PubMed HealthSearch

Biomedical subjects

G E Loeb

Publications and source records attributed to G E Loeb.

At least 19 recordsLinked to original sources

An intrinsic mechanism to stabilize posture--joint-angle-dependent moment arms of the feline ankle muscles.

The neuromuscular system can use sensory feedback to regulate motion, but these feedback loops involve relatively long delays (50-100 ms) and may produce undesirable oscillations. However, stabilizing changes in muscle force can also be provided intrinsically (i.e. without feedback) by 2 properties of the muscle itself, its force-length and force-velocity relationships. We have discovered another intrinsically stabilizing mechanism in the musculoskeletal architecture of the cat ankle joint. Many of its muscles have their predominant moment arms about the adduction/abduction axis, with smaller moment arms for inversion/eversion and about the principal axis of motion, dorsiflexion/extension. The magnitudes of the moment arms in ab/adduction and in/eversion depend strongly on joint angle, increasing for positions away from neutral that lengthen the muscles. Thus, co-activation of agonist-antagonist pairs, known to occur in these muscles, would provide immediate stabilizing changes in torque when the ankle is perturbed away from neutral position.

Anesthesia

Neuromuscular organization of feline anterior sartorius: II. Intramuscular length changes and complex length-tension relationships during stimulation of individual nerve branches.

The feline anterior sartorius is a long strap-like muscle composed of short muscle fibers. Nerve branches that enter this muscle contain the axons of motor units whose constituent muscle fibers are distributed asymmetrically within the muscle. In the present study, twitch and tetanic isometric contractions were evoked by stimulating individual nerve branches while muscle force was recorded and intramuscular length changes were monitored optically by the movement of reflective markers on the muscle. Contractions elicited by stimulating the parent nerve produced little change in the positions of the surface markers. Contractions elicited by stimulating the proximally or distally directed nerve branches caused the muscle to shorten at the end closest to the nerve branch and lengthen at the opposite end. Some muscles were supplied by a centrally directed nerve branch whose stimulation produced variable effects: in some cases a portion of the muscle shortened whereas the rest lengthened, but in other cases, the positions of the surface markers showed little change. The intramuscular length changes produced by stimulating single nerve branches were greater during isometric contractions at short whole-muscle lengths than at long whole-muscle lengths. The twitch and tetanic length-tension relationships obtained by stimulating the individual nerve branches were not congruent with the length-tension relationship produced when the parent nerve was stimulated. At short whole-muscle lengths, stimulation of a single nerve branch generated only a small fraction of the force that could be generated by the muscle when the parent nerve was stimulated. As whole-muscle length increased, an increased fraction of total muscle force could be generated by stimulating a single nerve branch. The results suggest that a complex relationship between passive and active elements contributes to the total muscle force and depends on the distribution of active and passive muscle units throughout the muscle.

Animals

Electromyographic studies of neck muscles in the intact cat. I. Patterns of recruitment underlying posture and movement during natural behaviors.

Natural head movements in alert, unrestrained cats were studied using video-filming, videofluoroscopy and electromyographic (EMG) recording methods. In each cat, up to sixteen neck muscles or neck-muscle compartments were implanted with recording electrodes. Patterns of muscle recruitment were examined during systematically-selected behavioral epochs in which the cat held a range of stationary postures, and when it performed volitional and exploratory behaviors such as flexion-extension or turning, grooming, eating, or headshaking. Patterns of muscular activity were interpreted with reference to simultaneous video images of head and neck movements. In separate, videofluoroscopic analysis, flexion-extension movements were examined to gain insight into the underlying movements of the skull and cervical vertebrae. These and other movements were found commonly to depend upon changes in joint angles between lower as well as upper cervical joints. Stationary postures in which the neck was held vertically were consistently associated with tonic EMG activity in only two long dorsal muscles, biventer cervicis and occipitoscapularis. Less consistent activity was also present in dorsal intervertebral muscles crossing lower cervical joints. When the neck was held horizontally, the long dorsal muscles increased their EMG activity and moderate activity was also recorded in deeper intervertebral and suboccipital muscles. When flexion-extension occurred around upper cervical joints, greatest activity was recorded in rectus capitis posterior and complexus, but when it involved the lower cervical joints, large changes in EMG activity could also be detected in biventer cervicis, occipitoscapularis, and the intervertebral muscles crossing lower cervical joints. During specialized, sagittal-plane movements such as grooming, well-defined patterns of synergy could be recognized that varied according to the degree of involvement of upper and lower cervical joint-sets. Movements in the horizontal plane were associated with EMG activity in a largely different subset of neck muscles including splenius, longissimus capitis and obliquus capitis inferior. The levels of EMG activity during flexion-extension or turning movements were much lower than those observed during other more vigorous behaviors, such as head shaking. Some neck muscles, such as clavotrapezius and sternomastoideus, could only be recruited during forceful or ballistic head movements. Results showed that the patterns of muscular activation were linked not only to the speed and trajectory of the movements of the skull, but also to the kinematics of the motion occurring across different parts of the cervical column.

Animals

Electromyographic studies of neck muscles in the intact cat. II. Reflexes evoked by muscle nerve stimulation.

Short-latency reflexes were studied in the neck muscles of four alert cats following electrical stimulation of nerves supplying biventer cervicis (BC), splenius (SP) or rectus capitis posterior (RCP). Reflexes were assessed by comparing levels of EMG activity of muscles before and after each stimulus, as the cats lapped milk, licked their paws or walked on a treadmill. When BC or SP nerves were stimulated at 1.5-4 times threshold (T) for their motor axons, no short-latency heteronymous reflexes could be identified in most neck muscles. However, stimulation of RCP nerves produced inhibitory effects as early as 3-4 ms in the ipsilateral BC, CM, and SP muscles and 6 ms in contralateral BC. At stimulus strengths above 4xT, a more complex pattern of inhibitory or excitatory effects was observed in CM, SP and the intervertebral muscle spinalis dorsi. The reflex effects were attenuated or abolished by partial or complete C1 dorsal rhizotomy (2 cats). Cervicocollic reflex data may need to be reevaluated to consider the possible effects of disinhibition rather than excitation in short-latency reflex pathways.

Afferent Pathways

Effect of velocity and mechanical history on the forces of motor units in the cat medial gastrocnemius muscle.

1. Two fundamental aspects of the dynamic behavior of motor units of the cat medial gastrocnemius (MG) muscle were measured. Force-velocity (FV) relationships were measured with the use of constant velocity shortening and lengthening movements. Effects of mechanical history were assessed via comparisons of forces immediately after or during slow movements with standard isometric forces. Isometric force-length (FL) relations were also measured, and the effect of different stimulation rates on both FV and FL data was assessed. 2. Prior or concurrent movement greatly potentiated motor-unit force, but this movement potentiation was highly dependent on the amplitude of the unit's force. The smallest twitch forces of type S units (< 10 mN) were potentiated more than threefold, but no potentiation occurred for unit forces > 200 mN. It was tentatively concluded that movement potentiation may play little role in normal movements because it does not occur at forces > 1% of maximal isometric force of the MG. 3. During shortening, the normalized FV relations of type S units were relatively steeper than those of type FR or FF units. For lengthening, there was no evident relation between FV steepness and motor-unit type. 4. Stimulation rate affected both the FV and FL relationships of the motor units. The peak of the FL relationship (Lo) clearly shifted to shorter muscle lengths as stimulation rate was increased. The steepness of the FV relationship for shortening was decreased by increasing stimulation rate, but this effect was modest. 5. The shift in motor-unit Lo and the differences in motor-unit FV relationships were hypothesized to play significant roles during normal motor behavior. Realistic computer simulations of FL and FV functions for a population of motor units undergoing normal steady-state recruitment and rate modulation supported these hypotheses. As the level of simulated neural drive increased, the population Lo shifted to considerably shorter lengths, and the normalized FV function became much less steep. The significance of these results for models of muscle are discussed.

Animals

Distribution of motoneurons supplying cat sartorius and tensor fasciae latae, demonstrated by retrograde multiple-labelling methods.

Sartorius (SART) and tensor fasciae latae (TFL) in the cat hindlimb are functionally heterogeneous muscles with regions that differ in their skeletal actions and electromyographic recruitment during normal activity. The topographical organization of motoneurons supplying different regions of SART or TFL has been investigated by exposing cut nerve branches supplying different peripheral territories to a combination of retrograde tracers, including Fast Blue (FB), Fluorogold (FG), and horseradish peroxidase (HRP). Motoneurons supplying medial, central, and anterior regions of SART were intermixed extensively throughout a single columnar nucleus located in the ventrolateral part of segments L4 and L5. With this column, motoneurons supplying medial SART tended to lie more rostrally than those supplying anterior regions, but the gradient was modest and showed some cat-to-cat variation. Two major branches entered anterior SART at different proximodistal levels. When these two branches were exposed to different tracers, most motoneurons contained a single tracer; only a few double-labelled cells were apparent. The labelling suggests that anterior SART may contain two separate, in-series divisions of motor units. In TFL, motoneurons supplying nerve branches to posterior, central, and anterior parts of the muscle were intermingled indiscriminately in a single ventrolateral cell column in L6 and rostral L7. These results suggest that topographical organization in lumbar motor nuclei does not always reflect the highly ordered biomechanical and functional specialization evident in the peripheral organization of the muscles themselves.

Amidines

Functionally complex muscles of the cat hindlimb. I. Patterns of activation across sartorius.

The cat sartorius (SA) can be divided functionally into an anterior (SAa), knee extensor portion and a medial (SAm), knee flexor portion; it can be further subdivided anatomically by multiple nerve branches into parallel longitudinal columns that terminate in a distributed insertion at the knee with a continuous range of moment arms. Thus, SA may be controlled by a discrete number of motoneuron task groups reflecting a small number of central command signals or by a continuum of activation patterns associated with a continuum of moment arms. To resolve this question, the activation patterns across the width of the SA were recorded with an electrode array during three kinematically different movements--treadmill locomotion, scratching and paw shaking, in awake, unrestrained cats. Uniformity of activation along the longitudinal axis was also examined because individual muscle fibers do not extend the length of the SA. In addition, the cutaneous reflex responses were recorded throughout all regions of the SA during locomotion. Two fascial surface-patch arrays, each carrying 4-8 pairs of bipolar EMG electrodes, were sutured to the inner surface of the SA, one placed proximally and the other more distally. Each array sampled separate sites across the anterior to medial axis of SA. During locomotion, two basic EMG patterns were observed: the two burst-per-step-cycle pattern typical of SAa and the single burst pattern typical of SAm. There was an abrupt transition in the pattern of activation recorded in the two parts of SA during locomotion, and no continuum in the activation pattern was observed. Stimulation of both sural and saphenous cutaneous nerves during locomotion produced reflex responses that were uniformly distributed throughout SA, in contrast to the regional differences noted during unperturbed walking. Similarly, during scratching and paw shaking all parts of the SA were active simultaneously but with regional differences in EMG amplitude. The abrupt functional border between SAa and SAm coincided with the division of the SA into a knee flexor vs. a knee extensor. In all cases, the quantitative and qualitative differences in SAa and SAm EMGs were uniformly recorded throughout the entire extent of SAa or SAm; i.e., there was no segregation of activity within either SAa or SAm. Furthermore, the time course of EMG from each proximal recording site was nearly identical to the corresponding distal site, indicating no segregation of function along the longitudinal axis of SA. These results indicate that SAa and SAm constitute the smallest functional modules that can be recruited in SA.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Functionally complex muscles of the cat hindlimb. II. Mechanical and architectural heterogenity within the biceps femoris.

The goal of this study was to analyze the architecture of the cat biceps femoris (BF), a multifunctional hamstring muscle, and to evaluate the relationships between muscle architecture, limb position, and muscle function during natural movement. The BF muscle consists of three neuromuscular compartments: anterior (BFa), middle (BFm) and posterior (BFp). Each compartment is innervated by a separate nerve branch. Nerve branch stimulation and 2-dimensional surface EMG recordings showed that individual compartment territories were discrete and non-overlapping with well-defined borders. Comparisons of the three compartments revealed consistent differences in architecture, relationship to the skeleton, and function. The BFa crossed only the hip joint and appears to function as a pure hip extensor. The BFm had equal lever arm lengths to the hip and knee joints, appears to function as a hip extensor, and may contribute to knee flexion or femoral rotation. The BFp had a greater lever arm to the knee, functions as a knee flexor, and may contribute to hip extension, femoral rotation or ankle extension. Measurements of individual fascicles from the three compartments revealed a surprising range of lengths, 3.3-12.0 cm. Microdissection of gold-stained tissue showed that fascicles from all compartments were comprised of interdigitated, short fibers (range: 0.6-5.0 cm; average 2.14 cm) arranged in-series in fascicles, running parallel to the origin-insertion axis of each muscle compartment. In regions of fiber interdigitation, the fiber endings were round and tapered (taper lengths: 1-11 mm) although flat, tapering endings like ribbons were occasionally found. As hip and knee joint angles were varied over physiological ranges corresponding to minimal to maximal muscle length, fascicles of the three compartments changed length disproportionately. Long BFa fascicles maximally lengthened 10-18%, consistent with in vivo length measures during treadmill locomotion. However, the long BFp fascicles lengthened 25-45%, and the relatively short fascicles near the BFm/BFp border maximally lengthened 45-53%. How do these unexpectedly large length changes affect sarcomere lengths? Using laser diffraction to measure sarcomeres, static fascicle and sarcomere lengths were compared in muscles that went into rigor mortis after fixing the hip and knee joint angles. Sarcomeres within the short BFm/BFp and long BFp fascicles consistently lengthened proportionately less than the whole fascicle. It remains to be determined how and where the fascicle length changes are dissipated in the connective tissue between the interdigitated muscle fibers and whether such a series-compliance operates during the large excursions over which this muscle normally works.

Animals

Functionally complex muscles of the cat hindlimb. IV. Intramuscular distribution of movement command signals and cutaneous reflexes in broad, bifunctional thigh muscles.

Similarities between the muscle synergies associated with the flexion reflex and locomotion in reduced preparations have suggested that spinal circuits subserving these two motor tasks might share common interneurons. To test this hypothesis in functionally complex muscles, we studied the interaction between low-threshold cutaneous afferents and the locomotor central pattern generator (CPG) during treadmill locomotion in awake, intact cats. Electrical stimuli were delivered via implanted nerve cuff electrodes at all phases of locomotion, and EMGs were recorded from fourteen intramuscular subregions in eight bifunctional thigh muscles (adductor femoris, biceps femoris, caudofemoralis, gracilis, semimembranosus, semitendinosus, tensor fasciae latae, and tenuissimus). In addition, the EMG patterns recorded during locomotion were compared with those recorded during two other centrally driven rhythmical behaviors, scratching and paw shaking, to determine whether the functional relationships among these intramuscular subregions were fixed or task dependent. Four of the five broad, bifunctional muscles studied (biceps femoris, gracilis, semimembranosus, and tensor fasciae latae) had functional subunits that could be differentially activated in one or more of the three movements studied; adductor femoris was consistently uniformly activated despite its distributed skeletal attachments. The pattern of recruitment of the intramuscular functional subunits was movement-specific. The locomotor CPG and cutaneous reflex pathways both similarly subdivided some bifunctional muscles, but not others, into intramuscular subregions. The results of the present study confirm that some combinations of muscle subregions and cutaneous nerves constitute simple reciprocal categories of flexors and extensors, as described originally by Sherrington (1910). "Typical" low threshold excitatory or inhibitory reflex responses were produced in muscles or muscle subregions that were recruited as "net" flexors of extensors, respectively. However, muscles with complex activation patterns during walking often had very individualized, complex reflex responses during locomotion that did not conform to the background locomotion synergies. All of the reflex responses observed were mediated by low threshold cutaneous afferents. These data indicate that there are multiple, low threshold, excitatory and inhibitory cutaneous reflex pathways that have highly specialized connections with flexor and extensor muscles and even their intramuscular subregions. It is also clear that the premotoneuronal circuits mediating these cutaneous reflex effects are not necessarily synonymous with those of the locomotor CPG. These two systems do interact powerfully, however, suggesting some convergence. The nature of the convergence between the CPG and the many independent subsets of spinal interneurons mediating cutaneous reflexes is specialized and muscle subregion-specific.

Afferent Pathways

Functionally complex muscles of the cat hindlimb. V. The roles of histochemical fiber-type regionalization and mechanical heterogeneity in differential muscle activation.

Several cat hindlimb muscles that exhibit differential activation (activity that is restricted to a specific region of muscle) during natural movements were studied to determine the possible roles of 1) non-uniform distribution of histochemically-identified muscle fiber-types (semitendinosus, ST; tibialis anterior, TA) or 2) mechanical heterogeneity (biceps femoris, BF; tensor fasciae latae, TFL). Using chronic recording techniques, electromyographic (EMG) activity was recorded from multiple sites of each muscle during treadmill locomotion, ear scratch, and paw shake. Standard histochemical analysis was performed on each muscle to determine fiber-type distribution. The histochemically regionalized muscles (ST and TA) were differentially active during slow locomotion; the deep regions (high in type I [SO] fibers) were active, but the superficial regions (high in type IIB [FG] fibers) were inactive. Vigorous movements (fast locomotion, ear scratch, paw shake) produced additional, synchronous activation of the superficial regions. In all movements, ST and TA activation patterns were consistent with the existence of identically timed synaptic inputs to all motoneurons within each motoneuron pool, resulting in an orderly recruitment of each whole pool. The differential activation recorded from ST and TA during slow locomotion was presumably a consequence of the non-uniform distribution of the different muscle fiber types. In contrast, differential activation of the histochemically nonregionalized, mechanically heterogeneous muscles (BF and TFL) resulted from non-synchronous activation of different muscle regions. The selective activation of BF or TFL compartments was indicative of differential synaptic inputs to, and selective recruitment of, subpopulations of the motoneuron pool, with each motoneuron subpopulation exclusively innervating physically separate regions of the muscle consistent with the regions defined by the neuromuscular territories of the major nerve branches supplying each muscle. Individual neuromuscular compartments of BF and TFL differ in their mechanical arrangements to the skeleton and in their contribution to mechanical action(s) at the hip and knee joints. Selective neural activation of mechanically distinct compartments within a mechanically heterogeneous muscle can provide highly advantageous mechanical "options" for animals that perform kinematically diverse movements. With regard to EMG recording techniques, the results of this study emphasize the need for carefully chosen EMG sampling sites and the value of knowing the muscle histochemistry, neuromuscular and musculoskeletal anatomy and possible mechanical functions prior to recording EMG.

Adenosine Triphosphatases

Injectable microstimulator for functional electrical stimulation.

A family of digitally controlled devices is constructed for functional electrical stimulation in which each module is an hermetically sealed glass capsule that is small enough to be injected through the lumen of a hypodermic needle. The overall design and component characteristics of microstimulators that receive power and command signals by inductive coupling from a single, externally worn coil are described. Each device stores power between stimulus pulses by charging an electrolytic capacitor formed by its two electrodes, made of sintered, anodised tantalum and electrochemically activated iridium, respectively. Externally, a highly efficient class E amplifier provides power and digitally encoded command signals to control the amplitude, duration and timing of pulses from up to 256 such microstimulators.

Electric Stimulation

Issues in cochlear prosthetics from an international survey of opinions.

Cochlear prostheses are beginning to be implanted regularly to restore hearing in profoundly deaf patients, but there is little agreement on the relative merits of the many different designs and rehabilitative procedures. We report on the responses to a technology assessment questionnaire that was sent to 120 researchers and clinicians worldwide who have been at the forefront of research in this field.

Attitude of Health Personnel

Morphometry of the human thigh muscles. A comparison between anatomical sections and computer tomographic and magnetic resonance images.

The present study examined the relative accuracy and precision of MR and CT procedures for determining the CSA of individual muscles from the human thigh. Serial AN, CT and MR cross-sections were obtained from three cadaveric lower limbs. The MR measurements provided accurate and precise values for the CSAs of most thigh muscles, generally within +/- 7.5% of the AN standard. In contrast, CT tended systematically to overestimate the AN CSA by 10-20%. Retest procedures indicated that highly reliable measurements could be obtained from both MR and CT images. However, subjective interpretations of boundaries between closely apposed muscle bellies, particularly for muscles with more than one head, were necessary for resolving entities in the imaging records and this decreased the relative accuracy of MR and CT measures. Interestingly, MR records demonstrated an incomplete septum between vastus lateralis and vastus intermedius more prominently than AN cross-sections. The novel cross-validation procedures used in this study also highlighted several system-based errors in the MR records that, if undetected and left uncorrected, would have seriously biased the morphometric data obtained with this technique. In general, MR provides high resolution images of the human thigh muscles which may be used to obtain valid measures of the CSA of these structures.

Cadaver

Monosynaptic and dorsal root reflexes during locomotion in normal and thalamic cats.

1. In normal and thalamic walking cats electrical stimulation of muscle nerves via chronically implanted electrodes produced electromyographic (EMG) and neurographic responses that were modulated in amplitude depending on the phase of the step cycle. These responses were examined for possible indications of effects of primary afferent depolarization (PAD) during stepping. 2. Monosynaptic reflexes (MSRs) produced by stimulating the lateral gastrocnemius (LG) and medial gastrocnemius (MG) nerves were recorded as EMGs in MG or LG muscles during treadmill locomotion in normal cats. These heteronymous MSR responses were greatest during the stance (extensor) phase. 3. In the same animals, after decerebration, similar modulation of the heteronymous ankle extensor MSRs occurred during spontaneous locomotion with the use of the same stimulus and recording sites. 4. In both normal and thalamic cats the amplitude of neurogram responses recorded from LG or MG nerve after stimulation of the other muscle nerve varied with phase of stepping but did not parallel the variations of the MSR measured as EMG amplitude in the same muscle. The nerve responses were largest during the flexion phase of the step cycle and had a calculated central latency of 0.6-1.0 ms. These are interpreted as arising from antidromic activity in large-caliber afferent nerve fibers (i.e., dorsal root reflexes). 5. Spontaneous antidromic activity in severed L7 dorsal rootlet fibers to triceps surae was observed in the thalamic cats during episodes of locomotion and was closely correlated with flexion phase EMG activity in semitendinosus, a bifunctional muscle. 6. In decerebrate cats, dorsal root reflexes (DRRs) in severed filaments of L4-L7 dorsal roots were produced by stimulation of saphenous and posterior tibial nerves. These DRRs were always smaller during locomotion than during rest and were smallest during the flexion phase. 7. The short-latency antidromic activity produced in muscle nerves by stimulating heteronymous muscle nerves thus appears to be a DRR produced in Group I terminal arborizations that are depolarized close to threshold during the flexion phase. Such PAD could account for changes in the MSR that do not always parallel the levels of recruitment of the motor pools as manifest by background EMG amplitude.

Animals