Kinematic invariants during cyclical arm movements

It has been observed that the motion of the arm end-point (the hand, fingertip or the tip of a pen) is characterized by a number of regularities (kinematic invariants). Trajectory is usually straight, and the velocity profile has a bell shape during point-to-point movements. During drawing movements, a two-thirds power law predicts the dependence of the end-point velocity on the trajectory curvature. Although various principles of movement organization have been discussed as possible origins of these kinematic invariants, the nature of these movement trajectory characteristics remains an open question. A kinematic model of cyclical arm movements derived in the present study analytically demonstrates that all three kinematic invariants can be predicted from a two-joint approximation of the kinematic structure of the arm and from sinusoidal joint motions. With this approach, explicit expressions for two kinematic invariants, the two-thirds power law during drawing movements and the velocity profile during point-to-point movements are obtained as functions of arm segment lengths and joint motion parameters. Additionally, less recognized kinematic invariants are also derived from the model. The obtained analytical expressions are further validated with experimental data. The high accuracy of the predictions confirms practical utility of the model, showing that the model is relevant to human performance over a wide range of movements. The results create a basis for the consolidation of various existing interpretations of kinematic invariants. In particular, optimal control is discussed as a plausible source of invariant characteristics of joint motions and movement trajectories.     

Representation of movements and muscles within the primate precentral motor cortex: historical and current perspectives

A major problem with the study of the control of movement and posture is to determine how specific brain areas contribute to the selection of those particular muscle patterns that underlie a coordinated movement. With this problem in mind, a selective review is presented of mapping studies of the primate motor cortex, whose results bear on the question of how the spatial organization of cortical efferent cells might contribute to the production of organized muscle synergies. More recent findings are also summarized, which appear to resolve previous controversies on the question of whether movements or muscles are the primary units of motor cortex organization. These same findings suggest also a form of spatial organization within the primate precentral gyrus that would allow spatially simple afferent inputs to evoke the muscle synergies that are necessary for a variety of simple movements of the arm and hand.     

Using voluntary motor commands to inhibit involuntary arm movements

A hallmark of voluntary motor control is the ability to stop an ongoing movement. Is voluntary motor inhibition a general neural mechanism that can be focused on any movement, including involuntary movements, or is it mere termination of a positive voluntary motor command? The involuntary arm lift, or ‘floating arm trick’, is a distinctive long-lasting reflex of the deltoid muscle. We investigated how a voluntary motor network inhibits this form of involuntary motor control. Transcranial magnetic stimulation of the motor cortex during the floating arm trick produced a silent period in the reflexively contracting deltoid muscle, followed by a rebound of muscle activity. This pattern suggests a persistent generator of involuntary motor commands. Instructions to bring the arm down voluntarily reduced activity of deltoid muscle. When this voluntary effort was withdrawn, the involuntary arm lift resumed. Further, voluntary motor inhibition produced a strange illusion of physical resistance to bringing the arm down, as if ongoing involuntarily generated commands were located in a ‘sensory blind-spot’, inaccessible to conscious perception. Our results suggest that voluntary motor inhibition may be a specific neural function, distinct from absence of positive voluntary motor commands.     

Mutual influences of upper and lower extrimities during cyclic movements

The possibility of muscle activation of passive arm during its cyclic movements, imposed by active movements of contralateral arm or by experimenter was studied, as well as the influence of lower extremities cyclic movements onto arm muscles activity. In addition to that the activity of legs muscles was estimated in dependence on motor task condition for arms. Ten healthy supine subjects carried out opposite movements of arms with and without stepping-like movements of both legs. The experiment included three conditions for arm movements: 1) the active movements of both arms; 2) the active movements of one arm, when other entirely passive arm participated in the movement by force; 3) passive arm movement caused by experimenter. In the condition 2) additional load on active arm was applied (30 N and 60 N). In all three conditions the experiment was carried out with arms movements only or together with legs movements. The capability of passive moving arm muscles activation depended on increasing afferent inflow from muscles of contralateral arm was demonstrated. Emerging electrical activity was modulated in the arms movements cycle and depended on the degree of active arm loading. During combined active movements of arms and legs the reduction of activity in the flexor muscles of shoulder and forearm was observed. Concomitant arms movements increased the magnitude ofelectromiographic bursts during passive stepping-like movements in the most of recorded muscles, and the same increasing was only observed in biceps femoris and tibialis anterior muscles during active legs movement. The increasing of loading of one arm caused essential augmentation of EMG-activity in the majority of recording legs muscles. The data obtained are the additional proof of existence of functionally significant neuronal interaction both between arms and between upper and lower extremities, which is evidently depend on the intraspinal neuronal connections.     

Rethinking Motor Lateralization: Specialized but Complementary Mechanisms for Motor Control of Each Arm

Motor lateralization in humans has primarily been characterized as “handedness”, resulting in the view that one arm-hemisphere system is specialized for all aspects of movement while the other is simply a weaker analogue. We have proposed an alternative view that motor lateralization reflects proficiency of each arm for complementary functions that arises from a specialization of each hemisphere for distinct movement control mechanisms. However, before this idea of hemispheric specialization can be accepted, it is necessary to precisely identify these distinct, lateralized mechanisms. Here we show in right-handers that dominant arm movements rely on predictive mechanisms that anticipate and account for the dynamic properties of the arm, while the non-dominant arm optimizes positional stability by specifying impedance around equilibrium positions. In a targeted-reaching paradigm, we covertly and occasionally shifted the hand starting location either orthogonal to or collinear with a particular direction of movement. On trials on which the start positions were shifted orthogonally, we did not notice any strong interlimb differences. However, on trials on which start positions were shifted orthogonally, the dominant arm largely maintained the direction and straightness of its trajectory, while the non-dominant arm deviated towards the previously learned goal position, consistent with the hypothesized control specialization of each arm-hemisphere system. These results bring together two competing theories about mechanisms of movement control, and suggest that they coexist in the brain in different hemispheres. These findings also question the traditional view of handedness, because specialized mechanisms for each arm-hemisphere system were identified within a group of right-handers. It is likely that such hemispheric specialization emerged to accommodate increasing motor complexity during evolution.     

Interlimb interactions during cyclic in-phase and antiphase movements of arms and legs and their dependence on afferent influences

Coordinated arm and leg movements imply neural interactions between the rhythmic generators of the upper and lower extremities. In ten healthy subjects in the lying position, activity of the muscles of the upper and lower extremities was recorded during separate and joint cyclic movements of the arms and legs with different phase relationships between the movements of the limbs and under various conditions of the motor task. Antiphase active arm movements were characterized by higher muscle activity than during the inphase mode. The muscle activity during passive arm movements imposed by the experimentalist was significantly lower than muscle activity during passive arm movements imposed by the other arm. When loading one arm, the muscle activity in the other, passively moving, arm increased independently from the synergy of arm movements. During a motor task implementing joint antiphase movements of both upper and lower extremities, compared to a motor task implementing their joint in-phase movements, we observed a significant increase in activity in the biceps brahii muscle, the tibialis anterior muscle, and the biceps femoris muscle. Loading of arms in these motor tasks has been accompanied by increased activity in some leg muscles. An increase in the frequency of rhythmic movements resulted in a significant growth of the muscle activity of the arms and legs during their cooperative movements with a greater rate of rise in the flexor muscle activity of the arms and legs during joint antiphase movements. Thus, both the spatial organization of movements and the type of afferent influences are significant factors of interlimb interactions, which, in turn, determine the type of neural interconnections that are involved in movement regulation.     

Mutual influences of upper and lower extremities during cyclic movements

The possibility for the activation of muscles in a passive arm during its cyclic movements imposed by active movements of the contralateral arm or by an experimenter and the effect that the movements of lower extremities have on the activity of the arm muscles have been studied. In addition, the activity of the leg muscles was studied as dependent on the motor task performed by the arms. Ten healthy subjects performed antiphase arm movements with and without stepping-like movements of both legs in the supine position. The experiment was performed under three conditions for the arm movements: (1) both arms performed active movements; (2) one arm performed active movements, and the contralateral arm, being entirely passive, was forced to participate in movements; (3) the movement of the passive arm was caused by an experimenter. Under condition (2), additional loadings of 30 and 60 N were applied to the active arm. Under all conditions, the arm movements were performed with and without leg movements. The possibility for the activation of muscles in the arm performing passive movements has been demonstrated. To a large extent, this is possible due to an increase in the afferent inflow from the muscles of the contralateral arm. The electrical activity was modulated during cyclic arm movements and depended on the level of loading of the active arm. During the combined active movements of the arms and legs, the reduction in the activity of the flexor muscles of the shoulder and forearm was observed. In the case of passive stepping-like movements, the concomitant arm movements increased the magnitude of electromyographic bursts in most of the examined leg muscles. During active leg movements, a similar increase in electromyographic bursts was observed only in the m. biceps femoris (BF) and the anterior tibial muscle. An increase in the loading of one arm caused a significant increase in the EMG activity in most examined muscles of the legs. The data obtained provide additional proof for the existence of a functionally significant neuronal interaction between the arms, as well as between the upper and lower extremities, which is probably due to intraspinal neuronal connections.     

Dietary reconstruction of the Amphipithecidae (Primates, Anthropoidea) from the Paleogene of South Asia and paleoecological implications

The primate family, Amphipithecidae, lived during the early Cenozoic in South Asia. In this study, the diet of late middle Eocene amphipithecids from the Pondaung Formation (Central Myanmar) is characterized using three different approaches: body mass estimation, shearing quotient quantification and dental microwear analysis. Our results are compared with other Paleogene amphipithecids from Thailand and Pakistan, and to the other members of the primate community from the Pondaung Formation. Our results indicate a majority of frugivores within this primate community. Pondaungia and “Amphipithecus” included hard objects, such as seeds and nuts, in their diet. Folivory is secondary for these taxa. Myanmarpithecus probably had a mixed diet based on fruit and leaves. Contrasting results and a unique dental morphology distinguish Ganlea from other amphipithecids. These render interpretation difficult but nevertheless indicate a diet tending towards leaves and fruit. However, the anterior dentition of Ganlea suggests that this taxon engaged in seed predation, using its protruding canine as a tool to husk hard fruits and obtain the soft seeds inside. Bahinia and Paukkaungia, two other Pondaung primates, are small (<500 g) and therefore would have depended on insects as their source of protein. As such, they occupied a very different ecological niche from Pondaung amphipithecids. This primate community is then compared with the Eocene-Oligocene primate communities of the Fayum from North Africa. Similarities between the late middle Eocene Pondaung primate community and extant equatorial and tropical South American primate communities are noted.     

<Emphasis Type="Italic">In vitro</Emphasis> fertilization and preimplantation development in the primate

Nonhuman primates represent a strong model for examining the chromosomal, biochemical, and temporal normality of embryos produced byin vitro fertilization. Morein vitro fertilized embryos from the squirrel monkey(Saimiri sciureus) have been produced and examined than with all other primate species combined. In studies over a 13 year period a fertilization rate approximating 60 % has been developed in this species with 30% of these embryos proceeding to the two cell stage and 50% of these to the three-four cell stage. Chromosomal abnormalities (primarily missing or extra chromosomes) at a level of nine to 16% have been found, a value corresponding to that found inin vivo mating andin vitro fertilization in other species. An incidence of triploidy of 16.7% was observed. RNA and protein synthetic rates appear comparable with those of laboratory species subjected toin vitro fertilization and indicate the initial stages of metabolic activity of the newly formed embryo. Similarly, increases in estrogen incorporation appear after fertilization but no effect is observed in progesterone incorporation. Utilizing 2-deoxy-glucose and insulin, it was determined that the glucose requirement as an energy source for early preimplantationin vitro fertilized primate embryos is very low.Of very great importance is the temporal relationship of the development ofin vitro fertilized squirrel monkey embryos compared with similar development in other primates (including humans) afterin vivo andin vitro fertilization. An analysis of over a decade of work with the squirrel monkey embryos demonstrates a pattern of temporal development that is comparable with all other primate species that have been examined (including the human) and comparable with development afterin vivo fertilization.     

Behavioral responses of free-ranging rhesus monkeys to food shortage

During early July 1968, a severe food shortage occurred on Cayo Santiago, an island colony of free-ranging rhesus monkeys (Macaca mulatta). The food shortage produced striking changes in the behavior of the monkeys. Within the one social group intensively studied, the total frequencies of grooming, play and fights decreased significantly; the frequency of matings also dropped; body contact, displacements by other groups, and non-displacement movements decreased, but not in statistically significant amounts. Changes in the percentage of total grooming attributable to related and unrelated monkeys reflected the stability of the rhesus matriline. Comparisons made between Cayo Santiago and other primate groups under analogous situations reveal similar responses to food shortage.     

Activation of interlimb interactions increases the motor output in legs of healthy subjects: Study under the conditions of arm and leg unloading

The effect of arm movements and movements of individual arm joints on the electrophysiological and kinematic characteristics of voluntary and vibration-triggered stepping-like leg movements was studied under the conditions of horizontal support of the upper and lower limbs. The horizontal support of arms provided a significant increase in the rate of activation of locomotor automatism by noninvasive impact on tonic sensory inputs. The addition of active arm movements during involuntary stepping-like leg movements led to an increase in the EMG activity of hip muscles and was accompanied by an increase in the amplitude of hip and shin movements. The movement of the shoulder joints led to an increase in the activity of hip muscles and was accompanied by an increase in the amplitude of hip and shin movements. Passive arm movements had the same effect on induced leg movements. The movement of the shoulder joints led to an increase in the activity of hip muscles and an increase in the amplitude of movements of knee and hip joints. At the same time, the movement of forearms and wrists had a similar facilitating effect on the physiological and kinematic characteristics of rhythmic stepping-like movements, but influenced the distal segments of legs to a greater extent. Under the conditions of subthreshold vibration of leg muscles, voluntary arm movements led to activation of involuntary rhythmic stepping movements. During voluntary leg movements, the addition of arm movements had a significantly smaller impact on the parameters of rhythmic stepping than during involuntary leg movements. Thus, the simultaneous movements of the upper and lower limbs are an effective method of activation of neural networks connecting the rhythm generators of arms and legs. Under the conditions of arm and leg unloading, the interactions between the cervical and lumbosacral segments of the spinal cord seem to play the major role in the impact of arm movements on the patterns of leg movements. The described methods of activation of interlimb interactions can be used in the rehabilitation of post-stroke patients and patients with spinal cord injuries, Parkinson’s disease, and other neurological diseases.     

Influence of Vibration on Motor Responses in the Upper Arm Muscles under Stationary Conditions and during Voluntary and Evoked Arm Movements under Limb Unloading Conditions

Locomotion of mammals, including humans, is based on the rhythmic activity of spinal cord circuitries. The functioning of these circuitries depends on multimodal afferent information and on supraspinal influences from the motor cortex. Using the method of transcranial magnetic stimulation (TMS) of arm muscle areas in the motor cortex, we studied the motor evoked potentials (MEP) in the upper arm muscles in stationary conditions and during voluntary and vibration-evoked arm movements. The study included 13 healthy subjects under arm and leg unloading conditions. In the first series of experiments, with motionless limbs, the effect of vibration of left upper arm muscles on motor responses in these muscles was evaluated. In the second series of experiments, MEP were compared in the same muscles during voluntary and rhythmic movements generated by left arm m. triceps brachii vibration (the right arm was stationary). Motionless left arm vibration led to an increase in MEP values in both vibrated muscle and in most of the non-vibrated muscles. For most target muscles, MEP was greater with voluntary arm movements than with vibration-evoked movements. At the same time, a similar MEP modulation in the cycle of arm movements was observed in the same upper arm muscles during both types of arm movements. TMS of the motor cortex significantly potentiated arm movements generated by vibration, but its effect on voluntary movements was weaker. These results indicate significant differences in the degree of motor cortex involvement in voluntary and evoked arm movements. We suppose that evoked arm movements are largely due to spinal rather than central mechanisms of generation of rhythmic movements.     

Twist limits for late twisting double somersaults on trampoline

An angle-driven computer simulation model of aerial movement was used to determine the maximum amount of twist that could be produced in the second somersault of a double somersault on trampoline using asymmetrical movements of the arms and hips. Lower bounds were placed on the durations of arm and hip angle changes based on performances of a world trampoline champion whose inertia parameters were used in the simulations. The limiting movements were identified as the largest possible odd number of half twists for forward somersaulting takeoffs and even number of half twists for backward takeoffs. Simulations of these two limiting movements were found using simulated annealing optimisation to produce the required amounts of somersault, tilt and twist at landing after a flight time of 2.0 s. Additional optimisations were then run to seek solutions with the arms less adducted during the twisting phase. It was found that 3½ twists could be produced in the second somersault of a forward piked double somersault with arms abducted 8° from full adduction during the twisting phase and that three twists could be produced in the second somersault of a backward straight double somersault with arms fully adducted to the body. These two movements are at the limits of performance for elite trampolinists.     

Uniqueness of primate forelimb posture during quadrupedal locomotion

Among the characteristics that are thought to set primate quadrupedal locomotion apart from that of nonprimate mammals are a more protracted limb posture and larger limb angular excursion. However, kinematic aspects of primate or nonprimate quadrupedal locomotion have been documented in only a handful of species, and more widely for the hind than the forelimb. This study presents data on arm (humerus) and forelimb posture during walking for 102 species of mammals, including 53 nonhuman primates and 49 nonprimate mammals. The results demonstrate that primates uniformly display a more protracted arm and forelimb at hand touchdown of a step than nearly all other mammals. Although primates tend to end a step with a less retracted humerus, their total humeral or forelimb angular excursion exceeds that of other mammals. It is suggested that these features are components of functional adaptations to locomotion in an arboreal habitat, using clawless, grasping extremities.     

Effects of coupled upper limbs movements on postural stabilisation

The preference for in-phase association of coupled cyclic limbs movements is well described (mirror-symmetrical patterns) and this is demonstrated by the ease of performing in-phase movements compared to anti-phase ones. The hypothesis of this study is that the easiest movement patterns are those with minor postural activity. The aim of this study was to describe postural activity in standing subjects in the sagittal and frontal planes during the execution of three upper limbs tasks (single arm, in-phase, anti-phase) at four different frequencies (from 0.6 to 1.2 Hz).We employed six infrared cameras for recording kinematics information, a force platform for measuring forces exerted on the ground, and a system for surface electromyography (SEMG). Outcome measures were: upper limb range of movement and relative-phase, centre of pressure displacement (COP), screw torque (T_z) exerted on the ground, and SEMG recordings of postural muscles (adductor longus, gluteus medius, rectus femoris, and biceps femoris).Our results show that in both the planes the in-phase task resulted in less COP displacement, torque production, and postural muscles involvement than the anti-phase and single arm tasks. This reduced need of postural control could explain the ease of performing in-phase coupled limb movements compared with anti-phase movements.     

The adaptive and classificatory significance of certain quantitative features of the forelimb in primates

Quantitative osteological features of the primate brachium and antebrachium can be defined in relation to functionally significant contrasts in the principal muscle blocks. Such features, as shown by univariate and multivariate statistical study of a total of 525 sets of arm bones from 39 primate genera, demonstrate a broad correlation with the extent to which the forelimb is subject to tensile forces during locomotion. Man, in different features, resembles contrasting groups of non-human Primates; in sum total of such characteristics, he is unique.
Other dimensional characters, defining more general features of the arm bones differentiate individually, but more especially in multivariate combination, between major taxonomic groups within both Prosimii and Anthropoidea.
In combination, the two groups of dimensions give a separation that is primarily taxonomic but which has a locomotor underlay.
When, in sum total, the dimensional characters of the arm are combined with those of the shoulder (where, as established in earlier work, the separation produced is almost exclusively locomotor), the initial division is in accordance with locomotor function of the forelimb—and especially the extent to which the limb is subjected to tensile forces. But there is a strong taxonomic underlay.
A method is thus provided for enquiry into both the taxonomic and functional significance of arm bones of fossil Primates.
The work, in addition to endorsing the classical concept of Man's retention of primitive mobility in the distal segments of his forelimb, also re-emphasizes his uniqueness through arm mobility being associated with shoulder action principally caudal and ventral to the joint.     

Muscle fatigue does not lead to increased instability of upper extremity repetitive movements

Muscle fatigue alters neuromuscular responses. This may lead to increased sensitivity to perturbations and possibly to subsequent injury risk. We studied the effects of muscle fatigue on movement stability during a repetitive upper extremity task. Twenty healthy young subjects performed a repetitive work task, similar to sawing, synchronized with a metronome before and after performing each of two fatiguing tasks. The first fatigue task (LIFT) primarily fatigued the shoulder flexor muscles, while the second fatigue task (SAW) fatigued all of the muscles of the arm. Subjects performed each task in random order on two different days at least seven days apart. Instantaneous mean EMG frequencies (IMNF) decreased over both fatiguing tasks indicating that subjects did experience significant muscle fatigue. The slopes of the IMNF over time and the decreases in maximum force measurements demonstrated that the LIFT fatigue task successfully fatigued the shoulder flexors to a greater extent than any other muscle. On average, subjects exhibited more locally stable shoulder movements after the LIFT fatigue task (p=0.035). They also exhibited more orbitally stable shoulder (p=0.021) and elbow (p=0.013) movements after the SAW fatigue task. Subjects also had decreased cocontraction at the wrist post-fatigue for both tasks (p=0.001) and at the shoulder (p<0.001) for the LIFT fatigue task. Therefore, increased dynamic stability of these repeated movements cannot be explained by increased muscle cocontraction. Possible alternative mechanisms are discussed.     

Effect of Tendon Vibration on Hemiparetic Arm Stability in Unstable Workspaces

Sensory stimulation of wrist musculature can enhance stability in the proximal arm and may be a useful therapy aimed at improving arm control post-stroke. Specifically, our prior research indicates tendon vibration can enhance stability during point-to-point arm movements and in tracking tasks. The goal of the present study was to investigate the influence of forearm tendon vibration on endpoint stability, measured at the hand, immediately following forward arm movements in an unstable environment. Both proximal and distal workspaces were tested. Ten hemiparetic stroke subjects and 5 healthy controls made forward arm movements while grasping the handle of a two-joint robotic arm. At the end of each movement, the robot applied destabilizing forces. During some trials, 70 Hz vibration was applied to the forearm flexor muscle tendons. 70 Hz was used as the stimulus frequency as it lies within the range of optimal frequencies that activate the muscle spindles at the highest response rate. Endpoint position, velocity, muscle activity and grip force data were compared before, during and after vibration. Stability at the endpoint was quantified as the magnitude of oscillation about the target position, calculated from the power of the tangential velocity data. Prior to vibration, subjects produced unstable, oscillating hand movements about the target location due to the applied force field. Stability increased during vibration, as evidenced by decreased oscillation in hand tangential velocity.