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Mirror Neurons and Empathic Resonance:
Action and Understanding
A Research Paper
Presented to
The Faculty of the Adler Graduate School
_____________________________
In Partial Fulfillment of the Requirements for
The Degree of Master of Arts in
Adlerian Counseling and Psychotherapy
_____________________________
By:
Elizabeth M. Nash
October 2006
Action and Understanding
Running Head: Mirror Neurons and Empathic Resonance
Mirror Neurons and Empathic Resonance:
Action and Understanding
A Research Paper
Presented to
The Faculty of the Adler Graduate School
_____________________________
In Partial Fulfillment of the Requirements for
The Degree of Master of Arts in
Adlerian Counseling and Psychotherapy
_____________________________
By:
Elizabeth M. Nash
October 2006
1
Action and Understanding
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Abstract
To survive we must understand the actions of others. With this action understanding social
organization and empathy become possible. This empathic resonance requires abilities that are
provided by the mirror neuron system. Empirical evidence supports this system’s role in action
recognition and predication. Implications of the evidence are discussed in relation to our ability
to associate self and others. Understanding failures of the mirror neuron system may help us
understand social and communication problems seen in autism spectrum disorders. This
understanding also emphasizes the importance of therapists being emotionally, intellectually and
behaviorally congruent models for clients.
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Mirror Neurons and Empathic Resonance: Action and Understanding
This paper discusses recent evidence that describes a system, a mirror neuron system
(MNS), which allows for our ability to understand and imitate actions of others. Investigations of
this system are providing information on how the brain interprets action, recognizes the purpose
of actions and comprehends the meaning of language. This neural system has intriguing
implications regarding how the brain interprets and recognizes the meaning of other’s actions.
The MNS also may explain much about imitation and language acquisition.
Typically, we are linked with the actions, emotions and sensations of others. This
resonance allows us to understand others. Effortless for most, we are able to understand others as
goal oriented persons like us. There are basic brain/ body functions by which we can understand
and model interactions in the world. These functions allow us to resonate empathically with
others. Conceptualization of the implementation of mental functions within the brain has been
changed in some basic ways in the last five years. This paper discusses this new
conceptualization and suggests applications of the MNS in the therapeutic process.
Mirror Neurons
Our Relationship with Objects
Much of our waking hours are spent interacting with objects in the environment. One
way of conceptualizing consciousness may emerge out of an understanding of an individual’s
relationship to objects that make up our world known as “object proxy” (Gallese, 1996).
The study of learning, perception, memory and language has provided much information
regarding the sensory processes that are used to interpret the world. The MNS provides a motor
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interpretation based on the actions of self and others in addition to sensory interpretation (Gallese
& Metzinger, 2003).
Our interactions with objects are represented in the brain through neural mechanisms that
map the object’s characteristics (Gallese, 2003) . The knowledge of an object can then be
interpreted through these neural mechanisms creating a motor vocabulary. A motor vocabulary is
a vast schemata set which themselves are sets of actions that accomplish various goals. The
knowledge is stored to memory so it can be recalled or recognized. It is also categorized
conceptually or in linguistic terms. The MNS provides a mechanism by which objects are
understood.
Monkey see, Monkey do
Mirror neurons are a particular class of visuomotor neurons. These neurons are found to
discharge when an individual does a particular action and when they observe another doing a
similar action. Mirror neurons were first discovered in monkey’s ventral premotor cortex known
as F5 (Gallese, Fadiga, Fogassi., Rizzolatti, 1996). Further study of motor functions in humans
has provided evidence of a similar group of neurons in humans (Fadiga, Fogassi, Pavesi,
Rizzolatti, 1995).
Research with monkeys was able to identify the involvement of the ventral motor cortex
(F5) when monkeys reach and when they grasp an object (Gallese, Fadiga, Fogassi, Rizzolatti,
1996; Murata, Fadiga, Fogassi, Gallese, Rizzolati, 1997). The research described served to
differentiate various aspects of our interaction with objects and also provided information about
the visuomotor properties of single neurons.
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Figure 1: F5 and F1 area of monkey 1
Fff
Figure 1: F5 and F1 area of monkey 1
__________________________________________________________________________________________________________________
1. From ilab.usc.edu/classes/2002cs54/lecture_notes/24-Re. retrieved 10-08-2006.
Single neuron recordings were made while monkeys did behavioral tasks. The data found
activation of the F5 area when the monkeys reached and grasped an object and surprisingly, also
when they observed another individual doing the same action. The researchers found different
areas in the F5 area responsible for different parts of the motor movement. The F5 bank area was
found to be involved with the motor control during the action. The F5 convexity was found to be
the area by which the hand is shaped (grasp). By inactivating neuron groups, the investigators
sought to discover the distinct function of these areas. Inactivation of the F5 area caused the
monkeys to be distinctly impaired in their grip (Fogassi, Gallese, Buccino, Fadiga, Rizzolatti,
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2001). The monkeys had difficulty shaping their hand to conform to the objects shape and size.
When the F5 bank area was inactivated, the monkeys became impaired in their movement
toward the object. Inactivating the F5 area located on the convexity area produced only some
deficits. There was a slowing of the movement, but the ability to shape the hand was not
affected. Inactivation of F1, known for control finger movements, caused severe paralysis (figure
1). No neurons in F1 responded to the observation task. The study showed that areas of the
ventral premotor cortex have specificity as to what their functions are. They are involved in
distinct distal and proximal organization of movement. It has been shown that all mirror neurons
show congruence between the observation of visual action and the motor responses coded by
them (Fogassi et al., 2001; Gallese et al., 1996). This allows us to copy what we’ve seen on a
motor level, an action level, without cognitive mediation.
There is a relationship between mirror neuron’s visual and motor properties that is
important to their function. All areas of motor cortex and MNS would be important for action
imitation but there are some areas that posess only motor movement ability. Motor areas are used
simply to program motor neurons. The mirror neurons interpret motor function and sensation.
Motor response, sensation and mirror neurons react to observations and program motor action, so
they can both respond to the external world and arrange meaningful responses to it.
Mirror neurons are responsive when a monkey grips an object and when they see another
individual (human or monkey) doing the same task (Gallese et al., 1996). To be effective in
movement, motor action and observation overlap. They coincide in reaching a goal (grasping)
and how the goal is achieved (grip).
The previous research focused on hand movements. The next investigation sought to
discover if the congruence found in mirror neurons could be expanded to the mouth region
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(Ferrari, Gallese, Rizzolatti, & Fogassi, 2003). The location of the set of mirror neurons involved
with mouth movements was located in the lateral part of F5 (figure 1).
Firing rates of neurons were measured while ingestive actions were performed by the
monkeys and when the monkeys observed the investigators doing the same action. The ingestive
action tasks were grasping food with the mouth, sucking from a syringe, food preparation, lip
protrusion and lip smacking. Seventy-four mirror neurons were studied. Ten trials of each action
were measured per monkey. To monitor where the monkeys were looking and what they were
observing, the investigators monitored and tracked their gaze and eye movement. Most of the
time the monkeys gaze was on the investigator’s mouth during the observation of ingestion tasks.
To control for the ingestion actions, neurons were also measured while the monkeys observed
the investigators during a neutral gaze. Electomyograph (EMG) recordings of face muscles
during observation of ingestive actions did not measure any muscle movement. No difference in
discharge was seen when the same gestures were made by different experimenters.
Ferrari et. al. showed 25% of the neurons have mirror properties. Ingestive and
communicative mirror neurons were distinguished (Ferrari, et. al. 2003). Communicative mirror
neurons responded to lip smacking and lip protrusion. The ingestive neurons respond when the
monkeys observed action related to ingestive functions (grasping food, breaking it and sucking).
Almost all ingestive mirror neurons fired whether watching the action or doing the action. This
shows a correspondence between the action and the action’s goal in both observation and action.
About one third of the mirror neurons measured were classified as “strictly congruent”
mirror neurons. Strictly congruent mirror neurons are those in which the observation and effect
of the action corresponds to a goal and a means of reaching the goal. In the experiment, the effect
observed and firing neurons were almost identical. The remaining neurons were classified as
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“broadly congruent” mirror neurons. Broadly congruent mirror neurons do not require the exact
observation of the same action to fire. This allows us to have flexibility in figuring out the goal
of an action.
The existence of mouth mirror neurons is of interest because they may support the
cortical mechanisms (e.g. auditory, sensory motor cortex) by which we understand speech.
Communicative gestures resulting from ingestive actions may be the mechanism by which
humans developed speech.
As previously found, neurons were highly congruent with the specific task. Different
ingestive actions caused different location of neurons and number of neurons to respond. The
differences in the measured discharge over specific areas indicated that the monkeys were able to
differentiate the different gestures that were presented to them. For instance communicative
mouth neurons responded to lip smacking.
Audio-Visual Mirror Neurons
Another interesting finding is that action understanding can be accomplished without
actually seeing the object. Object-related actions can be recognized by their sound. Neurons in
the monkey premotor cortex were found to discharge when the animal performs an action and
when a related sound is heard (Kohler, Keyser, Ultima, Fogassi, Gallese & Rizzolatti, 2002). In
addition, most of the neurons discharged when the animal observed the same action. In three
macaque monkeys a total of 497 neurons were recorded. Properties were studied that responded
to the experimenter’s action related and non-action related sounds. Investigators first exposed the
monkeys to the site and sound of the paper ripping. Neuron one responded to the sound and sight
of paper ripping. Then the sound of paper ripping was presented out of the sight of the monkey.
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Neurons measured equal discharge as the other two observable events. Non-action related sounds
showed no response (white noise). In neuron two, the neurons discharged on the sound and sight
of a stick dropping. Once again, non-action related sounds did not produce consistent discharge.
The goal of the action triggers mirror neuron firing.
Neurons that respond to sound have been found in several areas of the brain. Previously,
neurons have been found that respond only to the location or direction of a sound. These neurons
code sensory information. Researchers located neurons in the F5 area that discharge on execution
and observation of a specific action and interestingly, when the action could only be recognized
through its sound (Keysers, Kohler, Ultima, Nanetti, Gogassi & Gallese, 2003). These “audiovisual mirror neurons” have a different function. Audio-visual mirror neurons help with the
understanding of action ideas in terms of their goal. When you hear a doorbell you don’t just
locate where the sound is coming from or that you heard it. You know that someone rang the
doorbell. In addition you realize that someone is at the door and you might want to answer the
door. By hearing a sound, motor ideas are evoked. The sound represents the actions contents.
Audiovisual mirror neurons allow for action understanding without seeing the action.
A similar experiment was conducted to understand if mirror neurons could discriminate
action when different modalities are used (visual or auditory) (Keysers, et al., 2003). In the
experiment ten slightly different recordings were played for monkeys during the tasks. The
audio-recordings were slightly different to avoid the monkey hearing the same thing repeatedly
and becoming primed for the response. The recordings were somewhat varied in frequency,
distance from the animal, amplitude and pressure.
The monkeys could discriminate the sounds 97% of the time, almost perfectly. In
addition audio-visual mirror neurons discharged at the same intensity whether the action was
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seen or heard, or both heard and seen. This implies that the monkeys were coding the action in an
abstract way. For the monkeys, ripping a piece of paper is just that, whether it is seen or heard.
These findings suggest a mechanism by which humans acquire and use abstract thought.
Mirror Neurons in Humans
Unlike research in monkeys, there are no single cell mirror neuron recordings from
humans. Neurophysiological and brain imaging experiments provide the evidence of a mirror
neuron system in humans. While this evidence is not as direct as single cell recordings it is
compelling.
In humans, seeing actions by another performed by others activates three areas of the
brain which include the superior temporal sulcus (STS), the inferior parietal lobule (IPL) and
inferior frontal gyrus (IFG) (Rizzolatti & Craighero, 2004). The STS contains neurons that
respond to moving body parts and the IPL and IFG areas in humans correspond to areas in
monkey’s preventral cortex and premotor cortex (including the F5 area) where motor neurons
have been recorded in monkeys (Rizzolatti & Craighero,2004).
Research using transncranial magnetic stimulation (TMS) provided most of the direct
evidence of a mirror neuron system in humans (Gallese, 1996). TMS is used to stimulate an area;
however it does not allow specific location of an effect. Cortical magnetic signals were recorded
using a 122-channel SQUID neurogradiometer to locate the areas response. Research conducted
on humans aimed to identify activation of the precentral cortex when an action was executed and
when an individual observed those movements in another (Hari, Forss, Kirveskari & Salenius,
1998). Instructions were given to the participants to view the actions. The ten participants in the
study did three tasks while undergoing TMS. The three tasks were performance of an action,
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manipulation of an object and observation of another doing the same action. Instructions were
given to the participants to view only the actions but it cannot be ruled out that the participants
used motor imagery (imagining the action) components during the observation. As a control the
participants also observed stationary or moving stimuli as additional tasks.
The investigator’s results were in agreement with other TMS studies (Hari, et al., 1998).
The study concluded that there is an execution matching system in humans similar to those found
in monkeys. Activity recorded over the precentral cortex (15-25 HZ) indicated the primary motor
cortex activates during action observation and action execution in humans. An interesting
additional finding showed no increase in muscle activity during observation but a small but
significant increase in muscle activation was seen during active motor imagery, imagining a
movement. This might imply activation of additional areas that are involved with only motor
control in addition to the mirror areas which involve a goal directed motor movement during
imagining of a movement.
Electrophysiological Evidence
Electroencephalography (EEG) in humans has differentiated between two rhythms at rest
in the alpha frequency range (8-13 Hz); a posterior alpha rhythm and a central mu rhythm
(Niedermeyer, Da Silve, 2005). Both the alpha rhythm and mu rhythm are seen at about 10 Hz.
They are an idling rhythm of neurons. When an individual performs an action the neurons fire
asynchronously, thereby reducing the power of the mu-band oscillations. The posterior alpha
rhythm is present when sensory systems, in particular vision, are not activated. Close your eyes
and this rhythm occurs. During motor rest the mu rhythm appears, while during motor activity it
disappears. Desynchronization of mu rhythm is known to occur during observed movements.
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It is important to note that it is impossible to differentiate mirror neurons from their
adjacent motor neurons, or sensorimotor neurons using EEG. It is likely that mu wave
suppression is the result of activation of neuronal systems in the premotor and sensorimotor
cortices (Neidermeyer & Da Silva, 2005). During hand movement the mirror neuron system is
known to be the only one activated (Pfurtscheller, Neuper, Andrew & Edlinger, 1997). Therefore
the mu rhythm is a useful method for monitoring this brain activity non-invasively in humans
(Mcfarland, Miner, Vaughn & Wolpaw, 2000; Muthukumarmawamy & Johnson, 2004).
It is proposed that mu rhythm reflects the operation of an information processing system
that links perception and the transformation of seeing into doing (Pineda 2005). Mu rhythms are
sensitive to differences in observed motor action. Research was conducted while subjects were
recorded using a 128-channel EEG during three tasks. The tasks were observation of an object,
imitation of an object being gripped (no object) and precision gripped an object
(Muthukumaraswamy et al. 2004). Mu rhythms were lower when the subjects observed an object
being gripped than during the imitation of an object being gripped (no object). Lower mu rhythm
amplitude indicates desynchronization of the cells. This reflects active processing during
observation of grip. Mu rhythm is measured on the surface of the skull so it is unlikely that they
directly measure the activity of cells. EEG does measure cell system coordination or coherence.
Muthukumaraswamy et. al. concluded likely the EEG measurement is sensitive to the mirror
neuron system since they have similar functional properties (Muthukumaraswamy et al. 2004). In
addition EEG recordings showed that the effect was separate from muscular activity that might
be evident if the participant were actually physically gripping the object (Muthukumaraswamy
et al. 2004). The study provides evidence of sensorimotor desynchronization during execution,
observation and imitation of biological movements. An interesting additional finding shows that
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the sensorimotor cortex desynchronizes during imagined hand movements (Pfurtscheller et al.,
1997). It is also known that that individuals can learn to volitionally control mu rhythms. These
findings might provide the mechanism by which paraplegics learn to execute motor acts such as
powering a motor to propel a wheel chair by imagining it so.
EEG recordings in 10 high functioning autistic individuals showed impairment in their
mu rhythm (Oberman, Hubbard, McCleery, Altschular, Ramachandran & Pineda, 2005). As
discussed previously, Mu rhythm is suppressed during action and action observation. Recordings
were made using controls and children with autism as they opened and closed their hand and
observed a video of the same movement. The hand moved at an approximate rate of 1 Hz. An
additional video was observed of a bouncing ball. Observation of a video of white noise was
used as a control. The viewing distance was constant. In the experiment the direction of the gaze
was not monitored so it is unclear what they were looking at or for how long. Since children with
autism are known to have impairment of gaze future experiments should include a control for
this.
During observation of action and execution of action, the control group showed a normal
response; suppression of the mu wave rhythm during observation of an action and execution of
an action (Oberman et al. 2005). In the children with autism there was suppression of the mu
wave rhythm only with their own movement. Suppression of the mu wave rhythm did not occur
during observation. Oberman (2005) concludes that this could indicate a dysfunction of their
mirror neuron system. It fits with the behavioral symptoms of autism spectrum disorders where
deficits of communicative and social skills, imitation, pragmatic language and empathy are
found.
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There is evidence of mu rhythm attenuation in children under eleven years of age, during
observation of hand movements as is seen in adults (Lepage & Theoret, 2006). Failure of mu
attenuation during observational learning may be of assistance in the understanding and
diagnosis of developmental disorders.
These investigations in mu blocking could provide a basis for early diagnosis for autism
(Oberman, et al., 2005). However the repetitive movements of low functioning autistic
individuals could cause artifact in the EEG and make it difficult to interpret. Early practice in
imitation might enhance these children’s abilities. There are many programs that work with
these children from a young age on these skills. However, the mirror system defect might be just
part of the dysfunction. There may be sensory problems as well. However if it were purely a
problem with gaze they might be inclined to bump into things which is not commonly seen. They
are known however to do repetitive motions and behaviors that imply a motor abnormality in
keeping with a mirror neuron deficit rather than purely a sensory system deficit.
Characteristics of Mirror Neurons
One of the most intriguing qualities of mirror neurons is that they only activate when
there is movement by a biological effector (hand or mouth) to an object. A tool moving an object
does not suffice. A biological effector’s movement alone without an object does not activate
mirror neurons. Mirror neurons are activated during a goal directed action. They activate in
addition to the purely motor neurons that guides the movement.
Mirror neurons have basic properties. There are two classifications of neurons in
monkeys and humans, canonical and mirror (Grezes, Armony, Rowe & Passingham, 2003).
Canonical neurons respond when viewing an object that can be grasped. It is as if the brain is
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coding a future action, an item that could be grasped. Mirror neurons activate when goal-directed
observation occurs and during goal-directed execution of an action. The sight of the object alone
or an agent mimicking an action does not cause an effect in mirror neurons.
It is understood that mirror neurons fire when there is an interaction between a biological
effector and an object (Umilta and Kohler, 2001). Umilta et. al. addressed whether the effector
and object always need to interact to invoke a mirror response. He found that monkeys, like
humans can infer the goal of an action even when visual information is incomplete.
Mirror neurons are triggered by visual stimuli only when a biological effector interacts
physically with the object (Fogassi, et al., 2001). However, vastly different stimuli are effective
in activating mirror neurons as long as the stimuli represent the same action. The distance from
the observation (size of the hand viewed) does not affect the response. Bigger arms or bigger
objects do not change the response, so it is not merely a sensory response. Neuronal activity is
seen in mirror neurons without the action being rewarded. Non-significant items (a geometric
solid) and significant items (food) generate an equal response in mirror neurons in monkeys.
Mirror neurons are involved in perception and comprehension of motor acts, but they
may play a role in higher cognitive processes (Gallese and Metzinger, 2003). Mirror neurons are
indistinguishable from their neighboring neurons in terms of their motor properties. Both
canonical and mirror neurons generate an internal representation of the action or action
observation. The interaction of these neurons might provide the basis for how we understand
what another person is doing.
The mirror neuron system could be called a motor resonance system (Gallese &
Metzinger, 2003). In humans mirror neurons have been found to possess qualities not seen in
monkeys (Gallese et al. 1996). In humans, intransitive actions, not just goal directed actions,
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determined motor resonance. In addition, human mirror neurons can be activated during action
imitation and imagination. It is not known if monkeys posses this ability.
Imitation
Imitation is understood to be an automatic replication. It can occur with or without
understanding of the meaning (goal) of an action. Many mechanisms have been postulated that
contribute to imitation; instrumental learning, associative learning and more complex cognitive
processes.
Copying of actions is assumed to be based on simple neural mechanisms.
Infants are known to be excellent imitators of the facial expressions of those around them (Chen,
Striano & Rakoczy, 2004). A large array of behaviors including hand gestures, eye blinking and
head movements are mimicked by infants as young as a few hours old. It appears as though the
brain possesses the ability to imitate from birth. Whether these behaviors by infants are goal
directed and voluntary is still being debated. Further research to determine whether infant
monkeys have a mirror neuron system would provide valuable information on the role of
imitation in infant learning.
Imitation is a mechanism by which humans learn and become social, empathic beings.
Imitation by infants draws them in to relationships where they learn that others are like them and
it draws others to them. It is part of the bonding of an infant to a care giver and vice versa.
Imitation is a way of learning social behavior. This social behavior is based on understanding the
intentions of others. Mirror neurons could provide the mechanism for doing this.
To further understand mirror neuron’s role in action understanding, brain functions were
measured using fMRI. Subjects were measured during three observation and execution tasks and
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three observation conditions (Iacoboni, Woods, Brass, Bekkering, Mazziotta, Rizzolatti, 1999).
Iacoboni and his colleagues proposed that cortical areas that have motor properties should
become more active when the action to be executed is elicited by the observation of that action
(imitation).
Results of the experiment showed the imitation task produced reliably larger signal
intensity when compared with the two observation execution tasks (Iacoboni, et al. 1999).
Participants knew the task was to move a finger or refrain from moving their finger. It was
important that the participants knew this information because the investigators were interested in
the mental imagery of moving the finger or not moving it. When the participants observed finger
movement the scan showed intensified activity in area 44 (Broca’s region) and the right parietal
cortex (PE/PC). Broca’s region is considered to be the equivalent of the F5 region in monkeys
and is thought to posses similar characteristics (Gallese et. al. 2003). Broca’s region has been
demonstrated to be one of the few regions where finger movements are represented in humans.
The ability to make gestures and interpret them comes from this region. The motor region for
speech is also located in Broca’s region. This region is known to be crucial to language
acquisition (Gallese et al. 2003). This finding is important because it provides a basis for many
who believe that there is a motor basis to speech.
Language
Although language is associated with sound, speech might be better understood as a
motor activity. Many researchers are investigating the relationship between the areas of the brain
that allow controlled hand movement (gestures) and movement of the mouth (articulation).
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Evolutionarily, associations between gestures, mouth movements and sounds might be the
mechanisms by which speech evolved. When an individual began to make a noise, “mm yumm,
mm” while eating, others recognized they were eating and enjoying it. Soon the sound became
an abstraction for this motor act and was observed by another. Mirror neurons in Broca’s region
might provide the mechanism by which creation, observation and understanding of another’s
vocalizations became possible (Gallese et al. 2003).
Language requires a sequencing of sounds, a motor act of imitation, and an abstraction of
goals. These capabilities are possible through the motor cortex and mirror neurons. Audiovisual
mirror neurons are of particular importance in interpreting input and output of sound (Keysers et
al. 2003).
Abstraction of gesture or vocalization (representation of meaning by a sound or a gesture)
is not readily seen in animals. However, African Diana monkeys are known to vary their calls to
indicate if a leopard or an eagle is approaching (Prum & Brush, 2003). Whale songs are unique
to their social grouping. Parrots and dolphins can mimic or imitate. Imitation is part of how
babies learn to speak. Human’s ability to use syntax is unique in all species. It is possible
because of the evolved anatomical ability found in the larynx not seen in other animals. Deaf
individuals have been able to readily adapt to a manual mode of communication perhaps
indicating the gestural origins of speech. Speech might have become evolutionarily preferred to
manual gestures when man acquired the ability to use tools and needed to hold tools in their
hands (Jarvelainen, Schurmann & Hari, 2004; Ferrari, Gallese, Rizzolatti & Fogassi, 2005). As
previously discussed, a tool moving toward an object does not activate mirror neurons thereby
differentiating manual gestures leading to speech and tool use (Jarvelainene, Schurman & Hari
2004).
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Predicting the Future
Context
What is the mechanism by which humans became symbolic thinkers that made the
abstraction of speech possible? To investigate whether mirror neurons play a role in action
recognition and abstraction Iacoboni et al. (2005) investigated the neural and functional
mechanism by which intention and understanding might occur. The investigators measured
mirror neurons using functional magnetic resonance imaging (MRI) during observation of
context related actions. The context in which an action is performed might provide clues as to the
intention of the action. Different meanings might be attached to the same action done in two
different contexts. If the mirror neuron system codes the observed action and the goal, then there
will be no difference observed by the changing of context.
To understand the relationship of context to mirror neurons, three types of movie clips
were presented to subjects containing “context,” “action” and “intention”. By subtracting fMRI
activation patterns from stimuli, brain regions that are activated in response to stimuli are then
apparent. Areas that are involved both action and intention become evident in the fMRI
“intention minus context”. “Intention minus action” revealed areas involved in recognizing
context and intention. Iacoboni and his colleagues compared the results and found a set of
neurons of the inferior frontal cortex that they believe codes the “why” of an action. The area is
in the right frontal cortex. This area is known to be involved in the mirror neuron system. It was
activated only by scenes in which intention could be inferred by the participants. These mirror
neurons code some aspect of future actions. This would indicate that the mirror neuron system
can understand context and this assists in the understanding of another’s behavior and in
predicting future behavior.
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Intention
These neural mechanisms might be part of the way human’s code and use abstract
thought. This abstract coding seen in mirror neurons is similar to how we use verbs, action
words. The meaning of the verb does not change depending on the context it is used in. Whether
we say, “I dropped my keys” or “I dropped a dish,” the meaning of the verb dropped remains the
same. In addition the location of these neurons corresponds to the Broca’s area in humans, a
speech and language center.
Language requires the capacity to bring abstract representation together with auditory
output similar to what audio-visual neurons do with actions. Language provides a great
evolutionary advantage. It allows individuals to communicate mentalistic concepts. The ability to
store symbols externally is important to all we do today.
The mirror neuron system mediates action to understanding. When an individual sees an
action by another individual the premotor cortex is activated. This happens automatically and
seems to correspond to a goal that which is generated during active or observed action. The
outcome of this action is then known to the individual. The visual information received is then
transformed into knowledge, an understanding of the action.
The mirror neuron system seems to be a categorization system of motor acts. It is known
that neurons in the motor cortex possess great specificity. Specific neurons in this system
activate for specific actions. If mirror neurons operate in this manner it seems to indicate a
recognition process. There is an understanding of the goal of the action. Observation of a grasp
activates certain neurons as if to say “that’s a grasp”. This seems to indicate that the mirror
neurons provide a way to recognize an action rather then an intention coding system, “I am going
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to grasp”. The system appears to code actions by their goal. This is seen in both observation of
action, execution of an action and imagining the action.
Part of recognizing an action is predicting what the other person intends to do. When I
see a person reach for a piece of fruit I see a hand grasping a fruit. But what I also recognized is
that a person is grasping the fruit. There is a link to an object that implies intention not merely a
motor act. Why are they grasping the fruit? Is the individual hungry? Is it a gift for someone else
to eat? Will they make a pie with it? It helps predict the future.
From an early age, children begin naming the world. They look for the stories that will
connect the parts. The child will imagine, play, pretend and anticipate, with stories what they
will do when they grow up. The actions of others will be tracked and noted.
Mirror neuron systems are important in developing children. Developing children must
learn to recognize opportunities for action and what the body can do. Human children have a
prolonged period of dependency as compared to other species, this leads to conditions that allow
for complex social learning. When this doesn’t occur development is impacted. Success depends
on the attention of the caregiver to the child and attention of the child to the caregiver. This cycle
of turn taking imitation and the small group of almost innate gesturing is important is important
in establishing this turn taking. Falk (2004) calls it “motherese”. Social reciprocity may be an
essential precursor to complex imitation. Motherese may be important by setting up the basic
tenant of “I am like the other” (Zurkow-Goldring, 2005). Infant mother imitation may provide
the basis for more complex imitation required as the child matures.
Mirror neurons provide the ability to discern context, action and intention in others.
Mirror neurons have this ability by coding current actions and interpreting them by looking for
logically related actions or schema.
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Action Schema
As was previously discussed, imitation is understood to be an automatic replication. It
can occur with or without understanding of the meaning (goal) of an action. Action
understanding is a conscious understanding. Imitation and understanding are two different and
related mechanisms of the mirror system. They are both involved in man’s ability to act. Having
a future goal state of going to work does not require conscious knowledge of every motor step
required to arrive at work. They are packaged into action schema or motor vocabulary that has a
chronological order.
These action schemas might be organized according to what is of value to the individual
(Gallese & Metzinger, 2003). Having this reward system allows the individual to evaluate if they
are doing the right thing. This organization allows complex interactions to be grouped
systematically and expanded upon. The organization of these interactions can be automatic or
deliberate. This high level of integration allows for flexibility and expresses an individual’s logic
of life. Adler’s private logic is comparable to these physiologically findings (Adler, 1956).
This ability to predict the future and organize goal states has an important evolutionary
component. An individual is able to size up a situation. “What’s this person up to? Are they
friend or foe?” By being able predict actions and their meaning humans have enhanced their
ability to survive.
Sensation and Emotions
The findings discussed up until this point do not exclude other areas from being involved
in biological movement and action understanding. Connections from Broca’s region to PF and
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superior temporal sulcus (STS) provide a circuit that attaches to the amygdale and the orbital
frontal cortex. Wicker and his colleagues located a connection in the circuitry by which we
observe disgust in others and feel it ourselves (Wicker, Keysers, Pially & Yoyet, 2003). During
fMRI participants smelled a disgusting odor and then were measured when they observed a video
of another showing the facial expression of disgust. Observation and feeling of disgust was
shown to activate motor areas in the anterior insula and to a smaller degree the anterior cingulate
cortex. This study provides a unifying circuitry which link action representation with an
emotional understanding of others. Interestingly, the orbital frontal cortex is an area involved in
taste. This might be the mechanism by which human’s have a moral sense and know when
someone is acting in “bad taste”.
Emotions make available a way to acquire knowledge about a situation. Emotions are one
of the earliest ways that man used to acquire this knowledge. Activity of the affective neural
systems and sensory motor system result in a simple and automatic way to obtain knowledge of a
situation and reorganize the information based on the acquired knowledge. The speed or
automatic quality of these responses is important. The probability of survival is increased if
behaviors that are likely to promote survival are highly automated.
There is a pairing between affect recognition and sensorimotor integration. One hundred
brain-damaged patients were reviewed by Adolphs et al. (Adolphs, Damasio, Tranel, Cooper,
2000).The results of this review showed that those who suffered the most damage to the
sensorimotor cortex and amygdale were also the group that performed the worst on when
requested to name and rate a facial expression.
Evidence suggests that the same neural mechanisms that are activated during sensations
and emotions are activated when these sensations and emotions are seen in others (Adolphs, et
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24
al., 2000). Results of experiments seem to indicate that when the ability to experience an emotion
or sensation are lost (as in the case of stroke), one also loses the ability to detect these sensations
in others. A patient who suffered a stroke that damaged cortical and sub cortical structures
(including the insula and putamen) became selectively impaired in his ability to distinguish
disgust. This impairment occurred not just in one modality but in many including non-verbal
emotional sounds, facial expressions and emotional prosody (Calder, 2000).
The mirror system provides an understanding of movement, but we also use neuronal
activity to interpret intention, sensation and emotions of those around us. These understandings
are providing fuel to further our understanding of empathy.
Sensation and Perception
The superior temporal sulcus (STS) region contains neurons that have visual properties.
One hypothesis of understanding an action is visual; our senses provide us information from
which we interpret our surroundings. This visual hypothesis contrasts with direct matching
hypothesis which states that understanding cannot occur without motor involvement (Gallese
2003). A representation is mapped directly onto our motor representation of the same act. The
description of the motor event is sufficient for action understanding. In this argument the
properties of F5 mirror neurons are sufficient; a visual description of action is not needed.
Complex imitation has a perceptual side. Observation of a complex action requires
recognizing another’s action while comparing it against stored variants of that action. Perception
adds information by which the observed action can be compared. Complex imitation depends on
perceptual comparison. Experience further tunes the understanding. This can be seen when
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children are learning to speak and they repeat words that are spoken to them while more
experienced adults understand the complexities of what is being said.
The mirror system mechanism depends on the part of the motor system that resonates.
The PF and premotor areas become active when an action requires understanding (Iacoboni, et
al. 2005). When an action does not require understanding other regions are active. When an
action requires understanding and imitation then the activity would be shared between those
regions. If the action has emotional content then other areas might be called to action.
The amygdale is another structure known to be important in emotional behaviors and the
recognition of facial expression in others. Patients with Mobeius Syndrome are congenitally
incapable of moving their facial muscles (Cattaneo, Chierici, Bianchi, Sesanna & Pavesi, 2006).
They also have difficulty understanding facial expressions and the emotions they represent in
others. Impairment in motor movement and the inability to understand others are seen in this
disorder. This might indicate impairment of the mirror system.
The role of the limbic system was investigated to further understand the relationship
between action and emotion (Carr, Iacoboni, Dubeau, Mazziotta, & Lenzi, 2003). In this
experiment, an imitative task was used. Randomly presented facial expressions were presented to
eleven participants. fMRI was used to monitor and record cortical activity. Three stimulus tests
were randomly assigned. The first stimulus task contained full faces and the other two contained
only eyes or mouths from the full face images. The participants were instructed to observe and
imitate the images.
Results revealed that there was no difference in activation between the three sets of
images. Observation of emotional expression caused strong activation over the motor areas.
Activity in the amygdala increased while subjects imitated behaviors, when compared to only
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observation. Representation of the actions associated with the emotions is necessary in empathy.
The insula provides the mechanism for communication between action representation networks
and limbic areas that makes empathic resonance possible.
Understanding Others
Agency, Intention
Social behavior is based on understanding the intentions of others. Mirror neurons could
provide a mechanism for doing this. To be able to empathize with another, one must be able to
understand their feelings and interpret them through yourself while one maintains their own
agency. An interesting deficit in agency is seen in anarchic hand syndrome (Schwoebel, Boronat
& Coslett, 2002). Patients with this syndrome are aware, well aware of carrying out acts with
their non-dominant hand at the same time experience the movement with no conscious volition.
Functionally and subjectively the arm is out of control. Interestingly, the patient attributes the
action of the hand to having a mind of its own. There is no sense of volition or goal
representation, no sense of agency. The anterior insula is made up of slow-conducting
unmyelinated fibers. These fibers respond to light caress like touch. It is thought that they are
important for emotional and affiliative behavior between individuals (Ollausson, Lamarre,
Backlund, Morin, Wallin, Starck, Ekhom, Strigo, Worsely, Vallbo & Bushnell, 2002). It is also
important in the understanding of agency, an ownership of actions.
But how does an individual tell whose body is moving during action imitation. How is
the sense of agency maintained during imitation? The parietal operculum is a sensory area that
may assist with that. Activation of the right hemisphere during imitation was discovered. Lesions
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to the area are known to be associated with body schema disorders (Schwoebel, Boronat &
Coslett, 2002). The increase activation over the right hemisphere parietal cortex could be the
means for knowing whose body is doing the action. This might be the neural mechanism used by
the mirror neuron allow for empathy while maintaing a sense of who you are.
Humans may have numerous mirror systems that have nothing to do with movement
planning (Keysers, Wicker, Gazzola, Anton,Fogassi & Gallese, 2004). Researchers studied the
response of subjects while they underwent fMRI and had their bare leg gently stroked by a rod or
a brush. The participants were also measured during observation of a video of another being
stroked on the bare leg and during observation of a video of the same movement where the brush
or rod never touched the bare leg. During the observation of the video where the brush did not
actually touch the skin, this area was not activated. The actual touch and the observation of the
actual touch activated similar activity in the secondary somatosensory cortex. This area is known
to be involved in the sensation of touch. The brain did translate an observation of touch into an
understanding of the sensation. This might be the mechanism by which we flinch at the sight of
another’s blood or when they hurt themselves.
Researchers studied women’s response when they received a painful shock or saw their
romantic partner receive the shock (Singer, Seymour, O'Doherty, Kaube, Dolan & Frith, 2004).
Bilateral anterior insula (A1), rostral anterior cingulated cortex (ACC), brainstem and
cererbellum were activated on receiving pain and observation of their partner receiving pain.
These areas are known to process the emotional content of pain. The somatosensory cortex
known to register the sensation and location of pain remained quiet during observation of the
shock. Most interestingly, those women who showed the greatest brain responses in A1 and ACC
also scored highest on two empathy scales, the Balanced Emotional Empathy Scale and the
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Empathic Concern Scale. The mirror system, acting with the motor system, may provide the
empathic resonance that is required to understand another’s pain. The automatic nature of this
reaction may provide the mechanism necessary for the social learning of reactions to pain.
A relationship between the motor system and an individual’s ability to understand the
actions of others is seen in children. Children with impaired motor coordination perform poorly
on scales that measure their ability to recognize static and changing facial expressions
(Cummins, Piek & Dyck, 2005). A child’s motor ability was a significant predictor of social
behavior even when visual-spatial processing, age, sex and emotion recognition was controlled
for.
The studies are consistent with findings of abnormalities of the mirror system in people
with autism and other disorders that impair the ability to empathize. People with impaired
emotional experience or learning also experience difficulty in interpreting others. There might be
a key time where mirror neurons learn their job. Abnormalities in certain behaviors that occur
together such as imitation, language and the ability to interpret the intentions of others might be
due to an impaired mirror system.
Social Interest and Empathy
Self and other relationships are the basis for the development of sophisticated
interpersonal relations. It is through this identification with others that we are able to articulate
complex social interactions.
Empathy is an experience. In this experience one must be able to understand and
distinguish many states of mind. These states of mind include desires, beliefs and the emotions
of others. Empathic resonance requires understanding the emotions of another without confusing
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them with your own. It is ambiguous whether the process of empathy requires imagining being
the other person or just being in the situation.
There is probably more to empathy then the mirror system. This mechanism may only
provide a gut level that lies beneath our understanding of empathy. If you see someone joyfully
hurting another, hopefully you don’t feel joy as well.
Embodied Simulation
In order to penetrate the world of another, we predict and model their emotions and
experience. Embodied simulation describes this joining. This modeling is what enables a person
to resonate with another’s experience. Action observation becomes action simulation.
Embodied simulation establishes a direct link between agent and observer. It is an
automatic, direct and unconscious process. This link between agent and observer is direct and
neutral, waiting to be interpreted.
The same logic of self functioning is applied when observing another’s actions. Gallese
describes this “shared Manifold” as operating at three different levels: (i) a phenomenological
level; (ii) a functional level; and (iii) a sub-personal level (Gallese, 2003).
In this shared manifold, at the phenomenological level we acquire perceptual
information. Then, at the functional level, one would experience an action sequence. The subpersonal level is characterized by matching a series of neural states with assistance through the
mirror neuron system. Activation of this system is tied to changes in bodily states.
Mirror neurons allow for a conceptual shared space between self and others. These are
the spaces that allow us to understand, appreciate and experience others. It is important to
understand that we do not experience others as we experience ourselves. The “shared Manifold”
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as Gallese describes it only enables the basics of mutual intelligibility.
By means of simulation we automatically link to others. From birth forward we share our
semantic space. When we perceive others and observe their array of expression we share an
intersubjective space. The operation of the mirror neuron system is automatic and unconscious. It
could be the mechanism by which the effects of hypnosis and EMDR are attained. It could also
be the conceptualization of Adler’s ideas of fictional finalism, lifestyle and early recollections.
This simulation is also described by Brandt and Stark who demonstrated when research
participants listen to sentences containing right and left , they moved their eyes in primarily a
horizontal orientation ( as cited in Baranek, Foster & Berkson, 1997). Further, the participants
moved their eyes vertically when listening to above or below. Spivey describes research
participants looked up when listening to a vignette about a skyscraper and down when listening
to a vignette describing the bottom of a canyon (Spivey & Spirn, 2000). Humans tend to
accompany their understanding of sentences or imaginative actions with bodily reactions that
simulate the real experience.
Embodied simulation is not merely motor responses although it is driven in part by them.
It is a more general and basic brain function. It applied to actions and emotions where the
sensory motor system is involved but also gains information from sensations, like vision or touch
or hearing. It is embodied because of the body model or map realized through the sensory motor
system. This helps explain why early childhood learning is so well incorporated into our adult
style.
Intentional attunement requires the collapsing of other’s intentions into the observers.
Gallese understands this to be the process by which a person experiences a curious sense of
familiarity in encounters with other persons. Two bodies that function by the same rules whereby
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objectual other becomes another self. Mind reading can be seen occurring in this manner and is
seen in the manifestation of empathy (Singer, 2006).
Simulations
Human cognition may rest on sensoriomotor simulations (Gallese, 2003). These
sensoriomotor simulations represent and manipulate data. There are many types of simulations
seen in humans. On line simulation runs parallel to the event being simulated, in real time and is
used for tracking, predicting and interacting with the external world. Off line simulation recruits
the same sensoriomotor brain resources but disconnects them from input and output. Secondorder simulation exploits human’s capacity for seeing analogies, so as to represent
nonsensorymotor information (abstract concepts). In the service of representing something else,
a sensorimotor event is recreated (like off line simulation). For example, mentally picturing a
sundial to represent time. Formalisms occur when second order simulations become automatic;
the analogical link no longer needs to be activated in order to perform the computation. This is
how symbols are manipulated according to rules without mentally grounding their meaning.
Formalisms can be unpacked when needed in order to retrieve their conceptual content. This
might be the realm of the unconscious. Second order (automated) formalisms might be what has
been described as the unconscious mind. All simulations have a role in cognition.
Experiments show that neither gaze, nor objects, nor hands are automatically simulated in
the observer. It is the task context that dominates the relation between the modeled action and the
action to be executed in the observer (Ultima et al., 2001). Simulation is a form of experiential
understanding that is achieved by modeling behaviors as intentional experiences. Side by side
with sensory description, observed sensory information is evoked in the observer. This modeling
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mechanism is embodied simulation (Gallese, 2003). Coupled with descriptive sensory
information of the observation, internal representations of the actions, emotions and sensations
are induced in the observer. Mirror neurons and other multimodal mirroring areas in the brain are
the possible mechanism.
Perception and Prediction
Humans hold implicit certainties about each other. Unavoidably, people interpret each
others actions. These interpretations are made in terms of cloaked mental states like desires or
beliefs which must be understood by others for success in social situations. Successful human
development depends on these abilities to be learned and understood.
Actions can be viewed as goal-directed or referential (Greeley & Csiro, 2003). When an
action is goal directed, intentions, beliefs and desires are assigned to the observed action or
agents. A specific stance is assigned the action. A goal directed action is understood through its
end state or goal. Young infants construe the goal of many actions based on this teleological
understanding. Many researchers believe the infants rely on detection of motion cues to
understand the goal of an action. One way this is accomplished is through self motion.
Referential understanding of actions allows assignment of an attentional state, referential intents,
and communicative messages. Infants are known to do referential responses to actions. This can
be seen in young infant’s preference for faces with eye contact and objects that react to them
contingently. Later, these are the responses through which gaze and pointing are elaborated.
Referential actions are about the state of the world; interpreting an action in this way connects
the world to an agent. To understand language, referential understanding of actions is required
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Infants at about 12 months of age tend to follow the gaze of other persons. This is called
joint attention. If you make eye contact with a twelve month old and then look at something else,
the infant will follow your gaze to the object of your attention (Gregely, Nadasdy, Csibra & Biro,
1995). As the child matures, the accuracy of this ability increases.
Researchers hypothesize that referential understanding requires that actions are supported
by two cues (Gergely et al., 2003). The first cue indicates a communicative situation. The second
cue is perceptual (spatial orientation). So a child learns to look for the communicative situation
and then must follow it (orient toward it). Although a child orients toward a gaze, it is not until
they are 16 months old or older do they start making sense out of it. The MNS system is an
important part of how one makes sense of experiences.
Referential learning does not specify the intent of an action. Many believe the purpose of
referential learning is to assist in language acquisition. It is assumed that referential gaze
between an infant and agent help establish the link between words and objects.
Meltzoff (1995) found that 18 month old toddlers were able to successfully complete a
target act failed by the experimenters. In the experiment the toddler observes the experimenter
fake his failing to pull a dumbbell apart. The toddler, when handed the dumbbell, pulls it apart,
some with an enthusiastic smile. The toddler saw the unrealized target act, mentally simulated
the completed target act (through the mirror neurons) and then was then able to successfully
complete the target act.
A strategy young children take when confronting a novel situation is known as social
referencing (Moses, 2001). A child will check with their parents or other adults when consulting
a novel situation and modulate their behavior accordingly. In the same novel situation the child
may behave one way if the mother looks worried and another way if the mother is smiling.
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Successful perception requires the capacity to predict upcoming sensory events.
Successful action necessitates envisioning the likely consequences of the action. These two types
of prediction describe unconscious and automatic states that are described as a simulation
process.
Understanding others intentions isn’t purely a simulation response. Social cognitive skills
are complex. The same actions performed in different context can lead to different
interpretations. Social stimuli provide information based on previous experience, context and
cognitive elaboration. But Gallese feels that these two ideas are not exclusive. As he explains it,
embodied simulation is experience based while the second mechanism is a cognitive description
of an external state of affairs. When both mechanisms are not functioning properly, as in autism,
this mind reading ability is pale and shallow.
Balancing Sensory Information
A disassociation in mind reading ability is seen in children with autism, a developmental
disorder where the person has significant difficulty attributing mental states to others (Happen,
2003). The literature on ASD has been considered sufficient to assume the impact of sensory
differences with those with ASD. However, the mirror neuron is at a more basic level and may
provide a better explanation of the differences seen in autism spectrum disorder. Perhaps in these
individuals, the Mirror Neuron system is not functioning adequately or at all.
The literature describing the differences among persons with autism spectrum disorder
(ASD) is vast. As early as 1943, Kanner described sensation seeking behavior and sensation
differences in children with ASD. A variety of unusual behaviors are associated with ASD and
that are known to be associated with difficulties in registering and processing sensory
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information.
The sensory differences in ASD are well documented. Adreien et. al. retrospectively
analyzed video of children later diagnosed to have ASD (Adrien, Lenoir, Martineau, Perrot,
Haneury & Larmande, 1993). His study suggested that there was a qualitative difference in the
sensory functioning of children who were later diagnosed with ASD when compared with
normal children. Videos analysis of infants between the ages of 9 and 12 months show that
children with ASD respond slower or not at all to attention-getting behaviors (being called by
name) when compared to both developmentally delayed and typically developing infants. In
addition children later diagnosed with ASD showed poor visual orientation, poor attention to
sensory stimuli and social touch aversion (Baranek, 1999). Unusual sensory responses were
demonstrated in 42%- 80% of individuals with ASD. Many of the sensory differences seen in
autism may be from a malfunctioning MNS. The sensory differences may be simply the
manifestation of this.
Unusual eye gaze is commonly seen in ASD. Some parents report that their infants and
young children have an empty gaze (Laundry & Bryson, 2004) and seem to look through objects
and people. Individuals with ASD tend to focus on physical features of the environment (e.g. a
lamp). They have a tendency to use peripheral vision and to look at objects and fail to attend to
social features (faces, eyes) although most individuals with ASD have strengths in visual
processing (Brian, Tipper, Weaver & Bryson, 2003). Those with ASD tend to focus on the
details of the environment (Baylis & Tipper, 2005) which are irrelevant in terms of interactive
function. The amount of eye contact by these individuals was shown to be influenced by
structure and task demands (Volkmar, Carter, Grossman & Kiln, 1997).
Hearing is also altered in ASD. Part of hearing consists of the ability to screen out
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irrelevant stimuli and orient themselves in the environment to relevant sound. Individuals with
ASD show auditory responses that differ from others as early as the first year of life (Hayes &
Gordon, 1977). They also show a preference for nonhuman noises over human voices (Kiln,
1992).
The three variables that predict the development of spoken language in children with and
without disabilities are: motor imitation, joint attention (JA) and object play (Mundy, Sigman &
Kasari, 1990). The development of symbolic play and language relies on a child’s ability to use
one object to represent another (Piaget, 1962). Motor imitation requires a child to attend to
another person and form a mental representation of that person’s action so it can be imitated. JA
occurs when a child is able to recognize what another person is attending to. JA is also seen
when a child draws another’s attention toward something that interests them.
Crucial information is provided by the senses that are necessary to function. The
integration of these responses, leads to goal directed behavior. Some Sensory information will
lead to arousal which would then generate a response. Other sensory information leads to arousal
which reduces a response.
For an individual to do an activity, a certain level of arousal must be maintained.
Achieving the best level of arousal is key to optimum functioning. Sensory information is
abundant and the demands on the CNS to register integrate and modulate relevant information
depends on the assistance of the mirror neuron system. For some this ideal level is difficult to
achieve. Part of the over arousal seen in ASD may stem from an overabundance of input from
the senses.
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Sensation and Movement; Generating New Behavior
Even in basic movement innovation is present. Studies have shown that a movement that
is goal directed is not necessarily reproduced exactly as observed. When moving toward a goal,
corrections are made and the movement may not exactly resemble another individual’s
movement toward the same goal. This explains why children are able to imitate despite the
radical difference in size.
Individuals with ASD are unable to translate movements that require mirroring. When
you wave at a person with ASD they may return the wave with their palm facing themselves
(Laundry & Bryson, 2004) . They are unable to interpret the goal directed movement and rely
only on direct perception. Ballistic movements are the exception to goal directed movement and
can be viewed as movement for movement’s sake. Striking a nail is an example of a ballistic
movement. These ballistic movements are usually quick and require no innovation although they
do require planning. When lifting a bag of groceries and placing it on the counter you balance
your path and make changes based on the weight of the package, height of the counter etc.
Sensory System
The sensory systems (SS) include olfactory, gustatory, tactile, vestibular, proprioceptive,
visual and auditory. The SS brings information to the central nervous system (CNS). This
information is processed in conscious and unconscious ways and serves to modulate behavior
(Kandel, Schwartz & Jessell, 2000). Much information is received from the senses and a person
does not acknowledge all the sensory input they receive for some of it is unnecessary. To do so
could be overwhelming.
While the sensory system is an important way that people receive information about the
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environment, the mirror neuron system is how they “make sense” of it. The mirror neuron
system provides a vocabulary by which to organize sensation and perception. The MN system
can decide what is important to attend to. Without the MN system, the perceptual system
becomes the primary mode by which individuals interpret the environment. Uncensored it
provides an overabundance of stimuli.
So what happens to individuals whose mirror neuron system is not functioning
adequately or not at all? When sensory input cannot be organized into a useful hierarchy, it
becomes chaotic and overwhelming and compensated for by avoidance or stimulation
(distraction and calming strategies).
Central Nervous System
Sensory Information is provided through the central nervous system and configured so
that this sensory data becomes linked to responses and experiences. Small mammalian brains
cope with this by using automatic responses. This type of automatic process capacity limits the
flexibility with which events can be interpreted.
When the bonds between sensation and action are instinctual and automatic they are
resistant to change. Negative consequences of this resistance can result. For instance, a turkey
hen’s protective maternal instinct is to attack any moving object that does not utter the
characteristic peep of her chicks, will peck her own chick to death if she is made deaf (Schleidt
and Schleidt, 1960 as cited in Mesulam, 1998) Humans with more advanced CNS systems are
less vulnerable to the appearance of inflexible patterns.
With the exception of autonomic responses (as seen in the brainstem and spinal reflexes)
identical sensory events can generate many different reactions depending on context. Humans
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are able to practice much biological freedom because of the loosening of stereo-typed stimulus
response connections. This bias pushes human’s toward the pursuit of novelty and flexibility
rather than sameness and stereotypy. Neural systems bridge the path from sensation to cognition
through “intermediary” and “integrative” processes. These processes, also known as cognition,
are memory, emotion, attention, language, thought and consciousness. These intermediary
processes activate different responses depending on the situational context, past experience,
present needs and considered consequences.
There are many different ways people use and register their sensory information.
Sensation seekers (Baranek, Foster & Berkson, 1997) actively seek out additional sensation.
They have a very high threshold of sensory registration. Many people with Pervasive
Developmental disorder (PDD), an autism spectrum disorder, have been found to be sensation
seekers (Kootz, Martinelli & Cohen, 1981). Other individuals, with a low threshold, register
more sensory information than is comfortable. When compared with those with more typical
thresholds, these individuals may avoid sensory input (Kinnealey, Olliver & Wilbarger, 1995).
It is known that individuals with autism look to the periphery of an object during a gaze, perhaps
this is a compensation to prevent of stimulation of perception that is un-modulated by the mirror
neuron system. Perhaps this is seen in the difficulty seen in integrating sensory information by
those with ASD. 40% of those with ASD have temporal lobe seizures; fovel vision feeds directly
into this probably damaged region (the amygdale). It might be upsetting to make direct eye
contact if your amygdale isn’t functioning properly.
Many adults who avoid sensation have reported coping strategies such as; avoiding the
stimuli, maintaining predictability, preparing mentally, talking themselves through the situation
and doing something to mitigate the discomfort associated with the stimuli. This is called
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simulation. Simulation is the process by which the mind uses its off line ability to model or
predict external physical or social events. The act of simulation requires singling out some target
event and actively fitting into place an internal evaluation or predictive consideration and then
projecting the result to the target user.
Applied Behavior Analysis can be seen as a simulation externally applied. These are the
very things that the mirror neuron system could assist with when operating as expected. Action
schema would provide experience so that sensations and experiences do not overwhelm, need to
processed individual or require being experienced anew. While repeated exposure to a stimuli is
known to mitigate response this is possible in two different ways. The more a person experiences
a stimulus the more they become desensitized to them. However these new sensory experiences
need to be stored and categorized so that they can organize in a hierarchical fashion and
habituated to. Habitation is a learned response diminishes the recognition of sensory stimuli
habituation (Dunn, 1997).
Children with fragile X and sensory modulation disorders are less able to habituate
sensory stimuli (McIntosh, Miller, Shyu & Hagerman, 1999). Most people tend to have lower
threshold to sensory stimuli when not psychologically or physiologically comfortable. So when a
person is hungry or tired they are more reactive to sounds, smells and movement. Information
from different sensory input is processed, organized and combined to produce adaptive motor
responses (Bundy & Murray, 2002). Sensory information, from eyes, ears, touch etc., combine to
provide information. The central nervous system can provide protective responses if a stimuli is
perceived to be threatening. A loud boom will cause a person to jump even when engrossed in
something else. Protective responses are automatic. However, when the sensory input is not
threatening then modulation of the response tales place. A carefully modulated process can be
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41
seen in the act of writing a check at a grocery store. Consider the following scenario. A person is
writing a check at the grocery market when all of a sudden a large crash is heard. Another person
in the store dropped a large jar. The person writing the check, startles and their handwriting is
interrupted and becomes illegible. A protective response will startle or interrupt their carefully
modulated response (handwriting).
Proprioception
The Proprioception is the neural system that allows us to know the position of our body
parts in reference to each other and to sense internal space (Damasio, 1999). It also allows us to
sense fatigue and discomfort. Proprioception allows one to sense direction and the velocity of
movement. It enables an individual to know how much effort is required to grasp and lift objects.
An example of Proprioception is when one lifts and carries a bag of objects, and it becomes
heavier as time goes by. There is a conscious sense of awareness that proprioception is occurring
in this example. You realize your muscles are getting fatigued; the bag is not actually getting
heavier.
The proprioceptive system works in conjunction with the vestibular system to accomplish
movements such as walking, sitting, holding, chewing. The proprioceptive system functions
without looking. Most activities that involve the hands require proprioception. The temporal
mandibular joint (TMJ) is the most sensitive of proprioceptors. Chewing and biting and walking
on tiptoe give increased proprioceptive feedback.
Fine and gross motor difficulties are seen in individuals with proprioceptive difficulties.
These individuals may experience difficulty getting into and out of chairs, climbing stairs or
navigating climbing equipment. In addition they might have poor posture, bump into things and
Action and Understanding
42
get confused when looking into a mirror. Through experience an individual’s proprioceptive
system is refined. This allows for development of a body scheme and the ability to plan
movement (Lane, 2002).
Mirror Neuron System and Organization of Perception
The mirror neuron system provides sensory organization; action understanding,
prediction of action outcomes and perceptual organization. This action system provides functions
that the perceptual system cannot provide. Learning to perform new movements provides a
sequential frame that supports more accurate categorization of movement.
All sensory systems, except the proprioceptive system, have the ability to discriminate. If
you are served a meal that smells bad a protective response would be to refuse the meal. But
sometimes social convention causes us to override the protective response. The person’s
response might be disgust (to gag or make a face). However, if it is important not to insult the
host the individual might accept the meal by consciously overriding the protective response. One
can also ignore pain and fatigue when necessary. This is the work of mirror neurons.
Those individuals with ASD have difficulty with social conventions. Deficits in the
sensory system may not be the cause. Deficits in the sensory system may occur because this
system becomes overwhelmed by an inability to handle the input through the mirror neuron
system. As a result they are unduly influenced by protective responses and these responses are
not able to be modified by action schema or social convention. The integration of the sensory
input through mirror neuron system is necessary.
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43
Culture
To survive we must understand the actions of others. With action understanding social
organization and empathy becomes possible. Our ability to learn through imitation is the
foundation of human culture.
Understanding how the brain interprets social interactions has significant implications.
Mirror neuron theory challenges the view that execution of actions and the interpretation of the
cause of the action result from different mechanisms. The mirror neuron system appears to
combine events that include fundamental behaviors (response facilitation) to higher order
cognitions such as imitation, understanding and mind reading.
Animals have cognitive abilities that permit them to excel but only in their unique
habitats. Humans are able to solve novel problems. According to the Machiavellian intelligence
hypothesis, cultivating the most profitable relationships and being ale to read social relationships
allows for the most success in these relationships (Collison, 1998). Successful social
relationships allow the most intelligent individuals (who are making self protective choices) to
survive and pass on their genes to the generations that follow. The ability to figure out how to
proficiently gain nutrition and sex partners allows for genes to be passed on. The ability to be
clever and adapt to changing environmental influences would be rewarded as well.
Social learning is of prime importance as well. A child relies on their parents for many
things, guidance being one of them. Social or cultural training is passed on through parent to
child. Intelligence becomes a cultural phenomenon that promotes intelligence.
Innovations are learned and taught. They are able to be spread in a population with close
social ties through direct observation. A long childhood makes more time for these social lessons
to be observed. Adults other than parents are important in this social education. The child can
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learn from others in the social group. This is important because sometimes parents are unable to
provide this information.
Populations that observe others in action have more learning opportunities and greater
repertoires. Animals exposed to the fewest educational experiences have the smallest number of
cultural variants. Proximity and tolerance that allows others who are less skilled to observe
promotes intelligence. Although most skills are learned thru parenting, individuals will only be
able to pass these skills on with other tolerant role models. This allows for varying levels of skill
within the population.
Learning skills does not come automatically. A more connected and tolerant social
network allows a greater number of such behaviors. Through this social learning, greater
intelligence evolves over time.
Slowly developing animals in tolerant societies can grow a culture through improvements
learned and passed to present and future generations. Innovative abilities will increase. This
thinking explains why animals in captivity in close proximity to human behavior make use of
tools and even make tools while those in the wild do not. Two individuals of the same species
can differ dramatically based on the environment in which they grew up.
Intelligence by culture is most likely in communities where the larger group adopts
innovations. Social learning and the propensity toward innovation work together. Human’s
innate ability when working together with cultural history improves performance. Culture helps
grow our brainpower.
Evolution exploited the opportunity, imitation, observation, embodied simulation,
empathy. “Expatation” is an evolutionary term used to describe characteristics that arise in one
context but later are exploited by another (Brum & Bush, 2003). Feathers were first seen in
Action and Understanding
45
animals as an insulator, only later were expatated when feathers on animals were adapted to
flight. Mirror neuron system can be seen in this way.
The ability to hold abstract ideas in the mind, not just perceptual categorization or
generalization allow humans to reflect, play and plan. Humans are not locked into literal and
immediate perception. Humans are able to juggle images, hypothesis and possibilities and enter
the realm of imagination and the figurative. On the darker side, humans are able to thoroughly
mistake their projections and fantasies for reality.
Cultural Transmission
Cultural transmission requires representing the intentions and actions of others. It does
not necessarily require understanding in the intellectual sense. Examples of this can be seen in
innovation and creativity as well as “group think”. This group think can be seen in the ease with
which humans share mental representations which is essential for the transmission of cultural
information from one person to another.
One way this is seen in the aspects of “catchy memes” and “migratory mannerisms”
(Morrison, 2006). Catchy memes are bits of cultural information that one can get easily caught
up with. They are seen in the tune that you can’t get out of your head or a popular catch phrase.
Migratory mannerisms are gestures or mannerisms that are picked up by others that did not
originate them. These stuck or picked up mannerisms seem resistant to the frontal and motor
areas ability to get rid of them. They are notoriously persistent and easy to translate to action.
Mannerisms are often enacted without any conscious awareness. These memes and mannerism
are reproduced easily and with great fidelity with little exposure and with out any of the usual
learning mechanism employed (Heyes, 1994).
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Catchy memes and migratory mannerisms lack goal direction. A meme is an
informational entity that exploits neural resources as a means of reproducing itself in the manner
of a parasite or virus (Blackmore, 1999; Morrison, 2006) uses the term memes as a useful word
for certain phenomena that are familiar to everyone.
Memory, patterns of excitation and inhibition and emotion are physiological factors that
can influence the likelihood that an observed action will be repeated by the observer. Memory
allows for the meme or mannerism to be remembered and later reproduced. Inhibition or
excitation provides for the motor systems susceptibility to it being reproduced. A catchy meme
that is a song will likely be coded automotorically and represented as a motor image (Jeannerod,
1997). It is interesting that mannerism that are more commonly seen in cultural expression are
gestures of the hands or face (rather than the knee or shoulder) which suggests mirror neuron
involvement. Social goals can then be transmitted somatosensorily, quickly with out any
conscious awareness.
Filmed research shows that interactions between two individuals are highly organized
and synchronized (Kenden, 1970). It is also interesting that the degree of synchrony is correlated
to the comparative status of the two. The closer in status the two individuals are the more closely
timed their movements are. This synchrony is a form of social signaling showing parity or
disparity. The MNS system might play a role in this. MNS probably allows for us to produce a
cascade of responses which couple with perception and action and action disposition and
memory.
Morrison expands on Damasio’s term “somatic markers” when she says that emotional
action dispositions are somatically marked (Damasio, 1998). This means that individuals are
very responsive to what and who is happening around them. Social movement is very structured
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and coordinated and is more likely to be reproduced when coupled with emotion. Objects and
symbols can become part of this, as seen in how someone wears a cap. It becomes a
representation and does not need intellectually understanding.
Innovation and Creativity
Cultural innovations ignited the potential for symbolic processes that existed all along
through cultural diffusion. Symbolic processes allow humans the ability to abstract elements of
our experience and represent them with discrete symbols. Elements can then be divided,
categorized and given separate names. In doing so, humans are essentially recreated their
experience, categorizing it, etc. in their minds. Humans string together these experiences; words
into language, melodies to song; movement to dance; rules to games. The ability to communicate
mentalistic concepts and the ability to pass these concepts to others is key to all that we do. What
gives these mentalistic, abstract symbols of experience their power is their origin in the motoric,
experiential and automatic encoding in the MNS.
We symbolize in our minds our experience of the world. Intuitive reasoning,
reactions to stimuli can then be recombined. An intuitive appreciation of the relationships
between objects and ideas is the basis of creativity that is seen uniquely in humans. Art, science
and culture becomes possible. Innovation can take place, we can ask “what if?”
Perceiving More Than Perception Allows
The classical view of perception as essentially an imaging system that allows
reasonable spatial and physical negotiations with the physical world has been extended and
transformed. The separateness of the internal senses - the somatosensory processes - from the
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visual and auditory distance receptors has been struck down. We now have a way to understand
the commonplace, what it really means when we say someone is "very perceptive," for example.
We surely never meant that their feature detectors for line, color and movement seemed to work
well. The common language always knew that something much deeper was going on. This helps
account for various phrases like “grasp a concept” or “I see what you mean”. The discovery that
premotor neurons - and through them neurons in the sensorimotor cortex and limbic system respond to integrated visual input carrying high order perceptual information has deepened and
broadened our understanding of "perception."
Common Sense
Common sense is what is shared with others. An example of this is the ability to speak
and the similarities of semantics, metaphors in all cultures. Part of this common sense is
consensuality, we all agree to these shared preconceptions and those which may be fiction
(Vailhinger, 1911/1965) 25. An example is our use of a red stop light. It is only a light, but
symbolically we have all agreed that it means to stop.
Adler saw individuals as creating their own private consensus or private logic by which
they operated. Private logic is described by Adler as a pre-conscious striving toward a fictional
goal that are not clearly understood by the individual. “Acting As if,” imagination, intention and
belief all require the ability of the individual to formulate an intention. Intention becomes action
potential. This intention is made up of motor, sensory and proprioceptive; conscious and
pre-coconscious components packaged and unpackaged by and for the individual. These motor
schemas provide information that guide us and direct our movement based on a
phenomenological interpretation of experience (Mosak, 1995). Packaged memes and action
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49
schema can be unpackaged and made more conscious during the counseling process.
Humans are creative interpreters that cannot help but respond to the societal pressures of
the environment. Adler was correct when he said “first thing we discover in psychic trends is that
movement is directed toward a goal” (Adler 1927/1957). The physiological mechanism of this
movement toward a goal is in part through the MN system.
Translation of Recent Research To Clinical Practice
Post- mortem studies of amygdala show increased cell density and abnormally small cells
as well as a smaller number of neurons in the amygdalas of 9 males with autism versus 10 males
with no disorders (Bauman & Kemper 1985, 1994). There are also abnormalities in the amygdala
volume (both increased volume and decreased volume) that are revealed in structural imaging
studies (Howard, Cowell, Boucher, Broks, Mayes, Farrant, Roberts 2000; Pierce et. Al. 2001). It
is known that autism is associated with over-development of the amygdala in children with
autism by around age 6 (Salmond, De Haan, Friston, Gadian, Varfgha-Khadem 2003).
Salmond’s research leads one to speculate that anxiety may cause the deaths of unusually large
numbers of cells. This might account for the lack of cells in later years. As previously discussed,
a large percentage, perhaps 40%, of people with autism have seizure disorders that focus in the
amygdala. This may result in destruction of the cells as well, but the sample examined by
Schumann & Amaral apparently did not have epilepsy.
In addition, the research of Salmond et. al. describes that the foveal visual fields – the
center of the gaze – feed directly into the amygdala (2003). The peripheral visual fields are
processed in parietal regions concerned with establishing the overall “scene” and locating the
observer in space. When accurate and rapid acquisition of depth occurs as one looks around, a
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spatial "map" is formed in the perceptual mind. This map will be relatively stable, also implicit one doesn't usually notice it. With it one knows accurately how far objects are from one's central
point of view and how far objects are from each other. Without accurate judgment of this spatial
dimension there is always an implicit, likely unnoticed uncertainty, a sensory confusion. This can
lead to anxiety borne of not knowing exactly where things are in relation to the Self. Lack of
accurate perception also leads to clumsiness in catching a ball, approaching a person and
negotiating space in general. Conversely, one cannot know where one is "coming from" if one is
not placed firmly in the center of this three-dimensional spatial map. Deficits in spatial
perception fuel ontological insecurity on a fundamental, precognitive - that is to say, perceptual level. A similar argument could be made for accurate time perception, the fourth essential
dimension of direct experience. These differences may account for the deficits seen in ASD.
While imitating facial expressions and observing emotional expression high functioning
children with autism and matched controls underwent fMRI. Both groups performed equally but
the autism group showed no MN activity in the inferior gyrus. The activity recorded was
inversely related to the severity of symptoms (Dapretto, Davies, Pfeifer, Scott, Sigman &
Bookheimer, 2005).
The amygdala is more sensitive to seizure induction that most other parts of brain. This is
why the Japanese cartoon triggered over 250 cases of epilepsy several years ago. A character’s
eyes began flashing brightly and repetitively. Children looking directly at the eyes – hence
channeling this rhythmic photic stimulation directly to their amygdala – frequently developed
seizures. The seizures, unfortunately, did not disappear after the exposure and had to be treated
medically. The amygdala has neurons that are sensitive to gaze direction and autistic children
have difficulty interpreting this information (Hirstein, Iverstein & Ramachandran, 2001).
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51
These recent observations help make sense of the tendency of people with autism to
avoid direct eye contact. Children with autism pay little attention to faces. When they do look at
faces it triggers a threat response (Amaral & Corbet 2003; Dawson, Web, Carver, Panagiotides
& Mcpartland, 2004). The findings also suggest that the typical behavior modification treatment
that includes shaping direct eye contact might have long term negative consequences, potentially
damaging the amygdala. Perhaps this should be approached more carefully using a systematic
desensitization paradigm, emphasizing acquisition of a substantial relaxation skill prior to
attempting to improve eye contact.
Development of generalized relaxation would be an appropriate primary and ongoing
target of any attempt to ameliorate the symptoms of autism. It is well known that excess systemic
arousal will cause deterioration of more complex thinking and behavior. This is well described
by decades of research into human performance (Prisniakova 2004; Dickman 2002) and is well
summarized by the Yerkes-Dodson Law of Performance (Broadhurst 1957).
The Yerkes-Dodson Law describes an inverted U-shaped function relating performance
on a given task to arousal level. If arousal is too low or too high, performance will suffer and in
fact cease. As arousal levels increase, simpler performances will be optimized. More complex
tasks will be performed at optimal levels at low levels of arousal. That is why there are no
roaring crowds, cheerleaders or brass bands at chess matches, whereas these and other methods
of inducing high levels of arousal are seen at football games.
An increased ratio excitation/ inhibition may be the cause of some forms of autism
(Rubenstein & Merzenich, 2003). If one is attempting to shape social behavior in autistic
children one would want to start by developing strong capacities for cue controlled relaxation.
This can be done most easily by methods that require minimal conscious, deliberate
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52
participation, such as neurofeedback or peripheral biofeedback (e.g., heart rate variability, electro
dermal response).
The MNS also provides valuable information that can inform Adlerian Therapy. The
Adlerian notion of encouragement can be seen as modeling useful memes. The transferring of
motor ideas such as a smile or nod of the head can be understood as encouragement and empathy
appearing in part from the MNS.
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53
References
Adler, A. (1956). The Individual Psychology of Alfred Adler (H.L. Ansbacher & R. R.
Ansbacher, Eds.). New York: Basic Books.
Adler, A. (1957). Understanding Human Nature (W. B. Wolfe, Trans.). Greenwich, CT: Premier
Books. (Original work published 1927).
Adler, A. (1964). Social interest: A Challenge to Mankind (J. Linton & R. Vaughan, Trans.).
New York: Capricorn. (Original work published 1933).
Adolphs, S.,Damasio, H., Tranel, D., Cooper, G., Damasio, A. (2000). A role for somatosensory
cortices in visual recognition of emotion as revealed by three dimensional lesion
mapping. Neuroscience. 20 (7), 2683-90.
Adrien, J. L., Lenoir, P., Martineau, J., Perrot, A., Haneury, L., Larmande, C. (1993). Blind
ratings of early symptoms of autism based upon family home movies. Journal of the
American Academy of Child and Adolescent Psychiatry, 33, 617-626.
Amaral. D. G., Corbet, B. (2003). The amygdala, autism and anxiety. Novaritis Foundation
Symposium. 251 (177-87), 281-107.
Baranek, G. T., (1999). Autism during infancy a retrospective video analysis of sensory motor
and social behaviors at 9-12 months of age. Journal of Autism and Developmental
Disorders 29, 213-234.
Baranek, G. T., Foster, L., Berkson G. (1997). Tactile defensiveness and stereotyped behaviors.
American Journal of Occupational Therapy. 51, 91-95.
Action and Understanding
54
Bayliss, A., Tipper, S., (2005). Gaze and arrow cueing of attention reveals individual differences
along the autism spectrum as a function of target context. British Journal of Psychology,
96, 95-114.
Baunam, M., Kemper, T. (1985). Histoanatomic observations of the brain in early infantile
autism. Neurology, 35, 866-874.
Bauman, M., Kemper, T. (1994). Neuroanatomical Observations of the Brain in Autism (ed. M.
Bauman & T. L. Kemper), Baltimore, MD. Johns Hopkins University Press. 119-145.
Blackmore, S. (1999) The Meme Machine. Oxford: Oxford University Press.
Brian, J. A., Tipper, S., Weaver, B., Bryson, S. (2003). Inhibitory mechanisms in autism
spectrum disorders: typical selective inhibition of location versus facilitated perceptual
processing. Journal of Child Psychology and Psychiatry and Allied Disciplines, 44, 552556.
Brothers, L. (1995). Neurophysiology of the perception of intentions by primates. In M.
Gazzaniga (Eds.), The Cognitive Neurosciences. Cambridge, MA: MIT Press.
Broadhurst, P. L. (1957), Emotionality and the yerkes-dodson law. Journal of Experimental
Psychology. 54 (5), 345-52.
Carr, L., Iacoboni, M., Dubeau, M.-C., Mazziotta, J. C., & Lenzi, G. L. (2003). Neural
mechanisms of empathy in humans: a relay from neural systems for imitation to limbic
areas. Proceedings of the National Academy of Science, 100(9), 5497-5502.
Cattaneo, L., Chierici, E.,Bianchi, B., Sesenna, E.,Pavesi, G. (2006). The localization of facial
motor impairment in sporadic mobius syndrome. Neurology. 66(12), 1907 - 1912.
Chen, X., Striano, T., Rakoczy, H. (2004). Ausitory-oral matching behavior in newborns.
Developmental Science. 7(1), 42 - 47.
Action and Understanding
55
Collison, D. (1998). Fifty Major Philosophers a Reference Guide. London. Routledge.
Cummins, A., Piek J., Dyck, M. (2005). Motor coordination, empathy and social behavior in
school-aged children. Developmental Medicine and Child Neurology, 47, 437-442.
Damasio, D. (1999). The Feeling of What Happens. Harcourt, Inc. San Diego.
Damasio, A. (1998). The somatic marker hypothesis and the possible function of the prefrontal
cortex. In A. C. Roberts et al. (Eds). The Prefrontal Cortex. Oxford: Oxford University
Press.
Dapretto, M., Davies, M., Pfeifer. J., Scott, A., Sigman, M., Bookheimer, S., Iacoboni, M.
(2006). Understanding emotions in others: mirror dysfunction in children with autism
spectrum disorder. Nature Neuroscience. Jan.9(1), 28-30.
Dawson, G., Web, S., Carver, L., Panagiotides, H., McPartland, J. (2004). Young children with
autism show atypical brain responses to fearful versus neutral facial expressions of
emotion. Developmental Science. 7(3), 340-359.
Dickman, S. J., (2002). Dimensions of arousal: Wakefulness and vigor. Human Factors, 44 (3),
429-442.
Dunn, W. (1997). The impact of sensory processing abilities on the daily lives of young children
and their families: a conceptual model. Infants and Young Children. 9, 23 -35.
Fadiga, L., Fogassi, L., Pavesi, G.., Rizzolatti, G. (1995). Motor facilitation during action
observation: a magnetic stimulation study. Journal of Neurophysiology, 73, 2608-2611.
Ferrari, P. F., Rozzi, S., Fogassi, L. (2005). Mirror neurons responding to the observation of
actions made with tools in monkey ventral premotor cortex. Journal of Cognitive
Neuroscience. 17(2), 212-226.
Action and Understanding
56
Ferrari, P. F., Gallese, V., Rizzolatti, G., & Fogassi, L. (2003). Mirror Neurons responding to the
observation of ingestive and communicative mouth actions in the monkey ventral
premotor cortex. European Journal of Neuroscience, 17(8), 1703-1714.
Fogassi, L., Gallese, V., Buccino, G., Craighero, L., Fadiga, L., & Rizzolatti, G. (2001). Cortical
mechanism for the visual guidance of hand grasping movements in the monkey. Brain, 124(3),
571-586.
Gallese, V. M. (2003). The manifold nature of interpersonal relations: the quest for a common
mechanism. Philosophical Transcripts of the Royal Society of London. 358, 517-528.
Gallese, V. M., Metzinger, T. (2003). Motor ontology: the representational reality of goals,
actions and selves. Philosophical Psychology. 16 (3).
Gallese, V., Fadiga, L., Fogassi, L., Rizzolatti, G. (1996). Action recognition in the premotor
cortex. Brain, 119, 593-609.
Gergely, G., Csibra, G., (2003). Teleological reasoning in infancy: the nalive theory of rational
action. Trends in Cognitive Neuroscience. 7 (7): 287-292.
Gergely, G., Nadasdy, Z., Csibra, G., S. Biro. (1995). Taking the intentional stance at 12 months
of age. Cognition. 56 (2), 165-193.
Grezes, J., Armony, J., Rowe, J., Passingham, R. (2003). Activations related to mirror and
canonical neurones in the human brain: an fMRI study. Neuroimage, 18, 928-937.
Happe, F., (2003). Theory of mind and the self. Annuals of the New York Academy of Science.
1001, 134-144.
Hari, R., Forss, N., Avikainen, S., Kirveskari, E., Salenius, S., & Rizzolatti, G. (1998).
Activation of human primary motor cortex during action observation: a neuromagnetic
study. Proceedings of the National Academy of Science, 95(25), 15061-15065.
Action and Understanding
57
Hayes, R. W., Gordon, A. (1977). Auditory abnormalities in autistic children. Lancet, 2,767.
Heyes, C. (1994). Social learning in animals. categories and mechanism. Biological Review. 69,
207-231.
Hirstein, W., Iverstein, P., Ramachandran, V.. (2001). Autonomic responces of autistic children
to people and objects. Philosophical Transcripts of the Royal Society of London. 268,
1883-1888.
Howard, M. A., Cowell, P., Boucher, J., Broks, P., Mayes, A., Farrant, A., Roberts, N. (2000).
Convergent neuroanatomical and behavioral evidence of amygdale hypothesis of autism.
Neuroreport 11, 2931-2935.
Iacoboni, M., Molnar-Szakacs, I. (2005). Grasping the intention of others with one's own mirror
system. PloS Biology, 3(3), 529-535.
Iacoboni, M., Woods, R. P., Brass, M., Bekkering, H., Mazziotta, J. C., & Rizzolatti, G. (1999).
Cortical mechanism of human imitation. Science. 286(5449), 2526-2528.
Jarvelainen, J., Schurmann, M., Hari, R. (2004). Activation of the human primary motor cortex
during observation of tool use. Neuroimage. 23(1),187-192.
Jeannerod, M. (1997). The Cognitive Neuroscience of Action. Oxford: Blackwell Publishers.
Kandel, E., Schwartz, J., Jessell, T. (2000). Principles of Neural Science. New York. Mc-Graw
Hill.
Kanner, L. (1943). Autistic disturbances of affective contact. The Nervous Child 2, 217-250.
Kendon, A. (1970). Movement coordination in social interaction: some examples described. Acta
Psychologia. 32, 101-125.
Keysers, C., Wicker, B., Gazzola, V., Anton, J., Fogassi, L., Gallese, V. (2004). A touching site:
SII/PV activation during the observation and experience of touch. Neuron. 42, 335-346.
Action and Understanding
58
Keysers, C., Kohler, E., Umilta', M., Nanetti, L., Gogassi, L., Gallese, V. (2003). Audiovisual
mirror neurons. Experimental Brain Research. 153, 628-636.
Kiln, A. (1992). Listening preference in regard to speech; a possible characterization of the
symptom of social withdrawal. Journal of Autism and Developmental Disorders. 21, 2942.
Kinnealy, M., Oliver, B.,Wilbarger, P. (1995). A phenomenological study of sensory
defensiveness in adults. American Journal of Occupational Therapy. 49, 444-451.
Kohler, E., Keysers, C., Umilta, M. A., Fogassi, L., Gallese, V., Rizzolatti, G. (2002). Hearing
sounds, understanding actions: action representation in mirror neurons. Science.
297(5582), 846-848.
Kootz, J. P., Martinelli, B., Cohen, D. (1981). Sensory receptor hypersensitivity in autistic
children. Archives of General Psychiatry. 38, 271-273.
Lane, S. J., (2002). Structure and function of the sensory systems and sensory modulation., A. C.
Bundy, S. J. Lane, & E. A. Murray (Eds), Sensory integration: Theory and Practice (2nd
ed.). Philadelphia: F. A. Davis. 35 -70, 101 -122.
Laundry, R., Bryson, S. (2004). Impaired disengagement in young children with autism. Journal
of Child Psychology and Psychiatry. 45, 1115-1122.
Lepage, J. F., Theoret, H. (2006). EEG evidence for the presence of an action observationexecution matching system in children. European Journal of Neuroscience. 23 (9), 25052510
McFarland, D., Miner, L., Vaughn, T., Wolpaw, J. (2000). Mu and beta rhythm topographies
during motor imagery and actual movements. Brain Topography. 12(3), 177 - 186.
Action and Understanding
59
McIntosh, D., Miller, L., Shyu, V., Hagerman, R. (1999). Sensory modulation, disruption,
electordermal responses and functional behaviors. Developmental Medicine and Child
Neurology. 41 608 -615.
Mesulam, M., (1998). From sensation to cognition. Brain. 121,1013-1052).
Meltzoff,. (1995) Understanding the intention of others: re-enactment of intended acts by 18
month old children. Developmental Psychology. 31, 838-850.
Morrison, I. (2006). Mirror neurons and cultural transmission. In M. Staminov et. al. (Eds.),
Mirror Neurons and the Evolution of Brain and Language. Amsterdam: John Benjamins
Publishing.
Mundy P., Sigman, M., Kasari, C. (1990). A longitudidinal study of joint attention and language
development in autistic children. Journal of Autism and Developmental Disorders. 20,
115-129).
Murata, A., Fadiga, L., Fogassi, L., Gallese, V., Raos, V., & Rizzolatti, G. (1997). Object
representation in the ventral premotor cortex (area F5) of the monkey. Journal of
Neurophysiologia. 78(4), 2226-2230.
Muthukumaraswamy, S., Johnson, B. (2004). Changes in rolandic mu rhythm during observation
of a precision grip. Psychophysiology. 41, 152-156.
Niedermeyer, E., Da Silva, F. L. (2005). Electroencephalography; Basic Principles, Clinical
Applications and Related Fields. Fifth Edition. Lippincott Williams and Wilkins.
Philadelphia, Pa.
Oberman, L. M., Hubbard, E., McCleery, J., Altschular, E., Ramachandran, V., Pineda, J. (2005).
EEG evidence for mirror neuron dysfunction in autism spectrum disorders. Cognitive
Brain Research. 24, 190-198.
Action and Understanding
60
Olausson, H., Lamarre, Y., Backlund, H., Morin, C., Wallin, B., Stark, G., Ekholm, S., Strigo, I.,
Worsley, K., Vallbo, A., Bushnell, M. (2002). Nature Neuroscience. 5(9), 900-904.
Piaget, J. (1962). Play, Dreams and Imitation of Children in Childhood. New York: Norton .
Pierce, K., Muller, R., Ambrose, J., Allen, G., Courchesne, E. (2001). Face processing occurs
outside the fusioform 'face area' in autism; evidence from functional MRI. Brain. 124,
2059-2073.
Pineda, J. (2005). The functional significance of mu rhythms: translating seeing and hearing into
doing. Brain Research Brain Research Review. 1;50(1), 57 - 68.
Prisniakova, L. M. (2004). Fitness to work of astronauts in conditions of action of the extreme
emotional factors. Advanced Space Research. 33(8), 1381-1385.
Pfurtscheller, G., Neuper, C., Andrew, C., Edlinger, G. (1997). Foot and hand mu rhythms.
International Journal of Psychophysiology. 26(1-3), 121-135.
Prum, R. O., Brush, A. (2003). Which came first the feather or the bird? Scientific American.
March.
Rapin, I. (Ed). (1996). Preschool children with inadequate communication; developmental
language disorders, autism, low IQ. Clinics in Developmental Medicine, 139. London.
Mac Keith Press.
Rizzolatti, G., Craighero, L. (2004). The mirror-neuron system. Annual Review of Neuroscience.
27, 169 - 192.
Rubenstein, J. L., Merzenich, M. (2003). Model of autism: increased ration of excitation/
inhibition in key neural systems. Genes, Brain and Behavior. 2, 255-267.
Action and Understanding
61
Salamond, C. H., de Haan, M., Friston, K., Gadian, D., Varfgha-Khadem, F. (2003).
Investigating individual differences in brain abnormalities in autism. Philosophical
Transcripts of the Royal Society of London. 358 (405-413).
Schleidt and Schleidt, (1960) as cited in Mesulam, 1998 Brain, 121,1013-1052) From Sensation
to Cognition. Oxford University Press.
Schwoebel, J., Boronat, C., Branch Coslett, H. (2002). The man who executed imagined
movements: evidence for dissociable components of the body schema. Brain and
Cognition. 50(1), 1-16.
Singer, T., The neuronal basis and ontogeny of empathy and mind reading: review of literature
and implications for future research. (2006). Neuroscience Biobehavioral Review. 30(6)
855-863.
Singer, T., Seymour, B., O'Doherty, J., Kaube, H., Dolan, R. J., Frith, C. D. (2004). Empathy for
pain involves the affective but not sensory components of pain. Science. 303(5661),
1157-1162.
Spivey, M., Spirn, M. (2000). Selective visual attention modulates the direct tilt aftereffect.
Perceptual Psychophysiology. 62(8), 1525-1533.
Umilta, M., Kohler, E., Gallese, V., Fogassi, L., Fadiga, L., Keysers, C., Rizzollati, G. (2001). I
know what you are doing: a neurophysiological study. Neuron, 31, 155-165.
Valinger, H. (1965). The philosophy of "as if." (C. K. Ogden, Trans.). London: Routledge &
Kegan Paul. (Original work published 1911).
Volkmar, E. R.,Carter, A.,Grossman, J., Kiln, A. (1997). Social development in autism. In
D. Cohen & F. Volkmar (Eds), Handbook of Autism and Pervasive Developmental
Disorders (2nd edition., pp. 173-194). New York, Wiley.
Action and Understanding
Wicker, B., Keysers, C., Pially, J., Yoyet, J. (2003). Both of us disgusted in my insula: the
common neural basis of seeing and feeling disgust. Neuron, 40 (October 30), 655-664.
62