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7. Homologous Laminar Cortical Circuits for All Biological Intelligence  43




                     Fig. 2.7 also shows that bottom-up signals from the LGN use the same modulatory
                  on-center, off-surround network to activate layer 4 in V1 that is used by the top-down
                  attentional feedback pathway. In addition, there is a direct bottom-up excitatory
                  pathway from LGN to layer 4 so that the LGN can activate V1 in response to inputs
                  from the retina. Taken together, the direct LGN-to-4 pathway and the LGN-to-6-to-4
                  modulatory on-center, off-surround network ensure that bottom-up inputs from the
                  LGN to V1 are contrast-normalized at layer 4 cells.
                     The sharing of the layer 6-to-4 modulatory on-center, off-surround network by
                  bottom-up and top-down pathways converts this network into a decision interface
                  where preattentive automatic bottom-up processing and attentive task-selective
                  top-down processing can cooperate and compete to choose the combination of
                  signals that is most salient at any given moment.
                     Such a cooperative-competitive decision interface exists in every granular
                  neocortical area. As a result, a top-down task-selective priming signal from a higher
                  cortical area can propagate through multiple lower cortical areas via their layer 6,
                  which can then activate their layer 6-to-4 modulatory on-center, off-surround
                  networks. In this way, an entire cortical hierarchy may get ready to process incoming
                  bottom-up signals to accommodate the bias imposed by the prime.
                     Fig. 2.7 also shows that layer 2/3 in each cortical area also projects back to layer 6,
                  and then up to layer 4 via the folded feedback network. The horizontal connections in
                  layer 2/3 carry out a variety of functions in different cortical areas. In V2, they carry
                  out perceptual grouping and boundary completion [29], a process whose so-called
                  bipole grouping properties were predicted before the neurophysiological data of
                  von der Heydt et al. were reported [30,31] and which were subsequently extensively
                  modeled by LAMINART (e.g., Refs. [27,28,32,33]). In cognitive processing regions,
                  such as the ventrolateral prefrontal cortex, it has been suggested that such horizontal
                  connections enable learning of categories, also called list chunks,that respond
                  selectively to sequences of items that are stored in working memory [34,35].
                     The development of these horizontal connections begins before birth and
                  continues in response to the statistics of visual environments after birth. The fact
                  that the layer 2/3-to-6-to-4-to-2/3 pathway satisfies the ART matching rule enables
                  this development, as well as that of other cortical circuits, to dynamically

               =
                  circular reaction [17e19], is learned from spatial-to-motor and motor-to-spatial
                  representations at the two adaptive pathways in the model, which are denoted by
                  hemispherical synapses. In particular, the spatial direction vector (DV s ) is adaptively
                  mapped into the motor direction vector (DV m ), thereby carrying out the transformation from
                  visual direction into joint rotation that gives the DIRECT model its name.
                   (Left panel) Adapted with permission from D. Bullock, S. Grossberg, Neural dynamics of planned arm move-
                  ments: emergent invariants and speed-accuracy properties during trajectory formation, Psychological Review 95
                  (1988) 49e90. (Right panel) Reprinted with permission from D. Bullock, S. Grossberg, F.H. Guenther, A self-
                    organizing neural model of motor equivalent reaching and tool use by a multijoint arm, Journal of Cognitive
                                                               Neuroscience 5 (1993) 408e435.
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