srget

nitiothernickh-motor regions. Recent work, in contrast, has established a role for anteriorntral srecogniare assoods orday ap

2013 Elsevier Inc. All rights reserved.

y cortes inski et1), haguagePoep

y betwing in

word recognition (Geschwind, 1970; Peneld & Roberts, 1959),

terior ST, on the other hand, was found to be selective for soundlocation in monkeys (Rauschecker & Tian, 2000; Recanzone,2000; Tian et al., 2001). This paradox was noted in an early paperon dual-stream concepts in audition and language:

Speech perception in humans is traditionally associated with theposterior portion of the [superior temporal] region, often referred

man and mmonkey d

human anterior ST in word recognition. Still, apparent contween classical neurological models and the monkey worka spectrum of conclusions about the relative involvement orior and posterior ST in word recognition (Binder et al., 2000; Hick-ok & Poeppel, 2000; Price, Winterburn, Giraud, Moore, & Noppeney,2003; Scott et al., 2000; Thierry, Giraud, & Price, 2003; Wise et al.,2001). This enigma has been partially resolved by the meta-analysisof DeWitt and Rauschecker (2012), which, based on a large amountof data, clearly associates word-form recognition with anterior ST.What, if anything, the dorsal stream contributes to language com-prehension is now emerging as a key question. Increasingly, the

Corresponding authors. Fax: +1 202 687 0617.E-mail addresses: id32@georgetown.edu (I. DeWitt), rauschej@georgetown.edu

Brain & Language 127 (2013) 181191

Contents lists availab

La

w.e(J.P. Rauschecker).but results from monkeys showed anterior, not posterior, ST tobe most selective for communication calls (Tian et al., 2001). Pos-

generally substantiated comparisons between huauditory cortex, afrming the role implied by the0093-934X/$ - see front matter 2013 Elsevier Inc. All rights reserved.http://dx.doi.org/10.1016/j.bandl.2013.09.014onkeyata forict be-led tof ante-identication in humans is immediately apparent and has led to ahierarchical model of speech processing in the auditory ventralstream that is now almost universally accepted (DeWitt &Rauschecker, 2012; Hickok & Poeppel, 2007). Adoption of the mod-el, however, was not without controversy. Classical neurologyidentied posterior superior temporal cortex (ST) as the site of

evolution). However, the selectivity observed in macaque posteriorST for the location of sound sources was subsequently also observedin humans by numerous studies using functional magnetic reso-nance imaging (fMRI), as well as electro- and magneto-encephalog-raphy (Ahveninen et al., 2006; Arnott, Binns, Grady, & Alain, 2004;Deouell, Heller, Malach, DEsposito, & Knight, 2007; Krumbholzet al., 2005; Tata & Ward, 2005; Zimmer & Macaluso, 2005). This1. Introduction

The Dual Stream model of auditorbasis of neurophysiological studie(Rauschecker, 1997, 1998b; RomanDurham, Kustov, & Rauschecker, 200ence on current understanding of lancortex (Binder et al., 2000; Hickok &Rosen, & Wise, 2000). The similaritnisms of communication-call processx, rst proposed on thethe macaque monkeyal., 1999; Tian, Reser,s had a profound inu-organization in humanpel, 2000; Scott, Blank,een single-cell mecha-monkeys and phoneme

to as Wernickes area. In rhesus monkeys. . .neurons in thisregion. . .are highly selective for the spatial location ofsounds. . .Neurons in the anterior belt regions, on the other hand,are most selective for [monkey calls] (Rauschecker & Tian, 2000,pp. 1180411805).

Initially, one could have taken this apparent dissociation betweenhuman and monkey cortex as grounds for dismissing the applicabil-ity of the monkey model to human speech processing (i.e., divergenttwo cortical modules, an auditory word-form area (AWFA) in the auditory ventral stream and an innerspeech area in the auditory dorsal stream.Wernickes area revisited: Parallel stream

Iain DeWitt , Josef P. Rauschecker Laboratory of Integrative Neuroscience and Cognition, Department of Neuroscience, Geo3790 Reservoir Road NW, Washington, DC 20007, USA

a r t i c l e i n f o

Article history:Available online 23 October 2013

Keywords:Dual-stream modelWord recognitionLanguage comprehensionPure word deafnessWernickes aphasia

a b s t r a c t

Auditory word-form recogporal gyrus (STG), later furcically paraphasia), Weroutput from frontal speecSTG, part of the auditory veprimates and word-formspeech and motor controlwithout quantitative methlocalized functions that to

Brain &

journal homepage: wwtream, in the recognition of species-specic vocalizations in nonhumantion in humans. Recent work also suggests monitoring self-producedciated with posterior STG, part of the auditory dorsal stream. Workingevidence of sensory cortex hierarchical organization, Wernicke co-pear dissociable. Wernickes area thus may be better construed asand word processing

own University Medical Center, New Research Building, WP19,

n was originally proposed by Wernicke to occur within left superior tem-specied to be in posterior STG. To account for clinical observations (spe-e proposed his sensory speech center was also essential for correcting

le at ScienceDirect

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computational role of posterior ST in language is understood to per-tain to its role in sensorimotor integration and control (Hickok &Poeppel, 2007; Rauschecker & Scott, 2009). Recent proposals furtheremphasize a role for the dorsal stream in sequence processing andsyntax, particularly with respect to the computation of sentence-internal relations for syntactically complex sentences (Bornkessel-Schlesewsky & Schlesewsky, 2013; Friederici, 2012; Rauschecker,2011).

Here, we present an analysis of speech processing within thedual-stream architecture of auditory cortex with the aim of clarify-ing the neural substrates of auditory word-form recognition. Thepresent work builds on a previous study from our lab (DeWitt &Rauschecker, 2012). We extend that work by formal dissection ofthe roles proposed for Wernickes area and by extensive critical re-view of results from clinical neuroscience. First, we consider word-form recognition within the auditory ventral stream (see Section 2).Emphasis is given to outstanding questions, particularly with re-spect to the relationship between ndings from functional imaging(DeWitt & Rauschecker, 2012) and contemporary quantitativendings from aphasiology and neurosurgery (see Section 3). Next,

processing pathways: a dorsal stream optimized for sensorimotorintegration, including spatial processing, and a ventral stream,optimized for object (or pattern) recognition (see Fig. 1) (Kaas &Hackett, 1999, 2000; Rauschecker, 1997, 1998a, 1998b; Rauscheck-er & Scott, 2009; Rauschecker & Tian, 2000; Rauschecker, Tian, &Hauser, 1995; Romanski et al., 1999; Tian et al., 2001). This dual-stream organization resembles the functional organization of vi-sual cortex (Goodale & Milner, 1992; Ungerleider & Mishkin,1982; Van Essen & Gallant, 1994) and suggests greater homologiesbetween the sensory systems than could previously be assumed.

Current perspectives on speech processing have incorporatedthe dual-stream architecture of auditory cortex derived from non-human primate work (Binder et al., 2000; Hickok & Poeppel, 2007;Scott & Wise, 2004; Wise et al., 2001), making it the consensusview (but see Nelken, Fishbach, Las, Ulanovsky, & Farkas, 2003;Whalen et al., 2006). The precise course of the auditory ventralstream, however, remained a question of debate: some authors in-cluded in it posterior STS (Hickok & Poeppel, 2007; Wise et al.,2001), a site consistent with ndings from classical neurology(Geschwind, 1970; Peneld & Roberts, 1959); others rejected pos-

elds& Sauditlite, Monacom

182 I. DeWitt, J.P. Rauschecker / Brain & Language 127 (2013) 181191we perform a historical review of Wernickes (1874) characteriza-tion of his sensory speech center, which helps to clarify what func-tions should be accounted for in the localization of Wernickes areaand evaluation of the Wernickes area construct (see Section 4).The review also highlights some early misconceptions and over-simplications about auditory processing that, while reasonablefor the time, continue to color contemporary conceptions of Wer-nickes area and speech processing. Lastly, we discuss how the dis-parate functions Wernicke assigned to his sensory speech center,namely word-form recognition, supervision of speech productionand inner speech, segregate and embed within the dual-streammodel (see Section 5). Consistent with nonhuman primate electro-physiology and neuroanatomy, we conclude that word-form recog-nition, the principal attribute of Wernickes area, should beassigned to the auditory ventral stream, whereas the regulationof speech production and inner speech are associated with theauditory dorsal stream.

2. Word-form recognition and the auditory ventral stream

Concurrent with early functional imaging, work in nonhumanprimate electrophysiology made breakthroughs into the functionalorganization of nonprimary auditory cortex, identifying two main

A

Fig. 1. A composite illustration of human auditory cortex and macaque auditory posterior axis of the superior temporal plane (Fullerton & Pandya, 2007; Galaburda[core, Brodmanns area (BA) 41] located along Heschls gyrus (HG) and secondary apolare (PP) and planum temporale (PT). To facilitate comparisons with the macaquehuman anatomy (core: A1, R, RT, RTp; lateral belt: CL, ML, AL, RTL; medial belt: CMmacaque, scaled with respect to the volume of human core (Penhune, Zatorre, MacDtuning characteristics (Rauschecker, Tian, & Hauser, 1995; Chevillet et al., 2011). The

superior temporal plane. Fields exhibiting heightened selectivity for monkey calls are shoarea RTp (Kikuchi, Horwitz, & Mishkin, 2010). For orientation, the cortical patch shown inAdditional points of reference include the circular sulcus (CS), insular cortex (Ins), supraterior STS, concluding word-form recognition occurs in anteriorSTG (Binder et al., 2000; Mesulam, 1998; Scott & Wise, 2004).While posterior STS happens to be ventral to the Sylvian ssure,the ventral and dorsal streams are dened by cortico-cortical con-nections originating in the lateral belt areas of auditory cortex andby histoarchitectonic criteria (Kaas & Hackett, 2000; Rauschecker &Tian, 2000). These criteria characterize the posterior ST region inhumans as part of the dorsal stream and anterior STG as part ofthe ventral stream (see Fig. 2A and B).

In a recent paper, we leveraged the observation of large-scalesimilarity in the auditory and visual systems functional architec-tures to address the problem of localizing auditory word-form rec-ognition within the ventral stream (DeWitt & Rauschecker, 2012).We reviewed and synthesized literature bearing on the hypothesisthat auditory and visual word recognition are equivalent problemswith similar cortical solutions. This led us to hypothesize, as haveothers (Cohen, Jobert, Le Bihan, & Dehaene, 2004; Mesulam, 1998),that a cortical region supporting sensory aspects of auditory wordrecognition (i.e., an AWFA) should exist with properties compara-ble to those identied for the visual word-form area (VWFA)(Dehaene, Cohen, Sigman, & Vinckier, 2005) and, more generally,for pattern recognition in the visual ventral stream (DiCarlo,Zoccolan, & Rust, 2012; Riesenhuber & Poggio, 2002;Wallis & Rolls,

B

. Relative to the macaque, human auditory cortex is rotated 45 off the anterior-nides, 1980; Hackett, 2011; Rademacher et al., 2001) with primary auditory cortexory cortex (lateral and medial belt, BA 42 and 52, respectively) located in planumrature, names of functionally-dened macaque subelds are shown on a atmap ofM, RM, RTM) (A). Subeld delineation is estimated from relative eld sizes in theld, & Evans, 1996; Rademacher et al., 2001) and functionally localized according toposite gure implies a course for the human ventral and dorsal streams along thewn in yellow: lateral belt eld AL (Tian et al., 2001; Tsunada, Lee, & Cohen, 2011) andatmap (A) is outlined on the cortical surface (dashed line with scissor markers) (B).marginal gyrus (SMG) and STG.

n &A

I. DeWitt, J.P. Rauschecker / Brai1997). Specically, this AWFA should demonstrate selectivity forauditory words (i.e., it should respond more to auditory words thanto other sounds). Further, it should demonstrate invariance to cer-tain acoustical changes (i.e., its response should be more sensitiveto acoustical differences that affect the phonetic content of utter-ances than to acoustical differences which do not).

In our analyses, we assessed selectivity with respect to eitheracoustically matched articial stimuli or non-speech natural stim-uli. Invariance was assessed with respect to adaptation phenomena(Miller, Li, & Desimone, 1991), which can be used to probetolerance for non-categorytransformative physical stimulus

B

C

D

STS

PT

Fig. 2. Anatomical predictions for the site of auditory word-form recognition. (A) Inthe macaque, communication call processing is strongly associated with anterior-lateral portions of the superior temporal plane (circled) (adapted from Rauschecker& Tian, 2000). (B) The putatively homologous human site resides at the anterior-lateral aspect of Heschls gyrus (circled) (adapted from Galaburda & Sanides, 1980).(C) This site is within the territory originally proposed by Wernicke (shaded regionmarked x) (adapted from Wernicke, 1881) but (D) is inconsistent with the locationgiven for Wernickes area by Geschwind (shaded region marked 4) (adapted fromGeschwind, 1969).deformations and sensitivity to category-transformative deforma-tions (Grill-Spector & Malach, 2001). Further, the hierarchical orga-nization of auditory cortex implies increasing representationalcomplexity along the auditory ventral stream (Binder et al.,2000; Kaas & Hackett, 2000; Rauschecker & Scott, 2009; Raus-checker & Tian, 2000; Rauschecker et al., 1995; Chevillet, Riesenh-uber, & Rauschecker, 2011) similar to that found along the visualcortical hierarchy (Hubel & Wiesel, 1962; Riesenhuber & Poggio,2002). Therefore, where possible, we assessed processing for pho-neme, word, and phrase stimuli separately. As phoneme recogni-tion is a prerequisite of word recognition, we hypothesized peakphoneme processing to localize to an area proximate to primaryauditory cortex, relative to the site of peak processing for words.Phrase processing, in contrast, includes phoneme and word recog-nition, but it also strongly engages semantic and syntactic process-ing. Accordingly, we hypothesized phrase processing to engage thesites associated with phoneme and word recognition as well ashigher-order regions of ST. Separate consideration of phoneme,word and phrase processing, therefore, made the assessment of ef-fects pertaining to word recognition both more precise and moretractable.

To quantitatively assess our predictions, we focused on resultsfrom functional brain imaging. To systematically evaluate prior re-sults, we used an anatomically unbiased coordinate-based meta-analytic approach (Turkeltaub, Eden, Jones, & Zefro, 2002). Themethod found functional imaging results of auditory word recogni-tion to be consistent with principles of hierarchical processing (seeFig. 3) (DeWitt & Rauschecker, 2012). Results supported a left-biased, three-stage model, with analysis of phonemes occurringin mid-STG, lateral to Heschls gyrus, word recognition occurringin anterior STG, and phrase processing beginning in anterior STS(c.f., Miglioretti & Boatman, 2003). This diverged from the classicalmodel of language organization (Geschwind, 1970; Peneld & Rob-erts, 1959) and some contemporary perspectives (Hickok & Poep-pel, 2007; Wise et al., 2001), which locate word recognition inposterior ST.

Prior to the advent of contemporary imaging methods, infer-ence that posterior ST was the site of auditory word recognitionwas warranted by available evidence and methodology. Lesionsresulting in auditory comprehension decits (as well as poor ver-bal repetition and paraphasiainaccurate word selection duringspeech; i.e.,Wernickes aphasia) show greatest overlap in posteriorSTG (Robson, Sage, & Lambon-Ralph, 2012). Simple lesion-overlap(density) mapping, however, is spatially biased, owing to arterialanatomy. For instance, the middle cerebral artery bifurcates andnarrows as it progresses along the Sylvian ssure, likely increasingthe probability of posterior infarcts. Thus, while the center of massof most lesions that produce auditory comprehension decits maybe in posterior STG, this could be an epiphenomenon and compre-hension decits might be better explained by the anterior extent ofthese lesions (c.f., Dronkers, Wilkins, van Valin, Redfern, & Jaeger,2004). Contemporary methods mitigate spatial bias through theinclusion of control samples (Bates et al., 2003; Rorden & Karnath,2004). These methods utilize variance in symptom severity to fac-tor out lesion sites that are shared across aficted individuals butwhich do not contribute to task-specic impairments (for addi-tional discussion, see Section 3).

In the rst decade and a half of functional imaging, as the eldand its methodology matured, reservation in interpretation anddeference to well-established theories was prudent. Increasingly,however, functional imaging indicated that revisions to theclassical model were required (Belin, Zatorre, & Ahad, 2002; Belin,Zatorre, Lafaille, Ahad, & Pike, 2000; Binder, Frost, Hammeke, Rao,

Language 127 (2013) 181191 183& Cox, 1996; Binder et al., 1994; Binder et al., 1997, 2000; Dmonetet al., 1992; Mazziotta, Phelps, Carson, & Kuhl, 1982; Mummery,Ashburner, Scott, & Wise, 1999; Petersen, Fox, Posner, Mintun, &

n &184 I. DeWitt, J.P. Rauschecker / BraiRaichle, 1988; Scott et al., 2000; Wise et al., 1991, 2001). As dis-cussed, results from nonhuman primates were prompting revisionsin understanding of the functional and anatomical organization ofauditory cortex (Kaas & Hackett, 2000; Rauschecker & Tian, 2000;Tian et al., 2001). These results provided a framework for amend-ment of models of speech processing (Binder et al., 2000; Boatman,2004; Hickok & Poeppel, 2000, 2004, 2007; Mesulam, 1998; Scott,2005; Scott & Wise, 2004; Wise et al., 2001). While the revisedmodels of speech processing generally adopted a dual-streamframework, discrepancy persisted about the site of word-form rec-ognition within the auditory ventral stream. Some authors main-tained a site close to canonical Wernickes area (Hickok &Poeppel, 2000, 2007; Wise et al., 2001). Others adopted an anterior

and neurosurgical studies to conclude that anterior STGs involve-ment is causal. Some reports provide compelling evidence in sup-

Fig. 3. Meta-analyses of auditory-word processing. Analyses of studies comparingbrain response to speech stimuli versus matched control sounds (AC), indicative ofselectivity for speech sounds, found a leftward bias and an anterior progression inpeak effects with phoneme-length studies peak focus density in left mid-STG (A),word-length studies peak density in left anterior STG (B), and phrase-lengthstudies peak density in left anterior STS (C). Peak density for studies investigatingphonetically specic adaptation (D), indicative of invariant representation, wasfound in left mid- to anterior STG. Peak density for areal specialization studies (E),which compared brain response to speech stimuli versus other natural non-speechsounds, also indicative of selectivity for speech sounds, was greatest in left STG.Intensity represents ALE value. Adapted from DeWitt and Rauschecker (2012).port of causality (Boatman, 2006; Dronkers et al., 2004;Hamberger, Goodman, Perrine, & Tamny, 2001; Hamberger, McC-lelland, McKhann, Williams, & Goodman, 2007; Hamberger, Seidel,Goodman, Perrine, & McKhann, 2003; Hamberger, Seidel, McKh-ann, Perrine, & Goodman, 2005; Kmmerer et al., 2013; Malowet al., 1996; Matsumoto et al., 2011; Miglioretti & Boatman,2003; Rogalski et al., 2011). A direct relationship, however, hasyet to be demonstrated between the anatomical location of audi-tory word-form recognition (indicated by single-subject brainimaging) and behavioral impairment resulting from surgical proce-dures, such as reversible electrical interference or clinical resec-tion, as has been shown for the VWFA (Gaillard et al., 2006) andthe fusiform face area (Parvizi et al., 2012).

The relative scarcity of causal evidence for anterior STG involve-ment in word recognition is partly attributable to the typical reli-ance of intraoperative language mapping on outcome measuresthat assess non-auditory processing, namely single word reading,visual object naming and speech arrest (Hamberger et al., 2007; Sa-nai, Mirzadeh, & Berger, 2008). Those studies that assessed auditoryprocessing typically investigated acousticphonetic feature detec-tion (Boatman, 2006; Boatman, Hall, Goldstein, Lesser, & Gordon,1997; Boatman, Lesser, & Gordon, 1995; Miglioretti & Boatman,2003) or sentence comprehension (Hamberger et al., 2001, 2003,2005, 2007; Malow et al., 1996; Matsumoto et al., 2011; Miglioretti& Boatman, 2003). Though important levels of inquiry, neither levelspecically assesses word-form recognition. The former assessesthe stage prior to word-form recognition (i.e., phoneme recogni-STG localization (Binder et al., 2000; Scott & Johnsrude, 2003; Scott& Wise, 2004; Wise, 2003). Although evidence accumulated on theside of anterior localization, skepticism remained (Hickok, 2010).Our meta-analysis systematically and quantitatively weighedtwo-decades of published ndings with bearing on the site of audi-tory word-form recognition and concluded the preponderance ofevidence supports anterior STG localization.

In the same time period, work on the visual systems analogousproblem, visual word-form recognition, progressed more effec-tively. There, an area within the visual ventral stream, the VWFA,has come to be widely regarded and intensively studied as the cru-cial site for visual word-form recognition (Cohen et al., 2000; Deh-aene et al., 2005; McCandliss, Cohen, & Dehaene, 2003). Althoughinterpretational questions remain (Baker et al., 2007; Dehaene &Cohen, 2011; Price & Devlin, 2011), localization of the VWFAwithinventral occipito-temporal cortex (VOT) is now largely uncontrover-sial. Identication of this VOT site, analogous to macaque infero-temporal cortex, permitted detailed, mechanistic investigations toproceed, producing a prolic literature (Baker et al., 2007; Binder,Medler, Westbury, Liebenthal, & Buchanan, 2006; Braet, Wage-mans, & Op de Beeck, 2012; Cohen et al., 2004; Dehaene et al.,2010; Gaillard et al., 2006; Glezer, Jiang, & Riesenhuber, 2009;Rauschecker, Bowen, Parvizi, &Wandell, 2012; Turkeltaub, Flowers,Lyon, & Eden, 2008; Vinckier et al., 2007; Wandell, Rauschecker, &Yeatman, 2012). Resolving debate about the AWFAs location with-in STmay similarly position the eld tomake advances in unlockingthe nature of representation within the auditory ventral stream.

3. Causal involvement of anterior STG in word recognition

Although an unprecedented amount of evidence is nowamassed indicating the involvement of anterior STG in word recog-nition, there remains a paucity of direct evidence from neurological

Language 127 (2013) 181191tion) while the latter assesses phrase comprehension, which in-cludes semantic and syntactic processing. More rened methods(Bormann & Weiller, 2012; Goll et al., 2010; Hickok et al., 2008;

documented cases of patients with circumscribed cortical lesions are simi-

n &Miglioretti & Boatman, 2003; Rogalski et al., 2011; Thothathiri,Kimberg, & Schwartz, 2012) will be required in future investiga-tions for the specic evaluation of auditory word-form recognition.It should be noted, however, that resection of sites implicated inauditory sentence comprehension by electrical interference doesincrease the incidence of post-operative impairment in auditorycomprehension (Hamberger et al., 2005). Although the sitesresected in that study were not reported in detail, similar studiesreport a greater likelihood of impairment on auditory sentencecomprehension from stimulation of anterior ST (Hamberger et al.,2001, 2003, 2007; Miglioretti & Boatman, 2003).

Anterior temporal lobectomies are relatively common. Rarely,however, do studies report post-operative language decline. Thismight be attributable to three factors. First and foremost, the can-didate AWFA extends from 45 mm distal of the temporal pole to70 mm distal (DeWitt & Rauschecker, 2012). Standard resectionstypically remove 3555 mm of the anterior temporal lobe, withthe majority of resections removing 45 mm or less, sparing muchof the area in question (Alpherts et al., 2008; Bidet-Caulet et al.,2009; Binder et al., 2011; Helmstaedter et al., 2008; Hermann, Wy-ler, & Somes, 1991; Kho et al., 2008; Pataraia et al., 2005; Schwartz,Devinsky, Doyle, & Perrine, 1998; Seidenberg et al., 1998). Further,resections are sometimes performed differentially, sparing a great-er portion of STG relative to the middle and inferior temporal gyri,also decreasing the likelihood of resections including the candidateAWFA (Alpherts et al., 2008; Bidet-Caulet et al., 2009; Binder et al.,2011; Pataraia et al., 2005; Schwartz et al., 1998). Second, intraop-erative language mapping may indicate language function and,thereby, spare the candidate AWFA from resection. Third, there isa dearth of reported outcomes at short-term follow-up(t < 6 weeks). Researchers tend instead to report outcomes forlonger recovery durations (t > 6 months) (Bidet-Caulet et al.,2009; Davies, Risse, & Gates, 2005; Hermann et al., 1991; Pataraiaet al., 2005; Schwartz et al., 1998). Given the relative competencyof the non-dominant hemisphere during the incapacitation of thedominant hemisphere (Hickok et al., 2008), compensatory plastic-ity in the contralateral hemisphere could account for a low inci-dence of post-operative impairment at long-term follow-up, evenwhen resections include the candidate AWFA. Interestingly, classi-cal models have a similar evidentiary problem. There is a dearth ofevidence associating posterior ST resection with auditory compre-hension decits. Indeed, when studies report posterior ST resec-tion, they often argue they observe an absence of languagedecline (Petrovich, Holodny, Brennan, & Gutin, 2004; Sarubboet al., 2012).

Analogous to the temporal lobectomy literature, studies ofaphasia lesion mapping have not traditionally emphasized therole of anterior ST in auditory word comprehension. Simple den-sity mapping of Wernickes aphasia lesions nds the center ofmass of lesions to be in posterior ST, but the lesions commonlyextend into anterior STG (Ogar et al., 2011; Robson et al., 2012).Similarly, lesion mapping that utilizes both control samples andcontinuous symptom severity data implicates both anterior andposterior ST in auditory sentence comprehension (Dronkerset al., 2004; Saygin, Dick, Wilson, Dronkers, & Bates, 2003). Fur-ther, with respect to comprehension decits, this work expresslydissociates posterior STG from surrounding regions: lesions ofposterior STG were not found to affect comprehension. Impor-tantly, work specically investigating auditory word recognition(as opposed to sentential comprehension) exclusively implicatesanterior ST (Rogalski et al., 2011). In patients for whom auditoryword recognition is spared, decits in auditory sentence compre-hension, which can therefore be attributed to decits in syntactic

I. DeWitt, J.P. Rauschecker / Braiprocessing, are associated with lesions of posterior ST and inferiorparietal lobule (IPL) (Thothathiri et al., 2012). This result is con-sistent with the view that anterior ST must be spared for auditorylarly rare. Recent literature, utilizing modern brain imaging, pro-vides two general impressions (Clarke, Bellmann, Meuli, Assal, &Steck, 2000; Engelien et al., 1995; Fung, Sue, & Somerville, 2000;Iizuka, Suzuki, Endo, Fujii, & Mori, 2007; Kaga, Nakamura, Takay-ama, & Momose, 2004; Kim et al., 2011; Miceli et al., 2008; Palma,Lamet, Riverol, & Martnez-Lage, 2012; Praamstra, Hagoort, Maas-sen, & Crul, 1991; Slevc, Martin, Hamilton, & Joanisse, 2011; Stefa-natos, Gershkoff, & Madigan, 2005; Suh et al., 2012; Wang, Peach,Xu, Schneck, & Manry, 2000). First, while bilateral ST lesions arecommon (Buchman et al., 1986; Geschwind, 1965; Poeppel,2001), left hemisphere lesions can be sufcient (Palma et al.,2012; Slevc et al., 2011; Stefanatos et al., 2005). Second, whilesome cases involve lesions of mid- to posterior ST (Kim et al.,2011; Slevc et al., 2011) and others involve mid- to anterior ST(Engelien et al., 1995; Iizuka et al., 2007; Palma et al., 2012; Stefa-natos et al., 2005), the commonly affected region appears to be midSTG, lateral to Heschls gyrusthe putative site of phoneme recog-nition (Boatman, 2006; Boatman et al., 1995, 1997; DeWitt & Raus-checker, 2012; Liebenthal, Binder, Spitzer, Possing, & Medler, 2005;Liebenthal et al., 2010; Miglioretti & Boatman, 2003). In sum, clin-ical results are highly suggestive of a causal role for mid- to ante-rior STG in word recognition. What remains to be demonstrated,however, is direct correspondence between results from fMRIand behavioral impairment following lesion or functionalinactivation.

4. A brief history of Wernickes area

In the 1860s, Paul Broca established the presence of a motorspeech center in the left inferior frontal gyrus (IFG) (for review,see Dronkers, Plaisant, Iba-Zizen, & Cabanis, 2007). Reecting onBrocas observations, Carl Wernicke (1874) postulated a comple-mentary sensory speech center, for the storage and collection ofauditory images (representations) of speech soundsreferred totoday as Wernickes area. Initially, Wernicke recognized STG in totoas the site of auditory imagery and did not attempt to specicallylocalize auditory word representations within STG. Rather, henoted only that a circumscribed region ought to exist somewherewithin STG (see Fig. 2C), analogous to the circumscribed speech-motor region within IFG:

The rst temporal gyrus [STG], which is sensory in nature, may beregarded as the center of acoustic images. . . [It] may be regarded asthe central terminal of the acoustic nerve, and the rst frontalword recognition to be intact. When either single-word auditorycomprehension is factored out (Fridriksson et al., 2010) or generalauditory comprehension is spared (Buchsbaum, Padmanabhan, &Berman, 2011), word repetition decits are associated with le-sions of posterior ST and IPL. Again, this is consistent with theview that sensory aphasias that spare auditory word recognitionshould spare anterior ST. These results also dissociate posteriorST lesions with auditory word recognition decits. Finally, whenconsidering subcortical lesions, the integrity of tracts associatedwith the auditory ventral stream is closely associated with audi-tory comprehension, whereas the integrity of tracts associatedwith the dorsal stream is associated with vocal repetition (Km-merer et al., 2013).

While there is a sizable literature associated with pure worddeafness (see Appendix A), it is nonetheless a rare condition(Buchman, Garron, Trost-Cardamone, Wichter, & Schwartz, 1986;Poeppel, 2001). Consequently, there are no group-level lesion-mapping studies of the disorder (i.e., only case studies). Well-

Language 127 (2013) 181191 185gyrus [IFG], including Brocas area, as the central terminal of thenerves controlling the speech musculature (Wernicke, 1874/1977, p. 103).

the benet of far grater understanding of neuroanatomy and corti-cal processing than either Wernicke or Geschwind had access to

186 I. DeWitt, J.P. Rauschecker / Brain &This view is claried and reiterated in subsequent passages.As details of the ascending auditory tracts and the hierarchical

organization of auditory cortex were not yet known in 1874, Wer-nicke assumed direct innervation of the greater extent of STG byascending bers. His model, therefore, lacked an equivalent to pri-mary auditory cortex and a theory of representational transforma-tion along auditory cortex, resulting in large inaccuracies.Wernicke, for reasons supported by behavioral observations butanatomically awed, nonetheless posited that only a portion ofSTG functions as a sensory speech center:

The area containing acoustic imagery. . .is not identical to the broadradiation of the acoustic nerve itself, since complete loss of acousticimagery with intact bilateral hearing has been observed in apha-sia. . .In spite of destruction of the central acoustic radiation, whichcarries the sounds of words, perception of noise and musical tonewould still be intact (Wernicke, 1874/1977, p. 105).

Wernicke implies the functional consequence of incomplete deaf-ferentation of STG is auditory agnosia. Within the context of sen-sory aphasia, according to Wernicke, the prominent feature isverbal auditory agnosia (word deafness). Wernicke describes simi-lar consequences for cortical lesions:

When. . . the cortex of the rst temporal convolution [STG] isdestroyed, memory for the acoustic images designating. . .objectsis erased, though memory for concepts may continue existing in fullclarity. This is because the acoustic image of the name for the con-cept of an object is generally incidental to the concept, whereas pal-pable, tangible imagery is intrinsic (Wernicke, 1874, p. 22).1

Wernicke is clearly dissociating word-form representation fromsemantic representation. He, therefore, principally characterizeshis sensory speech center as an AWFA.

Wernicke subsequently ascribes a secondary function to thesensory speech center: a corrective role in the activation of motorrepresentations during speech production:

Apart from lack of understanding, the patient [with sensory apha-sia] has aphasic phenomena in speaking, owing to an absence ofunconscious correction exerted by the speech sound image (Wer-nicke, 1874, p. 23).2

The aphasic phenomena in speaking to which Wernicke refers areparaphasiasKussmaul (1877) had yet to coin the termwhichcommonly co-occur with auditory comprehension decits. At thetime, as is clear from later writings (Wernicke, 1886/1977), all thecases of sensory aphasia that Wernicke had seen to date includedparaphasia. Thus, Wernickes ascription of a corrective role to hissensory speech center reects both a clinically motivated desidera-tum and the assumption that only a single functional module is le-sioned in Wernickes aphasia.

Later works include ve main addenda (Wernicke, 1886/1977,1906/1977). First, responding to Kussmaul (1877) and Lichtheim(1885), Wernicke discussed pure word deafness (see AppendixA), which he referred to as subcortical sensory aphasia. He attrib-uted pure word deafness to deafferentation of the sensory speechcenter. Second, he developed his notion of the corrective inuence(during word selection) exerted by speech sound imagery onspeech motor imagery. Over development, he argued, the repeatedassociation of auditory and motor word representations conjoinsthem into word-concept representations (c.f., Lichtheim, 1885),which form the basis of inner speech (reviewed by Geva et al.,2011). Wernicke believed the inner-speech faculty is spared in ac-1 Authors translation.2 Authors translation.wemight conclude that the functions Wernicke subsumes within asingle area are actually performed by multiple cortical areas (c.f.,Goldstein, 1927, 1948; Mesulam, 1998; Wise et al., 2001). Thehypothesis most strongly supported by available empirical datafor the location of Wernickes AWFA is anterior STG (DeWitt &Rauschecker, 2012). This region, however, is neither a strong can-didate site for encoding representations that resemble Wernickesword-concepts (i.e., inner speech) nor a strong candidate site forperforming the corrective function Wernicke ascribes to them.

Cortical monitoring of self-produced speech and the correctionof speech motor programs is most parsimoniously viewed as a dor-sal-stream function (Hickok & Poeppel, 2007; Rauschecker & Scott,2009; Wise et al., 2001). To coordinate speech production, motorcontrol theory (Golnopoulos, Tourville, & Guenther, 2010; Guen-ther, 1994; Hickok, 2012; Rauschecker, 2011; Rauschecker & Scott,2009) posits the mapping of auditory representations of self-pro-duced speech sounds into the frame of reference of the speecharticulators (Cohen & Andersen, 2002; Dhanjal, Handunnetthi,Patel, & Wise, 2008). Multimodal articulator-encoded speech rep-resentations are then reconciled with expectations, derived fromefference copy, of the intended consequences of the activated mo-tor representation. Finally, the difference between expectation andfeedback (error) is transmitted to frontal cortex and used in updat-ing motor output. The temporo-parietal sites most strongly associ-ated with auditory feedback and speech production are posteriorPT, posterior STG, and SMG (Golnopoulos et al., 2010, 2011; Ham-berger et al., 2003; Takaso, Eisner, Wise, & Scott, 2010; Towle et al.,2008; Zheng, Munhall, & Johnsrude, 2010), regions associated withquired pure word deafness, explaining the absence of paraphasia.Third, he expressly localized his sensory speech center to leftSTG, something only implied previously. He also, however, allowedthat transient aphasia (recovery) might be explained by plasticityin right STG. Fourth, he circumscribed the portion of STG positedto contain his sensory speech center. Citing numerous pathologi-cal ndings at hand, but without identifying them, he describesthe center as being conned to the posterior third of half of[sic] STG (p. 235) and an adjoining strip of medial temporalgyrus (Wernicke, 1906/1977, p. 225). As Wernicke reproducedand endorsed the anatomical diagrams of Von Monakow and Dj-rine in his section on neuroanatomy (p. 272), numerous patholog-ical ndings may have been an allusion to their work. Lastly, hisviews of the relevance of his sensory speech center to written com-prehension, which were ambivalent in 1874, evolved (see Appen-dix B). Wernickes ultimate position was that the sensory speechcenter was essential for orthography-to-phonology mapping (i.e.,phonological reading) and that this was attributable to the centersrole in inner speech.

In the century following Wernickes observations, Wernickesarea was increasingly understood to be limited to the posteriorthird of STG with various formulations about which adjacent corti-cal regions should be included as well (for reviews, see Bogen & Bo-gen, 1976; Rauschecker & Scott, 2009). In the 1960s, Geschwindrevived the WernickeLichtheim model of aphasia (reviewed byCatani & Mesulam, 2008; Eling, 2011), presenting the most focalinterpretation, including only the most posterior aspect of STG(see Fig. 2D).

5. Paraphasia, inner speech and the auditory dorsal stream

Where is Wernickes area? Answering this question todaywith

Language 127 (2013) 181191the auditory dorsal stream.Accordingly, paraphasia could result from dorsal-stream lesions

that disrupt circuitry involved in rectifying unintended output,

1977), Kussmaul (1877) coined the term word deafness (Wort-

ciability of components. Wernicke (1886/1977) attributed the rst

n &hypothetically, even prior to overt speech production (c.f., Licht-heim, 1885). Consistent with this, lesion mapping associates para-phasia with posterior ST and IPL (Buchsbaum, Baldo, et al., 2011).Wernickes theory of paraphasia is that word selection requiresintegrated auditory-motor representations (word-concepts), whichdevelop through repeated association during speech production(c.f., Garagnani, Wennekers, & Pulvermuller, 2007; Pulvermller,1999). This is reminiscent of the articulator-encoded speechrepresentations posited for the dorsal stream. Wernicke viewedword-concept representation as the basis for inner speech.Localizing inner speech on the basis of articulatory rehearsal inthe phonological loop (Baddeley, 2003) indicates a posterior STlocus (Buchsbaum, Olsen, Koch, & Berman, 2005). Similarly, locali-zation based on covert rhyme and homophone judgment indicatesan IPL locus (Geva et al., 2011). Thus, the qualities Wernicke asso-ciated with paraphasia (i.e., word-concepts and inner speech) sug-gest dorsal-stream localization. Further, phonological reading,which Wernicke also associates with his speech center via innerspeech, also localizes to posterior ST and IPL (see Appendix B).

In a dispute with Kussmaul over terminology for what is nowcalled Wernickes aphasia, Wernicke said:

Word-deafness describes only one part of that which we seeas an indivisible, unitary picture: for in addition to theirword-deafness, such patients are also always aphasic [parapha-sic] (Wernicke & Friedlander, 1883/1977, p. 171).

Importantly, Wernicke is speaking of sensory aphasia resulting froma cortical lesion. When Wernicke later acknowledged pure worddeafness (Wernicke, 1886/1977), he referred to it as subcorticalsensory aphasia. Thus, Wernicke never entertained the possibilitythat there could be multiple speech centers within ST, each opti-mized for different functionsand furthermore that sensory apha-sia (i.e., Wernickes aphasia) might result from extensive lesions,disrupting multiple cortical modules (see Appendix B).

Freud (1891), citing a case in which a meningioma adjacent toSTG caused pure word deafness, concluded the disorder was notdue to subcortical lesion. This, he argued, could be reconciled withcases in which cortical lesions produced Wernickes aphasiathrough the assumption that pure word deafness was attributableto incomplete lesions of Wernickes area. Goldstein (1927, 1948)recognized a cortical locus for pure word deafnessthough heacknowledged subcortical loci as welland dissociated cortical re-gions specialized for auditory word-form representation and innerspeech. From consideration of historical cases (Henschen, 1918;Poetzl, 1919), Goldstein (1948) attributed pure word deafness tolesions of the middle part of the left rst temporal convolution[STG]. . .a region close to Heschls area (p. 222). Localization of in-ner speech, he felt, could not yet be decided. Nonetheless, he spec-ulated posterior STG and adjacent areas (i.e., planum temporale,insula and IPL) were involved. While acknowledging Goldsteinsobservations, Geschwind (1970) rejected Goldsteins dissociationof auditory word-form recognition and inner speech. Instead, whileGeschwind (1965) correctly maintained mid- (or anterior) STG wasa major outow of primary auditory cortex, he surmised that itslesion would merely disconnect posterior STG from primary audi-tory cortex, a view which lacks support from modern neuroanat-omy and is impoverished with respect to representationaltransformation in cortical processing.

6. Conclusions

Wernicke originally proposed a site within left STG to subserve

I. DeWitt, J.P. Rauschecker / Braiauditory word-form recognition. On the basis of post-mortem casestudies, classical neurology came to understand the location ofWernickes area to be within posterior STG (and adjacent areascase description of pure word deafness (though not referring to itas such) to Lichtheim (1885), whose writing appeared subsequentto Kussmauls (1877) (for discussion, see Eling, 2011). Lichtheimvariously described the condition as isolated word deafnessand outer commissural word deafness. Pure word deafnesswas in use by 1889 when Starr (1889) used it to describe auditorycomprehension decits unaccompanied by impairments in read-ing, writing and speaking, consistent with what was implied byKussmauls usage. Liepmann (1898)who provided the rst ana-tomical description of pure word deafnessis also sometimes citedas coining the term. The chronology and his exact terminology(reine Sprachtaubheit), however, are incorrect.

Initially, Wernicke (Wernicke & Friedlander, 1883/1977) dis-puted the existence of pure word deafness, arguing that word deaf-ness (assuming a lesion of his sensory speech center) was alwaysaccompanied by paraphasia. Consequently, he conjectured lesionsprior to his sensory speech center would cause primary deafnesswith no trace of aphasia (p. 104)though other remarks suggesttaubheit). He used it to describe selective decits in auditory wordcomprehension, as distinct from generalized deafness. Contrary tocommon citation (e.g., Auerbach, Allard, Naeser, Alexander, & Al-bert, 1982; Coslett, Brashear, & Heilman, 1984), Kussmaul neitherused the term pure word deafness (reine Worttaubheit) nor,as noted by Wernicke (Wernicke & Friedlander, 1883/1977), de-scribed a case that was uncomplicated by other maladies (e.g., par-aphasia). Kussmauls usage, however, implied what cameclassically to be regarded as pure word deafness. For instance, hedescribed word deafness with paraphasia, which implies a disso-of cortex). Wernicke posited a secondary function for his sensoryspeech center, namely the maintenance of correct motor output.In contrast, work on speech processing in humans with functionalneuroimaging (consistent with electrophysiological work on theprocessing of species-specic vocalizations in nonhuman prima-tes), has increasingly come to implicate left anterior STG as the siteof auditory word-form recognition. Although causal involvementin word-form recognition is yet to be specically demonstratedfor this site, quantitative neurological and neurosurgical investiga-tions support such a role. Similarly, contemporary understandingof auditory cortex associates speech-motor control with posteriorST. Wernickes area, functionally dened, therefore appears to con-sist of two areas: an AWFA in anterior STG and an inner-speecharea in posterior STG/IPL. This critical reappraisal of speech pro-cessing in auditory cortex and, specically, of the Wernickes areaconstruct suggests a new framework for the assessment and diag-nosis of sensory aphasias, as well as new procedures for the intra-operative mapping of language function.

Acknowledgments

We thank Anna Seydell-Greenwald for assistance with histori-cal research. This work was supported by an award from the Wil-liam Orr Dingwall Foundation (to I.D.), National ScienceFoundation Grants BCS-0519127 and OISE-0730255 (to J.P.R.), Na-tional Institute on Deafness and Other Communication DisordersGrant 1RC1DC010720 (to J.P.R.) and National Institute on Neuro-logical Disorders and Stroke Grant 2R56NS052494 (to J.P.R.).

Appendix A. Further historical notes on pure word deafness

If we are to believe Wernicke (Wernicke & Friedlander, 1883/

Language 127 (2013) 181191 187auditory agnosia would result from cortical deafferentation (seeSection 4) (Wernicke, 1874/1977). Subsequent to Lichtheims case,Wernicke (1886/1977) claimed to have never doubted the

often associated with Wernickes aphasia (Geschwind, 1970). We,

n &theoretical possibility of pure word deafness (p. 185). Wernickeslater works (1886/1977, 1906/1977) also revised the theoreticalconsequence of lesions prior to his speech center. He now theorizedsuch lesions could result in pure word deafness from selectivedestruction of ascending bers, hypothetically affecting only thoseprojections into left temporal cortex that carry the limited portionof the auditory spectrum over which speech sounds are conveyed.Owing to Wernickes initial hesitation, Kussmaul (1877) andLichtheim (1885) may be credited with conception of the disorder,if not the term itself.

Notably, pure has taken a different emphasis in contemporaryusage (Buchman et al., 1986; Pinard, Chertkow, Black, & Peretz,2002; Poeppel, 2001; Polster & Rose, 1998). Today, pure is often re-garded as expressly connoting the sparing of non-verbal soundcomprehension, as opposed to connotation of a lack of additionalaphasic complications. This is chiey a matter of emphasis but itcarries a subtle distinction. By either usage, patients with pureword deafness have auditory word comprehension decits; theydo not present with other language decits (e.g., paraphasia oralexia); and, they have residual hearing. Modern usage further dis-tinguishes between residual hearing that simply involves the abil-ity to detect and discriminate sounds (auditory agnosia) andresidual hearing in which non-verbal sound comprehension isspared (pure word deafness). Wernicke, Kussmaul and Lichtheimsconsideration of residual sound processing did not overtly distin-guish between low-level perception and the comprehension ofspectro-temporally complex non-verbal sounds (e.g., environmen-tal sounds or music). Indeed, Wernickes theory of pure word deaf-ness includes cortical deafness for the speech-related portion ofthe human frequency range. Therefore, while current usage is notstarkly inconsistent with classical usage, its emphasis and entail-ments are a modern innovation. Under contemporary usage, casesof pure word deafness are very rare (Buchman et al., 1986; Polster& Rose, 1998). In the present analysis, we are concerned merelywith the classical dissociation.

Appendix B. Written comprehension

Wernicke initially describes his sensory speech center to benon-essential for written comprehension in readers who have at-tained uent whole-word reading:

The individual who has been exposed to minimal training in read-ing may comprehend the written word only after it has been heard.But the educated person. . .may be able to grasp general meaningafter a glance at the page without awareness of the individualwords...The rst case presents symptoms of alexia apart from hisaphasia. The second. . .reveals intact comprehension of all writtenmaterial in striking contrast to his lack of comprehension of thespoken word (Wernicke, 1874/1977, pp. 108109).

Although lacking precision and nuance, Wernicke is clearly differ-entiating phonological reading (i.e., sounding words out) fromwhole-word reading. In the case of the former but not the latter,he posits the need for acoustic images to intermediate access tomeaning.

Subsequent to Grashey (1885), Wernicke (1886/1977) substan-tially revised his views on written speech. He now stated that,without qualication, lesion of the sensory speech center causesboth alexia and agraphia. Wernickes nal work (1906/1977), how-ever, amended his position again. He re-acknowledged whole-word reading, both for uent readers of alphabetic orthographiesas well as for readers of logographic orthographies. However, he

188 I. DeWitt, J.P. Rauschecker / Brairegarded whole-word reading as sufciently minor in contribution(relative to phonological reading) to be negligible and, therefore,dismissed it. Crucially, Wernicke viewed dependency of writtenhowever, are unaware of any empirical work that has specicallyinvestigated the likelihood of reading decits given auditory com-prehension decits, word repetition decits and paraphasia. Ascase reports show dissociability (Ellis, Miller, & Sin, 1983), readingdecits observed in individuals with Wernickes aphasia may re-ect the typically large lesion volume of middle cerebral arteryaccidents associated with Wernickes aphasia, which frequently in-volve anterior ST, posterior ST and IPL (Robson et al., 2012). That is,patients presenting with both auditory and reading comprehen-sion decits may have large lesions, disrupting multiple corticalmodules. Thus, it remains unclear whether lesions disrupting audi-tory word-form recognition or inner speech necessarily also dis-rupt reading comprehension.

In summary, the aspects of reading comprehension Wernickeassociated with his sensory speech center relate to phonologicalreading via inner speech. Both phonological reading and innerspeech are functions neuroanatomically associated with the audi-tory dorsal stream. Though decits in auditory and written com-prehension are often observed together, the dissociability of theirneural substrates (or aspects of them) and their precise neuroanat-omy requires further investigation.

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Wernickes area revisited: Parallel streams and word processing1 Introduction2 Word-form recognition and the auditory ventral stream3 Causal involvement of anterior STG in word recognition4 A brief history of Wernickes area5 Paraphasia, inner speech and the auditory dorsal stream6 ConclusionsAcknowledgmentsAppendix A Further historical notes on pure word deafnessAppendix B Written comprehensionReferences