Добавил:
kiopkiopkiop18@yandex.ru t.me/Prokururor I Вовсе не секретарь, но почту проверяю Опубликованный материал нарушает ваши авторские права? Сообщите нам.
Вуз: Предмет: Файл:
Скачиваний:
1
Добавлен:
28.03.2026
Размер:
2.5 Mб
Скачать

what percept the network is “seeing” means that a solution is assured for all iterations in terms of one of ten categorical cells. The second input feature is that of habituation. While this factor may not have been necessary to achieve rivalry in some cases, its presence can be argued as logical on the basis of single-cell electrophysiological studies where habituation is a common observation of recordings from cortex.

The issue of bottom-up or top-down theories of rivalry has not been directly addressed by this neural network attempt. An inspection of figure 18.1 shows that there is segregation of the visual information through the first couple of stages and that interaction between the two eyes really occurs only at the penultimate stage, just before the Kohonen classifier. In the current model it is the activity in the early extrastriate cortex that causes rivalry for dominance at the late extrastriate level. The result of the late extrastriate decision on the winner is fed back to the activity levels at the early extrastriate layer. Thus the current model embraces aspects of both theories. Also, the model does not really mimic the human visual system, where binocular interaction occurs at the level of striate cortex and where pipelining of information to “categorical cortex” in the inferotemporal regions occurs via several more synaptic steps utilizing predominantly binocular neurons.

The model does not give a prediction regarding cortical localization or hemispheric switching (Miller et al., 2000), largely because there is only one higher cortical region in the model. However, it is clear from the way in which attention was (somewhat artificially) implemented that the model would respond to neural firing changes accompanying such a hemispheric stimulation. Also, it is clear that rivalry does not require the presence of two hemispheres to compete, because the properties of the categorical cells are sufficient.

The one feature of this model that was not an input was not engineered, and yet gave plausibility to the top-down approach was the emergence of a principle of prediction of rivalry which was validated with high reliability. Consider the system from the bottom up and from the top down. A priori there is nothing from the bottom-up direction that allows a prediction of rivalry in particular choices of stimuli presented to the two eyes. While one might guess that stimuli with like contours are less likely to rival, it is only when the system has actually learned and categorized the set of stimulus images into one of ten categorical cells that one can predict with almost complete certainty which pairs of stimuli will not rival. The human visual system also shows a preference for images that it has learned. Yu and Blake (1992) showed that in rivalry, a face stimulus

350

D. P. Crewther and colleagues

predominated more than a pattern stimulus equated for spatial frequency, luminance, and contrast.

The Kohonen classifier is interesting in that neighboring classifiers are closer in terms of similarity than are more distant ones. This was seen in some of the rivalry cases where the expected categorical cells did not participate, but a neighboring categorical cell was involved. Also, rivalry comparison at high and low contrast showed, for example, that a stimulus may have been classified as GC#3 at low contrast but GC#4 at high contrast.

In principle, the model could be extended to provide an account of other bistable phenomena, where the low-level part of the network receives invariant input. Practically, this could be achieved through the training of images with binocular disparity (e.g., the two alternate 3D views of the Necker cube) and then testing with 2D (zero disparity) stimuli presented to the two eyes.

In conclusion, the results of this and other computational studies of binocular rivalry indicate that both bottom-up and top-down approaches yield rivalry behaviors that mimic well parts of the experimental literature. Thus it is likely that composite models, where rivalry may emerge from interactions at several levels of neural processing, need to be investigated.

REFERENCES

Alais, D., and Blake, R. (1999). Grouping visual features during binocular rivalry. Vision Research, 39, 4341–4353.

Andrews, T. J., and Blakemore, C. (1999). Form and motion have independent access to consciousness. Nature Neuroscience, 2, 405–406.

Blake, R. (1977). Threshold conditions for binocular rivalry. Journal of Experimental Psychology: Human Perception and Performance, 3, 251–257.

Blake, R. (1989). A neural theory of binocular rivalry. Psychological Review, 96, 145–167.

Bullier, J. (2001). Integrated model of visual processing. Brain Research Review, 36, 96–107.

Dayan, P. (1998). A hierarchical model of binocular rivalry. Neural Computation, 10, 1119–1135.

Epstein, R., and Kanwisher, N. (1998). A cortical representation of the local visual environment. Nature, 392, 598–601.

Fox, R., and Herrmann, J. (1967). Stochastic properties of binocular rivalry alternations.

Perception and Psychophysics, 2, 432–436.

Fox, R., and Rasche, F. (1969). Binocular rivalry and reciprocal inhibition. Perception and Psychophysics, 5, 215–217.

351

A Neural Network Model of Top-Down Rivalry

Hupé, J. M., James, A. C., Girard, P., Lomber, S. G., Payne, B. R., and Bullier, J. (2001). Feedback connections act on the early part of the responses in monkey visual cortex. Journal of Neurophysiology, 85, 134–145.

Kanwisher, N., McDermott, J., and Chun, M. M. (1997). The fusiform face area: A module in human extrastriate cortex specialized for face perception. Journal of Neuroscience, 17, 4302–4311.

Kovács, I., Papathomas, T. V., Yang, M., and Fehér, A. (1996). When the brain changes its mind: Interocular grouping during binocular rivalry. Proceedings of the National Academy of Sciences of the United States of America, 93, 15508–15511.

Lamme, V. A., Super, H., Landman, R., Roelfsema, P. R., and Spekreijse, H. (2000). The role of primary visual cortex (V1) in visual awareness. Vision Research, 40, 1507–1521.

Lamme, V. A., Van Dijk, B. W., and Spekreijse, H. (1992). Texture segregation is processed by primary visual cortex in man and monkey. Evidence from VEP experiments. Vision Research, 32, 797–807.

Lee, S. H., and Blake, R. (2002). V1 activity is reduced during binocular rivalry. Journal of Vision, 2, 618–626.

Lehky, S. R. (1988). An astable multivibrator model of binocular rivalry. Perception, 17, 215–228.

Leopold, D. A., and Logothetis, N. K. (1996). Activity changes in early visual cortex reflect monkeys’ percepts during binocular rivalry. Nature, 379, 549–553.

Levelt, W. J. M. (1965). On Binocular Rivalry. Soesterberg, The Netherlands: Institute for Perception RVO-TNO.

Levelt, W. J. M. (1966). The alternation process in binocular rivalry. British Journal of Psychology, 57, 225–238.

Liu, L., Tyler, C. W., and Schor, C. M. (1992). Failure of rivalry at low contrast: Evidence of a suprathreshold binocular summation process. Vision Research, 32, 1471–1479.

Logothetis, N. K. (1998a). Object vision and visual awareness. Current Opinion in Neurobiology, 8, 536–544.

Logothetis, N. K. (1998b). Single units and conscious vision. Philosophical Transactions of the Royal Society of London, B353, 1801–1818.

Logothetis, N. K., Leopold, D. A., and Sheinberg, D. L. (1996). What is rivalling during binocular rivalry? Nature, 380, 621–624.

Logothetis, N. K., and Schall, J. D. (1989). Neuronal correlates of subjective visual perception. Science, 245, 761–763.

Lumer, E. D. (1998). A neural model of binocular integration and rivalry based on the coordination of action-potential timing in primary visual cortex. Cerebral Cortex, 8, 553–561.

Miller, S. M., Liu, G. B., Ngo, T. T., Hooper, G., Riek, S., Carson, R. G., and Pettigrew, J. D. (2000). Interhemispheric switching mediates perceptual rivalry. Current Biology, 10, 383–392.

Mueller, T. J. (1990). A physiological model of binocular rivalry. Vision Neuroscience, 4, 63–73.

352

D. P. Crewther and colleagues

Mueller, T. J., and Blake, R. (1989). A fresh look at the temporal dynamics of binocular rivalry.

Biological Cybernetics, 61, 223–232.

Polonsky, A., Blake, R., Braun, J., and Heeger, D. J. (2000). Neuronal activity in human primary visual cortex correlates with perception during binocular rivalry. Nature Neuroscience, 3, 1153–1159.

Puce, A., Allison, T., Gore, J. C., and McCarthy, G. (1995). Face-sensitive regions in human extrastriate cortex studied by functional MRI. Journal of Neurophysiology, 74, 1192–1199.

Seghier, M., Dojat, M., Delon-Martin, C., Rubin, C., Warnking, J., Segebarth, C., and Bullier, J. (2000). Moving illusory contours activate primary visual cortex: An fMRI study.

Cerebral Cortex, 10, 663–670.

Sugie, N. (1982). Neural models of brightness perception and retinal rivalry in binocular vision. Biological Cybernetics, 43, 13–21.

Super, H., Spekreijse, H., and Lamme, V. A. (2001). Two distinct modes of sensory processing observed in monkey primary visual cortex (V1). Nature Neuroscience, 4, 304–310.

Tanaka, K. (1993). Neuronal mechanisms of object recognition. Science, 262, 685–688.

Tanaka, K. (2003). Columns for complex visual object features in the inferotemporal cortex: Clustering of cells with similar but slightly different stimulus selectivities. Cerebral Cortex, 13, 90–99.

Tong, F., and Engel, S. A. (2001). Interocular rivalry revealed in the human cortical blind-spot representation. Nature, 411, 195–199.

Tong, F., Nakayama, K., Vaughan, J. T., and Kanwisher, N. (1998). Binocular rivalry and visual awareness in human extrastriate cortex. Neuron, 21, 753–759.

Tononi, G., Srinivasan, R., Russell, D. P., and Edelman, G. M. (1998). Investigating neural correlates of conscious perception by frequency-tagged neuromagnetic responses. Proceedings of the National Academy of Sciences of the United States of America, 95, 3198–3203.

Wang, Y., Fujita, I., and Murayama, Y. (2000). Neuronal mechanisms of selectivity for object features revealed by blocking inhibition in inferotemporal cortex. Nature Neuroscience, 3, 807–813.

Wolfe, J. M. (1983). Influence of spatial frequency, luminance, and duration on binocular rivalry and abnormal fusion of briefly presented dichoptic stimuli. Perception, 12, 447–456.

Yu, K., and Blake, R. (1992). Do recognizable figures enjoy an advantage in binocular rivalry?

Journal of Experimental Psychology: Human Perception and Performance, 18, 1158–1173.

353

A Neural Network Model of Top-Down Rivalry

APPENDIX: SAMPLE CASES OF RIVALRY AND NONRIVALRY

Given the overall hypothesis of the study concerning the chance of rivalry—that stimuli which attract the same categorical cell to respond should not rival, while stimuli which associate with different categorical cells during training will rival—several cases exhibit the success (and variations of success) of the hypothesis.

Expect Rivalry—Get Regular Rivalry

Stim 1 versus stim 14. In this situation, with stim 1 presented to one eye and stim 14 presented to the other, rivalry was expected because stim 1 associated with categorical cell 7, while stim 14 associated with categorical cell 3 (see figure 18.3a). The figure shows a graphical representation of the rivaling stimuli as well as a representation across the 100 iterations (Time) of the left and right strength modifiers. Notice the regular alternation between the modifiers, with reciprocal weakening and strengthening (0.1 per iteration) until the point at which the “winner” (bottom graph, triangle symbols) changes identity. At this point the strength of the winning eye is enhanced by a factor (perhaps analogous to attention), with a corresponding drop in strength for the suppressed stimulus.

The overall strength of this factor was arbitrary, but was fixed throughout all of the network calculations. The similarity measure for the left and right eyes is shown in the second trace. It also shows regularity in alternation that is highly correlated with the switch in Kohonen choice (categorical selection) (lower trace). The “winner” is shown as a series of triangular markers taking on one of two states (upper right, lower left). In this case the change in “winner” occurs at precisely the same time as the change in Kohonen choice.

Expect Rivalry—Get Rivalry with Erroneous Percept

Stim 1 versus stim 18. Given the two stimuli, rivalry was expected between categorical cell 7 (stim 1) and categorical cell 9 (stim 18). However, as figure 18.3b shows, though rivalry was observed, it was between categorical cells 7 and 8 (bottom graph—selection), rather than GC7 and GC9. Note that the strength modifiers for the left and right eyes are again mirror

354

D. P. Crewther and colleagues

images around the value 1. Again the rivalry is quite regular, after an initial longer settling period—this appears to be associated with the relatively large change in similarity measure for each eye between the one percept and the other, what one might call “strong rivalry.” It is interesting that while “false” percept was generated, the incorrect categorical cell (GC8 rather then GC9) is a neighboring Kohonen classifier of the expected one, with the implication of some similarity relation between the neighbors. Such erroneous percept (disagreeing with either monocular input) is possible experimentally (Kovács et al., 1996).

Expect Rivalry—Get Irregular Rapid Rivalry Stim 5 (GC6) versus stim 0 (GC0). The lowest graph of figure 18.3c shows a rapid alternation between GC6 and GC0. While the rivalry is between the expected categorical cell representations, the frequency of alternation is irregular. This irregularity possibly arises because of the close similarity measures for the two eyes.

Expect Rivalry—Get Irregular, Infrequent Rivalry Stim 1 (GC7) versus stim 6 (GC0). While the strength modifiers overlap in a fashion very similar to that exhibited in figure 18.3a, the rivalry results are dramatically different. The similarity measures for the two eyes are not at all similar in shape, and the “winner” graph at the extreme bottom of figure 18.3d shows an irregularity in switching. This irregularity is further teased out by the Kohonen selection graph (lowest solid curve, figure 18.3d). The switching between categorical cells is not well predicted by the switching in winner. This possibly is due to the relatively small difference in similarity measure between dominance and suppression states for the two eyes (roughly of amplitude 30, compared with over 100 for figure 18.3a). This might be termed “weak rivalry.”

Failed Rivalry—When Expected

Stim 0 (GC0) versus stim 20 (GC7). One switch in percept near the start of the iteration was observed. After this time, the percept remained stuck at categorical cell 7 (see figure 18.3e). This failure of rivalry (when expected from the difference in categorical cell preference for the two stimuli) can be identified as due to the fact that the similarity measures never approached a crossing point.

355

A Neural Network Model of Top-Down Rivalry

Contributors

David Alais

Department of Physiology and Institute for Biomedical Research School of Medical Science University of Sydney

Sydney, Australia

Timothy J. Andrews

University Laboratory of

Physiology

University of Oxford

Oxford, United Kingdom

Department of Psychology

University of Durham

Durham, United Kingdom

Randolph Blake

Department of Psychology

Vanderbilt University

Nashville, Tennessee

Colin Blakemore

University Laboratory of

Physiology

University of Oxford

Oxford, United Kingdom

Thomas Carlson

Department of Psychology

University of Minnesota

Minneapolis, Minnesota

O. L. Carter

Vision Touch and Hearing

Research Centre

School of Biomedical Sciences

University of Queensland

Brisbane, Australia

Miguel Castelo-Branco Faculty of Medicine Instituto Biomédico de

Investigacão de Luz e Imagem Coimbra, Portugal

Xiangchuan Chen

University of Science and

Technology of China

Department of Neurobiology and

Biophysics

Hefei, Anhui, People’s Republic of

China

Tiffany Conway

Laboratory of Vision Research

Rutgers University

Piscataway, New Jersey

Paul M. Corballis

Center for Cognitive Neuroscience Dartmouth College

Hanover, New Hampshire

D. P. Crewther

Brain Sciences Institute

Swinburne University of

Technology

Melbourne, Australia

S. Crewther

School of Psychological Science

La Trobe University

Melbourne, Australia

Michal Eisenberg

Department of Psychology and Laboratory of Vision Research/Center for Cognitive Science

Rutgers University Piscataway, New Jersey

Andreas K. Engel

Institute of Neurophysiology

and Pathophysiology

University Hospital Hamburg-

Eppendorf

Hamburg, Germany

Robert Fox

Department of Psychology and

Department of Biomedical

Engineering

Kennedy Center Scientist

Vanderbilt University

Nashville, Tennessee

Alan W. Freeman

School of Biomedical Sciences

University of Sydney

Lidcombe, Australia

Itzhak Fried

Division of Neurosurgery and

Department of Psychiatry and

Biobehavioral Sciences

David Geffen School of Medicine

University of California

Los Angeles, California

Functional Neurosurgery Unit

Tel Aviv Medical Center and

Sackler School of Medicine

Tel Aviv University

Tel Aviv, Israel

Pascal Fries

F.C. Donders Centre for Cognitive Neuroimaging

Nijmegen, The Netherlands

Department of Biophysics

University of Nijmegen

The Netherlands

Sheng He

Department of Psychology

University of Minnesota

Minneapolis, Minnesota

Zijiang J. He

Department of Psychological and

Brain Sciences

University of Louisville

Louisville, Kentucky

Ian P. Howard

Centre for Vision Research York University

Toronto, Ontario, Canada

358 Contributors

Jean-Michel Hupé Center for Neural Science New York University New York, New York

R. Jones

School of Psychological Science

La Trobe University

Melbourne, Australia

Christof Koch

Computation and Neural Systems

Program

California Institute of Technology

Pasadena, California

Ilona Kovács

Department of Psychology and Laboratory of Vision Research/Center for Cognitive Science

Rutgers University Piscataway, New Jersey

Gabriel Kreiman

Division of Neurosurgery and

Department of Psychiatry and

Biobehavioral Sciences

David Geffen School of Medicine

University of California

Los Angeles, California

Computational and Neural

Systems Program

California Institute of Technology

Pasadena, California

David A. Leopold Department of Physiology of Cognitive Processes

Max Planck Institute for Biologic Cybernetics

Tübingen, Germany

Unit on Cognitive Neurophysiology and Imaging National Institutes of Health Bethesda, Maryland 20892

Nikos K. Logothetis Department of Physiology of Cognitive Processes

Max Planck Institute for Biologic Cybernetics

Tübingen, Germany

Alexander Maier Department of Physiology of Cognitive Processes

Max Planck Institute for Biologic Cybernetics

Tübingen, Germany

J. Munro

School of Psychological Science

La Trobe University

Melbourne, Australia

Vincent A. Nguyen

School of Biomedical Sciences

University of Sydney

Lidcombe, Australia

Teng Leng Ooi

Department of Basic Sciences

Pennsylvania College of

Optometry

Elkins Park, Pennsylvania

Robert P. O’Shea

Department of Psychology

University of Otago

Dunedin, New Zealand

359 Contributors

Thomas V. Papathomas

Department of Biomedical

Engineering and Laboratory of

Vision Research

Rutgers University

New Brunswick, New Jersey

J. D. Pettigrew

Vision Touch and Hearing

Research Centre

School of Biomedical Sciences

University of Queensland

Brisbane, Australia

T. Price

School of Psychological Science

La Trobe University

Melbourne, Australia

S. Pulis

School of Psychological Science

La Trobe University

Melbourne, Australia

Nava Rubin

Center for Neural Science New York University New York, New York

Frank Sengpiel

University Laboratory of

Physiology

University of Oxford

Oxford, United Kingdom

School of Biosciences

Cardiff University

Cardiff, United Kingdom

Wolf Singer

Max Planck Institut für

Hirnforschung

Frankfurt, Germany

Frank Tong

Department of Psychology

Princeton University

Princeton, New Jersey

Nicholas J. Wade

Department of Psychology

University of Dundee

Dundee, Scotland

Melanie Wilke

Department of Physiology of Cognitive Processes

Max Planck Institute for Biologic Cybernetics

Tübingen, Germany

Hugh R. Wilson

Centre for Vision Research York University

Toronto, Ontario, Canada

360 Contributors