Neurobiology of Language Steven L

CHAPTER 1 The Neurobiology of Language Steven L. Small1 and Gregory Hickok2 1Department of Neurology, University of California, Irvine, CA, USA; 2Department of Cognitive Sciences, Center for Language Science, Center for Cognitive Neuroscience, University of California, Irvine, CA, USA

1.1 HISTORY

For many centuries, the biological basis of human thought has been an important focus of attention in medicine, with particular interest in the brain basis of language sparked by the famous patients of Pierre Paul Broca in the mid 19th century (Broca, 1861a,c). The patient Louis Victor LeBorgne (Domanski, 2013)presentedtotheHoˆpital Biceˆtre in Paris with severe difficulty speaking, purport- edly only uttering the syllable “tan,” sometimes as a pair “tan, tan,” and often accompanied by gestures (Domanski, 2013). The diagnosis was not clear until autopsy, when Broca found on gross inspection that some neurological process (he reported a resulting collection of serous fluid) had destroyed a portion of the left posterior inferior frontal FIGURE 1.1 The exterior surface of the brain of LeBorgne (“tan”). gyrus (Broca, 1861c)(Figure 1.1). A subsequent patient, LeLong, had a similar paucity of speech output (five Wernicke (1874), the diagram-making of Lichtheim words were reported) with a lesion not dissimilar to that (1885) (Figure 1.2) and Grashey (1885), the anatomy of of LeBorgne (Broca, 1861a). Given the ongoing debates at De´jerine (1895), and of course many other contributors. the time about brain localization of language, including In the past century, Norman Geschwind recapitulated attribution of the “seat of language” to the frontal and added to the language “center” models that pre- lobes (Auburtin, 1861; Bouillaud, 1825; Gall & ceded him and presented a reconceptualized “connec- Spurtzheim, 1809)—which led Broca to investigate this tionist” view of the brain mechanisms of language case in the first place—he presented this patient with (Geschwind, 1965, 1970). Whereas the 19th century “aphe´mie” (LeBorgne) to the Socie´te´ d’Anthropologie de investigators relied on simple views of behavior and Paris in 1861 (Broca, 1861b). These brains remain pre- postmortem brain pathology, those of the mid to late served to this day, and brain imaging studies have con- 20th century were able to take advantage of significant firmed the original findings and extended them to advances in both the study of behavior (information pro- demonstrate lesions to deeper structures (Signoret, cessing psychology and formal linguistic theory), allow- Castaigne, Lhermitte, Abelanet, & Lavorel, 1984)andto ing much more robust characterizations of language white matter underlying the original descriptions performance than had been possible previously (Dronkers, Plaisant, Iba-Zizen, & Cabanis, 2007). (Caramazza & Berndt, 1978; Chomsky, 1965), and the technology of structural brain imaging with computed tomography and magnetic resonance imaging (MRI), 1.2 LESION ANALYSIS permitting the elucidation of brain pathology in vivo (e.g., Cappa & Vignolo, 1983; Metter et al., 1984). These The era of brain localization for language blossomed advances led to a blossoming of brain lesion analysis after this, with the famous doctoral dissertation of (neuropsychological) studies in the second half of the

Concept brain and behavior. Although the earliest functional ana- field tomical studies were conducted with positron emission tomography, the most recent work uses functional magnetic resonance imaging (fMRI), a noninvasive approach that does not require intravenous administration of 4 6 radioactive agents. These methods permit the investigation in vivo of brain regions that participate in the performance of any type of task that can be performed inside 3 Broca’s Wernicke’s of an imaging machine. By contrast, task-dependent area area electroencephalography (EEG), more commonly known 1 2 as “event-related potentials” (ERP), permits the charac- terization of temporal aspects of task performance. An 57important, if less commonly used, method for analyzing task-dependent brain function is magnetoencephalo- graphy (MEG), which can have finer spatial resolution Peripheral Peripheral than EEG and higher temporal resolution than fMRI, speech auditory and thus can play a particularly important role in charac- production input terizing brain processing over time. Besides these methods of activating the brain, the method of transcra- FIGURE 1.2 Lichtheim’s model. Note the emphasis on brain anatomy. nial magnetic stimulation (TMS) can be used to create “reversible lesions” in the brain. The majority of contemporary research in brain behavior relations for language À 20th century (for reviews, see Damasio & Damasio, 1989; uses either the older method of lesion analysis or one of Kertesz, 1983; Shallice, 1988a). Recent advances in image these newer methods of functional activation (fMRI, analysis (e.g., Ashburner & Friston, 2000; Bates et al., ERP, MEG) or ablation (TMS). Intracranial electrical 2003) have improved the lesion analysis method, and it recordings in humans undergoing elective brain surgery, continues to be a valuable method for biological inqui- including both surface (electrocorticography) and deep, ries in language. are becoming more commonplace. A host of other less By their very nature, lesion analysis studies tend to prevalent methods also play valuable roles. relate single foci of brain injury to single psychological Approximately 20 years ago, the burgeoning use of or linguistic phenomena. The goal of the enterprise is to these brain measurement techniques to study psychologi- “double dissociate” functions and brain regions, such cal and linguistic processes led to the creation of a new that an underlying focal substrate of brain can be tied field of cognitive neuroscience. During its existence, this unequivocally to a single function (and not another) field has evolved into an important discipline, with the (e.g., Damasio & Tranel, 1993) or to a specific “locus” in majority of top cognitive programs (and some neurosci- a cognitive or linguistic model (e.g., Garrard & Hodges, ence programs) incorporating cognitive neuroscience as 2000; Shallice, 1988b). Although not without its contro- an important component of curriculum and, in some versy (e.g., Plaut, 1995), this approach has been quite cases, degree-granting status. Importantly, the evolution successful in giving insights into the neurobiology of of this discipline has focused more on using biological language, that is, attributing functions to aspects of the methods than on asking biological questions, for exam- brain (in this case, brain areas damaged by vascular ple, addressing linguistic or psychological questions by lesions—or sometimes other types of lesions). measuring brain responses constitutes a significant portion—if not the majority—of scientific studies in this area. In other words, the fraction of studies that develop and 1.3 FROM NEUROPSYCHOLOGY TO test “linking hypotheses” between neural and computa- COGNITIVE NEUROSCIENCE tional systems is smaller than one would hope for in a field that targets an understanding of the relation between The advent of high-resolution functional brain mind and brain. Addressing biological issues is far less imaging in the past decade of the 20th century (e.g., Fox, common in this field, and most practitioners of this Raichle, & Thach, 1985; Raichle, Martin, & Herscovitch, discipline neither study biology nor concern themselves 1983), its initial applications to the study of language with biological questions. Students in this area are (Petersen, Fox, Posner, Mintun, & Raichle, 1988, 1989), typically not required to study cellular and molecular and its widespread acceptance for the study of brain/ neuroscience, and only a portion study systems neurosci- cognition relations in the early part of this century ence, neuroanatomy, neurophysiology, or neurology/ have dramatically changed the conduct of studies of neuropsychology. Of course, it is also true that from the

A. INTRODUCTION 1.5 SOME COMMON FALLACIES 5 biological side, students and researchers are not typically that the human brain evolved from that of nonhuman required to study linguistics or experimental psychology. primates, and thus the study of other species plays Despite this high prevalence of nonbiological studies of an important role in the investigations of the neurobiol- cognition using brain measurement, it has become ogy of language (Bornkessel-Schlesewsky, Schlesewsky, increasingly clear that the new methods of functional Small, & Rauschecker, 2015). brain imaging, along with other methods of human and A number of brain imaging methods are used for the animal neuroscience, provide new avenues to investigate neurobiology of cognition but do not play a significant the actual biological substrate and computations for role in cognitive neuroscience. An enormous amount of human language. information about the neurobiology of language has been gleaned from high-resolution quantitative structural imaging of both gray matter and white matter. For 1.4 THE NEUROBIOLOGY OF the gray matter, it is possible to measure precisely and LANGUAGE accurately whole brain and regional volumes, and for white matter, we can measure accurately anisotropic Thus, a neurobiology of language is now possible. diffusion (and several other diffusivity characteristics). Until several years ago, this term was barely used, if at In both cases, the anatomical measures can be related to all, with most research focusing on neuropsychology, behavior or to functional imaging measures. At a cognitive neuroscience, or vaguely “brain and lan- network level, white matter pathways can be recon- guage.” We define “neurobiology of language” as the structed from the diffusivity data and can be related to biological implementation and linking relations for regional correlations of gray matter characteristics or representations and processes necessary and sufficient functional brain activations. These relations between for production and understanding of speech and lan- structural connectivity and functional connectivity are guage in context. Biological disciplines that are highly leading to a preliminary understanding of brain circuits relevant to the neurobiology of language include the for language processing. anatomy and physiology of the human brain, the net- It is also possible to make direct measurements of work connectivity of the brain, and the multiple roles human neural cells when medically indicated, as in peo- of different brain areas (relevant cognitive areas are ple with intractable epilepsy who are undergoing abla- discussed later). We also note that the basic scientific tive brain surgery. Early work demonstrated that direct findings in this area will necessarily lead to a physiolog- cortical stimulation could lead to alterations in language ical approach to therapy for speech and language disor- performance in humans in vivo (Ojemann, Ojemann, ders (Small, Buccino, & Solodkin, 2013). Lettich, & Berger, 1989; Penfield & Roberts, 1959). Importantly, we define the neurobiology of language Although this technique remains viable (e.g., Duffau, as a subfield of neuroscience, requiring substantial 2008), recent work has used cortical recording (electrocor- knowledge of psychology and linguistics, sharing in its ticography) to begin to elucidate cellular and circuit primary assumptions, methods, and questions. By features of language representations at the level of small way of explanation, whereas psychology is the scientific neuronal populations (e.g., Chang et al., 2010; Flinker study of the human mind and its functions, especially et al., 2015). Many additional biological techniques, such those affecting behavior in particular contexts, and as genetic mapping, single cell recording, pharmacologi- linguistics is the scientific study of language and its cal manipulation, molecular imaging, and others, are structure, including the study of morphology, syntax, beginning to be viable for human neuroscience research. phonetics, and semantics, neurobiology is the study of the biology of the nervous system. Whereas cognitive neuroscience emphasizes experimental psychology and brain measurements, the neurobiology of cognition 1.5 SOME COMMON FALLACIES emphasizes cellular and molecular neuroscience, systems neuroscience, and other biological bases of From a philosophical point of view, several assump- language, such as genetics. We believe that both cogni- tions commonly made in contemporary research in tive neuroscience and the neurobiology of cognition brain behavior relations related to language are prob- À must address the links between cognitive and neurosci- lematic. We identify four such assumptions, which ence data, although they come from different directions. we call the (i) methodological fallacy; (ii) theoretical Although the neurobiology of language is primarily a fallacy; (iii) uniqueness fallacy; and (iv) the mind/ field involving human subjects—and has benefited from brain fallacy. These fallacies relate to research assump- the novel and highly robust methods of brain measure- tions by investigators in the cognitive sciences that ment discussed—it must also adopt the central tenets of lead to inappropriate biological conclusions from other- modern biology, including the evolutionary imperative wise well-conducted studies in cognition (not

A. INTRODUCTION 6 1. THE NEUROBIOLOGY OF LANGUAGE infrequently accompanying perfectly interesting and In summary, we are concerned with the notion that valid cognitive conclusions). We explain these in turn. linguistic computations are so unique that they The methodological fallacy assumes that if a brain bear no evolutionary resemblance to other systems. measurement technique is used in a study, that study However, we acknowledge that language uniqueness is a study of the brain. Such an assumption essentially cannot be ignored. Language ability is certainly unique focuses on the method that is used, for example, if a to our species, suggesting an evolutionary process in research study uses EEG or MRI, then it is a neurobio- which language computations developed on a frame- logical study. This is not the case. Among the most work of other computational structures in the brain. It common uses of ERP is to distinguish one linguistic or is thus critical that we take the biology seriously and psychological model from another, and fMRI is used search for both computational commonalities and dif- very commonly to distinguish individuals with one ferences between language and nonlanguage systems condition (medical or psychological) from another. In within our species and in our cousin species. neither case would an understanding of the biology be Finally, the most common of all the fallacies is the necessary to achieve the goals of the study. However, mind/brain fallacy, which argues that all studies of it is important to note that some EEG/MEG research human psychology are ipso facto studies of the brain, does aim to understand the relation between neural because the brain is the biological organ that subserves network rhythms and cognitive processes, and much thought. Of course, this is perfectly correct, to a point, fMRI research also addresses neural computations. In and the Society for Neuroscience includes human summary, studies of this sort do not address linking behavior among its disciplinary mandates. However, it hypotheses, although data from these studies could be is completely possible—even commonplace—for studies used to inform linking relations. of human behavior to completely ignore the constraints The theoretical fallacy assumes that a theoretical placed on the behavior by the physical aspects of the model developed exclusively from assumptions about brain or the plausibility of the behavioral conclusions in linguistic data or cognitive processing explains brain the context of brain structure and function. Among the function. Such an assumption essentially ignores data most common examples in language research are expla- from neuroscience while focusing on theoretical nations of experimental data in the form of “concep- aspects of the legal utterances (linguistics) or processes tual” or “cognitive” or “mental” models that could not (psychology) in one or more (or even “all”) languages. possibly be implemented in brain hardware. Even if a theory accounts for all legal utterances in all We certainly have no objection to cognitive models languages but does not say how brain structures actu- and indeed value them, whether or not they make imme- ally compute them, it is not a theory of the biology of diate contact with neural implementation. These models language. Of course, if a biological theory ignores the help us think about information flow (and/or dynamics) linkages between the neurobiology and cognition, it and computational descriptions of the process. As such, also is not a theory of the neurobiology of language. they can sometimes generate testable hypotheses related The uniqueness fallacy is perhaps the most important to the brain as well as provide prospective points of con- because it ignores evolution, which is one of the most tact for descriptions that relate brain to behavior. If we important principles of modern biology. This fallacy tried to build a neurobiology of language without typically arises in the form of arguments that human linguistics or psychology, as the classical neurologists language is so unique and so special that it is not com- were forced to do, we would not have been able to tran- putationally intertwined with other systems. But we scend the 19th century models (e.g., characterizations know that language systems of the brain are fundamen- such as “auditory word images”). But an important tally and inherently intertwined with those that process point here (for the cognocentric among us) is that the motor function, sensation, emotion, affect, and other constraints must work both ways. Knowledge of biology aspects of human experience, and that the human must constrain our models of cognitive and linguistic brain—including those parts of the brain that partici- function, just as knowledge of cognition must constrain pate in language processing—had to evolve from the our biological interpretations. Both are obviously critical brains of species (i.e., nonhuman primates) that do not in the development of linking hypotheses. use language. The evolutionary principles of phylogeny and ontogeny apply to the human brain as with other organ systems, and thus language representations in 1.6 HUMANS IN PARTICULAR the brain have necessarily co-opted neural structures with evolutionary history. Relevant examples include There is no question that the human brain is unique auditory object perception, motor sequence control and in its capacity to manage speech understanding and planning, action understanding, and social interaction, production, in addition to their evolutionarily younger all of which are found in other species. derivatives, reading and writing. That the human brain

A. INTRODUCTION 1.8 BRAIN DISEASE, TREATMENT, AND THE NEUROBIOLOGY OF LANGUAGE 7 can do this and other animal brains cannot is something The study of linguistics is therefore critical to a suc- that must be explained in a way that preserves the basic cessful neurobiology of language. tenets of biology. Certainly, humans have vocal commu- Equally important is characterizing the nature of nication skills of very high complexity, and these speech production and understanding in processing depend on a large and complex variety of sensory and terms. Without detailed knowledge about what people motor skills, memory capacities, and learning abilities, can understand, in what contexts, and under what con- as well as the capability to represent and process intri- straints, we cannot develop a solid neurobiology of cate messages of many types. Although it is not clear understanding. The same is true for production, that is, exactly what changed between nonhuman primates and we need to know what people can produce at a given humans to effectuate this, the expansion of the cerebral time and place, particularly with memories, emotions, cortex increased the human capacity for learning, and goals. Such investigations are the purview of exper- remembering, and executing motor sequences, complex imental psychology and, as such, research in such areas perceptions, and recursive processing (e.g., theory of as psycholinguistics and the psychology of memory has mind, syntax) (Hauser, Chomsky, & Fitch, 2002; fundamental importance to inform investigations in the Yoshida, Dolan, & Friston, 2008). It remains to be eluci- biological aspects of language. dated just how the processes of language work in the human brain, and through this—perhaps with concomi- tant work in nonhuman primates to test their limits— 1.8 BRAIN DISEASE, TREATMENT, AND we should find the answer to why humans have lan- THE NEUROBIOLOGY OF LANGUAGE guage and other species do not. There are many diseases and/or types of injury to the human brain that lead to language disturbances, and 1.7 COGNITION AND THE these are important to our enterprise for two reasons: NEUROBIOLOGY OF LANGUAGE (i) helping to understand the basic neurobiology and (ii) advancing knowledge useful for development of treat- The neurobiology of cognition will only be elucidated ments for conditions that affect language. Both of these by direct investigation of how the brain works, that is, endeavors have considerable importance to the field. the detailed implementation of understanding and pro- By understanding the nature of disease processes, it ducing speech in the hardware that is unique to the may be possible to gain insight into the fundamental human brain. At the same time, such direct investigation neurobiology of language itself. For example, the cellu- must be informed by scientific evidence about the nature lar and molecular biology of language has been given of cognition. For the neurobiology of language, this a first hearing with the distinct pathological markers entails understanding the nature of languages and its that differentiate the types of primary progressive processing, that is, investigations in linguistics and aphasia (Gorno-Tempini et al., 2011). The entire history experimental psychology. The first step must involve of research in the biological basis of language has spoken language, because that is the evolutionary basis depended significantly on inferences from individuals on which the brain architecture for written language sits. with ischemic stroke (e.g., Mohr, 1976). Patients Understanding the nature of human languages is surviving herpes simplex encephalitis, a multifocal crucial, particularly research into the wide variety of disease, have provided important fodder for infe- different types of languages and how they express rences about semantic representations in the brain thoughts (e.g., MacWhinney & Bates, 1989). There is (Warrington & Shallice, 1984). Language problems nothing in the basic brain structure of people born to arising in people with dementia, tumors, Parkinson’s parents who speak Basque, Farsi, or Navaho to distin- disease, schizophrenia, and many other diseases— guish them from speakers of English, Italian, or combined with the nature of the pathology specific to German. Yet, most research studies relating brain and that disease (particularly in the context of the pathol- language have been conducted in the latter three lan- ogy of each individual involved)—are highly informa- guages (and now, increasingly, Mandarin as well), and tive with respect to the neurobiology of language. rarely in the other approximately 7,000 languages of Additionally, people undergoing surgery for diseases the world (Lewis, Simons, & Fennig, 2009), some of such as epilepsy and brain tumors, even when they do which have properties that are quite different from not affect language, have generously allowed investi- those of the most commonly studied languages. Thus, gators to study their language intraoperatively. it is important in characterizing the biological basis of Of course, a very important purpose in understanding language to know just what needs to be accounted for the biology underlying language is to develop new both in terms of variation of language forms and the therapies (Small et al., 2013; Small & Llano, 2009). Novel computational mechanisms used to process them. treatments including pharmacotherapy, electrical and

A. INTRODUCTION 8 1. THE NEUROBIOLOGY OF LANGUAGE magnetic stimulation, and even behavioral (speech- the efforts of practicing scientists and their students to language therapy) interventions, have been developed delve directly into this challenging field. based on rationales that come from a better understanding of brain anatomy and physiology. References Ashburner, J., & Friston, K. J. (2000). Voxel-based morphometry— The methods. NeuroImage, 11, 805 821. À 1.9 SUMMARY Auburtin, E. (1861). Reprise de la discussion sur la forme et le volume du cerveau. Bulletins de la Socie´te´ d’Anatomie Paris, 2, 209 220. À In summary, investigation into the neurobiology of Bates, E., Wilson, S. M., Saygin, A. P., Dick, F., Sereno, M. I., Knight, language has a long history, beginning even before the R. T., et al. (2003). Voxel-based lesion-symptom mapping. Nature Neuroscience, 6(5), 448 450. seminal work of the 19th century neurologists, who asso- À Bornkessel-Schlesewsky, I., Schlesewsky, M., Small, S. L., & ciated brain lesions in certain focal areas (but not others) Rauschecker, J. P. (2015). Neurobiological roots of language in with language impairments. However, recently, this primate audition: Common computational properties. Trends in field has had a resurgence due to the advent of extraor- Cognitive Science. Available from: http://dx.doi.org/10.1016/j.tics. dinary new technologies that allow in vivo imaging of 2014.12.008. brain structure, structure/functional relations, and tem- Bouillaud, J.-B. (1825). Recherches cliniques propres a` de´montrer que la perte de la parole correspond a` la le´sion des lobules ante´rieurs poral processing features. Although lesion analysis du cerveau, et a` confirmer l’opinion de Gall, sur le sie`ge du remains quite important—and even more valuable in langage articule´. Archives of General Medicine (Paris), 8, 25 45. À conjunction with advanced imaging—the current era is Broca, P. P. (1861a). Nouvelle Observation d’Aphe´mie produite par marked by a predominance of investigations using brain une Lesion de la Partie Poste´rieure des Deuxie`me et Troisie`me imaging technologies (broadly speaking). These methods Circonvolutions Frontales. Bulletins de la Socie´te´ d’Anatomie Paris, 6, 398 407. À can be used to advance our understanding of linguistics Broca, P. P. (1861b). Perte de la parole, ramollisement chronique et or psychology, thus contributing in a different way to destruction partielle du lobe ante´rieur gauche du cerveau. the neurobiology of language, or to advance our knowl- Bulletins de la Socie´te´ d’Anthropologie, 2, 235 238. À edge of the anatomy, physiology, pharmacology, and Broca, P. P. (1861c). Remarques sur le sie`ge de la faculte´ du langage even cellular and molecular biology of language directly. articule´, suivies d’une observation d’aphe´mie (perte de la parole). Bulletins de la Socie´te´ d’Anatomie (Paris), 6(2e serie), 330 357. À Such direct investigations of the brain in conjuction with Cappa, S. F., & Vignolo, L. A. (1983). CT scan studies of aphasia. investigations into the representations and computations Human Neurobiology, 2(3), 129 134. À behind language processes represent basic prerequisites Caramazza, A., & Berndt, R. S. (1978). Semantic and syntactic pro- in building the future of this discipline. cesses in aphasia: A review of the literature. Psychological Bulletin, 85(4), 898 918. The two editors of this book—without a consensus on À Chang, E. F., Rieger, J. W., Johnson, K., Berger, M. S., Barbaro, N. M., all issues—have come to the many common understand- & Knight, R. T. (2010). Categorical speech representation in ings of the field articulated here. We are in agreement human superior temporal gyrus. Nature Neuroscience, 13(11), that it is important to have “theoretically precise, compu- 1428 1432. Available from: http://dx.doi.org/10.1038/nn.2641. À tationally explicit, biologically grounded explanatory Chomsky, N. (1965). Aspects of the theory of syntax. Cambridge, MA: models of the human brain’s ability to comprehend and The MIT Press. Damasio, A. R., & Tranel, D. (1993). Nouns and verbs are retrieved produce speech and language” (Poeppel & Hickok, with differently distributed neural systems. Proceedings of the 2004). We are also in agreement that the basic evolution- National Academy of Sciences of the United States of America, 90(11), ary biology of the brain must play an important role in 4957 4960. À such a theory, for example, by understanding the relation Damasio, H., & Damasio, A. R. (1989). Lesion analysis in neuropsychol- between computational implementations of sensory and ogy. New York, NY: Oxford University Press. De´jerine, J. (1895). Anatomie des centres nerveux (Vol. i ii). Paris: Rueff À motor functions (among others) to linguistic computa- et Cie. tions (Skipper, Nusbaum, & Small, 2005). By linking bio- Domanski, C. W. (2013). Mysterious “Monsieur Leborgne”: The mys- logical and cognitive hypotheses into well-designed tery of the famous patient in the history of neuropsychology is explained. Journal of the History of the Neurosciences, 22(1), 47 52. studies, it is now possible to build a true neurobiology of À language. Available from: http://dx.doi.org/10.1080/0964704x.2012.667528. Dronkers, N. F., Plaisant, O., Iba-Zizen, M. T., & Cabanis, E. A. The reviews in this book reflect a combination of (2007). Paul Broca’s historic cases: High resolution MR articles on the neurobiology of language per se and on imaging of the brains of Leborgne and Lelong. Brain, 130(Pt 5), the supporting knowledge from the neurosciences and 1432 1441. À from the cognitive sciences (including cognitive neuro- Duffau, H. (2008). The anatomo-functional connectivity of language science) that neurobiologists of language require to revisited. New insights provided by electrostimulation and tracto- graphy. Neuropsychologia, 46(4), 927 934. Available from: http:// À accurately characterize the neural structures and func- dx.doi.org/10.1016/j.neuropsychologia.2007.10.025. tions that implement language. Ultimately, we hope Flinker, A., Korzeniewska, A., Shestyuk, A. Y., Franaszczuk, P. J., that this reference book will encourage and support Dronkers, N. F., Knight, R. T., et al. (2015). Redefining the role of

A. INTRODUCTION REFERENCES 9

Broca’s area in speech. Proceedings of the National Academy of stimulation mapping investigation in 117 patients. Journal of Sciences of the United States of America, 112(9), 2871 2875. Neurosurgery, 71, 316 326. À À Available from: http://dx.doi.org/10.1073/pnas.1414491112. Penfield, W., & Roberts, L. (1959). Speech and brain mechanisms. Fox, P. T., Raichle, M. E., & Thach, W. T. (1985). Functional mapping Princeton, NJ: Princeton University Press. of the human cerebellum with positron emission tomography. Petersen, S. E., Fox, P. T., Posner, M. I., Mintun, M. A., & Raichle, Proceedings of the National Academy of Sciences of the United States of M. E. (1988). Positron emission tomographic studies of the cortical America, 82, 7462 7466. anatomy of single-word processing. Nature, 331, 585 589. À À Gall, F. J., & Spurtzheim, J. G. (1809). Recherches sur le syste`me nerveux Petersen, S. E., Fox, P. T., Posner, M. I., Mintun, M. A., & Raichle, en ge´ne´ral et sur celui de cerveau en particulier. Paris: F. Schoell. M. E. (1989). Positron emission tomographic studies of the proces- Garrard, P., & Hodges, J. R. (2000). Semantic dementia: Clinical, sing of single words. Journal of Cognitive Neuroscience, 1, 153 170. À radiological and pathological perspectives. Journal of Neurology, Plaut, D. C. (1995). Double dissociation without modularity: 247(6), 409 422. Evidence from connectionist neuropsychology. Journal of Clinical À Geschwind, N. (1965). Disconnection syndromes in animals and and Experimental Neuropsychology, 17(2), 291 321. Available from: À man. Brain, 88, 237 294. 585 644. http://dx.doi.org/10.1080/01688639508405124. À À Geschwind, N. (1970). The organization of language and the brain. Poeppel, D., & Hickok, G. (2004). Towards a new functional anatomy Science, 170(961), 940 944. of language. Cognition, 92(1 2), 1 12. À À À Gorno-Tempini, M. L., Hillis, A. E., Weintraub, S., Kertesz, A., Mendez, Raichle, M. E., Martin, W. R. W., & Herscovitch, P. (1983). Brain 15 M., Cappa, S. F., et al. (2011). Classification of primary progressive blood flow measured with intravenous H2 O II: Implementation aphasia and its variants. Neurology, 76(11), 1006 1014. Available and validation. Journal of Nuclear Medicine, 24, 790 798. À À from: http://dx.doi.org/10.1212/WNL.0b013e31821103e6. Shallice, T. (1988a). From neuropsychology to mental structure. Grashey (1885). Ueber aphasie und ihre Beziehungen zur Cambridge, England: Cambridge University Press. Wahrnehmung. Archiv fu¨r Psychiatrie und Nervenkrankheiten, 16. Shallice, T. (1988b). Specialisation within the semantic system. Hauser, M. D., Chomsky, N., & Fitch, W. T. (2002). The faculty of Cognitive Neuropsychology, 5(1), 133 142. À language: What is it, who has it, and how did it evolve? Science, Signoret, J. L., Castaigne, P., Lhermitte, F., Abelanet, R., & Lavorel, P. 298(5598), 1569 1579. Available from: http://dx.doi.org/ (1984). Rediscovery of Leborgne’s brain: Anatomical description À 10.1126/science.298.5598.1569. with CT scan. Brain and Language, 22(2), 303 319. À Kertesz, A. (Ed.), (1983). Localization in neuropsychology New York, Skipper, J. I., Nusbaum, H. C., & Small, S. L. (2005). Listening to talk- NY: Academic Press. ing faces: Motor cortical activation during speech perception. Lewis, M. P., Simons, G. F., & Fennig, C. D. (2009). (16th ed.). Ethnologue: Neuroimage, 25(1), 76 89. À Languages of the world, (Vol. 9). Dallas, TX: SIL international. Small, S. L., Buccino, G., & Solodkin, A. (2013). Brain repair after Lichtheim, L. (1885). On aphasia. Brain, 7, 433 484. stroke—A novel neurological model. Nature Reviews Neurology, 9 À MacWhinney, B., & Bates, E. (Eds.), (1989). The crosslinguistic study of (12), 698 707. Available from: http://dx.doi.org/10.1038/nrneurol. À sentence processing Cambridge, England: Cambridge University 2013.222. Press. Small, S. L., & Llano, D. A. (2009). Biological approaches to aphasia Metter, E. J., Riege, W. H., Hanson, W. R., Camras, L. R., Phelps, treatment. Current Neurology and Neuroscience Reports, 9(6), 443 450. À M. E., & Kuhl, D. E. (1984). Correlations of glucose metabolism Warrington, E. K., & Shallice, T. (1984). Category specific semantic and structural damage to language function in aphasia. Brain and impairments. Brain, 107(Pt 3), 829 854. À Language, 21(2), 187 207. Wernicke, C. (1874). Der aphasische symptomenkomplex. Breslau, ON: À Mohr, J. P. (1976). Broca’s area and Broca’s aphasia. In H. Whitaker, & Cohn & Weigert. H. Whitaker (Eds.), Studies in neurolinguistics (Vol. 1, pp. 201 233). Yoshida, W., Dolan, R. J., & Friston, K. J. (2008). Game theory of À New York, NY: Academic Press. mind. PLoS Computers in Biology, 4(12), e1000254. Available from: Ojemann, G., Ojemann, J., Lettich, E., & Berger, M. (1989). Cortical http://dx.doi.org/10.1371/journal.pcbi.1000254. language localization in left, dominant hemisphere: An electrical

A. INTRODUCTION