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THE LINGUISTIC
CEREBELLUM

Edited by

PETER MARIËN
Department of Neurology and Memory Clinic, ZNA
Middelheim General Hospital, Antwerp, Belgium; and
Clinical and Experimental Neurolinguistics (CLIN), Vrije
Universiteit Brussel, Brussels, Belgium

and

MARIO MANTO
Unité d’Etude du Mouvement
Université Libre de Bruxelles,
Brussels, Belgium; and Université de
Mons, Mons, Belgium

Amsterdam • Boston • Heidelberg • London

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CONTRIBUTORS

Hermann Ackermann
Department of Neurology and Stroke, Hertie Institute for Clinical Brain Research,
University of Tübingen, Tübingen, Germany
Michael Adamaszek
Department of Neurologic and Cognitive Rehabilitation, Bavaria Clinic Kreischa,
Kreischa, Germany
Louise Allen-Walker
School of Psychology, Bangor University, Wales, UK
Georgios P.D. Argyropoulos
Cognitive Neuroscience and Neuropsychiatry Section, Developmental Neurosciences
Programme, Institute of Child Health, University College London, London, UK
Lauren A. Barker
The Chicago School of Professional Psychology, Loyola University Chicago, Chicago, IL, USA
Lisa Bartha-Doering
Department of Pediatrics and Adolescent Medicine, Medical University Vienna,Vienna,
Austria
Alan A. Beaton
Department of Psychology, Aberystwyth University, Wales, UK; Department of Psychology,
Swansea University, Wales, UK
Florian Bodranghien
Laboratoire de Neurologie Expérimentale–ULB, Brussels, Belgium
R. Martyn Bracewell
School of Psychology, Bangor University, Wales, UK; School of Medical Sciences, Bangor
University, Wales, UK
Hyo-Jung De Smet
Clinical and Experimental Neurolinguistics (CLIN),Vrije Universiteit Brussel, Brussels,
Belgium
John E. Desmond
Departments of Neurology, Neuroscience and Cognitive Science, Johns Hopkins
­University School of Medicine, Baltimore, MD, USA
Thora Gudrunardottir
Oncological Department, Hilleroed Hospital, Copenhagen, Denmark; Posterior Fossa Society
Christophe Habas
Service de NeuroImagerie, Hôpital des Quinze-Vingts, Université Pierre et Marie Curie,
Paris, France
Ingo Hertrich
Department of Neurology and Stroke, Hertie Institute for Clinical Brain Research,
University of Tübingen, Tübingen, Germany
xi
xii Contributors

Laura Jansons
Private Practice, Arlington Heights, IL, USA
Kenneth C. Kirkby
Department of Psychiatry, School of Medicine, University of Tasmania, Hobart, Australia
Leonard F. Koziol
Private Practice, Park Ridge, IL, USA
Maria Leggio
I.R.C.C.S. Santa Lucia Foundation, Rome, Italy; Department of Psychology Sapienza
University of Rome, Rome, Italy
Mario Manto
Unité d’Etude du Mouvement, Université Libre de Bruxelles, Brussels, Belgium;
Université de Mons, Mons, Belgium
Peter Mariën
Department of Neurology and Memory Clinic, ZNA Middelheim General Hospital,
Antwerp, Belgium; Clinical and Experimental Neurolinguistics (CLIN),Vrije Universiteit
Brussel, Brussels, Belgium
Cherie L. Marvel
Departments of Neurology and Psychiatry, Johns Hopkins University School of Medicine,
Baltimore, MD, USA
Klaus Mathiak
Department of Psychiatry, Psychotherapy and Psychosomatics, University Hospital
Aachen, RWTH Aachen University, Aachen, Germany; Jülich-Aachen Research Alliance,
JARA-Brain, Jülich, Germany
Marco Molinari
I.R.C.C.S. Santa Lucia Foundation, Rome, Italy
Philippe Paquier
Clinical and Experimental Neurolinguistics (CLIN),Vrije Universiteit Brussel, Brussels,
Belgium; Department of Neurology and Neuropsychology, University Hospital Erasme,
ULB, Brussels, Belgium; Unit of Translational Neurosciences, School of Medicine and
Health Sciences, Universiteit Antwerpen, Antwerp, Belgium
Jeremy D. Schmahmann
Ataxia Unit, Cognitive Behavioral Neurology Unit, Laboratory for Neuroanatomy and
Cerebellar Neurobiology, Department of Neurology, Massachusetts General Hospital and
Harvard Medical School, Boston, MA, USA
Catherine J. Stoodley
Department of Psychology and Center for Behavioral Neuroscience, American University,
Washington, DC, USA
Kim van Dun
Clinical and Experimental Neurolinguistics (CLIN),Vrije Universiteit Brussel, Brussels,
Belgium
 Contributors xiii

Dorien Vandenborre
Cepos, Rehabilitation Centre, Rooienberg, Duffel, Belgium; Clinical and
Experimental Neurolinguistics (CLIN),Vrije Universiteit Brussel, Brussels, Belgium
Jo Verhoeven
Language and Communication Science, City University London, Northampton Square,
London, UK; Computational Linguistics & Psycholinguistics Research Center (CLiPS),
University of Antwerp, Antwerp, Belgium
Wolfram Ziegler
EKN—Clinical Neuropsychology Research Group, Clinic for Neuropsychology, City
Hospital Munich, Munich, Germany; Institute for Phonetics and Speech Processing,
Ludwig-Maximilians-University Munich, Munich, Germany
INTRODUCTION

Peter Mariën1,2, Mario Manto3,4

1
Department of Neurology and Memory Clinic, ZNA General Hospital, Middelheim, Antwerp, Belgium;
2
Clinical and Experimental Neurolinguistics (CLIN),Vrije Universiteit Brussel, Brussels, Belgium;
3
Unité d’Etude du Mouvement, Université Libre de Bruxelles, Brussels, Belgium; 4Université de Mons,
Mons, Belgium

In less than three decades, the concept of “cerebellar neurocognition”

has evolved from a mere afterthought to an entirely new and multifac-
eted area of neuroscientific research. A close interplay among three main
strands of contemporary neuroscience has induced a substantial modifi-
cation of the traditional view of the cerebellum as a simple coordinator
of autonomic and somatic motor functions. Indeed, the wealth of cur-
rently available evidence derived from (1) detailed neuroanatomical
investigations, (2) functional neuroimaging studies with healthy subjects
and patients, and (3) in-depth neuropsychological assessment of patients
with cerebellar disorders shows that the cerebellum plays also a cardinal
role in affective regulation, cognitive processing, and linguistic functions.
However, although considerable progress has been made in models
of cerebellar function, controversy remains regarding the exact role of the
“linguistic cerebellum” in a broad variety of nonmotor language processes.
This volume brings together a range of different viewpoints and opin-
ions regarding the contribution of the cerebellum to language function.
Recent developments and insights in the nonmotor modulatory role of
the cerebellum in language and some related disorders are discussed by
experts in the field. The role of the cerebellum in speech and language
perception, in motor speech planning including apraxia of speech, in ver-
bal working memory, in phonological and semantic verbal fluency, in
syntax processing, in the dynamics of language production, in reading,
and in writing will be addressed. In addition, the functional topography
of the linguistic cerebellum and the contribution of the deep nuclei to
linguistic function will be discussed. As such, with this volume, we hope
to offer a framework for debate and discussion. The reader interested in
the neuroscientific mysteries of this organ situated at the bottom of the
brain will find not only state-of-the-art contributions, but also novel
ideas that are being investigated in a growing number of laboratories
worldwide. The fact that the cerebellum contains more neurons than any

xv
xvi Introduction

other region of the brain and is characterized by a geometrical structure

makes it an excellent candidate for investigation of novel concepts in
neuroscience.

DEVELOPMENT OF CONCEPTS
Two centuries of research on cerebellar function have been dominated by
the role of the cerebellum in motor control (see Manto et al., 2012 for a
review). However, from time to time, clinical case descriptions and experi-
mental evidence from animal studies dating back to the early part of the
nineteenth century, already suggested an association between cerebellar
pathology and a variety of nonmotor cognitive as well as affective dysfunc-
tions (see Schmahmann, 1991, 1997). As early as 1831, Combettes described
in the Bulletins de la Société Anatomique de Paris, the postmortem findings
of an 11-year-old girl, Alexandrine Labrosse, who presented with neurode-
velopmental disorders including a range of cognitive, affective, and motor
impairments resulting from a complete absence of the cerebellum (Figure 1).
Nevertheless, a causal connection between cerebellar disease and cogni-
tive and affective disturbances was dismissed for decades. In the mid-1900s,
investigators started to examine a possible link between the cerebellum and
cognition and emotion, exemplified by the work of Snider and Eldred
(1948), Snider (1950), Snider and Maiti (1976), Dow (1974), Heath (1977,
1997), Heath, Franklin, and Shraberg (1979), Cooper, Riklan, Amin, and
Cullinan (1978) and others (see Schmahmann, 1991 for a review). This laid
a foundation for the rediscovery of this concept by Leiner, Leiner, and Dow
(1986, 1991), who hypothesized that more recently evolved parts of the
cerebellum contribute to learning, cognition, and language, and by
­Schmahmann (1991) and Schmahmann and Pandya (1987, 1989), who
introduced the dysmetria of thought hypothesis (Schmahmann, 1998).
These authors provided a historical, clinical, neuroanatomical, and theoreti-
cal framework within which a cerebellar role in higher cognitive and affec-
tive processes could be considered.That there may be a correlation between
the size of the cerebellum and aspects of general intelligence has been
known for some time (e.g., Allin et al., 2001; Ciesielski, Harris, Hart, &
Pabst, 1997; Mostofsky et al., 1998; Paradiso, Andreasen, O’Leary, Arndt, &
Robinson, 1997). From an evolutionary perspective, MacLeod, Zilles,
Schleicher, Rilling, and Gibson (2003) demonstrated a reliable linear regres-
sion contrast between volumes of whole brain, cerebellum, vermis, and
hemisphere of hominoids and monkeys and a striking increase in the lateral
cerebellum in hominoids (Beaton & Mariën, 2010). After controlling
 Introduction xvii

statistically for age and sex, Pangelinan et al. (2011) showed with school-
aged children that total cerebellar volume correlates significantly with cog-
nitive ability (as measured by overall intelligence quotient) (but see Parker
et al. (2008) for negative findings). Posthuma et al. (2003) reported that

Figure 1 Early description by Combettes (1831) of an 11-year-old girl with a complex of

cognitive, affective and motor developmental disturbances due to agenesis of the cerebel-
lum.

cerebellar volume in healthy adults (as well as total cerebral grey and white
matter volumes) correlates with working memory performance. Such find-
ings make it difficult to deny that the cerebellum is “an organ of cognition”
(Justus & Ivry, 2001).
xviii Introduction

SCHMAHMANN SYNDROME
Only a few years after the introduction of the dysmetria of thought theory,
Schmahmann and Sherman (1998) described in a seminal study of patients
with focal cerebellar lesions a consistent pattern of cognitive and affective
deficits and coined the term “cerebellar cognitive affective syndrome” to
describe this condition. Schmahmann syndrome, the eponym of cerebellar
cognitive affective syndrome (Manto & Mariën, 2015), was characterized as a
cluster of multimodal disturbances including: (1) executive deficits (deficient
planning, set-shifting, abstract reasoning, working memory, and decreased ver-
bal fluency), (2) disruption of visuospatial cognition (visuospatial disorganiza-
tion and impaired visuospatial memory), (3) personality changes (flattening or
blunting of affect, and disinhibited or inappropriate behavior), and (4) a range
of linguistic impairments among which were dysprosodia, agrammatism, and
mild anomia. However, analysis of the clinical data revealed that not all deficits
occurred in each patient, but that certain symptoms were particularly promi-
nent. Decreased verbal fluency, which did not relate to dysarthria, was said to
be present in 18 of the 20 patients.Visuospatial disintegration, mainly consist-
ing of disruption of the sequential approach to drawing and conceptualization
of figures was found in 19 cases. Eighteen of the 20 patients presented with
executive dysfunctions involving working memory, motor, and mental set-
shifting and perseverations of actions and drawing. In 15 patients, frontal-like
behavioral and affective changes were evident. Flattening of affect or disinhi-
bition occurred, taking the form of overfamiliarity, flamboyant and impulsive
actions, and humorous but inappropriate comments. Behavior was character-
ized as regressive and child-like in some cases and obsessive–compulsive traits
were occasionally observed. Deficits in mental arithmetic were evident in 14
patients. Visual confrontation naming was impaired in 13 patients. Eight
patients developed abnormal prosody characterized by high-pitched, whin-
ing, and a hypophonic speech quality. Mnestic deficits (verbal and visual learn-
ing and recall) were observed in some cases.The cluster of symptoms defining
Schmahmann syndrome was associated with a decrease of general intellectual
capacity. From an anatomoclinical perspective, cognitive and affective impair-
ments were more prominent and generalized in patients with large, bilateral,
or pancerebellar disorders, especially in a context of an acute onset of cerebel-
lar disease. Posterior lobe damage was particularly important in the genesis of
this novel syndrome. Damage of the vermal regions was consistently present
in patients with disruption of affect. Anterior lobe damage was found to be
less important to cause cognitive and behavioral deficits. Schmahmann
 Introduction xix

syndrome in patients with stroke improved over time, but executive function
remained abnormal. ­Schmahmann and Sherman (1998) pointed out that on
the basis of their observations, it was not possible to distinguish the contribu-
tion of the lesioned cerebellum to these abnormal behaviors from that of the
cerebral regions newly deprived of their connections with the cerebellum.
Indeed, the clinical features of the cognitive and affective impairments consti-
tuting Schmahmann syndrome are identical to those usually identified in
patients with supratentorial lesions affecting the cortical association areas and
paralimbic regions and their interconnections. Reciprocal connections link-
ing the cerebral association areas and paralimbic regions with the cerebellum
constitute the neuroanatomical basis to explain the pathophysiological mech-
anisms of the cerebellar induced cognitive and affective deficits. As pointed
out by Schmahmann and coworkers in an influential series of neuroanatomi-
cal studies, the cerebrocerebellar anatomical circuitry consists of a feedforward
limb (the corticopontine and pontocerebellar pathways) and a feedback limb
(the cerebellothalamic and thalamocortical systems) reciprocally connecting
the cerebellum with the supratentorial regions crucially implicated in cogni-
tive and affective processing. Since then, cerebellar involvement in linguistic
processes has been studied by advanced neuroimaging methods in healthy
subjects and several studies have been published focusing on a variety of lin-
guistic dysfunctions following cerebellar lesions of different etiologies in chil-
dren as well as adults. Reviews of the role of the cerebellum in nonmotor
language functions are provided by Gordon (1996), Mariën, Engelborghs,
Fabbro, and De Deyn (2001), Mariën et al. (2014), Paquier and Mariën (2005),
De Smet, Baillieux, De Deyn, Mariën, and Paquier (2007), Beaton and
Mariën (2010), Murdoch (2010), Highnam and Bleile (2011) and De
Smet, Paquier, Verhoeven and Mariën (2013).
With the introduction of Schmahmann syndrome, neuroscientists deal-
ing with the cerebellum have now a better idea of the topography of cere-
bellar deficits. Maps of lesion symptoms have been identified, clarifying the
roles of cerebellar lobules in motor, cognitive, or behavioral operations.

THE SUBSERVING NEURAL NETWORK

The neuroanatomical substrate of the recently acknowledged nonmotor
role of the cerebellum in cognitive and affective processing is a dense and
reciprocal network of crossed cerebrocerebellar pathways consisting of
corticopontocerebellar and cerebellothalamocortical loops that establish a
close connection between the cerebellum and the supratentorial motor,
xx Introduction

paralimbic, and association cortices subserving cognitive and affective pro-

cesses. Several contemporary lesion-behavior and neuroimaging studies
have demonstrated that, in addition to its somatotopic organization for
motor control, the human cerebellum is topographically organized for
higher order cognitive and affective functions as well. A meta-analysis of
neuroimaging studies investigating cerebellar involvement in motor, cogni­
tive, and affective processing paradigms has provided support for a dichot­
omy between the sensorimotor cerebellum—­geographically organized in
distinct regions in the anterior lobe—and the neurocognitive and affective
cerebellum—represented in distinct parts in the posterior lobe (for a review,
see Stoodley & Schmahmann, 2009). In addition, the majority of anatomo-
clinical studies of patients with linguistic impairments following focal cer-
ebellar lesions and the majority of neuroimaging studies employing
nonmotor language tasks typically show a lateralized involvement of lateral,
posterior cerebellar regions (including lobules VI and Crus I/II) in nonmo-
tor linguistic processes. Indeed, the patterns of lateralized (or even bilateral)
cerebral representation of language in dextrals and sinistrals seem to be
reflected at the cerebellar level by a “lateralized linguistic cerebellum,” sub-
served by crossed cerebellocerebral connections between the cortical lan-
guage network and the cerebellum.

THE THEORETICAL FOUNDATION

The theory that the cerebellum operates as an essential modulator of higher
level cerebral functions, including language and affect, currently attracts much
attention of the scientific community, yet no consensus exists about the exact
role and contributions of the cerebellum to the cognitive and affective domain.
Because of its uniform neuroanatomical structure and its dense connections
with the supratentorial association areas via cerebrocerebellar pathways (corti-
copontine-pontocerebellar-cererebellothalamic-thalamocortical), the cerebel-
lum is considered a functional entity that contributes in a unique and general
way to information processing (universal cerebellar transform; Schmahmann,
2000, 2001, 2004, 2010; Stoodley & Stein, 2013). One of the computational
models that supports this view on integrated cerebellar motor, cognitive, and
affective function is the dysmetria of thought theory that regards the cerebel-
lum as an oscillation dampener (realizing a maintenance of functioning around
a homeostatic baseline, to smooth out performance in all domains: cognitive,
motor, and emotional) (Schmahmann, 2000, 2001, 2004, 2010). Several
authors (Courchesne & Allen, 1997; Desmond, Gabrieli,
Wagner, Ginier, & Glover, 1997; Stoodley & Stein, 2013) regard the
 Introduction xxi

computational contribution of the cerebellum to motor, cognitive, and affec-

tive function as a predictor of future states.This functional role implicates that
the cerebellum generates internal neural “forward” models by means of opti-
mization of motor programs and mediation of cognitive functioning. In a
number of studies, Ito (2008) suggested that the corticonuclear microcom-
plexes of the cerebellum function as learning machines, performing a com-
parator role consisting of the formation and updating of internal models
through error learning (error predictions, processing, and correction). Accord-
ing to others, the cerebellum may act as an “internal clock” crucially involved
in the detection of deviations of an expected timing (control and regulation of
motor and cognitive functions) (Ivry, 1997; Ivry, Spencer, Zelaznik, & Died-
richsen, 2002) or as a detector of change and deviations of sequential events
(Ackermann, 2008; Leggio, Chiricozzi, Clausi, Tedesco, & Molinari, 2011;
Molinari, Chiricozzi, Clausi, Tedesco, De Lisa, & Leggio, 2008; Stoodley
et al., 2009; Stoodley,Valera, & Schmahmann, 2011). In other studies, the role
of the cerebellum is described as a control mechanism of shifts of attention,
priming and boosting activity in the extracerebellar system to operate rapidly
and efficiently (Courchesne & Allen, 1997; Salmi et al., 2009). More research
is needed to elucidate the theoretical underpinnings of cerebellar neurocogni-
tion. An integrated vision of the theoretical conceptualizations of the general
contribution of the cerebellum in cognitive functioning might be that of a
high-level operational device that does not subserve a specific cognitive or
affective function in itself but rather lends an active support to the central pro-
cesses in a variety of ways including prediction of the consequences of a motor,
affective or cognitive action or by error detection (in a sequence or in time).
The primary goal of this volume is to collect and summarize the key con-
cepts that have been proposed to explain the role of the cerebellar circuits in
linguistic processing.To this aim, we have gathered contributions from several
leading experts in various areas of cerebellar language processing, providing a
range of different, sometimes even controversial, viewpoints. As demonstrated
in this book, several dimensions of speech and language production and per-
ception seem to depend upon the modulatory role of the cerebellum: pho-
netic timing operations, auditory signal segregation and cross-modal binding
mechanisms, adaptive sensorimotor function of speech motor planning, artic-
ulatory control processes in verbal working memory, nonautomatic sequence
strategies in verbal fluency, temporal coordination and recall of established
explicit internal representations of sentence structures, high-level language
functions (e.g., figurative language, word association, antonym/synonym gen-
eration), planning and execution of manual production of letters, and reading
(development). Nonmotor linguistic function of the human cerebellum
xxii Introduction

seems to be topographically organized in a “lateralized linguistic cerebellum”

which reflects the patterns of cerebral language dominance in dextrals and
sinistrals. In the majority of dextrals the “lateralized linguistic cerebellum”
comprises, the right ventrocaudal part of the dentate nucleus and the right
posterior lateral cerebellum (involving lobules VI–VIII) that is reciprocally
linked to the supratentorial language networks of the dominant hemisphere.
However, anatomical studies providing direct evidence of reciprocal cerebel-
lar connections to language areas are still lacking. Although final agreement
has not yet been reached, we believe that a new consensus that draws on and
integrates the ideas presented here will eventually emerge to unravel the enig-
matic role of the cerebellum in nonmotor linguistic processing. At a time
where the field of ataxiology is being refined and keeps growing in popularity,
it is our hope that readers will find this book informative and useful.
We are particularly grateful to all the contributors who have found time
to prepare excellent chapters despite a busy schedule.We are thankful to the
anonymous reviewers for their constructive input and the staff of Elsevier
for their permanent support in the preparation of this book.

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CHAPTER 1

The Phonetic Cerebellum:

Cerebellar Involvement in Speech
Sound Production
Wolfram Ziegler1,2
1EKN—Clinical Neuropsychology Research Group, Clinic for Neuropsychology, City Hospital Munich,

Munich, Germany; 2Institute for Phonetics and Speech Processing, Ludwig-Maximilians-University Munich,
Munich, Germany

INTRODUCTION
The processes dealing with the production of speech sounds for words and
sentences are commonly broken down into several subcomponents.
Depending on which theoretical framework is chosen, different process-
ing stages are distinguished. A prominent psycholinguistic model devel-
oped by Levelt, Roelofs, and Meyer (1999), for instance, makes a distinction
between four discrete processing steps by which (1) word forms are
retrieved from a mental lexicon, (2) the phonemes constituting a word are
mapped onto the word’s rhythmical frame (“phonological encoding”), (3)
motor plans are retrieved for syllabified phonological forms (“phonetic
planning”), and (4) these motor plans are implemented in the neuromus-
cular system controlling speech movements (“articulation”). Otherwise,
models focusing on the lower levels of speech production are less elabo-
rate regarding abstract lexical and phonological processes, but are more
concerned with control subsystems of the speech motor apparatus (e.g.,
Guenther, Ghosh, & Tourville, 2006; Hickok, 2014). Clinically, distinc-
tions between different stages of speech production are reflected in a
widely accepted taxonomy of neurogenic speech sound disorders, with a
classification into lexical and postlexical phonological impairments (Schwartz,
2014), motor planning impairment (apraxia of speech; cf. Ziegler, Aichert,
& Staiger, 2012), motor execution disorders (dysarthria; cf. Duffy, 2013),
and impairments of motivational or emotional aspects of speaking and
speech initiation (Ackermann & Ziegler, 2010).
Assuming that different parts of the brain are engaged in the different
processing stages translating stored word forms into fully specified

The Linguistic Cerebellum © 2016 Elsevier Inc.

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2 The Linguistic Cerebellum

articulatory-motor patterns, the question that is pertinent in this vol-

ume is whether there is a contribution of the cerebellum to speech
sound production on one or several of these stages and how this
contribution can be framed in theories of speech production. Conven-
tionally, a major (and undisputed) role of the cerebellum in speech is
seen in the sensorimotor control and coordination of vocal tract and
laryngeal and respiratory movements during spoken language production
(Ackermann, 2008). In this vein, speech impairment after cerebellar
lesions has been characterized as a dysarthria syndrome, more specifically as
ataxic dysarthria (Kent et al., 2000). This perspective is consistent with
the traditional understanding of the cerebellum as an organ that is
primarily engaged in the coordination of movement (Holmes, 1917)
and the processing of sensory information during online motor control
(Gao et al., 1996).
Yet, along with a general revision of this view and with the discovery
of significant cerebellar contributions to linguistic, cognitive, and affective
processing (Schmahmann, 1991), the understanding of the cerebellum as
a purely sensorimotor coordination organ in speaking has been challenged
(Marien, Engelborghs, Fabbro, & De Deyn, 2001). An extensive number
of studies exists focusing on such higher order functions, both in the func-
tional imaging (e.g., Keren-Happuch, Chen, Ho, & Desmond, 2014) and
the clinical domain (Stoodley & Schmahmann, 2009a), and consensus
papers wrapping up the major findings have been published recently
(Koziol et al., 2014; Manto et al., 2012; Mariën et al., 2014). As com-
mented by Schlerf, Wiestler, Verstynen, and Diedrichsen (2014, p. 199),
“the cerebellum appears to play a role in nearly as many separate functions
as the neocortex.”1
Regarding the production of spoken language, this raises the question as
to whether the cerebellum has a role beyond the motor coordination of the
speech organs (i.e., whether it is also engaged in higher level processes of
speech sound production such as phonological encoding, phonetic plan-
ning, or the motivational and affective modulation of speech). In this chap-
ter, existing evidence on motor, linguistic, and emotional functions of the
cerebellum will be reviewed from the perspective of how they may contrib-
ute to speech production, from the phonological to the motor execution
and activation level.

1 However, there is no universal consensus about the view of the cerebellum as a cognitive organ (e.g.,
Glickstein, 2006; Schlerf,Verstynen, Ivry, & Spencer, 2010).
 The Phonetic Cerebellum 3

CEREBELLAR CONTRIBUTIONS TO SPEECH:

NEUROANATOMICAL BASIS
The understanding that the human cerebellum is an organ whose functions
are not limited to the coordination of movement is supported by new evi-
dence about its topographic organization and its connectivity with distinct
motor and nonmotor areas of the cerebral cortex (Stoodley & Schmah-
mann, 2010). Because many of these data stem from anatomical investiga-
tions in nonhuman primates, careful interpretation is warranted when it
comes to speech and language because monkeys and apes do not speak and
because the contribution of the cerebellum to precursor functions of speech
in these animals has not been examined extensively.

The Role of the Cerebellum in Nonhuman Primate

Vocalizations
The vocalizations of nonhuman primates are considered, by some researchers, as
a possible precursor of human speech (for references cf. Ackermann, Hage, &
Ziegler, 2014; Ackermann & Ziegler, 2010). Nonetheless, surprisingly little is
known about the role of the cerebellum in vocal communication in our closest
living relatives. Larson, Sutton, and Lindeman (1978) examined rhesus monkeys
which were trained to emit specific “coo” vocalizations. Experimental lesions
applied to the cerebellar nuclei and a partial removal of the cerebellar cortex
produced inconsistent changes in average fundamental frequency and/or loud-
ness and/or duration of the conditioned coo calls in some, but not all, of the five
animals. Unlike animals with lesions to mesiofrontal cerebral cortex (anterior
cingulate gyrus), monkeys with cerebellar lesions did not decrease their call
frequency. Larson et al. (1978) therefore concluded that the cerebellum is
involved in the modulation of pitch and loudness, but not in call initiation or
the modulation of the impulse to vocalize. Of note, however, the vocal altera-
tions mentioned in this study had nothing in common with the voice impair-
ments observed in humans with cerebellar dysfunctions (see the following
section). In a later study by Kirzinger (1985), experimental bilateral destruction
of the cerebellar nuclei of three squirrel monkeys had no effect on electrically
induced conspecific vocalizations in these animals, although their limb move-
ments became severely ataxic. Contrasting her results with those described by
Larson et al. (1978), Kirzinger (1985, p. 180) speculated that the cerebellum may
play a role in the volitional recruitment of the larynx for vocal communication
and the regulation of the “unspecific arousal level” underlying vocalization in
nonhuman primates, but not in the coordination of laryngeal movements.
4 The Linguistic Cerebellum

By conclusion, these data do not provide clear evidence that the (non-
human) primate cerebellum may serve as a model of human cerebellar
function in speaking. As a side note: In songbirds, which are considered to
constitute an important animal precursor model of especially the vocal imi-
tation and vocal learning aspects of human speech, there is also no evidence
so far of any significant contribution of the avian homologue of the cere-
bellum to birdsong (Ackermann & Ziegler, 2013). Therefore, speculating
about the possible specificity of a cerebellar contribution to vocal commu-
nication in our species, Ackermann (2008, p. 269) claimed that during
hominine evolution expression of the human-specific variant of the FOXP2
gene created a “permissive environment” within the primate cerebellar
motor circuit through which the extant motor computational resources of
the cerebellum could be harnessed for the vocal apparatus in the generation
of the sequential motor patterns involved in articulation (for a similar argu-
ment relating to the basal ganglia, see Ackermann et al., 2014).

Topographic Organization and Connectivity of the Human

Cerebellum From a Speech Perspective
Although current knowledge about the topography and connectivity of the
human cerebellum has to a large part been inferred from transneural tract
tracing studies in nonhuman primates (for a review, see Schmahmann,
2010), more recent studies using new in vivo diffusion tractography tech-
niques have added evidence on cerebrocerebellar connectivity in the intact
human brain that is by and large compatible with animal data (Catani &
Thiebaut de Schotten, 2008; Granziera et al., 2009).
In earlier anatomical models, the cerebellum was considered a motor
organ that funnels afferent information from different regions of the cortex
into output channels targeted primarily at the motor cortex. Modern theo-
ries, in contrast, assume that the cerebellum communicates through several
parallel closed-loop circuitries with motor, associative, and paralimbic corti-
cal areas of the “large brain” (Krienen & Buckner, 2009; Ramnani, 2006;
Stoodley & Schmahmann, 2010; Strick, Dum, & Fiez, 2009). It is claimed
that through this arrangement the cerebellum is implicated not only in
sensorimotor, but also in cognitive, linguistic, and affective functions.

The Motor Cerebellum

Among the 10 lobules that are distinguished within the cerebellar cortex,
those pertaining to the anterior lobe (lobules I–V) as well as lobule VIII and
parts of lobule VI of the posterior lobe are considered predominantly
 The Phonetic Cerebellum 5

sensorimotor. Hemispheric lobule VI is particularly relevant here because it

contains the lip and tongue area of a sensorimotor homunculus (Mottolese
et al., 2013). However, early electrophysiological investigations in animals
and more recent functional imaging studies in humans have revealed that
there is at least one further cerebellar homunculus in the posterior lobe,
with a tongue area located in lobule VIII, but still other somatotopic maps
(i.e., no less than four homunculi) are considered to exist in the anterior and
posterior lobes (Buckner, Krienen, Castellanos, Diaz, & Yeo, 2011; Grodd,
Hülsmann, Lotze, Wildgruber, & Erb, 2001; Manni & Petrosini, 2004;
Rijntjes, Büchel, Kiebel, & Weiller, 1999; Schlerf et al., 2010).
From animal work, it is known that the cerebellar cortical face areas in
lobule VI receive somatosensory afferent information from oral and facial
muscles via the trigeminal nuclei through the inferior cerebellar peduncles
(Stoodley & Schmahmann, 2010, p. 832).This afferent input is considered to
provide the cerebellar cortex with information about the sensory state of
the speech organs. Furthermore, the cerebellar hemispheres receive afferent
projections travelling through pontine nuclei from motor areas of the con-
tralateral cerebral cortex, especially from primary motor cortex, ventral pre-
motor cortex, and supplementary motor area (Brodal, 1978; Glickstein, May,
& Mercier, 1985; Schmahmann, Rosene, & Pandya, 2004).2 These projec-
tions are somatotopically organized throughout their pathway to the nuclei
in the basis pontis and further on through primarily the contralateral middle
cerebellar peduncle to the cerebellum (for references cf. Ramnani, 2006;
Stoodley & Schmahmann, 2010). From there, efferent fibers travel through
output channels in the deep cerebellar nuclei to the motor-nuclei of the
contralateral thalamus and back to the motor areas of the cerebral cortex,
still in a somatotopically organized fashion (Middleton & Strick, 2000).
Hence, the motor cerebellum is connected with cerebral motor cortical
areas via a two-stage feedforward limb through the basis pontis and via a
two-stage feedback limb through the thalamus (e.g., Kelly & Strick, 2003).
Several spatially distinct, somatotopically organized loops are considered to
project back to the primary motor, ventral premotor, and supplementary
motor areas from which they originate, thereby forming separate closed
feedforward-feedback motor circuits. Apart from these dentate-thalamico-
cerebral feedback projections, Mottolese et al. (2013) inferred from electro-
myographic latencies obtained during direct perioperative mapping in
2 Corticocerebellar projections are not restricted to the pontine nuclei. Other precerebellar nuclei in
the brainstem receive input from the cerebral cortex and serve as cerebrocerebellar relays (cf.
Glickstein, Sultan, & Voogd, 2011 for a comprehensive review).
6 The Linguistic Cerebellum

humans that efferent cerebellar signals effectuate motor responses not via a
cerebral cortical feedback loop, but rather more directly through a descend-
ing subcortical cerebelloreticular pathway.
By analogy to limb motor control, the described system constitutes the
substrate of the contribution of the cerebellum in oral motor control and
motor speech. A rather specific cerebellar mechanism in speech motor con-
trol was disclosed by Golfinopoulos et al. (2011), who performed a jaw-
perturbation experiment in the functional magnetic resonance imaging
(fMRI) scanner to examine the neural substrate of somatosensory feedback
control. Increased activation in bilateral lobule VIII was found in perturbed
versus unperturbed speech, which was interpreted as an indication that this
region facilitates somatosensory cortex in the monitoring and correction of
unexpected perturbations in the periphery.
The data described so far leave several unanswered questions, such as (1)
whether the motor loop involving the vocal tract and laryngeal muscles in
humans is lateralized for speech motor control; (2) whether the cerebrocer-
ebellar motor system composes two separate loops, one for the planning and
the other for the coordination of speech movements, separated by their
respective premotor and motor target regions in the cerebral cortex; or (3)
whether there are differential contributions of the repeated body represen-
tations in the cerebellum to speech. The functional imaging and clinical
data reported later will shed some light on these questions.

The Linguistic Cerebellum

The somatomotor maps in the anterior and posterior human cerebellum are
separated by an extended region within lobules VI–VIII that appears
uncharted for sensorimotor functions. Substantial evidence has been accu-
mulated that this neocerebellar region, especially two areas termed Crus I
and Crus II in lobules VI and VII, have dense connections with association
areas of the cerebral cortex, leading to the hypothesis that this expanded
part of the human cerebellum is implicated in cognitive functions and in
the regulation of affect and emotion (Schmahmann, 2010).3 In a meta-
analysis of functional imaging studies on the functional topography of the
human cerebellum, Stoodley and Schmahmann (2009b) localized spatial
processing capabilities, language functions, and executive functions to Crus
I and Crus II (i.e., to the areas connected with associative cerebral cortical
3 Opposing this theory, Schlerf et al. (2010) suggested that neocerebellar lobules VI and VII contain
further task-specific somatomotor representations dedicated to the control of particularly complex
motor acts.
 The Phonetic Cerebellum 7

regions). In this article, as in most others on this issue, linguistic processing

was included among the cognitive functions attributed to the posterior cer-
ebellar lobe and its cerebral connectivity (Mariën et al., 2014, p. 399; Stoodley &
Schmahmann, 2010). Keren-Happuch et al. (2014) identified significant
language-related activation also bilaterally in lobule VI.
Among the association areas of cerebral cortex projecting upon the pos-
terior cerebellum through the pontine nuclei, prefrontal, posterior parietal,
superior temporal, and parastriate areas have been established in nonhuman
primates (Schmahmann & Pandya, 1997b). Notably, studies reporting pre-
frontal-cerebellar connectivity are mostly committed to dorsolateral regions
which are not specifically implicated in speech (e.g., areas 9 and 46) (Brodal,
1978; Kelly & Strick, 2003; Ramnani, 2006), whereas the ventral prefrontal
region, especially areas 44 and 45, is mentioned less often (Schmahmann &
Pandya, 1997a). Leiner, Leiner, and Dow (1991) reviewed some pathoana-
tomic and experimental evidence that inferior frontal areas 44 and 45 ante-
rior to the premotor area (in the human left brain, this is termed Broca area)
send projections to the pontine nuclei and receive cerebellar input via the
ventrolateral thalamus.
Because in humans the “linguistic brain” in the cerebral hemispheres is
strongly lateralized, an equally strong lateralization—to the contralateral
hemisphere—would be expected to exist in the linguistic cerebellum. Jansen
et al. (2005) put the hypothesis of a crossed cerebrocerebellar link in lan-
guage processing to a particularly strong test by examining, in an fMRI
study, cerebellar activation during a word-generation task in participants
with (typical) left versus (atypical) right lateralized frontal lobe activation. A
crossed cerebrocerebellar activation was found in all participants (N = 14)
(i.e., right lateral cerebellum in participants with left hemisphere domi-
nance and left lateral cerebellum in those with the atypical right hemisphere
dominance). This outcome provides strong evidence for lateralized cerebel-
lar language processing, contralateral to the dominant cerebral hemisphere.
Interestingly, the paradigms subsumed under the term “linguistic” in
this literature rarely encompass plain production or comprehension of
words or sentences, but rather concentrate on meta-linguistic tasks testing
verbal fluency, verb generation, word stem completion, or verbal working
memory. As these tasks are known to be particularly sensitive to executive
functions involved in the specific retrieval strategies required for efficient
word generation (e.g., Leggio, Silveri, Petrosini, & Molinari, 2000), they
are not particularly appropriate to tax linguistic functions in speech
production.
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160

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CHAPTER to here

wild
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a portion seven

Egypt before

99

race is it