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Cognition 146 (2016) 81–89

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Cognition
journal homepage: www.elsevier.com/locate/COGNIT

Discussion

Towards a common framework of grounded action cognition: Relating


motor control, perception and cognition
Antje Gentsch a,b, Arne Weber c, Matthis Synofzik d,e, Gottfried Vosgerau c, Simone Schütz-Bosbach b,f,⇑
a
University College London, CEHP Research Department, London, UK
b
Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany
c
Department of Philosophy, Heinrich-Heine-University Düsseldorf, Germany
d
Department of Neurodegenerative Diseases, Hertie-Institute for Clinical Brain Research, University of Tübingen, Germany
e
German Research Center for Neurodegenerative Diseases (DZNE), Tübingen, Germany
f
Psychology Department, Ludwig-Maximilians-University, Munich, Germany

a r t i c l e i n f o a b s t r a c t

Article history: The relation between motor control and action cognition – including action-related thoughts and action-
Received 8 May 2015 related perception – has been subject to controversial discussions in the last three decades. During these
Revised 11 September 2015 decades, cognitive neuroscience has been increasingly confronted with a huge variety of different
Accepted 14 September 2015
accounts trying to understand and explain the relation between these systems, their interdependencies
Available online 24 September 2015
and the mediating mechanisms by establishing notions such as ‘‘internal models”, ‘‘simulation” or
‘‘shared representation”. These accounts, however, include a large array of partly overlapping, partly con-
Keywords:
tradictory theories using similar terms for different mechanisms and different terms for similar mecha-
Grounded cognition
Motor control
nisms. In the absence of a systematic work-up and comparison, this array of accounts and theories leads
Action perception to confusion in the field, duplication of experimental work, and unconnected parallelism of theory forma-
Action cognition tion within and between different disciplines. Here we provide a systematic comparison of current mod-
Simulation els and prospective theories that deal with the relation between cognition, perception and motor control
Internal models mechanisms. In a second step, we propose ‘‘grounded action cognition” as a comprehensive metatheoret-
Common coding ical framework which defines different hypothetical possibilities of the relations between these domains,
offers systematic insights into current models and theories and last but not least may help to increase
comparability of empirical research in the domain of action and action cognition.
Ó 2015 Elsevier B.V. All rights reserved.

1. Introduction ing the precise brain systems and neuronal dynamics underlying
the action–perception inter-linkage by means of internal forward/-
There is a long tradition and interest in many different disci- generative models (Friston, Daunizeau, Kilner, & Kiebel, 2010;
plines on the interplay between action and perception. This inter- Wolpert & Miall, 1996). The focus on the linkage between action
est is led by various aims: while some disciplines investigate this and perception has now been expanded to include also conceptual
interplay with the aim of understanding human ‘cognitive mecha- abilities and cognition in general. Specifically, growing interdisci-
nisms’ (cognitive psychology, philosophy), brain mechanisms (cog- plinary work has now begun to relate different theoretical
nitive neuroscience) or neurologic and psychiatric diseases approaches and empirical findings to explain also higher-
(neurology/psychiatry), others study this relation to model skillful cognitive skills like mind reading in social contexts (Gallese &
movements (robotics) or perceptual abilities (computational and Goldman, 1998) or mental imagery (Grush, 2004).
mathematical neuroscience). Some of these traditions date back Here we take an interdisciplinary point of departure with the
to 19th century experimental psychology and philosophical theory aim to provide a systematic comparison of current and established
of voluntary action (James, 1890/1981). Others were developed theoretical models and prospective theories that deal with the
more recently. For example, approaches from computational and relation between cognition, perception and motor control mecha-
cognitive neuroscience, starting in the early 1990s, aim at describ- nisms. We will in particular focus on the proposed internal repre-
sentational mechanisms governing mutual relations between
⇑ Corresponding author at: Psychology Department, General and Experimental perception and action. Accordingly, the selection criterion for the
Psychology, Ludwig-Maximilians-University Munich, 80802 Munich, Germany. theories to be compared in the present paper is that they make
E-mail address: simone.bosbach@gmail.com (S. Schütz-Bosbach).

http://dx.doi.org/10.1016/j.cognition.2015.09.010
0010-0277/Ó 2015 Elsevier B.V. All rights reserved.
82 A. Gentsch et al. / Cognition 146 (2016) 81–89

some substantial claim about the systematic connection between common coding theories as well as some internal model accounts,
the domains of cognition, perception and motor control.1 It is such as motor control theory, were explicitly developed from a
beyond the scope of the present endeavor to also take dynamic inter- motor control point of view. Therefore, the theoretical concepts
actions between mind/brain, body and world into account as pro- are grounded in notions of efferent and (re-)afferent information
posed by some more radical conceptions of embodied cognition (i.e., motor output and sensory feedback) which have been applied
(for an overview compare Shapiro, 2011; or Wilson, 2002; for certain not only to understand the mechanisms behind movement kine-
conceptions of these radical or dynamic views see Beer, 1995; Hutto matics (motor control theory) but also to describe the semantics
& Myin, 2013; Keijzer, 2002). Thus, we will here specifically consider of action (ideomotor theory). In contrast, another version of inter-
internal representations that appear to draw on motor-related pro- nal models, predictive coding, was originally developed to explain
cesses. We will use the term ‘grounding’ to refer to the general rela- visual perception and its underlying computational architecture. It
tion between motor processes and action perception/cognition, has been extended to other sensory modalities as well as to action
which seems underdetermined so far in current theories. Please also in terms of active inference (the latter being a more recent devel-
note that it is not our aim to provide a systematic review of those opment). Hence, the key concepts in this model are rooted in
single theories in terms of supporting or challenging empirical evi- assumptions around the organization of sensory input and the per-
dence (for a recent review see, for example, Engel, Maye, Kurthen, ception of the outside world. The ontogeny of simulation theories,
& König, 2013). Instead, we here aim to systematically compare the third main family of action theories, is most heterogeneous,
the empirical and explanatory foci those theories adopt with respect since the notion of ‘simulation’ has been used to account for a vari-
to the nature of the relation between motor control and action– ety of phenomena in the domains of action cognition and percep-
cognition. tion. For example, the notion of simulation has been used to
This paper comprises three sections. The first provides a brief explain a perceptual understanding of other’s actions (mirror neu-
overview of different action theories and suggests a classification ron theory) and the ascription of mental states to others (simula-
by target mechanisms. In the second section, we introduce a tion theory), as well as the imagination of action (motor
framework that will be used to illustrate our notion of grounded imagery) and even abstract conceptual abilities (perceptual sym-
action cognition as a metatheoretical view. Third and finally, our bols theory). Hence, understanding the primary phenomenon that
purpose is to show how existing theories can or cannot be classi- each theory is trying to account for is crucial for appreciating how
fied into genuine grounding theories. This new classification existing theories differ in relating the domains of cognition, per-
scheme shall offer new perspectives into commonalities and differ- ception and motor control mechanisms. The classification of the
ences as well as the explanatory scope with respect to the degree three families of theories according to their primary domain of
to which perceptual and cognitive abilities genuinely draw on explanation is illustrated by Fig. 1.
motor capacities; however, it is not meant to suggest an evaluation
of existing and established theories concerning their theoretical
and empirical adequacy. 3. Grounded action cognition: a metatheoretical view

In our approach towards a systematic framework, we will now


2. The three main families of action theories
specify common denominators of the manifold existing assump-
tions regarding the relation between the three domains of action
The large amount of highly heterogeneous, partly overlapping,
cognition, perception, control and concepts of action. Current the-
partly differing theories coming from very different disciplines is
ories and related experimental reports sometimes appear to use
often confusing. To facilitate a better overview and understanding,
the same terms to explain different phenomena or mechanisms,
we here classify the existing theories into three major theoretical
and different terms to explain the same phenomena or similar
frameworks/families of grounded action cognition accounts: (1)
mechanisms. For example, the term ‘simulation’ has been used to
Common Coding, (2) Internal Models, and (3) Simulation theories
refer to sensorimotor processes as a reactivation of neural motor
(Table 1; for a detailed description and analysis of each of these
circuitries not only when observing actions but also in the context
families, see next paragraph). These theories operate at different
of reasoning about mental states of others. Likewise, the finding of
levels and interfaces between domains of action cognition and
an activation of cortical areas involved in motor control during
use diverse conceptual tools (see Table 1). One shared assumption
action observation has been interpreted and labeled interchange-
of all major theories is that some kind of common representational
ably as ‘mirroring’, ‘simulating’ or ‘mentalizing’. Moreover, terms
ground for action and cognition exists. This assumption has been
such as ’embodiment’, for instance, have been applied in a narrow
developed, however, at different levels of action cognition, and
sense with reference to self-awareness as a form of bodily aware-
with crucial differences in the empirical and explanatory focus
ness (i.e., the embodied self) but also in a broader sense to explain
on motor control, perception or conceptual abilities (see Fig. 1).
a fully functional system of abstract concepts (cf. below, perceptual
This includes fundamental differences in the abilities these theo-
symbol systems theory).
ries try to account for: while some theories resort to a common
Moreover, study reports sometimes use vague or even over-
ground between action and cognition to explain action planning
stated wording when reporting their findings and interpreting the
or motor imagery, others refer to this notion to explain the under-
relation between the three domains. For instance, authors sympa-
standing of other individuals’ actions. The main assumptions
thetic with simulation theories often use various and quite strong
regarding the central thesis of grounded action cognition are sum-
expressions like cognition/perception (x) is ‘‘based” on a sensorimo-
marized in Table 1.
tor process/the sensorimotor system (y), x ‘‘arises” in y,2 x is ‘‘local-
In order to understand the differences between the theories in
ized” in y,3 x is ‘‘derived” from y,4 x is ‘‘specified” by y,5 x is
their attempt to link the domains of action, perception and cogni-
tion, it is helpful to consider the primary domain a theory was orig-
2
‘‘[P]erceptual states arise in sensory-motor systems.” (Barsalou, 1999, p. 577).
inally developed for and how it has been extended. For example, 3
‘‘[I]implicit memory has been localized in sensorimotor areas of the brain.”
(Barsalou, 1999, p. 589).
1 4
For this reason, the sensorimotor theory of visual consciousness by O’Regan and ‘‘[A]ffordances are derived from the perceptual symbols [i.e. a systematic
Noë (2001), for example, is not considered in the present framework: Although it activation pattern in the sensorimotor system].” (Glenberg & Kaschak, 2002, p. 599).
5
claims that visual perception is a form of action, it does not propose a specific linking ‘‘[C]onceptual processing of categories specified by motor and somatosensory
mechanism between perception and action (unlike ideomotor theory, for example). features [. . .].” (Barsalou, 1999, p. 585).
A. Gentsch et al. / Cognition 146 (2016) 81–89 83

Table 1
Systematic overview of theories of action cognition. These theories assume a tight, but differing interrelation between the domains of motor control, action perception and action
cognition. They can be grouped into three different families of grounded action cognition accounts: Common Coding, Internal Models, and Simulation theories.

Theory Explanatory target How do motor cognition, motor control and Origin Key references
action perception relate?
Common Ideomotor Action initiation A bidirectional sensory-motor link: perceptual Cognitive Greenwald (1970) and James (1890/
Coding Theory Cognitive processes behind representations automatically activate Psychology 1981)
voluntary intentional action response representations of associated motor (motor control)
actions, and vice versa
Theory of Action planning and A common code: the representational format Cognitive Hommel et al. (2001) and Prinz (1997)
Event perception for distal perceptual contents and motor Psychology
Coding Communication between plans is identical and formed through (action
motor control and action principles of temporal coupling and perception)
perception synchronized activation
Internal Motor Motor control & motor An internal forward model which represents Computational Wolpert and Miall (1996)
Models Control learning the dynamics of the motor system, including Modeling,
Theory Implementation of optimal all sensory states resulting from motor Cognitive
online control of actions, in order to predict future states Robotics (motor
movements and posture control)
Predictive Sensory perception & motor Active inference under the free-energy Computational Friston (2005) and Friston, Mattout, and
Coding control principle: sensory surprise automatically Modeling Kilner (2011)
Perception as prediction translates into motor control representations (perception in
(similar to motor reflexes) that serve to general)
minimize sensory prediction errors by
guiding sensory sampling
Emulation Motor control & motor Draws on and combines elements of internal Philosophy Grush (2004) and Hurley (2008)
Theory imagery models and mirror mechanisms central to (action
Interrelation of neural and motor control theory and mirror neuron cognition)
representational theory
mechanisms underlying
motor control and motor
imagery
Simulation Simulation Mind reading The concept of mental simulation: action Philosophy, Gallese and Goldman (1998), Goldman
Theories Theory Ascribing mental states of observation (i.e., the perceptual Cognitive (1992), and Gordon (1986)
others by observing their representation of a motor act of others) Neuroscience
actions involves the activation of mental state (social
sequences that are isomorphic to those cognition)
underlying self-generated actions
Mirror Action observation The notion of mirror neurons: a population of Neuro- Gallese et al. (1996) and Rizzolatti,
Neurons Neuroanatomy of observed neurons coding for both observed and physiology Fadiga, Gallese, and Fogassi (1996)
actions of others executed actions, that is, the same neural (action
system is used for representing motor acts perception)
and motor percepts
Motor Mental imagery The notion of covert actions: Motor images (i. Cognitive Decety, Jeannerod, and Prablanc (1989),
Imagery Neural mechanisms e., motor percepts) are covert actions, relying Neuroscience Jeannerod (2001), Jeannerod and
underlying overt and covert on simulation of neural mechanisms and (action Decety (1995), for review’, Vogt, Rienzo,
(imagined) actions dynamics of real, physical action in low level perception) Collet, Collins, and Guillot (2013)
motor centers
Perceptual Conceptual thinking The construct of simulators: Concepts act as Cognitive Barsalou (1999) and Barsalou, Kyle
Symbols Conceptual abilities simulators by relying on a re-activation of the Psychology Simmons, Barbey, and Wilson (2003)
Theory sensorimotor states involved in experiencing (cognition in
the actual entity general)

‘‘embodied”/”grounded” in y (passim), x is ‘‘constituted” by y,6 x ‘‘re- path towards an integrative understanding of current theories of
sides” in y,7 x/y ‘‘integrate(s)” y/x,8 y ‘‘enters” into x, y is ‘‘engaged” action. Within this new framework, a crucial distinction is drawn
in x, or y ‘‘essentially is” for x.9 All of these formulations are used to between constitution relation and acquisition relation, which offers
interpret data from experimental work usually showing a correlation a systematic classification tool for existing and future theories of
between two factors. However, the fact that two factors correlate can grounded action cognition.
be interpreted in very different ways, from mere unspecified
modulation of one factor by the other to causal dependence of the 3.1. Acquisition and constitution: specifying the relation between
one factor on the other. Here we propose grounded action cognition perception/cognition and action
as a metatheoretical framework that aims to define hypothetical
possibilities for the relations between different domains of action In order to specify the claim of grounded cognition, it is crucial
representation in order to provide a systematic synopsis and a to scrutinize the exact relation that is assumed to exist between
two domains of interest (e.g., between motor control and action
6
‘‘[O]ur ability to perceive not only depends on, but is constituted by, our
perception, or action cognition). So far, the question of whether
possession of [. . .] sensorimotor knowledge.” (Noë, 2004, p. 2). grounded or embodied cognition is meant to oppose ‘‘classical”
7
‘‘[C]ognitive and perceptual representations reside in a common system.” modular pictures of the mind (Fodor, 1975; Pylyshyn, 1984;
(Barsalou, 1999, p. 589). Smith & Medin, 1981; Tulving, 1972) is in fact not answered. There
8
‘‘[T]he sensorimotor integration supported by the premotor-parietal F5-PF
is certainly no doubt that action (i.e., motor control) and percep-
mirror-matching system [. . .] also to provide a meaningful account of the goals and
purposes of others’ actions, by means of their simulation.” (Gallese, 2003, p. 523).
tion/cognition are somehow interrelated. The question is thus
9
‘‘[M]otor behavior of an exploring animal is essential to perception [. . .].” (Hurley, not whether these processes are related, but how they are related.
2001, p. 11). Representatives of the ‘‘classical” picture of the mind even do not
84 A. Gentsch et al. / Cognition 146 (2016) 81–89

Fig. 1. Theories of ‘‘grounded action cognition” and their main domain of focus. The diagram relates theoretical accounts of action cognition and classifies them according to
their primary empirical and explanatory focus. All theories draw on a relation between action and cognition to explain a mental ability related to motor action. However, they
differ in the domain and process of action cognition they primarily refer to (action control, perception or concepts), and in the mental ability they are trying to account for.
Simulation Theories (green) are concerned with action perception and conceptual representations in order to understand socio-cognitive skills of mind-reading based on
mechanisms of motor control. Internal Models (blue) aim at describing primarily the interrelation between motor control and perception of action, by assuming that
perception arises with reference to motor control functions (cf. notions of ‘‘forward models” and ‘‘prediction error”) and motor planning arising with reference to perception
(cf. notions of ‘‘inverse models” or ‘‘active inference”). Common Coding (orange) accounts span all three areas of action perception, control and concepts by explaining (late)
event perception and (early) action planning through the existence of a shared sensorimotor representational code at a higher level.

explicitly deny a perception–action interrelation (although they acquire the ability to perceive but not necessary to maintain this
usually do not focus on it), but they typically claim this relation capacity once it is fully acquired.
to be rather weak, for example in the sense of mere modulating ‘‘Constitution”, in contrast, refers to a much tighter interpreta-
bi-directional influences. Thus, our reconstruction of the percep- tion of the relation between action and perception. An almost triv-
tion–action interrelation meant by the different authors above will ial example to illustrate a real constitution relation is that the
be such that their claims are indeed opposing more classical views ability to move one’s own legs is constitutive for the ability to walk,
in claiming a much stronger, thus functional determining relation while both abilities are not identical, since it is possible to be able
between action and perception in the sense of acquisition and/or to move one’s own legs without being able to walk. So, already in
constitution. In a first step, we will now introduce the technical this example it is obvious that constitution does not necessarily
terms of acquisition conditions and of constitution, respectively mean that there is no more to ability B than just ability A: indeed,
(cf., Weber & Vosgerau, 2012). We will illustrate how we under- there is much more to the ability to walk than just the ability to
stand these terms with the help of empirical examples and we will move one’s own legs. Nevertheless, you cannot do the latter with-
also discuss some empirical implications that follow from this new out being able to do the former. However, constitutive relations
perspective. can also be interpreted less strong. For example, if the complete
Acquisition can be illustrated by a classical experiment of Held loss of A only leads to some impairments of B, it is still valid to
and Hein (1963) in newborn animals. They divided kittens into two argue that A is a constitutive element of full-blown ability B. Thus,
groups; kittens in one group pull a carriage around a room in a hor- although the criterion for constitutive relations might read very
izontally rotating carousel, while kittens in the other group ride strong at first sight, it only requires a systematic dependence
this carriage. The carousel was set up such that both groups of kit- between break-downs in either ability.
tens have the same visual stimuli, the only difference being that So far only relatively few10 empirical studies were able to indi-
kittens in the first group are allowed to make active movements cate that motor control processes indeed play a constitutive role in
while the kittens in the other group were only passively moved.
After spending some time in the carousel, tests show that the rid- 10
Among notable exceptions are, for example, studies using brain stimulation such
ing kittens did not develop normal perceptual abilities as for exam- as Transcranial Magnetic Stimulation (TMS; Moore, Ruge, Wenke, Rothwell, &
ple depth perception. Obviously, these kittens were not able to Haggard, 2010, linking pre-SMA to both action preparation and action perception)
integrate their visual experiences with movements of their own or invasive, intracranial techniques such as direct electrical stimulation (DES;
Desmurget et al., 2009, linking parietal cortex to both representing motor intentions
bodies, which caused visual deficits in the long run (see also,
and forward modeling for movement monitoring). Although these studies attempt to
Gibson, 1969). These findings suggest that the ability to actively show causal relations, however, even these methods have some caveats. The effects
move (ability A) is among the acquisition conditions for the ability evoked by surface stimulation of the cortex are complex, mostly involving a large
to perceive depth (ability B). However, once B is acquired on the volume of brain tissue, and behavioral effects across studies are heterogeneous
basis of A, ability A can get lost without any negative impact on ranging from facilitation to inhibition of responses. For example, TMS stimulation of a
specific target area may induce effects in a larger network of interconnected brain
ability B. So, a kitten that already acquired normal sight will not regions depending on different physiological and morphological factors. Thus, these
lose its ability to perceive if we put it in a carriage (thus hindering techniques may not allow us to draw unequivocal conclusions about whether an
it from active movement). Active movement is only necessary to effect is indeed being generated in a stimulated brain area.
A. Gentsch et al. / Cognition 146 (2016) 81–89 85

action cognition, thus going beyond mere correlations between certain perceptual process as ‘‘grounded” in another basic process,
motor control processes and action cognition. A prominent example since then there is nothing left to be ‘‘grounded”; the former pro-
is a study that investigated patients diagnosed with amyotrophic lat- cess would be merely identical to the latter. Moreover, the differ-
eral sclerosis (ALS). ALS refers to a neurodegenerative condition that ent domains of motor control and action perception would not
is defined by progressive degeneration of motor tracts. This study be discriminable in empirical research. The other hypothetical
showed that ALS patients were significantly more impaired in con- extreme case would be that there is no constitution at all of one
cepts of actions than in concepts of objects, and that this difficulty process by another. This would then in the end lead to a picture
specifically correlated with atrophy of the motor cortex (Grossman similar to the classical conception of a modular mind (see above).
et al., 2008). These findings suggest that intact conceptual knowl- Then, however, there would be no novel and challenging way of
edge of action features depends on an intact motor cortex. If this talking about some cognitive processes potentially being
finding can be corroborated by future studies, confirming indeed a grounded. Such relations would simply be too weak to establish
specific relation between degraded action knowledge and degraded a picture opposing the classical one. Therefore, we argue that
motor systems (and not just degraded action knowledge as part of grounded action cognition should be understood and specified in
the more widespread extramotor cortical atrophy known in ALS terms of a partial constitution relation. The different types of con-
(Libon et al., 2012), this would suggest that motor systems are stitution relation are illustrated by Fig. 2.
indeed constitutive for action cognition. Intact motor systems would
here present the structural presupposition to intact action cognition.
4. Action cognition theories and implied constitution relation
In another domain, intact motor processes present the func-
tional presupposition for acquiring intact perception. Microsac-
In the first part of the present paper, we organized the estab-
cades, for example, allow restoring faded vision during fixation,
lished action cognition theories into three major families, Common
for both foveal and peripheral targets. Accordingly, it has been
Coding (CC), Internal Models (IM) and Simulation Theories (ST). We
shown already in the 1950s that when eye movements are com-
will now analyze them in more detail with respect to whether they
pletely suppressed (for instance, by techniques of retinal stabiliza-
can be considered genuine grounding theories; that is, they will be
tion (Ditchburn & Ginsborg, 1952), our perception of stationary
compared according to the implied constitution relation between
objects appears to fade completely (Martinez-Conde, Macknik,
motor control and action cognition. The aim shall be to put the
Troncoso, & Dyar, 2006; for review see, Martinez-Conde, Otero-
variety of existing accounts of action into a systematic frame of
Millan, & Macknik, 2013). For the more specific thesis that motor
grounded action cognition. Each family contains some theories that
processes are indeed constitutive elements for acquiring intact
implicitly assume an at least partial constitutive relation between
action perception, it would be necessary to demonstrate that sim-
action and perception, while other theories in contrast do not
ilar process are at work in the perception of actions.
imply a constitutive relation. In Table 2, we made an attempt to
organize those theories according to the constitution relation they
3.2. The partial constitution criterion
assume to hold. We will discuss the categorization of each theory
family in our table in turn.
Our proposal here is that the only genuine grounding relation is
that of a partial constitution relation between perception and
action, in the sense that there is more to ability B than just ability 4.1. Common Coding Theory
A. Thus, on this view the relation does not collapse into an identity
relation. To make this argument clearer, let us take the hypotheti- Common Coding theories are characterized by the assumption
cal case of a radical and full constitution of a certain perceptual/ that there is a bidirectional or common sensorimotor code. The
cognitive ability by motor processes. This would lead us to the ideomotor theory assumes that action and perceptual representa-
assumption that this ability is just a subclass of or even identical tions are separate representations in principle, but can be co-
to some (complex) motor ability. Moreover, if all perceptual/cogni- activated to generate and facilitate voluntary action and adaptive
tive abilities would be so fully constituted by motor processes, control of goal-directed behavior. Thus, the motor domain is here
some basic motor abilities would be constitutive of all action- not seen as constitutional for perceptual and cognitive abilities
perceptual and -cognitive capacities, and this implies that the (see Table 2). In contrast, the theory of event coding holds that
two domains would fall together and become the same. However, there is a common representational code for action and perception.
if perceptual processes or states are just kinds of motor processes This code structures action planning and action perception
or states, there would be no intelligible sense of understanding a and therefore action and perception are related through this

Fig. 2. Schematic representation of different types of constitution relation between two abilities, A and B. (i): Some B-ability correlates with some A-ability (dotted line),
while there is no constitution relation between the two. (ii): A B-ability is partially constituted by A-abilities if the loss of A leads to severe impairment of the B-ability but not
to a complete breakdown (since there is more to ability B than just A). (iii): The B-ability is fully constituted by A-abilities if there is no non-A-ability constitutive for it, and in
this case the B-ability is nothing but a (complex) A-ability itself.
86 A. Gentsch et al. / Cognition 146 (2016) 81–89

Table 2
The three major families of action cognition theories and the main assumptions regarding conditions of constitution.

Constitution Theory Central hypothesis


relation
Common Coding No Ideomotor theory Separate motor and perceptual representations can be co-activated
No Theory of event coding A common code is assumed, but only for representation of distal events and in addition to lower-
level sensory and motor representations
Internal Models Partial Motor control theory Internal forward models are thought to run in parallel to motor control processes
Partial Predictive coding In probabilistic models, processes of motor control and perceptual inference are identical upwards
of the spinal level
No Emulation theory Cognition (including motor imagery and motor planning) is based on the internal emulation of
motor and sensory processes
Simulation Full Mirror neuron theory; Specific, separate subpopulation of neurons in motor areas with mirror or simulation properties
Theories Motor imagery
Full Perceptual symbols theory Simulation requires a reenactment of sensorimotor mechanisms, no additional process is assumed

representational format. Critically, however, the code is indepen- 4.2. Internal Models
dent of executive action domains as well as of sensory modalities.
Thus, following this picture, the domain of motor control is not Internal Model theories are characterized by the proposed exis-
directly connected to (or even constitutional for) sensory modali- tence of internal models of the motor apparatus. These models are
ties. Instead, a common code enables an indirect communication postulated neural processes that extrapolate the commands of the
between the two domains. Moreover, motor control is not seen motor system in order to estimate and anticipate the outcome of a
as the primary system because the perceptual system and the motor command. One variant of Internal Model theories, motor
motor control system are coequal. control theory, has its origin in control theory and robotics and pri-
The main aim of Common Coding theories is to explain the rep- marily aims at explaining optimal online control of movements
resentations of action goal and action effects (a common represen- such as posture control and movement trajectory planning. More
tational domain for perception and action) with the cognitive precisely, motor control theory assumes different internal models:
antecedent of the action (prospective coding in event representa- a forward internal model predicts the sensory consequences from
tion’; Schütz-Bosbach & Prinz, 2007). The main thesis is that fea- efference copies of issued motor commands; additionally, an
tures relevant for motor control are involved already at the level inverse model calculates the necessary feedforward motor com-
of action perception and the perception of action effects, with mands from the desired movement trajectory (cf. Kawato, 1999).
abundant empirical evidence, for example from classical studies To explain perception, especially of self-generated actions, forward
on spatial stimulus–response compatibility effects (e.g., Elsner & models are sometimes also simply referred to as ‘‘efference copy”
Hommel, 2001; Hommel, Musseler, Aschersleben, & Prinz, 2001; (von Holst & Mittelstaedt, 1950) or ‘‘corollary discharge” (Bell &
Kunde, 2001; see Shin, Proctor, & Capaldi, 2010 for a review). The Grant, 1989; Sperry, 1950). Following this interpretation, percep-
typical observation is an advantage (in terms of speed and accu- tion and action are essentially dependent on motor intentions
racy) of responses that share ‘‘motoric” features with a target stim- and related motor signals, such as the efference copy and predic-
ulus (such as the relative location in space), supposedly due to the tions computed on their basis. Extensive evidence for this depen-
automatic activation of a common location code between stimulus dency comes from studies on the phenomenon that people
and response. An elaborated version of the theory of event coding cannot tickle themselves, for which predictions have been attribu-
assumes a common format of (late) perception and (early) motor ted a prominent role by attenuating the sensory consequences of
control which are coded in a common representational medium, self-generated actions (Blakemore, Frith, & Wolpert, 1999). How-
i.e. event codes that are both motoric and perceptual, commonly ever, the internal model itself, despite being a decisive part of
coded, ‘‘especially if the features of perceived and to-be- motor control, is not identical with the motor mechanism, other-
produced events overlap. [. . .] In other words, perceptual and wise a comparison between a predicted and actual state for opti-
action-planning processes only interact if the codes they operate mizing motor control (e.g. fast online corrections of movement
on refer to the same (kind of) feature of a distal event” (p. 862’, trajectories) would not be possible. That is, motor control theory
Hommel et al., 2001). postulates that there is one input signal (i.e. desired action out-
In contrast to Simulation Theories (as in the case of mirror neu- come) and two different output processes (i.e. motor command
rons), Common Coding – at least in its original version – is not a and efference copy), and motor control is emphasized as a consti-
framework for understanding actions of others but mainly for tutive condition. Therefore, according to our suggested framework,
action planning. In contrast to Internal Model theories, action con- a constitution of perception by motor control mechanisms can be
trol in Common Coding does not draw on a forward signal which is found but this constitution is not exhaustive (i.e. there are also
sent in parallel with the motor command to guarantee optimal other factors involved). Hence, in motor control theory, some
action performance. Whereas Common Coding emphasizes percep- motor representations are considered partially constitutive at least
tion because it supposedly shapes motor control (at least with for parts of action cognition.
respect to distal events) Internal Model theories focus primarily Another type of Internal Model theory, predictive coding, is
on motor control. To summarize, Common Coding assumes three based on the view of perception as prediction (Friston et al.,
different classes of representational formats: purely motoric, 2010). In contrast to motor control theory, the primary focus of this
purely sensory and a common code format. Thus, neither motor approach is on perceptual models and not on motor control mech-
control is grounded in perception nor the other way round, but anisms. However, active inference as a corollary of this perspective
both are linked through additional (common code) representa- also aims to account for many aspects of motor behavior, from ocu-
tions. None of the Common Coding theories can therefore be con- lomotor reflexes to goal-directed movements. It assumes a gener-
sidered, from a constitution point of view, as a genuine grounding ative forward model which maps from motor control to sensory
theory. consequences, and which forms part of a larger hierarchical model
A. Gentsch et al. / Cognition 146 (2016) 81–89 87

used for perceptual inference in general. In contrast to motor con- strict correspondence between motor control and action percep-
trol theory, it is thought to serve all aspects of motor control with- tion. It should be noted that mirror neurons are regarded as neuro-
out the need for a separate inverse model to provide control signals physiological instantiations of simulation and, therefore, mirror
(cf Friston, 2005). In this respect, forward-inverse models in motor neuron theories provide a mechanistic explanation of the involved
control are formally distinct from predictive coding accounts of neuronal processes, rather than offering a theoretical account on
action perception. In predictive coding accounts, motor commands their own.
are not constructed by an inverse model but arise from an inver- Similarly, the motor imagery account states that motor per-
sion of the forward model. That is, instead of a mapping from con- cepts (or ‘‘motor images”) are ‘‘covert actions”, i.e. a simulation
trol signals to sensory consequences and vice versa, motor control of neural mechanisms and dynamics of real, physical action in
signals are identical with descending sensory predictions of propri- low level motor centers before an action is executed (Jeannerod,
oceptive sensations. However, beyond internal proprioceptive con- 2001). Here, Jeannerod (2001, p. 106) argues that ‘‘lesions affecting
sequences of movements, exteroceptive consequences of actions structures located ahead of motor system [. . .] affect the ability to
(e.g., vision, audition) are considered corollary discharges, as in produce a given S-state”. By the term S-state, Jeannerod refers to
classical forward models. Thus, at least for exteroceptive modali- ‘‘those ‘mental states’ which involve an action content and where
ties of action perception in predictive coding mechanisms, the brain activity can be shown to simulate that observed during the
same holds true as for motor control theories: some motor repre- same, executed action” (Jeannerod, 2001, p. 103). But these simu-
sentations are constitutive for parts of action cognition. lations are ‘‘covert actions” that do not result in muscular activity
Some types of Internal Model theories incorporate elements as an overt movement.
from Simulation theories. For example, the emulation theory, as Another prominent example of Simulation Theories is the the-
proposed e.g. by Rick Grush (2004), combines constructs like effer- ory of perceptual symbol systems (Barsalou, 1999). According to
ence copy and motor prediction with mirror mechanisms. It seeks this theory, cognition exclusively consists of activation patterns
to explain how motor control operations in classical motor centers of various sensory modalities, here the perceptual symbol system.
can produce motor imagery and certain aspects of visual percep- In this case cognitive and perceptual mechanisms share the same
tion by driving the operation of a motor-visual emulator of the representational states because cognitive processing is essentially
musculoskeletal system. Beyond the motor domain, the theory a simulation, i.e. a re-activation of sensory states, including senso-
holds that imagery in general is based on both amodal and rimotor activation patterns underlying the original perceptual
modality-specific systems driving parallel emulators as real time state. This reactivation of the same neural circuits involved during
models of behavior and perception. Hence, the theory has been perception of and action towards those objects allows the re-
applied not only as an explanatory model of motor control and enactment of multimodal information. Consequently, also higher-
motor imagery, but also of several other cognitive functions, indi- order abilities such as thinking or categorization are ‘‘grounded”
cating dissimilarity between motor processes simpliciter and cogni- in low-level sensorimotor abilities. Empirical evidence for the the-
tive functions. Accordingly, in Grush’s discussion of emulation ory of perceptual symbol systems indicates that perceptual and
versus simulation accounts of motor imagery, he insists that ‘‘mere motor mechanisms are engaged when a cognitive system performs
operations of the motor center is not enough; to produce imagery conceptual processing. It is suggested that, for example, talking or
the motor centers must be driving an emulator of the body” and ‘‘ thinking about objects actually (partly) consists of the reactivation
[a] motor plan is one thing, a sequence of proprioception and of previous experiences stored in the perceptual symbol system
kinesthesis is another” (p. 385’, Grush, 2004). Thus, according to (for an overview cf. Barsalou, 2008; de Vega, Glenberg, &
Grush, the executed action and its effects are not sufficient (and Graesser, 2008; Pecher & Zwaan, 2005; Semin & Smith, 2008).
maybe not even necessary) to install action-related perception Based on mirror neuron theories, even cognitive abilities such
and cognition. as mind-reading and understanding mental states of other’s by
Taken together, all types of Internal Model theories postulate observing their actions are often ‘‘explained” by simulation mech-
that action perception and cognition are significantly informed anisms that shall help ‘‘put oneself in the ‘mental shoes’ of others”
and construed by predictions. In particular, exteroceptive predic- (cf. Gallese & Goldman, 1998).
tions are not identical with motor commands but are grounded All variants of simulation theories understand sensorimotor
in the latter. However, more than motor commands is necessary processes as being primary (in contrast to Common Coding). More-
to produce these predictions, namely sensory feedback, context over, in contrast to Internal Models, they focus on higher order cog-
cues, and context estimates. Thus, predictions and related action nitive abilities to explain conceptual thinking and mind-reading
perception/cognition cannot be exhaustively grounded in motor underlying social understanding and social communication, rather
commands. Therefore, according to Internal Model theories, motor than focusing solely on motor control functions. Following Simula-
abilities would be (at most) partially constitutive of action-related tion Theories in general, cognitive abilities such as motor imagery,
perceptual and cognitive capacities. conceptual abilities or understanding the mental states of others
rely on a reactivation of sensorimotor states. Because simulation
4.3. Simulation Theories is essentially a reenactment of sensorimotor modalities there is
nothing over and above those sensorimotor mechanisms. Thus,
The family of Simulation Theories is characterized by the gen- simulation theories postulate an identity relation (i.e., full consti-
eral assumption that cognition is essentially simulation-like (re-) tution) between motor abilities and action cognition. A meaningful
use of the brain’s modal systems, such as the sensorimotor system. use of the notion of ‘grounded cognition’, however, can only be
For example, the mirror neuron theory assumes a population of based on a relation of constitution and not of an identity between
neurons coding for both observed and executed actions, that is, the relevant domains of action cognition, perception and motor
the same neural sub-system is used for representing motor acts control, as claimed by simulation theories.
and motor percepts (for review see, Rizzolatti & Craighero, 2004).
This applies specifically to so-called ‘strictly congruent’ mirror
neurons (Gallese, Fadiga, Fogassi, & Rizzolatti, 1996), encoding 5. Conclusions
the kinematics of an action, in contrast to some class of ‘broadly
congruent’ mirror neurons (or ’logically-related’ mirror neurons; Following the modular picture of the mind, cognition and action
e.g. Iacoboni et al., 2005) which encode action goals with a less has commonly been assumed to operate in strictly different
88 A. Gentsch et al. / Cognition 146 (2016) 81–89

domains: cognition draws on abstract conceptual representations, represented and shape one another. This may help to determine
whereas motor control functions are thought to rely on relatively adequate empirical approaches to core questions around embodied
low-level and automatic processes. In recent years, however, cog- human cognition. For example, to which extent can motor control
nitive neuroscience has been confronted with an increasingly large deficits or motor constraints affect the perception of other’s actions
array of partly overlapping, partly contradicting theories that and socio-cognitive functions at higher levels? How does motor
diverge broadly with respect to the degree to which cognition is expertise or training determine action-related perceptual content?
thought to be grounded in systems for perception and action. Con- Which aspects of self-awareness in the domain of action, such as
sequently, the proposed conceptual relations of motor control and feelings and judgments of self-agency, are reducible to certain
action cognition are manifold. This has led to apparent duplication motor control functions? The present framework of constitution
of experimental work, a relative huge collection of unsorted exper- and constitution relations opens a new perspective for specifying
imental findings and unconnected development of different theo- the relation between motor processes and action cognition and
ries. Recently, there have been some attempts comparing and may help to find the right balance between reductionist point of
linking different theories pairwise. For example, connections have views on the one hand and purely associative theories on the other
been made between theories belonging to the family of Internal hand. It may help to further discriminate the different domains of
Models and Simulation Theory (Kilner, Friston, & Frith, 2007), action perception and motor states as well as gain further insights
between Common Coding accounts and Simulation Theory in common factors and interdependences of different domains.
(Iacoboni, 2009), or between Common Coding and Internal Models
(Chambon & Haggard, 2013). However, up to date, a comprehen- Acknowledgments
sive overall framework able to accommodate all the theories is still
missing. We here argued that ‘‘grounded action cognition” investi- This project was supported by a grant of the Volkswagen Stif-
gating constitution conditions between motor processes and action tung to all of the authors (‘‘Grounding thoughts in actions, Thin-
cognition may provide a comprehensive metatheoretical frame- kAct’’; Az. II/85 068/85 158/85 159) and a Heisenberg fellowship
work that allows bringing together this heterogeneous mix of the- (SCHU 2471/4-1) of German Research Foundation/Deutsche
ories and to analyze and classify them according to the specific Forschungsgemeinschaft (DFG) to SS-B.
relation they claim to hold between motor processes and action
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