Topographic representation of an occluded object and the effects of spatiotemporal context in human early visual areas
Open accessAbstract
Occlusion is a primary challenge facing the visual system in perceiving object shapes in intricate natural scenes. Although behavior, neurophysiological, and modeling studies have shown that occluded portions of objects may be completed at the early stage of visual processing, we have little knowledge on how and where in the human brain the completion is realized. Here, we provide functional magnetic resonance imaging (fMRI) evidence that the occluded portion of an object is indeed represented topographically in human V1 and V2. Specifically, we find the topographic cortical responses corresponding to the invisible object rotation in V1 and V2. Furthermore, by investigating neural responses for the occluded target rotation within precisely defined cortical subregions, we could dissociate the topographic neural representation of the occluded portion from other types of neural processing such as object edge processing. We further demonstrate that the early topographic representation in V1 can be modulated by prior knowledge of a whole appearance of an object obtained before partial occlusion. These findings suggest that primary "visual" area V1 has the ability to process not only visible or virtually (illusorily) perceived objects but also "invisible" portions of objects without concurrent visual sensation such as luminance enhancement to these portions. The results also suggest that low-level image features and higher preceding cognitive context are integrated into a unified topographic representation of occluded portion in early areas.
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Activity patterns in nonstimulated parts of early visual cortex contain content-specific information from the surrounding visual context, from objects presented in the periphery, and from visual stimuli solely held in working memory rather than being actively perceived.
“For example, activity patterns in nonstimulated parts of early visual cortex contain content-specific information from the surrounding visual context [31, 32], from objects presented in the periphery [33], and from visual stimuli solely held in working memory rather than being actively perceived [34, 35].”
From Decoding sound and imagery content in early visual cortex (2014) · cites this paper as [32]
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MethodSpatially selective responses to Kanizsa and occlusion stimuli in human visual cortex
de Haas, B., & Schwarzkopf, D. S. (2018) · Scientific Reports · doi:10.1038/s41598-017-19121-z · Cited by 17 · see citing works
Erlikhman, G., Gurariy, G., Mruczek, R. E. B., & Caplovitz, G. P. (2016) · NeuroImage · doi:10.1016/j.neuroimage.2016.03.044 · Cited by 18 · see citing works
“…examining the neural representation of occluded objects that are static (Edelman et al., 1998; Grill-Spector et al., 1998b; Kourtzi and Kanwisher, 2000, 2001), little work has been done to identify the neural correlates of spatiotemporal object perception (Yin et al., 2002; Ban et al., 2013).”
Revina, Y., Petro, L. S., Denk-Florea, C.-B., Rao, I. S., & Muckli, L. F. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.02.27.433018 · Cited by 18 · see citing works
Pre-Cueing Effects: Attention or Mental Imagery?
Fazekas, P., & Nánay, B. (2017) · Frontiers in Psychology · doi:10.3389/fpsyg.2017.00222 · Cited by 17 · see citing works
“…2001; Komatsu, 2006; Hedgé et al., 2008; Lommertzen et al., 2009; Vrins et al., 2009; Nanay, 2010a; Smith and Muckli, 2010; Bushnell et al., 2011; Shibata et al., 2011; Lee et al., 2012; Pan et al., 2012; Ban et al., 2013; Emmanouil and Ro, 2014; Hazenberg et al., 2014; Scherzer and Ekroll, 2015).”
BackgroundDecoding Visual fMRI Stimuli from Human Brain Based on Graph Convolutional Neural Network
Meng, L., & Ge, K. (2022) · Brain Sciences · doi:10.3390/brainsci12101394 · Cited by 13 · see citing works
“To sum up, the main brain response areas to visual stimuli include V1, V2, V3, V4, LOC, FFA, and PPA [1,12,26].”
BackgroundMethodResolving visual motion through perceptual gaps
Teichmann, L., Edwards, G., & Baker, C. I. (2021) · Trends in Cognitive Sciences · doi:10.1016/j.tics.2021.07.017 · Cited by 13 · see citing works
“Similar to representations during apparent motion [83,84], position-speci fi c information has been shown to be represented in the early visual cortex during perceptual gaps [85 – 87].”
Zbären, G. A., Meissner, S. N., Kapur, M., & Wenderoth, N. (2023) · Human Brain Mapping · doi:10.1002/hbm.26338 · Cited by 9 · see citing works
“The activations observed in early visual and parietal regions overlap with areas that have been shown to process occluded motion (Ban et al., 2013; Erlikhman & Caplovitz, 2017; Olson et al., 2004; Shuwairi et al., 2007).”
BackgroundGeorge, D., Lázaro-Gredilla, M., Lehrach, W. P., Dedieu, A., & Zhou, G. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.09.09.290601 · Cited by 9 · see citing works
“The dynamics of occlusion reasoning and filling in have limited studies in neuroscience [83–85], and predictions from Bio-RCN might be a useful tool in further explorations.”
BackgroundFemale túngara frogs do not experience the continuity illusion.
Baugh, A. T., Ryan, M. J., Bernal, X. E., Rand, A. S., & Bee, M. A. (2015) · Behavioral Neuroscience · doi:10.1037/bne0000115 · Cited by 7 · see citing works
“Cortical processing is also implicated in generating a coherent perception of partially occluded visual objects (Ban et al., 2013; Hegde, Fang, Murray, & Kersten, 2008; Kosai, El-Shamayleh, Fyall, & Pasupathy, 2014; Liu, Plomp, van Leeuwen, & Ioannides, 2006; Rauschenberger, Liu, Slotnick, &…”
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0013 · Cited by 7 · see citing works
Brain Functional Representation of Highly Occluded Object Recognition
Li, B., Zhang, C., Cao, L., Chen, P., Liu, T., Gao, H., Wang, L., Yan, B., & Tong, L. (2023) · Brain Sciences · doi:10.3390/brainsci13101387 · Cited by 5 · see citing works
“According to one functional magnetic resonance imaging (fMRI) study, low-level visual regions exhibited Brain Sci. 2023 , 13 , 1387 2 of 14 more activation in the occluded condition as compared to the non-occluded one [8].”
ResultHanakawa, T. (2015) · Springer eBooks · doi:10.1007/978-4-431-54541-5_5 · Cited by 5 · see citing works
Dissociation of perceived size and perceived strength in the scintillating grid illusion
Matsuno, T., & Sato, Y. (2019) · Journal of Vision · doi:10.1167/19.13.15 · Cited by 5 · see citing works
“…elements at the intersections of bars and overlapping items (Qian & Mitsudo, 2016), or on global mechanisms that relate to periodic gratings (Schrauf et al., 1997; von der Heydt, Peterhans, & Dursteler, 1992) or amodal-integrated representations of fragmented bars (Ban et al., 2013; Sugita, 1999).”
BackgroundAgostino, C. S., Merkel, C., Ball, F., Vavra, P., Hinrichs, H., & Noesselt, T. (2022) · Human Brain Mapping · doi:10.1002/hbm.26123 · Cited by 4 · see citing works
Representing dynamic stimulus information during occlusion
Maarseveen, J., Paffen, C. L. E., Verstraten, F. A. J., & Hogendoorn, H. (2017) · Vision Research · doi:10.1016/j.visres.2017.05.012 · Cited by 4 · see citing works
“This shows that dynamic properties of the stimulus can be represented automatically without any focused effort to maintain the information, similar to the results reported in earlier studies (Ban et al., 2013; Murray et al., 2006).”
InfluentialMomohara, S., & Aoki, S. (2020) · International Journal of Oral-Medical Sciences · doi:10.5466/ijoms.18.277 · Cited by 4 · see citing works
Framing Effects in Object Perception
Ivy, S., & Mroczko-Wąsowicz, A. (2025) · Review of Philosophy and Psychology · doi:10.1007/s13164-024-00763-8 · Cited by 3 · see citing works
“…(as opposed to cognitive) inso-far as amodal completion is performed in the early stages of perceptual processing through the primary visual cortex (Ban et al. 2013; Bushnell et al. 2011; Emmanouil and Ro 2014; Hazenberg et al. 2014; Lee et al. 2012; Pan et al. 2012; Scherzer and Ekroll 2015).”
Against the very idea of a perceptual belief
Helton, G., & Nánay, B. (2022) · Analytic Philosophy · doi:10.1111/phib.12277 · Cited by 3 · see citing works
A detailed theory of thalamic and cortical microcircuits for predictive visual inference
George, D., Lázaro-Gredilla, M., Lehrach, W. P., Dedieu, A., Zhou, G., & Marino, J. (2025) · Science Advances · doi:10.1126/sciadv.adr6698 · Cited by 3 · see citing works
Khazaie, V. R., Akhavanpour, A., & Ebrahimpour, R. (2020) · Iranian Conference on Machine Vision and Image Processing · doi:10.1109/mvip49855.2020.9116887 · Cited by 3 · see citing works
“It is found by using topographic mapping with simple geometric shapes that the occluded region of the shape is represented topographically in human primary visual areas like V1 and V2 [4].”
BackgroundInfluentialAmodal Completion: Mental Imagery or 3D Modeling?
Gauker, C. (2024) · Review of Philosophy and Psychology · doi:10.1007/s13164-024-00740-1 · Cited by 2 · see citing works
“Ban et al. (2013) show participants an animated display in which a colored wedge rotates around a center point, with the sharp point of the wedge pointed at the center.”
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0031 · Cited by 2 · see citing works
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0004 · Cited by 2 · see citing works
Amodal completion across the brain: The impact of structure and knowledge
Thielen, J., van Leeuwen, T. M., Hazenberg, S. J., Wester, A. Z. L., de Lange, F. P., & van Lier, R. (2024) · Journal of Vision · doi:10.1167/jov.24.6.10 · Cited by 1 · see citing works
“Nonetheless, other investigations did report EVC involvement (Rauschenberger et al., 2006; Hulme & Zeki, 2007; Ban et al., 2013; Erlikhman & Caplovitz, 2017; de Haas & Schwarzkopf, 2018).”
BackgroundImproving Animal Recognition Accuracy using Image De-fencing
Kawanaka, K., Felicia, R. M., Tanioka, H., Sano, M., Matsuura, K., & Ueta, T. (2019) · IIAI International Congress on Advanced Applied Informatics (IIAI-AAI) · doi:10.1109/iiai-aai.2019.00114 · Cited by 1 · see citing works
“It is known that human beings can recognize objects by removing obstacles and supplementing the removed areas in the brain [1].”
BackgroundMental Imagery versus Imagination
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0022 · Cited by 1 · see citing works
Bistable perception of symbolic numbers
Luo, J., Yokoi, I., Dumoulin, S. O., & Takemura, H. (2024) · Journal of Vision · doi:10.1167/jov.24.9.12 · Cited by 1 · see citing works
“This phenomenon of visual completion has been studied intensively in psychophysics (Nakayama, Shimojo, & Silverman, 1989; Shimojo & Nakayama, 1990; Tse, 1999), physiology (Albright & Stoner, 2002), and neuroimaging (Ban et al., 2013; Thielen, Bosch, van Leeuwen, van Gerven, & van Lier, 2019).”
MethodReport Decoding Sound and Imagery Content in Early Visual Cortex
Vetter, P., Smith, F. W., & Muckli, L. F. (2014) · Cited by 1 · see citing works
Testing for functional organization of three-dimensional surface tilt encoding within visual cortexPreprint
Rideaux, R., & Welchman, A. E. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.01.15.907683
Ushitani, T. (2020) · Springer eBooks · doi:10.1007/978-3-319-47829-6_1065-1
Ushitani, T. (2022) · Springer eBooks · doi:10.1007/978-3-319-55065-7_1065
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0002
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0008
Mental Imagery in Psychology and Neuroscience
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0001
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.002.0007
Top-Down Influences on Perception and Mental Imagery
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0011
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0006
Perception/Mental Imagery Mixed Cases
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0009
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0010
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0012
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0003
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0005
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0017
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0024
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0029
Sensory Substitution and Echolocation
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0015
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0018
Sense Modalities in Mental Imagery
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0014
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0028 · Cited by 1 · see citing works
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.002.0003
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0020
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0016
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0025
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0021
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0027
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0023
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0026
Clinical Applications of Mental Imagery
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0030
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0007
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.002.0006
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.002.0005
Nánay, B. (2023) · Oxford University Press eBooks · doi:10.1093/oso/9780198809500.003.0019
Brands, A. M., Lange, G., & Groen, I. I. A. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.04.03.646752
“…have been shown to benefit from temporal adaptation, namely object recognition under noise (Vinken et al., 2020; Brands et al., 2024), under occlusion (Ban et al., 2013) and in the context of novelty detection (Dragoi et al., 2002; Kohn, 2007; Whitmire and Stanley, 2016; Weber and Fairhall, 2019).”
An fMRI Dataset on Occluded Image Interpretation for Human Amodal Completion Research
Li, B., Tong, L., Zhang, C., Chen, P., Cao, L., Gao, H., Yu, Z., Wang, L., & Yan, B. (2025) · Scientific Data · doi:10.1038/s41597-025-05414-w
“This phenomenon, known as amodal completion (the unconscious perception of complete mental representations for occluded objects) 1–3 , is not only crucial in everyday life but also plays a significant role in specialized fields 4,5 .”
- Authors
- Ban, H., Yamamoto, H., Hanakawa, T., Urayama, S.-I., Aso, T., Fukuyama, H., & Ejima, Y.
- Affiliations
- Japan Society for the Promotion of Science (JP)
- Kyoto University (JP)
- National Center of Neurology and Psychiatry (JP)
- Kyoto Institute of Technology (JP)
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2013-10-23
- Volume
- 33
- Issue
- 43
- Pages
- 16992-17007
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC6618442
- OpenAlex
- W1998816141
- Cited by
- 80
- References
- 79
- Citations by year
- 2025: 4 · 2024: 4 · 2023: 38 · 2022: 5 · 2021: 3 · 2020: 6 · 2019: 4 · 2018: 3 · 2017: 4 · 2016: 2 · 2015: 2 · 2014: 5
- FWCI
- 1.64
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Journal of Neuroscience (publishedVersion, cc-by-nc-sa)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Institutional Repositories DataBase (IRDB) (submittedVersion)
- Kyoto University Research Information Repository (Kyoto University) (submittedVersion)
- Funders
- Ministry of Education, Culture, Sports, Science and Technology (23135517)
- Japan Society for the Promotion of Science
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Visual perception and processing mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Face Recognition and Perception
- Physical Sciences › Computer Science › Computer Vision and Pattern Recognition › Visual Attention and Saliency Detection
- Keywords
- occlusion, topographic representation, primary visual cortex, V2, object completion, spatio-temporal context, object rotation, illusory perception, luminance enhancement, early visual processing
- MeSH terms
- Adult, Brain Mapping, Female, Humans, Magnetic Resonance Imaging, Male, Visual Cortex, Visual Perception
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