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Grammar Processing Outside the Focus of

Attention: An MEG Study

ARTICLE in JOURNAL OF COGNITIVE NEUROSCIENCE · DECEMBER 2003

Impact Factor: 4.69 · DOI: 10.1162/089892903322598148 · Source: PubMed

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Yury Shtyrov

Aarhus University

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Friedemann Pulvermüller

Freie Universität Berlin


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Risto J Ilmoniemi

Aalto University


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http://www.researchgate.net/profile/Risto_Ilmoniemi?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_7http://www.researchgate.net/profile/Risto_Ilmoniemi?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_4http://www.researchgate.net/profile/Risto_Ilmoniemi?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_4http://www.researchgate.net/institution/Aalto_University?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_6http://www.researchgate.net/profile/Yury_Shtyrov?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_4http://www.researchgate.net/institution/Freie_Universitaet_Berlin?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_6http://www.researchgate.net/publication/8931460_Grammar_Processing_Outside_the_Focus_of_Attention_An_MEG_Study?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_3http://www.researchgate.net/?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_1http://www.researchgate.net/profile/Risto_Ilmoniemi?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_7http://www.researchgate.net/institution/Aalto_University?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_6http://www.researchgate.net/profile/Risto_Ilmoniemi?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_5http://www.researchgate.net/profile/Risto_Ilmoniemi?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_4http://www.researchgate.net/profile/Friedemann_Pulvermueller?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_7http://www.researchgate.net/institution/Freie_Universitaet_Berlin?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_6http://www.researchgate.net/profile/Friedemann_Pulvermueller?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_5http://www.researchgate.net/profile/Friedemann_Pulvermueller?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_4http://www.researchgate.net/profile/Yury_Shtyrov?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_7http://www.researchgate.net/institution/Aarhus_University?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_6http://www.researchgate.net/profile/Yury_Shtyrov?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_5http://www.researchgate.net/profile/Yury_Shtyrov?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_4http://www.researchgate.net/?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_1http://www.researchgate.net/publication/8931460_Grammar_Processing_Outside_the_Focus_of_Attention_An_MEG_Study?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_3http://www.researchgate.net/publication/8931460_Grammar_Processing_Outside_the_Focus_of_Attention_An_MEG_Study?enrichId=rgreq-53a343a1-8107-4ef2-b45e-fb81318e0369&enrichSource=Y292ZXJQYWdlOzg5MzE0NjA7QVM6MjczMzExNTM1OTg4NzM2QDE0NDIxNzM5NTY2Nzk%3D&el=1_x_2


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Grammar Processing Outside the Focus of Attention:an MEG Study

Yury Shtyrov 1, Friedemann Pulvermüller 1, Risto Näätänen3,4,and Risto J. Ilmoniemi2,4

Abstract

& To address the cerebral processing of grammar, we used

whole-head high-density magnetoencephalography to record

the brain’s magnetic fields elicited by grammatically correct

and incorrect auditory stimuli in the absence of directed

attention to the stimulation. The stimuli were minimal short

phrases of the Finnish language differing only in one single

phoneme (word-final inflectional affix), which rendered them

as either grammatical or ungrammatical. Acoustic and lexicaldifferences were controlled for by using an orthogonal design

in which the phoneme’s effect on grammaticality was inverted.

We found that occasional syntactically incorrect stimuli elicited

larger mismatch negativity (MMN) responses than correct

phrases. The MMN was earlier proposed as an index of pre-

attentive automatic speech processing. Therefore, its modu-

lation by grammaticality under nonattend conditions suggests

that early syntax processing in the human brain may take place

outside the focus of attention. Source analysis (single–dipole

models and minimum-norm current estimates) indicated

grammaticality dependent differential activation of the leftsuperior temporal cortex suggesting that this brain structure

may play an important role in such automatic grammar

processing. &

INTRODUCTION

For over a century, the processing of language in the

brain has remained one of the most intriguing issues in

science. Although its many aspects have been tackled by numerous studies, little is still known of how our centralnervous system goes about processing grammar. Gram-

mar is one of the main intrinsic properties of the human

language. The very presence of a grammatical system

distinguishes our ability of verbal communication fromall signaling systems used by animals. Earlier investiga-tions of the neurophysiology of syntactic processing

(e.g., Hagoort, Brown, & Groothusen, 1993; Osterhout

& Holcomb, 1992; Neville, Nicol, Barss, Forster, & Gar-

rett, 1991) have led to important insights into this issue.

Three major syntax-related phenomena have been

described on the basis of the brain’s electric responsesto language stimuli. The first one is the so-called early

left anterior negativity (ELAN) occurring, with a latency

of ¹125 msec, in response to function words1 whose

placement in a phrase violates sentence structure rules

(Neville et al., 1991). Secondly, grammatically relatedfrontal negativities with longer latencies (over 250 msec)

have been found, for example, for different morpholog-

ical or syntactic errors (Gunter, Friederici, & Schriefers,

2000; Münte, Schiltz, & Kutas, 1998). Finally, a late

positive shift (often termed P600), reaching its maxi-

mum at ¹600 msec and maximal at centro-parietal

recording sites, has been considered the most robustneurophysiological response to ungrammatical sen-

tences because it could be observed in many linguisticexperiments (for review, see Osterhout, McLaughlin, &

Bersick, 1997; Osterhout & Hagoort, 1999).

When studying the processing of grammar in the brain,

it seems important to control for a number of issues. For

instance, in such studies, subjects are typically asked toattend to presented sentences (e.g., Friederici, Wang,

Herrmann, Maess, & Oertel, 2000; Osterhout & Swinney,

1993; Neville et al., 1991); often, the task is to judge the

sentence grammaticality. In such cases, as attention is

required, one cannot be sure to what extent the regis-tered responses are influenced by brain correlates of

attention rather than by the language-related activity as

such. Attention-related phenomena are known to mod-

ulate a variety of the brain’s evoked responses in a

substantial span of time after stimulus onset and to

involve a number of brain structures including thoseclose to, or overlapping with, the core language areas

(see, e.g., Yamasaki, LaBar, & McCarthy, 2002; Yantis

et al., 2002; Escera, Alho, Winkler, & Näätänen, 1998;

Tiitinen, May, & Näätänen, 1997; Woods, Alho, & Algazi,

1993; Woods, Knight, & Scabini, 1993; Alho, 1992; Picton

& Hillyard, 1974). It also appears possible that subjectspay more attention to incorrect sentences as they try to

1 Medical Research Council, Cognition and Brain Sciences Unit,

Cambridge, UK, 2 Biomag Laboratory, Helsinki University

Central Hospital, 3 University of Helsinki, 4 Helsinki Brain

Research Center

© 2003 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 15:8, pp. 1195–1206


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make some sense of them or that they process grammat-

ical and ungrammatical items using different strategies.

These different strategies and attention variation may

find their reflection in the event-related measures, over-

lapping with true syntax-related activity. Therefore, lim-iting the attention-related effects seems to be necessary

at this stage for further investigation of the neural

processing of syntax.

A methodologically important aspect in such experi-ments is stimulus selection. Grammatically correct andincorrect phrases would inevitably differ in many other

features: e.g., sound onset times and durations in

acoustic stimulation, as well as visual geometry and

overall luminance in visual tasks, would change when

words or affixes are added, removed, or relocated to

modulate grammaticality. Differences even in basicphysical features may lead to differential brain activation

(Korth & Nguyen, 1997; Näätänen & Picton, 1987) that

could in principle overlap with or be misinterpreted as

language-related effects. For example, it was shown that

word-elicited responses are highly dependent on the word length (Osterhout, Bersick, & McKonnon, 1997;

Assadollahi & Pulvermüller, 2003), suggesting the ne-

cessity to control for such factors. Furthermore, the

physical stimulus parameters are of crucial importance

for obtaining ERP effects of syntactic processes (Gunter,Friederici, & Hahne, 1999). It seems most logical to us

to apply an experimental design in which physical

differences between the grammatical and ungrammati-

cal items are controlled for by presenting the same

physical contrasts to the subjects both in ungrammatical

and grammatical contexts; this sort of an orthogonaldesign would help to disentangle purely syntactic effectsfrom those related to the physical stimulus features.

So, in sum, it appears crucial to limit the influence of

attention and different processing strategies on the

brain response to grammatical and ungrammaticalstrings as well as to apply maximal control over stimulus

features using a counterbalanced stimulus design. Con-

trolling for these issues simultaneously in a single study

can be a challenging task, which, as to our knowledge,

has not been realized so far. The present study repre-sents an attempt to achieve this goal.

Here, we addressed brain activity related to grammat-ical processing in the absence of directed attention

towards stimuli and without an active task which would

possibly invite subjects to invent a strategy for dealing

with unusual stimuli. We used a counterbalanced stim-ulus design, which allowed us to control for effects of

the physical and lexical parameters of the stimuli. To

gain high temporal resolution necessary in such an

experiment, we opted for multichannel magnetoence-

phalographic (MEG) recording of brain responses, which can also provide information about the spatial

location of the registered activity.

We utilized the so-called mismatch negativity (MMN),a unique indicator of automatic cerebral processing of

acoustic stimuli, which can be used to investigate the

cerebral processing of speech and language (Shtyrov,

Kujala, Palva, Ilmoniemi, & Näätänen, 2000; Shtyrov &

Pulvermüller, 2002a, 2002b; Näätänen & Winkler, 1999;

Näätänen, 2001; Pulvermüller et al., 2001). MMN, withits major sources of activity in both the supratemporal

and frontal cortices, is a brain response elicited by rare

(deviant) stimuli occasionally presented in a sequence

of frequent (standard) stimuli (Opitz, Rinne, Mecklinger, von Cramon, & Schröger, 2002; Picton, Alain, Otten,Ritter, & Achim, 2000; Alho, 1995). It can be registered

as a negative deflection in the scalp electroencephalo-

graphic recording (EEG), or, as a component in the

brain’s magnetoencephalogram. Importantly, MMN (and

its magnetic counterpart, MMNm, or mismatch field),

can be elicited in the absence of the subject’s attention(Näätänen, 1995; Näätänen & Escera, 2000). It is con-

sidered to reflect the brain’s automatic discrimination of

changes in the auditory sensory input and, furthermore,

to provide an index of experience-dependent memory

traces in the human brain (Näätänen, 2001; Shtyrov et al., 2000; Kraus, McGee, Carrell, & Sharma, 1995).

Recent evidence suggests that the MMN reflects brain

processing of such language elements as phonemes

(Näätänen et al., 1997; Näätänen, 1999, 2001), syllables

(Shtyrov et al., 2000; Alho et al., 1998), and words(Shtyrov & Pulvermüller, 2002b; Korpilahti, Krause,

Holopainen, & Lang, 2001; Pulvermüller et al., 2001).

Importantly, we have recently found a specific pattern

of MMN responses to inflectional affixes (Shtyrov &

Pulvermüller, 2002a), which is best explained by the

activation of distinct cortical memory traces for suchaffixes realized as distributed strongly connected pop-ulations of neurons (Pulvermüller, 1999, 2001). Inflec-

tional (functional) affixes are important tools utilized in

many languages for realizing syntax and conveying

grammatical information. This suggests that the MMNcould be a sensitive tool for probing neural elements

responsible for the processing of grammar in the brain.

We have therefore set out to investigate the cerebral

processing of syntax using a whole-head high-density

MEG system, and recorded MMNm elicited by either grammatically correct or ungrammatical phrases, while

the subjects were distracted from the auditory stimula-tion by engaging in a different activity. The stimuli were

short pronoun–verb phrases differing only in their final

phoneme, which rendered them either as grammatical

or as grammatically incorrect (see Figure 1, Table 1, andMethods section for details). Any possible effects of

acoustic (phonetic) differences were ruled out by using

an orthogonal design reversing the effect of the pho-

neme and word combination on grammaticality.

RESULTS

Magnetic MMN responses were elicited in all conditions.The overall analysis of single equivalent current dipole

1196 Journal of Cognitive Neuroscience Volume 15, Number 8


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(ECD) sources showed that the grammaticality of the

deviant stimuli had a significant effect on the MMNm

dipole moment, F (1,8) = 6.35, p < .04. Separate

analyses for each hemisphere showed that the gram-

maticality had no effect for the right hemisphere alone

( p > .65), whereas in the left hemisphere its effect wassignificant, F (1,8) = 6.67, p < .04: syntactically incor-rect phrases elicited stronger MMNm activity than didthe correct ones (see Figures 2, 3, and 4a). This effect

also became manifest as a significant Pronoun £ Suffix

interaction ( p < .04). It neither depended on the

direction of the acoustic contrast between the suffixes

( p > .12) nor on the preceding pronoun ( p > .63) as

such, but solely on the correctness of the phrase as a

whole (Figure 4a).The L1 minimum-norm current estimate (MCE)

analysis of the MEG signal suggested activation of distributed cortical sources spread out over the left

Figure 1. Spectrograms of acoustic Finnish-language stimuli used in the four experimental conditions. The standard and deviant stimuli in each

pair are indiscriminable up to the divergence point (marked with white arrowheads) at their end. The standard–deviant contrasts are identical

across all experimental conditions. Grammatically incorrect phrases are marked with an asterisk (*). See Table 1 for more details.

Table 1. Auditory Finnish-Language Stimuli Used in the Four Experimental Sessions

Condition 1a Condition 1b Condition 2a Condition 2b

Deviant Mä tuo n * Mä tuot * Sä tuo n Sä tuot

(‘‘I bring’’) (‘‘*I bring’’) (‘‘*you bring’’) (‘‘you bring’’)

Standard * Mä tuot Mä tuo n Sä tuot * Sä tuo n

(‘‘*I bring’’) (‘‘I bring’’) (‘‘you bring’’) (‘‘*you bring’’)

English translations are given in parentheses, ungrammatical phrases are marked with (*). The standard and deviant stimuli in each pair areindiscriminable up to the divergence point at their end. The standard–deviant contrasts are identical across all experimental conditions. Criticalcontrasts are in bold.

Shtyrov et al. 1197


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temporal cortex (see Figure 5). This activation was

significantly stronger (Figure 4b) for the syntactically

incorrect than for correct deviant phrases, F (1,11) =

9.46, p < .011. Again, this difference was clearly

affected by the grammaticality, but neither by the pro-

noun context ( p > .25) nor by the suffix ( p > .8).

No differences between conditions were found in the

right hemisphere.

As one could notice in Figure 5 (upper left panel),

some additional activation could be suggested to appear

also in the left inferior frontal cortex, which would be

consistent with earlier results (Neville et al., 1991). We

Figure 3. Waveforms

(magnetic field gradient) of

MMNm responses in the left

hemisphere (grand average).

Both ungrammatical deviant

stimuli produced stronger

MMNm than the corresponding

correct phrases regardless of

the acoustic, phonetic,

phonological, or lexical

properties of the stimuli.

Temporal gradiometer with the

most prominent responses

(channel 0242) was used for

producing this diagram.

Figure 2. ECD models of MMNm activity in the left hemisphere (grand average). Both ungrammatical deviant stimuli produced stronger MMNm

source than the corresponding correct phrases regardless of the acoustic, phonetic, phonological, or lexical properties of the stimuli. Red and blue

contour lines indicate positive and negative values of the normal component of the magnetic field on the helmet-shaped MEG-array surface,respectively. The arrows show the dipole location and are proportional in relative size to the response magnitude. The view is from the left side of

the head.



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analyzed activity in this area as well, but although we

found that, similarly to the supratemporal source, it was

somewhat increased for the ungrammatical deviant

phrases, this effect was not significant.

In the current study, the grammaticality affected theMMNm responses, but had no significant effect on the

responses to the standard stimuli only (cf. Figures 3

and 6). We also found no effects of grammaticality on

the latency of the MMNm responses, the mean latency in

the left hemisphere being 213 msec after the divergencepoint (mean SEM 19 msec). We also looked for significant

differences between the conditions in later time intervals

(>350 msec) but did not find any. Specifically, we would

like to note that no late deflection of the magnetic

response, which could be related to the P3- or P600-like

components of the EEG signal (Osterhout, McLaughlin

et al., 1997; Donchin, 1981), could be detected (seeFigures 3 and 6). We also noticed that the [n]-ending

stimuli produced larger MMNm dipole moment than did

the [t]-ending ones. However, this independent of the

grammaticality effect (which could in principle be due

to differences in attack and voicedness between the twophonemes) was only significant for comparison between

Conditions 2a and 2b ( p < .02; but p > .95 for

Figure 4. Grand-average mean

and standard error of mean of

the MMNm dipole-moment

magnitude (nanoamperemeter,

nAm) in the left hemisphere:

(A) calculated for single

equivalent current sources

(ECD), (B) calculated for L1

MCE (calculated as a sum of

dipole moments of all

simultaneously activatedadjacent current sources). Both

analyses indicated that the

ungrammatical deviant stimuli

produced a stronger MMNm

source than the corresponding

correct phrases regardless of

the acoustic, phonetic,



Figure 5. Grand-average L1

MCEs of MMNm in the left

hemisphere. The MCE solutions

are projected unto

triangularized gray matter

surface. Both ungrammatical

deviant stimuli produced

stronger MMNm source than

the correct phrases regardless

of the acoustic, phonetic,



Shtyrov et al. 1199


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Conditions 1a and 1b) and only in ECD but not in MCE

analysis, so it does not seem to present a solid finding.

Finally, we found no topographical effects of any of the

factors.

DISCUSSION

In the present study, mismatch responses were elicited

by grammatically correct or incorrect pronoun–verb

phrases. The choice of the single final phoneme in allphrases rendered the whole structure as syntactically

correct or as ungrammatical. To exclude confounding

effects, the phonetic and lexical contrasts were counter-

balanced over the experimental conditions using a fully orthogonal stimulus design. The subjects were instructed

not to attend to the stimuli; they did not perform any stimulus-related language task.

We found that MMNm responses to the syntactically

incorrect phrases were always larger than those to the

correct phrases ending in the same word regardless of thedirection of the acoustic contrast or of the preceding

pronoun. That is, neither the choice of the final phoneme

nor that of the pronoun in the sentence altered this effect,

which was present in both conditions in which the

deviant stimuli were grammatically inconsistent.

The present data are consistent with earlier results onthe cerebral processing of syntactic violations (see In-

troduction), which suggested negative-going compo-nents of brain responses at time delays of 125 msec or

more following syntactic errors. The present modifica-

tion of the MMN may be related to this early negativity.There are, however, important differences between the

present study and the earlier ones. Importantly, we were

able to find syntactically related effects in the absence of

directed attention towards the stimuli as subjects were

engaged in a different task and were instructed to ignorethe stimulation. The violations in this study modulated

the MMN response, which is considered to reflect a pre-

attentive automatic level of auditory processing in the

cerebral cortex (Näätänen & Alho, 1995; Näätänen &Escera, 2000; Tiitinen et al., 1997). This suggests that the

brain may be capable of automatic syntactic analysis of

the incoming language signals already at relatively early

stages of speech processing. This conclusion is support-ive of the earlier suggestion that early negativities, such

as ELAN, may reflect automatic stages of syntactic pro-cessing (Friederici, 2002; Gunter et al., 2000; Hahne &

Friederici, 1999).

The ELAN/N125 component reported in the previous

studies was found in response to phrase structure

violations (examp le: mathematician’s *of proof thetheorem ) and has been difficult to replicate (Takazawa

et al., 2002; Neville et al., 1991), whereas in the present

study the MMN modulation was elicited at a similar

latency by agreement violations ( mä tuo*t, sä tuo*n ),suggesting that a larger variety of grammatical violations,

including agreement violations, could be reflected by early left-lateralized neurophysiological responses.

An established view in psycholinguistics is that sen-

tence comprehension is incremental, that is, a substantialamount of processing occurs immediately after a word in

a sentence is perceived, prior to the perception of the

following word (Pulvermüller, 2001; Traxler & Pickering,

1996; Marslen-Wilson & Tyler, 1975; Marslen-Wilson,

1987). This implies that by the time a violation occurs, a

certain amount of analysis has been carried out by the

parsing system, a position largely shared by severalsyntactic parsing models that may substantially differ

otherwise (e.g., Friederici, 2002; van Gompel & Pickering,2001; Spivey & Tanenhaus, 1998; MacDonald, Pearlmut-

ter, & Seidenberg, 1994; Trueswell, Tanenhaus, & Kello,

1993; Ferreira & Henderson, 1990; Frazier, 1987; Mitchell,1987). During such analysis, an expectation for syntac-

tically possible subsequent morphemes may have built

up. When this expectation fails to realize and the parser

therefore encounters a syntactic error (i.e., a mismatchbetween preceding and incoming morphemes), it sends

an ‘‘error signal’’. Such an error signal may be due to a

lack of neuronal links between lexical representations

which results in a failure to provide priming between the

representations of morphemes that do not match syntac-tically (Pulvermüller, 2002). In our case, the memory trace

Figure 6. Waveforms

(magnetic field gradient)

of auditory responses to

standard stimuli in the left

hemisphere (grand

average). Although the

MMNm responses were

affected by grammaticality,

there was no significant

effect on the standard

responses (cf. Figure 3).Temporal gradiometer

with the most prominent

responses (channel 0242)

was used for producing

this diagram.



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(neuronal assembly) representing the expected correct

affix (Shtyrov & Pulvermüller, 2002a) would have been

primed by the preceding context. Language-related

priming effects are known to reduce negative-going

components of the event-related potentials (e.g., Hol-comb & Neville, 1990; Bentin, McCarthy, & Wood, 1985).

In ungrammatical conditions, this priming would be

absent which should lead to an increased response as

compared with grammatically consistent stimuli. Thus,the parser’s error signal, the unprimed activation of morpheme-related neuron ensembles, may be the basis

of syntactically related early left-lateralized responses,

including the larger MMN response to grammatical viola-

tions observed here. Interestingly again, this putative

process seems to occur outside the focus of attention

(i.e., at least to some degree automatically).The strongest activation produced by the syntactic

abnormalities in the current study was located in the left

temporal cortex. We take this as an indication that there

is a distributed neuronal system in the left superior

temporal lobe that contributes to grammar processing.This area has been repeatedly suggested as a neural

substrate for a variety of language functions such as

phonetic/phonological processing, lexical access, seman-

tic and syntactic processing (e.g., Shtyrov et al., 2000;

Shtyrov & Pulvermüller, 2002a, 2002b; Martin & Chao,2001; Pulvermüller et al., 2001; Friederici et al., 2000;

Price, 2000; Helenius, Salmelin, Service, & Connolly,

1998; Näätänen et al., 1997; Binder et al., 1995), which

is in line with the current results. There probably could

be an overlap between these functions with respect to

underlying neuroanatomical structures, but it seemsreasonable to suggest that they are carried out by atleast partially distinct neuronal networks (Pulvermüller,

2002). It also appears justified to suggest that the left-

lateralized neural systems contributing to the grammat-

ically related MMN in the superior temporal lobe may bedistinct from those responsible for classical bilateral

acoustic-related MMNs.

Interestingly, although we found the strongest activa-

tion in the temporal cortex, a left anterior-frontal re-

sponse was reported earlier (Neville et al., 1991;Pulvermüller & Shtyrov, in press). As we found no

significant grammatically dependent variation for theinferior frontal activity, we conclude that in the present

data the frontal source did not seem to play a major role

in syntactic processing. There could be several possible

explanations for this: in principle, it could be that theadvanced source analysis algorithm used in this study,

MCE, provides more accurate source localization than

potential mappings used earlier. As a viable alternative

explanation, one could suggest that the anterior source

might not be detectable in MEG due to its spatialorientation (and/or, vice versa, that the posterior source

may be difficult to image using EEG). Both of these

explanations are supported by earlier MEG research, which has also demonstrated more temporal than frontal

activity to syntactic violations when subjects were asked

to judge grammaticality of auditory presented sentences

(Friederici et al., 2000). However, a frontal ELAN gener-

ator was reported in MEG, although this was also accom-

panied by a temporal one (Gross et al., 1998). Finally, it ispossible that the absence of directed attention to the

stimuli and of an active task may have reduced the frontal

activity. This would imply that the inferior frontal source

is of smaller importance for automatic syntactic parsing.To fully answer the question of the cortical generator(s)for the syntactic MMN modulation observed here and

their relation to ELAN activation, further studies using

similar paradigm would be needed.

Finally, we could not find brain dynamics that might

relate to the late centro-parietal positive shift (P600)

reported earlier in ERP studies of grammatical violations. Again, one could suggest that cortical generators of late

slow positive shifts are not optimal for MEG recordings,

although, among late positivities, at least the P3 was

successfully registered using MEG (Berg, Kakigi, Scherg,

Dobel, & Zobel, 1999; Mecklinger et al., 1998; Tarkka,Stokic, Basile, & Papanicolaou, 1995). However, the latter

still does not rule out the possibility that the syntactically

related P600 response has no clear magnetic correlate.

Another possible reason for the absence of a late gram-

maticality effect in our present data set is the absence of an attention-demanding task. It has been proposed

earlier that the positive shifts to grammatical violations

reflect attention-dependent processes, for example, at-

tempts at a structural reanalysis of a sentence fragment

(Osterhout & Holcomb, 1992). In this view, the present

absence of clear dynamics in P600 time range (alsoconfirmed in a separate EEG study, Pulvermüller &Shtyrov, in press) might suggest a role of attention in

the generation of the late positivity. Because subjects did

not focus their attention on language in this study, they

most likely did not engage in re-parsing the deviantstrings, and thus the centro-parietal positive shift did

not occur. This would be consistent with notions that

P600 reflects phrase reanalysis and repair (Friederici,

1997; Osterhout, Holcomb, & Swinney, 1994) and may

not reflect specific syntactic processes per se (Münte,Heinze, Matzke, Wieringa, & Johannes, 1998).

In conclusion, the present results suggest that the early syntax parsing of spoken language may not require

focused attention and therefore could probably be auto-

matic to a certain extent. The cortical structures in the left

superior temporal lobe appear to play an important rolein carrying out such automatic grammar processing.

METHODS

Subjects

Twelve healthy right-handed (handedness assessed

according to Oldfield, 1971; no left-handed family mem-bers) native Finnish speakers (age 21–29, 7 women)

Shtyrov et al. 1201


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wi th norm al hea ring an d no rec ord of neurol ogi-

cal diseases were presented with four sets of spoken

native-language stimuli in four separate experimental

conditions.

Stimuli

The four short-phrase stimuli (see Figure 1 and Table 1)

were prepared with requirements that the acoustic,phonetic, and phonological difference between thestandard and the deviant stimuli is identical in each

condition and that the stimuli themselves are as similar

acoustically as possible. These Finnish stimuli were:

(1a) Mä tuon (‘‘I bring,’’ syntactically correct),

(1b) * Mä tuot (‘‘*I bring,’’ syntactically incorrect: the verb tuot is in the second person inflection form, while

the pronoun mä, ‘‘I,’’ requires the first person); here

and throughout we use linguistic convention of marking

ungrammatical phrases with an asterisk (*),

(2a) * Sä tuon (‘‘*you bring,’’ syntactically incorrect:the verb tuon is in the first person form, while the

pronoun sä, ‘‘you,’’ requires the second person);

(2b) Sä tuot (‘‘you bring,’’ syntactically correct).2

Therefore, it is the single final phoneme, inflec-tional affix [n] or [t], which may render the whole phrase

as syntactically correct or as ungrammatical. Remarkably,

for ‘‘mä’’-stimuli (1a and 1b), the final [n] means that

the phrase is grammatical, while for the ‘‘sä’’-stimuli

(2a and 2b) the combination is exactly the opposite:

[t] is needed in the end for the whole phrase to becorrect. This orthogonal design is implemented inorder to control for purely acoustic, phonetic, and pho-

nological effects on mismatch responses as well as

for putative lexical and semantic differences between

the words. As seen in Table 1 presenting the complete stimulus

design, the stimuli starting with the same pronoun were

always presented in the same experimental condition,

one of them serving as deviant and the other one as

standard stimulus. Thus, the standard–deviant acoustic– phonetic contrast, the critical variable determining the

MMN (Näätänen & Alho, 1997), was the same in allconditions. However, in each pair of conditions (1a, b

and 2a, b) the responses were elicited either by correct

or incorrect deviant forms. Furthermore, while the

acoustic contrasts were identical in the corresponding

conditions ( tuon/tuot in 1a–2a and tuot/tuon in 1b–2b),the syntactic contrast (determined by the pronoun) was

the opposite: stimuli ending in tuon were correct in

conditions 1a and 1b, whereas tuot -ending phrases were

correct in Conditions 2a and 2b. This fully orthogonaldesign, therefore, allowed us to control not only for

possible acoustic/phonetic effects of the verb contrast,

but even for an unlikely late differential effect of the twopreceding pronouns.

For stimulus preparation, we recorded multiple rep-

etitions of each word uttered by a female native speaker

of Finnish. With great care we selected a combination of

recordings, whose vowels matched in their fundamental

frequency (F0) and whose overall length and maximalsound energy were identical both for sä and mä and for tuon and tuot . We then combined the four words in the

four abovementioned phrases with a 50-msec gap be-

tween the pronoun and the verb; all phrases were766 msec in duration. Thus, the two stimuli in eachcondition were perceptually identical up to a time point

(referred to as divergence point) shortly before their

end (see Figure 1). In order to determine this diver-

gence point, a gating experiment was performed, in

which initial fragments of increasing duration of bothtuon and tuot were presented to the subjects who wereasked whether they could perceive any difference be-

tween the two. In average, they could identify the

difference only when the first 336 msec ( SEM 9 msec)

of the words were presented (which equals to 636 msec

after the onset of the complete phrase). This perceiveddivergence point is therefore considered as the onset

of the perceptual standard/deviant contrast and a zero

time reference in the current study. Importantly, it is

the same in all pairs of stimuli due to their design.

All stimuli were normalized to have the same peaksound energy. For the analysis and production of the

stimuli, we used the Cool Edit 96 program (Syntrillium

Software, AZ).

Acoustic Stimulation

All four experimental conditions (Table 1) were per-formed with every subject, their order being counter-

balanced across the subject group. The stimuli were

binaurally presented at 50 dB above the individual

hearing threshold (determined using the experimentalstimuli) via headphones connected to an STIM setup

(Neuroscan Labs, VA). The interstimulus (stimulus

onset asynchrony, SOA) interval was 1450 msec. In each

condition, the deviant stimulus was presented with a

probability of 16.7% among the repetitive standardstimuli: A pseudorandom sequence of stimuli was cre-

ated so that there were always at least two standardstimuli between any two deviants. During the stimula-

tion, the subjects were seated in a magnetically shielded

chamber and instructed to watch a silent video film

of their own choice and to pay no attention to theauditory stimulation.

Magnetoencephalographic Recording

The evoked magnetic field was recorded (passband0.03–200 Hz, sampling rate 600 Hz) with 204 planar

gradiometer channels of a whole-head Neuromag Vec-

torview MEG system (Elekta Neuromag, Helsinki) duringthe auditory stimulation (Ahonen et al., 1993). The



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recordings started 100 msec before stimulus onset

and ended 1450 msec thereafter. The responses were

on-line averaged separately for the standard and devi-

ant stimuli in each condition. Epochs with voltage

variation exceeding 150 m V at either of two bipolar eye-movement electrodes or magnetic-field gradient

variation exceeding 3000 femtotesla per centimeter (fT/

cm) at any gradiometer channel were excluded from

averaging. The recordings for each condition containedat least 120 accepted responses to deviant stimuli (cor-responding to ¹600 standard responses) in order to

achieve satisfactory averaged signal quality. This require-

ment led to slight variations in overall recording dura-

tion, which in average constituted 2 hr per subject.

MEG Data Processing

The averaged responses were filtered off-line (passband

1–20 Hz). The period of 50 msec before the divergence

point was used as the baseline. The MMNm was ob-

tained by subtracting the averaged response to thestandard stimuli from that to the deviant ones. The

responses were evaluated separately for each subject

for all experimental conditions. Two approaches were

used for analyzing the neuromagnetic data.

Single-Dipole Fit

Fifty-four gradiometer channels on each side of the

magnetometer helmet were used for assessing cortical

responses in the left and right cerebral hemispheres. By

means of Neuromag sequential single-dipole fittingsoftware (Elekta Neuromag), the generator sources(equivalent current dipoles [ECDs]) of the MMNm were

estimated (Ilmoniemi, 1993). Only dipole models ex-

plaining more than 65% of the field gradients were

selected for statistical analysis; otherwise, the dipole-moment value was set to 0 nAm. The range of latencies

(as determined from peak dipole moment) accepted for

the MMNm responses was 100–350 msec after the

divergence point and the best fit (maximal goodness

of fit/dipole moment) was entered into analysis. Thedipole moments of the MMNm generators were calcu-

lated and compared between the experimental condi-tions. ECD fits were successfully constructed only for

nine subjects, suggesting that the single-dipole model

may not be optimal for explaining the syntactically

related activity. We therefore performed minimum-norm current analysis, which uses a distribution of

multiple current sources for modeling the neuromag-

netic signal.

L1 Minimum-Norm Current Estimate

The estimation of the cortical sources of the measured

neuromagnetic activity was performed using L1 MCEson the basis of recordings from all 204 gradiometer

channels equally distributed over the entire scalp. The

minimum-norm method provides a solution to the in-

ver se probl em of localizing neu ral activ ity in the

brain from its external recordings by revealing the

unique constellation of active neuronal current ele-ments that models the recorded magnetic field distri-

bution with the smallest amount of overall activity

(Uutela, Hämälä inen, & Somersalo, 1999; Ilmoniemi,

1993; Hämäläinen & Ilmoniemi, 1984, 1994). Among theinfinitely many current source combinations that canexplain a given scalp topography of the magnetic

field, this technique finds one that does so with the

least amount of overall activity. This minimal solution

should be preferred for the sake of parsimoniousness.

The L1 MCE minimizes the integral of the current

amplitude and reveals a realistic and robust constellationof localized generators. It is applicable when it can be

assumed a priori that the source distribution consists of

discrete areas of neuronal activity (in contrast with a

situation when no essential a priori information is

available, in which case L2 minimum norm estimate would be preferable). This assumption of local ized

sources is justified in the investigation of language

because neuroimaging results obtained with different

methodologies proved the activation of discrete cortical

areas during a variety of language tasks (Pulvermüller,2001; Price, 2000). A triangularized gray matter surface

was used for projecting MCE solutions for the each

subject individually and for the grand average magnetic

field. The dipole moments of the MMNm generators

were calculated (as a sum of dipole moments of all

simultaneously activated adjacent current sources) andcompared between the experimental conditions. TheMCE source analysis was successfully carried out for all

12 subjects, which suggests that it could be a better

method for analyzing distributed neural activity (as in

case of language-related phenomena) than ECD.

Statistical Analysis

The data were subjected to analyses of variance (AN-

OVA). Parameters of current sources constructed for each hemisphere were analyzed separately for ECD

and MCE. We compared them between conditions usingthe factors grammaticality (syntactically correct vs. in-

correct deviant stimulus), pronoun ( sä vs. mä ), and

suffix ([t] vs. [n]).

Ethical Considerations

All subjects gave their written informed consent to

participate in the experiments and were paid for

their participation. The experiments were performedin accordance with the Helsinki Declaration. Ethical

permission for the experiments was issued by the Re-

search Ethics Committee of Helsinki University CentralHospital.

Shtyrov et al. 1203


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Acknowledgments

We thank Olaf Hauk, Vadim Nikulin, Maritta Maltio-Laine, SimoMonto, Tuomas Murdoch, Christopher Bailey, Jussi Nurminen,

Johanna Salonen, Anthea Hills, Seppo Kähkönen, and WilliamMarslen-Wilson for their contribution at different stages of this

work. We would also like to thank three anonymous refereesfor their helpful comments and constructive critique.

Reprint requests should be sent to Dr. Yury Shtyrov, Medical

Research Council, Cognition and Brain Sciences Unit, 15Chaucer Road, CB2 2EF Cambridge, UK, or via e-mail: [email protected].

Notes

1. Function words are grammatical words, such as articles,conjunctions, auxiliary verbs, and so on and so forth, whichhave no concrete meaning by themselves and are utilized for grammatical purposes.2. These spoken-language stimuli represent the most usualcolloquial pronunciation used in the Helsinki area; therefore,

we only used Finnish subjects who had lived in the Helsinkiarea for a minimum of 6 years before the experiment.

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2003 Shtyrov Grammar Processing Outside the Focus of Attention