Postlexical Integration Processes in
Language Comprehension: Evidence
from Brain-Imaging Research
COLIN M. BROWN, PETER HAGOORT, AND MARTA KUTAS
ABSTRACT Language comprehension requires the activation,
coordination, and integration of different kinds of linguistic
knowledge. This chapter focuses on the processing of syntactic
and semantic information during sentence comprehension,
and reviews research using event-related brain potentials
(ERPs), positron emission tomography (PET), and functional
magnetic resonance imaging (fMRI). The ERP data provide
evidence for a number of qualitatively distinct components
that can be linked to distinct aspects of language understanding.
In particular, the separation of meaning and structure in
language is associated with different ERP profiles, providing a
basic neurobiological constraint for models of comprehension.
PET and fMRI research on sentence-level processing is at
present quite limited. The data clearly implicate the left perisylvian
area as critical for syntactic processing, as well as for aspects
of higher-order semantic processing. The emerging
picture indicates that sets of areas need to be distinguished,
each with its own relative specialization.
In this chapter we discuss evidence from cognitive neuroscience
research on sentence comprehension, focusing
on syntactic and semantic integration processes. The
integration of information is a central feature of such
higher cognitive functions as language, where we are
obliged to deal with a steady stream of a multitude of information
types. Understanding a written or spoken
sentence requires bringing together different kinds of
linguistic and nonlinguistic knowledge, each of which
provides an essential ingredient for comprehension.
One of the core tasks that faces us, then, is to construct
an integrated representation. For example, if a listener is
to understand an utterance, then at least the following
processes need to be successfully completed: (a) recognition
of the signal as speech (as opposed to some other
kind of noise), (b) segmentation of the signal into constituent
parts, (c) access to the mental lexicon based on
the products of the segmentation process, (d) selection
of the appropriate word from within a lexicon containing
some 30,000 or more entries, (e) construction of the
appropriate grammatical structure for the utterance up
to and including the word last processed, and (f) ascertaining
the semantic relations among the words in the
sentence. Each of these processes requires the activation
of different kinds of knowledge. For example, segmentation
involves phonological knowledge, which is largely
separate from, for instance, the knowledge involved in
grammatical analysis. But knowledge bases like phonology,
word meaning, and grammar do not, on their own,
yield a meaningful message. While there is no question
that integration of these (and other) sources of information
is a prerequisite for understanding, considerable
controversy surrounds the details.
Which sources of knowledge actually need to be distinguished?
Is the system organized into modules, each
operating within a representational subdomain and
dealing with a specific subprocess of comprehension?
Or are the representational distinctions less marked or
even absent? What is the temporal processing nature of
comprehension? Does understanding proceed via a
fixed temporal sequence, with limited crosstalk between
processing stages and representations? Or is comprehension
the result of more or less continuous interaction
among many sources of linguistic and nonlinguistic
knowledge? These questions, which are among the most
persistent in language research, are now gaining the attention
of cognitive neuroscientists. This is an emerging
field, with a short history. Nevertheless, progress has
been made, and we present a few specific examples in
this chapter.
A cognitive neuroscience approach to language might
contribute to language research in several ways. Neurobiological
data can, in principle, provide evidence on
the representational levels that are postulated by different
language models—semantic, syntactic, and so on (see
the section on PET/fMRI). Neurobiological data can
COLIN M. BROWN and PETER HAGOORT Neurocognition of
Language Processing Research Group, Max Planck Institute
for Psycholinguistics, Nijmegen, The Netherlands
MARTA KUTAS Department of Cognitive Science, University
of California, San Diego, Calif.
882 LANGUAGE
reveal the temporal dynamics of comprehension, crucial
for investigating the different claims of sequential and
interactive processing models (see the sections on the
N400 and the P600/SPS). And, by comparing brain activity
within and between cognitive domains, neurobiological
data can also speak to the domain-specificity of
language. It is, for example, a matter of debate whether
language utilizes a dedicated working-memory system
or a more general system that subserves other cognitive
functions as well (see the section on slow brain-potential
shifts).
Postlexical syntactic and semantic
integration processes
In this chapter we focus specifically on what we refer to
as postlexical syntactic and semantic processes. We do
not discuss the processes that precede lexical selection
(see Norris and Wise, chapter 60, for this subject), but
rather concern ourselves with processes that follow
word recognition. Once a word has been selected within
the mental lexicon, the information associated with this
word needs to be integrated into the message-level representation
that is the end product of comprehension. If
this integration is to be successful, both syntactic and semantic
analyses need to be performed.
At the level of syntax, the sentence needs to be parsed
into its constituents, and the syntactic dependencies
among constituents need to be specified (e.g., What is
the subject of the sentence? Which verbs are linked with
which nouns?). At the level of semantics, the meaning of
an individual word needs to be merged with the representation
that is being built up of the overall meaning of
the sentence, such that thematic roles like agent, theme,
and patient can be ascertained (e.g., Who is doing what
to whom?). These syntactic and semantic processes lie at
the core of language comprehension. Although words
are indispensable bridges to understanding, it is only in
the realm of sentences (and beyond in discourses) that
they achieve their full potential to convey rich and varied
messages.
The field of language research lacks an articulated
model of how we achieve (mutual) understanding. This
lack is not too surprising when we consider the problems
that confront us in devising a theory of meaning for
natural languages, let alone the difficulties attendant on
combining such a representational theory with a processing
model that delineates the comprehension process
at the millisecond level. However understandable,
the lack of an overall model has meant that the processes
involved in meaning integration at the sentential
level have received scant experimental attention. The
one area in which quite specific models of the relationship
between semantic representations and on-line language
processing have been proposed is the area of
parsing research. Here, a major concern has been to assess
the influence of semantic representations on the
syntactic analysis of sentences, with a particular focus on
the moments at which integration between meaning and
structure occurs (cf. Frazier, 1987; Tanenhaus and
Trueswell, 1995). Research in this area has concentrated
on the on-line resolution of sentential-syntactic ambiguity
(e.g., “The woman sees the man with the binoculars.”
Who is holding the binoculars?). The resolution of this
kind of ambiguity speaks to the separability of syntax
and semantics, as well as to the issue of sequential or interactive
processing. The prevailing models in the literature
can be broadly separated into autonomist and
interactive accounts.
In autonomous approaches, a separate syntactic knowledge
base is used to build up a representation of the syntactic
structure of a sentence. The prototypical example
of this approach is embodied in the Garden-Path model
(Frazier, 1987), which postulates that an intermediate
level of syntactic representation is a necessary and obligatory
step during sentence processing. This model stipulates
that nonsyntactic sources of information (e.g.,
message-level semantics) cannot affect the parser’s initial
syntactic analysis (see also Frazier and Clifton, 1996;
Friederici and Mecklinger, 1996). Such sources come
into play only after a first parse has been delivered.
When confronted with a sentential-syntactic ambiguity,
the Garden-Path model posits principles of economy, on
the basis of which the syntactically least complex analysis
of the alternative structures is chosen at the moment
the ambiguity arises. If the chosen analysis subsequently
leads to interpretive problems, this triggers a syntactic
reanalysis.
In the most radical interactionist approach, there are no
intermediate syntactic representations. Instead, undifferentiated
representational networks are posited, in which
syntactic and semantic information emerge as combined
constraints on a single, unified representation (e.g., Bates
et al., 1982; Elman, 1990; McClelland, St. John, and
Taraban, 1989). In terms of the processing nature of the
system, comprehension is described as a fully interactive
process, in which all sources of information influence
the ongoing analysis as they become available.
A third class of models sits somewhere in between the
autonomous and radical interactionist approaches. In
these so-called constraint-satisfaction models, lexically represented
information (such as the animacy of a noun or
the transitivity of a verb) but also statistical information
about the frequency of occurrence of a word or of syntactic
constructions play a central role (cf. MacDonald,
Pearlmutter, and Seidenberg, 1994; Spivey-Knowlton
BROWN, HAGOORT, AND KUTAS: BRAIN-IMAGING OF LANGUAGE COMPREHENSION 883
and Sedivy, 1995). The approach emphasizes the interactive
nature of comprehension, but does not exclude
the existence of separate representational levels as a
matter of principle. Comprehension is seen as a competition
among alternatives (e.g., multiple parses), based
on both syntactic and nonsyntactic information. In this
approach, as in the more radical interactive approach,
sentential-syntactic ambiguities are resolved by the
immediate interaction of lexical-syntactic and lexicalsemantic
information, in combination with statistical
information about the relative frequency of occurrence
of particular syntactic structures, and any available discourse
information, without appealing to an initial syntax-
based parsing stage or a separate revision stage (cf.
Tanenhaus and Trueswell, 1995).
Although we have discussed these different models in
the light of sentential-syntactic ambiguity resolution,
their architectural and processing assumptions hold for
the full domain of sentence and discourse processing.
Clearly, the representational and processing assumptions
underlying autonomous and (fully) interactive
models have very different implications for an account
of language comprehension. We will return to these issues
after giving an overview of results from the brainimaging
literature on syntactic and semantic processes
during sentence processing.
Before discussing the imaging data, a few brief comments
on the sensitivity and relevance for language research
of different brain-imaging methods are called for.
The common goal in cognitive neuroscience is to develop
a model in which the cognitive and neural approaches
are combined, providing a detailed answer to
the very general question of where and when in the
brain what happens. Methods like event-related brain
potentials (ERPs), positron emission tomography (PET),
and functional magnetic resonance imaging (fMRI) are
not equally revealing or relevant in this respect. In terms
of the temporal dynamics of comprehension, only ERPs
(and their magnetic counterparts from magnetoencephalography,
MEG) can provide the required millisecond
resolution (although recent developments in noninvasive
optical imaging indicate that near-infrared measurements
might approach millisecond resolution; cf.
Gratton, Fabiani, and Corballis, 1997). In contrast, the
main power of PET and fMRI lies in the localization of
brain areas involved in language processing (although
recent advances in neuronal source-localization procedures
with ERP measurements are making this technique
more relevant for localizational issues; cf. Kutas,
Federmeier, and Sereno, 1999). Recent analytic developments
in PET and fMRI research further indicate that
information on effective connectivity in the brain (i.e.,
the influence that one neuronal system exerts over another)
might begin to constrain our models of the language
system (cf. Büchel, Frith, and Friston, 1999;
Friston, Frith, and Frackowiak, 1993). However, localization
as such does not reveal the nature of the activated
representations: The hemodynamic response is a
quantitative measure that does not of itself deliver information
on the nature of the representations involved.
The measure is maximally informative when separate
brain loci can be linked, via appropriately constraining
experimental conditions, with separate representations
and processes. A similar situation holds for the ERP
method: The polarity and scalp topography of ERP
waveforms can, in principle, yield qualitatively different
effects for qualitatively different representations and/or
processes, but only appropriately operationalized manipulations
will make such effects interpretable (cf.
Brown and Hagoort, 1999; Osterhout and Holcomb,
1995). In short, whatever the brain-imaging technique
being used, the value of the data critically depends on its
relation to an articulated cognitive-functional model.
Cognitive neuroscience investigations
of postlexical integration
EVENT-RELATED BRAIN POTENTIAL MANIFESTATIONS
OF SENTENCE PROCESSING Space limitations rule out
an introduction on the neurophysiology and signalanalysis
techniques of event-related brain potentials (see
Picton, Lins, and Scherg, 1995, for a recent review). It is,
however, important to bear in mind that, owing to the
signal-to-noise ratio of the EEG signal, one cannot obtain
a reliable ERP waveform in a standard language experiment
without averaging over at least 20–30 different
tokens within an experimental condition. Thus, when
we speak of the ERP elicited by a particular word in a
particular condition, we mean the electrophysiological
activity averaged over different tokens of the same type.
Within the realm of sentence processing, four different
ERP profiles have been related to aspects of syntactic
and semantic processing: (1) A transient negativity
over left-anterior electrode sites (labeled the left-anterior
negativity, LAN) that develops in the period roughly
200–500 ms after word onset. The LAN has been related
not only to the activation and processing of syntactic
word-category information, but also to more general
processes of working memory. (2) A transient bilateral
negativity, labeled the N400, that develops between 200
and 600 ms after word onset; the N400 has been related
to semantic processing. (3) A transient bilateral positivity
that develops in the period between 500 and 700 ms.
Variously labeled the syntactic positive shift (SPS) or the
P600, this positivity has been related to syntactic processing.
(4) A slow positive shift over the front of the
884 LANGUAGE
head, accumulating across the span of a sentence, that
has been related to the construction of a representation
of the overall meaning of a sentence. Let us discuss each
of these ERP effects in turn.
Left-anterior negativities The LAN is a relative newcomer
to the set of language-related ERP effects. Both
its exact electrophysiological signature and its functional
nature are still under scrutiny. Some researchers
have suggested that the LAN is related to early parsing
processes, reflecting the assignment of an initial phrase
structure based on syntactic word-category information
(Friederici, 1995; Friederici, Hahne, and Mecklinger,
1996). Other researchers propose that a LAN is a reflection
of working-memory processes during language
comprehension, related to the activity of holding a
word in memory until it can be assigned its grammatical
role in a sentence (Kluender and Kutas, 1993a,b;
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