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4. Is Computational Neuroscience Separate From Neural Network Theory?    197




                  conditioning [63]. Yet this modeling approach in some ways betrays its one-size-
                  fits-all origins. It is based on distributed representations of concepts when there is a
                  basis for making some models use more localized representations (see Ref. [64]
                  for discussion). Also, the LEABRA equations use both associative learning and
                  error-driven learning at every synapse, rather than using different types of learning
                  for different processes.
                     A second source of many CCN models, particularly in the area of conditioning,
                  is the temporal difference (TD) model of Sutton and Barto [10,11].The TD idea
                  that learning occurs when there is an error inrewardprediction(i.e.,apreviously
                  neutral stimulus becomes rewarding or a previously rewarding stimulus becomes
                  unrewarding) obtained experimental support from results on responses of
                  dopamine neurons in the midbrain [65e67]. Reward prediction was thought to
                  be implemented via dopamine neuron connections to the basal ganglia, and several
                  later variants of the TD model exploited this connection [68e70].These models
                  built on the notion of reinforcement learning with an actor and a critic, a design
                  that is also popular in control engineering (e.g., Ref. [71]).
                     The TD approach is popular with conditioning researchers because it is built
                  around a single and easily understandable principle, namely, maximization of
                  predicted future reward. Yet its very simplicity, with the implication that there is
                  a unique locus in the brain that controls Pavlovian learning, limits the predictive
                  applicability of this approach unless it is extended to incorporate principles that
                  suggest roles for regions not included in these articles, such as amygdala and
                  prefrontal cortex (see Ref. [72]; for a review).
                     Other CCN models come not from previous simpler neural models but from
                  neural elaborations of previous nonneural models from mathematical psychology.
                  This approach has been particularly fruitful in the area of category learning
                  [73e75]. Interestingly Love and Gureckis [75] noted the kinship of their model
                  with the adaptive resonance theory of categorization [51], which was based on
                  associative learning combined with lateral inhibition and opponent processing.
                     Finally, there are a number of CCN models which are refinements or extensions
                  of more abstract models that arose before current data were available but embodied
                  network principles based on cognitive requirements. The cognitive requirements
                  these networks were designed to fulfill are frequently based on complementary
                  pairs, such as learning new inputs without forgetting old ones, or processing both
                  boundaries and interiors of visual scenes (e.g., [76]). One example of this type of
                  CCN model is Grossberg and Versace [77], which extends the previously more
                  abstract adaptive resonance model to incorporate neural data about corticothalamic
                  interactions and the role of acetylcholine. Another example is. [78] which is built
                  on previously more abstract conditioning models and incorporates neural data
                  about dopaminergic prediction error and different roles for the amygdala and
                  orbitofrontal cortex.
                     My answer to the question posed by the title of this section is, no, computational
                  neuroscience, or at least computational cognitive neuroscience, is not separate from
                  neural network theory. That is, CCN is not a fundamental conceptual break from
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