, 2001). The functional meanings of these are only beginning to be understood (Freed, 2000). A case in point is the expression of differing sets of regulation of G protein signaling (RGS) proteins, which control the kinetics of the response to synaptic input in ON bipolar cells

(Cao et al., 2012). Another is a type of bipolar cell that generates Na+ action potentials. Na+ currents have been known to occur from studies of many retinas, but their functions are unclear (Ichinose and Lukasiewicz, 2007; Ichinose et al., 2005; Ma et al., 2005; Zenisek et al., 2001). In the ground squirrel, the structurally defined bipolar cell termed Selleck Docetaxel cb5b has a large tetrodotoxin (TTX)-sensitive Na+ current. These cells

signal the onset of a light step with a few all-or-nothing action potentials (Figure 4). In response to a continually graded noise stimulus (more closely representing a natural scene), they generate both graded and spiking responses, the spikes occurring with millisecond precision. The cells select for stimulus sequences in which transitions to light are preceded by a period of darkness. Their axon terminals costratify with the dendrites of a specific group of ganglion cells, and these ganglion cells encode light selleck screening library onset with a short latency burst of spikes. It thus appears that this bipolar cell trades the bandwidth inherent in graded signaling for spikes that can elicit a rapid and reliable response in transient-type

ganglion cells (Saszik and DeVries, 2012). The central structural characteristic Parvulin that defines the ∼12 types of bipolar cells is the level of the inner plexiform layer at which their axons terminate. In other words, the bipolar cells receive input from all of the cones within their reach, as just described, but they terminate on very restricted sets of postsynaptic partners. Distinction of functional types on this basis is confirmed by molecular differences that correlate with types that have been defined in this way. The specificity is again confirmed by the fact that different sets of ganglion cells (as well as amacrine cells) costratify with them. These, too, represent distinct types: they have different central projections, different physiologies, and different molecular signatures. Although there is amacrine cell crosstalk between the layers (see below) the bulk of the inner retina’s connectivity occurs within the layers. The stalks of bipolar cell axons, and the proximal dendrites of ganglion cells, often pass through several laminae to reach their final level of stratification, but few synapses are made with these connecting processes en passant: the main work of synaptic connectivity is done within the layers. Indeed, the lamination of the inner plexiform layer is a fundamental guide to the retina’s wiring diagram.