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PubMed1992

Selective reduction of creatine kinase subunit mRNAs in striated muscle of diabetic rats.

Creatine kinase (CK) is important for energy transfer and is composed of mitochondrial (mitCK), muscle (MCK), and brain (BCK) subunits, each being the product of separate nuclear genes. The concentrations of MCK and BCK mRNAs have been shown to decrease in streptozotocin-hypoinsulinemic rat hearts, and in this report, we examined in detail the diabetic effect on CK gene expression in cardiac muscle and in two types of skeletal muscle. The level of sarcomeric mitCK mRNA was not altered in the diabetic myocardium, but was reduced by 86 and 67% in diabetic slow-twitch soleus muscle and fast-twitch extensor digitorum longus (EDL) muscle, respectively. MCK mRNA was also lowered in diabetic soleus muscle by 56%, while it remained at control levels in diabetic EDL. In both skeletal muscles, at either state, BCK mRNA was not detectable. There was a 33% decrease in total CK activity in diabetic cardiac and soleus muscle, but not in EDL. Diabetes thus exerts a widespread, muscle type-dependent adverse effect on CK expression that we found to be insulin therapy revertible. This study adds to our understanding of defective energy transduction in diabetic muscle.

Animals↗
PubMed1992

Comparison of protein lymph flux and extravascular uptake in skin during increased venous pressure.

The hypothesis that the albumin reflection coefficient is underestimated from measurements of lymph protein flux because of a prolonged interstitial washout was tested in the hindleg of anesthetized rabbits. In heel skin, the initial extravascular uptake for labeled albumin and immunoglobulin G (IgG) was compared with prenodal lymph flux for native albumin and IgG. Venous pressure was increased in one leg for 4 h, while the contralateral leg was the control. The extravascular uptake for labeled albumin was 67% greater than control during the fourth hour of increased venous pressure. Lymph albumin flux was 2.3 times control, indicating a small but significant washout of interstitial albumin. The magnitude of the interstitial washout for IgG was less than that for albumin. Using the relationship between the change in extravascular uptake and the change in lymph flow, the reflection coefficients for albumin and IgG were both 0.91. The reflection coefficient for albumin using lymph protein flux was lower because of a continued interstitial washout.

Animals↗
PubMed1992

Geometry of blood-tissue exchange in bat flight muscle compared with bat hindlimb and rat soleus muscle.

We investigated the relationship between capillary-to-fiber geometry and muscle aerobic capacity by comparing the bat flight muscle (pectoralis muscle), i.e., an ultimate case of extreme O2 demand in a mammalian skeletal muscle, with bat hindlimb and rat soleus muscles. At a given sarcomere length (2.1 microns), fiber cross-sectional area was considerably smaller in bat muscles (pectoralis 318 +/- 10 microns 2, hindlimb 447 +/- 35 microns 2) than in rat soleus muscle (2,027 +/- 125 microns 2). Capillary number per fiber cross-sectional area was much greater in bat pectoralis (6,394 +/- 380/mm2) than in bat hindlimb and rat soleus muscle (2,865 +/- 238 and 1,301 +/- 129/mm2, respectively; all values normalized to 2.1-microns sarcomere length). At the same sarcomere length (2.1 microns), the degree of tortuosity and branching of capillaries were significantly greater in bat pectoralis than in bat hindlimb and rat soleus muscle. In bat flight muscle, capillary length per fiber volume was very high (9,025 +/- 342/mm2). It was 2.2- and 5.4-fold larger than in bat hindlimb and rat soleus, respectively. Mitochondria occupied 35.3 +/- 1.2, 16.5 +/- 1.3, and 6.1 +/- 0.9% of the muscle fiber volume in bat pectoralis, hindlimb, and rat soleus muscles, respectively. There was a strong correlation between capillary length (as well as capillary surface) per fiber volume and mitochondrial volume density in all muscles. Considering capillary supply and mitochondrial volume on an individual fiber basis, we found that 1) the number of capillaries around a fiber was linearly related to mitochondrial volume per micron length of fiber in the muscles but that 2) capillary surface per fiber surface, at given mitochondrial volume per micron length of fiber, was about twice as large in bat pectoralis as in rat soleus muscle, whereas in bat hindlimb it was intermediate between that in bat pectoralis and that in rat soleus muscle. This was due to the differences in fiber size (rat soleus greater than bat muscles) and capillary-to-fiber ratio (bat pectoralis greater than hindlimb) between the muscles. It is notable that in the bat, the substantially greater O2 transfer capacity of the flight muscle compared with hindlimb was achieved by increasing the size of the capillary-to-fiber interface, i.e., capillary-to-fiber surface, via an increase in capillary number rather than by substantially reducing fiber size.

Animals↗
PubMed1993

Calcium sensitivity of skinned ferret EDL, soleus, and cremaster fibers.

The properties of the contractile system at different times of the year in the ferret extensor digitorum longus (EDL), soleus and cremaster muscles were examined by using chemically skinned (Triton X-100) preparations. The results show clear differences in calcium sensitivity between these skeletal muscles. The apparent calcium threshold for activation was lower in soleus than in EDL, while calcium concentrations ([Ca2+]) required to obtain the half-maximal tension, expressed as pCa50 (-log[Ca2+]), was lower in EDL than in soleus muscle. In fact, pCa50 obtained in fast and slow fibers by fitting the experimental data points by a modified Hill equation was 5.92 +/- 0.02 (n = 9) and 6.09 +/- 0.03 (n = 11) respectively. So EDL appears to be a typical fast-twitch muscle and soleus a typical slow-twitch muscle. Adult ferret cremaster muscle was composed of two types of fibers during the quiescent period similar to EDL and soleus, and only one type that was intermediate between EDL and soleus in the breeding period, as assessed by pCa50 values. These annual modifications in calcium activation of adult ferret cremaster muscle could be related to changes in the function of these muscles and may be correlated with seasonal variations of sexual activity.

Animals↗
PubMed1991

EMG patterns of rat ankle extensors and flexors during treadmill locomotion and swimming.

Intramuscular electromyography (EMG) was used to determine and compare the recruitment patterns of the rat soleus (Sol), tibialis anterior (TA), and a deep and a superficial portion of the medial gastrocnemius (MG) during treadmill locomotion at various speeds and inclines and during swimming. Raw EMG signals for 10-20 step or stroke cycles were rectified, averaged, and processed to determine cycle period (EMG onset of one cycle to EMG onset of the next cycle), EMG burst duration, and integrated area of the rectified burst (IEMG). Mean EMG per burst was calculated as IEMG/burst duration. IEMG/min was calculated as IEMG times the number of bursts (cycles) per minute. Cycle period and burst duration of the extensors decreased hyperbolically, while the TA burst duration was unchanged, with increased treadmill speed. With increased treadmill speed, IEMG was decreased in the Sol and unchanged in the MG and TA, whereas IEMG/min decreased in the Sol and increased in the MG and TA. An elevation in treadmill incline resulted in an increase in the activation levels of the MG but not in the Sol or TA. These data indicate that the additional power required at increased speeds and/or inclines of treadmill locomotion is derived from the recruitment of the fast extensors, e.g., the MG. The mean cycle period during swimming was similar to that observed during the fastest treadmill locomotion. EMG burst durations and amplitudes, however, were higher in the TA, relatively similar in the MG, and lower in the Sol during swimming than treadmill locomotion.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1991

Rostrocaudal pattern of fiber-type changes in an overloaded rat ankle extensor.

Our aim was to quantify the overload-induced hypertrophy and conversion of fiber types (type II to I) occurring in the medial head of the gastrocnemius muscle (MG). Overload of MG was induced by a bilateral tenotomy/retraction of synergists, followed by 12-18 wk of regular treadmill locomotion (2 h of walking/running per day on 3 of 4 days). We counted all type I fibers and determined type I and II mean fiber areas in eight equidistant sections taken along the length of control and overloaded MG. Increase in muscle weights (31%), as well as in total muscle cross-sectional areas (37%) and fiber areas (type I, 57%; type II, 34%), attested to a significant hypertrophic response in overloaded MG. An increase in type I fiber composition of MG from 7.0 to 11.5% occurred as a result of overload, with the greatest and only statistically significant changes (approximately 70-100%) being found in sections taken from the most rostral 45% of the muscle length. Results of analysis of sections taken from the largest muscle girth showed that it significantly underestimated the extent of fiber conversion that occurred throughout the muscle as a whole. These data obtained on the MG, which possesses a compartmentalization of fiber types, support the notion that all fiber types respond to this model with a similar degree of hypertrophy. Also, they emphasize the complex nature of the adaptive changes that occur in these types of muscles as a result of overload.

Animals↗
PubMed1993

Role of protein intake on protein synthesis and fiber distribution in the unweighted soleus muscle.

Protein turnover in skeletal muscle is very sensitive to protein intake. To examine whether protein intake is able to affect protein synthesis in the atrophied soleus muscle, the effects of a high-protein (30%, HP) and a medium-protein (15%, MP) diet were studied in rats after 21 days of hindlimb unweighting. Three weeks of unweighting induced a sharp decrease in food intake (30%). The fractional rate of protein synthesis (ks) was determined in vivo in the slow-twitch soleus muscle by use of a flooding-dose method. With respect to pair-fed animals, a significant reduction in ks occurred (33%) in MP non-weight-bearing rats, whereas it was of lesser magnitude and not significant in HP rats. In the atrophied soleus muscle of non-weight-bearing MP rats, a large decrease (42%) in type I fiber distribution was accompanied by an increase in intermediate and type IIa fibers. By contrast, a higher percentage of type I fiber was maintained with the HP diet. However, the HP diet had no beneficial effect in preventing the decrease in either type I fiber cross-sectional area (65%) or the average decrease in absolute myofibrillar and mitochondrial volumes (69 and 52%, respectively). These results demonstrate that an HP intake did not prevent soleus muscle atrophy but may sustain protein synthesis and partly preserve fiber type distribution without affecting the ultrastructural composition of fibers. Because the circulating level of free 3,5,3'-triiodothyronine was reduced by 14% with the HP diet, this effect on fiber type distribution, and possibly protein synthesis, may involve thyroid hormones.

Animals↗
PubMed2000

High-impact exercise and growing bone: relation between high strain rates and enhanced bone formation.

We investigated whether high-impact drop jumps could increase bone formation in the middiaphyseal tarsometatarsus of growing rooster. Roosters were designated as sedentary controls (n = 10) or jumpers (n = 10). Jumpers performed 200 drop jumps per day for 3 wk. The mechanical milieu of the tarsometatarsus was quantified via in vivo strain gauges. Indexes of bone formation and mechanical parameters were determined in each of twelve 30 degrees sectors subdividing the middiaphyseal cortex. Compared with baseline walking, drop jumping produced large peak strain rates (+740%) in the presence of moderately increased peak strain magnitudes (+30%) and unaltered strain distributions. Bone formation rates were significantly increased by jump training at periosteal (+40%) and endocortical surfaces (+370%). Strain rate was significantly correlated with the specific sites of increased formation rates at endocortical but not at periosteal surfaces. Previously, treadmill running did not enhance bone growth in this model. Comparing the mechanical milieus produced by running and drop jumps revealed that jumping significantly elevated only peak strain rates. This further emphasized the sensitivity of immature bone to high strain rates.

Animals↗
PubMed2003

Dihydropyridine and ryanodine receptor binding after eccentric contractions in mouse skeletal muscle.

The purpose of this study was to determine whether there are alterations in the dihydropyridine and/or ryanodine receptors that might explain the excitation-contraction uncoupling associated with eccentric contraction-induced skeletal muscle injury. The left anterior crural muscles (i.e., tibialis anterior, extensor digitorum longus, and extensor hallucis longus) of mice were injured in vivo by 150 eccentric contractions. Peak isometric tetanic torque of the anterior crural muscles was reduced approximately 45% immediately and 3 days after the eccentric contractions. Partial restoration of peak isometric tetanic and subtetanic forces of injured extensor digitorum longus muscles by 10 mM caffeine indicated the presence of excitation-contraction uncoupling. Scatchard analysis of [3H]ryanodine binding indicated that the number of ryanodine receptor binding sites was not altered immediately postinjury but decreased 16% 3 days later. Dihydropyridine receptor binding sites increased approximately 20% immediately after and were elevated to the same extent 3 days after the injury protocol. Muscle injury did not alter the sensitivity of either receptor. These data suggest that a loss or altered sensitivity of the dihydropyridine and ryanodine receptors does not contribute to the excitation-contraction uncoupling immediately after contraction-induced muscle injury. We also concluded that the loss in ryanodine receptors 3 days after injury is not the primary cause of excitation-contraction uncoupling at that time.

Animals↗
PubMed2004

Activity-driven synaptic and axonal degeneration in canine motor neuron disease.

The role of neuronal activity in the pathogenesis of neurodegenerative disease is largely unknown. In this study, we examined the effects of increasing motor neuron activity on the pathogenesis of a canine version of inherited motor neuron disease (hereditary canine spinal muscular atrophy). Activity of motor neurons innervating the ankle extensor muscle medial gastrocnemius (MG) was increased by denervating close synergist muscles. In affected animals, 4 wk of synergist denervation accelerated loss of motor-unit function relative to control muscles and decreased motor axon conduction velocities. Slowing of axon conduction was greatest in the most distal portions of motor axons. Morphological analysis of neuromuscular junctions (NMJs) showed that these functional changes were associated with increased loss of intact innervation and with the appearance of significant motor axon and motor terminal sprouting. These effects were not observed in the MG muscles of age-matched, normal animals with synergist denervation for 5 wk. The results indicate that motor neuron action potential activity is a major contributing factor to the loss of motor-unit function and degeneration in inherited canine motor neuron disease.

Animals↗
PubMed2002

Adaptive changes in locomotor activity following botulinum toxin injection in ankle extensor muscles of cats.

The present study investigated the adaptations made in motor behavior following a temporary reduction in ankle extensor activity in the walking cat. Temporary muscle weakness was induced by injecting botulinum toxin into the lateral gastrocnemius (LG), plantaris (PL), and soleus (SOL) muscles, or SOL alone. The medial gastrocnemius (MG) muscle was not injected. Adaptations in the level of muscle activity were recorded using chronically implanted electromyographic (EMG) electrodes. Serial recordings were made prior to botulinum toxin injections and for several days following the injections. Kinematic analysis of ankle joint movements was made from video records to assess the impact of the botulinum toxin injections on the function of the ankle joint during walking. Following injection of the LG, PL, and SOL muscles with botulinum toxin, the amplitude of the MG burst increased over a period of a few days to a week. This increase was similar to the previously reported changes produced in MG following transection of the nerves serving LG, PL, and SOL. Following the weakening of the ankle extensor muscles, there was a temporary deficit in ankle function during walking as evidenced by a marked increase in the amount of ankle flexion that occurred at stance onset. This functional deficit recovered relatively quickly and was not associated with a return of the EMG pattern to the preinjection pattern. After recovery from the initial injections, a second injection of botulinum toxin into SOL alone was performed. No functional deficits were observed in the ankle movements during walking following this second injection. However, weakening SOL produced increases in the burst amplitudes of the MG, LG, and PL muscles over a period of a few days. This suggests that normal movements at the ankle during walking can be generated with more than one pattern of ankle extensor activity and that there is flexibility in how the necessary torque is produced. A final procedure, transection of the nerves serving LG, PL, and SOL, failed to produce any functional deficits in ankle movements. The implication is that adaptations to the neural control of ankle extensor activity that were induced by the initial procedure persisted after the recovery of the injected muscles and were sufficient to compensate for the subsequent challenges.

Adaptation, Physiological↗
PubMed1989

Mutable and immutable features of paw-shake responses after hindlimb deafferentation in the cat.

1. Hindlimb paw-shake responses were assessed before and after unilateral deafferentation (L3-S1) in chronic-spinal cats (n = 5), spinalized at the T12 level 1 yr earlier. Selected ankle flexor [tibialis anterior (TA)] and extensor [lateral gastrocnemius (LG)] and knee extensor [vastus lateralis (VL)] muscles were surgically implanted with chronic electromyographic (EMG) electrodes to determine mutable features of cycle characteristics and muscle synergies that are modulated by motion-dependent feedback as opposed to immutable features that are centrally programmed and not modulated by limb afference. 2. Paw-shake responses were difficult to elicit in the extensively deafferented hindlimb; this was true particularly during the first recovery weeks after deafferentation. By the end of the first month, however, brief responses of 1 or 2 cycles were commonly elicited in four of five cats, and responses of 3-7 cycles were common by the end of the second month in three of five cats. Initially, responses in the deafferented limb were elicited by stimuli applied to the dorsolateral thigh, an oval patch of skin innervated by intact S2 afferents. Over the 4-mo recovery period, however, the receptive field of the largely denervated skin expanded, and responses were also elicited by stimuli applied to the lateral aspect of the knee and shank, but usually not the paw. 3. In addition to fewer average cycles per response (5 vs. 10 cycles), paw shaking evoked in the deafferented hindlimb was characterized by longer-than-average cycle periods (124 vs. 98 ms), but the average cycle period varied widely among responses, ranging from 99 to 239 ms. Before deafferentation, the temporal organization of consecutive cycles was stereotypic; cycle periods increased linearly throughout a response. After deafferentation, however, there was no systematic relationship between cycle period and cycle number, and approximately 14% of the records with greater than or equal to 3 cycles were characterized by arhythmical sequences of EMG bursts. 4. At the ankle, LG burst duration was not altered by deafferentation, but TA onset and burst duration were affected. Before deafferentation, TA onset was invariant with respect to the beginning of the cycle, and burst duration increased linearly with cycle period. After deafferentation, however, TA onset was delayed, and the delay increased linearly with cycle period. Consequently, the TA burst duration was brief and unrelated to cycle period.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗
PubMed1990

Reduction of Ib autogenetic inhibition in motoneurons during contractions of an ankle extensor muscle in the cat.

1. Triceps surae and plantaris (Pl) motoneurons were recorded intracellularly in chloralose or pentobarbital sodium (Nembutal)-anesthetized cats during unfused tetanic contractions of gastrocnemius medialis muscle (GM) produced by stimulating either a cut branch of the GM nerve or the muscle directly. 2. In alpha-motoneurons, during a series of GM twitches at 10/s, contraction-induced inhibitory potentials, probably the result of input from Golgi tendon organs (autogenetic inhibition), rapidly subsided before the end of the series. In contrast, excitatory potentials, probably the result of the activation of spindle primary endings during relaxation from contraction, persisted. 3. In gastrocnemius lateralis-soleus (GL-S) and Pl motoneurons lacking an excitatory connection with Ia afferents from GM, the sustained contraction of this muscle also elicited a declining inhibition. Rapid reduction of contraction-induced autogenetic inhibition was also observed in homonymous gamma-motoneurons. During unfused tetanic contractions lasting 0.5-4s, inhibitory potentials quickly subsided, but an abrupt increase in contractile force elicited a new series of decreasing inhibitory potentials. 4. The assumption that the inhibition induced by GM unfused tetanic contractions was due to activation of homonymous Ib afferents was supported by observations of the effects of electrical stimulation of the GM nerve. In Pl motoneurons lacking an excitatory connection with Ia afferents from GM, repetitive trains applied to the GM nerve, at a strength just above threshold for group I fibers, elicited rapidly declining inhibitory potentials similar to those produced by GM contraction. It was verified that during such stimulation, the amplitude of the group I afferent volleys did not decrease. 5. Reduction of contraction-induced Ib inhibition during sustained GM contraction was still present after a low spinalization of the preparation. As GM tendon organ discharges were verified to persist throughout prolonged contractions, the observed decline of autogenetic inhibition is likely to depend on a spinal mechanism, possibly involving presynaptic inhibition of Ib afferents and/or mutual inhibition of Ib-inhibitory interneurons.

Animals↗
PubMed1990

Segmental reflex action in normal and decerebrate cats.

1. The objective of this study was to evaluate the action of the stretch reflex on the ankle extensor muscles of normal and decerebrate cats. 2. Experiments were performed on nine freely standing, unrestrained cats and repeated after decerebration at the premammillary level. The length, force, and electromyograph (EMG) of the soleus (SOL) and lateral gastrocnemius (LG) muscles were recorded with the use of implanted transducers and electrodes. 3. The left ankle joint was unexpectedly and reproducibly dorsiflexed by briefly stimulating the common peroneal (CP) nerve with electrodes within an implanted nerve cuff. The ensuing twitch contractions of the ankle dorsiflexor muscles stretched the ankle extensor muscles by 0.3-2.0 mm. Lidocaine was infused into another nerve cuff proximal to the stimulation site, to reversibly block the central propagation of evoked volleys in the CP nerve. 4. Reflex action before and after decerebration was measured from the responses to perturbations of similar amplitude and duration delivered at approximately matched background values of muscle length and force. In most cats the temperature of the hindlimb was monitored with an implanted thermistor and was restored to normal values with radiant heat after decerebration. 5. A stretch imposed on the tonically active ankle extensor muscles immediately caused a considerable rise in the force recorded from the triceps tendon. Within 30-40 ms the triceps force peaked, reaching a value 10-20 N greater than background, and then rapidly declined while the extensor muscles were still lengthening. The initial rise in force preceded any change in triceps EMG. It was attributed to the intrinsic viscoelasticity of the stretched muscles and tendons. After decerebration the magnitude and timing of the initial force peak did not change. 6. A short-latency reflex EMG burst was typically recorded from both the SOL and LG muscles, starting 11-17 ms after stimulus onset. After decerebration the area of the reflex EMG burst increased in all nine cats, typically by a factor of 2 or 3. 7. After decerebration a second, smaller increase in force was typically observed starting 60-80 ms after onset of stretch. This later force rise, interpreted to be of reflex origin, was rarely apparent in normal cats. 8. Decerebration introduced consistent modifications in postural behavior that were revealed by pushing down on the back of quietly standing cats. In normal cats, after brief pushes that stretched the ankle extensor muscles by 1-2 mm, the EMG, force, and length quickly stabilized near their initial values.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1990

Factors determining segmental reflex action in normal and decerebrate cats.

1. In the companion paper the gain of the stretch reflex in the ankle extensor muscles of normal cats was shown to increase after decerebration. The objectives of this study were 1) to identify the origin of the increased reflex and 2) to evaluate the contribution from afferents other than ankle extensor muscle afferents to the short-latency reflex. 2. Six cats were trained to stand unaided on four pedestals. Three cats were also trained to control the force exerted with the left hindlimb. The left soleus (SOL) and lateral gastrocnemius (LG) electromyogram (EMG), length, force, and temperature were recorded by chronically implanted electrodes and transducers. Measurements were taken before and after decerebration at the premammillary level. After decerebration limb temperature was returned to its normal range by the use of radiant heat. 3. Reproducible ramp-and-hold stretches and releases of the ankle extensor muscles were produced by a servo-controlled motor that rotated the left rear pedestal about the ankle joint. The length of the ankle extensor muscles changed by 2-3 mm within 30-35 ms after the onset of a ramp perturbation. Reflex responses before and after decerebration were compared at matched background values of muscle length and force. 4. In both the SOL and LG muscles, a short-latency EMG burst appeared 8-12 ms after stretch onset and lasted approximately 20 ms. After decerebration the onset of the rectified and smoothed EMG burst remained unchanged, but its area was increased by 36-89%. 5. The lateral gastrocnemius-soleus (LG-S) electroneurogram (ENG) was chronically recorded in two cats with a nerve cuff recording electrode implanted on the LG-S nerve. LG-S ENG activity started to increase soon after stretch onset and remained high during the entire ramp phase. The stretch-evoked LG-S ENG burst started approximately 8 ms earlier than the short-latency SOL and LG EMG bursts. It was interpreted to reflect mainly an increase in the activity of Group Ia and Ib muscle afferents, caused by increases in both muscle length and muscle force during the stretch. After the cats were decerebrated, for matched postural conditions, the area of the stretch-evoked LG-S ENG burst was increased by 29-35%. Because the length and force changes sensed by the muscle receptors before and after decerebration were similar, this suggests that the sensitivity of muscle spindles was increased as a consequence of altered activity in fusimotor neurons after decerebration.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1992

Velocity of shortening of single motor units from rat soleus.

1. The force-velocity relationship of a motor unit can provide insight into the contractile proteins of its constituent fibers as well as fundamental information about the function and use of the motor unit. Although the force-velocity profiles of whole muscle and skinned mammalian fibers have been studied, technical difficulties have prevented similar studies on motor units. A technique is presented to directly measure the velocity of shortening of individual motor units from in vivo rat soleus muscle. 2. The soleus muscles of anesthetized rats were dissected free of surrounding tissue while their nerve and blood supplies were preserved. Both tendons were cut, and the distal tendon was attached to a servomechanism to control muscle length, whereas the proximal tendon was attached to a force transducer. Single motor units were stimulated via the ventral roots. 3. The major problem encountered in measuring the force-velocity profile of a motor unit was that the force from the large number of passive fibers and connective tissue in the soleus confounded the force produced by the small number of active fibers in the motor unit. This problem was minimized by measuring active motor unit tension during an isovelocity ramp. This allowed experimental measurement of the passive tension by shortening the muscle with an identical isovelocity ramp without, however, stimulating the motor unit. Active tension was estimated by subtracting the passive tension waveform from the waveform recorded when the motor unit was active. 4. The method substantially reduced the noise from the passive fibers; however, problems remained. The probable sources of error are discussed, with the most significant being the elasticity associated with the blood and nerve connections to surrounding tissue. The elasticity prevents uniform shortening velocities along the length of the active fibers, thereby introducing a systematic bias to measurements made at high velocities. These errors are most pronounced when the data are extrapolated to determine the maximum velocity of shortening (Vmax). Determination of velocity at peak power (Vpp) is a more robust measure; however, of the 34 motor units studied, only 19 exhibited a distinct peak in the power-force curve, indicating residual noise. 5. To assess the validity of using twitch contraction time as an index of the velocity of shortening, when possible, Vmax and Vpp of each motor unit were correlated with the inverse of its twitch contraction time. The correlation was poor (r less than 0.2), indicating that, although widely used, twitch contraction time is a poor index of contractile speed.

Animals↗
PubMed1996

Differential modulation of disynaptic cutaneous inhibition and excitation in ankle flexor motoneurons during fictive locomotion.

1. Intracellular recording from extensor digitorum longus (EDL) and tibialis anterior (TA) alpha-motoneurons during fictive locomotion was used to examine patterns of modulation of oligosynaptic postsynaptic potentials (PSPs) produced by electrical stimulation of the cutaneous superficial peroneal (SP) and medial plantar (MPL) nerves in unanesthetized, decerebrate adult cats. 2. In all 20 EDL motoneurons studied, electrical stimulation of the SP nerve with single pulses at about twice threshold for the most excitable fibers in the nerve (2xT) produced either no synaptic potentials or relatively small oligosynaptic excitatory or inhibitory PSPs (EPSPs or IPSPs), both at rest and during the extension phase of fictive stepping. However, at the onset of the flexion phase large, presumably disynaptic IPSPs (central latencies 1.7-2.0 ms) appeared in the SP responses. These IPSPs usually decreased in amplitude later in the flexion phase despite maintained membrane depolarization. 3. In most (7/8) TA motoneurons, SP stimulation produced oligosynaptic EPSPs at rest and during the extension phase of fictive stepping. These EPSPs were suppressed during flexion in a majority of TA cells studied (5/8) but no clearly disynaptic IPSPs were found in any TA motoneuron. 4. In most EDL and TA motoneurons, stimulation of the MPL nerve produced oligosynaptic EPSPs at rest and during the extension phase, most with latencies in the presumably disynaptic range (< or = 2.0 ms). When present, these MPL EPSPs were suppressed throughout the flexion phase of stepping in almost all EDL (18/ 20) and TA (6/8) motoneurons examined. 5. The available evidence suggests that these modulation effects during fictive stepping are due primarily to convergence of control information from the spinal central pattern generator (CPG) for locomotion onto segmental interneurons in the oligosynaptic cutaneous pathways. 6. These observations extend the evidence for precise differential control of transmission through cutaneous reflex pathways in the cat hindlimb by the locomotor CPG. Taken together with earlier evidence about locomotor modulation of cutaneous PSPs in flexor digitorum longus (FDL) motoneurons, the data suggest that cutaneous information from the dorsal surface of the foot, carried in part by the SP nerve, projects to digit motoneurons (FDL and EDL) through discrete sets of last-order interneurons that also receive powerful excitation from the locomotor CPG during flexion. In contrast, the last-order interneurons that convey excitatory information from the SP nerve to at least some TA motoneurons are inhibited by the CPG during flexion. 7. Another contrast resides in the fact that oligosynaptic cutaneous excitation from the plantar surface of the foot, via the MPL nerve, is suppressed in FDL, EDL, and TA motoneurons during the flexion phase of locomotion. The available information is consistent with the possibility that MPL effects may be delivered to these motor nuclei by common interneurons. 8. We suggest an interneuronal circuitry that could account for these observations and discuss possible functional implications of modulation of these sensory pathways during locomotion.

Animals↗
PubMed1998

Organization of recurrent inhibition and facilitation in motor nuclei innervating ankle muscles of the cat.

The distribution of recurrent inhibition and facilitation to motor nuclei of muscles that act at the cat ankle joint was compared with the locomotor activity and mechanical action of those muscles described in published studies. Emphasis was placed on motor nuclei whose muscles have a principal action about the abduction-adduction axis and the pretibial flexors: tibialis posterior (TP), peroneus longus (PerL), peroneus brevis (PerB), the anterior part of tibialis anterior (TA) and extensor digitorum longus (EDL). Most intracellular recordings in spinalized, unanesthetized decerebrate cats showed only inhibitory or excitatory responses to antidromic stimulation of peripheral nerves, but mixed effects were also seen. Recurrent effects among motor nuclei of ankle abductors and adductors were not distributed uniformly. TP motoneurons received recurrent inhibition from most other nuclei active in stance and stimulation of the TP nerve inhibited these motor nuclei. Although PerB motoneurons are also active during stance, they received primarily facilitation from most motor nuclei. PerL received mixtures of inhibition and facilitation from all sources. Stimulation of the nerves to PerL, PerB, and peroneus tertius (PerT) produced weak recurrent inhibition and facilitation, even in homonymous motoneurons and motoneurons of Ia synergists. The ankle flexors TA and EDL displayed different patterns of recurrent inhibition and facilitation. TA motoneurons received prominent homonymous inhibition and inhibition from semitendinosus (St). EDL, whose activity profile differs from TA and which also acts at the digits, did not receive strong recurrent inhibition from either TA or St, nor did stimulation of the EDL nerve produce much inhibition. The distribution of recurrent inhibition and facilitation is correlated with the pattern of locomotor activity, but with exceptions that suggest an influence of mechanical action, particularly in the antagonistic interactions between TP and PerB. The extended pattern of recurrent inhibition, the reduction or absence of inhibition produced by motor nuclei with individualized functions or digit function and the prevalence of facilitation suggest that the recurrent Renshaw system is organized into inhibitory and disinhibitory projections that participate in the control of sets of motor nuclei engaged in rhythmic and stereotyped movements.

Action Potentials↗