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Influence of handedness on motor unit discharge properties and force tremor.

Discharge properties of motor units (MUs) in the first dorsal interosseous muscle (FDI) were studied in the dominant and non-dominant hands of six right-handed (RH) and six left-handed (LH) individuals. MU discharge rates and variability were similar in each hand in RH (186 MUs) and LH (160 MUs) subjects. MU synchronization was less prominent in the dominant hand of RH subjects, with 51% (45/88) of cross-correlograms of MU discharge having significant central peaks, compared with 81% (90/111) for the non-dominant hand. The strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was weaker in the dominant hand of right-handers (0.23 +/- 0.03 s-1 vs 0.39 +/- 0.03 s-1), and synchronous peaks from that hand were slightly broader. Four of six RH subjects had significant differences in synchronization between hands (weaker in dominant hand). In contrast, left-handers had similar incidence (80 vs 82%, n = 161) and strength (0.41 +/- 0.03 s-1 vs 0.37 +/- 0.03 s-1) of MU synchrony in dominant and non-dominant hands. No LH subject had a significant difference in synchronization between hands. Force tremor was quantified in each hand in the same subjects during isometric abduction of FDI at 0.5 N and 3.5 N, and directly correlated with the extent of MU synchronization in the muscle. Tremor root mean square amplitude was similar in dominant and non-dominant hands. Power spectral analysis of the tremor force revealed that the peak frequency in the power spectrum was not influenced by handedness, but power at the peak frequency was higher in the non-dominant hand of RH subjects. Correlations between MU discharge variability and synchrony with measures of tremor amplitude were weak. The reduced MU synchronization in the dominant hand of right-handers may reflect a more restricted distribution of direct projections from motor cortical neurons within the FDI motoneuron pool, or reduced excitability of the cortical neurons during the task. These differences in MU synchronization, however, had an insignificant influence on the magnitude of physiological tremor in the FDI.

Adult↗

Symmetry, broken symmetry, and handedness in bimanual coordination dynamics.

The symmetrical dynamics of 1:1 rhythmic bimanual coordination may be specified by an order parameter equation involving the relative phase between rhythmic components, and an interlimb coupling which determines the relative attractiveness of in-phase and anti-phase patterns. Symmetry breaking of these dynamics can occur via the difference in the natural frequencies, delta omega, of the left and right rhythmic components, or by the intrinsic asymmetrical dynamics of the body. The latter is captured by additional terms that render the symmetrical coupling slightly anisotropic. A major prediction resulting from this step is that although delta omega = 0, as the frequency of coordination is increased, the asymmetrical coupling will increase and the symmetrical coupling will decrease. This results in a greater left-limb bias in left-handers and right-limb bias in right-handers. This "increased handedness" prediction was confirmed in an experiment in which 20 left-handed and 20 right-handed individuals performed 1:1 coordination with hand-held rigid pendulums. Manipulations of left and right pendulum lengths controlled delta omega, and the coupled frequency was determined by a metronome. Also confirmed was the prediction that the small shift in equilibria from in-phase and anti-phase due to the intrinsic asymmetry should be amplified in left-handers when delta omega > 0 and in right-handers when delta omega < 0. Further, the bias in left-handers was more consistent than the bias in right-handers, and a subgroup of right-handers was identified who performed similarly to left-handers. The coordination dynamics of functional asymmetry provides insights into the elementary synergy between the limbs, the dynamical mechanism that modulates it, and the nature of the asymmetry in left-handed and right-handed individuals.

Adult↗

"Handedness" of spiral fractures of the tibia.

"Handedness" is a basic property of spirals. In many cases it is possible to determine whether spiral fractures are right or left handed by analysis of radiographs. Spiral fractures of the tibia show striking bilateral symmetry, with right-handed spirals predominating on the left side of the body, and left-handed spirals on the right.

Humans↗

Concerning the origin of handedness in humans.

The origin of right-handedness as the predominant chirality in humans seems to be related to the tendency of human (and presumably prehuman) mothers to hold infants on the left side. The latter practice has previously been ascribed to imprinting and the soothing sound of the mother's heartbeat on the infant. Given the practice of holding the child in this manner, dextral mothers will be more skillful at manipulation of objects and selectively favored.

Biological Evolution↗

Influence of handedness on calcaneal ultrasound: implications for assessment of osteoporosis and study design.

Calcaneal ultrasound has been increasingly studied for its potential in the assessment of osteoporotic fracture risk. The accuracy of such an assessment is, in part, dependent on the reproducibility of the measurement. This study examines the impact of handedness on ultrasound measurements [broadband ultrasound attenuation (BUA) and velocity of sound (VOS)] in the calcaneus. Two hundred and sixty-four subjects (57 men and 297 women) aged 51.1 +/- 13.6 years (mean +/- SD) were studied. For each subject, calcaneal ultrasound measurements were performed on both heels with a McCue CUBA ultrasound densitometer. Right-handed dominance (94.7%) was determined by structured interview. In men, BUA measurements were significantly higher on the dominant side: mean difference 4.1 +/- 1.5 dB/MHz (mean +/- SD; p = 0.009), equivalent to 4.2 +/- 1.5% and more than 4 times the average rate of annual change in BUA. The difference between sides was greater in young (< 50 years) than old men (> 50 years). Among the women, the difference was not statistically significant (0.7 +/- 0.9 dB/MHz; p = 0.4); however, it was significant in younger women (20-30 years) (99 +/- 4 vs 90 +/- 4 dB/MHz, p = 0.01). By contrast VOS did not differ between sides in either men or women irrespective of age. Within-subject standard deviation of BUA was 9.8 dB/MHz for men and 8.6 dB/ MHz for women and the component due to right and left difference was 8.4 dB/MHz for men and 6.9 dB/MHz for women. This variability of BUA between right and left heels could increase the false-positive rate by up to 28% for a cut-off of 2 SD below the mean. These data indicate that variation between left and right heel measurements of BUA is higher than that of random error measurements, particularly in men and younger, presumably more physically active subjects. Although VOS measurements were not side dependent, in the smaller number of studies examining VOS and fracture risk, VOS appears to have a weaker predictive power than BUA. Clinical and epidemiological studies involving calcaneal BUA measurements should standardize the side measured to either the dominant or non-dominant heel, to reduce within-subject variation and increase their power.

Adult↗

On the origin of molecular "handedness" in living systems.

Elementary particle effects (beta-decay) provide at best only a weakly handed radiation in the biologically effective energy ranges. Global magnetic effects coupled to sunlight are randomized by paleomagnetic reversals. Hence a persistent terrestrial handed bias at possible local biopoetic sites offers a more promising explanation for the origin of the "handedness" of the molecules found among living systems on earth. Magnetite in lava flows maintains a handed bias for surface catalysis through many magnetic reversals. Magnetite contaminated with sulfur has already been proposed by Granick as a biopoetic site because it provides a weak source of chemical energy derived by photochemical conversion. Indirect evidence for this hypothesis has been provided by the molecular structure of ferredoxin - a single strand of the 14 primordial amino acids wrapped around an FeS core. Lava flows have been suggested as biopoetic sites by Fox, since their temperature and chemical composition might allow for the rapid synthesis of prebiotic compounds at the surface of the primitive earth. The additional fact that magnetite in lave flows also provides a persistent handed site for surface catalysis offers a further argument for the experimental investigation of this specific biopoetic environment.

Biological Evolution↗

Amplification of chirality based upon the association of nucleic acid strands of opposite handedness.

It is proposed that nucleotide strands of opposite handedness may strongly associate and thereby provide the key step of a mechanism for the amplification of a small enantiomeric excess in an initially near-racemic mixture of poly- or oligonucleotides. This hypothesis, if confirmed by experimentation, may have important implications for the question of the origin of biomolecular chirality. The results of preliminary NMR experiments are given, which do show evidence of a strong association between pentanucleotide RNA strands whose monomers have opposite chirality. Simple kinetic equations are solved to demonstrate the conditions under which such association can produce amplification of chirality.

DNA↗

Force synergies for multifingered grasping: effect of predictability in object center of mass and handedness.

To grasp with five digits of the hand requires an efficient parceling of contact forces in order to maintain static equilibrium as an object is lifted and held. In a previous study, subjects were asked to reach, grasp and lift a five-digit grip apparatus whose center of mass (CM) location was changed for each block of trials. Despite a modulation of force sharing patterns among the digits as a function of center of mass location, consistent in-phase and out-of-phase relationships between normal forces were found in the frequency domain. In the present study, we have used the same task to assess the effect of (a) predictability of an object's CM location (random vs blocked presentation) and (b) handedness (dominant vs non-dominant hand). Contrary to our original expectations, we found a similar modulation of normal forces to CM location during the hold phase across all conditions. Specifically, the force sharing pattern, i.e., the rank order of force contributed by each digit, emerged very early in the grasp sequence, remaining relatively stable throughout the duration of the lift and hold. Nevertheless, the extent to which force sharing patterns could be discriminated as a function of CM location was lower in the random than in the blocked conditions. Lastly, normal forces exerted by pairs of digits tended to be synchronized, both in-phase (thumb and fingers) and out-of-phase (pairs of digits) across a large proportion of the functional frequency range (up to 10 Hz) in all conditions. The composite of these findings suggests that the central nervous system uses stereotyped control strategies for coordinating multiple grip forces during grasping. Specific aspects of these schemes appear to be affected by predictability of object CM location, but not by hand dominance.

Adult↗

The effects of attention and handedness on coordination dynamics in a bimanual Fitts' law task.

Two experiments were conducted to investigate the effects of attention and handedness on bimanual coordination in the context of a dynamical model of coordinated movements. Participants performed a bimanual, rhythmic Fitts' law task in which the relative amount of attention directed to each task was manipulated by the relative difficulty associated with the pair of targets that each hand tapped. In both experiments, participants tended to lead with their preferred hand. The effects of attention, though, were mixed, which suggested that there was a combined effect of an attentional asymmetry and an asymmetry in the hands' uncoupled frequency, both of which are captured in the dynamical model.

Adult↗

Interlimb transfer of visuomotor rotations depends on handedness.

We previously reported that opposite arm adaptation to visuomotor rotations improved the initial direction of right arm movements in right-handers, whereas it only improved the final position accuracy of their left arm movements. We now investigate the pattern of interlimb transfer following adaptation to 30 degrees visuomotor rotations in left-handers to determine whether the direction of transfer depends on handedness. Our results indicate unambiguous transfer across the arms. In terms of final position accuracy, the direction of transfer is opposite to that observed in right-handers, such that transfer only occurred from the left to the right arm movements. Directional accuracy also showed the opposite pattern of transfer to that of right-handers: initial movement direction, calculated at peak tangential acceleration, transferred only from right to left arms. When movement direction was measured later in the movement, at peak tangential velocity, asymmetrical transfer also occurred, such that greater transfer occurred from right to left arms. However, a small, but significant influence of opposite arm adaptation also occurred for the left arm, which might reflect differences in the use of the nondominant arm between left- and right-handers. Overall, our results indicate that left-handers show a mirror-imaged pattern of interlimb transfer in visuomotor adaptation to that previously reported for right-handers. This pattern of transfer is consistent with the hypothesis that asymmetry in interlimb transfer is dependent on differential specialization of the dominant and nondominant hemisphere/limb systems for trajectory and positional control, respectively.

Adolescent↗

Handedness-related asymmetry in transmission in a system of human cervical premotoneurones.

The possibility was investigated that human handedness is associated with an asymmetrical cortical and/or peripheral control of the cervical premotoneurones (PreMNs) that have been shown to mediate part of the descending command to motoneurones of forearm muscles. Heteronymous facilitation evoked in the ongoing voluntary extensor carpi radialis (ECR) electromyographic activity (EMG) by weak (0.8 times motor threshold) stimulation of the musculo-cutaneous (MC) nerve was assessed during tonic co-contraction of biceps and ECR. Suppression evoked by stimulation of a cutaneous nerve (superficial radial, SR) at 4 times perception threshold in both the voluntary EMG and in the motor evoked potential (MEP) elicited in ECR by transcranial magnetic stimulation (TMS) was investigated during isolated ECR contraction. Measurements were performed within time windows or at interstimulus intervals where peripheral and cortical inputs may interact at the level of PreMNs. Results obtained on both sides were compared in consistent right- and left-handers. MC-induced facilitation of the voluntary ECR EMG was significantly larger on the preferred side, whereas there was no asymmetry in the SR-evoked depression of the ongoing ECR EMG. In addition, the suppression of the ECR MEP by the same SR stimulation was more pronounced on the dominant side during unilateral, but not during bilateral, ECR contraction. It is argued that (1) asymmetry in MC-induced facilitation of the voluntary EMG reflects a greater efficiency of the peripheral heteronymous volley in facilitating PreMNs on the dominant side; (2) asymmetry in SR-induced suppression of the MEP during unilateral ECR contraction, which is not paralleled by a similar asymmetry of voluntary EMG suppression, reflects a higher excitability of cortical neurones controlling inhibitory spinal pathways to cervical PreMNs on the preferred side.

Adult↗

Motor performance: normative data, age dependence and handedness.

Quantitative, apparatus-based assessment of motor performance is a useful tool when addressing questions of therapy evaluation, quality control and early detection of movement disorders. Until now, few such methods have been sufficiently standardised to allow their routine employment in the clinic. Before assessing the impaired performance of patients, the first stage of standardisation is the determination of criteria of normal performance in healthy subjects. We have standardised a multidimensional test battery for motor performance of the upper extremities (Schoppe's "motorische Leistungs-Serie" = MLS). On the basis of the performance of 80 healthy controls, normative algorithms which allow correction of performance with reference to age were calculated. Further, our investigation provided information concerning the complexity of normal motor performance and the question of laterality (handedness). Standardised apparatus-based quantification of performance promises to be a valuable supplement to rating scales in the evaluation of patient motor skills, particularly in the early stages of their illness.

Adult↗

The effect of handedness on spinal and supra-spinal reflex excitability.

Twenty-four subjects, 8 right-handers, 8 left-handers, and 8 familial left-handers, participated in a study aimed at understanding the relations between handedness, cerebral asymmetry, and a possible asymmetry in spinal excitability. This was accomplished by employing 3 different techniques of investigating the H reflex, measuring 3 different parameters of spinal excitability. The recovery curve height (RCH) of left-handers was significantly higher than the RCH of right-handers, and, as well, familial left-handers exhibited higher RCH than non-familial left-handers. The recovery ability was not found to be significantly more inhibited on the dominant side, as would be expected from previous work. Instead, recovery ability was associated with the fractional involvement of the motoneuron pool in the H process (H/M ratio). RCH was highly correlated with both hand preference and hand performance. The above results suggest involvement of a long-loop transcortical reflex in lateral asymmetries.

Adolescent↗

Depicting topology and handedness in jellyroll structures.

The jellyroll structure is a special case of the Greek key topology and, to date, has only been observed in complete form in one of its four possible arrangements. Like other elements of super-secondary structure involving the beta-strand (e.g. the beta alpha beta unit) the known structure forms a right-handed superhelix. The possibility of losing such tertiary information and other problems associated with representing these structures by two-dimensional topology diagrams are discussed. A series of rules are presented which allow this three-dimensional information to be represented in two-dimensional topology diagrams from which the handedness of a jellyroll structure can be determined.

Protein Conformation↗

Pathologic left-handedness: does it exist?

The concept of pathologic left-handedness is reviewed, both from a historical and an empirical point of view. It is suggested that there is no adequate evidence to justify its continued use. The fact that the concept is still much used may be the result of a desire to restore to the brain its lost symmetry, by allowing Dax's Law once more to be true.

Adult↗

Sex, handedness and asymmetry of hand and foot length.

Measurements of the length of the hands of 156 4-9 yr old children revealed a significant association among sex, handedness and direction of asymmetry of hand-length. Right-handed males have longer right than left hands while non-right-handed males have longer left than right hands (P less than 0.005). Females have a significant association also (P less than 0.005), with the direction of asymmetry the reverse of that found in males. No significant associations were found among the classification variables for foot-length measured at ages 4-9 or at birth.

Child↗

Spatial lateralization: the relationship among sex, handedness and familial sinistrality.

Spatial lateralization was examined in 80 subjects by use of a strictly balanced design contrasting sex, handedness and familial sinistrality (absent and present). A modified version of Witelson's dichaptic stimulation task (verbal and nonverbal) was employed. The verbal task showed a significant main effect for the right-hand score. On the nonverbal task only two groups displayed a significant difference between hand scores: left-handed males with familial sinistrality and right-handed females with no familial sinistrality. Results are discussed in relation to previously reported data.

Adolescent↗

Sex, handedness and weight discrimination.

Manual asymmetries in weight discrimination were investigated by testing 50 subjects with their left hand and 52 with their right hand, the groups being approximately balanced for sex and hand preference. Performance was better for the left-tested group, and for strongly handed males tested with the dominant hand. Female performance was unrelated to handedness. There was no overall superiority due to sex, males being superior with the dominant hand and females with the nondominant hand. The results are interpreted in terms of spatial hemisphere involvement, hand dominance, and the greater hemispheric specialization of males.

Adult↗