Structures in sensory region of snake spindles and their displacment during stretch.

1. Structures within the sensory region of short- and long-capsule snake muscle spindles have been visualized using differential interference contrast microscopy. Profiles seen with Nomarski microscopy have been identified by electron microscopy of the same preparations. 2. Sensory nerve terminals, nuclei and other cytoplasmic inclusions in the intrafusal fiber, collagen bands, and capsular cells may be seen in the living preparation. 3. The length changes of various elements within the sensory region in response to stretch of the spindle have been measured using high-speed ciné photomicrography. This has been corrleated with the impulse response from sensory endings in short-and long-capsule spindles. 4. Short-capsule spindles, which have a high dynamic sensitivity, show length changes in the sensory region in response to ramp-and-hold stretch, which are not parallel to the changes in impulse frequency. The implications for mechanical models of spindle behavior are discussed.     

Changes in motor commands, as shown by changes in perceived heaviness, during partial curarization and peripheral anaesthesia in man

1. The responses to stretch have been studied in living, isolated Golgi tendon organs (GTOs) from tail muscles of cat. Experiments were performed in vitro and consisted of subjecting single GTOs to controlled ramp-and-hold stretch while recording the response from their sensory axons raised in oil. 2. The threshold force required for sustained afferent discharge was measured directly, and found to be between 8 and 170 dynes at 24 degrees C for nine GTOs tested. Beyond threshold, the discharge frequency is approximately proportional to applied static tension over a wide range. Sensitivy to tension varies among different GTOs and appears to be inversely correlated with mechanical stiffness. 3. With impulse activity blocked by tetrodotoxin, graded receptor potentials could be recorded whose amplitude varied in proportion to applied static tension. All GTOs examined showed in addition a dynamic response, which became larger with increasing velocity of ramp stretch. This dynamic sensitivity appears in the receptor potential and is then augmentd by an apparent accommodative process at the impulse initiating site. 4. Based on the above findings, possible mechanical models are discussed for the sensory transduction mechanism.     

Facilitation effect of auxiliary noise stimuli on response of isolated frog muscle spindle to sinusoidal movements

Receptor potentials and impulse patterns were recorded from isolated frog muscle spindles using sinusoidal and superimposed random stretches as stimuli with different sinus-to-noise ratios. The entire dynamic amplitude range of the spindle receptor was evaluated by measuring the sensory response at different levels of static stretch. Auxiliary random stimuli provoked rectified fast depolarizing receptor potential transients; their amplitude and slope grew larger with increasing intensity of the noise stimulus and with increasing prestretch level. Due to this strongly nonlinear behavior of the transducing site the frequency and size of the receptor potentials evoked by the auxiliary input signal increased during the stretching phase of the sinusoidal movement. Since the fast depolarizing receptor potential transients provided a powerful trigger for the action potential encoding site, auxiliary random stimuli effectively enhanced the afferent discharge rate, especially during the stretching phase of the sinusoidal movement. Auxiliary noise stimuli could even activate the afferent discharge to an otherwise subthreshold sinusoidal stretch. It is assumed that by the same mechanism the transfer characteristic of the receptor is broadened towards higher frequencies. Since auxiliary random stimuli increased the nonlinear properties of all receptor response components, a "linearizing" approximation technique only partially describes the receptor's transfer properties. The facilitation effect recorded in the differentiated muscle spindle when random stimuli were superimposed on sinusoidal displacements closely resembled the excitation of afferent firing when passive stretching interacted with active fusimotor innervation. A hypothesis is proposed to explain both effects by the same mechanism acting upon the transducing sensory endings: Since passive random stretches as well as active twitching of the intrafusal muscle fibers exhibited almost the same range of frequency components, we propose that both stimuli also generate the same kind of receptor potentials; namely, those fast-rising depolarization transients of the receptor potential, which vigorously drive the encoding site. In general, these experiments explain how the specific response of a neuron can be facilitated by an additional unspecific (noisy) input.     

Responses of primary and secondary endings of isolated mammalian muscle spindles to sinusoidal length changes.

1. Responses of primary and secondary endings of isolated cat spindles to sinusoidal length changes have been recorded before and after block of impulse activity by tetrodotoxin. 2. Primary endings may discharge with each cycle of sinusoidal stretch at 25-50 Hz, with stretch amplitudes applied to the spindle poles as small as 1 micron. Thresholds are higher at lower frequencies. 3. In primary endings, amplitude of the receptor potential varies with frequency and magnitude of sinusoidal stretch. At a given stretch amplitude, the receptor-potential response increases markedly between 1 and 10 Hz. At a fixed frequency, for example, at Hz, the response to graded amplitude of sinusoidal stretch is highly nonlinear, sensitivity decreasing with large amplitudes. 4. Secondary endings show a much higher threshold than primary endings to sinusoidal stretch. Thus, at 25 Hz, secondary endings required stretch amplitudes of 50-100 micron to evoke discharge. Relatively large amplitudes of stretch were also required to evoked detectable receptor potentials. Over the range studied, the receptor potential varied more linearly with stretch amplitude in secondary than in primary endings.     

Impulse activity and receptor potential of primary and secondary endings of isolated mammalian muscle spindles.

1. An isolated muscle spindle preparation from a tail muscle of cat is described. The afferent response to a ramp-and-hold stretch was recorded in individual axons from identified primary and secondary endings. 2. Primary endings exhibit a prominent dynamic response, including an initial burst. They also show a well-maintained static discharge. Secondary endings also show a well-sustained static discharge but generally have a much lower dynamic sensitivity. The response of primary and secondary endings of the isolated spindle are similar to the typical responses seen in vivo in groups Ia or group II afferent fibres respectively. 3. Following impulse blockade by tetrodotoxin, the receptor potential was recorded from primary and from secondary endings in response to ramp-and-hold stretch. 4. During the dynamic phase the receptor potential of primary endings consists of a depolarization which has two components. (a) An initial component occurs early during ramp stretch, depends in rate of rise and amplitude on velocity of stretch and is reduced on repetitive stretch; it appears to be responsible for the initial burst. (b) A late dynamic component, which follows, is also dependent on stretch velocity and produces the late dynamic discharge. At the end of ramp stretch the receptor potential falls, and may undershoot, the static level. There is a subsequent adaptive fall during hold stretch, then a maintained static level of receptor potential. On release from stretch the membrane is hyperpolarized. 5. Secondary endings usually show a smaller dynamic response, lacking the initial component seen in primary endings. They also generally lack an undershoot following the ramp and have less of a post-release hyperpolarization. 6. Static levels of receptor potential in both primary and secondary endings are related to amplitude of stretch. 7. The receptor potentials of primary and secondary endings account for the major features of the impulse responses of these endings to ramp-and-hold stretch. In primary endings the dynamic frequencies may also depend upon a sensitivity of the impulse initiating site to rate of change of receptor current.     

Adaptation of the discharge of frog muscle spindles following a stretch

1. Stretching a frog muscle spindle evoked a discharge of action potentials in its sensory axon. As the rate of this discharge decreased during the adaptation that followed the dynamic phase of a stretch, the variability of the interspike intervals of the impulse train increased.2. Adaptation occurred in two phases. At first the impulse train was almost regular and adapted rapidly, but later this gave way to a phase of slower adaptation where the variability of the discharge was much increased. In the second phase of adaptation the interspike intervals increased in length less than half as quickly as in the first phase.3. When the rate of adaptation changed from the more rapid to the slower phase there was often an abrupt change in the character of the discharge and the relationship between the mean interspike interval and the variability changed. The interspike interval at which this change-over occurred was relatively constant in records of the discharge from one afferent fibre even though stretches of different amplitude were employed, though it differed from one afferent fibre to another.4. These features of the discharge during adaptation suggest that the two sections of the impulse trains were derived from different spike generators by a process of probabilistic mixing.     

Responses of isolated Golgi tendon organs of cat to sinusoidal stretch

1. Receptor potential and tension have been recorded from isolated Golgi tendon organs in response to sinusoidal stretch. Responses depended on amplitude and frequency of stretch and on the initial (resting) tension of the preparation. 2. Both tension and receptor potential behaved as power functions of stretch amplitude over most of the range corresponding to physiological tendon strains. However, for very small stretch amplitudes (less than 8 microns), a more linear response was seen. Those characteristics of responses that depended on stretch amplitude behaved similarly at all frequencies examined. 3. Frequency dependence of tension was slight. Its character, a gradual monotonic increase in response with increasing stretch frequency and a constant phase lead of a few degrees, did not change over the examined frequency range from 0.12 to 80 Hz. In contrast, receptor potential displayed a marked frequency dependence, increasing rapidly with increasing frequency of stretch in the range from approximately 1 to 20 Hz, then slowly declining as frequency was further increased. 4. Changes in initial tension of the preparation produced marked parallel changes in the amplitude dependence of tension and receptor potential. Frequency response was not significantly affected. 5. By comparing tension and receptor potential responses, the relative contributions of mechanical and electrical properties of the receptor to the sensory transduction process was examined. The present results suggest that in tendon organs the observed nonlinear dependence on amplitude of stretch originates primarily in the mechanical stage of transduction. Dynamic sensitivity, however, seems largely attributable to ionic processes within the sensory terminal membranes.     

Initial burst of primary endings of isolated mammalian muscle spindles.

The initial burst has been studied in primary endings of isolated mammalian muscle spindles subject to controlled ramp-and-hold stretch. Near the onset of ramp stretch the primary ending discharges at a frequency dependent on stretch velocity. The initial burst is reduced or abolished by repetitive stretch. After block of impulse activity by tetrodotoxin, the receptor potential of primary endings shows an initial component, a rapid depolarization which occurs near the onset of ramp stretch at the same time as the initial burst. This initial component depends, in rate of rise and amplitude, on stretch velocity. It is also reduced or abolished by repetitive stretch. Recording of tension development by the isolated spindle in response to ramp-and-hold stretch shows an early rise in tension associated with the initial burst and the initial component of the receptor potential. This tension rise is also dependent on stretch velocity and is reduced or abolished by repetitive stretch. The results provide direct evidence that the initial burst results from mechanical factors, probably from cross bridge formation between thick and thin filaments as has been suggested (3).     

Antidromic potential spread modulates the receptor responses in the stretch receptor neurons of the crayfish

The effects of antidromic potential spread were investigated in the stretch receptor neurons of the crayfish. Current and potential responses to conductance changes were recorded in the dynamic clamp condition and compared to those obtained by using some conventional clamp methods and a compartmental neuron model. An analogue circuit was used for dynamic calculation of the injected receptor current as a function of the membrane potential and the given conductance change. Alternatively, receptor current responses to a mechanical stimulus were recorded and compared when the cell was voltage clamped to a previously recorded impulse wave form and the resting potential, respectively. Under dynamic clamp, the receptor current had an oscillating waveform which contrasts with the conventional recordings. Frequency, amplitude and sign of the oscillations were dependent on the applied conductance level, reversal potential and electrotonic attenuation. Mean current amplitude and frequency of the evoked impulse responses were smaller under dynamic clamp, especially for large conductance increases. However, firing frequency was larger if plotted against the mean current response. Recorded responses were similar to those calculated in the model. It was not possible to evoke any adaptation in the slowly adapting neuron by using the dynamic clamp. Evoked potential change served as a self limiting response, preventing the depolarization block. However, impulse duration was significantly shorter in the rapidly adapting neuron when the dynamic clamp was used. It was concluded that, in the stretch receptor neurons during a conductance increase, antidromic potential spread modulates the receptor responses and contributes to adaptation.     

Receptor potentials of isolated frog muscle spindle evoked by sinusoidal stimulation

Receptor potentials in response to sinusoidal stimulation have been recorded from isolated muscle spindles of the frog. Sinusoidal displacements of different amplitudes (20-120 micron) and frequencies (0.1-100 Hz) were used. The mean static stretch level was adjusted between resting length (L0) and L0 + 400 micron, so that the amplitude and phase-response characteristics were measured at different operating points. Depending on the amount of static prestretch, there is a well-defined dynamic range, which limits the receptor potential by nonlinear compression of either its positive or negative half-cycle. For each point on the static operating curve there exists a dynamic operating curve with a sigmoidal shape. The range of each dynamic curve is approximately 80 micron, independent of the static displacement, and the maxima of all dynamic curves are the same. Therefore the dynamic curves are not symmetrical about their static operating point. The slope of the steepest portion is 10% of the maximum elicitable receptor potential per 10-micron dynamic displacement. For stimulus frequencies greater than 2 Hz the receptor potential deviates from a sinusoidal waveform, exhibiting a fast depolarization transient during stretch and a prolonged repolarization transient during release of stretch. The steepness of the depolarization transient increases with increasing stimulus frequency, amplitude, and prestretch level. As a result, the interval from trough to peak of the receptor potential shortens to less than 90 degrees instead of half a cycle. The repolarization transient has an exponential decay with a time constant of approximately 40 ms that remains constant during the various stimulus conditions. As a result of this slow decay time, individual receptor potentials summate, so that the response divides into a modulated receptor potential (AC component) and a maintained depolarization (DC component). The amplitude response characteristic of the stationary AC component increases with increasing stimulus frequencies up to a peak at 2 Hz, after which it declines with a slope of -3 dB/octave. Provided large sinusoidal stretches and/or extended prestretch levels are used, this high-frequency decline of the AC component is compensated for by the proportional increase of the DC component, so that the peak depolarization values remain constant from 2 to 100 Hz. Stimulus and response are in phase for stimulus frequencies less than 2 Hz and reverse to phase lag at higher stimulus frequencies.(ABSTRACT TRUNCATED AT 400 WORDS)     

Unilateral suppression of pharyngeal motor cortex to repetitive transcranial magnetic stimulation reveals functional asymmetry in the hemispheric projections to human swallowing

Age-related physiological and morphological changes of muscle spindles were examined in rats (male Fischer 344/DuCrj: young, 4–13 months; middle-aged, 20–22 months; old, 28–31 months). Single afferent discharges of the muscle spindles in gastrocnemius muscles were recorded from a finely split dorsal root during ramp-and-hold (amplitude, 2.0 mm; velocity, 2–20 mm s⁻¹) or sinusoidal stretch (amplitude, 0.05–1.0 mm; frequency, 0.5–2 Hz). Respective conduction velocities (CVs) were then measured. After electrophysiological experimentation, the muscles were dissected. The silver-impregnated muscle spindles were teased and then analysed using a light microscope. The CV and dynamic response to ramp-and-hold stretch of many endings were widely overlapped in old rats because of the decreased CV and dynamic response of primary endings. Many units in old rats showed slowing of discharge during the release phase under ramp-and-hold stretch and continuous discharge under sinusoidal stretch, similarly to secondary endings in young and middle-aged rats. Morphological studies revealed that primary endings of aged rat muscle spindles were less spiral or non-spiral in appearance, but secondary endings appeared unchanged. These results suggest first that primary muscle spindles in old rats are indistinguishable from secondary endings when determined solely by previously used physiological criteria. Secondly, these physiological results reflect drastic age-related morphological changes in spindle primary endings.     

Encoder response of isolated frog muscle spindle elicited by pseudorandom noise stimuli

The present experiments investigated the signal transfer in the isolated frog muscle spindle by using pseudorandom noise (PRN) as the analytical probe. In order to guarantee that the random stimulus covered the entire dynamic range of the receptor, PRN stimuli of different intensities were applied around a constant mean length, or PRN stimuli of the same intensity were used while varying the mean length of the spindle. Subthreshold receptor potentials, local responses, and propagated action potentials were recorded simultaneously from the first Ranvier node of the afferent stem fiber, thus providing detailed insight into the spike-initiating process within a sensory receptor. Relevant features of the PRN stimulus were evaluated by a preresponse averaging technique. Up to tau = 2 ms before each action potential the encoder selected a small set of steeply rising stretch transients. A second component of the preresponse stimulus ensemble (tau = 2-5 ms) opposed the overall stretch bias. Since each steeply rising stretch transient evoked a steeply rising receptor potential that guaranteed the critical slope condition of the encoding site, this stimulus profile was most effective in initiating action potentials. The dynamic range of the muscle spindle receptor extended from resting length, L0, to about L0 + 100 microns. At the lower limit (L0) the encoding membrane was depolarized to its firing level and discharged action potentials spontaneously. When random stretches larger than the upper region of the dynamic range were applied, the spindle discharged at the maximum impulse rate and displayed no depolarization block or "overstretch" phenomenon. Random stretches applied within the dynamic range evoked regular discharge patterns that were firmly coupled to the PRN. The afferent discharge rate increased, and the precision of phase-locking improved when the intensity of the PRN stimulus was increased around a constant mean stretch; or the mean prestretch level was raised to higher values while the intensity of the PRN stimulus was kept constant. In the case when the PRN stimulus covered the entire dynamic range, the temporal pattern of the afferent discharge remained constant for at least 10 consecutive sequences of PRN. A spectral analysis of the discharge patterns averaged over several sequences of PRN was employed. At the same stimulus intensity the response spectra displayed low-pass filter characteristics with a 10-dB bandwidth of 300 Hz and a high-frequency slope of -12 dB/oct. Increasing the mean intensity of the PRN stimulus or raising the prestretch level increased the response power.(ABSTRACT TRUNCATED AT 400 WORDS)     

A `late supernormal period' in the recovery of excitability following an action potential in muscle spindle and tendon organ receptors

1. Discharge patterns have been recorded from five types of stretch receptor; frog muscle spindles, lizard tendon organs, cat soleus tendon organs and primary and secondary endings of cat soleus muscle spindles.2. The fully adapted discharge of each type of receptor is irregular, especially for frog spindles and primary endings of cat spindles as compared with the other three types (the ;regularly firing' receptors). Frog spindles and some cat spindle primary endings would maintain a discharge at very low mean rates (1/sec or less) while the remaining receptors would stop suddenly, as soon as their rate of discharge fell below a critical value characteristic for each individual ending.3. This pattern of discharge suggests that there is a peak in the excitability of ;regularly firing' receptors at a time following a preceding impulse, which corresponds to the intervals between impulses at each particular receptor's slowest rate of maintained firing, and that the excitability subsequently falls again. Primary endings of cat muscle spindles also showed some evidence of such a ;late supernormal period', but frog spindles did not.4. Direct evidence for the ;late supernormal period' was obtained from experiments in which a maintained discharge was restarted by an antidromic action potential in a receptor which had stopped firing, and to which had been applied a stretch just too small to restart the discharge.5. It is shown in an Appendix that a model receptor in which the recovery of excitability following an impulse has a hyperbolic time course, and in which Gaussian distributed noise is superimposed on the generator potential, can have a discharge pattern very closely resembling that of a frog spindle (cf. Buller, 1965).6. After addition of a late supernormal period to the model, its discharge pattern could mimic closely that of a lizard or cat tendon organ, or of a secondary ending of a cat spindle.     

Transducer action of isolated frog muscle spindle evoked by pseudorandom noise stimuli

The dynamic response properties of the isolated frog muscle spindle receptor were investigated by recording the receptor potential evoked by pseudorandom noise (PRN) stimuli. The entire dynamic range of the receptor was determined by measuring the sensory response either at different intensities of the PRN stimulus (sigma = 8-30 microns) around a constant mean length or at the same intensity while varying the mean length from resting length L0 up to L0 + 150 microns. The 3-dB bandwidth of the test signal was 130 Hz. Random stimuli often evoked brief receptor potentials with variable size but characteristic shape. This shape contained a fast depolarization transient of the receptor potential during the stretching phase of the stimulus and a slowly decaying repolarization transient during release of stretch. The depolarization transient rose faster in proportion to the increasing amplitude of the receptor potential, so that larger receptor potentials were more phasic in character than smaller ones. The repolarization transient exhibited two segments of different exponential decay: The first brief repolarization phase lasted for 5 ms; its decline (tau = 2-5 ms) was faster for larger receptor potentials. The second slowly decaying repolarization transient was the same for different receptor potential amplitudes (tau = 47 ms). Consequently, the slow repolarization transients of succeeding receptor potentials displayed temporal summation. Since the amplitude and shape of the receptor potential remained constant during repeated sequences of PRN stimuli, this test stimulus was the most appropriate for the investigation of dynamic response properties under stationary conditions. Long-term stimulation caused a small shift of the mean membrane voltage towards hyperpolarizing values. This finding together with the marked "off effect" after termination of the stimulus indicate the action of an electrogenic pumping mechanism. The dynamic range of the muscle spindle receptor extended from resting length L0 up to L0 + 100 microns. Within this range static prestretches placed a bias upon the transducing site and effectively enhanced the amplitude of the receptor potential. Further prestretch beyond the dynamic region kept the receptor potential constant at its maximum amplitude. The receptor potential amplitude distribution was not symmetrical about the mean but was skewed in favor of depolarization values responding to the stretch trajectories of the PRN stimulus. Variation of the operating point by increasing the static prestretch also shifted the mode of the response distribution towards depolarization.(ABSTRACT TRUNCATED AT 400 WORDS)     

The response to stretch of human intercostal muscle spindles studied in vitro.

1. The discharge properties of human muscle spindles have been studied in vitro in a preparation based on the biopsied external intercostal muscle. 2. The static and dynamic responsiveness of thirty-six endings in twenty visualized and histologically identified spindles have been investigated using amplitudes and velocities of stretch likely to encompass those occurring in vivo. 3. The dynamic index, measured at a stretch velocity of 3 mm/sec, ranged from 3 to 40 impulses/sec and was distributed bimodally, consitent with the presence of primary and secondary endings. 4. The relationship between the dynamic index and the velocity of stretch was approximately linear both for primary and secondary endings up to the maximum velocity tested (10 mm/sec). 5. The frequency/extension relationship was approximately linear for both primary and secondary endings. The mean values of the slope for primary and secondary endings were 16-1 +/- 8-3 S.D. of the observation and 12-1 +/- 6-5 impulses/sec per five per cent extension. 6. The slopes of the frequency/extension relationship for endings lying in the same spindle were positively correlated, significant at the 10% level. 7. It was estimated from the results in vitro that the position sensitivity of human intercostal spindles in vivo ranges from 2 to 21 impulses/sec per millimetre.     

Regularity in the generation of discharge patterns by primary and secondary muscle spindle afferents,as recorded under a ramp-and-hold stretch

The discharge frequency of primary (Ia) and secondary (II) muscle spindle afferents from the tibial anterior muscle of the cat were recorded under a rampand-hold stretch of the host muscle. The rate of ramp stretch and the prestretch of the muscle were varied systematically. The degree of stretch was kept constant. For a discharge pattern recorded at a ramp rate of 10 mm/s, the peak dynamic discharge, the maximum static value and the final static value were determined. These three discharge rate values were plotted against the maximum static value. In the resulting charts the II afferents presented themselves as a homogeneous group of spindle afferents, whereas the Ia fibers separated into three subgroups. The existence of three subpopulations of Ia fibers was verified by the method of Hald. Furthermore, it is shown that each subpopulation generated its discharge patterns in its own regularly systematic manner. It was concluded that, as one of the three Ia subpopulations exhibits much the same dynamic and static stretch properties as the II fibers, the encoder of this subpopulation must receive its receptor current from the sensory terminals of passive intrafusal chain fibers. The encoder of a second Ia subpopulation indicates its action potentials using the receptor current stemming from the bag₁ sensory terminals, these Ia fibers eliciting a slow adaptation component of a high magnitude which is assumed to be the consequence of a high level of creep in the passive intrafusal bag₁ fiber. The third Ia subpopulation initiates its action potential sequences by means of the receptor current stemming from the passive bag₂ fiber, producing behavior patterns that lie between those of the other two Ia subpopulations.     

Ion conductance changes associated with spike adaptation in the rapidly adapting stretch receptor of the crayfish

The time course of the repetitive impulse discharges has been investigated for two high intensities of maintained depolarizing currents, 30 nA and 50 nA, for which the receptor adaptation was complete within 70 msec. The changes in sodium and potassium conductance associated with the decline in spike activity have been analyzed at different instances of time by interrupting in successive experiments the various action potentials in the pulse trains either at the early phase by holding the potential at about -60 mV and recording the inward current (upstroke-gNa) or by evaluating the delayed outward current flowing as the result of a depolarizing voltage pulse which at the end of the action potential re-increased the membrane potential by mV (after potentialgK). At the higher current intensity of 50 nA the discharge frequency was increased, while larger reductions in upstroke-gNa and after potential-gK during receptor adaptation became apparent. The progressive decrease in pulse amplitude from 99 mV to 63 or 55 mV is paralleled by a gradual reduction in upstroke-gNa from 97 mmho/cm-2 to 37 or 27.5 mmho/cm-2 and in after potential-gK from 11.5 mmho/cm-2 to about 7 mmho/cm-2. When under a stimulus of 30 nA the sodium conductance decreases to an average value of 37 mmho/cm-2 only a distorted spike can be elicited, while the spike activity was completely suppressed at upstroke-gNa equals 27.5 mmho/cm-2 was essentially the same under both conditions. The results have been interpreted in terms of the model for impulse generation formulated by Michaelis and Chaplain (1973). According to the model both sodium and potassium inactivation reduce the pulse amplitude. However, while Na-inactivation reduces the frequency of impulse discharge, the K-inactivation actually leads to an increase in spike frequency. As the frequency of the short train of pulses recorded under high-intensity current stimulation remained essentially unaltered, it is suggested that the coupling between Na- and K-inactivation actually leads to an increase in spike frequency. As the frequency of the short train of pulses recorded under high-intensity current stimulation remained essentially unaltered, it is suggested that the coupling between Na- and K-inactivation ensures a constancy of the information-carrying parameter, i.e. the average impulse density.     

A model of spindle afferent response to muscle stretch

1. A unified model of the properties of stretch responses of mammalian spindle endings is proposed. This model encompasses the disparity between sensitivity of spindle endings to small and to large stretch of the muscle as well as the disparity in their dynamic responsiveness for different amplitudes of stretch. 2. In the model the mechanical properties of intrafusal fibers include a property akin to friction, which is hypothesized on the basis of reported observations on amphibian muscle. Transducer and encoder processes are modeled in the light of recent observations on isolated spindles. The model involves five unknown parameters whose values are selected by reference to certain reported observations on deefferented primary and secondary endings. The model can be used to predict responses to length changes of arbitrary time course. 3. Predicted responses to large ramp-and-hold stretch are quantitatively comparable to observations over a wide range of stretch velocities. The quantities compared include the increment in response during ramp stretch as well as the dynamic index, which is a measure of adaptation at stretch plateau. 4. At a fixed frequency of sinusoidal stretch, the relation between amplitudes of stretch and response is predicted in quantitative agreement with measurements. As the frequency of stretch is decreased, the predicted phase lead decreases and then increases, while the sensitivity decreases monotonically, in accord with observations. 5. In the model the high sensitivity for small stretch is not specific to any particular length of the muscle. When stretch is large, the region of high sensitivity is gradually reestablished at the new length, a phenomenon referred to as resetting. The dynamic response to a large stretch can be seen as arising, for the most part, from the dynamic process of resetting. 6. The influences of static or dynamic fusimotor activation on stretch responses of the primary ending are simulated by modifying the parameter values in the model. The modifications are such that static (dynamic) fusimotor activity speeds up (slows down) the resetting of the high-sensitivity region. The predictions mimic qualitatively the observed fusimotor effects not only on the response to large ramp stretch but also the contrasting effects seen with smaller, sinusoidal stretch.     

Processing vibratory stimuli in isolated frog muscle spindle

Summary Experiments were performed on isolated frog muscle spindle receptors to study the particular transducer and encoder mechanisms involved in the signal transfer of high frequency sinusoids (vibration). In order to systematically investigate the signal transfer over the entire dynamic range of the receptor, vibration stimuli were applied to the intrafusal muscle bundle at different prestretch levels, so that the isolated receptor potential or the afferent impulse train were recorded at different operating points. The vibration-induced receptor potential displayed severe distortion, because the depolarization during stretch rose steeply, whereas the repolarization transient during release of stretch declined more slowly. The positive peak velocity values of the depolarization transient increased with increasing stimulus frequency, although the ac-component of the receptor potential decreased. The negative peak velocity values of the repolarization transient remained constant throughout the frequency range. The amplitude of the receptor potential grew larger when vibration of constant amplitude was applied at increasing levels of prestretch, revealing another non-linearity of the transducer. These two types of non-linearity were influential in determining the afferent discharge pattern. Each fast depolarization transient facilitated the generation of a single action potential, which therefore could be firmly phase-locked to a small segment of the vibratory movement. Due to its short risetime, the depolarization transient tended to prevent multiple firing during one stimulus cycle. The prolonged depolarizing afterpotential of the evoked action potential operated in the same direction. Increasing prestretch greatly enhanced the responsiveness of the spindle to vibration. Thus, under appropriate conditions, the afferent discharge was driven in 11 synchrony with the vibration. An analysis is given of the after-effects of repetitive activity at the receptor site. The progressive decline of the mean membrane voltage during long lasting stimulation and the post-tetanic hyperpolarization (off-effect) on termination of the vibration suggest the action of an electrogenic pumping mechanism. As a consequence, the afferent impulse train possessed a complex structure segmented into several transient and steady states, which differed in impulse rate, phase response, and in the degree of phaselocking     

The contribution of mechanical factors to the early adaptation of the spindle response

1. Adaptation in terms of the early fall of the receptor potential was studied in isolated frog spindles. The contribution of gross mechanical changes to the decline of the response was determined by comparing the responses obtained under constant length and under constant tension.2. It was found that the early adaptation under constant stretch increased with increasing lengthening of the spindle for stretches up to 25-30% of the resting length and decreased with still stronger stretches. When the spindle was stretched by 100% or more the static phase of the receptor potential reached nearly the same height as the dynamic peak and the early adaptation approached zero.3. The early adaptation decreased with decreasing velocity of linearly rising stretch and approached zero for stretches below about 0.5 mm/sec.4. For different strengths of a steplike stretch the amount of early adaptation was linearly related to the fall in tension over the same period. The relative amount of tension fall, however, was always less than the corresponding fall of the response.5. The early adaptation was 15-20% smaller under constant tension than under constant length for stretches below the level giving the maximum dynamic peak.6. The results suggest that a comparatively small amount of the early adaptation of the spindle response to constant stretch is related to gross alterations in length in different regions of the spindle. The main part of the adaptive fall of the response is probably related to functional properties of the sensory membrane and to the ionic mechanism underlying the production of the receptor potential.