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PubMed1980

Response of forelimb guard hair afferent units to air-jet stimulation of entire receptive field.

1. In anesthetized, immobilized cats, individual axonal responses to movement of forelimb G hairs were recorded in the ipsilateral cuneate fasciculus at C1-C2 with a glass-insulated tungsten microelectrode. Several test criteria were used to identify primary afferent axons among the recorded neurons. 2. The output discharge in the parent sensory axon during continuous air-jet stimulation of the entire receptive field was irregular and was usually maintained. Criteria for statistical stability were introduced, permitting an analysis of the response as a stable point process. 3. The main interspike interval during the response to maximal air-jet stimulation ranged in different units from 4 to 100 ms, with a transition zone near 20 ms (50/s) between short and long mean interval units. Long and short mean interval units tended to be related to G2- and G1-type hair receptors, respectively. 4. In both short and long mean interval units, the interspike-interval distribution (IID) had a dead time (usually 2-4 ms), a sigmoidal rising limb, a single mode, and a monotonic falling limb, a single mode, and a monotonic falling limb. In short mean interval units, the decrement of the falling limb was described by a single exponential. 5. The value of the standard deviation of the IID was a little less than that of the mean interval minus dead time, the inequality probably resulting from the finite duration of the rising limb and, in long mean interval units, the nonexponential falling limb. 6. Joint interval analysis revealed that successive intervals were independent except in short mean interval units, where the shortest intervals tended to be followed by intervals shorter than the mean. 7. The expectation density function did not reveal any periods in the output discharge. In short mean interval units, a constant mean (planteau) level was attained after a finite rise time and usually one or more overshooting-undershooting sequences. In long mean interval units, the expectation density function monotonically rose to the plateau level. 8. Reducing the air-jet intensity caused an increase in mean interval of discharge and in standard deviation of the IID and slowed the rise time of the expectation density function to the plateau level. 9. Similarities and differences between the whole-field response to air-jet stimulation and the Poisson process are discussed.

Action Potentials↗
PubMed1983

Dynamics of neck-to-forelimb reflexes in the decerebrate cat.

We have studied the neck-to-forelimb reflex evoked by head rotation around the longitudinal axis (roll) in the long and medial heads of triceps brachii of decerebrate, acutely labyrinthectomized cats. Reflexes were measured by recording mass electromyogram (EMG). As expected from the work of others, they were reciprocal in the two limbs, with excitation in the limb toward which the chin rotates. The reflex was sufficiently linear for a sinusoidal analysis. Although there was sometimes adaptation at stimulus frequencies of 0.1 Hz and below, response phase at these frequencies was usually in phase with position, and gain was flat. At higher frequencies there was some sensitivity to the velocity of the stimulus: gain increased with a slope of 10 dB/decade and phase advanced in some cats but not in others. Gain at low frequencies of head rotation, expressed as percent modulation of EMG, was typically 1%/deg or less. Reflexes evoked by head rotation in triceps and in the neck extensor splenius capitis have different dynamics. It remains to be determined whether this difference is due to activation of different receptors. We compared the dynamics of roll reflexes evoked by stimulation of neck receptors with those of vestibular reflexes evoked by tilt of the whole animal (23). Taking into account dynamics and gain, the two reflexes should cancel at low frequencies, as predicted by others. Above 0.2 Hz, cancellation becomes less effective.

Animals↗
PubMed1985

Patterns of facilitation and suppression of antagonist forelimb muscles from motor cortex sites in the awake monkey.

Patterns of excitatory and inhibitory effects were produced in antagonistic forelimb muscles by single intracortical microstimuli (S-ICMS) applied to motor cortex sites in macaque monkeys performing ramp-and-hold wrist movements. Stimulus-triggered averages (stimulus-TAs) of rectified electromyographic (EMG) activity revealed poststimulus facilitation and/or suppression in identified flexor and extensor muscles of the wrist and fingers. At 22 cortical sites the action potentials of single cells were also recorded and used to compute spike-triggered averages (spike-TAs) of covarying muscles. The set of muscles activated during the movement in which the cell was active are referred to here as "agonists"; those muscles active during wrist movement in the opposite direction are called "antagonists." (At sites where cells were not isolated the muscles showing poststimulus facilitation were called agonists.) Poststimulus effects in agonist muscles typically consisted of facilitation in a subset of the agonists. For 48 sites from which poststimulus effects were tested on both flexors and extensors, the following combinations of effects were observed: 1) pure facilitation of agonist muscles with no effect on antagonists; 2) facilitation of both agonists and antagonists; 3) facilitation of agonist muscles with reciprocal suppression of antagonists; 4) "mixed" facilitation and suppression of synergist muscles; and 5) pure suppression of some muscles with no effect on their antagonists. The suppression effects appeared most commonly in flexor muscles; conversely, facilitation was generally stronger in extensors. Cortical sites eliciting pure suppression of flexor muscles with no facilitation of extensor muscles were found in two monkeys. These purely suppressive effects were observed not only in stimulus-TAs but also in spike-TAs computed from single cells at these sites. Some of these cells increased their activity during wrist extension (but had no detectable effect on the extensor muscles); others discharged during flexion. Several observations suggest that the cortically evoked suppression is mediated by polysynaptic relays. The mean onset latency of the postspike suppression (7.4 ms) produced by inhibitory cells was longer than the mean onset latency of postspike facilitation (6.7 ms) produced by CM cells. Similarly, the mean onset latency of poststimulus suppression (8.9 ms) was longer than that of poststimulus facilitation (8.0 ms). Moreover, suppression was usually weaker than facilitation in the spike-TAs, as well as in stimulus-TAs compiled for the same stimulus intensity.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1986

Spatial organization of neck and vestibular reflexes acting on the forelimbs of the decerebrate cat.

EMG recording was used to study the spatial organization of vestibular and tonic neck reflexes acting on forelimb and shoulder muscles of the decerebrate cat. Neck reflexes were studied in preparations with intact labyrinths as well as those with acute or chronic labyrinthectomies. Reflexes were described by response vectors whose orientation component is aligned with the optimal excitatory direction of tilt or head rotation. A muscle's vector orientation remained reasonably stable over a period of hours, although there was sometimes drift at the beginning or end of an experiment. Orientation of muscle response vectors did not change systematically with stimulus frequency of 0.05-2.0 Hz. For vestibular reflexes this is so, although their dynamics are consistent with convergent input from semicircular canals and otolith organs. Regardless of the preparation, a consistent reflex pattern emerged. Vestibular reflexes are characterized by response vector orientation near ear-down roll. Neck vector orientation lies in the opposite direction from the vestibular vector but typically lies further from the roll plane: Nose-up pitch is excitatory for the shoulder muscles supra- and infraspinatus, and for the medial and lateral heads of triceps, whereas nose-down pitch excites the long head of triceps. Our results generally agree with the pattern proposed by Roberts (28) for neck reflexes but disagree in part with his proposed pattern of vestibular reflexes; we did not see the expected consistent excitation by nose-down pitch.

Animals↗
PubMed1990

Efferent neurons and suspected interneurons in S-1 forelimb representation of the awake rabbit: receptive fields and axonal properties.

1. Receptive-field properties of antidromically identified efferent neurons within the cutaneous forelimb representation of primary somatosensory cortex (S-1) were examined in fully awake rabbits. Efferent neurons studied included callosal neurons (CC neurons, n = 52), ipsilateral corticocortical neurons (C-IC neurons, n = 48) that project to or beyond the second somatosensory cortical area (S-2), and corticofugal neurons of layer 5 (CF-5 neurons, n = 97) and layer 6 (CF-6 neurons, n = 59) that project to and/or beyond the thalamus. 2. An additional class of neurons was studied that was not activated antidromically from any stimulus site, but which responded synaptically to electrical stimulation of the ventrobasal (VB) thalamus with a burst of three or more spikes at frequencies of 600 to greater than 900 Hz. Most of these neurons also responded synaptically to stimulation of S-2 and the corpus callosum. The action potentials of these neurons were much shorter (mean = 0.45 ms) than those of efferent neurons (mean = 0.95 ms). Such properties have been associated with interneurons found throughout the central nervous system, and these neurons are thereby referred to as suspected interneurons (SINs). 3. CF-5 neurons differed from CC, C-IC, and CF-6 neurons in their spontaneous firing rates, axonal properties, and receptive-field properties. Whereas CF-5 neurons had a mean spontaneous firing rate of 5.5 spikes/s, CC, C-IC, and CF-6 neurons had mean values of less than 1/s. Axonal conduction velocities of CF-5 neurons were much higher (mean = 12.92 m/s) than either CC (mean = 2.15 m/s), C-IC (mean = 1.31 m/s), or CF-6 (mean = 2.53 m/s) neurons. A decrease in antidromic latency (the "supernormal" period) that was dependent on prior impulse activity was seen in the great majority of CC, C-IC, and CF-6 neurons but was either minimal or absent in CF-5 neurons of comparable conduction velocity. A higher proportion of CF-5 neurons (98%) responded to peripheral sensory stimulation than did either CC (75%), C-IC (71%), or CF-6 (51%) neurons. CF-6 and C-IC neurons that did not respond to sensory stimulation had significantly lower axonal conduction velocities and spontaneous firing rates than those that responded to such stimulation. 4. Cutaneous receptive fields were seen in most neurons that could be driven by peripheral stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1997

Effects of inactivating individual cerebellar nuclei on the performance and retention of an operantly conditioned forelimb movement.

These experiments were designed to examine the effects of inactivating separately each of the major cerebellar nuclear regions in cats on the execution and retention of a previously learned, operantly conditioned volitional forelimb movement. The experiments test the postulates that the cerebellar nuclei, and particularly the interposed nuclei, contribute substantially to the spatial and temporal features of the interjoint coordination required to execute the task and that the engram necessary for the retention of this task is not located in any one of the cerebellar nuclei. All cats were trained to perform a task in which they were required to reach for and grasp a vertical bar at the sound of a tone and move the bar to a reward zone through a template consisting of two straight grooves in the shape of an inverted "L." After the task was learned, the effects of inactivating separately each nuclear region (the fastigial, interposed, and dentate nuclei) using muscimol microinjections were determined. Data were analyzed by quantifying several features of the movement's kinematics and by determining changes in the organization of the reaching component of the movement using an application of dimensionality analysis, an analysis that examines the correlation among the changes in joint angles and limb segment positions during the task. The retention of the previously learned task also was assessed after each injection. Injections of each nuclear region affected temporal and spatial features of the learned movement. However, the largest effects resulted from inactivating the interposed nuclei. These effects included an increased length of the reach trajectory, an accentuated deviation of the wrist trajectory from a straight line, cyclic movement of the distal extremity as the target was approached, a difficulty in grasping the bar, altered temporal features of the movement, and a highly characteristic change in the dimensionality measurements. The changes in dimensionality reflected a decreased correlation (linear interdependence) of the joint angular velocities coupled with an increased correlation among the linear velocities of markers located on the joints themselves. Related but less consistent changes in dimensionality resulted from fastigial injections. The motor sequence required to negotiate the template could be executed after the nuclear microinjections, indicating that retention of the motor sequence was not affected by the inactivation of any of the cerebellar nuclei. However, in two of the five animals, some decreases in performance were observed after dentate injection that were not characteristic of changes related to an effect on retention. These data suggest that the cerebellum plays an important role in regulating the consistent, stereotypic organization of complex goal-directed movements, including the temporal correlation among joint angle velocities. The data also indicate that the retention of the task is not dependent on any of the individual cerebellar nuclear regions. Consequently, these structures are unlikely to be critical storage sites for the engram established during the learning of this task.

Adaptation, Physiological↗
PubMed1998

Effects of cerebellar nuclear inactivation on the learning of a complex forelimb movement in cats.

The purpose of this study was to determine the effects of inactivating concurrently the cerebellar interposed and dentate nuclei on the capacity of cats to acquire and retain a complex, goal-directed forelimb movement. To assess the effects on acquisition, cats were required to learn to move a vertical manipulandum bar through a two-segment template with a shape approximating an inverted "L" after the injection of muscimol (saline for the control group) in the interposed and dentate cerebellar nuclei. During training periods, they were exposed progressively to more difficult templates, which were created by decreasing the angle between the two segments of the template. After determining the most difficult template the injected animals could learn within the specified time and performance constraints, the retraining phase of the experiment was initiated in which the cats were required to execute the same sequence of templates in the absence of any injection. This stage of the experiment assessed retention and determined the extent of any relearning required to execute the task at criterion levels. Next, the animals were overtrained without any injection on the most difficult template they could perform. Finally, to determine the effects of nuclear inactivation on retention after extensive retraining, their capacity to perform the same template was determined after muscimol injection in the interposed and dentate nuclei. The findings show that during the inactivation of the dentate and interposed nuclei the animals could learn to execute the more difficult templates. However, when required to execute the most difficult template learned under muscimol on the day after injections were discontinued, the cats had to "relearn" (reacquire) the movement. Finally, when the cerebellar nuclei were inactivated after the animals learned the task in the absence of any injections during the retraining phase, retention was not blocked. The data indicate that the intermediate and lateral cerebellum are not required either for learning this type of complex voluntary movement or for retaining the capacity to perform the task once it is learned. Nevertheless, when the cerebellum becomes available for executing a task learned in the absence of this structure, reacquisition of the behavior usually is necessary. It is hypothesized that the relearning observed after acquisition during muscimol inactivation reflects the tendency of the system to incorporate the cerebellum into the interactions responsible for the learning and performance of a motor sequence that is optimal for executing the task.

Animals↗
PubMed1998

Topographic position of forelimb motoneuron pools is conserved in vertebrate evolution.

The neuromotor conservatism hypothesis predicts that neuromotor patterns in homologous tetrapod muscles are conserved evolutionarily despite the musculoskeletal modifications of vertebrate limbs. A complete description of the anatomical organization of the neurons innervating homologous limb muscles is a prerequisite to any test of the neuromotor conservatism hypothesis. This study uses the retrograde neuronal tracer WGA-HRP to selectively label the motor neuron pools of seven homologous forelimb muscles in mice (Mus musculus) and iguanas (Iguana iguana): Mm. pectoralis, spinodeltoideus, biceps brachii, lateral and long heads of triceps brachii, and the supraspinatus and infraspinatus (in mice) or their reptilian homolog, the supracoracoideus (in iguanas). In vertebrates, motoneurons are arranged in longitudinal columns of cells in the ventral horn of the spinal cord. Mouse motor pools average 1,952 microns in length, except the pectoralis pool which averaged 2,949 microns in length. Iguana pools average 3,196 microns in length. The number of neurons per pool ranged from 70-199 in mice and from 58-114 neurons in iguanas. In both iguanas and mice the motor pools for the spinodeltoids, biceps, and the supracoracoideus (or its mammalian homologs) lie anterior to the pectoralis and triceps motor pools. In the transverse plane, the pectoralis pool lies medial to those of the triceps. The pools of the biceps and spinodeltoids are located dorsal and lateral to those of the pectoralis and supracoracoideus (or its homologs in mammals). The resulting motor pool maps support the hypothesis that the anatomical organization of motoneurons in ancestral reptiles has been retained in these two tetrapod descendents.

Animals↗
PubMed1989

Failure to perform visually triggered forelimb movements after unilateral nigrostriatal damage in monkeys.

Unilateral injections of 6-hydroxydopamine into the substantia nigra of monkeys disrupted the initiation of learned movements performed with the contralateral forelimb. Failure to respond was observed in two types of behavioral tasks. The one involved a hand movement directed towards a target and the other, hand withdrawal without any spatial requirements at the end of the movement. The results suggest that the ability to perform the required response was recovered more quickly in the case of the task involving visually controlled movement.

Afferent Pathways↗
PubMed1993

In vivo tendon forces in the forelimb of ponies at the walk, validated by ground reaction force measurements.

The load distribution over tendinous structures in the equine forelimb was studied by computing forces from in vivo signals of implanted liquid-metal strain gauges in 5 ponies. For validation, these tendon forces were converted to joint moments, which were summed and compared to the calculated joint moments caused by the ground reaction force. Mean peak forces per kilogram body weight (n = 5) amounted to 5.2 N/kg for the superficial digital flexor tendon, 3.8 N/kg for the deep digital flexor tendon, 7.3 N/kg for the distal accessory (check) ligament and 8.4 N/kg for the third interosseous muscle (suspensory ligament). The maximal moment exerted by the tendons about the fetlock joint differed 11 +/- 7% (average +/- SD, n = 5) from the maximal ground reaction force moment, which difference amounted to 17 +/- 15% for the coffin joint moments. These differences appeared to result to a substantial extent from errors in the moment arms. Therefore, the computed tendon forces were considered to be sufficiently reliable.

Animals↗
PubMed1993

Shortening of the forelimb in the horse during the stance phase.

Compressions in the forelimb of a horse trotting at 3.7 m/s were studied using high-speed cinematography. From a diagram of shortenings it has been concluded that successive loadings in the longitudinal direction from below signify longer impact time and smaller impact forces compared to a theoretically stiff limb. It is proposed that the limb takes up the impact force passively until the onset of loading at the elbow joint, i.e. 30 ms following first contact.

Acceleration↗
PubMed1974

Retardation of progressive forelimb regeneration by treatment with neuropharmacological agents.

Intraperitoneal administration of hormones or of certain neuropharmacological agentsdoes not affect initial blastema formation in adult newts (Notophthalmus viridescens). Hormonal treatment accelerates the onset of the morphogenetic phase of regeneration. In contrast, treatment with chlorpromazine (alone or with growth hormone), guanethidine, or atropine retards or represses the onset of morphogenesis. These results suggest that some interaction between the nervous and endocrine systems is required in promoting the proliferative and the morphogenetic stages of forelimb regeneration.

Animals↗
PubMed2002

SAGE profiling of the forelimb and hindlimb.

A recent study has used serial analysis of gene expression to compare mouse forelimb and hindlimb gene-expression profiles. The method successfully identified known regulators of limb identity and has generated a candidate set of differentially expressed genes that may regulate limb identity.

Animals↗
PubMed2003

The zebrafish fgf24 mutant identifies an additional level of Fgf signaling involved in vertebrate forelimb initiation.

The development of vertebrate limb buds is triggered in the lateral plate mesoderm by a cascade of genes, including members of the Fgf and Wnt families, as well as the transcription factor tbx5. Fgf8, which is expressed in the intermediate mesoderm, is thought to initiate forelimb formation by activating wnt2b, which then induces the expression of tbx5 in the adjacent lateral plate mesoderm. Tbx5, in turn, is required for the activation of fgf10, which relays the limb inducing signal to the overlying ectoderm. We show that the zebrafish fgf24 gene, which belongs to the Fgf8/17/18 subfamily of Fgf ligands, acts downstream of tbx5 to activate fgf10 expression in the lateral plate mesoderm. We also show that fgf24 activity is necessary for the migration of tbx5-expressing cells to the fin bud, and for the activation of shh, but not hand2, expression in the posterior fin bud.

Animals↗
PubMed2006

Prdm1 acts downstream of a sequential RA, Wnt and Fgf signaling cascade during zebrafish forelimb induction.

Vertebrate limb induction is triggered in the lateral plate mesoderm (LPM) by a cascade of signaling events originating in the axial mesoderm. While it is known that Fgf, Wnt and retinoic acid (RA) signals are involved in this cascade, their precise regulatory hierarchy has not been determined in any species. tbx5 is the earliest gene expressed in the limb bud mesenchyme. Recently, another transcription factor, Prdm1, has been shown to be crucial for zebrafish forelimb development. Here, we show that Prdm1 is downstream of RA, Wnt2b and Tbx5 activity. We find that RA activity, but not Fgf signaling, is necessary for wnt2b expression. Fgf signaling is required for prdm1 expression in the fin bud, but is not necessary for the initiation of tbx5 expression. We propose a model in which RA signaling from the somitic mesoderm leads to activation of wnt2b expression in the intermediate mesoderm, which then signals to the LPM to trigger tbx5 expression. tbx5 is required for Fgf signaling in the limb bud leading to activation of prdm1 expression, which in turn is required for downstream activation of fgf10 expression.

Animals↗
PubMed1994

Multiple developmental defects in Engrailed-1 mutant mice: an early mid-hindbrain deletion and patterning defects in forelimbs and sternum.

During mouse development, the homeobox-containing gene En-1 is specifically expressed across the mid-hindbrain junction, the ventral ectoderm of the limb buds, and in regions of the hindbrain, spinal cord, somites and somite-derived tissues. To address the function of En-1 during embryogenesis, we have generated mice homozygous for a targeted deletion of the En-1 homeobox. En-1 mutant mice died shortly after birth and exhibited multiple developmental defects. In the brains of newborn mutants, most of the colliculi and cerebellum were missing and the third and fourth cranial nerves were absent. A deletion of midhindbrain tissue was observed as early as 9.5 days of embryonic development and the phenotype resembles that previously reported for Wnt-1 mutant mice. In addition, patterning of the forelimb paws and sternum was disrupted, and the 13th ribs were truncated. The results of these studies suggest a cell autonomous role for En-1 in generation and/or survival of mid-hindbrain precursor cells and also a non-cell autonomous role in signalling normal development of the limbs and possibly sternum.

Animals↗
PubMed1995

Postaxial polydactyly in forelimbs of CRABP-II mutant mice.

The cytoplasmic retinoic acid (RA)-binding protein CRABP-II is expressed widely throughout early morphogenesis in mouse embryo, but its expression becomes more restricted as organogenesis progresses. CRABP-II expression remains strong in the developing limb bud suggesting a role for this protein in limb patterning. Here, we show that the CRABP-II promoter can direct expression of a lacZ transgene in a specific posterior domain during limb bud development. In order to investigate in more detail the role played by CRABP-II in RA signal transduction, we have also generated mice homozygous for a null mutation of this gene. CRABPII-/- mice are viable and fertile but show a developmental defect of the forelimb, specifically an additional, postaxial digit. This digit is generally, but not exclusively, limited to a single forepaw of an individual animal. The penetrance of the phenotype varies according to the genetic background, occurring most frequently on the inbred 129Sv background (50%), less frequently on the C57Bl/6 background (30%) and rarely on the outbred CD1 background (10%). This developmental abnormality implies a role for CRABP-II in normal patterning of the limb.

Animals↗
PubMed1987

Dorsal root ganglia grafts stimulate regeneration of denervated urodele forelimbs: timing of graft implantation with respect to denervation.

Amphibian forelimb regeneration is a nerve-dependent process; nerves presumably release one or more neurotrophic factors that stimulate blastema cell division. To date several candidate molecules/factors have been shown to stimulate macromolecular synthesis and/or mitosis but sustained cell cycle activity and blastema development have not been achieved. Because dorsal root ganglia (DRG) implants are capable of promoting regeneration of denervated adult newt limbs (Kamrin & Singer, 1959), we have evaluated the DRG stimulation of regeneration in denervated limbs of adult newts and larval axolotls; two alternative timing strategies were tested as a step toward defining bioassay parameters that best reflect neurotrophic activity. The frequency of regeneration in denervated adult newt limbs was compared after providing DRG before or at the time of denervation (to maintain neurotrophic and cell cycle activity) versus DRG implantation at various postdenervation times (to resupply neurotrophic activity and restimulate suppressed cell cycle activity). The results show that denervated adult newt limbs regenerated most frequently using the maintenance strategy, but as the denervation interval was extended in the restimulation strategy, the frequency of regeneration declined. Larval axolotl limbs responded positively in both maintenance and restimulation DRG-grafting protocols. These results suggest that the efficacy of DRG stimulation of regeneration in adult newts was related to the relative number of blastema cells present at the time of denervation and the proliferative status of the blastema cells; bioassays with denervated adult newt limbs should be designed with these constraints in mind. Because such constraints are not as problematic with the larval axolotl, this species may provide the best opportunity for further defining bioassay parameters related to the neurotrophic stimulation of regeneration.

Animals↗