REVIEW

CURRENT
OPINION The neuroscience of action semantics in
neurodegenerative brain diseases
Thomas H. Bak

Purpose of review
Within the last decade, the neurobiology of action processing has moved from an obscure topic of
specialist interest to one of the most popular themes in modern neuroscience. However, the wealth of
literature and the diversity of approaches and theoretical models can make the field complex and, at times,
bewildering. This review presents the main currents of research, examining their theoretical underpinnings
in an interdisciplinary context.
Recent findings
The presence of specific deficits in verb and action processing has been documented in a wide range of
neurodegenerative diseases, including parkinsonian syndromes and motor neuron disease. Interestingly,
most of these disorders affect the motor system, suggesting a systematic relationship between motor
functions and their cognitive and linguistic representations. Action processing has been explored with a
whole spectrum of methodologies, from neuroimaging to transcranial and intracranial stimulation. The
findings have been integrated with other influential concepts and theories, including mirror neurons and
embodied cognition.
Summary
Converging evidence from patient and imaging studies links the concepts of actions and their processing
with the execution of actions through the motor system. The theory of embodied cognition remains
influential as well as controversial. However, the points of criticism have changed, reflecting recent
paradigm shifts.
Keywords
action semantics, embodied cognition, motor neuron disease, Parkinson’s disease, verb processing

INTRODUCTION MND and frontotemporal dementia (FTD) as polar

&&

When at the end of the last century Bak and Hodges
[1] described selective verb impairment patients
with motor neuron disease (MND), the results were
treated with disbelief. Firstly, selective deficits in the
processing of specific word classes, although already
well documented in single case studies of neurologi-
cal patients [2,3], have been perceived as a rarity.
Secondly, MND was seen at that time (despite
already existing strong evidence to the contrary
[4]) as the prime example of a purely motor disorder,
sparing higher cognitive functions. Sixteen years
later, at the time of this review, the landscape could
not be more different. Deficits in the processing of
verbs and actions are not only well recognized and
widely reported, but also form one of the major
arguments in support of the influential theories
&&
such as embodied cognition [5 ]. As for MND, it
became the prime example of a condition combin-
ing motor and cognitive features, with the classical
phenotypes of a single entity [6 ].
It would be tempting to believe that such a
profound change must have been brought about
by new evidence, with advanced, sophisticated
methodology leading to changes in theoretical
thinking. However, this review argues that the
major changes in our understanding of action proc-
essing are not a direct consequence of new empirical
findings. Empirical evidence for selective deficits in
verb and action processing is arguably amongst the
School of Philosophy, Psychology and Language Sciences (PPLS),
Centre for Clinical Brain Sciences (CCBS), University of Edinburgh,
Edinburgh, UK
Correspondence to Thomas H. Bak, Department of Psychology, Univer-
sity of Edinburgh, 7 George Square, Edinburgh, EH8 9JZ, UK. Tel: +44
131 6503441; fax: +44 131 6503461; e-mail: thomas.bak@ed.ac.uk
Curr Opin Neurol 2013, 26:671–677
DOI:10.1097/WCO.0000000000000039

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 Degenerative and cognitive diseases
KEY POINTS
 Specific deficits in action processing (with object
processing relatively preserved) are well documented in
neurological patients.
 They occur most frequently in association with the
disorders of motor functions, such as motor neuron
disease (MND), Parkinson’s disease and atypical
parkinsonian syndromes, such as progressive
supranuclear palsy (PSP).
 Accordingly, impairments in action semantics are
usually interpreted in the theoretical framework of
embodied cognition.
 Opposite pattern of deficit (impaired object, preserved
action semantics) has been reported in some patients
with semantic dementia, but the evidence is
less conclusive.
oldest in neuropsychology [7] and has accumulated
in a continuous and incremental way since the
1970s [8]. What has changed is the theoretical
framework into which such dissociations can be
integrated. Accordingly, we will present first the
major theoretical developments in the field, then
the current status of the empirical evidence and
finally the main points of criticism.
ACTION SEMANTICS IN THE CONTEXT OF
MODERN NEUROSCIENCE
The current research into action semantics in neuro-
degenerative diseases, in all its bewildering richness
and complexity, is very much a continuation of the
methods, approaches and concepts developed con-
tinuously over the previous decades. Hence, in order
to understand it, it is helpful to examine the main
underlying ideas and theories in current neuro-
science, into which the new insights from the study
of action deficits are being integrated. We will focus
on four major and interlinked developments in the
field: the shift from modules to networks, the emer-
gence of concepts of motor and embodied cognition
and the growing appreciation of the role played by
the mirror neurons. Although all have their roots
in the late 20th century, they have continuously
grown in importance to become the dominating
theories in modern cognitive neuroscience. All four
exert a strong influence on our understanding of
action semantics.
From modules to networks
Much of the current research in the neuropsychol-
ogy of actions has its roots in the late 20th century.
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At that time, the predominant paradigm was that
of modularism, proposing that different cognitive
functions are subserved by highly specialized,
encapsulated modules. With the advent of neuro-
imaging techniques, the modules, originally con-
ceived as functional units, became increasingly
identified with specific brain areas. The observation
of selective word class deficits in the 1970s and
1980s seemed to fit very well into this framework.
The presence of specific verb impairments in neuro-
logical, mainly stroke, patients was seen as a proof of
existence of a verb-processing module, usually local-
ized in the frontal lobes [2,3,8].
With the growing recognition of the import-
ance of brain connectivity and plasticity, the
static and somewhat rigid modules became replaced
by more complex and dynamic functional net-
works. This was accompanied by a gradual shift of
emphasis from stroke to neurodegenerative diseases
[3]. Whereas the majority of cases of verb impair-
ment in the late 20th century were stroke patients
[8], most of the recent examples come from
neurodegenerative conditions. This difference in
pathology has important implications. Unlike in
stroke, in which the pattern of deficits is dictated
by the vascular topography of the brain, neurode-
generative processes target functional systems [9].
Studies examining selective deficits in processing
of a certain word class or semantic category
assume, therefore, that the observed patterns can
tell us something about the underlying functional
systems.
Both modular and network approaches co-exist
in the current literature on action processing. A fair
number of studies set out to determine which areas
of the brain are associated with verb production and
comprehension [10,11]. However, an increasing
number of researchers take a different approach,
linking action processing not to a specific brain
region but to an interconnected network, often
including a subcortical as well as cortical com-
&&
ponent [12 ,13].
Motor cognition
Until the 1990s, neurodegenerative disorders of
movement and cognition tended to be viewed as
separate phenomena. Diseases such as Parkinson’s
disease or MND were classified as disorders of the
motor system. Accordingly, the cognitive status was
either not assessed at all or assessed only with the
Mini-Mental State Exam (MMSE), a singularly inap-
propriate cognitive screening tool in this context, as
it fails to assess precisely the most affected cognitive
domains [14]. On the other hand, there was equally
little interest in the motor aspects of cognitive
Volume 26  Number 6  December 2013

disorders/dementias, with the assessment often
confined to memory testing.
The radical change in perspective was brought
about by converging evidence from clinical obser-
vation, neuroimaging and basic sciences. Clinical
studies assessing cognition in the so-called move-
ment disorders found regularly the evidence of cog-
nitive dysfunction [15]. Imaging studies confirmed
the involvement of cortical structures in diseases
previously considered as subcortical [16]. Advances
in basic sciences demonstrated that the same
proteins (e.g. TDP-43) and genes (e.g. C9ORF72)
can be involved in classical ‘motor’ (e.g. MND) as
well as in classical ‘cognitive’ (e.g. FTD) diseases [17].
The term ‘motor cognition’ [18] has been intro-
duced to refer to the aspects of cognition involved
in planning, executing, understanding and even
imaging of actions. Accordingly, this type of cogni-
tive process is predominantly affected in ‘movement
disorders’. Hence, current studies do not need any
more to prove the existence of cognitive symptoms
in ‘motor’ disorders; instead, they elucidate their
frequency and severity, and, above all, determine
their exact nature [19]. This change in perspective
has had profound consequences for the field of
action semantics, as most deficits in action process-
ing have been described in diseases traditionally
classified as motor.
Embodied cognition
There are arguably few areas of modern science in
which ancient philosophical debates interact so
fruitfully with modern empirical findings as that
of embodied cognition. The relationship between
mind and body, and, consequently, between know-
ledge and action, has been for centuries a central
theme of human thought. In the West, the notion
of embodied cognition [20], with its roots going
back to Cato the Elder [21], can be found in the
thinking of Kant, Merleau-Ponty, Unamuno and
Heidegger, to name just a few. In the East, the idea
of the unity of knowledge and action, put forward
by the neo-confucian scholar Wang Yangming, had
a profound influence on China as well as Korea
and Japan [22].
The modern theory of embodied cognition pro-
poses that cognitive processes are rooted in the
experience of our body, in perception and in action.
Formulated by philosophers and linguists [23], it
was enthusiastically received by many disciplines.
For philosophers, it offers the possibility to practice
‘experimental philosophy’ and examine their
theories with the methods of modern neuroscience
[20]. For neuroscientists, it provides an appealing
and coherent framework to interpret studies of
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The neuroscience of action semantics Bak
&&
normal and abnormal action processing [5 ,24].
The strength of embodied cognition movement
comes from its interdisciplinary perspective and
the variety of employed methodologies: it brings
together converging evidence from behavioural and
neuroimaging studies, from studies in children and
adults, in healthy individuals and in patients with
neurological disorders. For instance, a seminal study
by Hauk et al. [25] demonstrated that listening to
action words related to specific body parts (e.g. ‘kick’
to legs or ‘lick’ to mouth) modulates the activity in
the areas of the brain related to the movement of the
respective body parts. Another iconic imaging study
from the same period, comparing capoeira and clas-
sical ballet dancers [26], showed that brain acti-
vation when observing dance movements depends
on the personal experience of performing the dance.
Subsequent studies explored the functional import-
ance of such phenomena. A behavioural study in
adults showed that movements of hands and
feet interact with the processing of related action
words [27]. A study in children demonstrated that
imitating movements improves learning of related
new verbs [28]. Only few studies failed to find
&
motor–cognitive interactions [29 ], raising the
question whether a confirmation bias might work
in favour of the embodiment theory. Indeed, so
great is the popularity of the embodied cognition
that an exponential growth of articles makes the
field increasingly difficult to follow. Several recent
reviews attempt to offer a systematic overview
&& & &&
[5 ,30,31 ,32 ].
Mirror neurons
The first description of mirror neurons in the 1990s
set in movement a whole chain of discoveries con-
tinuing to this day and establishing the mirror
neuron system [33] as one of the most relevant
factors for language development [30,34] and action
understanding [35,36]. The crucial feature of mirror
neurons is that they fire when a certain object is
grasped not only by the experimental individual,
but also by other individuals. It provides, therefore,
a possible neural mechanism explaining the close
relationship between performing, observing and
understanding an action. In recent years, the mirror
neurons have been particularly often used as an
argument for embodied cognition in the context
of the evolution of language and the emergence of
concepts out of actions. However, although the
discovery of the mirror neurons has been undoubt-
edly a major achievement of modern neuroscience,
the popularity of the concept has led to its extensive
and often speculative use well beyond the empirical
evidence available.
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 Degenerative and cognitive diseases
ACTION DEFICITS IN
NEURODEGENERATIVE DISEASES
The neuroimaging studies presented above demon-
strate beyond reasonable doubt the involvement of
‘motor’ areas of the brain in the processing of action
words and action concepts. However, is this acti-
vation just an incidental co-activation of intercon-
nected areas or is it also functionally relevant? Do we
really need the motor system for action semantics?
This question has indeed been posed by the critics of
embodied cognition [37]. It is in this context that
patient studies assume a crucial role. The finding of
an action deficit associated with a motor disease
would provide a powerful argument in favour of a
functionally relevant connection between action
knowledge and its motor execution. Current
research in this field focusses on two major groups
of neurodegenerative diseases: MND and parkinso-
nian syndromes.
Motor neuron disease
The idea that verb and action deficits in motor
disorders are a logical extension of motor disorder
into the area of higher level representation of
actions was first formulated in the context of
MND [1,38–40]. It linked clinical observations in
MND patients with the data from neuroimaging
studies, suggesting a direct involvement of motor
cortex in the processing of action words [25]. Both
strands of evidence have been strengthened through
&
the recent studies: Taylor et al. [41 ] confirmed the
presence of action deficits in a large cohort of MND
patients, whereas a magnetoencephalography study
&
by Boulenger et al. [42 ] showed motor cortex
involvement in the processing of action-related
idioms. Moreover, recent models of the spread of
neurodegeneration along reinforced synaptic con-
nections (what fires together dies together) provide
a possible pathological mechanism for the involve-
ment of higher level action representations in a
&&
disease of the motor system [6 ].
Parkinson’s disease and related disorders
The second group of diseases in which action proc-
essing has been systematically studied in recent
years is Parkinson’s disease and related disorders.
The relatively high prevalence of Parkinson’s dis-
ease, the better general health of the patients and
the opportunity of therapeutic interventions allow
Parkinson’s disease-based studies to address ques-
tions that would be difficult to explore in the con-
&
text of MND. Fernandino et al. [43 ] addressed the
perennial debate between the grammatical versus
semantic/conceptual deficit by comparing action
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and abstract verbs in Parkinson’s disease patients.
The selective impairment of action but not abstract
verbs is interpreted as an indication of an action-
semantic deficit. The same research group compared
literal and idiomatic action-related expressions
&
in Parkinson’s disease [44 ], and found that both
were equally affected, strengthening the idea of
an involvement of motor representation in the
processing of action-related sentences. A promising
new type of experiment examines the effect of
therapeutic interventions in Parkinson’s disease
&&
on action processing. Silveri et al. [45 ] found
that stimulation of the subthalamic nucleus in
Parkinson’s disease patients improved action
naming more than object naming. Herrera et al.
& &&
[46 ] and Péran et al. [12 ] compared action process-
ing in on and off stages of dopaminergic medication,
finding a specific positive effect of medication on
verb processing and thus a support for the motor
theory of verb knowledge. The work of Cardona
&&
et al. [13] and Ibáñez et al. [47 ] examined patients
with Parkinson’s disease as well as with epilepsy (the
latter being examined with intracranial recordings)
and found evidence of action–sentence compatibil-
ity effect in both conditions. Selective action proc-
essing deficits have also been described in other
diseases from the parkinsonian spectrum, such as
&
progressive supranuclear palsy (PSP) [48 ] and corti-
cobasal degeneration (CBD) [49].
Semantic dementia as a necessary counter-
example
The evidence for the presence of action processing
deficits in motor disorders is overwhelming and
unlikely to be entirely attributable to artefacts. How-
ever, it could be that the verbs and actions are
genuinely more complex and difficult, and there-
fore more prone to be affected by any type of brain
disease. In order to claim a specific association
between action processing and motor system, we
need to show that neurodegenerative diseases that
do not target motor functions are not associated
with predominant verb impairment (or might even
lead to predominant involvement of nouns and
objects). So far, such type of evidence has been much
weaker than that for verb and action deficits. In stark
contrast to the large number of diseases character-
ized by the action deficits, only one so far has been
repeatedly associated with predominant noun and
object deficits and relative preservation of verbs
and actions: semantic dementia. Using a specially
designed test of action associations (Kissing and
Dancing Test; KDT) and comparing it to the estab-
lished Pyramid and Palmtrees Test (PPT), Bak and
Hodges [50] found a dissociation between patients
Volume 26  Number 6  December 2013

with behavioural variant FTD and semantic demen-
tia. Behavioural variant FTD patients were more
impaired on actions than on verbs – a similar
pattern has been documented using the same test
in patients with primary progressive aphasia [51],
&& &
Parkinson’s disease [47 ] and MND [41 ]. In con-
trast, the semantic dementia patients showed an
opposite pattern: a slightly better performance on
actions than objects. A similar result (an action
advantage in semantic dementia) has been con-
&
firmed in a more recent study [52 ]. What is needed,
however, is a large, systematic comparison of
semantic dementia patients with those with dis-
orders of the motor system.
LIMITATIONS, CRITICISMS AND NEW
DIRECTIONS
How far can we take embodied cognition? Does the
involvement of the motor system when confronted
with an action word spread as far as actual muscle
activity or the relevant muscular units? A recent
study using electromyography (EMG) found that
reading verbs did not lead to a subliminal muscle
& &
activity [53 ]. In contrast, Foroni and Semin [54 ]
using the same technique of EMG found that sen-
tences negating actions (e.g. ‘I am not smiling’) lead
to an inhibition of the respective muscles, a result
well in line with the embodied theories of language.
The Bayesian challenge
Until recently, the main criticisms of the theories
of embodied cognition came from two directions.
One drew attention to potential confounding
variables, unequally distributed between verbs and
actions and nouns and objects: frequency, image-
ability, age of acquisition, word length, syntactic
and morphological and syntactic complexity, etc.
With an exponentially growing number of studies
paying increasing attention to careful matching of
variables, it becomes less and less likely that all
the reported effects could be because of confound-
ing. The second type of criticism has its roots in
the modularist view of the representation of word
classes. Although such views continue to be
expressed by prominent figures in the field
&
[10,37,55 ], the described paradigm shift from
modules to networks [9] favours the interpretations
rooted in the theory of embodied cognition.
However, the comfortable dominance of
embodied cognition might soon get challenged
from the opposite direction: the increasingly pop-
ular Bayesian models of cognition. Fundamentally
opposed to modularism and seen by some as a
reincarnation of behaviourism through modern
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The neuroscience of action semantics Bak
computers, Bayesian approach assumes that cogni-
tive processing is essentially based on statistical
learning and estimation of probability. De Zubicaray
&
et al. [56 ] achieved an activation of the motor
cortex through the presentation of nonwords. They
argue that such an effect of nonwords cannot be
attributable to semantics and propose instead that it
reflects implicit processing of ortho-phonological
statistical regularities. However, their argument is
critically dependent on the assumption that non-
words do not evoke semantic associations, a view
not shared by all researchers.
Language revisited (and revived?)
The early descriptions of verb deficits tended to
interpret them through the lens of language, think-
ing in terms of word-class differences and grammar.
The following decades have seen a continuous shift
in emphasis from verbs as parts of speech to actions
as part of the general cognitive (and indeed motor)
system. The last years have witnessed a revival of
interest in linguistic characteristics and distinctions,
such as negation, tense or inflections. The role of
negation, already mentioned in the context of the
&
study by Foroni and Semin [54 ], has been explored
further by Liuzza et al. [57] with the help of neuro-
imaging and transcranial magnetic stimulation
(TMS) – brain responses were modulated by affir-
mative and negative sentence structures, respect-
&
ively. Pulvermüller et al. [58 ] found that the
pattern of cerebral activation was determined by
the morphological cues of the inflectional endings
as much as by the semantic properties. Another
aspect, widely researched in psycholinguistics but
so far not applied to action semantics, is that of
&
perspective taking [59 ]. Finally, one of the recent
foci of interest is the relationship between the
impairment of verb and action processing in
brain-damaged patients and the acquisition of verb
&
and action understanding in infants [60 ] – which
brings us back to the very beginning of our journey,
the 1744 book by Vico [7], in which the selective
impairment of verbs in a stroke patient is interpreted
in the context of verb acquisition in infancy.
CONCLUSION
Within the last decade, the field of action semantics
moved from an outsider area to one of the most
popular and prolific topics in modern cognitive
neuroscience. Abundant evidence from behavioural
and neuroimaging studies points to a close relation-
ship between the conceptual representation of
actions in the brain and their execution by the
motor system. Action words have been shown to
www.co-neurology.com 675
 Degenerative and cognitive diseases
influence the activity of motor areas of the brain and
to interact and interfere with the performance of
actions themselves. The findings tend to be inter-
preted within the broader framework of embodied
cognition, often with reference to the mirror
neurons.
Specific deficits in action processing have been
well documented in neurodegenerative diseases
characterized by a combination of motor and cog-
nitive impairment, above all MND and Parkinson’s
disease. This strong association between motor dis-
orders and deficits in action–cognition suggests a
functional importance of the motor involvement in
action processing, thus supporting the notion of
embodied cognition. However, whereas action
deficits with relatively intact object processing are
well documented, the opposite pattern (object
deficits with relatively preserved action processing)
has only been reported in a handful of semantic
dementia patients.
The embodied cognition movement has been
extraordinarily successful in bringing together
different disciplines, such as linguistics, philosophy,
psychology, neurology, computer science and
neuroscience. It has dealt successfully with the
criticism from the modularist school of thought
and seems well equipped to deal with the emerging
Bayesian challenge. However, in view of this wide
consensus and agreement, more controversy and
debate could indeed be beneficial.
Acknowledgements
The author would like to thank Friedemann Pulvermuller
and Agustin Ibanez for fruitful discussions, and
Blanca Ramirez Ruiz for her advice and help with the
manuscript.
Conflicts of interest
There are no conflicts of interest.
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