The Neural Correlates of Grammatical Gender: An fMRI Investigation

The Neural Correlates of Grammatical GenderAn fMRI Investigation

Gabriele Miceli1 Patrizia Turriziani2 Carlo Caltagirone25Rita Capasso2 Francesco Tomaiuolo2 and Alfonso Caramazza34

Abstract

amp In an fMRI experiment subjects saw a written noun andmade three distinct decisions in separate sessions Is itsgrammatical gender masculine or feminine (grammaticalfeature task) Is it an animal or an artifact (semantic task)Does it contain a tch or a k sound (phonological task)Relative to the other experimental conditions the grammaticalfeature task activated areas of the left middle and inferiorfrontal gyrus and of the left middle and inferior temporal gyrusThese activations fit in well with neuropsychological studiesthat document the correlation between left frontal lesions and

damage to morphological processes in agrammatism and thecorrelation between left temporal lesions and failure to accesslexical representations in anomia Taken together these datasuggest that grammatical gender is processed in a left fronto-temporal network In addition the observation that thegrammatical feature task and the phonology task activatedneighboring but distinct regions of the left frontal lobeprovides a plausible neuroanatomical basis for the systematicoccurrence of phonological errors in aphasic subjects withmorphological deficits amp

INTRODUCTION

To know a word is to know three types of things itsmeaning its form and its grammatical properties Howare these three types of information represented in thebrain More specifically what relationship is there be-tween a wordrsquos grammatical properties and its form andmeaning Here we present evidence that the grammat-ical properties of a word activate a neural circuit that ispartially independent of the networks activated by thatwordrsquos form and meaning

Neuropsychological studies have shown that meaningand form are functionally dissociable and that they arerepresented in distinct brain structures Evidence to thiseffect comes from complementary patterns of perform-ance in brain-damaged subjects demonstrating the dou-ble dissociation of meaning and form Thus there havebeen reports of selective damage to meaning in thepresence of spared lexical form in subjects with focalbrain damage (eg Hillis Rapp Romani amp Caramazza1990) and in subjects with lsquolsquosemantic dementiarsquorsquo whosuffer from a degenerative disease of the temporal lobes(eg Patterson amp Hodges 1992) The reverse dissocia-tion has been documented in cases of selective damagein accessing lexical form in the face of spared meaning

as a consequence of focal lesions of the left hemisphere(eg Caramazza amp Hillis 1990 Kay amp Ellis 1987) Thedissociation of meaning and form is also supported byneuroimaging investigations PET and fMRI studies havefound that semantic processing involves activation of aneural network distributed over several lobes of the lefthemisphere or both hemispheres (eg VandenberghePrice Wise Josephs amp Frackowiak 1996) Other reportshave found that phonological processing activates amore restricted network including the posterior por-tions of the left superior and middle temporal gyri andthe left inferior frontal gyrus (eg Xu et al 2001Hagoort et al 1999 Fiez 1997)

Neuropsychological observations have also providedevidence on the functional and neural organization ofthe grammatical properties of words For example thereare many reports of aphasic patients with selectivedifficulties in processing either verbs or nouns (egBerndt Haendiges Burton amp Mitchum 2002 TranelAdolphs Damasio amp Damasio 2001 Bak OrsquoDonovanXuereb Boniface amp Hodges 2001 Cappa et al 1998Hillis amp Caramazza 1995 Daniele Giustolisi SilveriColosimo amp Gainotti 1994 Caramazza amp Hillis 1991Zingeser amp Berndt 1988 Miceli Silveri Villa amp Caramaz-za 1984) These reports link the processing of nouns andverbs to distinct neural substratesmdashpoor performancewith verbs is frequently associated with damage to theleft middle frontal gyrus poor performance with nounsis most frequently linked to lesions of the temporal (andparietal) structures of the left hemisphere The existence

1 Universita Cattolica Rome Italy and Istituto di PsicologiaCNR Rome Italy 2Fondazione Santa Lucia Rome Italy3 Harvard University 4SISSA Trieste Italy 5Dipartimento diNeuroscienze Universita di Tor Vergata Rome Italy

copy 2002 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 144 pp 618ndash628

of separate neural networks in the processing of nounsand verbs is also supported by neuroimaging studies(eg Perani et al 1999 Warburton et al 1996 MartinHaxby Lalonde Wiggs amp Ungerleider 1995 PetersenFox Posner Mintun amp Raichle 1988) and by electro-physiological investigations (Federmeier Segal Lombro-zo amp Kutas 2000 Dehaene 1995) However theinterpretation of the nounverb dissociation as a specif-ically grammatical effect remains controversial This isbecause grammatical categories correlate with semanticproperties (eg nouns typically denote objects whereasverbs typically denote actions nouns are usually moreconcrete verbs usually more abstract etc) It is notimplausible therefore to suppose that the observeddissociations reflect the organization of the semanticsystem rather than grammatical properties of words(eg Bird Howard amp Franklin 2000 Breedin Saffranamp Schwartz 1998 but see Berndt et al 2002 Caramazzaamp Shapiro in press for arguments and evidence thatchallenge the strong version of this hypothesis)

One way to study the neural representation of thegrammatical properties of words is to focus on a lexicalfeature that is independent from meaning The gram-matical feature gender meets this requirement in manylanguages (Corbett 1991) This property of nouns playsan important role in determining agreement amongwords in noun phrases in the selection of pronominalforms and even in subjectndashverb agreement (eg specify-ing agreement between the subject of a sentence andthe past-participle form of the verb in Italian) Forexample in Italian all the words in a noun phrasemdashdeterminers adject ives and nounsmdashmust agreein number and gender1Consider the case of the phrases(masc = masculine fem = feminine sg = singularpl = plural) lsquolsquoilmasc sg piccolomasc sg pontemasc sgrsquorsquo(the small bridge) lsquolsquoimasc pl piccolimasc pl pontimasc plrsquorsquo(the small bridges) lsquolsquolafem sg piccolafem sg navefem sgrsquorsquo(the small ship) and lsquolsquolefempl piccolefempl navifemplrsquorsquo (thesmall ships) As may be seen the definite determinertakes different forms depending on the gender and thenumber of the nounmdashil and i for masculine singular andplural nouns respectively and la and le for femininesingular and plural nouns respectively Similarly theadjective lsquolsquosmallrsquorsquo is inflected appropriately for numberand gender -o or -i for masculine singular and pluralrespectively and -a and -e for feminine singular andplural respectively As another example consider thecase of clitics In Italian the sentence lsquolsquogive it to mersquorsquo isproduced with cliticized forms in which both the dativepronoun (lsquolsquoto mersquorsquo) and the direct object pronoun lsquolsquoitrsquorsquoare cliticized onto the verb However the specific formof the clitic for the direct object depends on the gram-matical gender and number of the noun for which thepronoun stands If the noun is singular masculine theclitic would be -lo as in dammelo (literally give-me-it)and it would be respectively -li -la and -le for mascu-line plural feminine singular and feminine plural nouns

As these examples illustrate the retrieval of a nounrsquosgrammatical gender is an essential part of the sentenceproduction process

Crucially for our purposes a nounrsquos semantic contentand grammatical gender have only an arbitrary relation-ship This can be readily appreciated by considering thefact that closely related semantic coordinates havedifferent grammatical genders For example the Italiantranslation of the first member of each of the followingpairs of words is masculine while the second is femi-nine lemonorange grapefruitapple tablechair sheetblanket platecup glassbottle spoonfork traincarelephantgiraffe liontiger sunmoon and so on Fur-thermore the grammatical gender of a word is languagespecific While the Italian and French translations ofpencil desk carriage summer and flag are feminineand masculine respectively the opposite is true for seatable limit river and tomato If grammatical genderswere determined by a nounrsquos semantic properties wewould expect semantically related words to have thesame gender and that words would have the samegender across languages Neither condition holds Thusthe results of investigations of the neural mechanismsinvolved in processing grammatical gender can be in-terpreted to reflect grammatical as opposed to semanticaspects of language processing

Although grammatical gender and meaning are re-lated only arbitrarily the same is not true for the relationbetween gender and form In many languages includingItalian Russian and Hebrew for example words of thesame grammatical gender tend to share specific phono-logical properties In the case of Italian a majority ofmasculine singular nouns end in the vowel o (plural i)and a majority of feminine singular nouns end in thevowel a (plural e) However the correlation is farfrom perfect Thus among the 3000 most frequentwords in Italian only 66 of nouns have a lsquolsquoregularendingrsquorsquo (De Mauro Mancini Vedovelli amp Voghera1993) The nonregular nouns consist either of wordsending in e (and a few in i) which are associatedequally frequently with masculine and feminine nounsor masculine nouns that end in a and feminine nounsthat end in o Thus although it has been shown thatsubjects are sensitive to the correlation between genderand word ending in word recognition tasks (eg BatesDevescovi Pizzamiglio DrsquoAmico amp Hernandez 1995) adefinitive decision about a wordrsquos grammatical gendercan only be taken after lexical access has occurred

The independence of grammatical gender from mean-ing and the fact that lexical access is necessary in orderto determine the gender of a noun has made thisgrammatical feature an appropriate tool for investigatingthe process of lexical access in normal language produc-tion (see Caramazza Miozzo Costa Schiller amp Alario inpress Schriefers 1993) These same properties of gram-matical gender make it suitable for the study of therepresentation of grammatical word properties in the

Miceli et al 619

brain As already noted the crucial aspect of this gram-matical feature is that it is independent of meaning andtherefore any effect of grammatical gender cannot beascribed to the organization of conceptual knowledge inthe brain

It is possible to formulate plausible expectationsabout the areas of the left hemisphere that are likelyto be involved in processing grammatical gender Sincegrammatical gender is a property of nouns and sincethere is considerable evidence that damage to the lefttemporal lobe results in the impairment of lexical accessof nouns we might expect the temporal lobe to beimplicated in processing grammatical gender Anotherarea that is likely to be involved in processing grammat-ical gender is the left inferior frontal lobe It is wellestablished that lesions in this area result in difficulties inmorphological processing in sentence production(agrammatism Goodglass 1976 Tissot Mounin amp Lher-mitte 1973) Since morphological processing is con-trolled by information about grammatical category andthe various grammatical features associated with nounsand verbs we would expect the left frontal cortex to playa crucial role in processing this type of information Inshort the expectation is that the processing of gram-matical features of nouns will implicate a left fronto-temporal network (Shapiro amp Caramazza submitted)

In the neuroimaging investigation reported here weasked normal subjects to process isolated nouns incontexts that accentuated distinct properties of eachnounmdashmeaning form or grammatical gender The pat-terns of activation observed in the three experimentaltasks were compared to those observed in a baselinetask during which subjects were asked to silently readpseudowords while alternately pressing response but-tons The pseudowords were matched to words inlength and consonantndashvowel structure Performanceon each experimental condition was also compared toperformance on the other two experimental conditionsWe were interested in seeing how the processing of aspecific lexical property influenced the activation ofdifferent neural structures during fMRI scanning andwe specifically focused on grammatical gender Based onavailable neuropsychological and neuroimaging obser-vations we expected relatively distributed (possibly bi-lateral) activation in response to semantic features andleft frontal activation in response to segmental phono-logical properties of lexical representations More im-portantly we expected activation of a fronto-temporalnetwork in processing grammatical gender

RESULTS

Behavioral Results

The experimental stimuli were administered to eightvolunteers in the context of a reaction time task Foreach condition (semantic syntactic and phonological)

subjects were asked to press a button as soon as theyhad made their decision Reaction time and responseaccuracy were comparable across conditions (Table 1)The average reaction time was 669 plusmn 646 msec for thegrammatical gender condition 7098 plusmn 624 msec forthe phonological condition and 6826 plusmn 381 msec forthe semantic condition F(214) = 189 p = 188 Onaverage control subjects produced 362 plusmn 2 correctresponses for the semantic condition 366 plusmn 28 forthe phonological condition and 37 plusmn 24 for the gram-matical condition F(214) = 59 p = 569

Activation Results

Experimental Conditions Versus Baseline

When compared to silent reading of pseudowordswhile alternately pressing buttons placed in the leftand in the right hand all experimental conditionsyielded significant activations ( p lt 001) at the highestthreshold level (voxel correction lt01 cluster correc-tion lt001) (Table 2) In the grammatical gendercondition there was significant activation in the leftfrontal lobe involving BA 9 and BA 45 The left BA 7was also activated The phonological task resulted insignificant activation of the left inferior frontal gyrusinvolving the frontal operculum (BA 44) and extendinginferiorly to BA 47 and posteriorly to the precentralgyrus (BA 6) There was also bilateral activation of thesuperior parietal lobule (BA 7) extending into theuppermost portion of BA 40 in the left hemisphereThe semantic task activated bilaterally the frontal andparietal structures and the cerebellum Frontal activa-tion involved the middle and inferior frontal gyri on theleft (BA 9 BA 946 and BA 47) and the middle frontalgyrus on the right (BA 4546 BA 9 and BA 68) sym-metrical activation was observed in the superior parietallobule (BA 7)

Thus all experimental conditions resulted in signifi-cant activation of the frontal lobe Activation was bilat-eral in the semantic condition and was restricted to theleft hemisphere in both the gender and the phonologyconditions Gender processing activated the more infe-rior portions of the inferior frontal gyrus to a lesserextent than did the phonological processing task

Significant activation of the superior parietal lobule(BA 7) was observed in all experimental conditions when

Table 1 Behavioral Results Average Reaction Times (msec)and Response Accuracy (Correct Responses) and StandardDeviations in the Three Experimental Conditions

Reaction Time(Average ndash SD)

Response Accuracy(Average ndash SD)

Semantic condition 6826 plusmn 381 362 plusmn 20

Syntactic condition 6690 plusmn 646 370 plusmn 25

Phonological condition 7098 plusmn 624 366 plusmn 28

620 Journal of Cognitive Neuroscience Volume 14 Number 4

compared to the baseline task This pattern of activationmost likely results from the different demands of theexperimental and baseline conditions in terms of bothattention and memory The control task required thesilent reading of pseudowords and alternately pressingresponse buttons with no need to maintain the stimuliactive in working memory The experimental tasks in-volved a binary decision to items presented in random

sequence requiring the subject to attend carefully toeach stimulus and to keep active a representation of thestimulus until a decision could be made This interpre-tation is supported by neuroimaging studies showingincreased activation in the inferior and superior parietallobules in tasks involving selective and divided attentionand working memory (for a review see Cabeza ampNyberg 2000) Since this activation appears to be

Table 2 Stereotactic Coordinates (Talairach amp Tournoux 1988) of the Foci of Activation for the Comparisons Between EachExperimental Condition and the Baseline Task ( p lt 001)

x y z Z

Gender gt pseudowords (voxel correction lt01 cluster correction lt001)

L inferior frontal gyrus BA 9 iexcl40 16 26 454



L superior parietal lobule BA 7 iexcl30 iexcl56 48 471


L inferior parietal lobule BA 7 iexcl38 iexcl42 36 358

Phonology gt pseudowords (voxel correction lt01 cluster correction lt001)


L inferior frontal gyrus BA 47 iexcl34 26 iexcl8 360

L prefrontal gyrus BA 6 iexcl34 2 28 424



L supramarginal gyrus BA 40 iexcl44 iexcl40 40 369

R superior parietal lobule BA 7 24 iexcl50 44 400


R inferior parietal lobule BA 7 32 iexcl60 46 354

Semantics gt pseudowords (voxel correction lt01 cluster correction lt001)

L middle frontal gyrus BA 9 iexcl44 iexcl32 66 477

L middle frontal gyrus BA 946 iexcl36 26 26 438


L superiorinferior parietal lobule BA 7 iexcl22 iexcl62 36 392



R middle frontal gyrus BA 4546 38 32 12 471




R precuneus BA 7 12 iexcl70 50 334

L cerebellum 6 iexcl66 iexcl18 460

R cerebellum iexcl6 iexcl54 iexcl8 441

Miceli et al 621

unrelated to the linguistic dimensions under scrutiny itwill not be discussed further

Comparisons Between Experimental Conditions

The activation observed in each experimental conditionwas also compared with the activation observed in each

of the other two experimental conditions (Table 3 andFigure 1)

The gender-versus-semantics comparison yielded sig-nificant values at low thresholds (voxel correction lt05cluster correction lt05) Significant left-sided activationwas observed in the inferior frontal gyrus (BA 45) and in

Table 3 Stereotactic Coordinates (Talairach amp Tournoux 1988) of the Foci of Activation for the Comparison Between Pairs ofExperimental Tasks ( p lt 001)

x y z Z

Gender gt phonology (voxel correction lt05 cluster correction lt001)

L inferior temporal gyrus BA 21 iexcl42 iexcl10 iexcl22 468

L middle temporal gyrus BA 2021 iexcl52 iexcl32 iexcl12 397



L cingulate gyrus BA 31 iexcl6 iexcl32 42 358

Phonology gt gender (voxel correction lt05 cluster correction lt05)


Gender gt semantics (voxel correction lt05 cluster correction lt001)


Lenticular nucleus iexcl14 6 iexcl8 430

Semantics gt gender (voxel correction lt05 cluster correction lt001)

R Heschlrsquos gyrus BA 41 42 iexcl20 4 397

R cingulate gyrus BA 31 16 iexcl24 40 360

R cerebellum 20 iexcl66 iexcl22 385

Semantics gt phonology (voxel correction lt01 cluster correction lt001)

R inferior frontal gyrus BA 8 28 10 30 386

L middlesuperior frontal gyrus BA 9 iexcl18 34 26 366



L fusiform gyrus BA 20 iexcl40 iexcl12 iexcl30 370

L cingular gyrus BA 32 12 14 30 368

L cerebellum iexcl10 iexcl54 iexcl8 466


R thalamus 10 iexcl40 10 403


Phonology gt semantics (voxel correction lt01 cluster correction lt001)



L precentral gyrus BA 6 iexcl50 2 42 319


the lenticular nucleus The reverse comparison (seman-tics vs gender) yielded significant activation at an inter-mediate threshold (voxel correction = 05 clustercorrection = 001) in the right hemisphere structuresthat are outside of the typical language areas (Heschlrsquosgyrus cingulate gyrus and the cerebellum)2

The gender-versus-phonology comparison yielded sig-nificant activations at an intermediate threshold (voxelcorrection lt05 cluster correction lt001) in the leftinferior and middle temporal gyrus (BA 2021 and BA 21)Patchy activation was observed also in a large region thatincludes the left supramarginal gyrus (BA 40) and the leftposterior cingulate gyrus (BA 31) At a lower threshold(voxel correction lt05 cluster correction lt05) thephonology-versus-gender comparison yielded significantactivation in the left inferior frontal gyrus (BA 44)

Comparisons between semantics and phonologyyielded significant differences at high threshold (voxelcorrection lt01 cluster correction lt001) The seman-tics-versus-phonology comparison yielded significant left

hemisphere activation in the inferior temporal gyrus andin the fusiform gyrus (BA 20) small areas of activationwere also observed in almost symmetrical structures ofthe left (BA 9) and right (BA 8) frontal lobe and in theleft anterior paracingulate cortex (BA 32) Additionalsmall activations of unclear significance involved theright thalamus and the cerebellum The phonology-versus-semantics comparison yielded significant activa-tion of the left middle (BA 8) and inferior (BA 446)frontal gyrus extending posteriorly to the precentralgyrus (BA 6)

DISCUSSION

We had predicted that processing of grammatical genderwould activate structures in the left frontal and temporallobes Both predictions were borne out Decidingwhether a noun is masculine or feminine resulted inthe significant activation of (1) the left inferior frontalgyrus (BA 45) relative to the baseline condition and

Figure 1 Lateral see-throughof the activations observedwhen comparing pairs ofexperimental conditions

Miceli et al 623

to the semantic task (2) the left middle frontal gyrus(BA 9) relative to the baseline and (3) the left middleand inferior temporal gyrus (BA 2021 and BA 21)relative to the phonological condition In addition tothe predicted areas the gender decision task also re-sulted in the significant activation of the left parietal lobe(BA 40) and of the left posterior cingulate gyrus (BA 31)when compared to the phonological task

Our reason for expecting left frontal activation isthat grammatical gender is involved in the control ofmorphological processes which are known to beimpaired in agrammatism and that lesion analysis inagrammatic speakers has frequently demonstrateddamage involving (but often not limited to) the leftfrontal lobe (eg Vanier amp Caplan 1991 Miceli SilveriRomani amp Caramazza 1989) Another reason to expectleft frontal lobe activation is that recent neuroimagingstudies have found activation of these structures dur-ing tasks requiring the ability to process sentences thatviolate morphological constraints (Moro et al 2001Ni et al 2000)

Activation of the left inferior (BA 45) and middle(BA 9) frontal gyrus during the gender decision task isintriguing also in the light of recent studies that impli-cate these regions in processing grammatical categoryinformation Results consistent with this possibility havebeen reported in studies of aphasics (Bak et al 2001Shapiro Shelton amp Caramazza 2000 Cappa et al 1998Daniele et al 1994) in neurophysiological investiga-tions (Shapiro Pascual-Leone Mottaghy Gangitano ampCaramazza 2002 Federmeier et al 2000 Dehaene1995) and in neuroimaging studies (Friederici Oppitzamp von Cramon 2000)

The gender decision task resulted also in the signifi-cant activation of portions of the left middle and inferiortemporal gyrus (BA 2021 and BA 21) when comparedto the phonological condition Our reason for expectingactivation in the left temporal lobe was based in part onreports of aphasic subjects with lsquolsquopure anomiarsquorsquo Thesepatients typically present with selective difficulty inaccessing lexical information about nouns despite pre-served ability to process the meaning of nouns Thelesion site most often associated with this pattern ofperformance is the left temporal lobe (Lambon RalphSage amp Roberts 2000 Miceli Giustolisi amp Caramazza1991 Kay amp Ellis 1987 Gainotti Silveri Villa amp Miceli1986) suggesting that the left temporal lobe is a crucialcomponent in accessing nouns Since grammatical gen-der is a property of nouns its retrieval presupposesaccess to the lexical node of the to-be-produced wordThe findings of left temporal lobe activation in normalsubjects engaged in performing a gender decision taskand of left temporal damage in pure anomia provideevidence for the role of these left hemispheric structuresin lexical access of nouns

The remaining two foci of activation in the leftparietal lobe (BA 40) and in the left cingulate gyrus

(BA 31) cannot be related straightforwardly to theprocessing of grammatical gender either as a propertyof lexical representations or as a feature involved inmorphological processes This is because deficits oflexical access or morphological processing are notassociated with damage (exclusively) to these regionsA more likely basis for the observed foci of activationis the metalinguistic nature of the gender decisiontask When debriefed after scanning most participantsreported solving the gender task by silently generatingthe determiner of the noun For example whendeciding the gender of say lsquolsquofalcefemrsquorsquo (sickle) theywould silently say to themselves lsquolsquolafem falcefemrsquorsquo Thisstrategy may have resulted in the involvement ofworking memory which has often been associatedwith left or bilateral activation of the supramarginalgyrus in PET and fMRI studies (eg Paulesu Frith ampFrackowiack 1993 for an exhaustive review see Ca-beza amp Nyberg 2000) The activation of BA 31 hasbeen documented in other metalinguistic languagetasks such as fluency tasks (eg Frith Friston Liddleamp Frackowiack 1991) and is also probably aspecificConsistent with these possibilities the activation of theleft BA 40 and BA 31 was observed in the comparisonbetween the grammatical gender task and the phono-logical task In the gender task the decision must bemade on a feature (grammatical gender) that cannotbe predicted by the written word whereas the deci-sion required by the phonological task can be reachedon the basis of the segmental information provided inthe written stimulus

In short the results of our neuroimaging experimentconverge with evidence from neuropsychological neu-rophysiological and neuroimaging studies in suggestingthat the grammatical features of nouns are representedin a network that includes frontal (BA 45 and BA 9) andtemporal (BA 2021 and BA 21) structures of the lefthemisphere The same results in association with neu-ropsychological studies of agrammatism and with neuro-imaging studies of sentence processing in cognitivelyunimpaired subjects confirm the critical role of the leftfrontal lobe in morphological processes (and in repre-senting grammatical category information)

The close proximity of the areas activated by thephonological task and by the gender task is also worthstressing BA 44 was activated by the phonological taskmore than the baseline condition the gender task andthe semantic task and BA 45 was activated more by thegender task than the baseline and the semantic tasksThis pattern of activation is intriguing in the light of theclinical observation that subjects who make morpholog-ical errors in single-word processing tasks almost invar-iably also make phonological errors in the same tasks(eg Miceli amp Caramazza 1988) The reasons for thisseemingly systematic co-occurrence are unclear Theresults reported here suggest that it might be due tothe proximity (or even the partial overlap) of the neural


substrates involved in processing the morphophonolog-ical (eg the computation of the gender and numberagreement inflections in adjectives) and phonologicalproperties of words In particular our results are con-sistent with the hypothesis that the left inferior frontalgyrus participates in distinct but at least partially inter-twined neural networks involved in phonological andmorphophonological processing On this view the ad-jacent but separate foci of activation observed in thepresent study would indicate a higher density of neuraltissue representing phonological processing in BA 44and higher density of neural tissue representing mor-phophonological processing in BA 45 The fact thatcontiguous brain regions are involved in processingthe morphology and the phonological properties of aword does not necessarily lead to the conclusion thatthese two types of information are functionally insepa-rable It is possible that morphophonological processes(ie those processes involved in specifying the phono-logical form of polymorphemic words) are inextricablyintertwined with lexical phonological informationwhereas morphosyntactic processes (ie those pro-cesses that manipulate inflectional morphology at thesentence level) may be clearly separable from phono-logical processes

In conclusion grammatical gender (and probablyother properties of the lexical representations of nouns)appears to be represented in the left hemisphere in aneural network that includes the inferior and middlefrontal gyri and the inferior and middle temporal gyri Inaddition the results obtained for gender converge withother neuropsychological and neuroimaging investiga-tions in showing that the left inferior and middle frontalgyri play a critical role in the representation of gram-matical category information and in the processing ofmorphological information The contiguity of the struc-tures activated by the phonological and the grammaticalgender condition indicates both functional overlapand functional specialization within the left inferiorfrontal gyrus

METHODS

Subjects

Nine right-handed (based on the Edinburgh Handed-ness Inventory Oldfield 1971) participants 6 men and3 women aged 23ndash29 native speakers of Italian gaveinformed consent to participate in the study All subjectshad normal or corrected-to-normal vision Exclusioncriteria included a history of prolonged use of prescrip-tion or recreational drugs a neurological or psychiatriccondition claustrophobia and significant prior radiationexposure Subjects were advised not to use alcoholwithin 24 hr and tobacco within 3 hr prior to the scanThe Ethics Committee of Fondazione Santa Lucia ap-proved the experimental protocol Subjects were paidfor their participation in the study

Materials

The experimental stimuli consisted of 40 concrete nounsand of 40 pseudowords Word stimuli were counter-balanced for grammatical gender semantic categoryand phonological content There were 20 masculinenouns (15 ending in -o and 5 ending in -e) and 20feminine nouns (15 ending in -a and 5 ending in -e) 20animal names and 20 artifact names and 20 nounscontaining a tch sound (eg uncino unrsquotchinohook) and 20 nouns containing a k sound (eg caprarsquokapra goat) The position of the target sound wasbalanced across stimuli Regardless of the parameterover which nouns were counterbalanced (meaninggrammatical gender phonology) the frequency of usagewas comparable in the two 20-item subsets The 40nouns selected as experimental stimuli were used toconstruct six 20-word lists Each list contained equalnumbers of masculine and feminine nouns of animaland artifact names and of words containing the tch orthe k phoneme (n = 10 each) Each word was includedin three sublists In one sublist participants were askedto decide whether the word was of masculine or femi-nine gender in another sublist they were asked todecide whether the word was the name of an animalor of an artifact and in the other sublist they were askedto decide whether the word contained a tch or a ksound In this last condition the phonological contrastcorresponds to a difference between the orthographiccontexts for the letter hci in the written string In Italianthe letter hci is pronounced as k when followed by thevowel hoi hai or hui or by the sequence hhii or hhei andit is pronounced tch when followed by the vowel hii orhei The fact that the phonological task could be per-formed accurately simply by inspecting the writtenstimulus may have allowed subjects to base their re-sponses in this condition on the segmental features ofthe stimulus and not on the retrieval of the correspond-ing lexical form as was necessary in the grammaticalgender and in the semantic conditions The putativeeffects of this difference across tasks are considered inthe Discussion

Pseudowords consisted of pronounceable letterstrings They were identical to the 40 experimentalwords in number of letters and syllables and in conso-nantvowel structure For each pseudoword the n countcorresponded to 0 (in other words changing one letterat a time did not result in a word) Pseudowords weredivided in two lists that were alternately repeated duringthe experiment

Procedure for the Behavioral Task

This task was administered to eight subjects (5 men3 women) comparable in age to those who participatedin the fMRI experiment A practice session was run toacquaint subjects with the experimental tasks Each

Miceli et al 625

subject was presented with the six sublists of words Foreach sublist subjects were given explicit informationabout the dimension they were going to be asked toevaluate (grammatical gender semantic category pho-nological content) and were instructed to press one oftwo computer keys as soon as they decided whetherthe stimulus was of masculine or feminine gender ananimal or an artifact or whether it contained a tch or ak sound A fixation point appeared at the center of thecomputer monitor for 400 msec followed by the stim-ulus which remained on the screen for 1200 msec

Procedure for the fMRI Task

The study consisted of three experimental tasks and acontrol task All tasks involved the presentation of awritten word at the center of the subjectrsquos field ofvision with the same timing as in the behavioral taskThe gender task required subjects to indicate whetherthe stimulus was masculine or feminine The semantictask required participants to judge whether the stimulusword corresponded to an animal or an artifact Thephonological task required participants to decidewhether the stimulus contained when pronounced atch or k sound Subjects responded by pressingbuttons held in the right and left hand The controltask consisted of alternately pressing buttons whilesilently reading pseudowords This task was clearly lessdemanding than the experimental tasks in terms ofprocessing (attention and memory) resources How-ever the stimuli were closely matched in visual com-plexity to the experimental noun lists and the taskprovides a good control for the motor component ofthe response in the experimental tasks Prior to scan-ning each participant was acquainted with the exper-imental tasks during a practice session during whichdifferent sets of words from those included in theexperimental conditions were presented

The four experimental tasks were organized in a blockdesign administered in one scanning session lastingapproximately 15 min Each experimental conditionwas presented in alternation with the control task(ct)mdashAndashctndashBndashctndashCndashct During the scanning sessionthis block was repeated four times The order of taskpresentation (AndashctndashBndashctndashCndashct) was fixed for eachsubject but counterbalanced across subjects (egCndashctndashBndashctndashAndashct) Each experimental block beganwith a slide showing instructions about the dimensionof the judgment (gender semantic category phonology)to be performed in that block of trials and which key hadto be pressed for each judgment (eg masculine gen-der left key feminine gender right key)

The stimuli were projected via mirroring to a backprojection screen using an LCD video projector (ModelVPL-351QM Sony Tokyo) located inside the MRI roomand connected to a Macintosh computer (G3266 AppleComputer Cupertino CA) located outside the MRI

room Images were obtained by using MATLAB 53(MathWorks Natick MA) and the MATLAB stats toolboxrunning on UNIX workstations (O2 Silicon GraphicsMountain View CA)

fMRI Procedures and Data Analysis

A Siemens Vision Magnetom MR system (Siemens Med-ical Systems Erlangen Germany) operating at 15 T andequipped for echo-planar imaging was employed inorder to acquire functional MR images Head movementwas minimized by mild restraint and cushioning Usingan EPI gradient-echo sequence (TR = 3200 msec TE =40 msec) 240 functional images were collected for eachdata acquisition run Thirty-four adjacent oblique-axialslices starting from the superior convexity so as to coverthe whole brain were acquired in an interleaved se-quence with 3-mm slice thickness an FOV = 192 pound 192mm and a 64 pound 64 matrix size resulting in in-planeresolution of 3 pound 3 pound 3 mm

Functional data were analyzed using the SPM99 (Well-come Department of Cognitive Neurology LondonEngland) Prior to any statistical analysis all volumesfor each subject were realigned using the first volume asreference and resliced using sync interpolation In orderto perform intersubject averaging all images were trans-formed into a standard space (Talairach amp Tournoux1988) matching to a template image As a final prepro-cessing step functional data were smoothed using a6-mm FWHM isotropic Gaussian kernel to compensatefor residual variability after spatial normalization and topermit application of Gaussian random field theory toprovide for correct statistical inference (Friston et al1995) Tasks were modeled as boxcar functions andconvolved with a synthetic hemodynamic responsefunction Head motion parameters were added as re-gressors (Friston et al 1995 Worsley amp Friston 1995)

The contrast between each activation condition versusrest was examined first Each experimental task was alsodirectly compared with each of the other two Random-effects analyses were performed on the group data Fociof activation were characterized in terms of spatialextent (k) and peak height (u) The correct significanceof each region was estimated in terms of the probabilitythat a region of the observed number of voxels couldhave occurred by chance and that the observed peakheight could have occurred by chance over the entirevolume analyzed

Different thresholds were retained for the variousanalyses The t statistic images for each contrast werecreated first by retaining regions surviving a threshold ofp lt 01 at the voxel level with a cluster-level correctioncorresponding to p lt 001 In the absence of significantactivations images were created by retaining regionsthat survived a higher threshold ( p lt 05 at voxel level)In this case the significance of each region was esti-mated in terms of spatial extent (cluster-level) applying a


threshold of p lt 001 or p lt 05 This strategy waspreferred to the selection of a uniformly low significancelevel so that for each contrast we could evaluate theregions showing the greatest activation

Acknowledgments

This project was supported in part by grants from MURST andfrom Fondazione Santa Lucia to G M and by NIH Grant NoDC 04542 to A C

Reprint requests should be sent to either Gabriele MiceliNeurologiamdashUniversita Cattolica Largo A Gemelli 8 00168Rome Italy (e-mail gmicelimclinkit) or Alfonso CaramazzaCognitive Neuropsychology Laboratory Harvard University 33Kirkland Street Cambridge MA 02138 USA or via e-mailcaram wjhharvardedu

The data reported in this experiment have been deposited inThe fMRI Data Center (httpwwwfmridcorg) The accessionnumber is 2-2002-112NH

Notes

1 In Italian most adjectives occur postnominally lsquolsquoil leoneferocersquorsquo literally means lsquolsquothe lion ferociousrsquorsquo For expositorypurposes we have used an adjective that can be usedprenominally as in English2 Failure to demonstrate significantly different activation ofthe language areas of the left hemisphere in this comparisonmay reflect the fact that since access to grammatical genderpresupposes lexical access which in turn requires the retrievalof meaning the gender and the meaning condition activatedoverlapping structures Greater activation of right hemispherestructures in the semantic condition may result from the factthat grammatical gender information is lateralized to the lefthemisphere whereas semantic information is representedmore bilaterally

REFERENCESBak T OrsquoDonovan D Xuereb J Boniface S amp Hodges J

(2001) Selective impairment of verb processing associatedwith pathological changes in Brodmann areas 44 and 45 inthe motor neuron diseasendashdementiandashaphasia syndromeBrain 124 103ndash120

Bates E Devescovi A Pizzamiglio L DrsquoAmico S ampHernandez A (1995) Gender and lexical access in ItalianPerception and Psychophysics 57 847ndash862

Berndt R S Haendiges A H Burton M W amp MitchumC C (2002) Grammatical class and imageability in aphasicword production Their effects are independent Journal ofNeurolinguistics 13 353ndash371

Bird H Howard D amp Franklin S (2000) Why is a verb likean inanimate object Grammatical category and semanticcategory deficits Brain and Language 72 246ndash309

Breedin S Saffran E amp Schwartz M (1998) Semantic factorsin verb retrieval An effect of complexity Brain andLanguage 63 1ndash31

Cabeza R amp Nyberg L (2000) Imaging cognition II Anempirical review of 275 PET and fMRI studies Journal ofCognitive Neuroscience 12 1ndash47

Cappa S Binetti G Pezzini A Padovani A Rozzini Lamp Trabucchi M (1998) Object and action naming inAlzheimerrsquos disease and frontotemporal dementiaNeurology 50 351ndash355

Caramazza A amp Hillis A E (1990) Where do semantic errorscome from Cortex 26 95ndash122

Caramazza A amp Hillis A (1991) Lexical organization of nounsand verbs in the brain Nature 349 788ndash790

Caramazza A Miozzo M Costa A Schiller N amp Alario F-X(in press) Lexical selection A cross-language investigation ofdeterminer selection In E Dupoux (Ed) Language brainand cognitive development Essays in honor of JacquesMehler Cambridge MIT Press

Caramazza A amp Shapiro K (in press) Categories in the brainIn L Rizzi amp A Belletti (Eds) Structures and beyondOxford Oxford University Press

Corbett G (1991) Gender Cambridge Cambridge UniversityPress

Daniele A Giustolisi L Silveri M Colosimo C amp GainottiG (1994) Evidence for a possible neuroanatomical basis forlexical processing of nouns and verbs Neuropsychologia32 1325ndash1341

Dehaene S (1995) Electrophysiological evidence for category-specific word processing in the normal human brainNeuroReport 6 2153ndash2157

De Mauro T Mancini F Vedovelli M amp Voghera M(1993) Lessico di frequenza dellrsquoItaliano parlatoMilano Etas

Federmeier K Segal B Lombrozo T amp Kutas K (2000)Brain responses to nouns verbs and classmdashambiguouswords in context Brain 123 2552ndash2466

Fiez J A (1997) Phonology semantics and the role of the leftinferior frontal cortex Human Brain Mapping 5 79ndash83

Friederici A D Oppitz B amp von Cramon D Y (2000)Segregating semantic and syntactic aspects of processing inthe human brain An fMRI investigation of different wordtypes Cerebral Cortex 10 698ndash705

Friston K Holmes A P Poline J B Grasby P J WilliamsS C amp Frackowiak R S J (1995) Analysis of fMRItime-series revisited Neuroimage 2 45ndash53

Frith C D Friston K Liddle P F amp Frackowiack R S J(1991) A PET study of word finding Neuropsychologia 291137ndash1148

Gainotti G Silveri M C Villa G amp Miceli G (1986) Anomiawith and without lexical comprehension disorders Brainand Language 29 18ndash33

Goodglass H (1976) Agrammatism In H Whitaker amp H AWhitaker (Eds) Studies in neurolinguistics (pp 237ndash260)New York Academic Press

Hagoort P Indefrey P Brown C Herzog H Steinmetz Hamp Seitz R J (1999) The neural circuitry involved in thereading of German words and pseudowords A PET studyJournal of Cognitive Neuroscience 11 383ndash398

Hillis A E amp Caramazza A (1995) Representation ofgrammatical categories of words in the brain Journal ofCognitive Neuroscience 7 396ndash407

Hillis A E Rapp B Romani C amp Caramazza A (1990)Selective impairment of semantics in lexical processingCognitive Neuropsychology 7 191ndash243

Kay J amp Ellis A W (1987) A cognitive neuropsychologicalcase study of anomia Implications for psychological modelsof word retrieval Brain 110 613ndash629

Lambon Ralph M A Sage K amp Roberts J (2000) Classicalanomia A neuropsychological perspective on speechproduction Neuropsychologia 38 186ndash202

Martin A Haxby J Lalonde F Wiggs C amp Ungerleider L(1995) Discrete cortical regions associated with knowledgeof color and knowledge of action Science 270 102ndash105

Miceli G amp Caramazza A (1988) Dissociation of inflectionaland derivational morphology Brain and Language 3524ndash65

Miceli G Giustolisi L amp Caramazza A (1991) The

Miceli et al 627

interaction of lexical and non-lexical processing mechanismEvidence from anomia Cortex 27 57ndash80

Miceli G Silveri M Villa G amp Caramazza A (1984) On thebasis for the agrammaticrsquos difficulty in producing main verbsCortex 20 207ndash220

Miceli G Silveri M C Romani C amp Caramazza A (1989)Variation in the pattern of omissions and substitutions ofgrammatical morphemes in the spontaneous speech ofso-called agrammatic patients Brain and Language 36447ndash492

Moro A Tettamanti M Perani D Donati C Cappa S Famp Fazio F (2001) Syntax and the brain Disentanglinggrammar by selective anomalies Neuroimage 13 110ndash118

Ni W Constable R T Mencl W E Pugh K R FulbrightR K Shaywitz S E Shaywitz B A Gore J C ampShankweiler D (2000) An event-related neuroimagingstudy distinguishing form and content in sentenceprocessing Journal of Cognitive Neuroscience 12 120ndash133

Oldfield R C (1971) The assessment and analysis ofhandedness The Edinburgh Inventory Neuropsychologia9 97ndash113

Patterson K amp Hodges J R (1992) Deterioration of wordmeaning Implications for reading Neuropsychologia 301025ndash1040

Paulesu E Frith C U amp Frackowiack R S J (1993) Theneural correlates of the verbal component of workingmemory Nature 362 342ndash345

Perani D Cappa S F Schnur T Tettamanti M CollinaS Rosa M M amp Fazio F (1999) The neural correlatesof noun and verb processing A PET study Brain 1222337ndash2344

Petersen S E Fox P T Posner M I Mintun M amp RaichleM E (1988) Positron emission tomographic studies of theprocessing of single words Nature 331 585ndash589

Schriefers H (1993) Syntactic processes in the productionof noun phrases Journal of Experimental PsychologyLearning Memory and Cognition 19 841ndash850

Shapiro K A amp Caramazza A (submitted) Neuralrepresentation of grammatical categories

Shapiro K A Pascual-Leone A Mottaghy F A GangitanoM amp Caramazza A (2002) Grammatical distinctions in theleft frontal cortex Journal of Cognitive Neuroscience 131ndash8

Shapiro K A Shelton J amp Caramazza A (2000) Grammaticalclass in lexical production and morphological processingEvidence from a case of fluent aphasia CognitiveNeuropsychology 17 665ndash682

Talairach J amp Tournoux P (1988) Co-planar stereotaxicatlas of the human brain A 3-dimensional proportionalsystem an approach to cerebral imaging New YorkThieme

Tissot R Mounin G amp Lhermitte F (1973) Lrsquoagramma-tisme Paris Dessart

Tranel D Adolphs R Damasio H amp Damasio A R(2001) A neural basis for the retrieval of action wordsCognitive Neuropsychology 18 655ndash670

Vandenberghe R Price C J Wise R J S Josephs O ampFrackowiak R S (1996) Semantic system(s) for words orpictures Functional anatomy Nature 383 254ndash256

Vanier M amp Caplan D (1991) CT-scan correlates ofagrammatism In L Menn amp L K Obler (Eds) Agrammaticaphasia (pp 37ndash116) Philadelphia John Benjamins

Warburton E Wise R J S Price C J Weiller C HadarU Ramsay S amp Frackowiak R S J (1996) Noun and verbretrieval by normal subjects Studies with PET Brain 119159ndash179

Worsley K J amp Friston K J (1995) Analysis of fMRItime-series revisitedmdashagain Neuroimage 2 173ndash181

Xu B Grafman J Gaillard W D Ishii K Vega-Bermudez FPietrini P Reeves-Tyer P Di Camillo P amp Theodore W(2001) Conjoint and extended neural networks for thecomputation of speech codes The neural basis of selectiveimpairment in reading words and pseudowords CerebralCortex 11 267ndash277

Zingeser L amp Berndt R (1988) Grammatical class andcontext effects in a case of pure anomia Implications formodels of language production Cognitive Neuropsychology5 473ndash516


of separate neural networks in the processing of nounsand verbs is also supported by neuroimaging studies(eg Perani et al 1999 Warburton et al 1996 MartinHaxby Lalonde Wiggs amp Ungerleider 1995 PetersenFox Posner Mintun amp Raichle 1988) and by electro-physiological investigations (Federmeier Segal Lombro-zo amp Kutas 2000 Dehaene 1995) However theinterpretation of the nounverb dissociation as a specif-ically grammatical effect remains controversial This isbecause grammatical categories correlate with semanticproperties (eg nouns typically denote objects whereasverbs typically denote actions nouns are usually moreconcrete verbs usually more abstract etc) It is notimplausible therefore to suppose that the observeddissociations reflect the organization of the semanticsystem rather than grammatical properties of words(eg Bird Howard amp Franklin 2000 Breedin Saffranamp Schwartz 1998 but see Berndt et al 2002 Caramazzaamp Shapiro in press for arguments and evidence thatchallenge the strong version of this hypothesis)

One way to study the neural representation of thegrammatical properties of words is to focus on a lexicalfeature that is independent from meaning The gram-matical feature gender meets this requirement in manylanguages (Corbett 1991) This property of nouns playsan important role in determining agreement amongwords in noun phrases in the selection of pronominalforms and even in subjectndashverb agreement (eg specify-ing agreement between the subject of a sentence andthe past-participle form of the verb in Italian) Forexample in Italian all the words in a noun phrasemdashdeterminers adject ives and nounsmdashmust agreein number and gender1Consider the case of the phrases(masc = masculine fem = feminine sg = singularpl = plural) lsquolsquoilmasc sg piccolomasc sg pontemasc sgrsquorsquo(the small bridge) lsquolsquoimasc pl piccolimasc pl pontimasc plrsquorsquo(the small bridges) lsquolsquolafem sg piccolafem sg navefem sgrsquorsquo(the small ship) and lsquolsquolefempl piccolefempl navifemplrsquorsquo (thesmall ships) As may be seen the definite determinertakes different forms depending on the gender and thenumber of the nounmdashil and i for masculine singular andplural nouns respectively and la and le for femininesingular and plural nouns respectively Similarly theadjective lsquolsquosmallrsquorsquo is inflected appropriately for numberand gender -o or -i for masculine singular and pluralrespectively and -a and -e for feminine singular andplural respectively As another example consider thecase of clitics In Italian the sentence lsquolsquogive it to mersquorsquo isproduced with cliticized forms in which both the dativepronoun (lsquolsquoto mersquorsquo) and the direct object pronoun lsquolsquoitrsquorsquoare cliticized onto the verb However the specific formof the clitic for the direct object depends on the gram-matical gender and number of the noun for which thepronoun stands If the noun is singular masculine theclitic would be -lo as in dammelo (literally give-me-it)and it would be respectively -li -la and -le for mascu-line plural feminine singular and feminine plural nouns

As these examples illustrate the retrieval of a nounrsquosgrammatical gender is an essential part of the sentenceproduction process

Crucially for our purposes a nounrsquos semantic contentand grammatical gender have only an arbitrary relation-ship This can be readily appreciated by considering thefact that closely related semantic coordinates havedifferent grammatical genders For example the Italiantranslation of the first member of each of the followingpairs of words is masculine while the second is femi-nine lemonorange grapefruitapple tablechair sheetblanket platecup glassbottle spoonfork traincarelephantgiraffe liontiger sunmoon and so on Fur-thermore the grammatical gender of a word is languagespecific While the Italian and French translations ofpencil desk carriage summer and flag are feminineand masculine respectively the opposite is true for seatable limit river and tomato If grammatical genderswere determined by a nounrsquos semantic properties wewould expect semantically related words to have thesame gender and that words would have the samegender across languages Neither condition holds Thusthe results of investigations of the neural mechanismsinvolved in processing grammatical gender can be in-terpreted to reflect grammatical as opposed to semanticaspects of language processing

Although grammatical gender and meaning are re-lated only arbitrarily the same is not true for the relationbetween gender and form In many languages includingItalian Russian and Hebrew for example words of thesame grammatical gender tend to share specific phono-logical properties In the case of Italian a majority ofmasculine singular nouns end in the vowel o (plural i)and a majority of feminine singular nouns end in thevowel a (plural e) However the correlation is farfrom perfect Thus among the 3000 most frequentwords in Italian only 66 of nouns have a lsquolsquoregularendingrsquorsquo (De Mauro Mancini Vedovelli amp Voghera1993) The nonregular nouns consist either of wordsending in e (and a few in i) which are associatedequally frequently with masculine and feminine nounsor masculine nouns that end in a and feminine nounsthat end in o Thus although it has been shown thatsubjects are sensitive to the correlation between genderand word ending in word recognition tasks (eg BatesDevescovi Pizzamiglio DrsquoAmico amp Hernandez 1995) adefinitive decision about a wordrsquos grammatical gendercan only be taken after lexical access has occurred

The independence of grammatical gender from mean-ing and the fact that lexical access is necessary in orderto determine the gender of a noun has made thisgrammatical feature an appropriate tool for investigatingthe process of lexical access in normal language produc-tion (see Caramazza Miozzo Costa Schiller amp Alario inpress Schriefers 1993) These same properties of gram-matical gender make it suitable for the study of therepresentation of grammatical word properties in the

Miceli et al 619




RESULTS

Behavioral Results



Activation Results















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DISCUSSION



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RESULTS

Behavioral Results



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DISCUSSION



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DISCUSSION



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The Neural Correlates of Grammatical Gender: An fMRI Investigation

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Transcript of The Neural Correlates of Grammatical Gender: An fMRI Investigation