Stability of motor-unit force thresholds in the decerebrate cat.

To further test the hypothesis that some fixed property of motoneurons determines their recruitment order, we quantified the variation in force threshold (FT) for motoneurons recruited in muscle stretch reflexes in the decerebrate cat. Motor axons supplying the medial gastrocnemius (MG) muscle were penetrated with micropipettes and physiological properties of the motoneuron and its muscle fibers, i.e., the motor unit, were measured. FT, defined as the amount of MG force produced when the isolated motor unit was recruited, was measured from 20 to 93 consecutive stretch trials for 29 motor units. Trials were selected for limited variation in base force and rate of rise of force, which have been shown to covary with FT, and in peak stretch force, which gives some index of motor-pool excitability. Under these restricted conditions, large variation in FT would have been inconsistent with the hypothesis. Analysis of the variation in FT employed the coefficient of variation (CV), because of the tendency for FT variance and mean to increase together. We found that CV was distributed with a median value of 10% and with only 2 of 29 units exceeding 36%. Some of this variation was associated with measurement error and with intertrial fluctuations in base, peak, and the rate of change of muscle force. CV was not significantly correlated with motor-unit axonal conduction velocity, contraction time, or force. In three cases FT was measured simultaneously from two motor units in the same stretch trials. Changes in recruitment order were rarely observed (5 of 121 stretch trials), even when FT ranges for units in a pair overlapped. We suggest that the large variation in recruitment threshold observed in some earlier studies resulted not from wide variation in the recruitment ranking of motoneurons within one muscle, but rather from variation in the relative activity of different pools of motoneurons. Our findings are consistent with the hypothesis that recruitment order is determined by some fixed property of alpha-motoneurons and/or by some unvarying combination of presynaptic inputs that fluctuate in parallel.

Recruitment of triceps surae motor units in the decerebrate cat. I. Independence of type S units in soleus and medial gastrocnemius muscles.

1. We tested the hypothesis that reflex inhibition of soleus motor units reflects selective inhibition of slow-twitch (type S) motor units throughout the triceps surae. Physiological properties including type, together with firing behavior, were measured from single motor units in the medial gastrocnemius (MG) muscle of decerebrate cats with the use of intra-axonal recording and stimulation. MG unit firing was contrasted during net inhibition or excitation of the slow-twitch soleus muscle produced by ramp-hold-release stretches of MG. 2. Stretch of the MG muscle increased the firing of type S motor units in the MG regardless of the reflex response of the soleus muscle. When stretch inhibited soleus, each of the 14 type S units sampled from MG either was newly recruited or exhibited increases in the rate of ongoing firing. Increased firing was observed in 320 of 321 stretch trials. For 8 of these 14 units, a total of 155 stretch trials evoked reflex excitation of soleus, and unit firing increased in all trials. 3. For the eight MG type S motor units studied during both reflex inhibition and excitation of soleus, firing rate tended to be higher during inhibition. The higher rates were also associated with the higher MG forces required to elicit soleus inhibition. For one MG type S unit it was possible to compare firing rates during soleus inhibition and excitation for trials of overlapping levels of MG force. For this unit, firing rate was similar, but still appreciably higher, during inhibition. 4. Soleus inhibition was also produced by stretch of the plantaris (PL) or lateral gastrocnemius (LG) muscles. Type S units in PL (n = 2) or in LG (n = 1) were recruited or increased firing rate even when stretch of these muscles produced soleus inhibition. 5. The firing behavior of 12 fast-twitch (type F) units was studied (11 from MG, 1 from PL). All type F units either were recruited or accelerated the rate of firing during soleus inhibition, as well as during soleus excitation. 6. These findings give evidence that reflex inhibition of type S motor units in the soleus muscle does not necessarily reflect an organizational scheme in which there is inactivation of type S units in other active muscles. In the DISCUSSION we point out the absence of direct evidence for selective inactivation of units on the basis of their type classification.

Cutaneous stimulation fails to alter motor unit recruitment in the decerebrate cat.

1. An attempt was made to repeat the observation that cutaneous input to the cat medial gastrocnemius (MG) muscle sometimes had the differential effect of inhibiting motoneurons with slow axonal conduction velocity while simultaneously exciting others with fast conduction velocity. Dual microelectrode recording from intact ventral root filaments was used to study the effects of cutaneous inputs on recruitment order and on firing frequency of physiologically characterized MG motor units in decerebrate cats. Motor responses to pinch of the skin over the lateral surface of the ankle as well as electrical stimulation of the caudal cutaneous sural (CCS) nerve were contrasted with the responses to static muscle stretch as well as muscle vibration. 2. In contrast to the prediction, recruitment order in pairwise tests was the same for skin pinch or CCS stimulation as it was for MG stretch or vibration in all 32 tested pairs of motor units. This sample included seven pairs comprising one slow-twitch (S) and one fast-twitch motor unit, where the predicted reversal of recruitment should have been most apparent. Regardless of the source of excitation, recruitment of motor units of the MG was consistent with Henneman's size principle in approximately 90% of trials. 3. Skin pinch increased the firing rate of 30 of 32 individual motor units previously activated by stretch or vibration, including 7 slow-twitch units. In the remaining two units, skin pinch transiently (100-400 ms) slowed the firing of an S unit in 11 of 13 vibration + pinch trials. The other unit (type unknown) showed one or two retarded spikes in each of four vibration + pinch trials. In three S units, including the lone inhibitable unit and two others that were only excited by skin pinch, there was a significant positive rank correlation between change in unit firing frequency and change in soleus integrated electromyographic activity.(ABSTRACT TRUNCATED AT 400 WORDS)

Rhodopsin-to-metarhodopsin II transition triggers amplified changes in cytosol ATP and ADP in intact retinal rod outer segments.

We have observed rapid, light-initiated changes in unbound cytosol ATP and ADP during the rhodopsin-to-metarhodopsin II transition in intact rod outer segments (ROS). Upon illumination of the ROS, ATP is rapidly removed from the unbound phase of ROS, accompanied by the concomitant release of ADP into the cytosol. The exchange process involves decreases of approximately equal to 0.5 mM ATP in ROS cytosol ATP content in response to a saturating flash. At levels of light well below saturation (less than 0.001% bleach), the process is highly amplified, with a decrease in cytosol ATP of approximately equal to 2,000 ATP molecules per absorbed photon per ROS. Rapid time-resolution techniques reveal that cytosol ATP content decreases rapidly, within 250 msec of a saturating flash. Bleaching rhodopsin to metarhodopsin II results in a decrease in cytosol ATP, accompanied by an increase in cytosol ADP, whereas photoreversal of metarhodopsin II by a blue flash reverses the process, increasing ATP concentration to its control level in the dark. The photoreversibility of the ATP decrease during the rhodopsin-to-metarhodopsin II transition establishes a direct link between the state of an early intermediate of photolyzed rhodopsin and the state of a nucleoside triphosphate in intact ROS. The results are consistent with a light-activated exchange of unbound ATP for bound ADP, and we propose, therefore, an ATP/ADP amplification cycle in which metarhodopsin II catalyzes the exchange of ATP for ADP on a nucleotide binding protein.