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Integration in descending motor pathways controlling the forelimb in the cat. 4. Corticospinal inhibition of forelimb motoneurones mediated by short propriospinal neurones.

A previously described inhibitory trisynaptic cortico-motoneuronal pathway (Illert et al., 1976a) was analysed in order to identify the two relay stations. Intracellular recording was made from motoneurones to elbow muscles. Corticospinal fibres were stimulated in the contralateral medullary pyramid. Pyramidal IPSPs were abolished by a transection of the corticospinal tract in C2 but remained after a corresponding lesion in C5. After a C5 lesion pyramidal volleys facilitated transmission in the Ia inhibitory pathway with a time course suggesting disynaptic excitatory action on the Ia inhibitory interneurones. The trisynaptic pyramidal IPSPs were depressed by volleys in the appropriate recurrent motor axon collaterals as would be expected if these IPSPs were mediated by Ia inhibitory interneurones. It is concluded that trisynaptic cortico-motoneuronal inhibition is evoked by consecutive activation of propriospinal neurones in C3--C4 and segmental Ia inhibitory interneurones.

Animals

Immediate and delayed changes of rat motor cortical output representation with new forelimb configurations.

These experiments examined motor cortical representation patterns after forelimb postural adjustments in rats. The experiments tested the hypothesis that postural adjustments that stretch muscles that are most strongly activated from the primary motor cortex (MI) enlarge their cortical representation. Intracortical electrical stimulation within MI, forelimb muscle activity and movements, and vibrissa movements were used to evaluate the border between the MI forelimb and vibrissa representations before and after forelimb position changes in anesthetized adult rats. The forelimb was originally maintained in retraction (wrist extension and elbow flexion) and then changed to protraction (wrist flexion and elbow extension). Movements and forelimb EMG evoked by electrical stimulation were evaluated during this period (up to 3 hr) through a set of four electrodes implanted in layer V of MI. Changing the forelimb configuration had both immediate and delayed effects on forelimb muscle activity evoked from MI. At some sites, the magnitude of evoked forelimb muscle activity immediately increased with forelimb protraction. At one-quarter of all sites, forelimb muscle activity was evoked where it was not previously detected following an average delay of 22-31 min after forelimb protraction. This change can be interpreted as an expansion of the forelimb area into the vibrissa representation. These data further support the hypothesis that motor cortical representations are flexible and show that sustained changes in somatic sensory input to MI are sufficient to reorganize MI output.

Animals

Coordination of movements of the kindlimbs and forelimbs in different forms of locomotion in normal and decerebrate cats.

The coupling of movements of the hindlimbs and forelimbs has been analysed in intact cats stepping overground and on a treadmill and during swimming, and in decerebrate cats stepping on a treadmill, immersed in water('swimming') and stepping suspended in the air. In the different preparations, and under different types of locomotion, two basic patterns of coupling have been observed. Both concern the hindlimb and forelimb of the same side of the body. The first pattern is found in the pacing gait where flexion of the forelimb precedes extension of the hindlimb, measured at the elbow and knee, respectively. The second pattern is typically found in the trot where flexion of the forelimb follows extension of the hindlimb. In decerebrate cats both patterns of coupling remain after bilateral deafferentation of the hindlimbs. In the alternate form of locomotion these patterns of coupling occurs symmetrically on both sides. In the rotatory and transverse gallop (examples of the in-phase form of locomotion) the coupling is asymmetrical: on one side it is comparable to pacing (forelimb flexion precedes hindlimb extension), and on the other side to trotting (forelimb flexion follows extension). These basic patterns of interlimb coordination simplify considerably the problem of neural control of the limbs in locomotion. Obersations of EMGs during the alternative forms of locomotion show that in the pacing type of coupling the extensor EMGs of forelimb and hindlimb overlap, with the hindlimb leading the forelimb by about 10% of a step cycle, while in the trotting type of coupling the forelimb flexor EMGs overlap the hindlimb extensor EMGs, the forelimb flexors leading the hindlimb extensors by about 10% of a step cycle. During acceleration the transition between the two forms of EMG occurs within one or two step cycles, and at some intermediate velocities the EMG coupling springs back and forth between the two different forms. These results further support the hypotesis of two basic forms of interlimb coupling in which long propriospinal pathways probably play a role.

Animals

Functional organization of vestibular and visual inputs to neck and forelimb motoneurons in the frog.

1. Intracellular responses in neck and forelimb motoneurons to electrical stimulation of the vestibular nerve, the optic tectum, and the optic nerve were studied in frog. 2. Stimulation of the anterior branch of the vestibular nerve typically produced EPSPs, bilaterally, in neck, shoulder (DOR), and forelimb extensor (TRI, RAD) motoneurons, and bilateral IPSPs in forelimb adductor (PED) and flexor (ULN, COR) motoneurons. 3. Latencies of PSPs recorded in neck, shoulder, and proximal extensor motoneurons (TRI) were mostly in the disynaptic range, whereas many of those recorded in distal extensor (RAD) and in adductor and flexor motoneurons involved three synapses. 4. Lesion of the vestibulospinal fibers greatly reduced the vestibular nerve-evoked field potentials in the spinal cord and the occurrence of PSPs in forelimb motoneurons. These results as well as the latency measurements suggest that the pathway linking vestibular nerve and forelimb motoneurons mainly consists of vestibulospinal fibers, though involvement of other structures for production of later PSPs could not be completely ruled out. Hemisection of the brain stem at its most caudal level showed that the pathway to the contralateral motoneurons crosses at the level of brain stem as well as in the spinal cord. 5. Stimulation of the optic tectum produced EPSPs, IPSPs, and a mixture of EPSPs and IPSPs in neck, shoulder, and forelimb motoneurons, bilaterally. Most frequently, a combination of an excitation and inhibition was observed. The pathway from the optic tectum to neck and limb motoneurons is at least dysnaptic in nature. 6. Stimulation of the optic nerve produced IPSPs and a mixture of EPSPs and IPSPs in neck and forelimb motoneurons. Impulses originating from the optic nerve descend as far as to lumbar motoneurons producing EPSP-IPSP sequences bilaterally. 7. Interaction studies suggested that the vestibular and optic pathways to neck and forelimb motoneurons are separate from each other so that the site of integration of vestibular and visual input occurs at the level of motoneurons. 8. Evidence for electronic coupling among forelimb motoneurons and electrical synaptic transmission in th pathway linking vestibular nerve and forelimb motoneurons is presented.

Animals

Interaction of transcapillary Starling forces in the isolated dog forelimb.

Three of the four Starling forces were measured in the intact dog forelimb after anesthetization and all four of the Starling forces were measured in the same forelimb which was surgically isolated yet innervated. In the isolated forelimb, isogravimetric capillary pressure (Pci) averaged 15.6 mmHg; colloid osmotic pressure of the plasma proteins (IIp) averaged 19.9 mmHg; mean interstitial fluid pressure (Pif) was +0.4 mmHg, and the average value of interstitial colloid osmotic pressure (IIif) was 4.9 mmHg. Thus the net imbalance in the Starling forces, i.e., (Pci - Pif) - (IIp - IIif), averaged 0.3 mmHg. Furthermore, the value of IIif was consistently decreased after isolation (average decrease of 1.2 mmHg) while Pif was always increased following isolation (average increase of 4.3 mmHg). In addition, it was found that if the forelimb was denervated during isolation, then Pif was increased by an average of 2 mmHg above Pif in the innervated, isolated forelimb. In summary, these studies show that the differences between the intact and isolated forelimb are that Pci averages 10-11 mmHg in the intact forelimb and 15-16 mmHg in the isolated innervated forelimb while interstitial fluid pressure is negative in the intact limb and positive in the isolated limb.

Animals

Functional morphology of forelimb joints in the woolly monkey Lagothrix lagothricha.

This gross anatomical study of embalmed forelimb joints of the South American woolly monkey Lagothrix lagothricha focuses on the problem of determining in osteoligamentous preparations how the disposition of the capsular apparatus and the geometry of the articular surfaces govern the amount and types of movement permitted at a joint, and then correlates these findings with the use of the forelimb in the positional capabilities of captive wooly monkeys observed by this author. Data collection was by dissection, quantitative range of motion studies on osteoligamentous preparations, and by qualitative manipulations of these preparations and of disarticulated bones. Supplemental evidence was obtained from radiographs and from an estimate of angular values of articular surfaces. Presented for the shoulder, elbow, radioulnar, and hand joint complexes are the functional anatomy of the capsular apparatus and articular surfaces, the quantitative range of motion data, and proposed mechanisms of movement that combine the functional morphology with the observed use of the forelimb in activities by the animal. The structurofunctional framework of MacConaill is the basis for the joint analysis. The evidence suggests that the functional anatomy of these joints correlates well with the possible positional capabilities of Lagothrix. The morphology of the capsular apparatus and joint surfaces reflect both the arboreal quadrupedalism and forelimb suspension of the woolly monkey. The positions of maximum congruency and close-pack approximate the forelimb's weight-bearing stance during palmigrade arboreal quadrupedalism. During forelimb suspension, the taut and twisted capsular apparatus of close-packed joints maintains the integrity of forelimb links in tension. This is the first known report of the internal band of the collateral ligament of the proximal interphalangeal joints of the second through fifth digits being incorporated into the terminal tendon of the extensor assembly. This author believes this is a mechanism not only for maintaining the integrity of these joints during forelimb suspensory activities when the body is supported by the middle phalanges, but also for helping to coordinate the flexion-extension actions of the phalanges for smooth grasping and release maneuvers in an arboreal environment. When sufficient comparative data have been collected, such correlations of the functional morphology of joints with positional behavior may be used to postulate positional capabilities of fossil primates.

Acromioclavicular Joint

Labyrinthine influence on cat forelimb motoneurons.

1. Intracellular responses in forelimb motoneurons to electrical stimulation of the whole labyrinth and of individual semicircular canal nerves were studied in decerebrated, unanesthetized cats. 2. Stimulation of the whole labyrinth typically produced EPSPs, usually bilaterally, in forelimb extensor (LON, LAT, MED) and shoulder (SI) motoneurons and bilateral IPSPs in forelimb flexor (BIC) motoneurons. 3. Latencies of PSPs indicated that most of those in extensor motoneurons were trisynaptic and many seen in flexor motoneurons may involve four synapses. 4. In the cells sampled, stimulation of the anterior, horizontal or posterior canal nerves often evoked EPSPs in extensor and IPSPs in flexor motoneurons, both ipsi- and contralaterally. Responses to canal stimulation were weaker and more variable than those to stimulation of the whole nerve. 5. Transection of the MLF in the lower medulla had no effect on PSPs evoked in forelimb motoneurons. Lesions in the medulla in the area of the LVST greatly reduced the occurrence of contralateral EPSPs in extensor and IPSPs in flexor motoneurons. The pathway linking labyrinths to forelimb motoneurons therefore appears to include the LVST. Hemisection shows that the pathway to contralateral motoneurons descends in the cord on the side of the stimulated labyrinth before crossing to influence these cells. 6. Labyrinthine control of forelimb motoneurons is less direct than control of neck and back motoneurons. It is suggested that the interneuron in the pathway to forelimb motoneurons is the site of integration of labyrinthine with other reflexes.

Animals

Reaching behavior in the rat: absence of forelimb peripheral input.

In order to test whether peripheral input from a moving forelimb is essential for reaching in rats, the effects of dorsal rhizotomy C5-Th2 were examined. Rats were trained to reach for a food pellet in the horizontal tube or on a tray. Reaching attempts before and after bilateral forelimb deafferentation were monitored by continual recording using magnetic induction. Deafferented animals were able to initiate and generate the motor program of reaching, but modulation of its ongoing execution was lost. Peripheral input from moving forelimb was necessary for the effective performance of grasping; the duration of the manipulative part of reaching was significantly prolonged and the success of grasping was markedly decreased. Also, the aiming of forelimb was impaired, probably by disturbing of body forelimb postural coordination. No significant changes were found in the execution of forelimb protraction. It is concluded that somesthetic feedback is not required for execution of forelimb protraction, but it is necessary for grasping.

Animals

Organization of the forelimb area in squirrel monkey motor cortex: representation of digit, wrist, and elbow muscles.

The EMG in 8 to 14 hand, forearm, and arm muscles evoked by intracortical electrical stimulation was recorded at 433 sites in layer V in the region of the forelimb area of the primary motor cortex (MI) of three squirrel monkeys during ketamine anesthesia. At each site, the EMG was recorded at movement threshold (T) and at 1.5T and 2T at each site (but less than or equal to 60 microA), and the threshold movement was noted. In the animals examined, the total MI forelimb area identified by movements or EMG occupied about 25 to 35 mm2. At most sites from which a forelimb movement was evoked, EMG activity was evoked in one or more of the recorded muscles. One group of sites located rostrolaterally to the main forelimb area was separated by an intervening zone largely related to the face. The average area from which digit, wrist, elbow, or shoulder movement was evoked at threshold was nearly the same, and their movement thresholds were not significantly different. Average movement thresholds across the anterior-posterior extent of MI were also similar. All muscles recorded could be activated by cortical stimulation. Most commonly more than one muscle was activated from a single site. The highest individual EMG levels were produced at sites from which more than one muscle was activated. These results suggest that small regions of MI influence multiple muscles. Individual muscles were typically activated at multiple, spatially separated locations. For many muscles, increasing the stimulation intensity revealed additional separate areas of activation. Spatial locations of different muscles showed considerable interanimal variation. The size of most muscle representations was relatively large. The smallest representations always included the intrinsic hand muscles and the largest included the proximal muscles. Orderly topographic relationships among forelimb joints or muscles within the MI forelimb area were not apparent. Although distal muscle activation tended to be found posteriorly in the forelimb area and proximal muscles tended to be activated from anterior sites, both could be activated from broadly distributed and overlapping areas. The broad, overlapping nature of the muscle representation supports the concept that a small region of cortex is involved in controlling functional groups of muscles. The intermingling of muscle representations may provide a substrate for local cortical interactions among territories representing various muscle synergies or for changing associations of muscle groups. The representation plan derived from these mappings contains elements of all previously described summaries of MI organization.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

The failure of double-half forelimbs to undergo distal transformation following amputation in the axolotl, Ambystoma mexicanum.

Although capable of initiating early regenerative responses, axolotl forelimb stumps which are composed of double-half limb tissues fail to undergo the events that normally lead to the replacement of missing parts. In the present study, the posterior halves of right forelimbs were exchanged with the anterior halves of left forelimbs, or the dorsal halves of right forelimbs were exchanged with the ventral halves of left forelimbs. Forelimbs were amputated through the graft region 30 days after grafting. Limb stumps bearing double-dorsal, double-ventral or double-posterior tissues either produced hypomorphic regenerates or failed to form any externally visible outgrowth. When the limb stump bore double-anterior tissues, no externally visible structures were formed. Normal and multiple regenerates were never formed by double-half limbs. These results are discussed in terms of the polar coordinate model and suggest that the regeneration blastema requires a complete circumference of positional values in order to complete distal transformation.

Ambystoma

The pathway from Ia forelimb afferents to the motor cortex: a new hypothesis.

Forelimb target-reaching and food-taking in cats depend on different interneuronal circuitry in the spinal cord. On the basis of previous findings regarding the effect of transection of the corticospinal tract in the spinal cord, of high dorsal column (DC) transection, of low pyramidotomy and of pyramidotomy after previous DC transection, it is proposed that the food-taking movement is temporally linked to target-reaching as follows: During target-reaching, the position of the paw is signalled by the pathway from forelimb proprioceptors (mainly Ia) to the motor cortex with a relay in the main cuneate nucleus. The command for food-taking is issued by the motor cortex only when the pathway from the forelimb signals that the paw approaches the target correctly, as may be determined by a comparison of the information from the forelimb with an efference copy of the motor program for target-reaching. The hypothesis is based on previous results regarding the organization of the pathway from the forelimb to area 3a and the motor cortex, and regarding the cortico-cuneate pathway with selective projection from area 3a and motor cortex to the basal caudal part of the cuneate nucleus, where the proprioceptive information from the forelimb is relayed. Results relevant to the present hypothesis regarding responses of precentral neurones during active and passive movements in awake animals are briefly discussed.

Animals

Relationship of intrinsic connections to forelimb movement representations in monkey motor cortex: a correlative anatomic and physiological study.

1. Intracortical microstimulation (ICMS) and horseradish peroxidase (HRP) histochemistry were combined to examine the relationship between intrinsic connections and intracortical microstimulation sites eliciting evoked movements in the forelimb representation of adult macaque monkey motor cortex. 2. The distribution of sites from which stimulation-evoked movements about individual forelimb joints were elicited under anesthesia varied considerably among animals. Identical movements could often be elicited from multiple, noncontiguous sites. 3. After single, small extracellular HRP injections at sites from which thumb movement was evoked, small groups of retrogradely labeled cells and dense patches of axon terminations were found scattered across a wide area of the forelimb representation. Terminal patches were discontinuous and arose from horizontal, intracortical axons. 4. Correlating the HRP labeling with the physiologically defined movement maps revealed a profuse set of intrinsic, bidirectional connections that connect digit representations and representations of movements about the wrist, elbow, and shoulder. 5. HRP injections placed in the forelimb representation close to the physiologically defined face representation resulted in virtually no retrogradely labeled cells or terminal fiber labeling that crossed into the face representation. A patch of anterolaterally placed label that was present may be the dissociated rostrolateral arm area of other authors. 6. Taken together, these data suggest that extensive, horizontally oriented, intrinsic axon collaterals provide inputs to many different forelimb movement representations and may be recruited during complex movements to coordinate the activity of motor cortical zones whose predominant output is to forelimb muscle groups acting synchronously.

Animals

Integration in descending motor pathways controlling the forelimb in the cat. 2. Convergence on neurones mediating disynaptic cortico-motoneuronal excitation.

With intracellular recording from forelimb motoneurones the spatial facilitation technique has been used to investigate interaction between descending pathways and forelimb afferents. As previously shown for the hindlimb, pyramidal volleys effectively facilitate interneuronal transmission in reflex pathways from different primary afferents. Evidence is presented suggesting disynaptic excitation from corticospinal fibres of interneurones in the reciprocal Ia inhibitory pathway. Interneurones of other reflex pathways from group I muscle afferents recieve monosynaptic pyramidal excitation. During pyramidal facilitation volleys in cutaneous afferents may evoke PSPs in motoneurones after a central delay of 1.3 ms suggesting that the minimal linkage is disynaptic. Information regarding convergence on the neurones intercalated in the disynaptic cortico-motoneuronal pathway was obtained by investigating the effect from primary afferents and from other descending pathways on the disynaptic pyramidal EPSPs. Volleys in cutaneous and group I muscle affferents facilitate transmission in the disynaptic cortico-motoneuronal transmission with a time course showing oligosynaptic (probably monosynaptic) action the intercalated neurone. Rubrospinal volleys likewise effectively facilitate disynaptic cortico-motoneuronal pathway with a time course showing oligosynaptic (probably monosynaptic) action on the intercalated neurone. Rubrospinal volleys likewise effectively facilitate disynaptic cortico-motoneuronal transmission with a time course showing monosynaptic action on the intercalated neurone. Spatial facilitation experiments involving three tests revealed that those intercalated neurones which receive convergent monosynaptic excitation from corticospinal and rubrospinal fibres are excited also from cutaneous forelimb afferents. Disynaptic cortico-motoneuronal transmission was also monosynaptically facilitated by stimuli in the dorsal mesencephalic tegmentum probably activating tectospinal fibres. Disynaptic, presumed tectospinal EPSPs were facilitated from cutaneous forelimb afferents. The convergence onto the neurones intercalated in the disynaptic excitatory cortico-motoneuronal pathway suggests that these neurones integrate the activity in different descending pathways and primary forelimb afferents.

Animals

Forelimb motor performance following cervical spinal cord contusion injury in the rat.

The purpose of this study was to examine the degree, persistence, and nature of forelimb behavioral deficits following cervical spinal cord contusion injury in the rat. Forelimb reaching and pellet retrieval, forehead adhesive sticker removal, and vibrissae-induced forelimb placing were examined for 16 weeks following a weight-drop injury (10.0 g-2.5 cm) at the C4-C5 spinal level. Nine of 13 rats studied were unable to perform the pellet retrieval task due to pronounced forelimb extension hypometria. However, these animals did carry out the forehead sticker removal and vibrissae-induced placing tasks. Therefore, the loss of reaching ability related to pellet retrieval was not due to generalized paralysis. This interpretation was further supported by evaluation of the rostrocaudal extent of relative motoneuron loss from 1-mm divisions through the lesion zone. The extent of motoneuron pathology ranged from 2 to 6 mm but was largely confined to the C4-C5 spinal segments. Morphometric assessments of axonal sparing revealed that pellet retrieval performance during the last month of observation was significantly correlated with fiber sparing in the dorsal columns and ventral white matter, whereas no significant correlation could be demonstrated with regard to dorsolateral white matter. While there were no conspicuous differences in qualitative assessments of damage to interneuron pools (i.e., laminae V to VII) between the nonreaching and retrieval-recovered rats, the possibility of combined white and gray matter pathology contributing to this deficit still exists. These initial findings thus demonstrate that the weight-drop contusion injury model can be adopted to studies of cervical spinal cord trauma in the rat. Such lesions yield permanent deficits in forelimb function lending to future studies of possible therapeutic interventions. Furthermore, performance deficits observed at 1 week postinjury in the placing and forehead sticker removal tasks can be predictive of any potential for long-range spontaneous recovery in pellet retrieval ability.

Animals

Analysis of the forelimb crossed extension reflex in thalamic cats during stepping.

Forelimb crossed extension reflexes were examined in 22 thalamic cats. These reflexes were elicited either by backward passive movement or by repetitive electrical stimulation of cutaneous and joint afferent nerves in the contralateral forelimb. Single stimulation of the superficial radial nerve evoked two types of reflex responses--early (ER) and late (LR)--from the triceps brachii muscle on the contralateral side. The latencies were about 7 and 16-25 ms, corresponding to the propriospinal (PSR) and spino-bulbo-spinal (SBS) reflexes of the ipsilateral flexor, respectively. Repetitive stimulation of the superficial radial nerve evoked the LR but not the ER. The crossed extension reflex and LR were abolished by lesions of the dorsolateral funiculus of the cervical cord on the side opposite to the recording. The tonic EMG activity, crossed extension reflex and LR in the extensor on the side of lesions were abolished by lesions of the ventrolateral funiculus of the cervical cord. During forelimb stepping, the amplitudes of both ER and LR fluctuated depending on the phase of the step cycle. The ER appeared during a narrow period in the early phase of the stance, whereas the LR was observed during a wide period from the middle of the swing to the middle of the stance. Both responses were absent from the middle of the stance to the middle of the swing. These observations suggest that forelimb crossed extension reflexes involve both spinal and supraspinal (SBS) loop mechanisms, and that these are utilized during stepping, with the latter mechanism in particular playing an important part in the extension phase of the forelimb forward movement.

Afferent Pathways

Monosynaptic excitation of motoneurons innervating forelimb muscles following stimulation of the red nucleus in cats.

We examined the rubrospinal projection to forelimb motoneurons (Mns) in cats, in comparison with the corticospinal projection. Under pentobarbital anesthesia, intracellular recordings were made from forelimb Mns following stimulation of the red nucleus (RN) and the cerebral peduncle (CP). Single pulse stimulation of the RN produced excitatory postsynaptic potentials (EPSPs) in the majority of forelimb Mns (58/80, 72%). Segmental latencies of RN-EPSPs were shorter than 1.2 ms in 14 out of 22 Mns in which RN-EPSPs were detected in the C8-T1 segments, and in 3 of 36 Mns in the C6-C7 segments. The results suggest monosynaptic rubro-motoneuronal connections in a substantial portion of forelimb Mns particularly in the C8-T1 segments. There was no evidence suggesting monosynaptic connections between cerebral cortex and forelimb Mns.

Animals

Enhanced c-myc gene expression during forelimb regenerative outgrowth in the young Xenopus laevis.

Analysis of the expression of the c-myc protooncogene has been carried out in the forelimb regenerate of the Xenopus laevis froglet. Northern blot hybridization analysis revealed the presence of a 2.5-kilobase c-myc transcript in the regenerate forelimb at a level at least 7-fold more than the one found in nonregenerating forelimbs or stumps of regenerating forelimbs. In situ hybridization analyses confirmed the relative abundance of c-myc RNA in the regenerate forelimb and provided evidence of spatial localization of high levels of c-myc RNA in specific cell layers. The deepest layers of the wound epithelium of epidermal origin showed a strong signal, whereas virtually no c-myc RNA was detected in the outermost layers. Labeling was also observed in mesenchymal cells of the blastema where it was relatively evenly distributed. This pattern of c-myc RNA in the regenerate might indicate that the expression of c-myc plays a role in the regulation of the continued proliferation of specific cells of the regenerate, whereas repression of this gene in the epidermis correlates with terminal differentiation of keratinocytes.

Amputation, Surgical

Reflex vascular responses in kidney, ileum, and forelimb to carotid body stimulation.

Reflex vascular responses to local carotid chemoreceptor stimulation with hypoxic-hypercapnic, hypoxic, or hypercapnic blood were investigated in pentobarbitalized dogs. Bilaterally isolated carotid chemoreceptors were perfused via an extracorporeal lung circuit. Oxygen and carbon dioxide tensions of blood perfusing the carotid bodies were altered by ventilating the isolated lung with various O2-CO2 mixtures. Ventilation of the whole animal maintained normal systemic O2 and CO2 tensions. Perfusion pressures of the isolated kidney, ileum, forelimb, gracilis and hindpaw were measured during constant-flow perfusion. Carotid chemoreceptor stimulation with hypoxic-hypercapnic blood before vagotomy increased renal vascular resistance but caused no change in intestinal or forelimb resistance. Following vagotomy, hypoxic-hypercapnic, hypoxic, or hypercapnic carotid body stimulation increased renal, intestinal, and forelimb vascular resistance. Forelimb skin and muscle vascular beds contributed about equally to the increase in forelimb resistance. Gracilis muscle and hindpaw resistance also increased during hypoxic-hypercapnic stimulation after vagotomy.

Animals