and that during normal muscle contractions, changes in the number of motor units acting on a single receptor must produce far more significant changes in firing rate than changes in the amount of force developed by any single unit. Changes in dynamic Ib sensitivity to single motor unit contractions are also shown to depend on length and in a similar way to the changes in static Ib sensitivity. During fatiguing contractions, a sigmoid relation was found between the dynamic Ib response and the rate of force development by single motor units.     

Projections of extraocular, neck muscle, and retinal afferents to superior colliculus in the cat: their connections to cells of origin of tectospinal tract.

Unit recordings were made in the superior colliculus of cats anesthetized with chloralose and with Pentothal. Electrical stimulation of extraocular muscle afferents and neck muscle afferents excited more units in the superior colliculus than did a variety of moving and stationary visual stimuli. Units responding to neck muscle afferent stimulation fell into three populations; one population firing with a short latency and following stimulus presentation up to 1/s, a second population with a long latency and following stimulus presentation at frequencies lower than 15/min, and a third population exhibiting paired firing. The latencies and firing patterns of the third population combined the characteristics of each of the first two patterns. It is suggested that these characteristics of unit discharges stem from the existence of two pathways from neck muscle afferents to the superior colliculus. The projection is predominantly bilateral. Units responding to neck muscle afferent stimulation are distributed throughout the superior colliculus on the basis of their latencies. Long-latency responses predominate in the superficial layers of the superior colliculus and short-latency responses, while more common in the intermediate and deep layers, predominate in the tegmentum. Extraocular muscle afferent projections to the superior colliculus constitute the single richest projection found in these experiments. While the response patterns and latencies are similar to those of the neck muscle afferents, long-latency responses are the most common and dominate in all collicular regions. Few units in the tegmentum could be excited by extraocular muscle afferents. Both extraocular muscle and neck muscle afferents show considerable convergence with one another and with retinal afferents within the superior colliculus. Cells of origin of the tectospinal tract were identified within the superior colliculus and tegmentum by antidromic excitation from the upper cervical cord. These cells were distributed predominantly within the intermediate and deep layers of the superior colliculus, and sparsely in the superficial layers and tegmentum. Almost 50% of the cells of origin of the tectospinal tract receive a convergent input from extraocular muscle and neck muscle afferents and from the retina. About 30% of the cells were inexcitable to the stimuli employed in these experiments. The significance of these projections is discussed with respect to superior collicular function in the cat and i     

Responses of muscle spindles following a series of eccentric contractions

To investigate the effects of eccentric exercise on the signalling properties of muscle spindles, experiments were done using the medial gastrocnemius muscle of cats anaesthetised with 40 mg/kg sodium pentobarbitone, i.p. Responses were recorded from single afferent nerve fibres in filaments of dorsal root during slow stretch of the passive muscle and during intrafusal contractions at a range of lengths, before and after a series of eccentric contractions. The sensitivity to slow stretch was measured as the average firing rate between muscle lengths 10.5 and 9.5 mm shorter than the physiological maximum (L_m), during stretch at 1 mm/s over the whole physiological range. The mean sensitivity of both primary and secondary spindle endings increased slightly, but not significantly, after a series of 20–150 eccentric contractions consisting of a 6 mm stretch, at 50 mm/s, to a final length of between L_m –7 mm and L_m, during stimulation of the whole muscle or sometimes of single fusimotor fibres. Discharges were recorded from primary endings during fusimotor stimulation at 100–150 pulses/s, and from secondary endings during static bag intrafusal contractures produced by i.v. injection of 0.2 mg/kg succinyl choline. Spindle responses were recorded, over a range of muscle lengths, in steps covering the whole physiological range. About half of the responses showed a peak in the relation between length and net increase in firing rate, while the remainder either progressively increased or progressively decreased over the physiological range. No large or consistent changes were seen after the eccentric contractions. It is concluded that the intrafusal fibres of muscle spindles are not prone to damage of the kind seen in extrafusal fibres after a series of eccentric contractions.     

Spinal interneurons that receive input from muscle afferents are differentially modulated by dorsolateral descending systems

The possibility that descending systems have differential actions on the spinal interneurons that receive input from muscle afferents was investigated. Prolonged, physiological inputs were generated by stretch of the triceps surae muscles. The resulting firing patterns of 25 lumbosacral interneurons were recorded before and during a reversible cold block of the dorsolateral white matter at the thoracic level in nonparalyzed, decerebrate preparations. The strength of group I muscle afferent input was assessed from the response to sinusoidal tendon vibration, which activated muscle spindle Ia afferents directly and tendon organ Ib afferents via the resulting reflex force. The stretch-evoked responses of interneurons with strong responses to vibration were markedly suppressed by dorsal cold block, whereas the stretch-evoked responses of interneurons with weak vibration input were enhanced. The cells most strongly activated by vibration received their primary input from Ia afferents and all of these cells were inhibited by the cold block. These results suggest that a disruption of the descending system, such as occurs in spinal cord injury, will lead to a suppression of the interneuronal pathways with group Ia input while enhancing excitability within interneuronal pathways transmitting actions from higher threshold afferents. One possible consequence of this suppression would be a decreased activity among the Ia inhibitory interneurons that mediate reciprocal inhibition, resulting in abnormal reciprocal relations between antagonists and promoting anomalous muscle cocontraction.     

Quantitative analysis of static strain sensitivity in human mechanoreceptors from hairy skin.

1. Microelectrode recordings from 15 slowly adapting (SA) cutaneous mechanoreceptor afferents originating in hairy skin were obtained from the radial nerve in humans. 2. Controlled skin stretch was applied to the back of the hand that encompassed the physiological range of skin stretch during movements at the metacarpophalangeal (MCP) joints. 3. Both SA Group I and II afferents showed exquisite dynamic and static sensitivity to skin stretch. The median static strain sensitivity was 1.0 imp.s-1 per percent skin stretch for SAI units and 1.8 for SAII units. 4. Translated into sensitivity to movements at the MCP joint, both SAI and SAII afferents in the skin of the back of the hand displayed a positional sensitivity that was comparable with that reported for muscle spindle afferents. 5. These data give quantitative support to suggestions that skin receptors in the human hairy skin provide information on nearby joint configurations and therefore may play a specific role in proprioception, kinesthesia, and motor control.     

Cortical mapping and laminar analysis of the cutaneous and proprioceptive inputs from the rat foreleg: an extra- and intra-cellular study

Summary The foreleg proprioceptive and cutaneous representations, in the Sm cortex of urethane-anesthetized rats was studied. Natural or electrical stimulations and stretches of single forearm muscles were used. Multiunitary, unitary or intra-cellular recordings were performed in the contra-lateral Sm cortex. The aims of the study were: 1 — to compare the proprioceptive and cutaneous maps 2 — to analyse the characteristics of the unitary responses and 3 — to study the laminar distribution of cutaneous and muscular inputs. It is shown that: 1 — the proprioceptive and cutaneous representations overlapped, except in the anterior part where only proprioceptive (mainly articular) responses were obtained. The representation of each stretched muscle extended over the whole cutaneous area, showing a total overlap between inputs from these muscles. 2–46% of the intracellularly recorded cells (n=215) responded to peripheral stimulation, and 30.7% were influenced by (at least) muscle stretch. The majority of excited cells showed cross-modal covergence, and among neurons responding to muscle stretch, 60% received inputs from the two muscles stretched. Two categories of EPSPs were found, and four neurons responded to cutaneous or muscular stimulation with a burst. 19% of the responding cells were inhibited by peripheral — mainly cutaneous — stimulations. 3 —Excited neurons were recorded in all layers, with just over half located in layer IV, whereas IPSPs were obtained mainly in layer V. The cells excited by cutaneous and muscular inputs (convergent neurons) were preponderant in layers IV to VI. This work shows that the cutaneous and muscular inputs reach the same area in Sm cortex, and that a majority of excited cells are convergent. The results are not in favor of an area 3a (by analogy with cats and monkeys) in the rat.     

The ‘initial burst’ of human primary muscle spindle afferents has at least two components

Ten muscle spindle primary afferents from the extensor digitorum communis muscle of man were studied with single unit afferent recordings. Responses to slow test stretches with three different pre-history conditions were assessed to investigate the contribution of rapid stretches to the stretch sensitization phenomenon. In two of the conditions, the slow test ramps were preceded by rapid stretch after which the parent muscle of the receptor was either (a) kept short for 5 seconds or (b) kept long for 3.2 seconds and then returned to the short muscle length for 5 seconds. The third condition (c) consisted of a slow stretch from short to long muscle length followed by a rapid return to the short muscle length, in turn followed by 5 seconds at the short muscle length. Afferent responses were depressed when the muscle had been kept at the long length after the rapid stretches (condition b) and enhanced when the muscle had been kept at the short length (conditions a & c). A prominent ‘initial burst’ was only present in the afferent discharge when the parent muscles of the primary endings had been kept short (condition a). A second, more prolonged burst was present for conditions (a) and (c) but was lacking or inconspicuous when the muscle had been kept long after rapid stretches (condition b). The rapid stretches in the stretch sensitization paradigm appear to be a primary factor not only for the enhanced responses of sensitized primary afferents but also for the depressed responses of desensitized primary afferents.     

Effects of changes in preoptic temperature on stretch response of muscle spindle endings in the cat's soleus muscle

1. The effects of changing the temperature in the preoptic region on the stretch responses of primary and secondary endings of the muscle spindle in the soleus muscle was investigated in urethane anesthetized cats. The local temperature of the preoptic region was controlled by implanting water perfused thermodes (32–42.5°C).
2. A standard ramp and hold stretch stimulus was repeatedly applied to the soleus muscle during changes of preoptic temperature from normal to hypo-or hyperthermic values. During each stretch the instantaneous firing rate was recorded and the static firing frequency and the dynamic index were electronically determined according to Crowe and Matthews (1964).
3. Of a total of 76 investigated muscle spindle afferents, the stretch responses of 49 (65%) were altered by changing preoptic temperature. The static stretch response was increased in all cases during cooling, whereas during preoptic heating the static stretch response increased in about half of the afferents and decreased in the rest. Increase in the static stretch response during both cooling and heating was usually combined with the appearence of spontaneous discharges at rest.
4. With regard to the static stretch response, primary and secondary muscle spindle endings responded similarly to cooling and heating.
5. Two types of preoptic temperature effects on the stretch responses of primary endings were observed. In the majority of afferents only the static stretch response was augmented during cooling, whereas the peak response was little changed resulting in a decrease of the dynamic index (‘static’ response type). In a minority of primary afferents the effect of preoptic cooling consisted of a greater increase of the peak response than of the static stretch response. i.e., the dynamic index also increased (‘dynamic-static’ response type). In secondary muscle spindle endings only the static type of response to preoptic cooling was observed.
6. It is concluded that changes of preoptic temperature influence the stretch response of muscle spindle afferents mainly by altering the activity of the static fusimotor innervation, its activation by preoptic cooling being the most consistent finding. In a minority of muscle spindles the dynamic fusimotor innervation appears to be additionally activated by preoptic cooling.

     

Vagal representation in the cerebellum of the cat

The cervical vagus nerve (VN) was electrically stimulated in Nembutal-anaesthetized cats. The responses recorded from the cerebellar surface were found in lob. V and VI in a bilateral sagittal strip perpendicular to the longitudinal axis of the folia. At a longer latency, potentials were also found in the paramedian lobule. Field potential analysis confirmed the existence of a sagittal strip in the deep parts of the lobules. The distribution of these potentials and their field potential profiles indicate that they are transmitted through the climbing fiber (CF) system. Experiments with local anaesthetics and deafferentation support our view that the potentials described in the vermis by Dell and Olson (1951) could be originated extracerebellarly. A strong parallelism was found between the amplitude of the cerebellar responses and the amplitude of the group B₁ component of the vagal afferent potentials, having a conduction velocity between 4 and 20 m/s. The pathway by which the vagal afferents reach the inferior olive and the functional significance of these afferents are discussed.     

Responses of cat mesencephalic reticulospinal neurons to stimulation of superior colliculus,pericruciate cortex,and neck muscle afferents

Summary Neurons were recorded extracellularly in the mesencephalic reticular formation outside the interstitial nucleus of Cajal in cerebellectomized cats anesthetized with chloralose. Reticulospinal neurons were identified by antidromic stimulation of the upper cervical segments. Stimulation in the deep layers of the ipsilateral superior colliculus evoked firing in 36% of reticulospinal neurons. For many neurons thresholds for activation were high in the intermediate tectal layers and declined as the electrodes entered the underlying tegmentum. However, low threshold points were found above the deep fiber layer within the superior colliculus for some cells. Stimulation of the contralateral superior colliculus excited 10% of neurons and thresholds for activation were high above the deep fiber layer for all neurons. Stimulation of the ipsilateral and contralateral pericruciate cortex excited 39 and 21% of neurons, respectively. The lowest threshold area was found in the frontal eye fields. Sixteen percent of neurons received excitation from neck muscle afferents (C₂ biventer-cervicis) bilaterally. Comparison of responses between mesencephalic reticulospinal neurons and interstitiospinal neurons (Fukushima et al. 1981) showed that responses of the two groups of neurons were similar when the pericruciate cortex and neck muscle afferents were stimulated. However, a difference was observed in tectal responses, since low threshold points were rarely observed above the deep fiber layer for interstitiospinal neurons.Supported in part by a Grant-in-Aid for Scientific Research (No. 477063) from The Ministry of Education, Science, and Culture of Japan     

Proprioceptive afferents survive in the masseter muscle of trkC knockout mice

Peripheral innervation patterns of proprioceptive afferents from dorsal root ganglia and the mesencephalic trigeminal nucleus were assessed in trkC-deficient mice using immunohistochemistry for protein gene product 9.5 and parvalbumin. In trkC knockout mice, spinal proprioceptive afferents were completely absent in the limb skeletal muscles, M. biceps femoris and M. gastrocnemius, as previously reported. In these same animals, however, proprioceptive afferents from mesencephalic trigeminal nucleus innervated masseter muscles and formed primary endings of muscle spindles. Three wild-type mice averaged 35.7 spindle profiles (range: 31–41), six heterozygotes averaged 32.3 spindles (range: 27–41), and four homozygotes averaged 32.8 spindles (range: 26–42). Parvalbumin and Nissl staining of the brain stem showed approximately 50% surviving mesencephalic trigeminal sensory neurons in trkC-deficient mice. TrkC −/− mice (n=5) had 309.4±15.9 mesencephalic trigeminal sensory cells versus 616.5±26.3 the sensory cells in trkC +/+ mice (n=4).These data indicate that while mesencephalic trigeminal sensory neurons are significantly reduced in number by trkC deletion, they are not completely absent. Furthermore, unlike their spinal counterparts, trigeminal proprioceptive afferents survive and give rise to stretch receptor complexes in masseter muscles of trkC knockout mice. This indicates that spinal and mesencephalic trigeminal proprioceptive afferents have different neurotrophin-supporting system during survival and differentiation. It is likely that one or more other neurotrophin receptors expressed in mesencephalic trigeminal proprioceptive neurons of trkC knockout mice compensate for the lack of normal neurotrophin-3 signaling through trkC.