Neuron 52, 941–952, December 21, 2006 ª2006 Elsevier Inc. DOI 10.1016/j.neuron.2006.12.
002
The Neural Basis of Language
Development and Its Impairment
Angela D. Friederici1,*
1
Max Planck Institute for Human Cognitive
and Brain Sciences
Leipzig
Germany
The neural correlates of early language development
and language impairment are described, with the adult
language-related brain systems as a target model.
Electrophysiological and hemodynamic studies indi-
cate that language functions to be installed in the
child’s brain are similar to those of adults, with lateral-
ization being present at birth, phonological processes
during the first months, semantic processes at 12
months, and syntactic processes around 30 months.
These findings support the view that the brain basis
of language develops continuously over time. Discon-
tinuities are observed in children with language im-
pairment. Here, the observed functional abnormalities
are accompanied by structural abnormalities in infe-
rior frontal and temporal brain regions.
Introduction
Language is acquired without much effort and seems to
develop as the brain matures. The milestones of normal
language development as evident from behavior can be
defined as follows. In the first days after birth, infants are
able to discriminate between different phonemes and
distinguish the sentence melody (prosody) of their
mother tongue from that of other languages. By the
age of 9 months, they have acquired the inventory of
phonemes as well as the specific stress patterns of their
mother tongue (for a review, see Jusczyk, 1997). The
child now understands the first words and starts to
produce first words between 11 and 13 months, with
a lexicon of about 50 to 75 items by the age of 16 months
and a clear vocabulary spurt between the age of 18 and
24 months (see Bates and Goodman, 1999). Syntactic
structures are acquired continuously in the second
and third year of life, with first productions of two-
word utterances at the age of 18 to 24 months, and later
more-word utterances. The basic knowledge of the syn-
tactic word order constraints is present around the age
of 2.5 years (Höhle et al., 2001).
The complexity of the language system is often only
detected when examined developmentally and realized
when language development derails. The way in which
language development can be impaired is multifaceted.
It can manifest itself in the inability to acquire phonolog-
ical, semantic, and syntactic information (specific lan-
guage impairment [SLI]) (Levy and Schaeffer, 2003), or
grammatical aspects selectively (grammatical-SLI)
(Van der Lely, 2005), the inability to read (Dyslexia)
(Shaywitz et al., 1990), or to understand others (autism)
(Baron-Cohen et al., 1985, 1997). As the former two
types of impairments are the most relevant when it
*Correspondence: angelafr@cbs.mpg.de
Review
comes to explaining early language development and
its impairment, the present review will focus on these.
The neural parameters of the neural basis of normal
language development, however, are still not under-
stood entirely, and the description of its impairment truly
remains incomplete. This is due to the fact that there are
dramatic constraints on the investigation of the neural
correlates of language development. Direct animal
model, cellular, and molecular approaches are impossi-
ble, and genetic approaches can only describe correla-
tions between phenotype and genotype. At the macro-
scopic level, functional magnetic resonance imaging
(fMRI) and structural imaging (diffusion tenor imaging
[DTI]) and magnetencephalography (MEG) are not
always applicable in young children and infants. Electro-
encephalography (EEG) and near-infrared spectroscopy
(NIRS) are the only methods that can readily be applied
to this age group without much constraint.
The review consists of two parts. The first part
discusses the neural correlates of normal language
development using adult data as the reference model,
and the second part covers the available neurophysio-
logical and neuroanatomical studies on specific lan-
guage impairment.
Neural Correlates of Normal Language Development
Studies on the neural basis of normal language develop-
ment have focused on different aspects of the language
system, namely (1) phonological processes concerning
suprasegmental information (i.e., prosody, sentence
melody) and segmental information (phonemes, i.e.,
speech sounds relevant for word meaning), (2) lexical-
semantic processes (i.e., processes that deal with
word forms and word meanings), and (3) syntactic pro-
cesses concerning the grammatical relation of different
words in a sentence.
Phonological Processes
An infant’s first exposure to language is based on
acoustic-phonetic and phonological information (for
a review, see Kuhl, 2004). As an initial step into language,
the infant must be able to differentiate speech from non-
speech sounds. When comparing forward to backward
played speech in a NIRS experiment, sleeping newborns
showed a larger increase in cerebral blood volume over
the left temporal brain regions for forward speech (Pena
et al., 2003). In an fMRI experiment with 3-month-olds,
Dehaene-Lambertz et al. (2002) also found that strong
left hemispheric activation comprised the superior tem-
poral gyrus for speech sounds as measured by forward
and backward speech compared to silence. This activa-
tion included Heschl’s gyrus and extended to the supe-
rior temporal sulcus and the temporal pole (see Figure 1).
Differences between forward and backward speech
were found in the left angular gyrus and precuneus. An
additional right frontal brain activation observed only
in awake infants was interpreted to reflect attentional
factors. Both studies suggest a left hemispheric domi-
nance for speech in early infancy which is similar to
that of adults.
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One language-relevant aspect of the speech input are
features of its intonational contour, which allow the seg-
mentation of the input into structural units. Crucially,
prosodic breaks (intonational phrase boundaries) signal
syntactic phrase boundaries, thus allowing an easy en-
trance into the target language. Using NIRS, Homae
et al. (2006) investigated the brain basis for prosodic
processes in 3-month-old infants. Examining the hemo-
dynamic response to normal speech and speech with
flattened intonational contours, they found bilateral
activation in the temporo-parietal and the frontal cor-
tices for both conditions. A direct comparison of normal
and flattened speech revealed the right temporo-
parietal cortex, suggesting a right hemispheric domi-
nance for the processing of sentential prosody (pitch
information) similar to adults (Meyer et al., 2004) to be
present already at 3 months of age.
Using a temporally more sensitive method, namely the
measurement of event-related brain potentials (ERP), it
was shown that the specific ERP component, identified
as an adult brain response to intonational phrase
boundaries (Steinhauer et al., 1999; Pannekamp et al.,
2005), was present in 8-month-olds (Pannekamp et al.,
2006). Its topographic distribution over the scalp was
similar to adults, but its peak was somewhat delayed.
This suggests that the brain systems involved in pro-
cesses of identifying intonational phrase boundaries
are similar in infants and in adults, but take more time
in earlier development.
The other language-relevant aspect in the speech
input is acoustic information concerning the different
phonemes and stress patterns of words in a given
language. The acoustic parameter of duration is most
Figure 1. Brain Activation in Response to
Speech Sounds in 3-Month-Olds
(A) Averaged brain activation in response to
speech sounds (forward speech and back-
ward speech against rest) in 3-month-old in-
fants. L, left hemisphere; R, right hemisphere.
(B) (Left panel) Averaged brain activation in 3-
month-old awake infants for forward speech
against backward speech. (Right panel) Aver-
aged hemodynamic responses of forward
speech and backward speech in awake and
asleep infants.
relevant during spoken language perception, as it differ-
entiates not only between different phonemes (i.e., short
vowel [full] versus a long vowel [fool]) but also between
words with different stress patterns (i.e., long first sylla-
ble [the ábstract] versus short first syllable [to abstráct]).
There are a number of ERP studies that provide
evidence for the early discrimination of different
language-relevant sounds and sound patterns. These
studies used the auditory oddball paradigm in which
participants are presented with a sequence of stimuli,
most of which are identical (standard) but interrupted
by a deviant stimulus differing on one or more acoustic
parameters.
ERP studies in adults have reported a mismatch
negativity (MMN) as a neural correlate of auditory
discrimination in such a paradigm (Näätänen, 1990; for
a review, see Näätänen et al., 2001). By means of
MEG, the MMN in adults was localized in the auditory
cortices bilaterally (e.g., Alho et al., 1998). fMRI studies
that applied a mismatch paradigm also reported activa-
tion in the auditory cortices bilaterally with stronger
activation in the right superior temporal gyrus to reflect
the discrimination of pitch and duration in nonspeech
sounds and additional frontal activations to be attribut-
able to attentional processes (Opitz et al., 2002; Doeller
et al., 2003; Schall et al., 2003; Molholm et al., 2005).
When directly comparing speech and music with
respect to duration discrimination, significantly more
activation was found in the left superior temporal gyrus
(BA 22/42) for speech and in the right Heschl’s gyrus
(BA 41) for music (Tervaniemi et al., 2006).
While the mismatch response in adults is always
expressed in a negativity, the response in infants is
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943
reported as either a negativity (Cheour et al., 1998; Kush-
nerenko et al., 2001) or a positivity (Dehaene-Lambertz
and Dehaene, 1994; Friederici et al., 2002; Leppänen
et al., 1999; Weber et al., 2004) and sometimes with dif-
ferent scalp distributions and latencies (for a review, see
Cheour et al., 2000).
Presenting a sequence of five syllables with the last
one either being the same or a different one, Dehaene-
Lambertz and Dehaene (1994) reported a mismatch
response in the form of a posterior positivity for the
deviant syllable (/ga/) compared to the standard syllable
(/ba/) in 2- to 3-month-old infants. An MMN-like negativ-
ity was found for vowel discrimination in the first months
of life (Cheour et al., 1997) and even in newborns
(Cheour-Luhtanen et al., 1995). Mismatch responses
either expressed as a negativity or as a positivity have
been observed in different languages, such as Finnish,
German, and English, for vowel contrasts (Cheour
et al., 1997; Leppänen et al., 1999; Pihko et al., 1999;
Friederici et al., 2002, 2004) and for consonant contrasts
(Dehaene-Lambertz and Baillet, 1998; Rivera-Gaxiola
et al., 2005) early during development.
Interestingly, evidence for a language-specific phone-
mic discrimination seems to establish between the age
of 6 and 12 months (Cheour et al., 1998; Rivera-Gaxiola
et al., 2005). While younger infants aged 6 and 7 months
show discrimination for the phonemic contrast relevant
and not-relevant for their target language, older infants
aged 11 and 12 months only display a discrimination
response for the phonemic contrast in their target
language.
The importance of word stress for word recognition
during speech perception was shown in a recent ERP
study with infants learning Dutch (Kooijman et al.,
2005). In this study, 10-month-olds recognized two-
syllable words with stress on the first syllable in contin-
uous speech after they had heard the words in isolation.
Recognition was reflected in a greater negativity
between 350 and 500 ms over the left hemisphere for
familiar words than for unfamiliar words.
Lexical-Semantic Processes
In adults, a particular ERP component, the N400 (i.e., a
negative going wave form peaking at around 400 ms),
has been identified to correlate with lexical-semantic
processes. The semantic N400 effect is reflected in
a larger amplitude for words that are semantically incon-
gruous to a given context than words that are congruous
and it is taken to indicate semantic integration difficul-
ties. Moreover, the N400 amplitude is found to be larger
for pseudowords than for words. This finding is inter-
preted to reflect the difficulty in identifying a lexical
representation for the pseudoword in the mental lexicon
(for reviews, see Kutas and Federmeier, 2000; Kutas and
van Petten, 1994).
Concerning the neural basis of semantic processes,
MEG studies with adults localized the sentential N400
in the auditory cortex bilaterally (Halgren et al., 2002;
Mäkelä et al., 2001) or with an additional left inferior
frontal source (Maess et al., 2006). fMRI experiments
with adults applying comparable sentential paradigms
indicate an involvement of the middle and superior
temporal gyri bilaterally with a larger activation in the
left hemisphere (Kuperberg et al., 2000; Ni et al., 2000;
Friederici et al., 2003) and sometimes the basal ganglia
bilaterally (Friederici et al., 2003).
The adequate description of the brain basis of the
N400 observed for pseudowords requires that the
paradigms used in the fMRI studies and the ERP ex-
periments are comparable. Such fMRI studies found
a variety of activations including anterior and inferior
portions of the left temporal lobe (Mummery et al.,
1999), anterior and middle portions of the left superior
temporal gyrus (Kotz et al., 2002), and the inferior frontal
gyrus/sulcus bilaterally (Mummery et al., 1999; Rossell
et al., 2001; Kotz et al., 2002). The frontal activations
are discussed as being related to task demands,
whereas the temporal activations are taken to reflect
lexical processes.
Lexical and Semantic Processes at the Word Level
An ERP study with 11-month-old infants suggests a dif-
ferential brain reaction to known and unknown words ex-
pressed as a negativity around 200 ms after word onset
with longer amplitude to familiar versus unfamiliar words
(Thierry et al., 2003). Testing 14- to 20-month-olds, Mills
et al. (2004) also found a negativity between 200 and 400
ms that was larger for known than for unknown words.
The distribution of which, however, seems to change
from bilateral at 13 months to left hemisphere dominant
at 20 months of age (Mills et al., 1997). These data dem-
onstrate that by the end of their first year infants are able
to differentiate between familiar and unfamiliar words,
but it is not clear whether infants at this age process
the semantics of words similar to adults.
In another word-processing study, the effects of word
experience (training) and vocabulary size (word produc-
tion) were tested (Mills et al., 1997). In this word-learning
paradigm, 20-month-olds acquired novel words either
paired with a novel object or without an object. After
training, the infants’ ERPs showed a repetition effect
indicated by a reduced N200-500 amplitude to familiar
and novel unpaired words, whereas ERPs indicated an
increased bilaterally distributed N200-500 for novel
paired words. This finding is taken to indicate that the
N200-500 is linked to word meaning. The interpretation
of this early effect as a semantic one is challenged, given
that semantic effects in adults are observed later, i.e., in
the N400.
Friedrich and Friederici (2004, 2005a, 2005b) used an
ERP paradigm appropriate to test for semantic knowl-
edge and knowledge about lexical forms in young
children between 12 and 19 months. In this paradigm,
a picture of an object is presented together with an
auditory stimulus that is either a word matching the
object’s name or not (semantic knowledge) or with
a pseudoword that is phonotactically legal or illegal
(phonological knowledge). In 12-month-olds, an early
fronto-central negativity between 100-400 ms was
found for words that were congruous with a picture
compared to words that were incongruous. This early
effect was interpreted as a familiarity effect reflecting
the fulfillment of a lexical expectation after seeing the
picture of an object. At the age of 14 months, an N400
effect was observed for semantically incongruous
words in addition to the early negativity effect for the
congruous words. By 19 months, an N400 was found
for semantically incongruous and for phonotactically
Neuron
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legal pseudowords, but not for phonotactically illegal
pseudowords. This indicates that at this age both real
words and phonotactically legal pseudowords are con-
sidered as word candidates, whereas phonotactically
illegal pseudowords are not. The observed semantic
N400 effect in 14- and 19-month-olds reached signifi-
cance later, lasted longer, and had more frontal distribu-
tion when compared to adults (compare Figure 3). The
latency differences suggest slower lexical-semantic
processing in children than in adults. The more frontal
distribution could mean either that children’s semantic
processes are still more image-based, as adults show
a frontal distribution when pictures instead of words
are processed (West and Holcomb, 2002), or that chil-
dren recruit frontal brain regions associated with in-
creased attentional processes (e.g., Opitz et al., 2002)
in addition to those subserving semantic processing.
There are a few fMRI studies with children investigat-
ing semantic processes at the word level. Using
a semantic judgment task requiring the evaluation of
the semantic relatedness of two auditory words, a study
with 9- to 15-year-old children revealed activation in the
temporal gyri bilaterally (BA 22), in the left middle tempo-
ral gyrus (BA 21), and in the inferior frontal gyri bilaterally
(BA 47/45) (Chou et al., 2006). Correlations with age were
observed in the left middle temporal (BA 21) and the
right inferior frontal gyrus (BA 47). The increased frontal
activation was interpreted to reflect a broader semantic
search and the temporal activation to relate to more
efficient access to lexical-semantic representations.
Using a semantic categorization task, a recent fMRI
study with 5- to 10-year-old children found activation
in similar frontal and temporal regions of the left hemi-
sphere and in the left fusiform gyrus (BA 37, BA 20), sug-
gesting language to be left-lateralized as early as 5 years
(Balsamo et al., 2006).
Semantic Processes at the Sentential Level
The processing of semantic information at the sentential
level has only recently been investigated in children
younger than 4 years of age (Silva-Pereyra et al.,
2005a; Friedrich and Friederici, 2005b). Previous studies
with 5- to 26-year-olds (Holcomb et al., 1992), 8- to
13-year-olds (Atchley et al., 2006), and 6- to 13-year-
olds (Hahne et al., 2004) reported N400-like negativities
for semantically anomalous sentences in children of all
age groups. In a study with 2.5-, 3-, and 4-year-olds,
English learning children listened passively to sentence
stimuli that were either semantically correct or anoma-
lous (e.g., My uncle will watch/blow the movie) (Silva-
Pereyra et al., 2005a, 2005b). Children at 2.5 years
showed an early starting frontally distributed negativity
peaking around 500 ms for semantically anomalous
compared to normal sentences (Silva-Pereyra et al.,
2005a). In contrast, 3- and 4-year-olds demonstrated a
negative slow wave negativity peaking at around 400
ms, 600 ms, and 800 ms. According to the authors, these
anteriorly distributed negativities may reflect either one
general semantic integration mechanisms or different
semantic mechanisms, which, however, are not speci-
fied further, except the late negativity, taken to reflect
sentence closure.
An N400-like semantic effect at the sentence level has
been reported even in younger German learning children
aged 19 and 24 months (Friedrich and Friederici, 2005c).
This study used sentences in which there was the
presence or absence of a semantic mismatch between
the verb and its object argument (e.g., The cat drinks
the ball/the milk). For 19-month-olds, a first negativity
was observed between 400 and 500 ms, followed by
a sustained negativity between 600 and 1200 ms. In
24-month-olds, the negativity was found to start at
300 ms and to last until 1200 ms, whereas in adult
listeners, the N400 effect was present between 300
and 800 ms. These data indicate that the processes
underlying the N400 effect in children are similar to
adults, but that the integration of the object noun into
the sentence context requires more processing time in
young children.
Semantic processes of word integration into senten-
tial context have been tested in English and German
children between the ages of 1 year and 26 years using
different sentence constructions. The studies available
suggest an early sensitivity to semantic anomalies as
reflected by an N400-like negativity (1) whose duration
decreases with age, as shown in children between
19 months and 2 years (Friedrich and Friederici, 2005c)
and between 5 and 15 years (Holcomb et al., 1992), indi-
cating faster integration processes, (2) whose amplitude
decreases linearly between 5 and 15 years, indicating
less reliance on sentential context with older age
(Holcomb et al., 1992), and (3) whose distribution is
wider in younger children than in older children or adults
(Friedrich and Friederici, 2005c; Atchley et al., 2006;
Holcomb et al., 1992). Its general morphology, however,
appears to remain similar across the age from childhood
to adulthood.
Given the ERP evidence, it is likely that the same brain
regions observed in adults are recruited during child-
hood. The available fMRI studies conducted with chil-
dren at the sentential level, however, have not looked
at semantic processes in isolation but rather at text com-
prehension. fMRI data from 6- to 10-year-olds listening
to short texts (compared to resting baseline) revealed
bilateral activation in the auditory cortices, specifically
in the superior temporal gyrus and Heschl’s gyrus, the
planum temporale, as well as in the inferior frontal gyrus,
the anterior cingulate regions, and parietal regions
(Ulualp et al., 1998). When listening to stories (compared
to reversed speech), 6-year-old children showed activa-
tion in the left superior temporal gyrus and sulcus
extending back to the angular gyrus and in the left middle
temporal gyrus (Ahmad et al., 2003). When listening to
stories in which content words were missing (compared
to resting baseline), 6- to 14-year-olds displayed activa-
tion in anterior superior temporal and posterior temporal
areas bilaterally and in the ‘‘classical inferior-frontal
language areas’’ (Wilke et al., 2005). In this study, the
inferior frontal activation was interpreted to result from
semantic memory retrieval necessary to fill in the missing
word. Future fMRI studies separating semantic and syn-
tactic processes at the sentential level will have to reveal
which of the activations found for text comprehension
are semantic in nature.
Syntactic Processes
ERP studies of syntactic processing in adults have
shown that violations of syntactic rules are associated
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945
with two ERP components: a late, centro-parietal
positivity (P600) preceded by a left anterior negativity
(LAN), or an early LAN (ELAN). The P600 is not only
seen for syntactically anomalous sentences requiring
syntactic repair, but also for temporally syntactic ambig-
uous sentences requiring syntactic reanalysis (Osterh-
out and Holcomb, 1992; Osterhout et al., 1994) and
moreover appears to modulate as a function of syntactic
complexity (Kaan et al., 2000). According to a recent
neurocognitive model of auditory language comprehen-
sion (Friederici, 2002), the ELAN component is viewed
to reflect difficulties in the process of initial local
structure building. The LAN is taken to reflect difficulties
in morpho-syntactic processing during the assignment
of grammatical relations in a second stage. The P600
is seen to reflect difficulties in the stage of integration
where syntactic and thematic structure have to be
mapped onto each other.
The brain basis of these processes has been investi-
gated in adults using space- and time-sensitive imaging
methods. Studies investigating syntactic processes
using fMRI usually report superior temporal activation
(including the anterior and posterior portion) as well as
activation in the inferior frontal gyrus (including Broca’s
area and/or the frontal operculum) (for reviews, see
Bookheimer, 2002; Friederici, 2002; Grodzinsky and
Friederici, 2006). The anterior portion of the superior
temporal gyrus and the frontal operculum have been
identified as those brain regions that are involved in
syntactic phrase structure building processes reflected
by the ELAN (Friederici et al., 2000, 2006).
Currently, no fMRI study and only very few ERP stud-
ies on syntactic processing in children are available.
Figure 2. Averaged Event-Related Brain Po-
tentials to Auditorily Presented Correct and
Syntactically Incorrect Sentences in 24-
Month-Olds, 32-Month-Olds, and Adults
Vertical line indicates the onset of the violat-
ing word. (A) ERPs at left anterior electrode
F7 for the early syntactic effect (ELAN). (B)
Distribution maps for the difference between
correct and incorrect sentences in the time
windows in which the ELAN effect is ex-
pected. Dark blue indicates negativity, which
is clearly present at left anterior site for 32-
month-olds and for adults. (C) ERPs at cen-
tro-parietal electrode PZ for the late syntactic
effect (P600).
Studies investigating morphosyntactic violations in
English (e.g., My uncle will watch/watching the movie),
which usually elicit an LAN-P600 pattern in adults,
reported no LAN effects but only a P600-like positivity
for 3- and 4-year-olds (Silva-Pereyra et al., 2005a). For
slightly younger 30-month-old children, the positivity
observed between 600 and 1000 ms did not reach signif-
icance (Silva-Pereyra et al., 2005b).
A recent ERP study in German investigating the
processing of local phrase structure violations (e.g.,
The lion in the roars instead of The lion in the zoo roars),
which elicits an ELAN-P600 in adults, found a biphasic
ERP pattern consisting of a delayed ELAN and a late
P600 in children at 2.5 years (Oberecker et al., 2005).
When applying the same phrase structure violation
paradigm to 2-year-olds, the ERPs revealed a late
P600, but no ELAN component nor any other left-
lateralized negativity preceding the P600 (Oberecker
and Friederici, 2006) (see Figure 2). The combined
results indicate that the neural basis for the ELAN and
the P600 follow a different developmental pace. What
could be the explanation for the finding that the ELAN
establishes later than the P600? Adult data indicate
that the ELAN reflects highly automatic processes of
initial structure building that are unaffected by
strategy-inducing factors such a instruction (Hahne
and Friederici, 2002) or a variation of the amount of
correct versus incorrect sentences in an experimental
set (Hahne and Friederici, 1999). In contrast, the P600
modulates as a function of these factors and thus seems
to reflect late controlled processes. On the basis of
these adult findings, the ERP data of the children would
suggest that those highly automatic processes reflected
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by the ELAN are not yet established in 2-year-olds,
whereas processes of syntactic and thematic integra-
tion are reflected by the P600 are. By pure logic, the find-
ing of a P600 in the absence of the ELAN would mean
that online syntactic structure building either was pres-
ent by not reflected in the ERP or that the build up of the
sentence’s structure was not entirely based on syntactic
features (as the preposition in not only constrains the
next word syntactically [i.e., noun instead of verb] but
also semantically [i.e., location instead of action]). The
presence of a delayed but adult-like ELAN component
at 2.5 years appears to indicate that the neural mecha-
nisms of online syntactic structure building are present
at this age, although the processes are clearly slower
in children than in adults.
In summary, the studies on normal language develop-
ment indicate that the language-related ERP compo-
nents reflecting (1) lexical-semantic processes (N400
by ERP) that involve the middle and superior temporal
gyri and possibly the left inferior frontal gyrus, (2) syntac-
tic processes (ELAN/LAN-P600 by ERP) which in adults
rely on the left superior temporal gyrus mainly its ante-
rior and posterior portion and the left inferior frontal
gyrus, and (3) prosodic processes (CPS by ERP) which
rely on the superior temporal and frontal opercular
cortex of the right hemisphere appear to change in their
latency and duration, but not in their basic morphology
from childhood to adulthood.
Neural Correlates of Language Impairment
Before turning to the studies on individuals diagnosed
with SLI, I will briefly discuss a unique family, spanning
four generations, in which about half of the members
of the first three generations are affected by a severe
disorder of language production.
The KE family was initially described as a severe form
of verbal apraxia (Hurst et al., 1990). Subsequently, the
family’s deficit has been described as a linguistic deficit
concerning inflectional morphology (Gopnik, 1990;
Gopnik and Crago, 1991). Further studies indicated
that the disorder is not specific to language, but affects
the processing of phonology and syntax as well as non-
linguistic oral praxia (Vargha-Khadem et al., 1995;
Watkins et al., 1999). Thus, it appears that the primary
deficit is a nonlinguistic one that affects language in
a secondary manner, and therefore, studies of this fam-
ily may not directly reflect true language abnormalities.
Members of the KE family have been investigated with
respect to abnormalities in brain function and brain
structure. The functional data available have used differ-
ent methods (PET, fMRI) and different tasks (word
repetition, verb generation). These report overactiva-
tions and underactivations for particular brain regions.
Liegeois et al. (2003) using verb generation tasks in the
fMRI reported underactivation for Broca’s area (BA
44/45), and Vargha-Khadem et al. (1998) found overacti-
vation in Broca’s area for a word-repetition task in a PET
study. The latter study registered underactivation in
the left posterior middle temporal gyrus, whereas the
former found overactivation in the superior and middle
temporal gyrus bilaterally. Abnormalities in the basal
ganglia (again over and underactivation) were reported
for the caudate nucleus and the putamen. As the func-
tional interpretation of overactivation and underactiva-
tion is not yet clear, our conclusion will only consider
deviances from the normal pattern and not their direc-
tion. Functional abnormalities are seen in speech/
language-relevant areas in the temporal and inferior
frontal cortex but also for those structures in the basal
ganglia, namely caudate and putamen known to be
crucial for controlling and selecting motor sequences
necessary for articulation (Volkmann et al., 1992).
Structural studies indicate reduced gray matter
volume and density of Broca’s area (BA 44/45) and
increased gray matter volume and density for the poste-
rior temporal gyrus/sulcus bilaterally (Vargha-Khadem
et al., 1998; Belton et al., 2002). All studies available
found reduced volume or gray matter density in the
head of the caudate nucleus (Vargha-Khadem et al.,
1998; Belton et al., 2002; Watkins et al., 2002). This univ-
ocal finding for the subcortical structures has led to the
suggestion that the genetic abnormality in the KE family
may selectively affect the development of the caudate
nucleus or the basal ganglia in general (Watkins et al.,
1999).
The genetic analyses conducted initially proposed
region 7q31 at chromosome 7 (Fisher et al., 1998) and
later more specifically FOXP2 located on chromosome
7q31 (Lai et al., 2001) as being functionally related to
the observed pathology of speech and language in the
KE family. In further studies, FOXP2 expression was
correlated with the development of motor-related brain
circuits in the mouse and the human (Lai et al., 2003).
This latter finding is in line with the phenotype of the
KE family, which is characterized by severe facial dys-
praxia, affecting an articulatory movements accompa-
nied by expressive and receptive language impairments
and which can clearly be differentiated from the SLI
phenotype. A differentiation of these two phenotypes
is supported by genetic evidence that FOXP2 is not
a major susceptibility gene for SLI (Newbury et al.,
2002; O’Brien et al., 2003) and therefore cannot be
considered to be ‘‘the language gene.’’
Specific Language Impairment
Specific language impairment (SLI) is defined as a devel-
opmental disorder that selectively affects the domain of
language processing. Children with SLI perform below
their age on language tasks requiring the processing
of phonological, semantic, or syntactic information
(Levy and Schaeffer, 2003), despite normal intelligence,
an adequate learning environment, and the absence of
peripheral hearing problems or emotional problems
(Bishop, 1992). Its genetic basis has been described
by the SLI Consortium (2002, 2004) as involving chromo-
somes 16q and 19q.
Different views on the underlying deficit of SLI have
been put forward. One view posits that children with
SLI, or at least certain subgroups, suffer from a deficit
specific to the domain of language or a subdomain,
i.e., grammar (Clahsen, 1989; Rice and Oetting, 1993;
Van der Lely, 1994). Another perspective holds that
SLI is not due to a processing deficit specific to lan-
guage but to a deficit in temporal auditory processing
(Merzenich et al., 1993; Tallal et al., 1993) or even to def-
icits that are general in nature, such as a reduced capac-
ity in processing (Bishop, 1994; Kail, 1994), or a deficit in
the procedural memory system (Ullman and Pierpont,
Review
947
Figure 3. Averaged ERP for Spoken Words Congruous and Incongruous with a Picture Presented
Vertical line indicates the word onset. (A) ERPs of typically developing children and adults showing a semantic N400 effect except for the
youngest group. (B) ERPs of 19-month-old infants who were diagnosed with risk for SLI at the age of 2.5 years.
2005). All of these views are based on empirical data—
mostly, however, from behavioral studies. Here, I will
not advocate one or the other view, but rather review
the neurophysiological studies conducted with children
with SLI.
Phonological and Lexical-Semantic Processes
Neurophysiologically, 2-month-old infants from families
at risk for SLI demonstrate a massively delayed brain
responses for the discrimination of two syllables with
vowels of different durations, i.e., long versus short
(Friedrich et al., 2004). Note that a mismatch discrimina-
tion response for long versus short vowels in CV sylla-
bles even differentiated newborns with risk for dyslexia
from those without risk (Leppänen et al., 1999; Pihko
et al., 1999).
By the age of 4–5 months, infants from families at risk
displayed a reduced brain response for the discrimina-
tion of the stress pattern of a two-syllable word (Weber
et al., 2004). A longitudinal study showed in a retrospec-
tive analysis that the reduced brain response observed
by the age of 4–5 months correlated with the children’s
impaired language abilities diagnosed at the age of
4 years (Weber et al., 2005). These data suggest
that early insufficiencies in the processing of duration
information can lead to language impairments in pro-
duction and comprehension later in life.
With respect to semantic processes, children who
were diagnosed as having an advanced risk for SLI
based on behavioral tasks at the age of 2.5 years already
by 19 months of age demonstrate ERP patterns for
lexical-semantic processes that differed from their age-
matched controls. These children did not show an N400
as found with normally developing children at this age
(Friedrich and Friederici, 2006) and even at the age of
14 months (Friedrich and Friederici, 2005b) (see Fig-
ure 3). The absence of the N400 effect in 19-month-old
SLI children may reflect insufficient lexical-semantic
representations that prohibit the normal detection of the
semantic mismatch between the word and the picture
context.
An ERP study with 10- to 12-year-old children diag-
nosed with SLI on the processing of semantic informa-
tion at the sentence level still reveals a pattern of brain
responses that are different from age-matched typically
developing children. The latter show an N400-P600
pattern in response to a semantic violation, whereas
the SLI children show no N400 component, but only
a P600 (see Figure 4). The absence of the N400 effect
suggests weaker lexical-semantic representation of
the critical words (Sabisch et al., 2006).
Thus, ERP studies with SLI children indicate that
those who are diagnosed with impaired verbal language
skills at the age of 3 or later fail to show normal age-
adequate ERP patterns already during their first months
of life. During the first months of life, infants at risk for
SLI are characterized by a massively delayed mismatch
response for the discrimination of syllable duration.
Such a delay may affect processes at the level of
a word’s phonological stress pattern, at the level of
word form and word semantics, and even semantic pro-
cesses at the sentential level. The studies reviewed indi-
cate that impairments can indeed be observed at all of
these levels. Whether the N400 deficits observed in the
latter two studies, however, are based on a primary
deficit in temporal auditory processing cannot be deter-
mined on the basis of the data presented.
The deficit in phonological and lexical-semantic
processes as revealed by the SLI children’s ERP pattern
can be related to a recent fMRI study with a Finnish
family diagnosed to suffer from SLI. Hugdahl et al.
(2004) conducted an auditory fMRI experiment with
five members of a family with SLI (10–70 years) and six
age-matched controls. The controls’ brain activation to
vowels, pseudowords, and words was bilateral in the
superior and middle temporal gyri with a slight laterali-
zation to the left, areas usually involved in auditory, pho-
nological, and word processing. There was also a small
Neuron
948
Figure 4. Averaged ERPs to Auditorily Presented Correct and Semantically Incorrect Sentences
Vertical line indicates the onset of the violating word. (A) ERPs of typically developing children and adults showing a semantic N400 effect. (B)
ERPs of 10- to 12-year-old children diagnosed with SLI.
activation in the right inferior frontal lobe, a region known
to support memory and attention (Demonet et al., 1994).
In the SLI family, the activation was also bilateral in the
temporal lobes, but it was smaller and weaker than in
controls. The reduction of the activation in the STS/
MTG, in areas usually involved in speech perception
and the mapping of acoustic-phonetic cues onto lexical
representations (Scott and Johnsrude, 2003), could
likely be related to the impairments in decoding the
phonological structure of pseudowords and words. This
interpretation would also be in line with the observed
ERP abnormalities in SLI children.
Syntactic Processes
With respect to syntactic processes, Van der Lely (2005)
hypothesizes a specific subgroup of SLI characterized
by selective impairment of a syntactic processing com-
ponent in the core language system. Others view the
grammatical impairment to result from a deficit in proce-
dural memory (Ullman and Pierpont, 2005).
In one of the first neurophysiological studies, Neville
et al. (1993) compared children with language impair-
ment with typically developing children. SLI children
presented a normal ERP pattern for the processing of
content words, but not for function words that carry
grammatical information. Function words that normally
elicit a LAN evoked a more bilateral or even right-
lateralized negativity in language-impaired children.
In an ERP study with 12- to 14-year-old children diag-
nosed with a selective grammatical impairment (G-SLI)
on the basis of behavior tests (e.g., Van der Lely et al.,
1998; Van der Lely, 2005), it was found that these chil-
dren do not demonstrate the syntax-related ELAN com-
ponent for violations of nonlocal syntactic violations
observed in normal age-matched controls. As these
children showed an N400 in relation to semantic aspects
of processing, the results were taken as evidence for
a selective impairment of grammatical aspects of lan-
guage processing (Van der Lely and Fonteneau, 2006).
Neuroanatomy of the SLI Brain
The brain basis of SLI is not well investigated. There are,
however, a number of MRI studies assessing structural
brain abnormalities in children with SLI (for an overview,
see Ullman and Pierpont, 2005). Regional abnormalities
have been reported for language-related and other
areas. Investigating 9-year-olds, Gauger et al. (1997)
found a volume decrease in the left pars triangularis as
part of the Broca’s area, and Jernigan et al. (1991)
reported a volume decrease in the left posterior tempo-
ral region. Moreover, it appears that children with SLI
have a less leftward structural asymmetry for frontal
and temporal language-related regions (Gauger et al.,
1997; Plante et al., 1991).
In addition to abnormalities in the temporal, orbito-
frontal, dorsolateral, and medial frontal cortex, abnor-
malities in the inferior frontal regions in SLI children
have been reported in some studies (Gauger et al.,
1997; Jernigan et al., 1991). In adults, the inferior frontal
gyrus (Broca’s area) has been functionally defined to
support phonological processes (dorsal portion) and
semantic processes (anterior ventral portion) as well
as syntactic processes (posterior ventral portion) (Book-
heimer, 2002; Friederici, 2002). In the absence of de-
tailed fMRI studies in normal children, it is not entirely
clear to which functional aspects the observed struc-
tural abnormalities should be related. There is, however,
a recent proposal which takes frontal abnormality to
support the notion of a deficit in procedural memory
(Ullman and Pierpont, 2005).
Leonard et al. (2002) compared the neuroanatomy of
children with SLI and those suffering from dyslexia. They
reported a reduced leftward asymmetry in the planum
temporale for the SLI group. Like most other anatomical
studies, this one suggests a reduced volume in the left
planum temporale. In adults, this area is known to be in-
volved in the segregation and identification of sound pat-
terns in speech and other acoustic patterns containing
spectotemporal information (Griffiths and Warren, 2002).
Review
949
A structural abnormality in the planum temporale could
explain the deficit in temporal auditory processing ob-
served behaviorally (Tallal et al., 1993) and in ERP studies
(Friedrich et al., 2004), which in turn could have major
impact on phonological and therefore on semantic and
syntactic processes.
Given the limited data available to explain SLI, we are
not in the position to decide which of the views put
forward is the most valid. It may well be that there are
at least two different subgroups, one that affects gram-
matical processes specifically, be it due to a deficit in
the core language system (Van der Lely, 2005) or to
a deficit in the procedural memory system (Ullman and
Pierpont, 2005), and one that affects all aspects of
language processing due to a deficit in auditory tempo-
ral processing (Tallal et al., 1993). If this is the case, the
former group should show abnormalities in the left
inferior frontal cortex, whereas the latter group should
demonstrate abnormalities in the superior temporal
gyrus, in particular the planum temporale.
Conclusions
The available studies on the neural basis of normal
language development suggest that the brain systems
underlying language processing are in place already in
early development. The processes supported by these
developing brain systems change quantitatively though
not qualitatively over time. In contrast, impaired lan-
guage development is correlated with abnormalities in
the neurophysiological patterns of different aspects
of language processing and with abnormalities in the
structures of areas known to support language pro-
cesses in the healthy adult brain.
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