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D J Crammond

Publications and source records attributed to D J Crammond.

11 recordsLinked to original sources

Prior information in motor and premotor cortex: activity during the delay period and effect on pre-movement activity.

In instructed-delay (ID) tasks, instructional cues provide prior information about the nature of a movement to execute after a delay. Neuronal responses in dorsal premotor cortex (PMd) during the instructed-delay period (IDP) between the CUE and subsequent GO signals are presumed to reflect early planning stages initiated by the prior information. In contrast, in multiple-choice reaction-time (RT) tasks, all motor planning and execution processes must occur after the GO signal. These assumptions predict that neuronal planning correlates recorded during the IDP of ID trials should share common features with early post-GO activity in RT trials, and that those response components need not be recapitulated after the GO signal of ID trials. These two predictions were tested by comparing activity recorded in RT and ID tasks from 503 neurons in PMd and caudal (MIc) and rostral (MIr) primary motor cortex. The incidence and strength of directionally tuned IDP activity declined progressively from PMd to MIc. The directional tuning of activity during the IDP of ID trials was more similar to that in the reaction-time epoch (RTE) of RT trials than after movement onset, especially in PMd. A modulation of post-GO activity was often observed between RT and ID trials and was confined mainly to the RTE. This effect was also most prominent in PMd. The most common change was a reduction in intensity of short-latency phasic responses to the GO signal between RT and ID trials, especially in PMd cells with a short-latency phasic response to CUE signals. However, the largest group of cells in each area showed no large change in peak RTE activity between RT and ID trials, whether they were active in the IDP or not. Since early phasic CUE-related responses are least likely to be recapitulated after the GO signal in ID trials, they may be a neuronal correlate of an early planning stage such as response selection. Tonic IDP responses, which are not as strongly associated with a post-GO reduction in activity, may be related to other aspects of motor planning and preparation. Finally, a major component of the movement-related activity in both MI and PMd is not susceptible to modification by prior information and is indivisibly coupled temporally to movement execution.

Analysis of Variance↗

Differential relation of discharge in primary motor cortex and premotor cortex to movements versus actively maintained postures during a reaching task.

The activity of cells in primary motor cortex (MI) and dorsal premotor cortex (PMd) were compared during reaching movements in a reaction-time (RT) task, without prior instructions, which required precise control of limb posture before and after movement. MI neurons typically showed strong, directionally tuned activity prior to and during movement as well as large gradations of tonic activity while holding the limb over different targets. The directionality of their movement- and posture-related activity was generally similar. Proximal-arm muscles behaved similarly. This is consistent with a role for MI in the moment-to-moment control of motor output, including both movement and actively maintained postures, and suggests a common functional relation for MI cells to both aspects of motor behavior. In contrast, PMd cells were generally more phasic, frequently emitting only strong bursts of activity confined mainly to the behavioral reaction time before movement onset. PMd tonic activity during different postures was generally weaker than in MI, and showed a much more variable relation with their movement-related directional tuning. These results imply that the major contribution of PMd to this RT task occurred prior to the onset of movement itself, consistent with a role for PMd in the selection and planning of visually guided movements. Furthermore, the nature of the relative contribution of PMd to movement versus actively maintained postures appears to be fundamentally different from that in MI. Finally, there was a continuous gradient of changes in responses across the rostrocaudal extent of the precentral gyrus, with no abrupt transition in response properties between PMd and MI.

Animals↗

Modulation of preparatory neuronal activity in dorsal premotor cortex due to stimulus-response compatibility.

1. Neuronal activity was recorded in the dorsal premotor cortex (PMd) of two monkeys performing a multidirectional, instructed-delay (ID) reaching task in which visuospatial cues signaled the direction of movement either congruent with the instruction cue ("direct-delay" trials, DD) or redirected 180 degrees opposite to the cue ("redirected-delay" trials, RD). Therefore, this task had two degrees of stimulus-response (S-R) compatibility because in one-half of the trials the spatial attributes of the visual cue were incongruent with those of the intended movement. 2. The majority of PMd cells discharged both at short latency to the RD or DD cues and subsequently with sustained activity during the remaining ID period (IDP). The earliest responses (< 250 ms) in both DD and RD trials covaried with cue location and so could be either a "visuospatial" response or a neuronal correlate of the selection of action with highest S-R compatibility, namely move to the stimulus. In contrast, later IDP activity usually covaried with the direction of movement signaled by the cues, independent of their spatial location, supporting the hypothesis that IDP discharge in PMd ultimately encodes attributes of intended reaching movements.

Animals↗

Cerebral cortical mechanisms of reaching movements.

Because reaching movements have a clear objective--to bring the hand to the spatial location of an object--they are well suited to study how the central nervous system plans a purposeful act from sensory input to motor output. Most models of movement planning propose a serial hierarchy of analytic steps. However, the central nervous system is organized into densely interconnected populations of neurons. This paradox between the apparent serial order of central nervous system function and its complex internal organization is strikingly demonstrated by recent behavioral, modeling, and neurophysiological studies of reaching movements.

Animals↗

Neuronal activity in primate parietal cortex area 5 varies with intended movement direction during an instructed-delay period.

A monkey was trained to make arm movements to visual targets immediately after presentation of a GO signal, either in a visual reaction-time paradigm (CONTROL task), or after an instructed-delay period of variable duration, during which a CUE stimulus signalled the direction of the impending movement (DELAY task). The activity of 98 area 5 cells recorded in 2 hemispheres varied with movement direction in the CONTROL task. This included 60 "early" cells which showed directional activity changes prior to movement onset. In the DELAY task, 54/98 cells (55%) showed activity changes during the instructed-delay period which varied with the direction of the impending movement. Most of these (45/54, 83%) were "early" cells. Forty proximal arm-related cells were recorded in adjacent area 2. In contrast to area 5, only 2/40 area 2 cells showed any evidence of changes in activity varying with intended movement direction during the instructed-delay period. The origin of area 5 activity changes during an instructed-delay period which are related to intended direction of a delayed movement is uncertain, but its presence is consistent with a number of proposed roles for area 5.

Animals↗

Neuronal correlates in posterior parietal lobe of the expectation of events.

During studies of response properties of single units in the posterior parietal cortex of 6 awake monkeys, 168 neurons were encountered (7.1% of examined units) which showed anticipatory types of activity. These neurons were found on either side of the intraparietal sulcus. In area 5, this expectation activity was expressed as a change in discharge rate whenever a specific body part (e.g. hand or shoulder) was approached by the investigator as though contact would be made. Invariably the neurons also responded to cutaneous and/or proprioceptive stimulation of the target body area. In area 7a the same type of response was also found but not always with a corresponding somatosensory receptive field. In addition, many neurons increased discharge rates (or rarely, decreased them) immediately prior to the expected occurrence of a reward, a visual task cue, or on hearing the approaching footsteps of a familiar person. None of these responses were correlated to eye movements, nor could they be attributed to any other body movement.

Action Potentials↗

Evoked potentials from passive elbow movements. II. Modification by motor intent.

Subjects were instructed to remain passive or to react to a forearm perturbation by opposing the imposed movement. Evoked potentials (EPs) were recorded at 8 scalp sites in both conditions. In the React condition, reflexes were observed in the EMG (mean onsets of 67 and 81 msec) and the EP was modified. Source derivation techniques revealed that the earliest cortical response (31 msec) across the central sulcus was not changed. Therefore the intention to react did not seem to affect afferent transmission to the primary sensorimotor cortex. Two periods of modulation were observed. In both, parietal and frontal potentials were modulated together, prior to the reflex components. After 70 msec, the pattern of potential gradients which occurred in the Passive case was accelerated and intensified in the React condition. The overall effect was to focus a larger zone of negativity over motor cortex at the time of triggered EMG output (109 msec). The earlier changes in cortical activity could be causally related to the appearance of the late stretch reflex. Since parietal and frontal areas were principally involved and not the motor area, it is suggested that the former exert a modulatory influence on spinal and brain-stem reflex centres.

Elbow↗

Measurements of human forearm viscoelasticity.

In human subjects, stiffness of the relaxed elbow was measured by three methods, using a forearm manipulandum coupled to a.d.c. torque motor. Elbow stiffness calculated from frequency response characteristics increased as the driving amplitude decreased. Step displacements of the forearm produced restoring torques linearly related to the displacement. The stiffness was very similar to that calculated from natural frequencies at amplitudes above 0.1 rad. Thirdly, elbow stiffness was estimated from brief test pulses, 120 ms in duration, by mathematically simulating the torque-displacement functions. Stiffness values in the limited linear range (under +/- 0.1 rad) were higher than in the linear range of the first two methods. A major component of elbow stiffness appears to decay within 1 s. The coefficients of viscosity determined from the simulation were, however, very similar to those calculated from the frequency response. Test pulse simulation was then used to determine joint impedance for different, actively maintained elbow angles. Joint stiffness and viscosity increased with progressive elbow flexion.

Adult↗

Evoked potentials from passive elbow movements. I. Quantitative spatial and temporal analysis.

Evoked potentials following perturbations of the forearm were recorded monopolarly at 8 scalp sites. Successive 10 msec bins of individual EPs were compared across subjects to determine intervals of consistent potential change. From this analysis it was possible to objectively sort subjects into two subpopulations with different common wave forms. Spatial vectors (potential gradient between two adjacent leads) were computed as well as Laplacian derivatives, to identify electrodes closest to source activity. Subsequently, temporal derivatives of the vectors were computed to define the timing of statistically significant response phases. To more precisely localize the largest potential gradients, component vectors (or their derivatives) in the Laplacians were resolved algebraically on a scaled representation of the scalp. Convergence of these resultant vectors from adjacent areas, identified zones of significant potential change which corresponded to known somatosensory areas. The two subpopulations had initial responses of similar topography localized to the central sulcal region. Subsequently, for one group the area of activated cortex expanded to include posterior parietal and more frontal areas. Prestimulus negative potential shifts had different distributions for the two groups and are described in relation to the poststimulus differences.

Adult↗

Deciding not to GO: neuronal correlates of response selection in a GO/NOGO task in primate premotor and parietal cortex.

Monkeys performed reaching movements in two opposite directions in a symmetrically rewarded GO/NOGO task with an instructed-delay period. Instructional cues were presented at the target locations. The decision not to move was clearly reflected in cell activity in dorsal premotor cortex, but not in parietal cortex area 5. In premotor cortex, the initial response (< 250 msec) of most cells to the appearance of the instructional cues in GO and NOGO trials was similar. However, by the end of the delay period, the responses of most cells were statistically different between the two trial types, and the population signals were much less directional in the NOGO trials than in the GO trials. In area 5, in contrast, single-cell and population signals were generally similar and strongly directional in both GO and NOGO trials. This result suggests a role for area 5 in visuomotor analysis for the guidance of limb movements. It further suggests that separate representations of potential motor responses to external inputs and of the intended response to that input can coexist in parietal and premotor cortex, respectively.

Animals↗