Activity of upper limb muscles during human walking

The EMG activity of upper limb muscles during human gait has rarely been studied previously. It was examined in 20 normal volunteers in four conditions: walking on a treadmill (1) with unrestrained natural arm swing (Normal), (2) while volitionally holding the arms still (Held), (3) with the arms immobilized (Bound), and (4) with the arms swinging in phase with the ipsilateral legs, i.e. opposite-to-normal phasing (Anti-Normal). Normal arm swing involved weak rhythmical lengthening and shortening contractions of arm and shoulder muscles. Phasic muscle activity was needed to keep the unrestricted arms still during walking (Held), indicating a passive component of arm swing. An active component, possibly programmed centrally, existed as well, because some EMG signals persisted when the arms were immobilized during walking (Bound). Anti-Normal gait involved stronger EMG activity than Normal walking and was uneconomical. The present results indicate that normal arm swing has both passive and active components.     

Dynamic arm swinging in human walking

Humans tend to swing their arms when they walk, a curious behaviour since the arms play no obvious role in bipedal gait. It might be costly to use muscles to swing the arms, and it is unclear whether potential benefits elsewhere in the body would justify such costs. To examine these costs and benefits, we developed a passive dynamic walking model with free-swinging arms. Even with no torques driving the arms or legs, the model produced walking gaits with arm swinging similar to humans. Passive gaits with arm phasing opposite to normal were also found, but these induced a much greater reaction moment from the ground, which could require muscular effort in humans. We therefore hypothesized that the reduction of this moment may explain the physiological benefit of arm swinging. Experimental measurements of humans (n = 10) showed that normal arm swinging required minimal shoulder torque, while volitionally holding the arms still required 12 per cent more metabolic energy. Among measures of gait mechanics, vertical ground reaction moment was most affected by arm swinging and increased by 63 per cent without it. Walking with opposite-to-normal arm phasing required minimal shoulder effort but magnified the ground reaction moment, causing metabolic rate to increase by 26 per cent. Passive dynamics appear to make arm swinging easy, while indirect benefits from reduced vertical moments make it worthwhile overall.     

Acquisition of the lateral inconsistency in involuntary behaviour of upper limbs in 12-year-old children during walking at moderate speed.

The aim of this work was to investigate possible lateralisation in the behaviour of periodic motion of the human upper limb, during normal walking at a comfortable speed of locomotion. Ten healthy pre-adolescent, strongly right-handed, 12-year-old males participated in the experiment. Participants were walking on a treadmill with a standardised velocity of 1.1m/s (comfortable speed for all of them). A video analysis system with Silicon software was used to synchronically measure various angles of arms and forearms. The initial, final and interim angular positions of both arms and forearms in 10 cycles of each participant were compared in terms of variations (cycle to cycle) between both upper extremities at corresponding phases of each cycle for distal and proximal segments, respectively. We compared the coefficients of variation in relation to the spatial and temporal data of both limbs and their angular velocities. In addition we investigated the level of cycle-to-cycle regularity (constancy) of behaviour in relation to various positions, periods and velocities of movement of upper extremities (specifically arms and forearms) using the Eta non-linear method of correlation. All participants exhibited a lower level of regularity for the distal segments. The spatial and temporal variations in the dominant limb were also greater than the non-dominant limb for all participants. This may be due to a larger contribution from the right-sided muscles that are considered to be the main contributing factor to the motion of the dominant upper limb during walking, rather than simply gravity force acting alone. A possible practical application of this information may be useful in the objective clinical identification of the level of dominance of the upper extremity (arm plus forearm), in addition to 'traditional' handedness.     

Combining muscle synergies and biomechanical analysis to assess gait in stroke patients

The understanding of biomechanical deficits and impaired neural control of gait after stroke is crucial to prescribe effective customized treatments aimed at improving walking function. Instrumented gait analysis has been increasingly integrated into the clinical practice to enhance precision and inter-rater reliability for the assessment of pathological gait. On the other hand, the analysis of muscle synergies has gained relevance as a novel tool to describe the neural control of walking. Since muscle synergies and gait analysis capture different but equally important aspects of walking, we hypothesized that their combination can improve the current clinical tools for the assessment of walking performance.To test this hypothesis, we performed a complete bilateral, lower limb biomechanical and muscle synergies analysis on nine poststroke hemiparetic patients during overground walking. Using stepwise multiple regression, we identified a number of kinematic, kinetic, spatiotemporal and synergy-related features from the paretic and non-paretic side that, combined together, allow to predict impaired walking function better than the Fugl-Meyer Assessment score. These variables were time of peak knee flexion, VAF_total values, duration of stance phase, peak of paretic propulsion and range of hip flexion. Since these five variables describe important biomechanical and neural control features underlying walking deficits poststroke, they may be feasible to drive customized rehabilitation therapies aimed to improve walking function.This paper demonstrates the feasibility of combining biomechanical and neural-related measures to assess locomotion performance in neurologically injured individuals.     

Effect of arm swinging on lumbar spine and hip joint forces

During level walking, arm swing plays a key role in improving dynamic stability. In vivo investigations with a telemeterized vertebral body replacement showed that spinal loads can be affected by differences in arm positions during sitting and standing. However, little is known about how arm swing could influence the lumbar spine and hip joint forces and motions during walking. The present study aims to provide better understanding of the contribution of the upper limbs to human gait, investigating ranges of motion and joint reaction forces.A three-dimensional motion analysis was carried out via a motion capturing system on six healthy males and five patients with hip instrumented implant. Each subject performed walking with different arm swing amplitudes (small, normal, and large) and arm positions (bound to the body, and folded across the chest). The motion data were imported in a commercial musculoskeletal analysis software for kinematic and inverse dynamic investigation.The range of motion of the thorax with respect to the pelvis and of the pelvis with respect to the ground in the transversal plane were significantly associated with arm position and swing amplitude during gait. The hip external-internal rotation range of motion statistically varied only for non-dominant limb. Unlike hip joint reaction forces, predicted peak spinal loads at T12-L1 and L5-S1 showed significant differences at approximately the time of contralateral toe off and contralateral heel strike.Therefore, arm position and swing amplitude have a relevant effect on kinematic variables and spinal loads, but not on hip loads during walking.     

The metabolic cost of walking in humans, chimpanzees, and early hominins

Bipedalism is a defining feature of the hominin lineage, but the nature and efficiency of early hominin walking remains the focus of much debate. Here, we investigate walking cost in early hominins using experimental data from humans and chimpanzees. We use gait and energetics data from humans, and from chimpanzees walking bipedally and quadrupedally, to test a new model linking locomotor anatomy and posture to walking cost. We then use this model to reconstruct locomotor cost for early, ape-like hominins and for the A.L. 288 Australopithecus afarensis specimen. Results of the model indicate that hind limb length, posture (effective mechanical advantage), and muscle fascicle length contribute nearly equally to differences in walking cost between humans and chimpanzees. Further, relatively small changes in these variables would decrease the cost of bipedalism in an early chimpanzee-like biped below that of quadrupedal apes. Estimates of walking cost in A.L. 288, over a range of hypothetical postures from crouched to fully extended, are below those of quadrupedal apes, but above those of modern humans. These results indicate that walking cost in early hominins was likely similar to or below that of their quadrupedal ape-like forebears, and that by the mid-Pliocene, hominin walking was less costly than that of other apes. This supports the hypothesis that locomotor energy economy was an important evolutionary pressure on hominin bipedalism.     

Interlimb Coordination during Forward and Backward Walking in Primary School-Aged Children

Previous studies comparing forward (FW) and backward (BW) walking suggested that the leg kinematics in BW were essentially those of FW in reverse. This led to the proposition that in adults the neural control of FW and BW originates from the same basic neural circuitry. One aspect that has not received much attention is to what extent development plays a role in the maturation of neural control of gait in different directions. BW has been examined either in adults or infants younger than one year. Therefore, we questioned which changes occur in the intermediate phases (i.e. in primary school-aged children). Furthermore, previous research focused on the lower limbs, thereby raising the question whether upper limb kinematics are also simply reversed from FW to BW. Therefore, in the current study the emphasis was put both on upper and lower limb movements, and the coordination between the limbs. Total body 3D gait analysis was performed in primary school-aged children (N = 24, aged five to twelve years) at a preferred walking speed to record angular displacements of upper arm, lower arm, upper leg, lower leg, and foot with respect to the vertical (i.e. elevation angle). Kinematics and interlimb coordination were compared between FW and BW. Additionally, elevation angle traces of BW were reversed in time (revBW) and correlated to FW traces. Results showed that upper and lower limb kinematics of FW correlated highly to revBW kinematics in children, which appears to be consistent with the proposal that control of FW and BW may be similar. In addition, age was found to mildly alter lower limb kinematic patterns. In contrast, interlimb coordination was similar across all children, but was different compared to adults, measured for comparison. It is concluded that development plays a role in the fine-tuning of neural control of FW and BW.     

‘Initial state’ coordinations reproduce the instant flexibility for human walking

An important feature of human locomotor control is the instant adaptability to unpredictable changes of conditions surrounding the locomotion. Humans, for example, can seamlessly adapt their walking gait following a sudden ankle impairment (e.g., as a result of an injury). In this paper, we propose a theoretical study of the mechanisms underlying flexible locomotor control. We hypothesize that flexibility is achieved by modulating the posture at the beginning of the stance phase—the initial state. Using a walking model, we validate our hypothesis through computer simulations     

An upper-body can improve the stability and efficiency of passive dynamic walking

The compass-gait walker proposed by McGeer can walk down a shallow slope with a self-stabilizing gait that requires no actuation or control. However, as the slope goes to zero so does the walking speed, and dynamic gait stability is only possible over a very narrow range of slopes. Gomes and Ruina have results demonstrating that by adding a torso to the compass-gait walker, it can walk passively on level-ground with a non-infinitesimal constant average speed. However, the gait involves exaggerated joint movements, and for energetic reasons horizontal passive dynamic walking cannot be stable. We show in this research that in addition to collision-free walking, adding a torso improves stability and walking speed when walking downhill. Furthermore, adding arms to the torso results in a collision-free periodic gait with natural-looking torso and limb movements. Overall, in contrast to the suggestions that active control may be needed to balance an upper-body on legs, it turns out that the upper and lower bodies can be integrated to improve the stability, efficiency and speed of a passive dynamic walker.     

Origins of primate locomotion: gait mechanics of the woolly opossum

The locomotion of primates differs from that of other mammals in three fundamental ways. During quadrupedal walking, primates use diagonal sequence gaits, protract their arms more at forelimb touchdown, and experience lower vertical substrate reaction forces on their forelimbs relative to their hindlimbs. It is widely held that the unusual walking gaits of primates represent a basal adaptation for movement on thin, flexible branches and reflect a major change in the functional role of the forelimb. However, little data on nonprimate arboreal mammals exist to test this notion. To that end, we examined the gait mechanics of the woolly opossum (Caluromys philander), a marsupial convergent with small-bodied prosimians in ecology, behavior, and morphology. Data on the footfall sequence, relative arm protraction, and peak vertical substrate reaction forces were obtained from videotapes and force records for three adult woolly opossums walking quadrupedally on a wooden runway and a thin pole. For all steps recorded on both substrates, woolly opossums always used diagonal sequence walking gaits, protracted their arms beyond 90 degrees relative to horizontal body axis, and experienced peak vertical substrate reaction forces on forelimbs that were significantly lower than on hindlimbs. The woolly opossum is the first nonprimate mammal to show locomotor mechanics that are identical to those of primates. This case of convergence between primates and a committed fine-branch, arboreal marsupial strongly implies that the earliest primates evolved gait specializations for fine-branch locomotion, which reflect important changes in forelimb function.     

Effects of arm swing on mechanical parameters of human gait

The aim of this study was to investigate the influence of the upper limb swing on human gait. Measurements were performed on 52 subjects by using the Elite system with two cameras and a Kistler force platform. The recording of trajectories of characteristic body points on the subjects and the measurement of ground reaction forces have been performed at normal walking and at walking with emphasised rhythmic upper limb swing. The trajectory of the whole body mass centre, central dynamic moments of inertia and ground reaction forces have been calculated for every subject and mean curves of the entire group have been determined for walking with the natural and the emphasised upper limb swing. The determined mean values of normalised mechanical parameters have been compared and differences between the gait with the natural and the emphasised upper limb swing have been described.     

Can altered muscle synergies control unimpaired gait?

Recent studies have postulated that the human motor control system recruits groups of muscles through low-dimensional motor commands, or muscle synergies. This scheme simplifies the neural control problem associated with the high-dimensional structure of the neuromuscular system. Several lines of evidence have suggested that neurological injuries, such as stroke or cerebral palsy, may reduce the dimensions that are available to the motor control system, and these altered dimensions or synergies are thought to contribute to impaired walking patterns. However, no study has investigated whether impaired low-dimensional control spaces necessarily lead to impaired walking patterns. In this study, using a two-dimensional model of walking, we developed a synergy-based control framework that can simulate the dynamics of walking. The simulation analysis showed that a synergy-based control scheme can produce well-coordinated movements of walking matching unimpaired gait. However, when the dimensions available to the controller were reduced, the simplified emergent pattern deviated from unimpaired gait. A system with two synergies, similar to those seen after neurological injury, could not produce an unimpaired walking pattern. These findings provide further evidence that altered muscle synergies can contribute to impaired gait patterns and may need to be directly addressed to improve gait after neurological injury.     

Association between arthritis score at the onset of the disease and long-term locomotor outcome in adjuvant-induced arthritis in rats

IntroductionTo investigate the connection between the intensity of initial symptoms of inflammation and locomotor outcome in rheumatoid arthritis, we examined the relationship between long-term locomotor abnormalities and signs of inflammation at the onset of the disease in adjuvant-induced arthritis (AIA) in rats.MethodsThe arthritis score and hind-paw diameter were followed from immunization to day 195 (~7 months). At this time, locomotion was recorded during forced treadmill walking using 3D motion technology before radiographic scoring of hind limb joint damage. Many locomotor parameters were analyzed including time and length parameters, limbs kinematics, lateral paw position at toe off, maximal hind-paw elevation and posture. Ankle mobility was assessed from range of motion (ROM) of the joint during locomotion. Experiments were run in AIA (n = 18) and age-matched non-AIA rats (n = 8).ResultsAll AIA rats exhibited signs of inflammation at day 14 with a peak of inflammatory symptoms at day 22 post-immunization. After the first episode of inflammation, 83 % of AIA rats demonstrated recurrent disease (from week 6 to week 23). The frequency of inflammatory episodes (1 to 5) was not linked to the arthritis score at day 22. At day 195 post-immunization, AIA rats showed significantly impaired locomotion and radiographic lesions as compared to control rats. Significant relationships were observed between most locomotion-related parameters and concurrent ROM of ankle, which correlated negatively with the radiographic score. ROM of ankle at day 195 correlated negatively with both the arthritis score and hind-paw diameter measured at day 14, 22 and 30 post-immunization.ConclusionDecreased ankle mobility can be considered a driver of locomotion impairment in AIA. In this model, the severity of the initial inflammatory symptoms had a good prognostic value for long-term locomotor outcome.     

Artificial neural network model for the generation of muscle activation patterns for human locomotion.

Skilled locomotor behaviour requires information from various levels within the central nervous system (CNS). Mathematical models have permitted researchers to simulate various mechanisms in order to understand the organization of the locomotor control system. While it is difficult to adequately characterize the numerous inputs to the locomotor control system, an alternative strategy may be to use a kinematic movement plan to represent the complex inputs to the locomotor control system based on the possibility that the CNS may plan movements at a kinematic level. We propose the use of artificial neural network (ANN) models to represent the transformation of a kinematic plan into the necessary motor patterns. Essentially, kinematic representation of the actual limb movement was used as the input to an ANN model which generated the EMG activity of 8 muscles of the lower limb and trunk. Data from a wide variety of gait conditions was necessary to develop a robust model that could accommodate various environmental conditions encountered during everyday activity. A total of 120 walking strides representing normal walking and ten conditions where the normal gait was modified in terms of cadence, stride length, stance width or required foot clearance. The final network was assessed on its ability to predict the EMG activity on individual walking trials as well as its ability to represent the general activation pattern of a particular gait condition. The predicted EMG patterns closely matched those recorded experimentally, exhibiting the appropriate magnitude and temporal phasing required for each modification. Only 2 of the 96 muscle/gait conditions had RMS errors above 0.10, only 5 muscle/gait conditions exhibited correlations below 0.80 (most were above 0.90) and only 25 muscle/gait conditions deviated outside the normal range of muscle activity for more than 25% of the gait cycle. These results indicate the ability of single network ANNs to represent the transformation between a kinematic movement plan and the necessary muscle activations for normal steady state locomotion but they were also able to generate muscle activation patterns for conditions requiring changes in walking speed, foot placement and foot clearance. The abilities of this type of network have implications towards both the fundamental understanding of the control of locomotion and practical realizations of artificial control systems for use in rehabilitation medicine.     

Simultaneous Scalp Electroencephalography (EEG), Electromyography (EMG), and Whole-body Segmental Inertial Recording for Multi-modal Neural Decoding

Recent studies support the involvement of supraspinal networks in control of bipedal human walking. Part of this evidence encompasses studies, including our previous work, demonstrating that gait kinematics and limb coordination during treadmill walking can be inferred from the scalp electroencephalogram (EEG) with reasonably high decoding accuracies. These results provide impetus for development of non-invasive brain-machine-interface (BMI) systems for use in restoration and/or augmentation of gait- a primary goal of rehabilitation research. To date, studies examining EEG decoding of activity during gait have been limited to treadmill walking in a controlled environment. However, to be practically viable a BMI system must be applicable for use in everyday locomotor tasks such as over ground walking and turning. Here, we present a novel protocol for non-invasive collection of brain activity (EEG), muscle activity (electromyography (EMG)), and whole-body kinematic data (head, torso, and limb trajectories) during both treadmill and over ground walking tasks. By collecting these data in the uncontrolled environment insight can be gained regarding the feasibility of decoding unconstrained gait and surface EMG from scalp EEG.     

Residual Upper Arm Motor Function Primes Innervation of Paretic Forearm Muscles in Chronic Stroke after Brain-Machine Interface (BMI) Training

Background

Abnormal upper arm-forearm muscle synergies after stroke are poorly understood. We investigated whether upper arm function primes paralyzed forearm muscles in chronic stroke patients after Brain-Machine Interface (BMI)-based rehabilitation. Shaping upper arm-forearm muscle synergies may support individualized motor rehabilitation strategies.

Methods

Thirty-two chronic stroke patients with no active finger extensions were randomly assigned to experimental or sham groups and underwent daily BMI training followed by physiotherapy during four weeks. BMI sessions included desynchronization of ipsilesional brain activity and a robotic orthosis to move the paretic limb (experimental group, n = 16). In the sham group (n = 16) orthosis movements were random. Motor function was evaluated with electromyography (EMG) of forearm extensors, and upper arm and hand Fugl-Meyer assessment (FMA) scores. Patients performed distinct upper arm (e.g., shoulder flexion) and hand movements (finger extensions). Forearm EMG activity significantly higher during upper arm movements as compared to finger extensions was considered facilitation of forearm EMG activity. Intraclass correlation coefficient (ICC) was used to test inter-session reliability of facilitation of forearm EMG activity.

Results

Facilitation of forearm EMG activity ICC ranges from 0.52 to 0.83, indicating fair to high reliability before intervention in both limbs. Facilitation of forearm muscles is higher in the paretic as compared to the healthy limb (p<0.001). Upper arm FMA scores predict facilitation of forearm muscles after intervention in both groups (significant correlations ranged from R = 0.752, p = 0.002 to R = 0.779, p = 0.001), but only in the experimental group upper arm FMA scores predict changes in facilitation of forearm muscles after intervention (R = 0.709, p = 0.002; R = 0.827, p<0.001).

Conclusions

Residual upper arm motor function primes recruitment of paralyzed forearm muscles in chronic stroke patients and predicts changes in their recruitment after BMI training. This study suggests that changes in upper arm-forearm synergies contribute to stroke motor recovery, and provides candidacy guidelines for similar BMI-based clinical practice.     

Joint moment work during the stance-to-swing transition in hemiparetic subjects

Following stroke many individuals are left with neurological and functional deficits, including hemiparesis, which impair their ability to walk. Our previous work reported that propulsion of the paretic leg during pre-swing is impaired and may limit gait speed and knee flexion during swing. To elucidate the mechanism of this impairment, we assessed the mechanical work produced by the hip, knee, and ankle moments during pre-swing of the paretic limb in a group of stroke subjects and compared it with the work produced by non-disabled controls walking at similar speeds. Kinematic and kinetic gait data were collected from 23 hemiparetic and 10 control subjects. The hemiparetic subjects walked at their self-selected speeds. The controls walked at their self-selected and two or three slower speeds. Even when compared to controls walking at slow speeds, ankle plantarflexor work during pre-swing was greatly reduced (-0.136+/-0.062J/kg) in the hemiparetic subjects. Differences in hip (+0.006+/-0.020J/kg) and knee (+0.040+/-0.026J/kg) moment work partially offset the reduction in ankle work, but net joint moment work was still significantly reduced (-0.088+/-0.056J/kg). The reduction in work accounts for the low energy of the paretic limb at the stance-to-swing transition previously reported. Future investigation is needed to determine if targeted training of the plantarflexors in the paretic limb improves swing-phase function and locomotor performance in hemiparetic individuals.     

The role of serotonin in sleep disordered breathing associated with Parkinson disease: a correlative [11C]DASB PET imaging study

Sleep dysfunction and excessive daytime sleepiness are common in Parkinson disease (PD). Several studies suggest that PD patients exhibit high prevalence of sleep-disordered breathing (SDB). PD has a complex profile of neurochemical deficits in which abnormalities of different neurotransmitter systems may play significant and differing roles in the development of non-motor features. In the present study, we investigated whether SDB in PD is related to serotoninergic neuron degeneration. We used a cross-sectional design to assess the correlation between SDB and measures of caudal brainstem serotonin neuron integrity. Fifty one PD participants with mean disease duration of 6.0 (SD 3.7) years and mean age of 63.9 (SD 6.2) years were studied. We measured caudal brainstem serotoninergic innervation with [(11)C]DASB positron emission tomography (PET) imaging and striatal dopaminergic innervation with [(11)C]DTBZ PET imaging. SDB was assessed with polysomnography (PSG) and sleepiness with multiple sleep latency tests. Greater than half of participants exhibited PSG evidence of significant SDB; 12 participants had normal PSGs, 6 had mild SDB, 20 had moderate SDB, and 13 had severe SDB. We found no association between severity of SDB and caudal brainstem serotoninergic innervation in PD participants. Striatal dopaminergic denervation did not correlate with severity of SDB. We did find significant correlations between measures of motor function impairment and sleep quantity and quality in PD. Neither serotoninergic nor dopaminergic neuron degeneration is likely to play a major role in SDB observed in PD patients.     

Ground reaction forces and lower-limb joint kinetics of turning gait in typically developing children

Turning is a common locomotor task essential to daily activity; however, very little is known about the forces and moments responsible for the kinematic adaptations occurring relative to straight-line gait in typically developing children. Thus, the aims of this study were to analyse ground reaction forces (GRFs), ground reaction free vertical torque (T_Z), and the lower-limb joint kinetics of 90° outside (step) and inside (spin) limb turns. Step, spin, and straight walking trials from fifty-four typically developing children were analysed. All children were fit with the Plug-in Gait and Oxford Foot Model marker sets while walking over force plates embedded in the walkway. Net internal joint moments and power were computed via a standard inverse dynamics approach. All dependent variables were statistically analysed over the entire curves using the mean difference 95% bootstrap confidence band approach. GRFs were directed medially for step turns and laterally for spin turns during the turning phase. Directions were reversed and magnitudes decreased during the approach phase. Step turns showed reduced ankle power generation, while spin turns showed large T_Z. Both strategies required large knee and hip coronal and transverse plane moments during swing. These kinetic differences highlight adaptations required to maintain stability and reorient the body towards the new walking direction during turning. From a clinical perspective, turning gait may better reveal weaknesses and motor control deficits than straight walking in pathological populations, such as children with cerebral palsy, and could potentially be implemented in standard gait analysis sessions.     

Regenerative ability of double-half and half upper arms in the newt, Notophthalmus viridescens

The upper arms of adult newts (Notophthalmus viridescens) were surgically manipulated to create double-half dorsal, double-half ventral, double-half anterior, and double-half posterior upper arms, and longitudinal half-dorsal, half-ventral, half-anterior, and half-posterior upper arms. Amputation through the double-half upper arms usually failed to elicit normal distal regeneration, despite the fact that an apparently normal regeneration blastema was initially formed. Instead, regeneration in these cases was limited to the formation of a variable number of small cartilage elements. On the basis of these results it is concluded that a complete limb circumference is required for distal transformation in newts, in addition to the well-established requirements for a wound epidermis, adequate innervation and dedifferentiation leading to blastema formation. A model for the sequential generation of new parts of the limb pattern during distal transformation from a complete circumference is presented. This model can also account for the occurrence of normal early stages of regeneration in double-half upper arms. Half upper arms which were amputated immediately were shown to develop single, complete regenerates. If amputation of half upper arms was delayed three or more weeks to permit complete wound healing, a supernumerary limb from the lateral wound surface sometimes developed in addition to a complete, single limb from the distal amputation surface.