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  NEURAL REPRESENTATION OF MOVEMENT TAU (τ)    by    HENG‐RU MAY TAN    A thesis submitted in partial fulfilment of the  requirements for the degree of    DOCTOR OF PHILOSOPHY    THE UNIVERSITY OF EDINBURGH  2007        THE UNIVERSITY OF EDINBURGH      DECLARATION    This thesis has been composed by the candidate   HENG‐RU MAY TAN.   The presented work is the candidate’s own,  and has not been submitted for any other degree or professional qualification.        SIGNED: __________________________________________________________    ON THE  _________________________________ OF THE YEAR  _________________        The presented work has been critically examined and appraised by    COLWYN TREVARTHEN, PhD.  Professor (Emeritus) of Child Psychology and Psychobiology   in the Department of Psychology, The University of Edinburgh, UK    GIUSEPPE PELLIZZER, PhD.  Associate Professor in the Department of Neuroscience,  The University of Minnesota, USA        ii   THE UNIVERSITY OF EDINBURGH  ABSTRACT  Neural representation of movement tau (τ)  Heng‐Ru May Tan    Supervisors:  Prof. David N. Lee1 and Prof. Apostolos P. Georgopoulos2   School of Philosophy, Psychology, and Language Sciences, Perception‐Movement‐Action Research Centre,  Perception‐in‐Action Laboratories, The University of Edinburgh.  2  Brain  Sciences  Center,  Departments  of  Neuroscience,  Neurology,  and  Psychiatry,  The  University  of  Minnesota Medical School, The University of Minnesota.  1        A  fundamental  aspect  of  goal‐directed  behaviour  concerns  the  closure  of  motion‐gaps  in  a  timely fashion. An influential theory about how this can be achieved is provided by the tau‐ theory (Lee, 1998). Tau (τ) is defined as the ratio of the current distance‐to‐goal gap over the  current  instantaneous  speed  towards  the  goal.  In  this  work  we  investigated  the  neural  representation  of  tau in  two  sets  of  experiments.  In  one  study  we  recorded  neuromagnetic  fluxes  (using  magnetoencephalography,  MEG)  from  the  whole  brain  of  human  subjects  performing  discrete  hand  movements  aimed  to  targets  in  space,  whereas  the  other  study  involved  recordings  of  single  cell  activity  from  prefrontal  and  posterior  parietal  areas  of  a  behaving  monkey  during  geometrical  shape‐copying  tasks.  These  two  studies  provided  complementary  information,  for  the  former  covered  the  whole  brain  (at  the  cost  of  weak  localization), whereas the latter used the finest neural grain (at the expense of limited brain  regions).  However,  the  two  studies  together  yielded  valuable  information  concerning  the  dynamic, time‐varying neural representation of tau, with respect to both integrated synaptic  events  in  neuronal  ensembles  (recorded  by  MEG)  and  neural  spike  outputs  (recorded  by  microelectrodes). The relations between neural signals and tau were analyzed using a linear  regression model where the time‐varying neural signal (magnetic field strength in fT or spike  density function) was the dependent variable and the corresponding value of movement tau  and speed were the independent variables. In addition, the model included an autoregressive  term  to  account  for  the  expected  correlated  errors,  given  the  time  series  nature  of  the  data.  The  neurophysiological  study  revealed  a  statistically  significant  (p 

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