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• W1973148612 abstract "Attentional selection plays a critical role in conscious perception. When attention is diverted, even salient stimuli fail to reach visual awareness [1Joseph J.S. Chun M.M. Nakayama K. Attentional requirements in a ‘preattentive’ feature search task.Nature. 1997; 387: 805-807Crossref PubMed Scopus (336) Google Scholar, 2Mack A. Rock I. Inattentional Blindness. MIT Press, Cambridge, MA1998Crossref Google Scholar]. Attention can be voluntarily directed to a spatial location [3Moran J. Desimone R. Selective attention gates visual processing in the extrastriate cortex.Science. 1985; 229: 782-784Crossref PubMed Scopus (1870) Google Scholar, 4Motter B.C. Focal attention produces spatially selective processing in visual cortical areas V1, V2, and V4 in the presence of competing stimuli.J. Neurophysiol. 1993; 70: 909-919PubMed Google Scholar, 5Reynolds J.H. Pasternak T. Desimone R. Attention increases sensitivity of V4 neurons.Neuron. 2000; 26: 703-714Abstract Full Text Full Text PDF PubMed Scopus (705) Google Scholar, 6Tootell R.B. Hadjikhani N. Hall E.K. Marrett S. Vanduffel W. Vaughan J.T. Dale A.M. The retinotopy of visual spatial attention.Neuron. 1998; 21: 1409-1422Abstract Full Text Full Text PDF PubMed Scopus (536) Google Scholar, 7Luck S.J. Chelazzi L. Hillyard S.A. Desimone R. Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex.J. Neurophysiol. 1997; 77: 24-42PubMed Google Scholar, 8Kastner S. Pinsk M.A. De Weerd P. Desimone R. Ungerleider L.G. Increased activity in human visual cortex during directed attention in the absence of visual stimulation.Neuron. 1999; 22: 751-761Abstract Full Text Full Text PDF PubMed Scopus (1177) Google Scholar, 9Desimone R. Duncan J. Neural mechanisms of selective visual attention.Annu. Rev. Neurosci. 1995; 18: 193-222Crossref PubMed Scopus (5540) Google Scholar] or a visual feature [9Desimone R. Duncan J. Neural mechanisms of selective visual attention.Annu. Rev. Neurosci. 1995; 18: 193-222Crossref PubMed Scopus (5540) Google Scholar, 10Treue S. Maunsell J.H. Attentional modulation of visual motion processing in cortical areas MT and MST.Nature. 1996; 382: 539-541Crossref PubMed Scopus (751) Google Scholar, 11Treue S. Martinez Trujillo J.C. Feature-based attention influences motion processing gain in macaque visual cortex.Nature. 1999; 399: 575-579Crossref PubMed Scopus (1004) Google Scholar, 12Martinez-Trujillo J.C. Treue S. Feature-based attention increases the selectivity of population responses in primate visual cortex.Curr. Biol. 2004; 14: 744-751Abstract Full Text Full Text PDF PubMed Scopus (459) Google Scholar, 13Saenz M. Buracas G.T. Boynton G.M. Global effects of feature-based attention in human visual cortex.Nat. Neurosci. 2002; 5: 631-632Crossref PubMed Scopus (420) Google Scholar, 14Rossi A.F. Paradiso M.A. Feature-specific effects of selective visual attention.Vision Res. 1995; 35: 621-634Crossref PubMed Scopus (76) Google Scholar] for facilitating the processing of information relevant to current goals. In everyday situations, attention and awareness are tightly coupled. This has led some to suggest that attention and awareness might be based on a common neural foundation [15O'Regan J.K. Noe A. A sensorimotor account of vision and visual consciousness.Behav. Brain Sci. 2001; 24: 939-973Crossref PubMed Scopus (1863) Google Scholar, 16Mitchell J.F. Stoner G.R. Reynolds J.H. Object-based attention determines dominance in binocular rivalry.Nature. 2004; 429: 410-413Crossref PubMed Scopus (182) Google Scholar], whereas others argue that they are mediated by distinct mechanisms [17Lamme V.A. Why visual attention and awareness are different.Trends Cogn. Sci. 2003; 7: 12-18Abstract Full Text Full Text PDF PubMed Scopus (704) Google Scholar, 18Dehaene S. Changeux J.P. Naccache L. Sackur J. Sergent C. Conscious, preconscious, and subliminal processing: a testable taxonomy.Trends Cogn. Sci. 2006; 10: 204-211Abstract Full Text Full Text PDF PubMed Scopus (1202) Google Scholar, 19Koch, C., and Tsuchiya, N. Attention and consciousness: Two distinct brain processes. Trends Cogn. Sci., in press.Google Scholar]. A body of evidence shows that visual stimuli can be processed at multiple stages of the visual-processing streams without evoking visual awareness [20He S. MacLeod D.I. Orientation-selective adaptation and tilt after-effect from invisible patterns.Nature. 2001; 411: 473-476Crossref PubMed Scopus (120) Google Scholar, 21Rajimehr R. Unconscious orientation processing.Neuron. 2004; 41: 663-673Abstract Full Text Full Text PDF PubMed Scopus (47) Google Scholar, 22Haynes J.D. Rees G. Predicting the orientation of invisible stimuli from activity in human primary visual cortex.Nat. Neurosci. 2005; 8: 686-691Crossref PubMed Scopus (606) Google Scholar]. To illuminate the relationship between visual attention and conscious perception, we investigated whether top-down attention can target and modulate the neural representations of unconsciously processed visual stimuli. Our experiments show that spatial attention can target only consciously perceived stimuli, whereas feature-based attention can modulate the processing of invisible stimuli. The attentional modulation of unconscious signals implies that attention and awareness can be dissociated, challenging a simplistic view of the boundary between conscious and unconscious visual processing. Attentional selection plays a critical role in conscious perception. When attention is diverted, even salient stimuli fail to reach visual awareness [1Joseph J.S. Chun M.M. Nakayama K. Attentional requirements in a ‘preattentive’ feature search task.Nature. 1997; 387: 805-807Crossref PubMed Scopus (336) Google Scholar, 2Mack A. Rock I. Inattentional Blindness. MIT Press, Cambridge, MA1998Crossref Google Scholar]. Attention can be voluntarily directed to a spatial location [3Moran J. Desimone R. Selective attention gates visual processing in the extrastriate cortex.Science. 1985; 229: 782-784Crossref PubMed Scopus (1870) Google Scholar, 4Motter B.C. Focal attention produces spatially selective processing in visual cortical areas V1, V2, and V4 in the presence of competing stimuli.J. Neurophysiol. 1993; 70: 909-919PubMed Google Scholar, 5Reynolds J.H. Pasternak T. Desimone R. Attention increases sensitivity of V4 neurons.Neuron. 2000; 26: 703-714Abstract Full Text Full Text PDF PubMed Scopus (705) Google Scholar, 6Tootell R.B. Hadjikhani N. Hall E.K. Marrett S. Vanduffel W. Vaughan J.T. Dale A.M. The retinotopy of visual spatial attention.Neuron. 1998; 21: 1409-1422Abstract Full Text Full Text PDF PubMed Scopus (536) Google Scholar, 7Luck S.J. Chelazzi L. Hillyard S.A. Desimone R. Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex.J. Neurophysiol. 1997; 77: 24-42PubMed Google Scholar, 8Kastner S. Pinsk M.A. De Weerd P. Desimone R. Ungerleider L.G. Increased activity in human visual cortex during directed attention in the absence of visual stimulation.Neuron. 1999; 22: 751-761Abstract Full Text Full Text PDF PubMed Scopus (1177) Google Scholar, 9Desimone R. Duncan J. Neural mechanisms of selective visual attention.Annu. Rev. Neurosci. 1995; 18: 193-222Crossref PubMed Scopus (5540) Google Scholar] or a visual feature [9Desimone R. Duncan J. Neural mechanisms of selective visual attention.Annu. Rev. Neurosci. 1995; 18: 193-222Crossref PubMed Scopus (5540) Google Scholar, 10Treue S. Maunsell J.H. Attentional modulation of visual motion processing in cortical areas MT and MST.Nature. 1996; 382: 539-541Crossref PubMed Scopus (751) Google Scholar, 11Treue S. Martinez Trujillo J.C. Feature-based attention influences motion processing gain in macaque visual cortex.Nature. 1999; 399: 575-579Crossref PubMed Scopus (1004) Google Scholar, 12Martinez-Trujillo J.C. Treue S. Feature-based attention increases the selectivity of population responses in primate visual cortex.Curr. Biol. 2004; 14: 744-751Abstract Full Text Full Text PDF PubMed Scopus (459) Google Scholar, 13Saenz M. Buracas G.T. Boynton G.M. Global effects of feature-based attention in human visual cortex.Nat. Neurosci. 2002; 5: 631-632Crossref PubMed Scopus (420) Google Scholar, 14Rossi A.F. Paradiso M.A. Feature-specific effects of selective visual attention.Vision Res. 1995; 35: 621-634Crossref PubMed Scopus (76) Google Scholar] for facilitating the processing of information relevant to current goals. In everyday situations, attention and awareness are tightly coupled. This has led some to suggest that attention and awareness might be based on a common neural foundation [15O'Regan J.K. Noe A. A sensorimotor account of vision and visual consciousness.Behav. Brain Sci. 2001; 24: 939-973Crossref PubMed Scopus (1863) Google Scholar, 16Mitchell J.F. Stoner G.R. Reynolds J.H. Object-based attention determines dominance in binocular rivalry.Nature. 2004; 429: 410-413Crossref PubMed Scopus (182) Google Scholar], whereas others argue that they are mediated by distinct mechanisms [17Lamme V.A. Why visual attention and awareness are different.Trends Cogn. Sci. 2003; 7: 12-18Abstract Full Text Full Text PDF PubMed Scopus (704) Google Scholar, 18Dehaene S. Changeux J.P. Naccache L. Sackur J. Sergent C. Conscious, preconscious, and subliminal processing: a testable taxonomy.Trends Cogn. Sci. 2006; 10: 204-211Abstract Full Text Full Text PDF PubMed Scopus (1202) Google Scholar, 19Koch, C., and Tsuchiya, N. Attention and consciousness: Two distinct brain processes. Trends Cogn. Sci., in press.Google Scholar]. A body of evidence shows that visual stimuli can be processed at multiple stages of the visual-processing streams without evoking visual awareness [20He S. MacLeod D.I. Orientation-selective adaptation and tilt after-effect from invisible patterns.Nature. 2001; 411: 473-476Crossref PubMed Scopus (120) Google Scholar, 21Rajimehr R. Unconscious orientation processing.Neuron. 2004; 41: 663-673Abstract Full Text Full Text PDF PubMed Scopus (47) Google Scholar, 22Haynes J.D. Rees G. Predicting the orientation of invisible stimuli from activity in human primary visual cortex.Nat. Neurosci. 2005; 8: 686-691Crossref PubMed Scopus (606) Google Scholar]. To illuminate the relationship between visual attention and conscious perception, we investigated whether top-down attention can target and modulate the neural representations of unconsciously processed visual stimuli. Our experiments show that spatial attention can target only consciously perceived stimuli, whereas feature-based attention can modulate the processing of invisible stimuli. The attentional modulation of unconscious signals implies that attention and awareness can be dissociated, challenging a simplistic view of the boundary between conscious and unconscious visual processing. In the first experiment, we examined whether spatial attention can target unconscious signals for invisible stimuli by using the tilt aftereffect (TAE), where prolonged viewing of an adaptation stimulus slightly tilted from the vertical causes a percept of an opposite tilt for a subsequently presented vertical stimulus [23Gibson J.J. Radner M. Adaptation, aftereffect and contrast in the perception of tilted lines.J. Exp. Psychol. 1937; 20: 453-467Crossref Scopus (508) Google Scholar]. A number of neurophysiological [3Moran J. Desimone R. Selective attention gates visual processing in the extrastriate cortex.Science. 1985; 229: 782-784Crossref PubMed Scopus (1870) Google Scholar, 4Motter B.C. Focal attention produces spatially selective processing in visual cortical areas V1, V2, and V4 in the presence of competing stimuli.J. Neurophysiol. 1993; 70: 909-919PubMed Google Scholar, 5Reynolds J.H. Pasternak T. Desimone R. Attention increases sensitivity of V4 neurons.Neuron. 2000; 26: 703-714Abstract Full Text Full Text PDF PubMed Scopus (705) Google Scholar] and brain-imaging [6Tootell R.B. Hadjikhani N. Hall E.K. Marrett S. Vanduffel W. Vaughan J.T. Dale A.M. The retinotopy of visual spatial attention.Neuron. 1998; 21: 1409-1422Abstract Full Text Full Text PDF PubMed Scopus (536) Google Scholar] studies have shown that spatial attention modulates cortical activity in a retinotopic fashion, even in the absence of visual stimuli [7Luck S.J. Chelazzi L. Hillyard S.A. Desimone R. Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex.J. Neurophysiol. 1997; 77: 24-42PubMed Google Scholar, 8Kastner S. Pinsk M.A. De Weerd P. Desimone R. Ungerleider L.G. Increased activity in human visual cortex during directed attention in the absence of visual stimulation.Neuron. 1999; 22: 751-761Abstract Full Text Full Text PDF PubMed Scopus (1177) Google Scholar]. A case study on a blindsight patient has shown that precueing a target location in his “blind” hemifield facilitates the processing of a stimulus invisible to the patient at that location [24Kentridge R.W. Heywood C.A. Weiskrantz L. Attention without awareness in blindsight.Proc Biol Sci. 1999; 266: 1805-1811Crossref PubMed Scopus (230) Google Scholar]. These lines of evidence suggest that spatial attention might increase the neural responses to invisible adapting stimuli and thus lead to greater TAE. Alternatively, if top-down spatial attention is limited to consciously perceived stimuli, attending to the location of an invisible stimulus should not affect adaptation at all. The design of the experiment is illustrated in Figure 1A. We rendered adapting stimuli invisible by presenting rapidly changing Mondrian patterns presented to the other eye, a technique called continuous flash suppression (CFS) [25Tsuchiya N. Koch C. Continuous flash suppression reduces negative afterimages.Nat. Neurosci. 2005; 8: 1096-1101Crossref PubMed Scopus (581) Google Scholar]. We manipulated spatial attention by instructing the observers to attend to one of the two spatial markers drawn on top of the Mondrian patterns (Figure 1A). The positions of the markers corresponded to the locations of the Gabor stimuli presented to the other, suppressed left eye (Figure 1A). The two adaptors had the same orientation so that the effects of feature-based attention remain constant. After adaptation of the left eye to the invisible Gabor with a tilt orientation from the vertical by 15° for 5 s, a test stimulus was presented to the eye, either at the attended or the unattended location. The observers were asked to judge whether the test Gabor stimulus with a variable orientation was tilted to the right or left. In a control experiment, we repeated the same experiment without the presentation of the Mondrian masks. In this case, both adaptors were visible to the observers. We included this control condition to confirm that the effect of spatial attention is obtained reliably with our stimulus parameters (see the Experimental Procedures in Supplemental Data available with this article online for further details). The results (Figure 1B) show that TAE was induced by invisible adaptors (paired t test against the baseline, T(3) = 4.73, p < 0.05 for the attended location; T(3) = 5.69, p < 0.05 for the unattended location), indicating that the orientation information was processed to a certain degree for the invisible stimuli. However, the TAE magnitude was not significantly different (T(3) = 1.45, p = 0.24) regardless of whether the TAE was tested at the attended location (1.98° ± 0.34°; hereafter the SEM are shown to indicate interobserver variability) or at the unattended location (2.15° ± 0.29°). On the other hand, when the adaptors were visible (Figure 1C), the TAE magnitudes were subject to attentional modulation (visible-attended, 4.71° ± 0.81°; visible-unattended, 3.86° ± 0.74°). With a two-way ANOVA, we confirmed the main effects of visibility (F(1,3) = 22.60, p < 0.05) and attention (F(1,3) = 33.36, p < 0.05) and a significant interaction (F(1,3) = 16.99, p < 0.05) (Figure 1D). The significant interaction between visibility and spatial attention indicates that spatial attention modulates the TAE only when the adaptors were consciously visible (visible-attended versus visible-unattended, T(3) = 5.51, p < 0.05). The attentional modulation in the visible condition ensures that the lack of attentional modulation in the invisible condition is not due to the particular parameter set used in the present experiment or to a failure of the observers to voluntarily allocate attention (more data to support this point are available in the Supplemental Data). In the second experiment, we examined whether feature-based attention can reach and modulate the TAE induced by invisible stimuli. In a previous study, it was shown that feature-based attention spreads to a brief, unattended stimulus [26Melcher D. Papathomas T.V. Vidnyanszky Z. Implicit attentional selection of bound visual features.Neuron. 2005; 46: 723-729Abstract Full Text Full Text PDF PubMed Scopus (83) Google Scholar], whose critical feature (i.e., motion) was presented at the subthreshold level. This suggests that feature-based attention might be able to reach processing levels where signals do not directly give rise to a conscious percept. The design of the experiment is illustrated in Figure 2A. While adapting to an invisible adaptor presented to the left visual field of the left eye, observers continuously attended to one of the two visible targets displayed in the right visual field of the same eye (Figure 2A). One of the visible attention targets was a Gabor patch with the same tilt orientation as the adaptor, and the other target had the opposite tilt orientation. After 5 s of adaptation, a test stimulus was presented at the location of the invisible adaptor. The observers judged whether the test stimulus was tilted to the left or right. In a control experiment, we conducted the same experiment without presenting the masks and left the adapting stimuli visible to the observers (see the Experimental Procedures in the Supplemental Data for details). The results are shown in Figure 2B. The TAE magnitude was larger (T(3) = 6.02, p < 0.01) when the attended stimulus share the same orientation as the invisible adaptor (3.35° ± 0.37°) than when it had the opposite orientation (2.54° ± 0.08°). As before, the invisible stimuli produced a significant TAE (attend-same, 3.35° ± 0.09°, paired t test against the baseline, T(3) = 33.6, p < 0.01; attend-opposite, 2.54° ± 0.37°, T(3) = 6.42, p < 0.01). As expected, attentional modulation was observed also in the visible condition (Figure 3C). Attending to the same-tilt target (4.95° ± 0.34°) resulted in a larger TAE (T(3) = 6.43, p < 0.01) than attending to the opposite-tilt target (4.13° ± 0.30°). A two-way ANOVA revealed the main effects of visibility of the adaptor (F(1,3) = 10.81, p < 0.05) and attention (F(1,3) = 14.51, p < 0.05) but no interaction between the two factors (F(3,3) < 1). The main effect of visibility replicates our finding in the experiments on spatial attention; TAE is reduced when adaptors are suppressed by Mondrian patterns. The main effect of attention together with the absence of interaction confirms that the TAE magnitude both for the visible and invisible adaptors can be modulated depending on the orientation of the distant attended target. This suggests that feature-based attention has the ability to reach and modulate unconscious signals arising from invisible adapting stimuli. The individual data for the spatial attention experiment and the feature-based attention are summarized in Figure 3. Although there is an interobserver variability in TAE magnitudes, the pattern of attentional modulation was consistent for all observers. This consistency corroborates our conclusions. Our present study demonstrates that top-down feature-based attention but not spatial attention can modulate the TAE magnitudes to unconsciously processed visual stimuli. Previous neurophysiological and brain-imaging studies showed that spatial attention enhances the neural activity at the attended location even in the absence of physical stimuli [7Luck S.J. Chelazzi L. Hillyard S.A. Desimone R. Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex.J. Neurophysiol. 1997; 77: 24-42PubMed Google Scholar, 8Kastner S. Pinsk M.A. De Weerd P. Desimone R. Ungerleider L.G. Increased activity in human visual cortex during directed attention in the absence of visual stimulation.Neuron. 1999; 22: 751-761Abstract Full Text Full Text PDF PubMed Scopus (1177) Google Scholar]. Moreover, one previous study found that attending to a spatial location invisible to a blindsight patient facilitated reaction times to the target presented at that location [24Kentridge R.W. Heywood C.A. Weiskrantz L. Attention without awareness in blindsight.Proc Biol Sci. 1999; 266: 1805-1811Crossref PubMed Scopus (230) Google Scholar]. Despite this, we found no evidence of attentional modulation by top-down spatial selection. According to population coding models of orientation perception, the TAE is thought to arise from an imbalance in population activity caused by selective adaptation of neurons responsive for that particular orientation [27Clifford C.W. Wenderoth P. Spehar B. A functional angle on some after-effects in cortical vision.Proc Biol Sci. 2000; 267: 1705-1710Crossref PubMed Scopus (160) Google Scholar, 28Jin D.Z. Dragoi V. Sur M. Seung H.S. The tilt aftereffect and adaptation-induced changes in orientation tuning in visual cortex.J. Neurophysiol. 2005; 94: 4038-4050Crossref PubMed Scopus (77) Google Scholar, 29Kohn A. Movshon J.A. Adaptation changes the direction tuning of macaque MT neurons.Nat. Neurosci. 2004; 7: 764-772Crossref PubMed Scopus (228) Google Scholar]. In this sense, it is possible that spatial attention produces a generic enhancement of the entire neural population but has no influences on the TAE magnitude. Spatial attention might increase the total activity level at the attended location, but the nonselective attentional effects on the whole population do not result in a modulation of TAE magnitudes. This idea may reconcile the generic influence of spatial attention and the lack of attentional modulation for the TAE from invisible stimuli. It should be noted that our control experiment (Figures 1D) as well as a past study [30Spivey M.J. Spirn M.J. Selective visual attention modulates the direct tilt aftereffect.Percept. Psychophys. 2000; 62: 1525-1533Crossref PubMed Scopus (57) Google Scholar] have shown that spatial attention as such has the ability to modulate the TAE when the adaptors are visible and feature-based attention is constant. This indicates that when adaptors are visible, spatial attention as such must have orientation-selective effects, but the orientation specificity appears to be lost for invisible adaptors. It remains to be seen whether the absence of effects of spatial attention to invisible stimuli is limited to the case of TAE or generalized to other types of adaptation effects. In contrast to spatial attention, feature-based attention did modulate adaptation to invisible stimuli and offered empirical evidence that top-down attention can be dissociated from awareness [17Lamme V.A. Why visual attention and awareness are different.Trends Cogn. Sci. 2003; 7: 12-18Abstract Full Text Full Text PDF PubMed Scopus (704) Google Scholar, 18Dehaene S. Changeux J.P. Naccache L. Sackur J. Sergent C. Conscious, preconscious, and subliminal processing: a testable taxonomy.Trends Cogn. Sci. 2006; 10: 204-211Abstract Full Text Full Text PDF PubMed Scopus (1202) Google Scholar, 19Koch, C., and Tsuchiya, N. Attention and consciousness: Two distinct brain processes. Trends Cogn. Sci., in press.Google Scholar]. Other studies have found that attention can affect priming from invisible stimuli [31Sumner P. Tsai P.C. Yu K. Nachev P. Attentional modulation of sensorimotor processes in the absence of perceptual awareness.Proc. Natl. Acad. Sci. USA. 2006; 103: 10520-10525Crossref PubMed Scopus (104) Google Scholar, 32Naccache L. Blandin E. Dehaene S. Unconscious masked priming depends on temporal attention.Psychol. Sci. 2002; 13: 416-424Crossref PubMed Scopus (367) Google Scholar]. These lines of evidence for the dissociation between awareness and attention implies a potential confound for neurophysiological studies searching for the neural correlates of visual awareness. Some studies use attentional modulation of neuronal activity as a marker of facilitated visual processing leading to a conscious percept (e.g., [9Desimone R. Duncan J. Neural mechanisms of selective visual attention.Annu. Rev. Neurosci. 1995; 18: 193-222Crossref PubMed Scopus (5540) Google Scholar]). Others look for neuronal correlates of perceptual changes in effects such as binocular rivalry [33Logothetis N.K. Schall J.D. Neuronal correlates of subjective visual perception.Science. 1989; 245: 761-763Crossref PubMed Scopus (425) Google Scholar, 34Leopold D.A. Logothetis N.K. Activity changes in early visual cortex reflect monkeys' percepts during binocular rivalry.Nature. 1996; 379: 549-553Crossref PubMed Scopus (666) Google Scholar, 35Tong F. Engel S.A. Interocular rivalry revealed in the human cortical blind-spot representation.Nature. 2001; 411: 195-199Crossref PubMed Scopus (347) Google Scholar, 36Wunderlich K. Schneider K.A. Kastner S. Neural correlates of binocular rivalry in the human lateral geniculate nucleus.Nat. Neurosci. 2005; 8: 1595-1602Crossref PubMed Scopus (213) Google Scholar, 37Haynes J.D. Deichmann R. Rees G. Eye-specific effects of binocular rivalry in the human lateral geniculate nucleus.Nature. 2005; 438: 496-499Crossref PubMed Scopus (262) Google Scholar]. However, attentional modulation of unconscious sensory signals implies that neural activities exhibiting a percept-dependent modulation may not necessarily be directly linked with visual awareness. Instead, they could reflect the effects of attention on unconscious signals via attended or consciously perceived stimuli. As has been emphasized in recent fMRI studies [38Huk A.C. Ress D. Heeger D.J. Neuronal basis of the motion aftereffect reconsidered.Neuron. 2001; 32: 161-172Abstract Full Text Full Text PDF PubMed Scopus (238) Google Scholar], the dissociation of attentional effects from awareness will become essential when one tries to determine the neural activity associated with conscious percept [19Koch, C., and Tsuchiya, N. Attention and consciousness: Two distinct brain processes. Trends Cogn. Sci., in press.Google Scholar]. Finally, the present study supports the notion that spatial attention and feature-based attention are based on distinct neural mechanisms [39McAdams C.J. Maunsell J.H. Attention to both space and feature modulates neuronal responses in macaque area V4.J. Neurophysiol. 2000; 83: 1751-1755PubMed Google Scholar, 40Hayden B.Y. Gallant J.L. Time course of attention reveals different mechanisms for spatial and feature-based attention in area V4.Neuron. 2005; 47: 637-643Abstract Full Text Full Text PDF PubMed Scopus (98) Google Scholar]. Electrophysiological studies have shown that the effects of spatial attention are delayed and lag behind the initial transient responses to a stimulus onset [40Hayden B.Y. Gallant J.L. Time course of attention reveals different mechanisms for spatial and feature-based attention in area V4.Neuron. 2005; 47: 637-643Abstract Full Text Full Text PDF PubMed Scopus (98) Google Scholar, 41Reynolds J.H. Chelazzi L. Desimone R. Competitive mechanisms subserve attention in macaque areas V2 and V4.J. Neurosci. 1999; 19: 1736-1753PubMed Google Scholar, 42Mehta A.D. Ulbert I. Schroeder C.E. Intermodal selective attention in monkeys. I: Distribution and timing of effects across visual areas.Cereb. Cortex. 2000; 10: 343-358Crossref PubMed Scopus (201) Google Scholar], whereas feature-based attention has both an early and continuing effect and modulates even the initial transient responses [40Hayden B.Y. Gallant J.L. Time course of attention reveals different mechanisms for spatial and feature-based attention in area V4.Neuron. 2005; 47: 637-643Abstract Full Text Full Text PDF PubMed Scopus (98) Google Scholar]. Interestingly, several lines of evidence suggest that the initial feedforward sweep elicited by a stimulus onset does not give rise to visual awareness, whereas later responses do correlate with the conscious percept [43Lamme V.A. Roelfsema P.R. The distinct modes of vision offered by feedforward and recurrent processing.Trends Neurosci. 2000; 23: 571-579Abstract Full Text Full Text PDF PubMed Scopus (1484) Google Scholar]. Our findings and these physiological results offer interesting parallels regarding feature-based attention and early unconscious processing versus spatial attention and late conscious processing. This research was partly supported by the Netherlands Organization for Scientific Research (NWO). We would like to thank Tom Carlson, Patrick Cavanagh, Christof Koch, David Leopold, Alex Maier, Farshad Moradi, Chris Paffen, Junghyun Park, Takao Sato, Jan Theeuwes, Keiji Tanaka, Masataka Watanabe, Melanie Wilke, and Daw-An Wu for discussion and comments on the manuscript. Download .pdf (.89 MB) Help with pdf files Document S1. Supplemental Results, Supplemental Experimental Procedures, and Five Supplemental Figures" @default.
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• W1973148612 date "2006-12-01" @default.
• W1973148612 modified "2023-10-11" @default.
• W1973148612 title "The Scope and Limits of Top-Down Attention in Unconscious Visual Processing" @default.
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