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Input organization of distal and proximal forelimb areas in the monkey primary motor cortex: a retrograde double labeling study.

The present double-labeling study was designed to demonstrate the morphological framework for motor control in coordinated distal and proximal forelimb movements, which may partly, at least, depend on corticocortical and thalamocortical inputs to the forelimb area in the primary motor cortex. After intracortical microstimulation mapping of the forelimb area in the primary motor cortex of four macaque monkeys, a retrograde tracing study with fluorescent dyes was attempted to label simultaneously neurons in cortical and subcortical sites projecting to the distal forelimb representation area and those projecting to the proximal representation area of the primary motor cortex. Neurons projecting to distal and proximal forelimb parts of the primary motor cortex were largely separate in the following areas: the premotor area, primary somatosensory area, secondary somatosensory area, area 5, and thalamus. In contrast, there was no precise topographic organization of labeled projection neurons in the following areas: the supplementary motor area, cingulate motor area, primary motor cortex adjacent to the injection sites, claustrum, and basal nucleus of Meynert. The present study revealed that the forelimb area of the primary motor cortex receives both segregated and intermixed inputs from cortical and subcortical sources. In particular, the fact that the forelimb area of the primary motor cortex receives topographically organized inputs from the premotor area and nontopographically organized inputs from the supplementary motor area and cingulate motor area indicates possible different functional roles of frontal premotor areas in control of coordinated distal and proximal forelimb movements.

Animals↗

Laminar-dependent dendritic spine alterations in the motor cortex of adult rats following callosal transection and forced forelimb use.

Previously, the authors found that partial denervation of the motor cortex in adult animals can enhance this region's neuronal growth response to relevant behavioral change. Rats with partial corpus callosum transections that were forced to rely on one forelimb for 18 days had increased dendritic arborization of layer V pyramidal neurons in the opposite motor cortex compared to controls. This was not found as a result of denervation alone or of forced forelimb use alone. However, it seemed possible that each independent manipulation (i.e., forced forelimb use alone and callosal transections alone) resulted in neural structural alterations that were simply not revealed in measurements of dendritic branch number and/or not inclusive of layer V dendrites. This possibility was assessed in the current study with a reexamination of the Golgi-Cox impregnated tissue generated in the previous study. Tissue was quantified from rats that received either partial transections of the rostral two-thirds of the corpus callosum (CCX) or sham operations (Sham) followed either by 18 days of forced use of one forelimb (Use) or unrestricted use of both forelimbs (Cont). Measurements of apical and basilar dendrites from pyramidal neurons of layer II/III and layer V were performed to detect spine addition resulting from either increased spine density or the addition of dendritic material. As hypothesized, significant spine addition was found following forced forelimb use alone (Sham+Use) and callosal transections alone (CCX+Cont). However, forced use primarily increased spines on layer II/III pyramidal neurons, whereas callosal transections primarily increased dendritic spines on layer V pyramidal neurons in comparison to Sham+Cont. A much more robust increase in layer V dendritic spines was found in animals with the combination of forced forelimb use and denervation (CCX+Use). In contrast to the effects of forced use alone, however, CCX+Use rats failed to show major net increases in spines on layer II/III neurons. These results indicate that while callosal denervation may greatly enhance the neuronal growth and synaptogenic response to behavioral change in layer V, it may also limit spine addition associated with forced forelimb use in layer II/III of the motor cortex.

Animals↗

Convergence of sensory inputs upon projection neurons of somatosensory cortex: vestibular, neck, head, and forelimb inputs.

Cortico-cortical neurons and pyramidal tract (PT) neurons of the cat cerebral cortex were tested for convergent inputs from electrically stimulated vestibular, neck, head and forelimb nerves. Neurons were recorded within forelimb and vestibular projection regions of cortical area 3a. Consideration was given to both suprathreshold and subthreshold inputs. Neither vestibular, neck nor head inputs were detected in the forelimb region of area 3a. In contrast, within the vestibular projection region of area 3a, 43% (6/14) of the cortico-cortical neurons and 63% (24/38) of the PT neurons received excitatory vestibular input. Inputs from the skin of the pinna (greater auricular nerve) were detected only for PT neurons (66%, 25/38). No inputs were detected from afferent nerves supplying the dorsal neck muscles biventer cervicis and complexus. Cortico-cortical and PT neurons receiving vestibular input also received convergent inputs originating from forelimb group I deep and low threshold cutaneous afferent fibers. Further, one half of the PT neurons with vestibular input (12/24) received input from three somatic sources: forelimb group I deep, forelimb low threshold cutaneous and greater auricular (head) nerves. The input connectivities suggest a role for these projection neurons of somatosensory cortex in the coordination of head and forelimb movements. The convergence of vestibular information with somatic input from the forelimb implies that vestibular-influenced neurons of area 3a projecting to the motor cortex or through the pyramidal tract would signal head position or movement with respect to proprioceptive feedback from the limbs.

Afferent Pathways↗

Monosynaptic rubrospinal projections to distal forelimb motoneurons in the cat.

In the present electrophysiological experiments, we examined whether or not there are short-latency excitatory postsynaptic potentials (EPSPs) suggesting monosynaptic rubro- and corticomotoneuronal connections in the cervical spinal segments in cats. Under pentobarbital anesthesia and immobilization with gallamine triethiodide, intracellular recordings were made from forelimb motoneurons (Mns) following stimulation of the red nucleus (RN) and the cerebral peduncle (CP). Mns comprising the musculocutaneous nerve (Musc. Mns), the radial nerve (Rad. Mns), the ulnar nerve (Uln. Mns), and the median nerve (Med. Mns) were identified by their antidromic activation. Single-pulse stimulation of RN produced EPSPs with a mean amplitude of 0.35 mV in the majority of forelimb Mns (88 of 122, 72%). Half of the forelimb Mns (23 of 46) of the C8-T1 segments in which RN-EPSPs were detected by single shocks exhibited segmental latencies of RN-EPSPs shorter than 1.0 ms, while only 3 of 43 (7%) forelimb Mns in the C6-7 segments showed such latencies. These results suggest that rubromotoneuronal connections are, at least in part, monosynaptic for forelimb Mns that innervate in particular the hand and digit muscles. When paired-pulse stimuli (interpulse interval of 3 ms) were applied to RN, the second EPSPs were potentiated moderately (mean amount of temporal facilitation 24%) for EPSPs with short segmental latencies (< 1.0 ms). By contrast, single shocks to CP rarely produced EPSPs in forelimb Mns (9 of 106, 8%). There was no obvious evidence to suggest monosynaptic connections between corticospinal fibers and the forelimb Mns. The present results suggest the presence of preferential direct rubrospinal projection to Mns innervating distal forelimb muscles in the cat.

Animals↗

Delayed reorganization of the shoulder representation in forepaw barrel subfield (FBS) in first somatosensory cortex (SI) following forelimb deafferentation in adult rats.

We previously reported that 6-16 weeks after forelimb amputation in adult rats, neurons in layer IV of rat first somatosensory cortex (SI) in the forepaw barrel subfield (FBS) associated with the representation of the forepaw became responsive to new input from the shoulder (Pearson et al. 1999). These new shoulder-responsive sites in deafferented FBS had longer evoked response latencies than did sites in the shoulder representation located in the posterior part of the trunk subfield, hereafter referred to as the original shoulder representation. Furthermore, projection neurons in the original shoulder representation in both intact and deafferented adults did not extend their axons into the FBS, and ablation of the original shoulder representation cortex and/or the second somatosensory cortex (SII) failed to eliminate new shoulder input in the deafferented FBS (Pearson et al. 2001). These results led us to conclude that large-scale reorganization in FBS quite likely involved a subcortical substrate. In addition, the time course for large-scale cortical reorganization following forelimb amputation was unknown, and this information could shed light on potential mechanisms for large-scale cortical reorganization. In the present study, we extended our previous findings of large-scale cortical reorganization in the FBS by investigating the time course for reorganization following forelimb amputation. The major findings are: a) deafferented forelimb cortex remained unresponsive to shoulder stimulation during the 1st week following forelimb amputation; b) new responses to shoulder stimulation were first observed in deafferented forelimb cortex 2-3 weeks after forelimb amputation; however, the new shoulder input was restricted to locations in the former forearm cortex; c) islet(s) of new shoulder representation were first observed in deafferented FBS 4 weeks after amputation; these islets occupied a larger percentage of FBS in subsequent weeks; d) portions of FBS remained unresponsive as many as 4 months after deafferentation (maximum time examined between amputation and recording); and e) the increase in total size of the shoulder representation appeared to result from the establishment of new shoulder representations that were often discontinuous from the original shoulder representation. These findings provide evidence that forelimb amputation results in delayed reorganization of the FBS and we describe possible mechanisms and substrates underlying the reorganization.

Age Factors↗

Functional organization of the spinal reflex pathways from forelimb afferents to hindlimb motoneurones in the cat. II. Conditions of the interneuronal connections.

The interneuronal conditions of the descending pathways from forelimb afferents to hindlimb motoneurones were investigated by testing spatial interactions in these pathways and between these pathways and segmental lumbar reflex pathways. In high spinal unanaesthetized cats hindlimb motoneurones were intracellularly recorded and spatial interactions were tested between effects evoked by stimulation of pairs of ipsi- and contralateral forelimb nerves or pairs of a forelimb and an ipsilateral hindlimb nerve. The excitatory and late inhibitory pathways from forelimb afferents projecting to most of the hindlimb motoneurone pools, showed an interactive pattern which was distinctly different to the fast inhibitory pathway projecting specifically from ipsilateral forelimb afferents to flexor digitorum and hallucis longus (FDHL) motoneurones. Stimulation of homonymous or heteronymous pairs of two forelimb nerves of both sides evoked generally a distinct spatial facilitation of the excitatory and late inhibitory effects, while the specific early IPSPs to FDHL motoneurones were not facilitated. Paired stimulation of two forelimb nerves of one side only produced spatial facilitation of EPSPs or late IPSPs if low strength stimuli were used, using higher strength which induced larger effects, generally caused occlusion instead. In case of large IPSPs this may be due to the vicinity to the equilibrium potential. Except for an inhibition of cutaneous reflex pathways, the spatial interaction of the excitatory and late inhibitory pathways onto segmental lumbar reflex pathways was weak and variable. The fast inhibitory pathway to FDHL motoneurones showed a partial spatial facilitatory interaction with lumbar reflex pathways from cutaneous and group II muscle afferents. The second IPSP wave evoked by this pathway was inhibited by antidromic stimulation of the ventral root L7S1 and of the alpha-efferents of the antagonistic peroneal nerve. From the results conclusions are drawn on the interneuronal organization of the descending pathways from forelimb afferents to hindlimb motoneurones.

Animals↗

Facilitation of motor skill learning by callosal denervation or forced forelimb use in adult rats.

Unilateral forelimb sensorimotor cortex lesions in adult rats produce a compensatory hyper-reliance on the forelimb ipsilateral to the lesion and temporally related glial and neural plasticity in the contralateral homotopic cortex. Recently, we found that these lesions enhance acquisition of a motor skills task with the ipsilateral, non-impaired, forelimb in comparison to shams. This effect might be related to a denervation-induced facilitation of neuroplastic changes in the motor cortex opposite the lesion and/or to the lesion-induced hyper-reliance on the non-impaired forelimb. The present study assessed whether increased forelimb use, denervation of motor cortical callosal afferents, or a combination of the two influences acquisition of a skilled reaching task. Adult rats with partial corpus callosum transections or sham procedures were either forced to rely on one forelimb or permitted normal forelimb use for 8 days. Rats were then trained for 14 days with their previously non-preferred forelimb (and the forced-use limb) on a unilateral pellet retrieval task. Compared to shams, transections produced a greater acquisition rate and asymptotic performance level on the task. Forced-use improved reaching performance relative to controls, but this effect was less enduring than the improvements produced by transections alone. The addition of forced-use to transections did not further enhance performance. These findings suggest that denervation-induced changes are likely to be a major contributor to the enhanced learning observed after unilateral sensorimotor cortex lesions.

Animals↗

How the horse moves: 1. Significance of graphical representations of equine forelimb kinematics.

The kinematics of 24 two-year-old Dutch Warmblood horses were recorded at the trot (4 m/s) on a high-speed treadmill to study the coordination of joints within the equine forelimb. Joint angle-time, angle-angle, stick, and marker diagrams were used to show forelimb motion graphically. Because the kinematic data referred to the joint angles of the horse standing squarely and were time-standardised to the duration of the stride cycle, mean joint curves could be calculated for the total group. The motion of each segment in the equine forelimb during a complete stride is described and its function in intralimb coordination evaluated. It appeared that the rotation of the scapula and the cranio-caudal movement of the distal forelimb are synchronous and pendular. The carpal joint rapidly snaps into overextension at the beginning of the stance phase to enable the forelimb to work as a propulsive strut. The fetlock joint acts as an elastic spring, thereby conserving energy and, at the same time, absorbs oscillations generated by initial ground contact. Furthermore, the coordination between carpal and fetlock joints in the swing phase appears to be strongly influenced by inertia. Using the graphic tools evaluated in this paper, we were able to visualise the kinematics of the equine forelimb and relate these to specific functions of the forelimb in locomotion. This information can be used to select kinematic variables for clinical studies in which equine forelimb function has to be described and quantified.

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↗

Temporally staggered forelimb stimulation modulates barrel cortex optical intrinsic signal responses to whisker stimulation.

Characterization of neurovascular relationships is critical to accurate interpretation of functional neuroimaging data. We have previously observed spatial uncoupling of optical intrinsic signal imaging (OIS) and evoked potential (EP) responses in rodent barrel cortex following simultaneous whisker and forelimb stimulation, leading to changes in OIS response magnitude. To further test the hypothesis that this uncoupling may have resulted from "passive" overspill of perfusion-related responses between functional regions, we conducted the present study using temporally staggered rather than simultaneous whisker and forelimb stimulation. This paradigm minimized overlap of neural responses in barrel cortex and forelimb primary somatosensory cortex (SI), while maintaining overlap of vascular response time courses between regions. When contrasted with responses to 1.5-s lone-whisker stimulation, staggered whisker and forelimb stimulation resulted in broadening of barrel cortex OIS response time course in the temporal direction of forelimb stimulation. OIS response peaks were also temporally shifted toward the forelimb stimulation period; time-to-peak was shorter (relative to whisker stimulus onset) when forelimb stimulation preceded whisker stimulation and longer when forelimb stimulation followed whisker stimulation. In contrast with OIS and EP magnitude decreases previously observed during simultaneous whisker/forelimb stimulation, barrel cortex OIS response magnitude increased during staggered stimulation and no detectable changes in underlying EP activity were observed. Spatial extent of barrel cortex OIS responses also increased during staggered stimulation. These findings provide further evidence for spatial uncoupling of OIS and EP responses, and emphasize the importance of temporal stimulus properties on the effects of this uncoupling. It is hypothesized that spatial uncoupling is a result of passive overspill of perfusion-related responses into regions distinct from those which are functionally active. It will be important to consider potential influences of this uncoupling when designing and interpreting functional imaging studies that use hemodynamic responses to infer underlying neural activity.

Animals↗

Vascular distribution of contrast medium during intraosseous regional perfusion of the distal portion of the equine forelimb.

OBJECTIVE: To describe the vascular distribution pattern of contrast medium during intraosseous regional perfusion (IORP) of the distal portion of the equine forelimb. SAMPLE POPULATION: 13 cadaveric forelimbs from 12 horses without forelimb diseases. PROCEDURES: Serial lateromedial radiographic views were taken of the distal portion of 10 heparinized cadaveric forelimbs at 0, 1, 2, 6, 15, and 30 minutes during IORP of the third metacarpal bone (MCIII) by use of iodinated contrast medium and a tourniquet placed over the proximal portion of MCIII. Vascular regions of interest (ROI) were created for each radiograph. Reviewers identified the presence or absence of contrast medium-induced opacified vessels in all ROI on radiographs. This information was summarized to identify vessel-filling patterns over time. Vessel identification was verified by use of computed tomography angiography and latex perfusion studies on the distal portion of separate cadaveric forelimbs. RESULTS: During IORP, contrast medium filled the medullary cavity of the MCIII; exited via transcortical vessels; and diffused distally to the remaining arteries and veins of the forelimb, distal to the tourniquet. Maximum vessel and soft tissue opacification occurred in most specimens at 6 and 30 minutes, respectively. Serial radiography vessel patterns matched those of computed tomography images and dissected specimens. CONCLUSIONS AND CLINICAL RELEVANCE: IORP provides a repeatable pattern of vascular distribution in the distal portion of the equine forelimb. To our knowledge, our study provides the first documentation of arterial perfusion by use of IORP; results of previous reports indicate that IORP delivers medications to only the venous vessels of the perfused forelimb.

Animals↗

Effects of bolus injections of leukotrienes and norepinephrine on forelimb vascular and lymphatic pressures.

Leukotrienes, lypoxygenase metabolites of arachadonic acid, have been reported to be potent vasoconstrictors in some organs. This study was undertaken to delineate the actions of leukotrienes on both vascular and lymphatic vessels in the canine forelimb. Bolus intra-arterial injections of 1 microgram and 10 micrograms of leukotriene B4, C4, and D4 and 1 microgram of norepinephrine were made into forelimbs perfused at constant flow. Norepinephrine significantly increased systemic, forelimb perfusion and small artery pressures. Lymphatic pressure was significantly increased from a control of 6.6 mmHg to a peak of 14.4 mmHg. Leukotriene B4 in either dosage, did not significantly affect vascular or lymphatic pressures. Leukotriene C4 (1 microgram or 10 micrograms) significantly increased systemic and forelimb arterial pressures but did not alter lymphatic pressure. Leukotriene D4 (1 microgram) significantly increased small artery pressure. Leukotriene D4 (10 micrograms) increased systemic and forelimb arterial pressures. Neither dosage of leukotriene D4 significantly affected lymphatic pressure. Repeat injection of norepinephrine after completion of all leukotriene injections again markedly increased systemic, forelimb arterial and lymphatic pressures. These data indicate that leukotrienes exhibit only mild constrictor effects on forelimb blood vessels and do not significantly affect forelimb prenodal lymphatic vessels.

Animals↗

Organization of adult motor cortex representation patterns following neonatal forelimb nerve injury in rats.

Somatotopic representation patterns in the motor cortex (MI) of rats that had a unilateral forelimb amputation on the first postnatal day were examined after 2-4 months of survival. Intracortical electrical stimulation and recording techniques were used to map the somatic representation in MI and in the somatic sensory cortex (SI). In normal rats, vibrissa, forelimb, and hindlimb areas comprise the bulk of the MI representation. Stimulation within the forelimb area elicits elbow, wrist, or digit movements at the lowest current intensities. The proximal limb representation appears to be contained within the distal forelimb area, since shoulder movements are nearly always evoked by stimulating at higher current intensities at some distal forelimb sites. In agreement with previous studies, the distal forelimb representation overlapped the adjacent part of the granular SI cortex. Following removal of the forelimb at birth, 3 novel features of MI organization were observed. First, the areas from which stimulation evoked movements of the vibrissa or the shoulder musculature were larger than normal. Stimulation thresholds were lower than those required for comparable movements in normal rats throughout these areas, suggesting that nerve section had not simply unmasked a high-threshold representation. Second, vibrissa movements were more commonly paired with movements of the proximal forelimb muscles at the same site. Third, stimulation in the adjacent granular SI cortex failed to evoke shoulder or trunk movements, although receptive-field mapping in this region showed that cells were responsive to cutaneous stimulation of the trunk and shoulder region. These results indicate that several organizational features develop differently in MI following perinatal nerve injury: certain remaining muscle groups have enlarged cortical representations, there is a strengthening of some normally weak connections from MI to the proximal musculature, and muscles are grouped in unusual combinations. These data demonstrate that the formation of MI representation patterns is strongly influenced by nerve injury during the perinatal period.

Amputation, Surgical↗

Abnormalities in skilled reaching movements are improved by peripheral anesthetization of the less-affected forelimb after sensorimotor cortical infarcts in rats.

Unilateral damage to sensorimotor cortical (SMC) regions can profoundly impair skilled reaching function in the contralesional forelimb. Such damage also results in impairments and compensatory changes in the less-affected/ipsilesional forelimb, but these effects remain poorly understood. Furthermore, anesthetization of the ipsilesional hand in humans with cerebral infarcts has been reported to produce transient functional improvements in the paretic hand [Floel A, Nagorsen U, Werhahn KJ, Ravindran S, Birbaumer N, Knecht S, et al. Influence of somatosensory input on motor function in patients with chronic stroke. Ann Neurol 2004;56:206-12; Voller B, Floel A, Werhahn KJ, Ravindran S, Wu CW, Cohen LG. Contralateral hand anesthesia transiently improves poststroke sensory deficits. Ann Neurol 2006;59:385-8]. One aim of this study was to sensitively assay the bilateral effects of unilateral ischemic SMC damage on performance of a unimanual skilled reaching task (the single pellet retrieval task) that rats had acquired pre-operatively with each forelimb. The second aim was to determine whether partially recovered contralesional reaching function is influenced by anesthetization of the ipsilesional forelimb. Unilateral SMC lesions were found to result in transient ipsilesional impairments in reaching success and significant ipsilesional abnormalities in reaching movements compared with sham-operates. There were major contralesional reaching impairments which improved during a 4 week training period, but movements remained significantly abnormal. Anesthetization of the ipsilesional forelimb with lidocaine at this time attenuated the contralesional movement abnormalities. These findings indicate that unilateral ischemic SMC lesions impair skilled reaching behavior in both forelimbs. Furthermore, after partial recovery in the contralesional forelimb, additional improvements can be induced by transient anesthetization of the ipsilesional forelimb. This is consistent with the effects of unilateral anesthetization in humans which have been attributed to the modulation of competitive interhemispheric interactions. The present findings suggest that such interactions are also likely to influence skilled reaching function in rats.

Anesthesia↗

Detection of lameness and determination of the affected forelimb in horses by use of continuous wavelet transformation and neural network classification of kinematic data.

OBJECTIVE: To investigate continuous wavelet transformation and neural network classification of gait data for detecting forelimb lameness in horses. ANIMALS: 12 adult horses with mild forelimb lameness. PROCEDURE: Position of the head and right forelimb foot, metacarpophalangeal (ie, fetlock), carpal, and elbow joints was determined by use of kinematic analysis before and after palmar digital nerve blocks. We obtained 8 recordings from horses without lameness, 8 with right forelimb lameness, and 8 with left forelimb lameness. Vertical and horizontal position of the head and vertical position of the foot, fetlock, carpal, and elbow joints were processed by continuous wavelet transformation. Feature vectors were created from the transformed signals and a neural network trained with data from 6 horses, which was then tested on the remaining 2 horses for each category until each horse was used twice for training and testing. Correct classification percentage (CCP) was calculated for each combination of gait signals tested. RESULTS: Wavelet-transformed vertical position of the head and right forelimb foot had greater CCP (85%) than untransformed data (21%). Adding data from the fetlock, carpal, or elbow joints did not improve CCP over that for the head and foot alone. CONCLUSIONS AND CLINICAL RELEVANCE: Wavelet transformation of gait data extracts information that is important for the detection and differentiation of forelimb lameness of horses. All of the necessary information to detect lameness and differentiate the side of lameness can be obtained by observation of vertical head movement in concert with movement of the foot of 1 forelimb.

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↗

Forelimb and hindlimb forces in walking and galloping primates.

One trait that distinguishes the walking gaits of most primates from those of most mammalian nonprimates is the distribution of weight between the forelimbs and hindlimbs. Nonprimate mammals generally experience higher vertical peak substrate reaction forces on the forelimb than on the hindlimb. Primates, in contrast, generally experience higher vertical peak substrate reaction forces on the hindlimb than on the forelimb. It is currently unclear whether this unusual pattern of force distribution characterizes other primate gaits as well. The available kinetic data for galloping primates are limited and present an ambiguous picture about peak-force distribution among the limbs. The present study investigates whether the pattern of forelimb-to-hindlimb force distribution seen during walking in primates is also displayed during galloping. Six species of primates were video-recorded during walking and galloping across a runway or horizontal pole instrumented with a force-plate. The results show that while the force differences between forelimb and hindlimb are not significantly different from zero during galloping, the pattern of force distribution is generally the same during walking and galloping for most primate species. These patterns and statistical results are similar to data collected during walking on the ground. The pattern of limb differentiation exhibited by primates during walking and galloping stands in contrast to the pattern seen in most nonprimate mammals, in which forelimb forces are significantly higher. The data reported here and by Demes et al. ([1994] J. Hum. Evol. 26:353-374) suggest that a relative reduction of forelimb vertical peak forces is part of an overall difference in locomotor mechanics between most primates and most nonprimate mammals during both walking and galloping.

Animals↗

Detection of insulin receptors in newt liver and forelimb regenerates and the effects of local insulin deprivation on epimorphic regeneration.

Previous in vivo and in vitro studies indicate that insulin is required in adult newt forelimb regeneration. The objectives of the current study were 1) to detect insulin receptors in the liver (a classical target organ for insulin) and once verified, detection of insulin receptors in the adult newt forelimb regenerate; and 2) to determine whether locally implanting insulin antibody-soaked hydrolyzed polyacrylamide beads (hypa beads) into a regenerating forelimb blastema would affect its growth and/or differentiation. The results show that insulin receptors are detectable in the plasma membranes of newt liver and forelimb regenerates. Radioiodinated bovine insulin binding is time-dependent and specific; unlabeled bovine insulin competes with labeled insulin for binding to NLPM more effectively than does insulin-like growth factor-I, guinea pig insulin, and glucagon. The newt hepatic insulin receptor binds insulin with high affinity (1.1 nM-1) and low capacity (63 +/- 8 fmoles/mg). The size of the alpha subunit of the newt insulin receptor is 130 kDA and that of the beta subunit is 95 kDa. The beta subunits undergo insulin-stimulated phosphorylation in response to insulin. An autoantibody against the human insulin receptor recognizes the newt receptor protein. Insulin receptors are also detectable in 15 and 20 day newt forelimb regenerates. Specific immunogold labelling of the receptor-bound antibody appears to be restricted to the cellular processes of the regenerate. Implanting hypa beads soaked with purified insulin antibody into regenerating adult newt forelimbs results in abnormal growth and differentiation of the regenerates, confirming that insulin plays an essential role in adult newt forelimb regeneration.

Affinity Labels↗