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Introduction | p. 1 |
Introduction and overview | p. 1 |
Neurons | p. 2 |
Neurons in a network | p. 2 |
Synaptic modification | p. 4 |
Long-Term Potentiation and Long-Term Depression | p. 7 |
Distributed representations | p. 11 |
Definitions | p. 11 |
Advantages of different types of coding | p. 12 |
Neuronal network approaches versus connectionism | p. 13 |
Introduction to three neuronal network architectures | p. 14 |
Systems-level analysis of brain function | p. 16 |
The fine structure of the cerebral neocortex | p. 21 |
The fine structure and connectivity of the neocortex | p. 21 |
Excitatory cells and connections | p. 21 |
Inhibitory cells and connections | p. 23 |
Quantitative aspects of cortical architecture | p. 25 |
Functional pathways through the cortical layers | p. 27 |
The scale of lateral excitatory and inhibitory effects, and the concept of modules | p. 29 |
Backprojections in the cortex | p. 30 |
Architecture | p. 30 |
Learning | p. 31 |
Recall | p. 33 |
Semantic priming | p. 34 |
Attention | p. 34 |
Autoassociative storage, and constraint satisfaction | p. 34 |
The primary visual cortex | p. 36 |
Introduction and overview | p. 36 |
Retina and lateral geniculate nuclei | p. 37 |
Striate cortex: Area V1 | p. 43 |
Classification of V1 neurons | p. 43 |
Organization of the striate cortex | p. 45 |
Visual streams within the striate cortex | p. 48 |
Computational processes that give rise to V1 simple cells | p. 49 |
Linsker's method: Information maximization | p. 50 |
Olshausen and Field's method: Sparseness maximization | p. 53 |
The computational role of V1 for form processing | p. 55 |
Backprojections to the lateral geniculate nucleus | p. 55 |
Extrastriate visual areas | p. 57 |
Introduction | p. 57 |
Visual pathways in extrastriate cortical areas | p. 57 |
Colour processing | p. 61 |
Trichromacy theory | p. 61 |
Colour opponency, and colour contrast: Opponent cells | p. 61 |
Motion and depth processing | p. 65 |
The motion pathway | p. 65 |
Depth perception | p. 67 |
The parietal cortex | p. 70 |
Introduction | p. 70 |
Spatial processing in the parietal cortex | p. 70 |
Area LIP | p. 71 |
Area VIP | p. 73 |
Area MST | p. 74 |
Area 7a | p. 74 |
The neuropsychology of the parietal lobe | p. 75 |
Unilateral neglect | p. 75 |
Balint's syndrome | p. 77 |
Gerstmann's syndrome | p. 79 |
Inferior temporal cortical visual areas | p. 81 |
Introduction | p. 81 |
Neuronal responses in different areas | p. 81 |
The selectivity of one population of neurons for faces | p. 83 |
Combinations of face features | p. 84 |
Distributed encoding of object and face identity | p. 84 |
Distributed representations evident in the firing rate distributions | p. 85 |
The representation of information in the responses of single neurons to a set of stimuli | p. 90 |
The representation of information in the responses of a population of inferior temporal visual cortex neurons | p. 94 |
Advantages for brain processing of the distributed representation of objects and faces | p. 98 |
Should one neuron be as discriminative as the whole organism, in object encoding systems? | p. 103 |
Temporal encoding in the spike train of a single neuron | p. 105 |
Temporal synchronization of the responses of different cortical neurons | p. 108 |
Conclusions on cortical encoding | p. 111 |
Invariance in the neuronal representation of stimuli | p. 112 |
Size and spatial frequency invariance | p. 112 |
Translation (shift) invariance | p. 113 |
Reduced translation invariance in natural scenes | p. 113 |
A view-independent representation of objects and faces | p. 115 |
Face identification and face expression systems | p. 118 |
Learning in the inferior temporal cortex | p. 120 |
Cortical processing speed | p. 122 |
Conclusions | p. 125 |
Visual attentional mechanisms | p. 126 |
Introduction | p. 126 |
The classical view | p. 126 |
The spotlight metaphor and feature integration theory | p. 126 |
Computational models of visual attention | p. 129 |
Biased competition - single cell studies | p. 132 |
Neurophysiology of attention | p. 133 |
The role of competition | p. 135 |
Evidence of attentional bias | p. 136 |
Non-spatial attention | p. 136 |
High-resolution buffer hypothesis | p. 139 |
Biased competition - fMRI | p. 140 |
Neuroimaging of attention | p. 140 |
Attentional effects in the absence of visual stimulation | p. 141 |
The computational role of top-down feedback connections | p. 142 |
Neural network models | p. 145 |
Introduction | p. 145 |
Pattern association memory | p. 145 |
Architecture and operation | p. 146 |
The vector interpretation | p. 149 |
Properties | p. 150 |
Prototype extraction, extraction of central tendency, and noise reduction | p. 151 |
Speed | p. 151 |
Local learning rule | p. 152 |
Implications of different types of coding for storage in pattern associators | p. 158 |
Autoassociation memory | p. 159 |
Architecture and operation | p. 160 |
Introduction to the analysis of the operation of autoassociation networks | p. 161 |
Properties | p. 163 |
Use of autoassociation networks in the brain | p. 170 |
Competitive networks, including self-organizing maps | p. 171 |
Function | p. 171 |
Architecture and algorithm | p. 171 |
Properties | p. 173 |
Utility of competitive networks in information processing by the brain | p. 178 |
Guidance of competitive learning | p. 180 |
Topographic map formation | p. 182 |
Radial Basis Function networks | p. 187 |
Further details of the algorithms used in competitive networks | p. 188 |
Continuous attractor networks | p. 192 |
Introduction | p. 192 |
The generic model of a continuous attractor network | p. 195 |
Learning the synaptic strengths between the neurons that implement a continuous attractor network | p. 196 |
The capacity of a continuous attractor network | p. 198 |
Continuous attractor models: moving the activity packet of neuronal activity | p. 198 |
Stabilization of the activity packet within the continuous attractor network when the agent is stationary | p. 202 |
Continuous attractor networks in two or more dimensions | p. 203 |
Mixed continuous and discrete attractor networks | p. 203 |
Network dynamics: the integrate-and-fire approach | p. 204 |
From discrete to continuous time | p. 204 |
Continuous dynamics with discontinuities | p. 205 |
Conductance dynamics for the input current | p. 207 |
The speed of processing of one-layer attractor networks with integrate-and-fire neurons | p. 209 |
The speed of processing of a four-layer hierarchical network with integrate-and-fire attractor dynamics in each layer | p. 212 |
Spike response model | p. 215 |
Network dynamics: introduction to the mean field approach | p. 216 |
Mean-field based neurodynamics | p. 218 |
Population activity | p. 218 |
A basic computational module based on biased competition | p. 220 |
Multimodular neurodynamical architectures | p. 221 |
Interacting attractor networks | p. 224 |
Error correction networks | p. 228 |
Architecture and general description | p. 229 |
Generic algorithm (for a one-layer network taught by error correction) | p. 229 |
Capability and limitations of single-layer error-correcting networks | p. 230 |
Properties | p. 234 |
Error backpropagation multilayer networks | p. 236 |
Introduction | p. 236 |
Architecture and algorithm | p. 237 |
Properties of multilayer networks trained by error backpropagation | p. 238 |
Biologically plausible networks | p. 239 |
Reinforcement learning | p. 240 |
Contrastive Hebbian learning: the Boltzmann machine | p. 241 |
Models of invariant object recognition | p. 243 |
Introduction | p. 243 |
Approaches to invariant object recognition | p. 244 |
Feature spaces | p. 244 |
Structural descriptions and syntactic pattern recognition | p. 245 |
Template matching and the alignment approach | p. 247 |
Invertible networks that can reconstruct their inputs | p. 248 |
Feature hierarchies | p. 249 |
Hypotheses about object recognition mechanisms | p. 253 |
Computational issues in feature hierarchies | p. 257 |
The architecture of VisNet | p. 258 |
Initial experiments with VisNet | p. 266 |
The optimal parameters for the temporal trace used in the learning rule | p. 274 |
Different forms of the trace learning rule, and their relation to error correction and temporal difference learning | p. 275 |
The issue of feature binding, and a solution | p. 284 |
Operation in a cluttered environment | p. 295 |
Learning 3D transforms | p. 301 |
Capacity of the architecture, and incorporation of a trace rule into a recurrent architecture with object attractors | p. 307 |
Vision in natural scenes--effects of background versus attention | p. 313 |
Synchronization and syntactic binding | p. 319 |
Further approaches to invariant object recognition | p. 320 |
Processes involved in object identification | p. 321 |
The cortical neurodynamics of visual attention - a model | p. 323 |
Introduction | p. 323 |
Physiological constraints | p. 324 |
The dorsal and ventral paths of the visual cortex | p. 324 |
The biased competition hypothesis | p. 326 |
Neuronal receptive fields | p. 327 |
Architecture of the model | p. 328 |
Overall architecture of the model | p. 328 |
Formal description of the model | p. 331 |
Performance measures | p. 336 |
Simulations of basic experimental findings | p. 336 |
Simulations of single-cell experiments | p. 337 |
Simulations of fMRI experiments | p. 339 |
Object recognition and spatial search | p. 341 |
Dynamics of spatial attention and object recognition | p. 343 |
Dynamics of object attention and visual search | p. 345 |
Evaluation of the model | p. 348 |
Spatial attention and object attention | p. 348 |
Translation-invariant object recognition | p. 350 |
Contributions and limitations | p. 351 |
Visual search: Attentional neurodynamics at work | p. 353 |
Introduction | p. 353 |
Simple visual search | p. 354 |
Visual search of hierarchical patterns | p. 358 |
The spatial resolution hypothesis | p. 358 |
Neurodynamics of the resolution hypothesis | p. 361 |
Visual search in the framework of the resolution hypothesis | p. 363 |
Visual conjunction search | p. 369 |
The binding problem | p. 369 |
The time course of conjunction search: experimental evidence | p. 371 |
Extension of the computational cortical architecture | p. 373 |
Computational results | p. 376 |
Conclusion | p. 381 |
A computational approach to the neuropsychology of visual attention | p. 383 |
Introduction | p. 383 |
The neglect syndrome | p. 383 |
A model of visual spatial neglect | p. 384 |
Spatial cueing effect on neglect | p. 388 |
Extinction and visual search | p. 390 |
Effect on neglect of top-down knowledge about objects | p. 392 |
Hierarchical patterns - neuropsychology | p. 398 |
Conjunction search - neuropsychology | p. 400 |
Simulations and predictions | p. 400 |
Experimental test of the predictions in human subjects | p. 401 |
Conclusion | p. 403 |
Outputs of visual processing | p. 404 |
Visual outputs to Short Term Memory systems | p. 406 |
Prefrontal cortex short term memory networks, and their relation to temporal and parietal perceptual networks | p. 406 |
Computational details of the model of short term memory | p. 409 |
Computational necessity for a separate, prefrontal cortex, short term memory system | p. 412 |
Role of prefrontal cortex short term memory systems in visual search and attention | p. 412 |
Synaptic modification is needed to set up but not to reuse short term memory systems | p. 413 |
Visual outputs to Long Term Memory systems in the brain | p. 413 |
Effects of damage to the hippocampus and connected structures on object-place and episodic memory | p. 414 |
Neurophysiology of the hippocampus and connected areas | p. 415 |
Hippocampal models | p. 418 |
The perirhinal cortex, recognition memory, and familiarity | p. 421 |
Visual stimulus-reward association, emotion, and motivation | p. 424 |
Emotion | p. 425 |
Reward is not processed in the temporal cortical visual areas | p. 429 |
Why the reward and punishment associations of stimuli are not represented early in information processing in the primate brain | p. 430 |
Amygdala | p. 434 |
Orbitofrontal cortex | p. 439 |
Effects of mood on memory and visual processing | p. 447 |
Output to object selection and action systems | p. 448 |
Visual search | p. 452 |
Visual outputs to behavioral response systems | p. 453 |
Multimodal representations in different brain areas | p. 453 |
Visuo-spatial scratchpad, and change blindness | p. 454 |
Conscious visual perception | p. 454 |
Principles and Conclusions | p. 456 |
Transform invariance in the inferior temporal visual cortex | p. 456 |
Representation of information in IT | p. 456 |
IT information processing is fast | p. 457 |
Continuous neuronal dynamics allows fast network processing | p. 457 |
Hierarchical feature analysis | p. 457 |
Trace learning rule for invariant representations | p. 459 |
Spatial feature binding by feature combination neurons | p. 460 |
IT provides a representation for later memory networks | p. 461 |
Face expression and object motion | p. 462 |
Attentional mechanisms | p. 462 |
Visual search | p. 464 |
Egocentric vs allocentric representations | p. 464 |
Short term memory as the controller of attention | p. 465 |
Output to object selection and action systems | p. 466 |
'What' versus 'where' processing streams | p. 466 |
Short term memory must be separated from perception | p. 467 |
Backprojections must be weak | p. 468 |
Long-term potentiation and short-term memory | p. 469 |
"Executive control" by the prefrontal cortex | p. 469 |
Reward processing occurs after object identification | p. 470 |
Effects of mood on memory and visual processing | p. 471 |
Visual outputs to Long Term Memory systems | p. 471 |
Episodic memory and the operation of mixed discrete and continuous attractor networks | p. 472 |
Visual outputs to behavioural response systems | p. 472 |
Multimodal representations in different brain areas | p. 472 |
Visuo-spatial scratchpad and change blindness | p. 472 |
Invariant object recognition and attention | p. 473 |
Conscious visual perception | p. 473 |
Attention - future directions | p. 473 |
Integrated approaches to understanding vision | p. 475 |
Apostasis | p. 475 |
Introduction to linear algebra for neural networks | p. 477 |
Vectors | p. 477 |
The inner or dot product of two vectors | p. 477 |
The length of a vector | p. 478 |
Normalizing the length of a vector | p. 479 |
The angle between two vectors: the normalized dot product | p. 479 |
The outer product of two vectors | p. 480 |
Linear and non-linear systems | p. 481 |
Linear combinations of vectors, linear independence, and linear separability | p. 482 |
Application to understanding simple neural networks | p. 484 |
Capability and limitations of single-layer networks: linear separability and capacity | p. 484 |
Non-linear networks: neurons with non-linear activation functions | p. 487 |
Non-linear networks: neurons with non-linear activations | p. 488 |
Information theory | p. 490 |
Basic notions | p. 490 |
The information conveyed by definite statements | p. 491 |
Information conveyed by probabilistic statements | p. 491 |
Information sources, information channels, and information measures | p. 492 |
The information carried by a neuronal response and its averages | p. 494 |
The information conveyed by continuous variables | p. 496 |
The information carried by neuronal responses | p. 498 |
The limited sampling problem | p. 498 |
Correction procedures for limited sampling | p. 500 |
The information from multiple cells: decoding procedures | p. 501 |
Information in the correlations between the spikes of different cells | p. 504 |
Information theory results | p. 507 |
Temporal codes versus rate codes within the spike train of a single neuron | p. 507 |
The speed of information transfer from single neurons | p. 509 |
The information from multiple cells: independent information versus redundancy across cells | p. 512 |
The information from multiple cells: the effects of cross-correlations between cells | p. 514 |
Conclusions | p. 517 |
Information theory terms--a short glossary | p. 518 |
References | p. 520 |
Index | p. 565 |
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