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Neurobiology of Language NEUROBIOLOGY OF LANGUAGE Edited by GREGORY HICKOK Department of Cognitive Sciences, University of California, Irvine, CA, USA STEVEN L. SMALL Department of Neurology, University of California, Irvine, CA, USA AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier SECTION A INTRODUCTION This page intentionally left blank 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 medi- cine, 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) presented to the Hoˆpital 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)andGrashey (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 Neurobiology of Language. DOI: http://dx.doi.org/10.1016/B978-0-12-407794-2.00001-8 3 © 2016 Elsevier Inc. All rights reserved. 4 1. THE NEUROBIOLOGY OF LANGUAGE Concept brain and behavior. Although the earliest functional ana- field tomical studies were conducted with positron emission tomography, the most recent work uses functional mag- netic resonance imaging (fMRI), a noninvasive approach that does not require intravenous administration of 4 6 radioactive agents. These methods permit the investiga- tion in vivo of brain regions that participate in the perfor- mance 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 contem- porary research in brainÀbehavior relations for language 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 por- tion—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
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