-
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:
[email protected] (I. DeWitt), [email protected]
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
nguage
l sevier .com/locate /b&l
-
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