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. 2018 Aug 2;9:1027. doi: 10.3389/fphys.2018.01027

FIGURE 2.

FIGURE 2

Wiring diagram for the inferior frontal system controlling tactile discrimination behaviors in Octopus as proposed by Young (1983, 1991). This representation neglects positive sensory input associated with acceptance behaviors and instead depicts input from noxious sensory fibers from the arms that enter the inferior frontal lobes from the cerebro-brachial connective (Budelmann and Young, 1985). The stylistic circuit is drawn to highlight similarities in neural connectivity as previously described for learning and memory circuits of the superior frontal and vertical lobes (Hochner and Shomrat, 2013; Shomrat et al., 2015). Large projection neurons are depicted as unfilled rounded rectangles and in the subfrontal lobe (sfl), lateral inferior frontal lobe (lifl) and posterior buccal lobe (pbl) there are two principal types – acceptance (A) and rejection (R) – which selectively project to one another. The projection neurons associated with tactile discrimination in the median inferior frontal lobe (mifl) project to both the sfl and lifl where they terminate on small amacrine interneurons (a). Although this projection neuron is represented as having axon branches this is simply diagrammatic and it is most likely that there are multiple types of these neurons that have different innervation densities in the sfl and lifl. Synapses are represented as filled circles (inhibitory) or unfilled circles (excitatory). Noxious sensory fibers selectively innervate projection neurons in the lifl and mifl. Electric shock would be expected to activate R projection neurons in the lifl and depending on their state of activation by local amacrine interneurons, these R neurons would activate R projection neurons in the pbl and lead to a reject behavior. Noxious sensory fibers projecting to the mifl would cause a cascade of modulatory activity in the sfl and lifl that would robustly regulate motor behavior. With tactile discrimination training (using either or both positive and negative reinforcements) this circuitry outlined here would enable behavioral learning and memory formation that could readily increase efficacy of motor actions.