What implications does the somatotopic organization of the primary motor

cortex have in the control of action?

Structure of the lecture:

1. MI and the output pathway (Corticospinal tract, CM cells)
2. MI somatotopy
3. What is the coding principle of MI
a. Ina-grained local somatotopy in MI is fractured
b. Many-to-many mapping
c. What is represented in primary motor cortex? (perhaps each
movement is represented once for each task or synergy? Perhaps
each muscle is presented several times so there is flexibility in how
it is controlled? Allow for learning: perhaps each muscle is
represented several times so there is plasticity for learning new
actions?
d. Marionette model vs Hebbian synergy model
4. Operant conditioning, plasticity, BMI

Introduction
 The primary motor cortex is located in the dorsal portion of the frontal
lobe and is the primary region of the motor system.
 The primary motor cortex is arranged somatotopically, meaning each part
of the body corresponds to a specific region of the motor cortex.
 Although it is known that the primary motor cortex is arranged in a
homunculus, the conclusions that can be drawn with respect to the
control of action is less clear.
 This essay discusses the theories relating the somatotopic organization of
the primary motor cortex to the control of action.

P1 – Early studies revealed the somatopic organization of the Primary Motor

Cortex
 Jackson (1870s) – partial seizures seemed to start in the hands and then
move systematically up the body towards the face – “Jacksonian march” –
caused Jackson to hypothesize that different areas in the cortex might be
responsible for movement in different parts of the body. Believed that
abnormal electrical activity in a partial seizure might e moving through
the cortex and at the same time causing abnormal movement to move
through the body in a corresponding fashion
 Fritsch & Hetzig (1870s) – electrical stimulation of motor cortex elicits
contractions of muscles on the contralateral side of the body by
stimulating motor cortex of live dogs. Also found that stimulation
produced movement in a predictable way – as if certain areas of the body
wre mapped onto the cortex. First widely recognized piece of evidence for
the motor cortex. Damaging areas of the cortex in dogs resulted in dogs
having difficulty with movement in the opposite side of the body.
 Work by Sherrington (apes) and later Penfield (humans). Penfield
(1930s) – found that human motor cortex contained a “map” of the body
such that stimulating an area of the motor cortex correlated with the
location of muscle contraction. Mapped out these correlations and found
 that areas associated with finer motor control occupied greater space in
the cortex. These were mirrored on the somatosensory cortex in the post-
central gyrus.
 Penfield mapped out homunculus = the most lateral areas control muscles
in the face, then the fingers and arm, and the legs are the most medial.

P2 – Larger areas are tuned to finer detail

 Elbert (1995) – increased cortical representation of the fingers of the left
hand in string players

P3 – However, this homunculus structure suggests one-to-one mapping

 However, one-to-one mapping seems inefficient
 Might work at the coarse level, but finer-level stimulation reveals that
muscles can be activated by applying electrical stimulation to many of the
areas.
 Furthermore – cortical microstimulation experiments have shown that a
single muscle is represented multiple times over a wide region of the
motor cortex (about 2-3 mm in primates) in a complex, mosaic fashion
 Microstimulation combined with recordings of muscle electrical activity
 even the smallest currents capable of eliciting a response initiated the
excitation of several muscles (and the simultaneous inhibition of others),
suggesting that organized movements rather than individual muscles are
represented in the map.
 Hlustik (2001) – overlapping regions in the movement of finger digits
o Interestingly, difference between 3-finger and 1-finger movements
more pronounced in S1 than M1
P4 – Instead, it is likely that we get many-to-many mapping
 But how does it work? Movements or muscle?
 In order to understand the implications, one must understand what the
neurons code for.
 Evarts 1968 – marionette model – the harder you pull, the more force.
Added weight to bar. Corticospinal tract neuron activity. Neuron codes
for muscle force. A resistive load requires more muscle contraction. No
coding of position.
P5 – However, it is not as simple as simply neurons coding for brute force
 Lemon (1993). A given M1 cell contributes to several muscles – one-to-
many mapping.
o These neurons are task specific – monkey trained to do precision
grip and power grip
 Movements, rather than muscles, are encoded by activity of the upper
motor neurons in the cortex
 Kakei (1999) – both movements AND muscles – trained monkey to use
three types of grips and different direction of movements. 28/88 were
tuned to muscles. 44/88 were tuned to differences in direction, regardless
of what muscles were used.
P6 – So, many-to-many mapping and movement coding.
 Hebbian synergy model – keep together the representations of muscles
you use together.
  Minimize wiring length because these neurons need to communicate with
each other

Conclusion

History of functional organization of the pimrary motor cortex

1. Fritsch and Hitzig showed that electrical stimulation of the motor cortex
elicits contractions of muscles on the contralateral side of the body
2. Hughlings Jackson – motor cortex contains a complete
representation/map of the body’s musculature (hypothesis) – evidence:
abnormal movements during some types of epileptic seizures “march”
systematically from one part of the body to another
a. Partial motor seizures may start with abnormal movements of a
finger, progress to involve the entire hand, then the forearm, the
arm, the shoulder, and final the face
3. Evidence for motor maps: Sherrington in 1930s: focal electrical
stimulation in great apes
4. 1930s – Penfield – extended Sherrington’s work by demonstrating that
human motor cortex also contains a spatial map of body’s musculature
a. Correlating the location of muscle contractions with the site of
electrical stimulation on the surface of the motor cortex 
Penfield mapped the representation of the muscles in the
precentral gyrus in over 400 neurosurgical patients. This motor
map shows the same disproportions observed in the somatic
sensory maps in the postcentral gyrus.
b. Thus, the musculature used in tasks requiring fine motor control
(such as movements of face and hands) occupies a greater amount
of space I the map than does the musculature requiring less
precise motor control
Behavioural implications:
What do motor maps represent?
 Electrical stimulation by Penfield and colleagues clearly demonstrated a
systematic map of body’s musculature in the primary motor cortex
 The fine structure of this map, however – continuing source of
controversy
 Is the map in the motor cortex a “piano keyboard” for the control of
individual muscles? Or is it a map of movements, in which specific sites
control multiple muscle groups that contribute to the generation of
particular actions?
 Initial experiments – implied the map in the motor cortex is a fine-scale
representation of individual muscles
 Thus, stimulation of small regions of the map activated single muscles,
suggesting that vertical columns of cells in M1 were responsible for
controlling the actions of particular muscles, much as columns in the
somatic sensory map are thought to analyse particular types of stimulus
information
 More studies using anatomical and physiological techniques, however, -
shown that the map in the motor cortex is far more complex than a
columnar representation of particular muscles.
 Individual pyramidal tract axons are now known to terminate on sets of
spinal motor neurons that innervate different muscles.
 This relationship is evident even for neurons in the hand representation
of the motor cortex, the region that controls the most discrete,
fractionated movements.
 Furthermore – cortical microstimulation experiments have shown that a
single muscle is represented multiple times over a wide region of the
motor cortex (about 2-3 mm in primates) in a complex, mosaic fashion
 Microstimulation combined with recordings of muscle electrical activity
 even the smallest currents capable of eliciting a response initiated the
excitation of several muscles (and the simultaneous inhibition of others),
suggesting that organized movements rather than individual muscles are
represented in the map.
 Within major subdivisions of the map (e.g. arm, forearm, finger regions), a
particular movement could be elicited by stimulation of widely separated
sites, indicating that nearby regions are linked by local circuits to
organize specific movements
 Regions responsible for initiating particular movements overall
substantially.
 Evarts () – implanted microelectrodes to record electrical activity in
individual motor neurons in awake, behaving monkeys. In these
experiments, monkeys were trained to perform a variety of motor tasks –
correlating neuronal activity with voluntary movements. He had
marionette view.
o Found that force generated by contracting muscles changed as a
function of the firing rate of upper motor neurons
o Firing rates of the active neurons often changed prior to
movements involving very small forces. Proposed that the M1
contributes to the initial phase of recruitment of lower motor
neurons involved in the generation of finely controlled
movements.