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Bernal S D_2010.pdf - University of Plymouth

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2.2. HSGH-LEVEL FEEDBACK<br />

feature analysis modules. Results show this type <strong>of</strong> network can learn invariance to translation,<br />

size, rotation and contrast, achieving good generalization to new objects even using only a small<br />

training datasel (Wiskott and Sejnowski 2(H)2, Malhias et al. 2008).<br />

The slow feature principle is closely related to the trace rule employed in the Visnet model<br />

(Rolls and Milward 2000) previously described. In contrast, the main advantage <strong>of</strong> slow feature<br />

analysis is thai it is not limited to extracting a single invariant representaiion. i.e. object iden­<br />

tity, but also maintains a structured representation <strong>of</strong> other parameters such as object position,<br />

rotation angles and lighting direction.<br />

2.2 High-level feedback<br />

The previous section acts as an introduction to the visual system and in particular to object<br />

recognition. This provides the context to discuss the role <strong>of</strong> high-level feedback in percep­<br />

tion, exposing many <strong>of</strong> the phenomena which remain unexplained and challenging some <strong>of</strong> the<br />

existing classical concepts. To avoid misinterpreialion, wc define feedback as activity origi­<br />

nating in a high-level region targeting a lower-level region, which therefore excludes inlralevel<br />

inlerluminar activity.<br />

2.2.1 Experimental evidence<br />

2.2.1.1 Anatomical pcrspeclivc<br />

From the anatomical point <strong>of</strong> view, feedback connections extensively outnumber feedforward<br />

sensory pathways (Felleman and Van Essen 1991, Macknik and Marlinez-C'onde 2007). The<br />

great majority <strong>of</strong> connections between regions shown in Figure 2. la are reciprocal, which pro­<br />

vides lower areas with massive feedback from higher cortical areas. For example, cat LGN<br />

intemeurons receive 25Cf <strong>of</strong> their inputs from the retina, while 37% come from cortex; for<br />

LGN relay cells, the corresponding percentages are 12% and 5S% (Montero 1991). The same is<br />

true for thalamic relay nucleus (TRN), which mediates the transfer <strong>of</strong> information to the cortex,<br />

where Ihe largest anatomical projection is from connections <strong>of</strong> cortical feedback and not die<br />

ascending collaterals <strong>of</strong> relay cells (Sillito and Jones 2008). For LGN relay cells it is generally<br />

believed that feedback exerts a mtnluiatory influence, whereas cortical feedback to TRN is more<br />

24

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