Brain and Cognition 42, 7881 (2000) doi:10.1006/brcg.1999.1167, available online at http://www.idealibrary.

com on

The When and Where of Reading in the Brain

Sara C. Sereno
University of Glasgow

and Keith Rayner

University of Massachusetts

Skilled readers scan a text without apparent effort and, from successive glimpses, construct meaningful representations. Reading involves far more than simply identifying individual words, yet word recognition remains an indispensable component of written language comprehension and is a prudent point of entry for understanding concurrent brain activity. Within this domain, there is still much uncertainty: When in our brains are individual words recognized? Where exactly does this occur? What factors inuence the word recognition process? Recording eye movements on-line is a methodology that has succeeded in uncovering facts about the time course of language processing without imposing articial constraints on normal reading. The eye movement technique offers several advantages over traditional behavioral techniques. First, no secondary task is required of the reader. Second, it is well documented that processing a word in text is reected in its xation time (see Rayner, 1998, for a review). Readers spend less time xating words that are shorter, occur more frequently, or are more predictable in context. Finally, xation time as a response measure (about 250 ms) is much shorter than comparable lexical decision or naming times (about 500800 ms) and less contaminated by response bias. Because eye movements reect moment-to-moment cognitive processes, the average xation duration of 250 ms constrains the time for lexical processing. Measured forward from the start of a xation, it takes about 60 ms for information about the xated word to travel from the retina to higher cortical areas where lexical processing begins. Measured backward from the
Address correspondence and reprint requests to Keith Rayner, Department of Psychology, University of Massachusetts, Amherst, MA 01003. E-mail: rayner@psych.umass.edu. 78 0278-2626/00 $35.00
Copyright 2000 by Academic Press All rights of reproduction in any form reserved.

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end of a xation, oculomotor latency (the time needed to program and execute the next eye movement) requires a minimum of 100 to 150 ms. This severely limits the interval during which lexical processing can inuence when the eyes will next move. Given that xation duration varies as a function of lexical difculty, lexical processing must be sufciently advanced within the rst 100 to 150 ms of a xation in order to intelligently trigger the next eye movement. Eye-contingent techniques (making changes in a text dependent on readers eye movements) have been developed to probe the time course of processing during an eye xation. The moving window (McConkie & Rayner, 1975) and boundary (Rayner, 1975) paradigms are two such techniques used to determine the amount and type of information extracted foveally and parafoveally from a single xation. Fast priming (Sereno & Rayner, 1992) is a recently developed procedure in which a prime, presented on the order of 30 ms at the onset of a target-region xation, is then immediately replaced by a target word. Fixation time on the target as a function of preceding prime type is the dependent measure. Studies using this paradigm have revealed very early time courses for activation of orthographic, phonological, semantic, and contextual information (Lee, Rayner, & Pollatsek, 1999; Rayner, Sereno, Lesch, & Pollatsek, 1995; Sereno, 1995). In the neuroanatomical and neurophysiological domain, three decades of research on monkeys have revealed much about the 30 or so visual areas of the brain. Unfortunately, monkeys are not procient readers, so there is a lack of information about psycholinguistic behavior. Various functional brain imaging technologies have emerged that are lling this gap. Positron emission tomography (PET) and functional magnetic resonance imaging (fMRI) are based on tracking blood ow to active sites in the brain and render general pictures of localized differences of activity in the human brain. They provide no on-line time course information, nor will they ever do so because they depend on changes in blood ow. It takes seconds (aeons in psychological time) for the blood to respond to a metabolic depletion caused by neural ring. Other imaging techniques are based on measuring electrical activity generated by the brain via densely mapped sensor arrays positioned at the scalp. These include the electroencephalogram (EEG) and the magnetoencephalogram which provide a ms-by-ms record of activity. Event-related brain potentials (ERPs) are stimulus-locked averages of the EEG across many presentations of stimuli. Unlike behavioral, eye movement, or blood-ow measures, they provide a continuous ms-by-ms record of electrical changes related to on-going perceptual and cognitive processing and can thus index changes related to word recognition in real-time. Traditional ERP studies of language have typically focussed on later (endogenous) components of the waveform, such as the N400 (a negative going wave occurring about 400 ms after stimulus onset). Such ndings are discrepant with eye movement models of word recognition in the timing of lexical and se-

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mantic events. In general, these ERP results occur after the horse has left the barn. By 400 ms, the eyes will have already moved onto a following word. Thus, it seems sensible to examine the early (exogenous) components of the ERP for lexical differences. A study by Sereno, Rayner, and Posner (1998) is one of only a few to exhibit early processing differences, indicating P100 sensitivity to word form and N150 sensitivity to word frequency. In that study, identical materials were used in parallel eye movement and ERP experiments in order to establish a precise time-line of processing during a single eye xation on a word in reading. However, localization of the neural generators, solving the inverse problem (nding sources given surface recordings), is problematic (Dale & Sereno, 1993). In sum, word recognition in reading occurs incredibly fast and involves intricate neural machinery. Unhappily, there is no single technique that combines the on-line record of eye movements in a normal reading situation with the real-time ms-by-ms record provided by ERPs and the precise brain localization supplied by fMRI. By comparing data across techniques, for example, using evidence from eye movements, we do know that the later, typically larger activations in ERPs reect postlexical processing. And, while PET and fMRI have begun to chart the psycholinguistic territories of the brain, we need to be cautious in interpreting results because, without ne temporal scaling, they may only reect later activations registered in ERPs. While different imaging methodologies hint at convergence, it is important to recognize their limits (see, e.g., Posner, Abdullaev, McCandliss, & Sereno, 1999). Results from related localization and time course studies cannot simply be meshed. Knowing the time course of activation is the key in determining the sequence of processing and, hence, in providing clues to the circuitry of language comprehension.
REFERENCES
Dale, A. M., & Sereno, M. I. 1993. Improved localization of cortical activity by combining EEG and MEG with MRI cortical surface reconstruction: A linear approach. Journal of Cognitive Neuroscience, 5, 162176. Lee, Y., Rayner, K., & Pollatsek, A. 1999. The time course of phonological, semantic, and orthographic coding in reading: Evidence from the fast-priming technique. Psychonomic Bulletin & Review, 6, 624634. McConkie, G. W., & Rayner, K. 1975. The span of the effective stimulus during a xation in reading. Perception and Psychophysics, 17, 578586. Posner, M. I., Abdullaev, Y. G., McCandliss, B. D., & Sereno, S. C. 1999. Neuroanatomy, circuitry and plasticity of word reading. NeuroReport, 10, R12R23. Rayner, K. 1975. The perceptual span and peripheral cues in reading. Cognitive Psychology, 7, 6581. Rayner, K. 1998. Eye movements in reading and information processing: 20 years of research. Psychological Bulletin, 124, 372422. Rayner, K., Sereno, S. C., Lesch, M. F., & Pollatsek, A. 1995. Phonological codes are automat-

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ically activated during reading: Evidence from an eye movement priming paradigm. Psychological Science, 6, 2632. Sereno, S. C. 1995. Resolution of lexical ambiguity: Evidence from an eye movement priming paradigm. Journal of Experimental Psychology: Learning, Memory, and Cognition, 21, 582595. Sereno, S. C., & Rayner, K. 1992. Fast priming during eye xations in reading. Journal of Experimental Psychology: Human Perception and Performance, 18, 173184. Sereno, S. C., Rayner, K., & Posner, M. I. 1998. Establishing a time-line of word recognition: Evidence from eye movements and event-related potentials. NeuroReport, 9, 21952200.