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Weiner Vol-4 c10.tex V1 - 05/18/2012 9:10pm Page 265 PART IV Attention and Action Processes Weiner Vol-4 c10.tex V1 - 05/18/2012 9:10pm Page 266 Weiner Vol-4 c10.tex V1 - 05/18/2012 9:10pm Page 267 CHAPTER 10 Selective Attention DOMINIQUE LAMY, ANDREW B. LEBER, AND HOWARD E. EGETH EARLY VERSUS LATE SELECTION 268 ATTENTION AND CONSCIOUSNESS 284 SOURCES OF ATTENTIONAL CONTROL 274 CONCLUSIONS 290 ATTENTION AND WORKING MEMORY 281 REFERENCES 290 A basic characteristic of human perception is its selective- goal-directed factors (e.g., Bundesen, 2005; Treisman & ness. At any given moment, we perceive only a fraction Sato, 1990; Wolfe, 2007). For instance, according to the of the myriad of stimuli reaching our sense organs. For biased competition model (e.g, Desimone & Duncan, example, while reading these lines, you ignore the pressure 1995) object representations compete for neural represen- of the chair on your thighs and the humming of the refrig- tation in our brains. Items that are highly salient enjoy a erator. In the words of Garner (1974, p.23), “the human competitive advantage relative to low-salience items, but organism exists in an environment containing many dif- the goals of the observer bias this competitive process, ferent sources of information. It is patently impossible for thereby ensuring that low-salience objects that are highly the organism to process all these sources, because it has relevant to the task at hand are also represented across the a limited information capacity, and the amount of infor- visual hierarchy. However, claims that attention selection mation available is always much greater than the limited might be guided exclusively by how salient an object is, capacity.” We experience these limitations on our process- irrespective of the observers’ intentions or, alternatively, ing capacity every day. Suppose two people are talking only by the goals adopted by the observers, have ignited a at once. It is easy to selectively attend to one of them. heated debate that has generated considerable research in However, it is very difficult to listen to both at once. At the past 20 years and are reviewed in the second section best, you will have to switch attention back and forth from of this chapter. speech stream to speech stream, missing chunks of one For goal-driven processing of relevant information to while attending to the other. be at all possible, representations of certain object charac- Because our information processing capacity is limited teristics, which meet the observer’s goal, must be activated (e.g., Broadbent, 1958; Neisser, 1967), we attend to some within long-term memory and maintained during the task. stimuli in our environment while ignoring others. As a This formulation of goal-directed attention highlights the result, we perceive the former while the latter gets lost. An close relationship between attention and working memory issue that has kept researchers busy for decades concerns (e.g., Awh, Vogel, & Oh, 2006; Cowan, 1995). In the third the fate of unattended stimuli: To what extent are they section of this chapter, we provide a selective review of processed? Do we register only their elementary perceptual the growing behavioral literature demonstrating interac- features such as their location, color, or shape, or do we tions between memory of spatial locations, features and also compute their meaning? The first part of this chapter objects and the deployment of attention and of neuroscien- is devoted to examining the current status of what has tific findings showing that the neural structures associated become known as the early versus late selection debate. with working memory and attention often overlap (see Another important question concerns the factors that also Nairne & Neath, this volume). determine which stimuli are selected at a given time. A The last issue covered by this chapter focuses on the re- core premise of most leading models of visual search lationship between attention and conscious awareness (see is that selection is guided by both stimulus-driven and also Banks & Farber, this volume). Although cognitive 267 Weiner Vol-4 c10.tex V1 - 05/18/2012 9:10pm Page 268 268 Attention and Action Processes psychologists have long been interested in the unconscious Renewed Support for Early Selection mind, and in particular in establishing how deeply infor- mation that is not consciously perceived can be processed Neurophysiological Studies (e.g., see Kouider & Dehaene, 2007 for a recent review), A spate of physiological studies, mainly during the 1990s, more recent research has shifted toward the study of con- has provided renewed support for early selection. In a pio- scious awareness. In particular, since Crick and Koch neering single-cell recording study, Moran and Desimone (1990) launched the search of the neural correlates of con- (1985; see also Reynolds, Chelazzi, & Desimone, 1999) sciousness, a large number of studies have contrasted con- recorded from neurons in area V4 of the monkey brain and scious and unconscious processing in order to elucidate showed that their activity could be modulated by attention. what specific mechanisms characterize conscious percep- The monkey was trained to attend to one of two objects tion. Within this framework, they have often used atten- presented simultaneously within the same receptive field tional manipulations in order to exclude parts of the visual of a V4 neuron. When the monkey attended to the inef- field from conscious awareness, implying that there might fective stimulus, the neuron’s response was weak, even be a causal relationship between attention and awareness. though the effective stimulus—which evoked a strong Here, we review the findings pertaining to the question of response by the neuron when presented alone—was also whether attention to an object is necessary and sufficient present in the cell’s receptive field. for conscious perception of this object to arise. Evidence for attentional effects in early visual areas A single chapter cannot completely cover the broad and was also obtained using various noninvasive techniques active field of attention, and we chose to focus on only four on humans. Although there was initially much debate of the issues that we deem to be particularly central in the regarding the earliest region in which spatial attention can current study of attention. Readers interested in pursuing modulate neural activity (e.g., Posner & Gilbert, 1999 for other aspects of attention may also consult a recent Annual a review), several functional magnetic imaging (fMRI) Review of Psychology on attention (Chun, Golomb, studies (e.g., Gandhi, Heeger, & Boynton, 1999) have Turk-Browne, 2011). demonstrated such modulation in area V1 (as well as in other retinotopic visual areas). Failures to find stri- EARLY VERSUS LATE SELECTION ate modulations during spatially directed attention (e.g., Mangun, Hopfinger, Kussmaul, Fletcher, & Heinze, 1997) Broadbent (1958; see also Neisser, 1967) proposed a fil- have been accounted for by the hypothesis that the target ter theory that laid the foundations of early perceptual must compete with nearby distractors in order for spatial selection theory. He suggested that there is a bottleneck attention to engage primary visual cortex (e.g., Worden & in the sequence of processing stages involved in percep- Schneider, 1996). tion. Whereas physical properties such as color or spatial However, as there are both feed-forward and feedback position can be extracted in parallel with no capacity lim- processing within the cortical hierarchy, finding attentional itations, further perceptual analysis (e.g., identification) modulation in anatomically early areas does not necessar- can be performed only on selected information. Thus, ily entail that such modulation occurs early in time. Thus, unattended stimuli, which are filtered out as a result of fMRI studies may be less appropriate for investigating the attentional selection, are not fully perceived. early versus late selection issue than tools that provide Subsequent research was soon to call filter theory into better temporal resolution, such as event-related poten- question, giving rise to the late-selection approach. One tial (ERP) recordings. This point is nicely illustrated by a striking example is the finding by Moray (1959) that study by Martinez et al. (2001) who used fMRI to localize when a message in the unattended ear is preceded by the attention-related changes in neural activity within V1 as subject’s own name, the likelihood of reporting the unat- well as within other retinotopic visual areas, while record- tended message is increased. This result suggests that the ings of ERPs traced the time course of these changes. unattended message had not been entirely excluded from These authors were able to show that stimulus-locked V1 further analysis. Thus, according to the late-selection view activity shows an early peak (50–90 ms post-stimulus) and (e.g., Deutsch & Deutsch, 1963; Duncan, 1980; Shiffrin & a later peak (160–260 ms), with only the latter being Schneider, 1977), perceptual processing operates in par- affected by spatial attention. In addition, they showed that allel and selection occurs after perceptual processing is extrastriate regions (V3, V4) show attentional modula- complete (e.g., after identification), with capacity limita- tion during a time window that occurs between the two tions arising only from later, response-related processes. phases of striate activation. These findings suggest that Weiner Vol-4 c10.tex V1 - 05/18/2012 9:10pm Page 269 Selective Attention 269 attentional modulation of V1 activity observed using fMRI presented for 200 ms. That duration is long enough for indeed reflects feedback to V1 from higher areas. They subjects to move attention from the target to one of the also indicate that the earliest modulation of neural activ- flankers, so that both target and flanker are attended. A ity by attention nonetheless occurred during the first flow shorter display followed by a mask would be required to of information through extrastriate regions, that is, before eliminate that problem.