PP 14

Local field potentials during decoupled visually-guided reach movements in the superior parietal lobule
P. F.Sayegh, K. M. Hawkins & L. E. Sergio .
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School of Kinesiology and Health Science, Centre for Vision Research, York University, Toronto, Ontario, Canada.

Introduction
The objective of our research is to understand how the brain plans and executes movements when the motions of the eye and hand must be decoupled. Our previous research has demonstrated that these decoupled movements are associated with activity in a network of brain regions that includes the superior parietal lobe (SPL) and dorsal premotor cortex 1,2 (PMd) . Here we examined the contribution of the local cell assemblies in SPL when the action of the eye and the hand are decoupled. We analysed local field potentials (LFP) recorded simultaneously from multiple electrodes. We observed changes in SPL when the eye and hand were decoupled which may reflect a signal allowing gaze and hand to decouple. We have 3 previously observed similar task dependent changes in PMd . Taken collectively, these results suggest that SPL and PMd are key players in our ability to decouple gaze direction from hand direction.
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Behavioural Results
A. Monkey A Standard B. Monkey A Non-Standard
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Preferred direction lost when decoupling eye and hand:

Standard vs. Non-standard (mean RT)

Mean Reaction Time (msec)

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Non-standard Condition
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C. Monkey B Standard D. Monkey B Non-Standard

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Error bars: +/- SE

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Figure 3: Mean reach trajectories. Purple lines: mean movement trajectories, blue tick marks: standard deviations. Yellow asterisks denote trajectory segments that were significantly (p < .05)more variable in comparison to the standard condition.

Figure 4: Mean reaction times. Mean reaction time comparison between the standard and nonstandard conditions (Monkeys A and B; N = 46). Vertical bars denote +/- 1 standard error. There was no significant difference (=0.01).

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Figure 9: Population time-frequency spectrogram`s directional tunning during instructed delay period (IDP): Population spectrograms have been aligned to preferred direction (0 degrees, right target). Mean spectrograms are aligned to cue onset (black line). A.) Shows results for monkey A demonstrating an increase in 0-50 Hz in the preferred direction following cue. This increase in power is lost when performing the non-standard condition, N = 26. B.) shows similar results for monkey B, N = 7.

Some LFPs respond to gaze while other follow hand:

Neurophysiological Results
Loss in power when eye-hand are decoupled during IDP period but not during movement Increased coherency when eye and hand are decoupled
Standard condition Non-standard condition IDP

Movement period Non-standard Standard Non-standard Standard Condition Condition Condition Condition 3 gaze hand 2

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Time from move onset (msec) 1 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2

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Methods
We examined eye and hand movement related LFP activity in the superior parietal lobule (SPL) during standard and non-standard situations (Figure 2). Monkeys (2 female macaca mulatta) were trained to fixate on a central target throughout the instructed delay period, then to move their eyes and hand to one of eight peripherally cued targets and hold them there throughout the target hold period.The full trajectory of the hand and eye were recorded to ensure that the motor task remained similar between conditions. Eye movements were monitored using the ISCAN-ETL 200 Eye Tracking System (ISCAN Inc, Burlington MA) at a sampling rate of 1KHz. Hand paths were monitored using a touch sensitive screen (100Hz, Touch Controls Inc, San Diego CA). A four electrode microdrive (FHC Inc.) was used in conjunction with a multi-unit recording system (Alpha-Omega Engineering, Israel) to collect single unit (12.5kHz) and waveform (1562.5 Hz) activity. Data were analyzed in Matlab (Mathworks, USA) using both custom written and open source (Chronux.org) programmes.

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Figure 9: Population time-frequency spectrogram`s directional tunning during movement (MOVE): Population spectrograms have been aligned to preferred direction (0 degrees, right target). Mean spectrograms are aligned to cue onset (black line). A.) Demonstrates an increase in 0-50 Hz power in the preferred direction during the standard condition and a 90 degree rotation in preferred direction during the non-standard task, N = 20. B.) Demonstrates a population of LFPs that did not show a rotation in preferred direction between tasks but maintained their preferred direction, N = 13.

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Summary and Conclusions

We did not observe topographical changes within the recorded area of SPL During the early planning phase we observe:
Decrease in oscillatory activity when the eye and hand were decoupled. Increase in power for the preferred direction during the standard condition.

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Frequency (Hz) 80 60 40 20
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MOVE

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Instructed delay period Movement period

z transformed coherence

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Eye and hand congruent

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These observations may reflect a change in the inputs to this area when vision and proprioception are no longer congruent. During the late planning we observe:
Some LFPs rotated their preferred direction when going from standard to non-standard mapping. Some LFPs did not rotate their preferred direction between conditions.

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Eye fixation, central target

Figure 2: Experimental setup and trial timing. A) Schematic of the standard and dissociated conditions. B) During each trial, one of eight equally spaced (45) peripheral targets were presented on either a touch-sentivie screen placed over the animals lap or on a monitor positioned vertially 40 cm away from the animals frontal plane. Arm movements were always made over the horizontal touch screen. Light grey circles represents the eight possible target locations (not illuminated before cue). Epochs - CHT: centre hold time, IDP: instructed delay period, RT: reaction time, MT: movement time, THT: target hold time. Red horizontal line represents the time in which the animal had to maintain fixation and hand inside the center target. The animal's head was fixed throughout the experiment.

Figure 5: Population time-frequency spectrograms of SPL activity during IDP and MOVE periods: A.) Population spectrograms are aligned to cue onset and show an increase in oscillations from 0-50Hz following cue presentation during both conditions with greater power during the standard task when compared to the non-standard task (N=26). Black line indicates cue onset. Gray dashed line indicates end of the baseline period. B.) Population spectrograms during the early movement period demonstrating an increase in power within the 0-30Hz range following movement onset for both conditions. Black line indicates movement onset.

Frequency (Hz)

Frequency (Hz)

Figure 6: Field-field coherency during instructed delay and movement epochs: A) Example field-field coherence aligned to cue onset showing an increase in coherency in the non-standard condition when compared to the standard condition. B) Population field-field coherence during the early planning phase (instructed delay period) showing an increasein coherence during the non-standard condition in the in the 0-25Hz and 74-100Hz frequency ranges. C) Population fieldfield coherence during early movement showing an increase in coherence during the non-standard condition in the 0-23Hz and 51100Hz range. * denotes significant difference (p < 0.05).

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The rotation in preferred direction may be related to the change in gaze direction that accompanies the non-standard mapping. Those cell assemblies that did not show this rotation may reflect the hand position, which remains constant between the two conditions. These data suggest that communication between the local cell assemblies in PPC plays a role in the visuomotor transformations required to decouple the eye and the hand.
References: 1. Gorbet et. al. (2004) Neuroimage, 23: 1100-1111. 2. Granek et. al. (2010) Cortex, 46(9): 1165-1177. 3. Sayegh et. al. (2009) SFN 455.0 4. Pesaren et. al. (2008) Nature, 453: 406-409.

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Adapted from Kalaska et. al., Curr. Op. Neurobiol. 1997.

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Figure 1: Cortical regions accessed by chamber placement: The red circle represents location of chamber (19mm diameter).

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Non-standard condition IDP

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