Brain and Cognition 50 (2002) 95–115 www.academicpress.com Visual field asymmetries and allocation of attention in visual scenes Dell L. Rhodesa,* and Lynn C. Robertsonb,1 a Reed College, 3203 S.E. Woodstock Boulevard, Portland, OR 97202, USA b Veterans Administration Medical Research, Martinez, CA 94553, USA Accepted 25 February 2002 Abstract Single items such as objects, letters or words are often presented in the right or left visual field to examine hemispheric differences in cognitive processing. However, in everyday life, such items appear within a visual context or scene that affects how they are represented and selected for attention. Here we examine processing asymmetries for a visual target within a frame of other elements (scene). We are especially interested in whether the allocation of visual attention affects the asymmetries, and in whether attention-related asymmetries occur in scenes oriented out of alignment with the viewer. In Experiment 1, visual field asymmetries were affected by the validity of a spatial precue in an upright frame. In Experiment 2, the same pattern of asymmetries occurred within frames rotated 90° on the screen. In Experiment 3, additional sources of the spatial asymmetries were explored. We conclude that several left/ right processing asymmetries, including some associated with the deployment of spatial attention, can be organized within scenes, in the absence of differential direct access to the two hemispheres. Ó 2002 Elsevier Science (USA). All rights reserved. Keywords: Hemispheric lateralization; Visual attention; Endogenous cuing; Frames of reference; Visual scenes 1. Introduction It is not news that there is functional asymmetry within the human brain. Motorically one hand, eye or foot is preferred over the other, and cognitively the hemispheres differ in a number of ways. The latter processing differences are often described in terms such as linguistic/spatial, part/whole, time/space, or local/global. Although the nature of the cognitive processes underlying hemispheric asymmetry may be debated, it is a fact that asymmetric function exists. Most of us are righthanded, would likely become aphasic following left hemisphere damage and would likely suffer hemineglect following right hemisphere damage. The neuropsychological * Corresponding author. E-mail addresses: dell.rhodes@reed.edu (D.L. Rhodes), lynnrob@socrates.berkeley.edu (L.C. Robertson). 1 Department of Psychology, Tolman Hall, University of California, Berkeley, CA 94720, USA. 0278-2626/02/$ - see front matter Ó 2002 Elsevier Science (USA). All rights reserved. PII: S 0 2 7 8 - 2 6 2 6 ( 0 2 ) 0 0 0 1 4 - 3 96 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 literature as a whole makes it undeniable that the hemispheres process information differently. Perhaps the strength of the clinical literature was one of the reasons psychology was fast to embrace the study of hemispheric differences in normals. Studies exploiting the decussation of the optic pathways, which directs the visual input of the left and right visual fields initially to the right and left hemispheres, respectively, produced an overall picture congruent with the clinical literature. For instance, when lateralized presentations of words produced a right visual field (left hemisphere) performance advantage (Mishkin & Forgays, 1952), it was hard to resist linking this finding to language deficits observed following left hemisphere damage. Such convergence produced a plethora of studies aimed at understanding functional hemispheric differences in normal participants through controlled visual field presentations. As successful as this method has been, there remain questions regarding when an obtained visual field difference reflects a difference in entry-level hemispheric function (i.e., in input-dependent processing) and when it reflects a lateralization of more controlled processing such as executive control or allocation of attention (e.g., Bryden & Mondor, 1991). The present studies were designed to examine the role of various components of automatic and controlled visual processing in producing visual field asymmetries by manipulating spatial attention and the orientation of spatial reference frames. A spatial reference frame refers here to a global coordinate system that defines the spatial layout of local parts (Ivry & Robertson, 1998). From studies of normal perception, we know that reference frames are necessary to account for how we process stimuli in the real world (see Palmer, 1999). For example, consider a photograph of a family dinner. The spatial coordinates of the scene that includes the table, chairs and diners are defined by the orientation of the photograph, not by the coordinates of the room in which we are standing, by our own orientation, or by the orientation of individual objects within the scene.2 Instead, the global spatial frame of the photograph defines the spatial relationships and orientations of objects within that scene. We see where the family portrait is hung in relation to the landscape painting, or that the chair in the corner has been tipped over. Consistent with many formal arguments about spatial reference frames (Hinton & Parsons, 1988; Schyns & Oliva, 1994; Quinlan, 1995), as well as with common sense, we use ‘‘scenebased’’ to refer to this type of environmental, contextual reference frame. Other types of environmental reference frames, such as ‘‘object-based’’ and ‘‘environment-based’’ (gravity-bound), as well as a variety of ‘‘viewer-centered’’ frames have been invoked to account for the influence of different types of information on performance. Particularly relevant to the present experiments are studies of attention-related deficits in patients with visual neglect (often called hemineglect) resulting from unilateral brain damage. Several investigators have dissociated viewer-, gravity-, and object-centered frames of reference, and, in general, hemineglect has been found to occur within all of these frames. For instance, Calvanio, Petrone, and Levine (1987) dissociated environment- and viewer-centered reference frames by turning patients on their sides while presenting stimuli upright as usual in environmental coordinates. Under such conditions, neglect was observed in both environment- and viewer-centered frames of reference. A more recent study by Behrmann 2 However, the spatial coordinates of the scene within a photograph could be defined by the configuration of the objects within the scene, if, for example, the vertical axis of the camera was not aligned with the vertical axis of the environment when the photograph was taken and the objects in the photograph unambiguously define gravitational up/down. A ‘‘scene’’ thus established by the configuration of a group of individual objects may be similar to the type of reference frame studied, for example, by Palmer and his colleagues (e.g., Palmer, 1980). D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 97 and Tipper (1999) showed that left-sided neglect occurred both in object-based and in environment/viewer-based frames (viewers were always upright). Importantly in this study, neglect of the left side of an object (i.e., the side initially located in the neglected left visual field) rotated with the object, even when this side ended up in the patient’s right visual field. What is surprising about these findings is that a deficit resulting from damage in one hemisphere was not confined to visual information projected directly to the damaged hemisphere. In other words, even a processing deficit with an almost certain hemispheric basis was not confined to viewer-centered coordinate systems. In the present work, we demonstrate that a scene-based reference frame such as that provided by the display itself can have powerful influences on visual processing asymmetries in normal viewers. We examine the sources of the affected asymmetries by rotating the display and by using spatial precues to direct attention. The studies follow up earlier studies (Robertson, 1995; Robertson & Lamb, 1988, 1989), which showed that some visual field asymmetries found when viewers are upright shift with transformations of display orientation (although certainly not all do—see for example, Hellige, Cowin, Eng, & Sergent, 1991). Specifically, Robertson (1995) showed that participants were faster to make a reflection judgment about letters presented in the right (vs left) visual field when a display orientation congruent with viewer upright was primed. Then, when the primed display was oriented 90° orthogonal to viewer-centered coordinates (with participants themselves remaining upright), the advantage continued to occur for targets presented on the right side of the display, whether this location was in the upper or lower hemifield in viewer-centered coordinates. The rightward advantage in Robertson’s 90° displays could not be attributed to differential access of the target stimuli to one hemisphere or the other. Moreover, the size of this rightward advantage did not vary as a function of display orientation, supporting the conclusion that it was occurring nearly exclusively in the scene-based frame. In an additional experiment, Robertson showed that participants were not mentally rotating the 90 scenes back to environment/viewer upright before making the response. If anything, they were mentally rotating a viewer-centered frame into alignment with the display. The cognitive mechanisms generating the rightward advantage in Robertson’s studies were not investigated. Robertson (1995) speculated that her scene-based rightward advantage was due to a directional advantage in covertly orienting attention. There is support for the existence of one or more rightward attention-related spatial advantage(s) in the literature (Heilman, 1995; Posner & Cohen, 1984; ReuterLorenz, Drain, & Hardy-Morais, 1996; Reuter-Lorenz, Kinsbourne, & Moscovitch, 1990). Further, the studies of hemineglect patients described above demonstrate that an attention-related spatial advantage can occur in object-based as well as in viewercentered coordinates. Finally, a large number of studies demonstrate that attention can be allocated in object-based frames in normal viewers (e.g., Egly, Driver, & Rafal, 1994; Gibson & Egeth, 1994; Olson & Gettner, 1996). Thus, it seems reasonable to propose that attention-related spatial advantages can occur within scenebased frames in normal participants. Nonetheless, the allocation of attention was not explicitly manipulated in Robertson’s experiments, and the rightward advantage could instead have resulted from asymmetries in target-related perceptual processes or in post-discrimination response selection. Knowing the source(s) of spatial advantages that are not tied to viewercentered coordinates, such as those described by Robertson, can help us to decide which lateralized processes require differential direct access to the hemispheres and which do not. In the present studies, a spatial precuing method was used to investigate possible attention-related sources of the scene-based rightward advantage in the letter 98 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 reflection task obtained by Robertson. Specifically, we used a precuing procedure generally thought to affect the endogenous (voluntary) allocation of attention, i.e., a central arrow (the precue) occurred before a letter target and predicted the target location with 75% accuracy (Posner, 1980). The precue-induced focusing of attention should help to distinguish among several different possible sources of the rightward advantage. 1. If the rightward advantage is due to an automatic process that is immune to the effects of endogenous attention, it should occur in all trials, independent of precue validity (e.g., Hardyck, Chiarello, Dronkers, & Simpson, 1985). 2. If the rightward advantage is due to a global asymmetry in the activation of attentional resources, such as that accompanying differential strategic engagement of the left hemisphere (Kinsbourne, 1970), it should be eliminated altogether, or at least reduced, by precuing (e.g., Mondor & Bryden, 1992). 3. The rightward advantage could reflect an asymmetry in the ability to move attention to letter targets appearing on the right as a result of a learned strategy, such as scanning from left to right during reading (Posner & Cohen, 1984). If so, it ought to be diminished following valid precues when there is a long enough stimulus onset asynchrony (SOA) between precue and target for attention to be engaged at the precued location before the target appears. However, the rightward advantage should still be present following invalid precues, because the appearance of the target itself will serve as the ‘‘cue’’ that activates the biased strategy. 4. If the rightward advantage is due to an attention-sensitive asymmetry in the ability to perceive or process the targets, precues should have a greater influence on targets presented on the disadvantaged side. In other words, left valid precues should counteract the superiority (producing a larger benefit for validly precued left targets) and right invalid precues should exacerbate the superiority (producing a larger cost for invalidly precued left targets). Such a pattern was obtained by Nicholls and Wood (1998), who reported a larger effect of precue validity for left visual field targets in a word recognition task for which there was a rightward advantage in neutral precue trials. 5. If the rightward advantage is due to an asymmetry in the consequences of focusing endogenous attention on the left and right sides, there should be a different asymmetry in the effect of precue validity. If there is a larger facilitatory effect (benefit) of endogenous attention on the right, there might be a rightward advantage in valid precue trials compared to neutral and invalid precue trials if focusing attention in response to the precue shortens the time to maximum benefit. Such a pattern was seen, for example, in a letter localization task in Egly and Homa (1984, see their Fig. 5). If there is also a larger inhibitory effect (cost) on the right of focusing attention on the left (i.e., a bi-directional asymmetry in the effects of attention), there should be a leftward advantage in invalid precue trials, compared to both neutral and valid precue trials. Thus, the pattern of spatial advantages obtained in the presence of predictive spatial precues should help to characterize the bases of the rightward advantage found in the letter reflection task. In addition, rotation of the display will determine which components are mapped in the scene-based frame. In Experiment 1, left/right asymmetries were examined with the display aligned with viewer/environment upright to determine the effects of endogenous spatial precues at short and long SOAs. In Experiment 2, we showed that the pattern of asymmetries observed in Experiment 1 occurred in scenes visibly rotated through 90° on the screen (before the precuing procedure), as predicted by Robertson’s (1995) results with primed scene orientations. Finally, in Experiment 3, a more detailed examination of possible sources of the scene-based spatial advantages was undertaken. D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 99 2. Methods: general The primary task adopted in the present experiments was the letter reflection task employed by Robertson (1995). Studies of mental rotation suggest that object representations rarely specify the object’s left/right sides (Hinton & Parsons, 1981; McMullen & Jolicoeur, 1990; Rollenhagen & Olson, 2000; Tarr & Pinker, 1990), so most handedness judgment tasks require rotation of the object into viewer-centered coordinates. However, several investigations have shown that an informative contextual frame can also be used to determine the left/right sides (Corballis, Nagourney, Shetzer, & Stefanatos, 1978; Hinton & Parsons, 1981). Thus, a handedness judgment can be used to induce reliance on a scene for the purposes of comparing the use of this type of environmental frame with that of the viewer-centered frame. In the present experiments, three letter targets were used: E, K, and G. Because two of these letters (E and K) are symmetrical around their horizontal axes, participants were required to use their knowledge of the scene orientation to do the letter reflection task in Experiment 2, when frames rotated to orientations 90° with respect to the viewer. However, previous studies (Robertson & Lamb, 1989) found no difference in response times between symmetric and asymmetric letters as a function of side or of frame orientation, and Robertson (1995) obtained a strong scene-based rightward advantage using only asymmetric letters. So, there’s no reason to expect that the effects we obtained in these studies depend on the specific letters used. Participants reported the reflection of the target letter by pressing left and right response keys with two fingers (index and middle) of one hand. Differential hemispheric activation associated with the use of one hand to respond could influence the rightward advantage. Thus, half the participants in each study used their right hand, the other half used their left hand, and response hand was included as a between subjects variable in the data analyses. For all participants, however, the right key press indicated a ‘‘normally-oriented’’ (henceforth: normal) target letter and the left key press indicated a ‘‘mirror-reflected’’ (henceforth: reflected) target letter. This mapping seemed much more natural to participants, presumably because the informative side of a normal letter is on its right and the informative side of a reflected letter is on its left.3 It did, however, set up the possibility that we would obtain a stimulus/response compatibility effect known as the Simon Effect (normal letters on the right side would be responded to faster than reflected letters on the right side and than normal letters on the left side). This possibility is examined in Experiment 3. Spatial precuing was accomplished using a standard Posner-type endogenous precuing paradigm (Posner, 1980). Specifically, a centrally located arrow indicated the side of the upcoming target with 75% accuracy. In Experiments 1 and 2, the arrow formed the arms of a stick figure with head and feet, to emphasize the orientation of the frame. In Experiment 3, more traditional arrow cues were used. In each experiment, two or three SOAs were used for three reasons: (1) to reduce the temporal predictability of target onset, (2) to allow us to confirm that we were obtaining the normal SOA-dependent validity effects with our precuing procedure, 3 This mapping could produce a response hand target side interaction, as the index finger might be faster than the middle finger. This pattern was in fact obtained in Experiment 2 (p < :05). However, for the right hand, the index finger signaled a reflected letter, whereas for the left hand, the index finger signaled a normal letter. Thus, this mapping should help to prevent asymmetries in stimulus/response compatibility effects. Furthermore, in Experiment 1, the response hand target side interaction was in the opposite direction, reflecting a weak hand-based spatial compatibility effect (p < :10), and in Experiment 3, the response hand target side interaction was not significant (p ¼ :61). Note also that because the informative sides of normal and reflected letters occur closer to and farther from fixation, depending on target side, possible effects of stimulus feature eccentricity on each target side should be controlled for. 100 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 and (3) because some sources of the spatial asymmetries were predicted to vary with SOA. Head position was constrained in Experiment 2, when display orientation was dissociated from viewer/environment upright. In all Experiments, the instructions emphasized the importance of maintaining fixation at the center of the display. Although eye movements were not monitored, it is unlikely that our findings are due to eye movements.4 Although most studies of hemispheric laterality employ brief stimulus presentations to insure that participants do not fixate the target, in these studies both the precue and the target remained on the screen until the participant responded. We adopted longer stimulus presentation times for two reasons: (1) precue disappearance might have increased the likelihood of eye movements from central fixation, and (2) briefly presented targets can produce across-trial inhibition of return (Klein, 2000). The ways in which these procedures may have affected our findings are addressed in Section 6. 3. Experiment 1: Spatial asymmetries in upright scenes 3.1. Participants Six men and 10 women, aged 18–47 (mean ¼ 23) participated. All but one were naive to the goals of the study. Participants were right-handed and had normal or corrected-to-normal vision. All gave written consent and were paid for their participation. 3.2. Stimuli and apparatus Stimuli were white on a black background. Each trial began with a 500 ms fixation display of a centered ‘‘A’’, 1.5° high. The ‘‘A’’ was replaced by ‘‘arrowperson,’’ a stick figure with a head, two legs, and an arrow for arms. The arrow precue predicted the upcoming target side with 75% accuracy. There were no target location markers. Following randomized SOAs of 150, 400, or 650 ms, a 1.5° target letter (E, K, or G) appeared 6° to the left or right of fixation. The target letter was normally oriented (with respect to the computer screen/display) or mirror-reflected around its vertical axis. Arrowperson and the letter remained on the screen for 1000 ms. Incorrect responses were followed by auditory feedback. Intertrial intervals were 517 ms. 3.3. Procedure Participants were seated 54 cm from the computer in low level illumination. Sample trials were provided, and the following aspects of the task emphasized: 1. fixate the center of the screen, i.e., look at the fixation ‘‘A’’ and then arrowperson, without moving your eyes, throughout each trial; 2. the arrow predicts the target location; 4 We have obtained spatial asymmetries of comparable directionality and magnitude in experiments in which eye movements have been monitored (Rhodes & Montgomery, 2000). It is also unlikely that the asymmetries we report here were due to eye movements to the target locations. Eye movement asymmetries cannot easily explain the dissociation from viewer-centered coordinates obtained in Experiment 2 and in Robertson (1995). Additionally, the target stimuli were presented for only 100 ms in Robertson’s studies, and the rightward advantage was about the same size as those obtained in the present studies. D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 101 3. report whether the target letter is normally oriented or mirror-reflected (backward) by pressing the right or left response buttons, respectively; 4. respond as rapidly as possible while not making many errors. There were one block of 24 practice trials and three blocks of 144 trials each. 3.4. Data analyses Reaction times (RTs) were not included if the response was incorrect, or if the RT was longer than the display duration or more than three standard deviations from the participant’s mean. Mean RTs were calculated collapsed over letter identity and letter reflection, as there were not enough invalid precue trials (18/cell combined over letter identity and letter reflection) to support inclusion of these variables in the overall analysis. Mean percent errors (incorrect responses/total responses) were also calculated. Mean RTs and percent errors were analyzed in separate 4-factor mixed effects ANOVAs with responding hand as a between groups variable, and precue validity, target side, and SOA as within subjects variables. 3.5. Results The mean RT over all conditions was 563 ms. Precue validity and SOA effects conformed to those normally obtained in an endogenous spatial precuing task. Responses were 25 ms faster to validly than invalidly precued targets, F ð1; 14Þ ¼ 25:9, p < :001, and RTs were inversely related to SOA, F ð2; 28Þ ¼ 21:2, p < :001. Responses were faster to validly than to invalidly precued targets at all SOAs, but the difference was largest at the 650 ms SOA (validity SOA interaction, F ð2; 28Þ ¼ 5:3, p ¼ :01Þ. The main effect of target side was not significant, F ð1; 14Þ ¼ :6, but the target side precue validity interaction was highly reliable, F ð1; 14Þ ¼ 34:6, p < :001. For validly precued targets, responses were 21 ms faster to right than to left targets, tð15Þ ¼ 2:6, p < :05. For invalidly precued targets, this spatial advantage was reversed, but the leftward advantage was only 8 ms, tð15Þ ¼ 1:0, n.s. As can be seen in Fig. 1, the pattern of spatial asymmetries was similar at all SOAs (target side precue validity SOA interaction, n.s.). On valid precue trials there was a right side advantage at all SOAs; on invalid precue trials, the mean RT advantage either reversed to a left side advantage or was absent. The pattern of spatial asymmetries was weakly related to responding hand (p < :10).5 The mean error rate was low (3.8%). In general, longer RTs were accompanied by higher error rates, arguing against a speed-accuracy trade-off. Errors did not vary significantly as a function of precue validity, target side, or precue validity target side (valid right 2.9%, valid left 3.8%, invalid right 3.9%, invalid left 4.9%). 3.6. Discussion The rightward spatial advantage observed by Robertson (1995) appeared only for validly precued targets in this study. For invalidly precued targets, the spatial advantage was leftward overall. This pattern of spatial asymmetries is not compatible with some of the options described in Section 1. First, because the rightward advantage was not eliminated by valid precues, it is unlikely to be due to a global 5 The rightward advantage in valid precue trials was due primarily to participants using their right hand to respond, and the leftward advantage in invalid cue trials was due primarily to participants using their left hand to respond. However, this pattern was not replicated in Experiments 2 and 3. 102 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 Fig. 1. Mean reaction times to report letter reflection as a function of target side (right/left) and cue validity (valid/invalid) at each SOA (150/400/650 ms) in Experiment 1 with all frames of reference upright. asymmetry in the activation of attentional resources, such as that accompanying differential strategic engagement of the left hemisphere (option #2; Kinsbourne, 1970). Second, the obtained pattern was the converse of that predicted by hypotheses of more rapid orienting of attention to targets on the right side (option #3; Posner & Cohen, 1984). Finally, the most consistent effect that can be seen in Fig. 1 is a larger difference between valid and invalid precue trials for targets on the right (40 ms) than for those on the left (11 ms). Theories suggesting that there are more attentional resources available for targets on the right side would predict a larger effect of precues on the left (option #4; Nicholls & Wood, 1998). The larger effect of precue validity on the right could reflect a larger benefit of right valid precues, a larger cost of left invalid precues (i.e., a larger cost for right targets appearing after left precues), or both (option #5). However, the rightward advantage in valid precue trials in the present study is of approximately the same magnitude as the rightward advantage obtained by Robertson (1995), who used no predictive precues. It thus seems likely that there is a rightward advantage that occurs whenever a target appears on the right, that is counteracted in invalid precue trials by a leftward advantage associated with a larger cost of invalid left precues. Whether the rightward advantage is attention-insensitive (option #1) or is instead a consequence of focusing attention on a right target (a version of option #5) cannot be determined from these data. Two other explanations of the findings in Experiment 1 are possible. First, rightward precues might produce an RT advantage for whatever target follows them.6 Such an advantage could be due, for example, to greater arousal following right precues (or, in the absence of precues, greater arousal in response to right targets). Second, there could be an interaction between the direction of cued attentional shifts and the relative strengths of stimulus–response spatial compatibility effects on the right and left sides of the display. For example, responses to normal letters could be especially fast on the right following valid (right) precues, whereas responses to reflected letters might be especially slow on the right following invalid (left) precues. These alternative explanations of the precue validity target side interaction will be explored in more detail in Experiment 3. Before considering these possibilities further, we first establish that the pattern of asymmetries observed in Experiment 1 occur following rotation of the scene out of alignment with viewer upright. 6 We thank Ray Klein for this suggestion. D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 103 4. Experiment 2: Spatial asymmetries in rotated frames Only the methods differing from those of Experiment 1 are described below. 4.1. Participants Nine men and seven women, aged 17–33 (mean ¼ 22.5) participated. All were naive to the goals of the study. 4.2. Stimuli and apparatus Fig. 2 shows the stimulus displays used in this experiment. In addition to the ‘‘A’’ at fixation, two vertical columns of ‘‘A’’s and ‘‘V’’s located 6.2° right and left of center emphasized the scene orientation. After 500 ms, the display equiprobably rotated around its center 90° clockwise (+90, top at right), rotated around its center 90° counterclockwise ()90, top at left), or remained upright. When rotation of the display occurred, it took about 230 ms and was perceptually smooth. Five hundred ms following rotation (or 730 ms after onset for upright displays), the fixation A was replaced by arrowperson, oriented upright in the scene. Either 150 or 650 ms after arrowperson appeared, the target letter appeared 4° to the left or right of arrowperson, i.e., in the upper or lower visual fields in the 90° displays. The top of the target letter was always oriented in the same direction as the top of the display frame and arrowperson’s head. Arrowperson and the letter remained on the screen for 1500 ms or until the participant responded. Intertrial intervals were 548 ms. 4.3. Procedure Participants’ heads were held upright by a chin rest, forehead bar, and side rings. Instructions emphasized that the reflection of the letter must be determined with Fig. 2. The stimulus displays in Experiment 2 for a +90° frame, validly cued, left target side, and normal target orientation trial. Drawing not accurately scaled. 104 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 respect to the orientation of the display and that fixation should be maintained at the center throughout each trial. There were one block of 72 practice trials and six blocks of 144 trials each. 4.4. Data analyses Mean RTs and percent errors were analyzed in separate 5-factor mixed effects ANOVAs with responding hand as a between groups variable, and frame orientation (upright/+90/)90), precue validity, target side (in scene-based coordinates) and SOA as within subjects variables. 4.5. Results Overall RTs in the upright frame were comparable to those in Experiment 1 (572 ms). RTs were 30 ms longer in rotated than upright frames, F ð2; 28Þ ¼ 12:4, p < :001. This was expected, because in rotated frames the viewer/environment and scene-based frames were in conflict. RTs indicated successful precuing of spatial attention. Responses were 28 ms faster to validly than invalidly precued targets, F ð1; 14Þ ¼ 32:6, p < :001, and were faster at the long than short SOA, F ð1; 14Þ ¼ 38:4, p < :001. The validity effect (invalid–valid precue trial RTs) was significant at both SOAs, but was larger at the 650 ms SOA (precue validity SOA interaction, F ð1; 14Þ ¼ 14:0, p < :005Þ. As in Experiment 1, the main effect of target side (left/right in scene-based coordinates) was not significant, F ð1; 14Þ ¼ :2, but the target side precue validity interaction was highly reliable, F ð1; 14Þ ¼ 31:9, p < :001. The scene-based spatial asymmetries were very similar to those in Experiment 1. In valid precue trials, responses were 25 ms faster to right than left targets, tð15Þ ¼ 3:9, p < :005. For invalid precue trials, responses were 13 ms faster to left than right targets, tð15Þ ¼ 2:2, p < :05. Furthermore, these asymmetries were independent of frame orientation averaged over SOA and of SOA averaged over frame orientation. However, there was also a rightward advantage that was independent of precue validity at the short SOA in this experiment, target side SOA, F ð1; 14Þ ¼ 13:5, p < :005. Averaged over precue validity and frame, responses were faster to targets on the right than left side at the short SOA (right ¼ 605 ms, left ¼ 620 ms), F ð1; 15Þ ¼ 2:5, p < :05, but not at the long SOA (right ¼ 573 ms, left ¼ 570 ms). Because a rightward advantage occurred in the absence of precues in Robertson (1995), this precue validity-independent rightward advantage will be referred to as a precue-independent rightward advantage. The asymmetries that depend on precue validity (i.e., that are precue-dependent) will be referred to as cue-validity dependent spatial advantages. The only influence of frame orientation also occurred at the short SOA, frame target side precue validity SOA, F ð2; 28Þ ¼ 3:6, p < :05. The target side precue validity RTs are plotted separately for the 150 and 650 SOAs for each frame orientation in Fig. 3. At long SOAs, the target side precue validity interactions were remarkably similar across frame orientations, frame target side precue validity interaction, F < 1. In each frame orientation, RTs were faster to validly precued right than left targets and slower to invalidly precued right than left targets. At the short SOA, however, the precue-independent rightward advantage enhanced the rightward advantage for targets on valid precue trials (i.e., the rightward advantage in valid precue trials was larger at the short SOA than at the long SOA). It also reduced the leftward advantage for invalid precue trials, but only in the upright and +90 frames (frame target side precue validity interaction, F ð2; 28Þ ¼ 3:9, p < :05 at the short SOA). D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 105 Fig. 3. Mean reaction times to report letter reflection as a function of target side and cue validity at each SOA (150/650 ms) and in each frame orientation (upright, rotated þ90°, rotated )90°) in Experiment 2. Target side is identified in scene-based coordinates. For example, the ‘‘right target’’ in the +90° frame was in the participants’ lower visual field and in the )90° frame it was in the upper visual field. The mean error rate was low (2.8%). As in Experiment 1, shorter RTs were generally associated with fewer errors. Error rates were lower in the upright frame (1.6%) than in the rotated frames (+90: 3.2%, )90: 3.5%), F ð2; 28Þ ¼ 8:32, p < :005. The main effects of precue validity and target side and the precue validity target side interaction were not significant (valid right 2.0%, valid left 2.8%, invalid right 3.1%, invalid left 3.1%), nor did the effects of these variables interact with frame orientation. 4.6. Discussion The pattern of cue validity-dependent spatial asymmetries obtained in this experiment was very similar to that obtained in Experiment 1. There was a consistent rightward RT advantage for target discrimination in valid precue trials. Although less consistent than the rightward advantage, there was a leftward advantage for target discrimination in invalid precue trials. The overall effect of precue validity (benefit of valid precues plus cost of invalid precues) was larger and more consistent for right targets than for left targets across both SOAs and all frame orientations. The mechanisms underlying these asymmetries were clearly organized within scene-based coordinates. If the left/right asymmetries included components mapped in viewer/environment coordinates, at the very least the target side precue validity interaction would have been reduced in the rotated scenes. Further, if an additional viewer/environment asymmetry (such as an upward advantage) had been exposed, a new set of frame orientation dependencies would have appeared. The one difference in the asymmetries obtained in Experiments 1 and 2 was the appearance of a precue-independent rightward advantage at the short SOA in Experiment 2. The leftward advantage for invalidly precued targets was numerically larger at the short than at the long SOA in Experiment 1, but was larger at the long than at the short SOA in Experiment 2. These findings at least tentatively suggest 106 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 that a precue-independent rightward advantage is replaced over time by cue-validity dependent asymmetries. In Experiment 1, the ‘‘short’’ SOA might not have been short enough to reveal the precue-independent advantage. In Mondor & Bryden’s (1992) experiments, for example, a rightward advantage in letter identification was overcome by predictive peripheral precues at SOAs as short as 67 ms. In Experiment 2, the addition of rotation to the displays may have added temporal and/or spatial uncertainties that extended the time course of events, such that the precue-independent rightward advantage was still strong at the 150 ms SOA. A precue-independent rightward advantage at short but not long SOAs could be explained by mechanisms such as a rightward advantage in the speed of orienting attention (option #3 in Section 1). As in Experiment 1, the cue validity-dependent spatial asymmetries could reflect larger effects of attention (facilitation or benefits and/or inhibition or costs) on the right side of the scene. In other words, the rightward advantage could reflect an enhanced facilitatory effect of attention directed to the right on valid precue trials and/or an enhanced inhibitory effect on the right when attention was directed to the left by the precue. A model that could account for both of these effects is a bi-directional gradient model that operates both to facilitate an attended location and inhibit an unattended location (option #5; e.g., Handy, Kingstone, & Mangun, 1996). To assess the cost/benefit predictions of this model and to compare it with the alternative explanation that right precues speeded RTs overall, in Experiment 3 we added neutral precue trials to a version of Experiment 1. The neutral precue trials allowed us to evaluate costs and benefits of cuing independently. In addition, we increased the number of trials to assess stimulus/response spatial compatibility effects (often called the Simon Effect). Hommel & Lippa (1995) demonstrated a Simon Effect within rotated visual contexts, so an examination of this spatial asymmetry is interesting both in its own right and as a possible source of the scene-based spatial asymmetries. 5. Experiment 3: Cost/benefit analysis Only the methods differing from those of Experiment 1 are described below. 5.1. Participants Eight men and nine women, aged 18–23 (mean ¼ 20.3) participated in Experiment 3. Three female participants were eliminated from the data analyses because the number of acceptable trials in one or more cell(s) of the design fell below 80% (see Section 5.4). Of the remaining participants, seven used their right hand and seven used their left hand to respond. 5.2. Stimuli and apparatus Each trial began with a 525 ms fixation display of a centered ‘‘A’’, 1° high. After 60 ms, the ‘‘A’’ was replaced by a diamond of the same height. In 80% of the trials, the two right or the two left lines of the diamond were white, forming an arrow that predicted the target location with 75% accuracy. The two lines forming the opposite side of the diamond were dark gray. In the remaining 20% of the trials, the entire diamond was light gray (neutral precue). Following randomized SOAs of 150 or 650 ms, the target appeared 5° to the left or right of the precue. Intertrial intervals were 555 ms. There was no feedback for incorrect responses. D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 107 5.3. Procedure The experiment was run at Reed College on a Macintosh rather than on the PCbased computer system used in the previous experiments. Participants reported whether the target was normal or reflected by pressing the K or J keys, respectively, on a computer keyboard. There were one block of 36 practice trials and eight blocks of 120 trials each. 5.4. Data analyses Trials were eliminated if the response was incorrect (2.1%), or if the RT was longer than 1000 ms (0–3% of all trials, mean ¼ .7%). Participants were excluded from analysis if the percentage of trials in any cell of the design fell below 80% (<20/ 24 for invalid and neutral precue trials, <58/72 for valid precue trials). Mean RTs and mean percent errors (incorrect responses/total responses) were calculated collapsed over letter identity and analyzed in separate 5-factor mixed effects ANOVAs with responding hand as a between groups variable, and precue validity, target location, SOA, and letter reflection as within subjects variables. The first analysis included only validly and invalidly precued trials, to confirm the effects obtained in Experiments 1 and 2. The second analysis included all three precue validity conditions, and separate analyses of the benefits (valid/neutral) and costs (neutral/invalid) of precuing were performed where indicated. There were no significant effects of responding hand in these analyses. 5.5. Results The mean RT over all conditions was 517 ms. Fig. 4 shows the RTs to targets on each side following valid, neutral and invalid precues at short and long SOAs, collapsed over reflection. Fig. 5 displays the RTs to normal and reflected target letters separately. Percent errors are shown above each bar. Valid/invalid trials. RTs to validly precued targets were 21 ms faster than to invalidly precued targets, F ð1; 12Þ ¼ 43:4, p < :001, comparable to the effects obtained Fig. 4. Mean reaction times to report letter reflection as a function of target side (right/left) and cue validity (valid/neutral/invalid) at each SOA (150/650 ms) in Experiment 3 with all frames of reference upright. 108 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 Fig. 5. Mean reaction times to report letter reflection as a function of target side and cue validity at each SOA (150/650 ms) for normal and reflected letters in Experiment 3. Numbers above each bar are mean percent errors. in Experiment 1 (25 ms) and Experiment 2 (28 ms). As expected, the validity effect was larger at the long SOA, F ð1; 12Þ ¼ 9:7, p < :01. RTs to targets on the right side were 10 ms faster overall. Although this difference was only 3 ms larger than in Experiments 1 and 2, it was more consistent, leading to a reliable main effect of target side, F ð1; 12Þ ¼ 15:6, p < :005. Most importantly, the target side precue validity effect resembled that obtained in the previous experiments, F ð1; 12Þ ¼ 14:0, p < :005. Specifically, there was a 16 ms rightward advantage for valid precue trials, tð13Þ ¼ 15:7, p < :001, but only a 4 ms rightward advantage for invalid precue trials (n.s.). This pattern of asymmetries was independent of SOA. Valid/neutral/invalid trials. Confirming the existence of a precue-independent rightward advantage, there was an 18 ms rightward advantage for neutral precue trials when analyzed alone, tð13Þ ¼ 17:6, p < :001. Costs and benefits were evaluated relative to this neutral condition. The main effect of precue validity, F ð2; 24Þ ¼ 27:6, p < :001 consisted of a 7 ms benefit, F ð1; 12Þ ¼ 5:8, p < :05 and a 15 ms cost, F ð1; 12Þ ¼ 27:5, p < :001. The benefit of valid precues was approximately the same for targets on the right (6 ms) and left (8 ms) sides, while the cost of invalid precues was much larger for targets on the right (22 ms) than on the left (8 ms), precue validity target side, F ð1; 12Þ ¼ 15:3, p < :005. The precue validity target side interactions for benefits and costs did not vary with SOA. Thus, the asymmetrical validity effects obtained in Experiments 1 and 2 were due entirely to a larger cost of invalid left precues (i.e., longer RTs to right targets following left precues). Not surprisingly, RTs to normal letters were 20 ms faster than to reflected letters overall, F ð1; 12Þ ¼ 7:7, p < :05. An important question is whether an asymmetry in D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 109 stimulus–response spatial compatibilities (i.e., the Simon Effect) can account for either the precue-independent or the cue validity-dependent spatial asymmetries. A Simon Effect should produce a significant target side letter reflection interaction, in which right keypresses to normal letters are faster when the letters appear on the right side and left keypresses to reflected letters are faster when the letters appear on the left side of the display. This interaction was not significant overall, but the SOA target side letter reflection interaction was highly reliable, F ð1; 12Þ ¼ 20:0, p < :001. At the short SOA, there was no evidence for a Simon Effect. There was a rightward advantage for both normal (12 ms) and reflected (16 ms) letters (target side letter orientation n.s.). These data make it particularly clear that the precueindependent rightward advantage was not due to an asymmetry in the Simon Effect. At the long SOA, there was a Simon Effect, target side letter reflection, F ð1; 24Þ ¼ 5:5, p < :05. The rightward advantage increased to 25 ms for normal letters, and reversed to a 3 ms leftward advantage for reflected letters. Thus, the precue-independent rightward advantage seen at the short SOA for both normal and reflected letters was symmetrically modulated by the Simon Effect at the long SOA, confirming that the precue-independent rightward advantage did not result from an asymmetry in the Simon Effect. Perhaps more important is the possibility that an interaction between the precuing procedure and the response mapping used in these studies could account for the precue validity target side interaction. Critically, the precue validity target side letter reflection interaction was not significant (p ¼ :38). However, consistent with the presence of a Simon Effect only at the long SOA, the precue validity target side letter reflection SOA interaction was significant, F ð2; 24Þ ¼ 5:5, p ¼ :01. This interaction was complicated, but the data in Fig. 5 make it reasonably clear that the longer RTs for invalidly precued right targets was not an artifact of the Simon Effect, as it occurred for normal letters at the short SOA as well as for reflected letters at the long SOA.7 The mean error rate was low (2.1%). The main effects of precue validity and target side and the precue validity target side interaction were not significant. However, the 4-way interaction (SOA precue validity target side letter reflection) was significant, F ð2; 24Þ ¼ 7:6, p < :005. As can be seen in Fig. 5, response accuracy varied across SOA especially for invalidly precued normal letters on the right, for which error rates were high at the short SOA and low at the long SOA. At the short SOA, errors were lowest for validly precued normal letters on the right side and highest for validly precued normal letters on the left side. At the long SOA, errors were lowest for validly precued reflected letters on the left, and were highest for invalidly precued normal letters on the left. Thus, the interaction reflects precue validity and SOA modulation of the Simon Effect, which just barely achieved significance averaged over SOA and precue validity, target side letter reflection, F ð1; 11Þ ¼ 4:8, p ¼ :05. 7 At the short SOA, costs and benefits of precuing were virtually identical for reflected targets on the right and left sides (validity main effect F ð2; 24Þ ¼ 17:1, p < :001, target side precue validity, n.s.). For the normal letters, however, there were no effects of precue validity on the left side, and a large cost of invalid precuing on the right (validity effect n.s., target side precue validity, F ð2; 24Þ ¼ 4:1, p < :05). At the long SOA, it was the normal letters that showed identical costs and benefits of precuing on the right and left sides (validity main effect, F ð2; 24Þ ¼ 29:3, p < :001, target side precue validity n.s.). For the reflected letters, one can see both the influence of the Simon Effect (rightward advantage reduced for neutral precue trials and reversed for valid precue trials compared to the rightward advantage for normal letters) and the exaggerated cost of invalid precuing of right targets (27 ms cost for right reflected letters, no cost for left reflected letters, target side precue validity, F ð2; 24Þ ¼ 6:1, p < :01). The detailed sources of this interaction are not understood. 110 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 5.6. Discussion Analysis of the valid and invalid precue trials in Experiment 3 revealed the same target side precue validity interaction observed in Experiments 1 and 2, i.e., a rightward advantage in valid precue trials that was reduced or eliminated in invalid precue trials (Fig. 4). The cost/benefit analysis afforded by the addition of neutral precue trials revealed that the benefit of valid precues (neutral—valid precue trial RTs) was about the same for right and left targets. Confirming the findings of Robertson (1995), who used no spatial precues, there was also a substantial rightward advantage following neutral precues (18 ms). Further, a smaller (10 ms), but significant, rightward advantage that was independent of precue validity was obtained in this study. Thus, there was strong evidence for a precue-independent rightward advantage that might reflect either an attention-insensitive mechanism (option #1 in Section 1) or an attention-related asymmetry engaged by all targets appearing on the right side (a version of option #5). These alternatives will be discussed in more detail in Section 6. Given that there was no spatial asymmetry in the benefit of valid precuing, the target side precue validity interaction was due entirely to an asymmetrical cost of invalid precues (invalid-neutral precue trial RTs). The cost of invalid precues was much larger for targets on the right than on the left, and this effect was present at both short and long SOAs. In Experiment 2, the rightward advantage in valid precue trials and the asymmetrical validity effect in upright displays was essentially unchanged in displays rotated 90° out of alignment with the viewer-centered frame. Thus, both the precueindependent rightward advantage and the larger inhibitory effect of invalid left precues on right targets were programmed in scene-based coordinates. Neither of these effects could be due to differential direct access of the target stimuli to the two hemispheres. There was no evidence of a Simon Effect at the short SOA in this study, but an appropriate target side letter reflection interaction occurred at the long SOA. Because both the precue-independent and cue-validity dependent spatial asymmetries were obtained at both short and long SOAs, it is unlikely that they result from asymmetrical stimulus/response compatibility effects. Mean RTs for normal and reflected letters in the invalid precue conditions of Experiments 1 and 2 must be calculated from a very small number of trials (9/cell), but the pattern of effects was quite similar to that seen in Experiment 3. In summary, the results of Experiment 3 suggest that there are three spatial asymmetries affecting performance in the task used here. (1) A rightward advantage that occurs when there is no directional precue and that is independent of precue validity when precues are provided. (2) Greater inhibition of the right than of the left side of the display by invalid spatial precues, revealed in greater costs for targets appearing on the right after left precues than for targets appearing on the left after right precues. (3) A Simon Effect evident only at longer SOAs. Most importantly, it seems likely that all three of these asymmetries occur in scene-based coordinates. 6. General discussion When participants discriminated the reflection of a target letter whose location was predicted by a central spatial precue, we obtained a pattern of spatial asymmetries that included a larger precue validity effect (invalid–valid precue trial RTs) on the right than the left side of the display. This pattern occurred at all SOAs across three experiments. Importantly, in Experiment 2 the pattern was unaltered when right and left target locations were defined within displays that had been rotated D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 111 90° out of alignment with the viewer and the environment. This is strong evidence that the spatial asymmetries generated by the cue-induced allocation of spatial attention occurred in scene-based coordinates and not in viewer- or environmentbased coordinates. Indeed, there was no evidence for a viewer-centered component, and thus no evidence that the cue validity-dependent spatial asymmetries were due to differential direct access of visual information to one or the other hemisphere. The cost/benefit analysis of Experiment 3 indicated that the cue validity-dependent component of the pattern of asymmetries was due specifically to a larger cost of invalid left precues. This effect is not easily explained by existing theories of attention-related asymmetries, such as those outlined as options #2–4 in the Introduction. However, it is consistent with a gradient theory of attention, in which the focusing of attention facilitates processing at the attended location and inhibits processing at non-attended locations (option #5 in Section 1; e.g., Handy et al., 1996). In its simplest form, an attentional gradient model might predict that both the facilitatory and the inhibitory effects of precuing attention would be amplified on the right side of a frame. In this case, one would expect to see a rightward advantage in RTs to validly precued targets and a leftward advantage in RTs to invalidly precued targets (because the inhibition of left targets by right invalid precues would be less than the inhibition of right targets by left invalid precues). This is close to the pattern we obtained, with one important exception—the rightward advantage we obtained was not dependent on precue validity. Although we obtained a rightward advantage in valid precue trials, it was not consistently larger than the rightward advantage obtained in neutral precue trials (Experiment 3) and in no precue trials (Robertson, 1995). Further, there was no asymmetry in the benefit of valid precues. And, finally, although the leftward advantage in invalid precue trials was consistently present in comparison to the rightward advantage in valid precue and neutral precue trials, it appeared to occur ‘‘on top of’’ a rightward advantage. As a result, the leftward advantage was not always apparent numerically, and, in some cases (Experiment 3 and short SOA in Experiment 2), a rightward advantage that was independent of precue validity (main effect of target side) appeared in the statistical analyses. Thus, it is not clear that the rightward advantage obtained in valid and neutral precue trials in this study, and in Robertson’s (1995) no precue trials, is related to an asymmetry in the effects of attention. It could, instead, be due to an attention-insensitive process (option #1 in the Introduction). However, an attention-related explanation seems more likely. One reason is that the benefits of precuing in Experiment 3 were numerically rather small (7 ms). This may have been because the effect of the valid precues was ‘‘masked’’ by a comparable process occurring in the neutral precue trials. Neutral precue trials (and no precue trials) can be thought of as trials in which the appearance of the target itself serves as the ‘‘cue’’ that initiates the movement and subsequent focusing of attention. If focusing attention on the right side of a scene facilitates processing more than focusing attention on the left side, this ‘‘benefit’’ should occur in neutral precue trials (and in no precue trials) as well as in trials with valid precues, and it does. The prediction that there ought to be a rightward advantage in only valid precue trials depends upon an assumption that precues speed up the benefit of focusing attention. In addition, the attention-insensitive option seems unlikely. There is little reason to expect stimulus-related or task-related processing asymmetries in this task, in which the target stimuli differed only in their reflection around their vertical axes. Further, an attention-insensitive account for the cue-independent rightward advantage would predict its disappearance in rotated displays as was observed by Hellige et al. (1991) in studies of CVC nonsense syllable identification. Instead, the precue-independent rightward advantage occurred in scene-based coordinates, as it appeared across frame orientations at the short SOA in Experiment 2, and in noprecue trials across frame orientations in Robertson’s experiments. 112 D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 Further, we obtained no evidence to suggest that response processes produced the precue-independent rightward advantage. In general, responding hand did not affect the asymmetries we obtained, and when it did, it did not do so in a consistent way across experiments3 . Our examination of stimulus/response compatibility effects (i.e., the Simon Effect) in Experiment 3 provided no evidence that an asymmetry in these effects could account for either the cue-validity dependent or the precue-independent spatial advantages. Thus, an especially appealing explanation of the spatial advantages obtained in these experiments is that both the facilitatory effect of focusing attention on an upcoming target location and the inhibitory effect of focusing attention on a contralateral (invalidly cued) location were larger on the right side of a scene than on the left. The inhibitory component was activated by an invalid precue. The facilitatory component, however, was activated whenever the target stimulus was attended, i.e., following a valid precue, but also following invalid and neutral precues. This explanation conforms to bi-directional gradient theories of the effects of attention (Handy et al., 1996), and attributes both the cue validity-dependent and the precueindependent spatial advantages obtained in these experiments to processes associated with the allocation of attention within scenes. It also suggests that the deployment of attention in environmental or scene-based spatial reference frames may account for some of the spatial asymmetries previously attributed to the lateralized projections of the retinal hemifields. Across experiments, the strength of the precue-independent rightward advantage was variable. Comparing procedures across experiments, one possible explanation is that the larger cost of invalid left precues (i.e., the leftward advantage in invalid precue trials) was reduced when the locations of upcoming targets became less predictable overall. In Experiment 1, there was a (not statistically significant) leftward advantage in invalid precue trials and no cue validity-independent rightward advantage. In Experiment 2, trial-by-trial frame rotation added a degree of temporal, and perhaps spatial,8 uncertainty to the process of localizing the predicted target location, especially at the short SOA. In this Experiment, there was a smaller leftward advantage in invalid precue trials at the short (than at the long) SOA, and a significant precue validity-independent rightward advantage at the short SOA. Finally, in Experiment 3 the overall proportion of valid precue trials was reduced (from 75% to 60%) by the presence of neutral precue trials, and the precue-independent rightward advantage dominated the leftward advantage in invalid precue trials at both SOAs. The notion is that, under conditions of greater temporal/spatial uncertainty, participants were less likely to focus all their attention at the precued location, thus reducing the asymmetry in the cost of invalid precues. The impact of these small differences in the predictive value of the precues may have been enhanced by the fact that the precue, and especially the target, remained on the screen until the participant responded. In addition to the attention-related spatial asymmetries, the findings in Experiment 3 suggest that stimulus/response compatibility effects can be added to the list of spatial asymmetries influenced by scene-based frames. As there were not enough trials in the invalid conditions to analyze the letter reflection variable as a function of scene orientation in Experiment 2, we cannot be certain that this spatial asymmetry occurred within scene-based coordinates. However, the consistency of the spatial asymmetries obtained across experiments is very suggestive, and support for the idea that the Simon Effect can occur in scene-based coordinates is found in the literature. For instance, Hommel & Lippa (1995) showed quite convincingly that a stimulus/ response compatibility effect can occur in the functional equivalent of a scene rotated 8 This problem may have been amplified by the lack of target location markers, especially in the 90° display orientations. D.L. Rhodes, L.C. Robertson / Brain and Cognition 50 (2002) 95–115 113 out of alignment with the viewer and environment. Although it might seem counterintuitive that an asymmetry related to response-selection processes might rely on representations of space independent of the viewer, there is considerable evidence that the division into left/right of the stimulus representations underlying the Simon Effect is either ‘‘attention-centered’’ or relies on the programming of attentional movements through space (Nicoletti & Umilta, 1989, 1994 Stoffer & Yakin, 1994). The evidence reported here showing that multiple attention-related processing mechanisms are organized in scene-based frames does not mean that the hemispheres are not asymmetrically involved in the computation or use of the spatial coordinate systems that define these frames. In fact, the literature on hemispatial neglect reviewed in the Introduction suggests that they are (see also Behrmann & Hamison, 1999). Spatial locations in higher-order frames ultimately must be calculated on the basis of retinal coordinates in combination with other coordinate systems, and the left and right sides of these higher-order frames may remain preferentially represented in the right and left hemispheres, respectively (see Driver, 1999). However, our findings demonstrate that it would be a mistake to draw conclusions regarding the roles of the hemispheres on the basis of left/right visual field asymmetries alone, as the asymmetries we obtained do not appear to be due to differential direct access of visual information to the two hemispheres. Electrophysiological and other functional imaging methods will be beneficial in assessing both the temporal and spatial aspects of differential hemispheric function in the future. In addition, our findings require that studies of environment-based spatial representations take into account not only gravity-bound reference frames and reference frames provided by individual objects, but also the influence of contextual reference frames that encompass objects and are themselves more global objects, or scenes. The present findings show that even when there are multiple spatial influences on response times to task-relevant objects, the mappings underlying these influences can operate together in scene-based frames, dissociated from both the viewer and the gravitybound environment. Much work remains to be done in order to determine the conditions in which scenes are utilized and how the brain contends with multiple spatial reference frames, selecting the one(s) most relevant for the task at hand. Nevertheless, there is increasing evidence that the perceptual organization of a display and the spatial frames that define different levels of organization within it are critical for object perception, response mapping and the control of attention (Humphreys, 1983; Lamberts, Tavernier, & d’Ydewalle, 1992; Palmer, 1989; Quinlan, 1995; Robertson & Kim, 1998; Rock, 1990; Schendel & Robertson, submitted), and that each of these can influence the visual field asymmetries observed in normal performance. Acknowledgments The research was supported by a Veterans Administration Medical Research Scientist Award and National Science Foundation Award SBR-9222118 to Lynn C. Robertson, supplemented by a Research Opportunity Award to support Dell Rhodes. We thank John Lackey for the development of computer software, and Dana Myers for help in running Experiment 3. Harold Pashler, Ray Klein, Bill Yund, Krista Schendel, and three anonymous reviewers provided helpful comments. 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