Chapter 25

Motor learning
Şermin Tükel
˙
Department of Physiotherapy and Rehabilitation, Izmir ˙
University of Economics, Izmir, Turkey

Introduction other sensory-perceptual and cognitive learning types

often interact to achieve a skilled performance.
Learning is a process that refers to changes in behavior
and in the central nervous system. Motor learning is
defined as: ‘a change in the capability of a person to per- Learning hypotheses
form a skill that must be inferred from a relatively perma-
Schmidt’s schema theory of discrete motor skill learning
nent improvement in performance as a result of practice
was originally presented in 1974 at a meeting of the
or experience’ (Magill and Anderson, 2007). Motor learn-
North American Society for Psychology of Sport and
ing covers three main types of learning procedure; 1)
Physical Activity (Schmidt, 1975). According to the
acquisition of a novel motor skill (novel task learning), 2)
model, there is a generalized motor program, a schema,
enhancement of performance of a learned or highly expe-
for a group of motor tasks. During learning of a motor
rienced motor skill (expertise), and 3) reacquisition of
skill, there are four important parameters to store in work-
skills that are difficult to perform or cannot be performed
ing memory. These are; 1) the conditions at the beginning
because of injury, disorder or disease (rehabilitation). In
of the movement, i.e. proprioceptive information about
order to understand motor learning, it is important to clar-
the posture, weight of a tool or object that is used, 2) the
ify the interrelated terms of motor skill and motor perfor-
specifications for the motor program such as, speed and
mance. Motor skill refers to a goal-directed activity or
force, 3) the knowledge of results, or actual outcome, and
task that requires voluntary control over movements of
4) the sensory consequences of the movement response,
the joints and body segments, whereas motor performance
how the movement felt, looked and sounded. These para-
refers to the execution of a skill at a specific time and in
meters compose the motor response schema. Important
a specific situation. Motor performance is an observable
point is that the schema is modified in every execution of
behavior, whereas motor learning cannot be observed
the movement based on the sensory feedback and knowl-
directly, but rather, can be inferred from performance.
edge of results. Schema model involves learning a recog-
Fig. 25.1A and B show the performance of rock climbing
nition schema that is an error detection mechanism, and
skill by professional athletes. Learning of this sport skill
learning a recall schema that is the selection of accurate
starts with observing and imitating an experienced
motor program.
climber. The novice climber closely monitors an experi-
Ecological theory of motor learning has been proposed
enced climber while experienced climber is ascending. In
recently by Newell (Newell, 1991). This model relies on
doing so, the novice might plan some explicit strategies
perception-action coupling, and defines that optimal
such as, the rock edges for gripping with hands and feet.
movement strategies are developed in the most efficient
When attempting climbing, the novice must rely on sen-
way given certain environmental constraints. Perception
sory feedback from proprioceptive and tactile, vestibular,
is important because the person needs to understand the
and visual organs for maintaining balance and movement
goal of the task, learn the movement and interpret the
control. In addition, trainers usually give explicit guid-
feedback, both during and after the movement. In this
ance, such as the direction of the next handling, or where
approach, the model highlights the structural and func-
to step. Despite all the explicit knowledge gathered by the
tional constraints related to the individual, task, and envi-
novice, the practice relies on implicit musculoskeletal and
ronment, and views motor skill as a dynamic exploratory
neural processes. As in this example, motor learning and
activity in perceptual-motor workspace.

Comparative Kinesiology of the Human Body. DOI: [Link]

© 2020 Elsevier Inc. All rights reserved. 453
454 PART | 7 Sensory-motor integration and performance

FIGURE 25.1 Rock climbing is an exciting and challenging sport. During learning of rock climbing, beginners practice handling and stepping tech-
niques, which exert pushing, pulling and compression forces to transfer body’s center of gravity. Beginners start with low grade top rope climbing or
bouldering routes that are easy therefore, more convenient for practising. In photos of rock climbing, performance of two Turkish National sport
climbing athletes Duru Güneş Yalçın in ’Kısa Samsun’ route (Grade 6c 1 ) (A) and Zorbey Aktuyun during a lead climbing in ‘Nessuono’ route
(Grade 8a 1 , 27 hours climbing) (B) are shown. Nessuono is a much more difficult route because of the structure of the rock that can be handled only
by pinch gripping beyond endurance for 27 hours of climbing (B). Performance at the expert level has an endurance of finer movements and coordi-
native structure in handling and stepping techniques that are acquired as a result of extensive practice.

FIGURE 25.2 Classification of long-term memory and different systems of learning are presented. There is an interaction between learning and
memory processes. Systems are not fully independent from each other, rather they depend on each other to some extent.

Behavioral approach that cannot be described verbally, unlike explicit learning,

defined as the conscious process of learning facts, infor-
Learning and memory are both theoretical constructs,
mation, and events (Byrne, 1997). Regarding the wider
which are interconnected and inferred from behavior (per-
theoretical framework of human learning, ‘motor learning’
formance). Learning is defined as ‘an enduring change in
is largely considered as an implicit learning process, and
the neural mechanisms of behavior that result from experi-
is used interchangeably with the term ‘procedural learn-
ence with environmental events’ (Domjan and Burkhard,
ing’. Fig. 25.2 shows different types of learning and mem-
1982). We can divide learning processes into two major
ory, and the place of ‘motor learning’ in this framework.
types; implicit and explicit. Implicit learning defines an
Below is the description of different learning types.
unconscious, yet extremely important process of learning
 Motor learning Chapter | 25 455

Perceptual learning consequences, such as pain or failure (negative reinforce-

This refers learning perceptual skills, such as differentiat- ment), it will decrease the likelihood of its occurrence. In
ing two colors, or the identification of different food rehabilitation science, operant conditioning of spinal
tastes. Examples can be expanded for other sensory reflex has been investigated as a promising tool for loco-
modalities; auditory, tactile, olfactory, and proprioception. motion. Simply stated, stimulus-induced muscle responses
Perceptual learning can be viewed as an essential part of (reflexes) are used to induce neuroplasticity. In a study by
complex cognitive functions. For example, being able to Wolpaw’s group, patients with incomplete spinal cord
discriminate close sounds is a clear advantage in language injury decreased H-reflex of soleus muscle with operant
learning (Ward, 2015). Similarly for motor learning, down-conditioning, which was associated with faster and
extracting and discriminating relevant sensory information more symmetrical locomotion (Thompson et al., 2013).
is essential (Wolpert and Flanagan, 2010). Experienced Similarly, operant up-conditioning has been led to
soccer players showed advanced visual discrimination of increase in motor-evoked potential of tibialis anterior in
an opponent’s postural orientation, which enables them to incomplete spinal cord injury (Thompson et al., 2018).
anticipate their next movement (Williams, 2009), and to
find relevant sensory information more quickly (van Nonassociative learning
Maarseveen et al., 2015). Nonassociative learning involves learning the properties
of a single stimulus. Simple forms of non-associative
Associative learning learning consists of habituation and sensitization.
Associative learning modifies the behavior via relating Habituation refers to diminishing response amplitude,
one stimulus with another, or relating a stimulus with a while sensitization refers to increasing response amplitude
particular behavior. In classical conditioning, a person to the repeated stimulus (Kandel et al., 2000). Habituation
pairs two stimuli, and therefore reflex response is modi- is important for minimizing the energy expenditure of the
fied. In operant conditioning, a person pairs his/her own motor system. Studies have shown that large automatic
behavior with the consequences of that behavior (Kandel responses to unexpected movements of a supporting plat-
et al., 2000). Classical conditioning is a simple form of form are progressively decreased due to a generalized
associative learning, where the behavioral response is habituation in the postural control system (Keshner et al.,
modified by conditioned stimulus. In the classical exam- 1987). Similar postural control responses occur very fre-
ple, developed by Ivan Pavlov, dogs produce reflex quently in everyday life, for example, while standing in a
response of salivation when conditioned with a sound moving bus, sudden braking results in larger postural
stimulus. In the experiment, dogs associated the sound reactions at first, but subsequent similar braking results in
stimulus with the food (natural stimulus leading to saliva- less dramatic postural reactions. In addition, habituation
tion response) after sufficient conditioning. Then, the dog may be an important factor to consider in orthopedic inju-
shows salivation response to the sound in the absence of ries. Stretch reflex response of the lateral ligament and
food. Classical conditioning is usually related to 1) emo- peroneal muscles are considered to have a role in preven-
tional responses and 2) skeletal muscle responses. Eye tion of ankle sprains. It has been shown that after ten con-
blink conditioning is a form of classical conditioning that secutive trials of ankle inversion (as if there is a lateral
has been studied in the investigation of neural mechan- ankle spraining force), stretch reflex response amplitude
isms underlying learning and memory. A mild air puff is decreased 20 50%, suggesting increased risk for injuries
a natural stimulus that results in an eye blink reflex. In (Jackson et al., 2009).
the experiment, the air puff is paired with visual or audi-
tory stimuli, so that, for example, eye blink reflex is seen
Cognitive approaches
when a person hears a sound. The eye blink conditioning
experiments showed the important role of cerebellum in A major part of human learning consists of interplay
associative learning; especially in acquisition and timing between cognitive functions, such as thoughts, memories
of motor actions (Gerwig et al., 2007). and expectations. Here, we consider observational learn-
Operant conditioning (also called trial-and-error learn- ing and learning through imitation. The concept of obser-
ing) is another type of associative learning in which a vol- vational learning was introduced by Albert Bandura. He
untary motor behavior is strengthened or weakened, demonstrated that children imitated the violent behaviors
depending on its favorable or unfavorable consequences. of adult models towards a toy (Bandura et al., 1963). This
When motor behavior is associated with desirable conse- type of learning is very important for learning clinical
quences such as a reward (positive reinforcement), there procedural skills, such as injection, intubation etc.
is a tendency to repeat the behavior. In the opposite situa- Imitation of a model is strengthened if the model is
tion, when the behavior results in unwanted rewarded for that particular behavior. In humans,
456 PART | 7 Sensory-motor integration and performance

observational learning is associated with mirror neurons, a limited capacity. It describes the work-bench of the
a group of neurons in the supplementary motor area and mind, where we hold the significant information active
in the medial temporal lobe, which show response during for ongoing mental process of comprehension, solving
both execution and observation of actions (Rizzolatti problems, and decision-making. Therefore, working mem-
et al., 1996; Mukamel et al., 2010). Mirror neurons have ory is an important capacity during motor skill acquisi-
been found to show response to specific type of goal- tion. Indeed, a stronger visuospatial working memory is
directed movements in upper extremity functions, such as associated with faster learning of manual skills in the
grasping and facial emotional gestures. Therefore, it has early phase of practice (Anguera et al., 2010, 2011;
been suggested that mirror neurons are important for Ruitenberg, De Dios et al., 2018). This association has
interacting with objects, and also for social cognition par- been related to individual differences in explicit strategies
ticularly for understanding the actions of others (Heyes, used to achieve the manual task (Christou et al., 2016).
2010). Long-term memory stores information on a relatively
permanent basis, and is considered to have unlimited
capacity within the inherent compounds of the brain. The
Memory declarative long-term memories can be consciously
Memory is created through learning. It is defined as ‘a accessed and declared and has two distinct types; seman-
theoretical term used to determine instances in which sub- tic memory and episodic memory (Fig. 25.2). An example
jects’ current behavior is determined by some aspects of for semantic memory is remembering the names of all
his previous experience’ (Domjan and Burkhard, 1982). objects associated with the color red. This gives us a
Remembering threatening situations or locations where semantic word tree, which is important in language learn-
food has previously been found has an evolutionary ing. The second clearly defined type is episodic memory.
importance. It is also meaningful for the energy system; if This covers episodes, events in a person’s life and con-
a motor behavior is repeated very often, it should be ceptual issues such as subjective time and consciousness
stored so when needed again, the neural and musculoskel- (Tulving, 2002). A type which cannot be declared is pro-
etal systems consume less energy. Adaptation to repeated cedural memory, referring to motor memory for skills like
actions results in change in nervous and musculoskeletal cycling. This is a type of long-term memory; e.g. we
systems. This ability to change is called plasticity. never forget how to ride a bike, swim or play tennis.
Neuroplasticity describes the change in synaptic proper- Four different stages in memory formation can be
ties of neurons in response to environmental stimulation, defined: encoding and storing newly acquired information
and in recovery from brain and spinal cord injuries (encoding), transferring encoded information from an
(Merzenich, 2013). Neuroplasticity is greatest during unstable state to a more stable state (consolidation),
childhood, when synaptic connections bloom and are retrieving the information when needed (retrieval), and
pruned, but it has been known to persist through the life updating the information and re-storing it (reconsolida-
span, although to a lesser extent. A similar tendency of tion). An important stage is consolidation, which stabi-
plasticity exists for skeletal muscles, which have great lizes information in our memory. Motor memory
adaptive and regenerative capacity in response to environ- significantly differs from declarative memory in consoli-
mental mechanical stimuli, exercise and injury throughout dation phase. In every trial of the task motor memory has
the life span, but particularly during early periods of ‘savings’, a term used to describe a more rapid rate of
development (Mersmann et al., 2014). relearning compared with the original learning (Krakauer
Memory may have several components, including and Shadmehr, 2006). If you have previously practiced a
visual, phonological or olfactorial. For example, memo- novel motor task, the next time you memorize the move-
ries of your mother may be triggered by hearing her ment pattern, meaning that you relearn it faster because
name, smelling her perfume, seeing her photo, and hear- of ‘saving’ in the motor memory. Savings of motor learn-
ing her voice. The memory can be divided into two cate- ing have been observed after one day (Bédard and Sanes,
gories; short-term and long-term memory (Fig. 25.2). 2011; Villalta et al., 2013; Seidler et al., 2016), one
Long-term memory may be further subdivided into differ- month (Della-Maggiore and McIntosh, 2005), five months
ent components in relation to how learning is acquired. (Shadmehr and Brashers-Krug, 1997), and even as much
This model is termed the multiple memory systems as one year after initial learning (Landi et al., 2011).
approach (Nyberg, 1996). Savings can be better understood with online and off-
Short-term memory holds information for 15 25 s, line processes during motor learning and formation of
and has a limited capacity (Feldman and Garrison, 1993). motor memory. Online process describes the gain or loss in
For example, you can hold a phone number in your short- motor performance during a practice session, and offline
term memory while dialing, but once dialed, it is forgot- process refers to performance changes occuring between
ten. Working memory is a short-term memory buffer with subsequent practice sessions (Dayan and Cohen, 2011).
 Motor learning Chapter | 25 457

Offline learning is related to neuroplasticity (Muellbacher initiate new programs for movement and introduce
et al., 2002), thereby the type of activity after the practice changes in programs that are in progress. In later stages
session might interact with the offline learning process. when the attention demand decreases, there is lower level
While sleep enhances offline learning (positive transfer) of prefrontal activation (Poldrack et al., 2005).
(Walker et al., 2003), performing interfering motor task The serial reaction time (SRT) task is a widely used
has been shown to abolish motor memory consolidation motor paradigm used to study neural correlates of motor
(negative transfer) (Brashers-Krug et al., 1996). learning. In the SRT task, a stimulus appears on a com-
puter screen, and the participant is instructed to press the
specific key assigned to that stimulus as quick as possible.
Physiological approach
The order of stimuli follows a predictable order, but the
Motor learning or training induces structural and func- participant does not initially know this. During the task,
tional changes at neuronal level. These changes include reaction time decreases as a result of motor learning. If
increase in the number of neurons, reorganization or the order of stimulus is changed, then the reaction time
expansion of cortical motor and sensory maps, modulation increases. In SRT, some individuals can verbally express
of neuron’s firing rate and synaptic transmission. These the order of the stimulus, but even for those who cannot
findings were shown in animal and human studies. reaction time still decreases (Willingham et al., 2002).
Learning new acrobatic movements, rats showed Neuroimaging studies make use of this paradigm to dif-
increased number of synapses per Purkinje cell in cerebel- ferentiate neural networks involved in implicit and
lum (Black et al., 1990). Also in motor cortex, rats trained explicit learning. The neural distinction between implicit
on a skilled reaching task showed enlarged cortical repre- and explicit learning comes from a case study, H.M.
sentations of wrist and digit movements, and more synap- Surgical removal of bilateral medial temporal lobes
ses per neuron (Kleim et al., 2002). Functional resulted in loss of explicit, but not implicit learning
reorganization of the human brain is shown in a study (Corkin, 2002). In most cases of motor skill acquisition,
using TMS, which showed piano practicing expanded the both implicit and explicit components are present; an
cortical representation of finger movements (Pascual- fMRI study neatly showed that basal ganglia is related to
Leone et al., 1995). Likewise, increased sensory- the implicit component of learning, and, the prefrontal
perceptual skills, for example auditory discrimination, is cortex to the explicit component (Destrebecqz et al.,
associated with the expansion primary sensory cortex 2005). These findings show the neural correlates of online
(Recanzone et al., 1992; Recanzone et al., 1993), and motor learning. A newly acquired motor skill is trans-
sharpening neural tuning (Schoups et al., 2001). In ferred from an unstable to a more stable state during con-
another study about piano practice, increased white matter solidation phase, or, in other words, during offline motor
fiber tracts were shown (Bengtsson et al., 2005). The learning. Primary motor cortex (M1) has been found to be
structure of white matter fiber tracts regulate the timing essential to early stage of motor consolidation. It has been
and speed of action potentials across axons. Neurons shown that acceleration and muscle force generation of
change their synaptic properties in response to transmis- newly learned finger movements have been disrupted
sion of action potentials. According to Hebb’s learning when M1 is stimulated by repetitive transcranial magnetic
rule, “neurons that fire together, wire together” (Hebb, stimulation (rTMS) (Muellbacher et al., 2002).
1949). Therefore, training-induced plasticity in the pri- Neural networks activated during motor learning are
mary motor cortex may occur through lasting modulations different from those activated during adaptation. During
in synaptic transmission, including synaptogenesis and the motor learning of, for example, typewriting, activation
coordinated strengthening, e.g. long-term potentiation, occurs in a large neural network of sensory and motor
and weakening, e.g. long-term depression, of synaptic cortices in the frontal and parietal lobes, basal ganglia
connections. and cerebellum. In motor learning, initially whole net-
Acquisition and retention of motor skill leads to struc- work is active but later activity in cerebellum, but not the
tural and functional changes at cortical and subcortical basal ganglia, decreases. In motor adaptation, however,
levels in the central nervous system. These changes the cortico-cerebellar network continues to be active,
evolve over time and engage different brain regions. while the activity in the basal ganglia decreases
Functional magnetic resonance imaging (fMRI) studies (Forssberg, 2008). The role of cerebellum during motor
suggested that prefrontal, parietal and partly hippocampal adaptation is very prominent, since it was found to be
brain regions, in addition to sensorimotor cortical-striato- important for predicting the state of the body during
cerebellar networks, are active during the initial stage of movement (Miall et al., 2007). The cerebellum estimates
motor learning (Kami et al., 1995; Honda et al., 1998; the sensory consequences of movement, and the response
Floyer-Lea and Matthews, 2005; Albouy et al., 2008). of the cerebellar neurons has been shown during motor
The premotor area seems to play an important role to learning of head movements. The difference between
458 PART | 7 Sensory-motor integration and performance

motor command and motor output led to robust response face deficits of perceptual and motor functions on return
of cerebellar neurons, demonstrating sensitivity of neu- to Earth. The average de-adaptation time for functional
rons to cerebellar predictions (Brooks et al., 2015). mobility of astronauts following six-month stay in space
Further knowledge about the neural correlates of sensory was found to be 15 days (Mulavara et al., 2010). Recent
motor adaptation is given in Chapter 27, The effects of research in this field attempted to enhance sensorimotor
weightlessness on human body: spatial orientation, adaptability through training that is dependent on practic-
sensory-integration and sensory-compensation. ing sensory variations of the task (Bloomberg et al.,
2015). In this practice, astronauts use a special treadmill,
whose walking surface can be manipulated to provide
Motor adaptation challenges to gait stability. Also, discordant visual stimuli
Motor skill performance is dependent on the person, per- is provided by a virtual visual scene, ensuring that gait
formance environment and the skill level interaction. and posture need to be adjusted to several sensory varia-
Context differs every time we perform the task. tions (Bloomberg et al., 2015). In this and similar
Performance may be influenced by level of stress, emo- attempts, the ultimate aim is learning to learn, in other
tional state, environmental factors, such as the ground, words, to train nervous system to adapt to changing situa-
change in temperature, and even wind speed. Motor adap- tions faster and more efficiently.
tation, usually termed sensorimotor adaptation, is the abil-
ity to adjust behavior to changing environmental or
internal demands to execute appropriate, goal-directed
Stages of motor learning
motor performance. Briefly, the difference between motor All of those who are introduced a novel skill are very
learning and motor adaptation is that in the former a new naive at the beginning, but improve their performance
motor program is learned and, in the latter, a pre-existing with practice. In the skill acquisition process, we observe
motor program is adapted. Because of adaptation, move- differences of the following performance characteristics:
ment may change in pattern of force or direction. It improvement, consistency, stability, persistence, adapt-
requires the modification of the movement based on error ability, and reduction of attention demand (Magill and
signal (Martin et al., 1996). The error signal is the differ- Anderson, 2007). First of all, over period of time, perfor-
ence between the brain’s predicted outcome of the move- mance improves. Second, as learning progresses, we see a
ment and the observed movement; the nervous system more consistent and stable performance from one trial to
needs to reorganize itself to reduce this signal (Tseng another. Persistency describes the capability of showing
et al., 2007). Once adapted to the new condition, indivi- improved performance over increasing periods of time;
duals cannot easily readapt their prior behavior; instead, e.g. days, weeks or months. Adaptability or generalizabil-
they must ‘de-adapt’ it with practice (Bastian, 2008). ity is an important aspect of performance related to skill
A good example for adaptation and de-adaptation is a learning. It refers to adaptation to changes in personal,
‘backwards’ bicycle (Sandlin, 2015), i.e. one in which task, and/or environmental characteristics. Lastly, motor
turning the handlebar to the left makes the wheel go to learning leads to reduction in attention demand, allowing
the right, and vice versa. So, riding the backwards bike an individual to show dual-task performance, such as con-
require adaptation of balance and postural control in a versing while juggling balls.
totally opposite way to normal. In the video of this exper- Most of the motor tasks require optimization of both
iment, riders struggle with their balance and fail to prog- speed and accuracy. For experts such as professional ath-
ress even a few meters. The inventor of the backwards letes, the goal is to perform a task as quickly and accu-
bike, Mr. Sandlin, reported that he learned to ride it in rately as possible. According to the well-known Fitts’
eight months, while his six year-old son took only two law, movement time (speed) can be predicted when the
weeks. Both father and son were used to ride ‘normal’ distance to the target, and the width of the target are
bikes, but the son adapted to utilizing ‘backwards’ bike known (Fitts, 1954). The formula of Fitts’ law is
much faster because of higher level of neuroplasticity dur- MT 5 a 1 b*log2 (2D/W). Movement time (MT) scales as
ing childhood. Once adapted to the backwards bike, Mr. a logarithmic function of the ratio of movement distance
Sandlin reported that he had difficulty of de-adaptation to (D) to the target width (W), where a and b are
normal bike again and needed to practice (Sandlin, 2015). coefficients.
Sensorimotor adaptation and de-adaptation processes are Derived from the Fitts’ law, there is speed-accuracy
probably more experimented in astronauts. Astronauts are trade-off, which defines the decrease in the movement
unique group, exposed to microgravity and disrupted sen- speed when the task requires more accuracy; for example,
sory information during their mission in space. The ner- to thread a needle, a certain degree of slowness is needed
vous system must organize sensorimotor information in to push the thread through the eye. If you speed the move-
space and reorganize it upon arrival to Earth. Astronauts ment up, the thread would target a larger target area, as
 Motor learning Chapter | 25 459

can be derived from the formula. The question is whether Ann Gentile proposed another two-stage model with a
Fitts’ law holds its predictive value during motor learning, perspective on the learner’s goal. In the initial stage of
since during learning, movements become faster and learning , movement pattern is acquired and the condi-
more accurate while target distance and target width tions in the environmental context are learned and identi-
remain the same. The answer is that Fitts’ law is still fied in order to achieve the action goal. In later stages,
found to be valid, but with changed coefficients (Latash, she highlighted improvement in three characteristics of
2012). the action: adaptation, consistency and economy of effort.
The distinct stages of motor learning have been She also proposed that learner’s goals depend on the type
defined by certain models. Despite different terminology of skill practiced (Gentile, 1972). In closed skills, learner
and stages defined, all models essentially agree on the shows fixation on the movement pattern learned in the
main characteristics of the learning process. Below, the initial stage, and refines it to achieve consistency and
models are briefly described. economy. An example of a closed skill is writing with the
Fitts and Posner three-stage model proposes a process same type of pen on the same type of surface. Open skills
of cognitive, associative and autonomous stages. This require the learner show diversification of the basic move-
model highlights that in the early stages of learning there ment pattern learned in the initial stage, thereby motor
is much higher cognitive demand, such as problem solv- adaptation. Gentile’s perspective provided a framework
ing, working memory and attention until the skill for goal-directed training for rehabilitation settings
becomes automatic (Magill and Anderson, 2007). (Mastos et al., 2007).
Bernstein’s three-stage model focuses on the The final model is Halsband’s three-stage model,
changes in motor control in consecutive stages of motor defined in neuropsychological terms as follows: In the ini-
learning. In order to understand his theory, it is impor- tial stage, performance is under close sensory guidance,
tant to understand the degrees of freedom problem. The and the accuracy and speed of performance varies greatly
number of independent components and how each com- from trial to trial. In the intermediate stage, sensory-
ponent can vary creates numerous degrees of freedom motor connections become stronger; movements become
for the motor system. For example, in order to reach faster, and more accurate, and are executed with smaller
object, many muscles and joints have to be controlled variance. In the advanced stage, performance is fast, auto-
by the nervous system. During motor learning, mated, and skillful, with isochronous movements and
Bernstein suggested that the motor system has to solve whole sensory field control (Halsband and Freund, 1993).
this problem. At the first stage, he proposed an inhibi- Halsband’s model was demonstrated in a study where
tory pattern that there is an elimination of, or freezing, individuals were trained with auditory information for
redundant degrees-of-freedom to simplify the control of reach-and-grasp task. This task was designed to give audi-
the movement. In the second stage, control becomes tory information regarding the size of the objects to grasp
more comfortable, which is associated with releasing to allow individuals to establish a new audiomotor map.
degrees of freedom that were previously frozen. In the They found no overt signs of learning in the first approxi-
last stage, there is an optimal interaction with external mately 10 15 trials; subjects used the maximum grip
forces that leads refinement of control strategies, mean- aperture to achieve the task. This was followed by a
ing that a person learns to use, rather than fight against period of fast learning, where individuals adjusted the
external forces (Latash and Turvey, 1996). Bernstein’s hand grip according to the size of the object via auditory
model was experimentally supported; a recent finding in information, and in the final stage they reached a plateau
transtibial amputees confirmed the freezing and releas- of learning (Säfström and Edin, 2006).
ing stages during lower limb joint flexion and extension
(Wurdeman et al., 2014).
There is also a two-stage model proposed by Latash,
Learning skill in different life stages
which relies on the changes of motor synergies. Not all behavioral changes result from learning.
According to this model, variance of performance charac- Phylogenetic memory was shaped during evolution, and
teristics can be viewed as having two components; good results in some innate behavioral responses, such as fear.
and bad variance . In the first stage, bad variance Recent study in mice showed the strong memory traces of
decreases while good variance shows almost no change, fearful experiences in subsequent generations. Fear
which strengths the motor synergy. In the second stage, response to conditioned odor was found to enhance neuro-
bad variance decreases as much as possible, while good anatomical representation of the odor in the brain and,
variance decreases to the optimum level, which weakens more importantly, this was selectively transmitted to next
the motor synergy. With practice, bad variance decreases generations (Dias and Ressler, 2014). What about human
so, for example, accuracy of performance increases behaviors? How much genetics and experience contribute
(Latash, 2012). to learning? This question touches the controversy of the
460 PART | 7 Sensory-motor integration and performance

effect of ‘nature versus nurture’ on development. with participants aged in 6 to 89 years showed the perfor-
Developmental studies, especially on infants, have inves- mance of older adults was comparable with those of chil-
tigated whether genes predetermine motor development, dren and young adults. The study showed the best
or whether it is a result of practice and interaction with outcome for teenagers and young adults between 15 and
the environment. 29 years (Voelcker-Rehage and Willimczik, 2006).
According to the developmentalist Gesell (1934)
motor development or acquisition of motor skills such as
Motor learning in expertise
rolling, sitting, standing, walking appear in a sequence
and within specific periods during infancy, emphasizing Experts are a special group of people who develop skillful
the role of genes in unfolding development of motor skills actions above the level of normal performance. The area
(Gesell et al., 1934). On the other hand, recent dynamic of expertise can be in cognitive domains like science,
systems theory suggests that motor development is not a mathematics, chess-playing, or in motor domains such as
simply passive process in which genes dictate the devel- athletics, piano playing or ballet. There are some key
opmental milestones over time. Rather, infants actively characteristics of experts. First, they deliberately practice
perceives and acts upon the constraints set by their bodies the skill for extensive periods. Second, they have a strong
and environment to achieve a goal. Therefore, according interest in the skill, and thereby, motivation. Lastly, they
to other developmentalists, infants assemble motor skills prioritize skill training over other significant daily life
for perceiving and acting (Keen et al., 2014). However, activities. In motor learning, experts are a valuable group
experiments such as the “visual cliff”, suggest that infants for investigating enhancement of performance of a
are born with depth perception that directs their motor learned or highly experienced motor skill.
skill acquisition (Feldman and Garrison, 1993). Musical expertise in piano players has been exten-
Myelination is very important for motor skill devel- sively studied by Fredrik Ullen’ s research group. They
opment during childhood. Training or practice during measured the amount of practice (total hours of practice)
childhood has very prominent effects on myelination. during different developmental periods; childhood, ado-
Motor practice induces the neural activity in fiber tracts, lescence and adulthood, both in piano players and in non-
and therefore induces myelination. In a study by musicians. Longer practice time and white matter changes
Bengtsson et al., in which they compared the neural cor- were found in all groups, regardless of age. Age was gen-
relates of extensive piano practicing during childhood, erally found as a strong predictor for brain development.
adolescence and adulthood, they found regional specific However, this finding may show that practice may hinder
changes in pyramidal tract. Most importantly, largest the age-development relationship (Bengtsson et al., 2005).
number of brain regions correlated with childhood prac- Macro anatomical changes have been observed in musi-
ticing, even compared to much more intensive practice cians. Increase in cortical gray matter volume and higher
in adulthood. This example explains the interaction fractional anisotropy in white matter have been reported.
between nature and nurture, and indicates a develop- These changes have been shown in the corpus callosum,
mental window in which white matter plasticity is motor cortex, cerebellum and planum temporale of musi-
highly favored (Bengtsson et al., 2005). cians (Schlaug, 2015). Cortical excitability has also been
Older adults is another group for whom it is important shown after long-term motor training; musicians showed
to consider for motor learning. Increase in aging popula- less interhemispheric inhibition (Nordstrom and Butler,
tion and longer life expectancy in many countries high- 2002).
light the motor learning capacity and plasticity in this Expertise in professional athletes has been one of the
group. With increasing age, musculoskeletal and nervous most intriguing areas in the field of motor learning.
systems, and sensory organs are negatively affected. Recent technological advances in methods to study kinesi-
Seeing, hearing, touching are important sensory functions ology provided valuable data. Video-based analysis of
for motor system, and decline in these sensory functions technical movements showed that famous football player
are significant in individuals 75 years and older Lionel Messi makes more use of dribbling and feint of
(Santrock, 2006). According to a study, older adults take change of direction than Cristiano Ronaldo, meaning that
longer to move than young adults, and this difference Messi showed variability, and was more
remains for both easy and difficult movement tasks unpredictable (Castañer et al., 2017). It has been shown
(Ketcham and Stelmach, 2001). With these information, it that humans predict others’ actions by using bell-shaped
is no surprise that perceptual-motor coupling is declining, velocity profile (Stadler et al., 2012). However, data that
and leading problems for already learned motor skills movement of Messi is unpredictable, i.e. he does not fit
such as driving (Santrock, 2006). On the other hand, into this bell-shaped velocity profile. A study by
learning new motor skills is intact, but with a slower rate. Anderson and Sidaway showed that as an acute result of
In a study, six days of training in a novel task, juggling, practice, novice soccer players increased linear velocity
 Motor learning Chapter | 25 461

of the foot in relation to changing pattern of coordination organs. More intriguingly, it can be seen during childhood
underlying the movement (Anderson and Sidaway, 1994). development without any apparent physical or intellectual
Trial-to-trial variability in the execution of movements is problem. Developmental Coordination Disorder (DCD) is
usually considered a drawback of the noisy sensory motor one of these developmental disorders. In DCD, children
system, however, recent studies have suggested that motor fail to learn complex coordinated movement skills, such
variability may also be a feature of how sensorimotor sys- as riding a bike, or handwriting. Motor learning problems
tems operate and learn. This view stems from the rein- recently proposed as an explanation of co-occurence of
forcement learning theory, and sees motor variability as neurodevelopmental disorders such as ADHD, dyslexia,
purposeful exploration of motor space that can drive language impairment and DCD. The “procedural deficit
motor learning (Dhawale et al., 2017). hypothesis” suggests that commonly observed clumsiness
The variable practice, rather than repeating the same and learning deficits in these mildly effected groups are
task, is favored in athlete training programs (Li and Lima, results of procedural (or motor) learning deficit (Nicolson
2002; Shoenfelt et al., 2002) because sensorimotor adap- and Fawcett, 2007). We use the motor learning system to
tation leads to a high level of plasticity and learning acquire the movement patterns to produce speech sounds
effect, which in effect teaches the system to learn to learn. and motor actions. During development, most affected by
Schmidt’s schema theory of motor learning suggests that motor learning deficit are speed and accuracy of manual
the motor system creates a library of motor actions (sche- movements during complex task, e.g. drawing or hand-
mas) for classes of motor problems (Schmidt, 1975). In writing, and articulation of complex speech sounds.
each trial, the performer stores the initial condition, spe- Attention mechanisms are known to be important in the
cific surrounding, sensory consequences and the outcome initial phase of motor learning. Consequently, impair-
of the movement. Over trials, varying conditions of prac- ments in speech and motor domains, which require learn-
tice allows the motor schema to become optimized. When ing of complex sequence of movements, are commonly
a novel motor task is presented, the person, if trained with observed in neurodevelopmental disorders. Childhood
variable practice, is able to retrieve the appropriate move- Apraxia of Speech is considered as a neurodevelopmental
ment patterns more efficiently (McCracken and Stelmach, disorder that exhibits the deficit in motor learning,
1977; Shea and Morgan, 1979; Catalano and Kleiner, because affected children showed lower performance both
1984; Sherwood, 1996). in speech and complex manual skills such as drawing
Another focus in athlete training has been on whether (Tükel et al., 2015). Autism Spectrum Disorders (ASD) is
implicit or explicit processes lead superior performance, another group with defined motor learning difficulties,
and accordingly, which components of performance bene- despite the absence of any brain lesion or malformation
fits. It has been found that implicitly learned motor skills (Ming et al., 2007).
tend to be stable under performance pressure (Hardy
et al., 1996; Mullen et al., 2007) and physiological fatigue
(Poolton et al., 2007; Masters et al., 2008), and also
Motor learning in rehabilitation
demand less attention (Maxwell et al., 2003; Poolton The primary goal of rehabilitation is to induce neuro-
et al., 2006). Due to difficulties in designing experiments plasticity from cortex to spinal cord for restoring motor
involving implicit and explicit learning, there are only and cognitive skills. Motor learning principles are
small number of studies. In one study (Lam et al., 2009), used in the rehabilitation context to facilitate neuro-
in modified basketball shooting task, three groups were plasticity and motor recovery. Experiments in healthy
compared after receiving three-day training. The first subjects highlighted that practicing the task under var-
group received explicit learning via eight technical ied sensory stimuli, rather than repeating the same task
instructions, the second group received implicit learning in stable conditions, leads to better outcomes that can
via single analogical instruction, and the final group be described as faster adaptation (Roller et al., 2001).
received no instructions. No performance differences This means that the sensorimotor system learns to
were found between groups in retention tests, indicating adapt to changing conditions better, in other words, the
that there was no difference between learning models. system is more plastic. This principle has been trans-
However, in transfer test, despite the lack of any kine- ferred to therapeutic approaches in Down syndrome
matic changes, performance deteriorated for both explicit (Latash et al., 2002), and to neurorehabilitation of
and control groups, but not for the implicit group. stroke patients (Krakauer, 2006).
Rehabilitation programs in stroke patients showed the
potential variation in groups with different severity of
Disability of motor learning dysfunctions. In a study by Hardwick et al. 4-day training
Disability of motor learning can be seen in diseases that was given to stroke patients and healthy controls. Stroke
affect nervous and musculoskeletal systems, and sensory survivors were divided into two groups according to
462 PART | 7 Sensory-motor integration and performance

motor impairment: mild-to-moderate and moderate-to-

severe. Groups practiced isometric contractions of elbow
flexors to navigate an on-screen cursor to different targets
for 4 days. The speed-accuracy trade-off function was
assessed for each group before and after the training, and
controlled for differences between individuals in self-
selected movement speeds. All groups were able to
improve their performance through skill acquisition. The
moderate-to-severe group reached the baseline perfor-
mance of the mild-to-moderate group, and the mild-to-
moderate group reached the baseline performance of the
control group (Hardwick et al., 2017). Robot-assisted
therapy seems to have strong potential for improving
upper limb function in stroke patients. It was shown more
effective than usual care when applied for 12 weeks and FIGURE 25.3 A child with unilateral cerebral palsy (CP) is shown dur-
36 weeks; however, intensive therapy outperformed ing handwriting test. Learning handwriting refers to copying readable
robot-assisted therapy (Lo et al., 2010). letters therefore; it requires speed and accuracy of fine motor movements
along with visual perception and visual working memory. Children with
There is a greater amount of research on motor learn- unilateral CP usually use their non-effected hand for handwriting while
ing during development in the case of brain lesion in chil- using their effected hand to stabilize the paper as shown in the picture.
dren with cerebral palsy (CP) conducted by Ann-Christin Motor learning of copying readable letters take longer time period for
Eliasson’s research group. Sensorimotor reorganization of children with CP compared to those of their typically developing peers.
the central nervous system, in case of early brain lesion, Learning rate in CP group can be predicted by age and IQ.
shows an impressive capacity for compensation and pres-
ervation of functional motor skills. In unilateral CP, con- and facilitate subsequent movement execution. The inten-
tralateral corticomotor projection pattern was found tion to imitate movements shown on a video resulted bet-
important for the speed and accuracy of reaching, grasp- ter outcome for functional upper limb dexterity for stroke
ing, releasing and functional tasks (Holmström et al., patients (Franceschini et al., 2012). Similarly, for
2010). Two weeks of motor training with constraint Parkinson’s disease, imitation of motor actions seems a
induced movement therapy (CIMT) improved manual promising rehabilitation tool (Caligiore et al., 2017). In
skills irrespective of corticomotor projection pattern and healthy individuals, observational motor learning has
brain lesion characteristics (Islam et al., 2014). Motor been found very effective for implicit learning of sequen-
learning of handwriting skill has also been investigated in tial complex motor skills. Findings from SRT task showed
children with unilateral CP, and findings indicated a that subjects can acquire sequence information by watch-
slower rate of learning in motor skill acquisition, which ing another person performing the task, and observation
can be predetermined by chronological age and IQ score results in as much sequence learning as task practice. This
(Fig. 25.3) (Tükel Kavak and Eliasson, 2011). study also showed that sequence knowledge acquired by
Observational learning and the so-called ‘mirror neu- model observation can be encoded motorically (Heyes
ron system’ has become one of the most promising and Foster, 2002).
approaches in rehabilitation. During the last decade, vari- Since 1994; when Salinas & Abbot showed that the
ous studies were carried out regarding the clinical use of direction of resulting limb movement can be computed
action observation for motor rehabilitation of sub-acute from measurements in fewer than 100 primary motor cor-
and chronic stroke patients. Mirror neurons are a specific tex cells (Salinas and Abbott, 1994), the activation of pri-
class of neurons that are activated and discharge both dur- mary motor cortex has been used to guide prosthetic limb
ing observation of another individual’s motor act, and in prosthetic rehabilitation, suggesting applications for
during the execution of the same or similar motor act patients with amputated or paralyzed limbs. One study
(Rizzolatti et al., 1996). By using TMS and fMRI, differ- demonstrated that two tetraplegic patients could exert
ent studies have demonstrated the presence of the mirror some control over the speed and direction of movement
neuron system and their mechanism in humans. Studies of a computer cursor based on recording of 96 neurons in
have demonstrated the activation of brain areas involving the dominant hand area of the primary motor cortex (Kim
inferior parietal lobe and the ventral premotor cortex, as et al., 2008). In a more recent study, Hochberg et al.
well as the caudal part of the inferior frontal gyrus, when showed that tetraplegic patients are able to move a robotic
individuals learn motor actions via execution, imitation, arm for reaching and grasping, and in one case, use the
observation and motor imagery. This means that observ- robotic arm to drink from a bottle (Hochberg et al., 2012).
ing or imagining an action can affect cortical excitability Of course, there are high levels of variation between
 Motor learning Chapter | 25 463

individuals in these experiments. The impressive finding Byrne, J.H., 1997. Neuroscience online: an electronic textbook for the
is that individuals show neuroplasticity and can learn to neurosciences. Chapter 7. Learning and Memory. 2018. From
fire their motor neurons to control the robotic arm. [Link]
In rehabilitation setting, studies also highlight the Caligiore, D., Mustile, M., Spalletta, G., Baldassarre, G., 2017. Action
observation and motor imagery for rehabilitation in Parkinson’s dis-
strong relationship between motor learning and emotional
ease: A systematic review and an integrative hypothesis. Neurosci.
and motivational factors, thus are often use the words
Biobehav. Rev. 72, 210 222.
‘goal-directed’ or ‘targeted’ to describe a movement, Castañer, M., Barreira, D., Camerino, O., Anguera, M.T., Fernandes, T.,
because all movement has an aim. Motion, cognition and Hileno, R., 2017. Mastery in goal scoring, T-pattern detection, and
emotion are interconnected, faster and more efficient polar coordinate analysis of motor skills used by Lionel Messi and
motor learning can be facilitated by motivation and cogni- Cristiano Ronaldo. Front. Psychol. 8, 741.
tion. This interaction is well-known by clinicians working Catalano, J.F., Kleiner, B.M., 1984. Distant transfer in coincident timing
with patients during the recovery phase, but without tools as a function of variability of practice. Percept. Mot. Skills 58 (3),
to motivate the patients, is often overlooked. It is impor- 851 856.
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