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Feedback control during voluntary motor actions
Stephen H Scott1,2,3, Tyler Cluff1, Catherine R Lowrey1 and
Tomohiko Takei1
Humans possess an impressive ability to generate goal-
oriented motor actions to move and interact with the
environment. The planning and initiation of these body
movements is supported by highly distributed cortical and
subcortical circuits. Recent studies, inspired by advanced
control theory, highlight similar sophistication when we make
online corrections to counter small disturbances of the limb or
altered visual feedback. Such goal-directed feedback is likely
generated by the same neural circuits associated with motor
planning and initiation. These common neural substrates afford
a highly responsive system to maintain goal-directed control
and rapidly select new motor actions as required to deftly move
and interact in a complex world.
Addresses
1
Centre for Neuroscience Studies, Queen’s University, Kingston, ON,
Canada
2
Department of Biomedical and Molecular Sciences, Queen’s
University, Kingston, ON, Canada
3
Department of Medicine, Queen’s University, Kingston, ON, Canada
Corresponding author: Scott, Stephen H (steve.scott@queensu.ca)
Current Opinion in Neurobiology 2015, 33:85–94
This review comes from a themed issue on Motor circuits and action
Edited by Ole Kiehn and Mark Churchland
For a complete overview see the Issue and the Editorial
Available online 28th March 2015
http://dx.doi.org/10.1016/j.conb.2015.03.006
0959-4388/# 2015 Published by Elsevier Ltd.
Introduction
Over the last two decades, most research on voluntary
motor control has focused on the planning and initiation
of motor actions, with little emphasis on afferent feed-
back from the moving limb. The ideas of optimal feed-
back control and other advanced control theories have
refocused attention on the importance of sensory feed-
back in biological control. The result is a renaissance in
exploring the complex ways in which somatosensory and
visual feedback can influence ongoing motor actions.
Here we review studies demonstrating that small correc-
tive responses during voluntary motor actions possess
most, if not all, of the complexities associated with motor
planning and movement initiation. Further, we discuss
how these sophisticated corrective responses reflect
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processing in a highly distributed network of cortical
and subcortical circuits.
There have been several recent reviews to discuss the
principles of feedback control and their application to
biological control [1,2], with several in-depth reviews
focused on the importance of somatosensory feedback
[3–7], visual feedback [8,9], or the integration of vision
and somatosensory feedback during voluntary actions
[10]. Related topics of interest are the importance of
sensory feedback in motor decisions [11] and the use
of dynamical systems theory to interpret the cortical
control of movement [12].
Task-dependent corrective responses during
voluntary motor actions
Somatosensory feedback: upper limb
A common paradigm to explore how somatosensory
feedback influences voluntary control is to observe
how subjects respond to small mechanical perturbations
[5–7]. Recent studies have shown that corrective
responses are faster when the urgency to respond is
increased by manipulating the return time, or size and
shape of the spatial goal [13,14]. Rapid corrective
responses occur even for very small disturbances that
approach the natural variability of limb motion [15],
illustrating that feedback processes are omnipresent in
voluntary motor actions. Sophisticated corrective
responses are also observed when a muscle is unloaded
by a mechanical perturbation [16].
A striking example of the sophistication of human correc-
tive responses is the ability to quickly respond to a me-
chanical perturbation and re-route a movement to avoid
obstacles in the environment [17] (Figure 1a). The selec-
tion to move to the right or left side of the obstacle
appeared to depend on the hand’s position at the time
the perturbation was applied. If the hand position was
deviated slightly towards the left/right, the subject was
more likely to go leftward/rightward around the obstacle.
The rapid selection of movements to avoid an obstacle
suggests that multiple potential strategies can be prepared
and launched as required during motor actions.
Detailed examination of electromyographic (EMG) ac-
tivity can provide an important clue as to the exact time
the motor system can generate task-dependent feedback
responses. The fastest of these responses occurs from
25 to 50 ms (short latency response, SLR). The SLR
engages spinal circuits, is relatively small and considers
Current Opinion in Neurobiology 2015, 33:85–94
86 Motor circuits and action

Figure 1

(a) Obstacle Avoidance (b) Lateral Triceps

R1 R2 R3 EV
3 * * * *

ΔEMG [a.u.]
2

1
Load
On 0
Between obstacles
2 cm 0 100 200
Around obstacles
Time [ms]

(c) One Target Multiple Targets (d) Lateral Triceps

R1 R2 R3 EV
4 * *

ΔEMG [a.u.]
3

1
Load Load
On On 0

Correct to initial target 0 100 200

2 cm Correct to new target
Time [ms]
Current Opinion in Neurobiology

Rapid online decisions following mechanical perturbations. (a) Subjects made reaching movements from a start position (bottom) to an end target
(top) while avoiding obstacles (filled circles) in their workspace. In some trials, a perturbation deviated their hand directly towards an obstacle.
Subjects either corrected back between the obstacles (blue) or deviated around the obstacles (red). (b) Perturbation-related muscle activity
(DEMG relative to unperturbed trials) in which subjects moved between (blue) or around (red) the obstacles. Data are aligned to perturbation onset
(solid vertical line). R1 (20–45 ms), R2 (45–75 ms), and R3 (75–105 ms) denote standard time epochs of the muscle stretch response, and EV
indicates the early voluntary component (105–135 ms). Note that differences in muscle activity are evident in the R2 epoch (45–75 ms after
perturbation onset). (c) Hand trajectories from a representative subject when there is one goal (left) or multiple potential goals (right). (d)
Perturbation-evoked response of lateral triceps (mean  SEM). Differences in muscle activity were not evident until the R3 epoch (75–105 ms after
perturbation onset). *P < 0.05.
Adapted from Nashed et al. [17].

how much the muscle is lengthened as well as back-
ground muscle activity (i.e., ‘gain scaling’ [18–20]). In
contrast, task-dependent responses are observed during
the long-latency time period (LLR; 50–105 ms post-per-
turbation [1,10]). In the Nashed et al. study [17], the
choice to redirect to the left or right around the obstacle
was reflected in EMG activity at 60 ms (Figure 1b; R2
epoch). Variations of this task demonstrated that when
subjects encountered a perturbation while reaching to
one of multiple goal targets, they sometimes switched to
reaching towards a secondary goal target (Figure 1c). The
selection of this alternate goal target occurred a little later
at 75 ms after perturbation onset (Figure 1d; R3 epoch).
This 15 ms shift in timing suggests different brain path-
ways or processes may be engaged in launching a response
to attain a target versus selecting a new goal target during
an ongoing action (how to attain the selected goal versus
what goal to select).
Indeed, recent evidence indicates that the neural pro-
cesses that generate LLRs are tightly coupled to ongoing
decisional processes [21]. In this study, subjects viewed a
dynamic random dot motion display and were asked to
indicate the direction of stimulus motion by moving to a
left or right target. Leftward and rightward force pulses
were randomly interleaved to probe the ongoing deci-
sional processes. LLRs scaled with the strength of the
stimulus (i.e., coherence) and duration of viewing, sug-
gesting that LLRs track an evolving decision variable
based on accumulated sensory evidence. Collectively,
these findings highlight a striking parallel linking the
computations involved in classic decision-making tasks
with online motor control [22].
Somatosensory feedback: lower limb
In standing subjects, perturbations to the support surface
evoke short, medium and long latency responses in lower
limb muscles. Short and medium-latency bursts in lower
limb EMG are thought to involve spinal and brainstem
pathways, whereas LLRs are linked to cortical processing
(see [23] for review). Recent work highlights that sensory
feedback drives the continuous modulation of leg muscle

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activity during dynamic balance control tasks [24,25].
During trains of platform accelerations applied in multi-
ple directions, LLRs better reflected the control of a
‘global’ task variable (center of body mass) than control
of ‘local’ variables such as individual joint motion patterns
[24].
Although corrective responses typically vary with proper-
ties of the task, there may be some functional differences
between upper and lower limb LLRs. For example, when
subjects controlled a virtual inverted pendulum with their
foot, a reduction in the stability of the virtual load was
associated with reductions in both the short [26] and
medium-latency response [27]. The authors argued that
feedback gains may be reduced in favour of a muscle co-
contraction strategy, which may increase joint stiffness
and the amplitude of the SLR to minimize displacement
caused by the perturbation [26,27,28]. In contrast, com-
pliant (i.e., unstable) loads applied to the upper limb are
associated with increased LLRs [29], suggesting different
control strategies may be engaged by certain upper and
lower limb motor tasks. However, it is difficult to inter-
pret these discrepancies in feedback modulation because
subjects were instructed not to respond to the perturba-
tion in both the upper and lower limb tasks (i.e., ‘do not
intervene’). Further work may be required to delineate
the task-relevancy of this feedback modulation.
Somatosensory feedback: interlimb
Many behavioural tasks require coordination between
limbs. For example, when the two hands control a single
cursor, which is presented at the spatial midpoint of the
hands, perturbations to one arm lead to fast, feedback-
driven responses in most muscles in the opposite arm
within 75–100 ms [30,31]. Remarkably, participants can
independently learn and switch between control policies
acquired for different force fields during unimanual and
bimanual movements [32,33], although reoptimization
may be incomplete when switching between two control
policies for unimanual arm movements [34]. Similarly,
perturbations applied to one leg during locomotion can
evoke rapid responses in muscles of the contralateral leg
in less than 75 ms [35]. Critically, these inter-limb
responses only occur when the contralateral limb is
weight bearing (i.e., stance phase), and a stabilizing
response is necessary to maintain standing balance.
Feedback responses are also highly coordinated between
the upper and lower limbs. Perturbations to standing
balance can generate muscle activity in the upper limbs
within 90 ms, which is faster than voluntary reaction
times [36]. These rapid arm movements appear to be
influenced by the direction of the perturbation even on
the first exposure [37]. Similarly, perturbations applied to
a handle held by subjects during treadmill walking evoke
muscle responses in the upper and lower limbs that are
modulated by verbal instructions (resist or do not resist
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Feedback control during voluntary motor actions Scott et al. 87
perturbation) and phase of the gait cycle [38]. However, it
is difficult to determine the latency of these responses
as EMG activity was averaged for 200 ms following
the earliest onset of perturbation-evoked muscle activity,
which likely incorporates voluntary contributions.
Further work is necessary to establish how quickly
task-dependency from upper limb tasks is incorporated
into corrective responses in the lower limbs.
Visual feedback
The importance of visual feedback in voluntary control
has received a lot of attention in motor neuroscience.
Visuomotor corrections have been addressed with the
‘double-step’ paradigm, in which the target unexpectedly
changes location before or during the course of movement
(‘target jump’ [39–42]), or by displacing the hand-aligned
cursor (‘cursor jump’ [43]) during reaching (see [8,9]).
Recently, Dimitriou and colleagues [44] used visual
shifts of the target or hand-cursor to test whether the
motor system is able to update the control policy during
reaching. By perturbing hand-cursor feedback either si-
multaneously with or 100 ms after the target jump, the
authors showed that feedback responses took 100 ms to
consider the new target position (see also [45]). Visual
feedback of target information is influenced by visual
attention, such that environmental distractors reduce the
amplitude and increase the latency of corrective
responses following a sudden shift in target position. In
contrast, there appears to be a more dedicated link for
visual feedback of the limb as responses to cursor jump
perturbations are less affected by environmental distrac-
tors [46]. This dedicated binding mechanism for visual
feedback of the moving hand may be generated by the
presence of faster somatosensory feedback used to esti-
mate limb motion.
Feedback responses and learning
A key feature of the voluntary motor system is the ability
to learn new motor skills. Many studies have demonstrat-
ed that the voluntary motor system updates its motor
commands when exposed to novel environments that
alter the physical properties of our limbs (e.g., using
sports equipment, walking on sand or in water [47]). As
described above, numerous studies have highlighted
flexible, goal-directed feedback responses across a broad
range of voluntary motor actions. Thus, it seems reason-
able that there is a link between motor learning and
corrective responses.
We recently tested this hypothesis by examining how
corrective responses were altered when subjects learned
to make reaching movements in the presence of resistive
elbow loads (velocity dependent [48]). In random trials,
mechanical perturbations to the elbow were used to probe
LLRs during the learning phase. We found that LLRs
adapted during learning and scaled with the strength of
Current Opinion in Neurobiology 2015, 33:85–94
88 Motor circuits and action
the novel resistive loads applied at the elbow. These
altered LLRs correlated with reaching errors during
learning, showing that subjects who adapted more to
the novel load (i.e., had smaller reach errors) displayed
greater modulation of their LLRs in perturbation trials.
These results highlight an important link between the
adaptation of feedforward and feedback control, and
suggest a key part of motor adaptation is to adjust feed-
back responses when the physical properties of the limb
or environment are altered. In addition, LLRs compen-
sate for novel loads when a perturbation is applied before
movement, suggesting the adapted control policy is en-
gaged while preparing a voluntary action [49]. Finally,
both voluntary actions and corrective responses are
adapted when the strength of the force field is consistent
across trials. In contrast, exposure to variable force fields
appears to only modify corrective responses [50], indicat-
ing the adaptation of voluntary actions and feedback
responses may be partly dissociated in some circum-
stances.
A key challenge with feedback control is dealing with
sensory and motor delays. Recent work illustrates that
forward models might help remove this delay [51].
Further, this study demonstrated that state estimation
processes were altered on a trial-by-trial basis. These
rapid trial-by-trial updates in state estimation are broadly
consistent with two-timescale models of motor learning
[52] and the recent suggestion that fast learning is associ-
ated with forward models (i.e., state estimation) whereas
slow learning is related to inverse models (i.e., feedback
gains [53]).
Corrective responses to visual cursor shifts also update
during force and kinematic adaptation [54–56]. Visuomo-
tor corrective responses are modulated by the direction
and structure of visual rotations [56], and selectively
account for the relevance of cursor-jump perturbations
[54]. Remarkably, corrective responses can be learned
independently based on the relevance of visual perturba-
tions towards the left or right side of the workspace. For
example, subjects can learn to make vigorous corrective
responses when visual perturbations persist until the end
of movement on the left side of their workspace, but learn
to disregard transient perturbations on the right side of
their workspace [54]. These flexible visuomotor
responses highlight the specificity of how the motor
system evaluates and responds to sensory feedback dur-
ing ongoing actions. Collectively, the adaptation of cor-
rective responses highlighted in recent studies establishes
they play an important role in the basic mechanisms of
generating skilled actions.
Neural implementation of feedback control
The voluntary motor system is hierarchically organized,
and correspondingly, corrective responses reflect contri-
butions from many levels of this hierarchy (Figure 2). It
Current Opinion in Neurobiology 2015, 33:85–94
Figure 2
Neural control
SMA
PMd
M1 S1 5 7
PMv
BG V1
C
Motor behaviour
Spinal cord
Musculoskeletal mechanics
Current Opinion in Neurobiology
Neural basis of feedback processing during voluntary motor actions.
Spinal feedback provides a fast default response for somatosensory
feedback. However, goal-directed corrective responses probably
involve cortical and subcortical circuits already ascribed to feed-
forward planning and initiating motor actions (M1, primary motor
cortex; S1, primary somatosensory cortex; PMd, dorsal premotor
cortex; PMv, ventral premotor cortex, SMA, supplementary motor
area; 5, parietal area 5; 7, parietal area 7; V1, primary visual cortex;
BG, basal ganglia; C, cerebellum).
Adapted from [99].
has been assumed that co-contraction of antagonist mus-
cles plays an important role in minimizing the effect of
disturbances. However, estimates for muscle stiffness
commonly used in the field are almost an order of magni-
tude too high, as they include contributions from neural
feedback processes [28]. The main benefit of co-con-
traction may be to increase the amplitude of spinal stretch
responses (i.e., short latency) that scale with the level of
muscle activity, although the link between co-contraction
and spinal feedback gains may be altered in some situa-
tions [26].
Spinal feedback
There is considerable debate regarding the relative con-
tribution of spinal versus supraspinal feedback during
voluntary motor tasks, with some concluding spinal feed-
back plays a dominant role [57] and others a minimal role
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[6]. Neural networks inspired by spinal cord circuitry can
generate reaching movements or respond to perturbations
assuming simple descending motor signals [57]. Howev-
er, neural networks without spinal circuitry can also
generate good control [58]. In all cases, these models
inevitably ignore a major problem in biological control —
sensory and motor delays. Such delays are devastating for
controlling fast movements or corrective responses [28],
but can be mitigated by using state estimation to predict
the sensory consequences of our actions [51]. The com-
plexity of state estimation during voluntary actions may
explain why goal-directed muscle responses are only
observed at 60 ms when transcortical feedback can
contribute [1,6,10]. Further experimental work, notably
at the spinal level, is essential to resolve this issue, as only
a limited number of studies have recorded spinal neural
activity in non-human primates during single-joint motor
tasks (for review see [59]) or prehension tasks [60,61].
The development of novel molecular and genetic tech-
niques provides an exciting new approach to understand
the function of spinal circuits during voluntary actions
[62–64]. Fink et al. [63] selectively ablated GABAergic
interneurons in mice cervical spinal cord that mediate
presynaptic inhibition of proprioceptive afferents. The
mice could still walk and maintain posture after selective
removal of these interneurons, but displayed pronounced
forelimb oscillations during reaching (Figure 3). These
forelimb oscillations suggest that feedback gains were too
high during movement, which could involve many factors
from changes in muscle stiffness to an imbalance between
spinal and supraspinal feedback processes [28,65].
Figure 3
(a) (c)
Cutaneous
Reach trajectories
Pre-DT
Proprioceptor 10
Post-DT
vGluT1
z (mm)
GAD2
0 Pellet
GAD1
Motor neuron
(b)
Camera –10
0
20 10 0 5 y (mm)
Camera x (mm)
Current Opinion in Neurobiology
Behavioural effect of ablation of presynaptic inhibition on primary
afferents. (a) Gad2-expressing interneurons, which form axo–axonic
contacts with primary afferent terminals, were the target of the genetic
manipulation and selectively ablated after the administration of
diphtheria toxin (DT) in mice spinal cord. (b) Mice reached for and
retrieved a food pellet through a narrow access window. (c) Before DT
administration mice displayed smooth paw trajectories (pre-DT). In
contrast, after DT administration the reaches lost smoothness and
exhibited pronounced oscillation (post-DT).Adapted from Fink
et al. [63].
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Feedback control during voluntary motor actions Scott et al. 89
Genetic tools have provided new insight that spinal
interneurons send efference copies of descending motor
commands to the cerebellum [66,67]. Azim et al. [67]
recently illustrated that ablation or modulation of the
activity of V2a neurons that project back to the cerebel-
lum disturbed reaching behaviour in mice. Taken togeth-
er, the use of genetic and molecular techniques is helping
to shed new light on how the spinal cord contributes to
voluntary motor actions.
Transcortical feedback
It is generally accepted that transcortical feedback plays
an important role in LLRs (for review see [1,3,5–7,10]).
The use of transcranial stimulation techniques has dem-
onstrated the involvement of M1 in modulating LLRs
related to knowledge of limb mechanics [68], and phase-
dependent responses between the legs during locomotion
[35]. However, inhibition of motor cortex did not alter
task-dependent LLRs when subjects interacted with stiff
versus compliant loads [69], which opposes previous
observations by this group [70]. It may be worthwhile
to re-examine this issue for tasks where subjects attain a
quantifiable goal following the perturbation rather than
the commonly used ‘do not intervene’ instruction.
Interest in exploring sensory feedback in M1 has
returned. Studies have shown that neural activity is
broadly tuned to mechanical perturbations applied to
the shoulder and elbow [71], and reflects knowledge of
multi-joint limb dynamics [68]. The prevalence of so-
matosensory feedback in M1 is also highlighted by its
ability to enhance the performance of Brain Machine
Interfaces generated from M1 activity [72].
The role of M1 in task-dependent feedback processing
was first demonstrated in the seminal experiments by
Evarts and Tanji [73], in which monkeys were instructed
to either ‘push’ or ‘pull’ a lever triggered by a mechanical
perturbation (Figure 4a [73]). An updated version of this
task was recently performed where monkeys responded
to a mechanical perturbation by moving their hand to a
spatial goal [74]. The perturbation either pushed the
monkey’s hand towards or away from the target, and a
background load was applied before the perturbation to
prevent subthreshold modulation of motoneuron pools.
As in Evarts and Tanji [73], the initial perturbation
response occurred as early as 20 ms and was largely
insensitive to the location of the spatial goal. This early,
non-specific response was followed by a task-dependent
response beginning at 50 ms after perturbation onset
(Figure 4b).
The presence of an early non-specific response might
indicate that it takes time for M1 to generate task-
dependent responses. However, it may also reflect ongo-
ing postural control before the perturbation was applied,
and that it takes time to disengage postural control
Current Opinion in Neurobiology 2015, 33:85–94
90 Motor circuits and action

Figure 4

(a) Evarts and Tanji (1976) (b) Pruszynski et al. (2014) (c) Omrani et al. (2014)

ins-push/ pert-pull
E L
Posture
60 OUT

Firing Rate (Hz)

40 IN
Movie
ins-pull/ pert-pull
20 Difference
Difference

–500 0 500 –40 0 40 80 120 –40 0 40 80 120

Time (ms) Time (ms) Time (ms)
Current Opinion in Neurobiology

Task-dependent modulation of perturbation responses in M1. (a) Perturbation response of a single pyramidal tract neuron (PTN) for lever ‘pull’
perturbation under different instructions. When the monkey was instructed to ‘push’ the lever (instruction-push/perturbation-pull condition, top),
the PTN showed clear sustained activity until monkey attained the target (thick dot in each row). When the monkey was instructed to ‘pull’ the
lever responding to the same ‘pull’ perturbation (instruction-pull/perturbation-pull condition, bottom), the PTN showed suppression of activity
following the initial excitation. (b) M1 population activity in a spatial target task (n = 104). Monkeys were trained to move to spatial targets
following a mechanical perturbation. The disturbance either moved the hand into (IN, blue) or away from the target (OUT, red). Although M1
neurons responded to the perturbation as early as 20 ms (E, early epoch), target-dependent responses (OUT–IN, gray) began at 50 ms (L, late
epoch). (c) M1 population response in posture and movie tasks (n = 129). M1 activities were recorded when the monkey responded to a
mechanical perturbation to maintain its arm position (posture task, red) or when it was watching a movie and not required to respond to the
perturbation (movie task, blue). Black arrow denotes population response to perturbation and gray arrow denotes task-dependent response. All
data are aligned to perturbation onset (solid vertical line).
Panels (a), (b) and (c) are adapted from [73], [74] and [75], respectively.

before generating a response to the spatial goal. Omrani
et al. [75] tackled this question by comparing M1 per-
turbation responses when the monkey was engaged
(positioning its hand at a spatial target) or not engaged
in postural control (watching a movie, Omrani et al. [75]).
The results again indicated the initial M1 response was
identical across behavioural contexts with task-depen-
dent responses beginning only 40 ms post-perturbation
(Figure 4c). A similar response pattern was observed in
M1 when a monkey was required to convert patterns of
limb motion into the appropriate motor responses to deal
with limb dynamics [68]. This delay may reflect the need
to wait for multiple sources of sensory feedback before
M1 expresses the appropriate motor response.
Transcortical feedback beyond primary motor cortex
A key somatosensory pathway to M1 is through primary
somatosensory cortex (S1) where neurons respond to
either active or passive arm movements [76]. Move-
ment-related activity in S1 can occur before movement
onset, suggesting that some activity in S1 reflects an
efference copy of descending motor commands and
may arise from connections with M1 [77,78]. Transmis-
sion of sensory information to S1 is reduced during but
not prior to movement, as demonstrated by reductions in
sensory evoked potentials (SEPs) [79].
There is growing evidence that transcortical feedback
involves cortical motor regions beyond primary motor
cortex. M1 along with dorsal premotor cortex (PMd) show
a reduction in SEPs during movement [79]. Moreover,
SEPs in M1 and PMd are also reduced during the delay
period prior to movement onset. Inhibition of supple-
mentary motor area (SMA) using transcranial magnetic
stimulation (TMS) diminishes the typical modulation of
the LLR observed for a ‘resist’/do not resist’ paradigm.
Reduction of LLR modulation following SMA inhibition
suggests SMA may have a direct influence on the LLR
and/or that SMA ‘presets’ M1 activity according to task
instruction prior to perturbation onset [80]. Finally, a
recent study found that task-dependent LLRs were
absent in chronic stroke patients with unilateral cortical
lesions [81]. Importantly, although strokes were unilat-
eral, deficits in goal-directed LLRs were observed bilat-
erally in both the affected and unaffected arms. Bilateral
impairments in LLRs suggest ipsilateral cortex plays a
direct or indirect role in task-dependent motor correc-
tions. However, a follow-up study in healthy subjects
found that inhibition of ipsilateral M1 with a strong TMS
pulse did not alter the amplitude of LLRs [69]. Further
work is required to understand the role of ipsilateral
cortex in the generation of task-dependent LLRs in
healthy and patient groups.

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The involvement of cerebellum in LLRs was established
in the seminal studies by Strick [82] and Villis and Hore
[83]. Recent work confirms that cerebellar dysfunction is
associated with altered scaling of the LLR with either
reduced [84] or exaggerated [85] amplitude. Although
cerebellar damage impairs active compensation for inter-
action torques during multi-joint arm movements [86],
cerebellar patients appear to maintain some knowledge of
limb mechanics in the LLR [84]. Difficulty in dealing
with limb mechanics may be linked to deficits in estimat-
ing active limb motion [87,88] as knowledge of limb
dynamics seems to be compromised by cerebellar damage
[89].
The basal ganglia may also modulate the transcortical
pathway, as a hallmark of stretch responses in patients
with Parkinson’s disease (PD) is an exaggerated LLR to a
mechanical perturbation [90]. There is also evidence that
PD patients lose the ability to scale responses to features
of the task [91]. However, recent work with a non-human
primate model of PD found no overt change in M1
activity associated with the observed increase in reflex
gain [92], which implicates subcortical circuits in goal-
directed feedback responses.
Neural basis of visual feedback
Visual feedback during voluntary motor actions is as-
sumed to follow the dorsal visual stream towards PMd
and M1 [9]. PMd appears to play an important role in
online visual feedback as TMS inhibition in this area
disrupts online corrections and reduces the ability to learn
visuomotor rotations [93]. Sudden changes in target loca-
tion at movement onset elicit corrective activity in PMd
and M1 [94]. The earliest change in activity following a
shift in target location occurs in PMd, followed by M1 and
then only later in parietal area 5 [95]. Although parietal
area 5 responds after motor cortical regions during online
adjustments, this cortical region appears to be important
for online control as bilateral muscimol injections delays
corrective responses following a change in target position
[96].
Recent evidence from patients with damage to cortical
areas has delineated hemisphere-specific contributions to
the use of this visual feedback. For example, damage to
the right frontal lobe delays the onset of corrective
responses [97,98], whereas left hemisphere damage
interferes with inter-limb coordination during rapid on-
line corrections [97].
Conclusions
A growing body of literature highlights the surprising
sophistication of rapid corrective responses generated
by visual and somatosensory feedback. Long-latency
stretch responses are highly context-dependent reflecting
the behavioural goal and many other factors essential for
moving and interacting in the world. These corrective
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Feedback control during voluntary motor actions Scott et al. 91
responses share most, if not all, of the complexities
typically ascribed to voluntary control. Considerable work
remains to understand how these feedback processes are
generated by the brain and spinal cord. It is likely that the
role of different brain regions is similar for planning,
initiating and correcting motor actions, blurring the dis-
tinction between feed-forward and feedback control.
Conflict of interests
SHS is associated with BKIN Technologies, which com-
mercializes the KINARM robot used in some studies
described in this review. TC, CRL and TT do not have
any conflicts of interest.
Acknowledgements
This work was supported by grants from the Natural Sciences and
Engineering Research Council of Canada (NSERC) and the Canadian
Institutes of Health Research (CIHR). SHS is supported by a GSK-CIHR
Chair in Neuroscience. TC is supported by a Banting Postdoctoral
Fellowship.
References and recommended reading
Papers of particular interest, published within the period of review,
have been highlighted as:
 of special interest
 of outstanding interest
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