The visual system is intelligent—it is capable of recovering a coherent surface from an incomplete one, a feat known as perceptual completion or filling in. Traditionally, it has been assumed that surface features are interpolated in a way that resembles the fragmented parts. Using displays featuring four circular apertures, we showed in the study reported here that a distinct completed feature (horizontal motion) arises from local ones (oblique motions)—we term this process emergent filling in. Adaptation to emergent filling-in motion generated a dynamic motion aftereffect that was not due to spreading of local motion from the isolated apertures. The filling-in motion aftereffect occurred in both modal and amodal completions, and it was modulated by selective attention. These findings highlight the importance of high-level interpolation processes in filling in and are consistent with the idea that during emergent filling in, the more cognitive-symbolic processes in later areas (e.g., the middle temporal visual area and the lateral occipital complex) provide important feedback signals to guide more isomorphic processes in earlier areas (V1 and V2).
BrainardD. H. (1997). The psychophysics toolbox. Spatial Vision, 10, 433–436.
3.
BrittenK. H.ShadlenM. N.NewsomeW. T.MovshonJ. A. (1992). The analysis of visual motion: A comparison of neuronal and psychophysical performance. Journal of Neuroscience, 12, 4745–4765.
4.
CookE. P.MaunsellJ. H. (2004). Attentional modulation of motion integration of individual neurons in the middle temporal visual area. Journal of Neuroscience, 24, 7964–7977.
5.
CornelissenF. W.WadeA. R.VladusichT.DoughertyR. F.WandellB. A. (2006). No functional magnetic resonance imaging evidence for brightness and color filling-in in early human visual cortex. Journal of Neuroscience, 26, 3634–3641.
6.
CulhamJ.NishidaS.LedgewayT.CavanaghP.von GrünauM.KwasM.. . . RaymondJ. (1998). Higher-order effects. In MatherG.VerstratenF. A. J.AnstisS. (Eds.), The motion aftereffect: A modern perspective (pp. 85–124). Cambridge, MA: Massachusetts Institute of Technology.
7.
DavisG.DriverJ. (1994). Parallel detection of Kanizsa subjective figures in the human visual system. Nature, 371, 791–793.
8.
DavisG.DriverJ. (1997). Spreading of visual attention to modally versus amodally completed regions. Psychological Science, 8, 275–281.
9.
DavisG.DriverJ. (1998). Kanizsa subjective figures can act as occluding surfaces at parallel stages of visual search. Journal of Experimental Psychology: Human Perception and Performance, 24, 169–184.
10.
DennettD. C. (1992). Filling in versus finding out: A ubiquitous confusion in cognitive science. In PickH. L.Jr.van den BroekP.KnillD. C. (Eds.), Cognition: Conceptual and methodological issues (pp. 33–49). Washington, DC: American Psychological Association.
11.
De WeerdP.GattassR.DesimoneR.UngerleiderL. G. (1995). Responses of cells in monkey visual cortex during perceptual filling-in of an artificial scotoma. Nature, 377, 731–734.
12.
FioraniM.RosaM. G.GattassR.Rocha-MirandaC. E. (1992). Dynamic surrounds of receptive fields in primate striate cortex: A physiological basis for perceptual completion?Proceedings of the National Academy of Sciences, USA, 89, 8547–8551.
13.
GerritsH. J.VendrikA. J. (1970). Simultaneous contrast, filling-in process and information processing in man’s visual system. Experimental Brain Research, 11, 411–430.
14.
GilbertC. D. (1992). Horizontal integration and cortical dynamics. Neuron, 9, 1–13.
15.
GilbertC. D.DasA.ItoM.KapadiaM.WestheimerG. (1996). Spatial integration and cortical dynamics. Proceedings of the National Academy of Sciences, USA, 93, 615–622.
16.
GregoryR. L. (1972). Cognitive contours. Nature, 238, 51–52.
17.
GrossbergS. (1994). 3-D vision and figure-ground separation by visual cortex. Perception & Psychophysics, 55, 48–121.
18.
HalgrenE.MendolaJ.ChongC. D. R.DaleA. M. (2003). Cortical activation to illusory shapes as measured with magnetoencephalography. NeuroImage, 18, 1001–1009.
19.
HeZ. J.NakayamaK. (1992). Surfaces versus features in visual search. Nature, 359, 231–233.
20.
KanizsaG. (1979). Organization in vision: Essays on gestalt perception. New York, NY: Praeger.
21.
KellmanP. J.ShipleyT. F. (1991). A theory of visual interpolation in object perception. Cognitive Psychology, 23, 141–221.
22.
KomatsuH. (2006). The neural mechanisms of perceptual filling-in. Nature Reviews Neuroscience, 7, 220–231.
23.
LakA. (2008). Attention during adaptation weakens negative afterimages of perceptually colour-spread surfaces. Canadian Journal of Experimental Psychology, 62, 101–109.
24.
LalanneC.LorenceauJ. (2006). Directional shifts in the barber pole illusion: Effects of spatial frequency, spatial adaptation, and lateral masking. Visual Neuroscience, 23, 729–739.
25.
LeeT. S.NguyenM. (2001). Dynamics of subjective contour formation in the early visual cortex. Proceedings of the National Academy of Sciences, USA, 98, 1907–1911.
26.
LiZ. (1998). A neural model of contour integration in the primary visual cortex. Neural Computation, 10, 903–940.
27.
MarrD.UllmanS. (1981). Directional selectivity and its use in early visual processing. Proceedings of the Royal Society of London B: Biological Sciences, 211, 151–180.
28.
MattingleyJ. B.DavisG.DriverJ. (1997). Preattentive filling-in of visual surfaces in parietal extinction. Science, 275, 671–674.
29.
MendolaJ. D.DaleA. M.FischlB.LiuA. K.TootellR. B. (1999). The representation of illusory and real contours in human cortical visual areas revealed by functional magnetic resonance imaging. Journal of Neuroscience, 19, 8560–8572.
30.
MengM.RemusD. A.TongF. (2005). Filling-in of visual phantoms in the human brain. Nature Neuroscience, 8, 1248–1254.
31.
MengX.MazzoniP.QianN. (2006). Cross-fixation transfer of motion aftereffects with expansion motion. Vision Research, 46, 3681–3689.
32.
MichotteA.ThinesG.CrabbeG. (1964). Les complements amodaux des structures perceptives [Amodal complements of perceptual structures]. Louvain, France: Studia Psycologica Louvain, Publications Universitaires de Louvain.
MurrayM. M.FoxeD. M.JavittD. C.FoxeJ. J. (2004). Setting boundaries: Brain dynamics of modal and amodal illusory shape completion in humans. Journal of Neuroscience, 24, 6898–6903.
35.
NishidaS.AshidaH.SatoT. (1994). Complete interocular transfer of motion aftereffect with flickering test. Vision Research, 34, 2707–2716.
36.
PackC. C.BornR. T. (2001). Temporal dynamics of a neural solution to the aperture problem in visual area MT of macaque brain. Nature, 409, 1040–1042.
37.
PalmerS. E.NeffJ.BeckD. (1996). Late influences on perceptual grouping: Amodal completion. Psychonomic Bulletin & Review, 3, 75–80.
38.
PelliD. G. (1997). The VideoToolbox software for visual psychophysics: Transforming numbers into movies. Spatial Vision, 10, 437–442.
39.
PernaA.TosettiM.MontanaroD.MorroneM. C. (2005). Neuronal mechanisms for illusory brightness perception in humans. Neuron, 47, 645–651.
40.
PessoaL.ThompsonE.NoeA. (1998). Filling-in is for finding out. Behavioral and Brain Sciences, 21, 781–802.
41.
PillowJ.RubinN. (2002). Perceptual completion across the vertical meridian and the role of early visual cortex. Neuron, 33, 805–813.
42.
PriceN. S.GreenwoodJ. A.IbbotsonM. R. (2004). Tuning properties of radial phantom motion aftereffects. Vision Research, 44, 1971–1979.
SeghierM. L.VulleumierP. (2006). Functional neuroimaging findings on the human perception of illusory contours. Neuroscience & Biobehavioral Reviews, 30, 595–612.
49.
ShimojoS.KamitaniY.NishidaS. (2001). Afterimage of perceptually filled-in surface. Science, 293, 1677–1680.
50.
SinghM. (2004). Modal and amodal completion generate different shapes. Psychological Science, 15, 454–459.
51.
SmithA.OverR. (1975). Tilt aftereffects with subjective contours. Nature, 257, 581–582.
52.
SnowdenR. J.MilneA. B. (1997). Phantom motion aftereffects: Evidence of detectors for the analysis of optic flow. Current Biology, 7, 717–722.
53.
StanleyD. A.RubinN. (2003). fMRI activation in response to illusory contours and salient regions in the human lateral occipital complex. Neuron, 37, 323–331.
54.
TynanP.SekularR. (1975). Moving visual phantoms: A new contour completion effect. Science, 188, 951–952.
55.
UllmanS. (1976). Filling-in gaps: Shape of subjective contours and a model for their generation. Biological Cybernetics, 25, 1–6.
56.
WeissteinN.MaguireW.BerbaumK. (1977). A phantom-motion aftereffect. Science, 198, 955–958.
Supplementary Material
Please find the following supplemental material available below.
For Open Access articles published under a Creative Commons License, all supplemental material carries the same license as the article it is associated with.
For non-Open Access articles published, all supplemental material carries a non-exclusive license, and permission requests for re-use of supplemental material or any part of supplemental material shall be sent directly to the copyright owner as specified in the copyright notice associated with the article.
