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1.  Interlimb differences of directional biases for stroke production 
Directional preferences during center-out horizontal shoulder–elbow movements were previously characterized for the dominant arm. These preferences were attributed to a tendency to actively accelerate one joint, while exploiting largely passive motion at the other joint. Since the non-dominant arm is known for inefficient coordination of inter-segmental dynamics, here we hypothesized that directional preferences would differ between the arms. A center-out free-stroke drawing task was used that allowed freedom in the selection of movement directions. The task was performed both with and without a secondary cognitive task that has been shown to increase directional biases of the dominant arm. Mirror-symmetrical directional preferences were observed in both arms, with similar bias strength and secondary task effects. The preferred directions were characterized by maximal exploitation of interaction torques for movement production, but only in the dominant arm. The non-dominant arm failed to benefit from interaction torques. The results point to a hierarchical architecture of control. At the higher level, a movement capable to perform the task while satisfying preferences in joint control is specified through forward dynamic transformations. This process is mediated for both arms from a common neural network adapted to the dominant arm and, specifically, to its ability to exploit interaction torques. Dynamic transformations that determine actual control commands are specified at the lower level of control. An alternative interpretation that strokes might be planned evenly across directions, and biases emerge during movement execution due to anisotropic resistance of intrinsic factors that do not depend on arm dominance is also discussed.
PMCID: PMC3697123  PMID: 22076406
Arm movements; Optimal control; Interaction torque; Multijoint; Internal model; Secondary task
2.  Load emphasizes muscle effort minimization during selection of arm movement direction 
Directional preferences during center-out horizontal shoulder-elbow movements were previously established for both the dominant and non-dominant arm with the use of a free-stroke drawing task that required random selection of movement directions. While the preferred directions were mirror-symmetrical in both arms, they were attributed to a tendency specific for the dominant arm to simplify control of interaction torque by actively accelerating one joint and producing largely passive motion at the other joint. No conclusive evidence has been obtained in support of muscle effort minimization as a contributing factor to the directional preferences. Here, we tested whether distal load changes directional preferences, making the influence of muscle effort minimization on the selection of movement direction more apparent.
The free-stroke drawing task was performed by the dominant and non-dominant arm with no load and with 0.454 kg load at the wrist. Motion of each arm was limited to rotation of the shoulder and elbow in the horizontal plane. Directional histograms of strokes produced by the fingertip were calculated to assess directional preferences in each arm and load condition. Possible causes for directional preferences were further investigated by studying optimization across directions of a number of cost functions.
Preferences in both arms to move in the diagonal directions were revealed. The previously suggested tendency to actively accelerate one joint and produce passive motion at the other joint was supported in both arms and load conditions. However, the load increased the tendency to produce strokes in the transverse diagonal directions (perpendicular to the forearm orientation) in both arms. Increases in required muscle effort caused by the load suggested that the higher frequency of movements in the transverse directions represented increased influence of muscle effort minimization on the selection of movement direction. This interpretation was supported by cost function optimization results.
While without load, the contribution of muscle effort minimization was minor, and therefore, not apparent, the load revealed this contribution by enhancing it. Unlike control of interaction torque, the revealed tendency to minimize muscle effort was independent of arm dominance.
PMCID: PMC3538625  PMID: 23035925
Movement cost; Interaction torque; Multi-joint; Dominant arm; Muscle energy
3.  Control of Human Limb Movements: The Leading Joint Hypothesis and Its Practical Applications 
The leading joint hypothesis (LJH) offers a novel interpretation of control of human movements that involve multiple joints. The LJH makes control of each multijoint movement transparent. This review highlights effective applications of the LJH to learning of new motor skills and to analysis of movement changes caused by aging and motor disorders.
PMCID: PMC2965031  PMID: 20871237
arm movement; intersegmental dynamics; torque analysis; coordination; motor learning; motor disorder
4.  Biased Wrist and Finger Coordination in Parkinsonian Patients during Performance of Graphical Tasks 
Neuropsychologia  2009;47(12):2504-2514.
Handwriting impairments in Parkinson’s disease (PD) have been associated with micrographia, i.e. diminished letter size. However, dyscoordination of the wrist and fingers may also contribute to handwriting deterioration in PD. To investigate this hypothesis, right-handed PD patients and controls were tested in performance of three types of cyclic wrist and finger movements: drawing of two lines and a circle. The line drawing was performed with either simultaneous flexion and extension of the wrist and fingers (equivalent pattern resulting in a right-tilted line) or with wrist flexion/extension accompanied with finger extension/flexion (nonequivalent pattern resulting in a left-tilted line). Circle drawing required a specific phase difference between wrist and finger motions. Movements were performed with an inkless pen on a digitizer-tablet at two frequency levels. Consistent deformations of the circle into right-tilted ovals and lower variability in equivalent compared with nonequivalent lines revealed preference to produce right-tilted shapes. This preference became more apparent with increased movement speed and it was amplified in PD patients. Analysis revealed that the circle deformation emerged mainly due to reduction in relative phase, while wrist and finger amplitudes remained unchanged. The results suggest that PD causes deficit characterized by strong tendency to produce certain coordination patterns between wrist and finger motions. This deficit may significantly contribute to handwriting impairments in PD by reducing the dexterity in the production of the variety of shapes of the cursive letters. Furthermore, the deficiency revealed in wrist and finger coordination may represent a more general deficit affecting control of various multi-joint movements in PD.
PMCID: PMC2712583  PMID: 19410590
5.  Origins of Submovements during Pointing Movements 
Acta psychologica  2008;129(1):91-100.
Submovements that are frequently observed in the final portion of pointing movements have traditionally been viewed as pointing accuracy adjustments. Here we re-examine this long-lasting interpretation by developing evidence that many of submovements may be non-corrective fluctuations arising from various sources of motor output variability. In particular, non-corrective submovements may emerge during motion termination and during motion of low speed. The contribution of these factors and the factor of accuracy regulation in submovement production is investigated here by manipulating movement mode (discrete, reciprocal, and passing) and target size (small and large). The three modes provided different temporal combinations of accuracy regulation and motion termination, thus allowing us to disentangle submovements associated with each factor. The target size manipulations further emphasized the role of accuracy regulation and provided variations in movement speed. Gross and fine submovements were distinguished based on the degree of perturbation of smooth motion. It was found that gross submovements were predominantly related to motion termination and not to pointing accuracy regulation. Although fine submovements were more frequent during movements to small than to large targets, other results show that they may also be not corrective submovements but rather motion fluctuations attributed to decreases in movement speed accompanying decreases in target size. Together, the findings challenge the traditional interpretation, suggesting that the majority of submovements are fluctuations emerging from mechanical and neural sources of motion variability. The implications of the findings for the mechanisms responsible for accurate target achievement are discussed.
PMCID: PMC2600723  PMID: 18550020
arm kinematics; discrete; continuous; accuracy; variability
6.  Origins of submovements in movements of elderly adults 
Slowness is a well-recognized feature of movements in aging. One of the possible reasons for slowness suggested by previous research is production of corrective submovements that compensate for shortened primary submovement to the target. Here, we re-examine this traditional interpretation and argue that the majority of submovements in older adults may be a consequence rather than the cause of slowness.
Pointing movements in young and older adults were recorded. Conditions for submovement emergence were manipulated by using small and large targets and three movement modes: discrete (required stopping on the target), reciprocal (required reversal on the target), and passing (required crossing the target and stopping after that). Movements were parsed into a primary and secondary submovement based on zero-crossings of velocity (type 1 submovements), acceleration (type 2 submovements), and jerk (type 3 submovements). In the passing mode, secondary submovements were analyzed only after crossing the target to exclude that they were accuracy adjustments.
Consistent with previous research, the primary submovement was shortened and total secondary submovement incidence was increased in older adults. However, comparisons across conditions suggested that many submovements were non-corrective in both groups. Type 1 submovements were non-corrective because they were more frequent for large than small targets. They predominantly emerged due to arm stabilization and energy dissipation during motion termination in the discrete and passing mode. Although type 2 and 3 submovements were more frequent for small than large targets, this trend was also observed in the passing mode, suggesting that many of these submovements were non-corrective. Rather, they could have been velocity fluctuations associated predominantly with low speed of movements to small targets.
The results question the traditional interpretation of frequent submovements in older adults as corrective adjustments. Rather, the increased incidence of submovements in older adults is directly related to low movement speed observed in aging, whereas the relationship between submovement incidence and target size is a result of speed-accuracy trade-off. Aging-related declines in muscular control that may contribute to the disproportional increases in submovement incidence during slow movements of older adults are discussed.
PMCID: PMC2628348  PMID: 19014548
7.  The Role of Vision in the Control of Continuous Multijoint Movements 
Journal of motor behavior  2006;38(1):29-44.
The authors investigated whether visual fixations during a continuous graphical task were related to arm endpoint kinematics, joint motions, or joint control. The pattern of visual fixations across various shapes and the relationship between temporal and spatial events of the moving limb and visual fixations were assessed. Participants (N = 16) performed movements of varying shapes by rotating the shoulder and elbow joints in the transverse plane at a comfortable pace. Across shapes, eye movements consisted of a series of fixations, with the eyes leading the hand. Fixations were spatially related to modulation of joint motion and were temporally related to the portions of the movement where curvature was the highest. Gathering of information related to modulation of interactive torques arising from passive forces from movement of a linked system occurred when the velocity of the movement (a) was the lowest and (b) was ahead of the moving limb, suggesting that that information is used in a feedforward manner.
PMCID: PMC1941686  PMID: 16436361
coordination; limb control; oculomotor control

Results 1-7 (7)