There is growing evidence that sonic hedgehog (Shh) signaling regulates ventral neuronal fate in the vertebrate central nervous system through Nkx-class homeodomain proteins. We have examined the patterns of neurogenesis in mice carrying a targeted mutation in Nkx6.1. These mutants show a dorsal-to-ventral switch in the identity of progenitors and in the fate of postmitotic neurons. At many axial levels there is a complete block in the generation of V2 interneurons and motor neurons and a compensatory ventral expansion in the domain of generation of V1 neurons, demonstrating the essential functions of Nkx6.1 in regional patterning and neuronal fate determination.

Intracellular recordings from the dorsal cochlear nucleus have identified cells with both simple and complex action potential waveforms. We investigated the hypothesis that cartwheel cells are a specific cell type that generates complex action potentials, based on their analogous anatomical, developmental, and biochemical similarities to cerebellar Purkinje cells, which are known to discharge complex action potentials. Intracellular recordings were made from a brain slice preparation of the guinea pig dorsal cochlear nucleus. A subpopulation of cells discharged a series of two or three action potentials riding on a slow depolarization as an all-or-none event; this discharge pattern is called a complex spike or burst. These cells also exhibited anodal break bursts, anomalous rectification, subthreshold inward rectification, and frequent inhibitory postsynaptic potentials (IPSPs). Seven complex-spiking cells were stained with intracellular dyes and subsequently identified as cartwheel neurons. In contrast, six identified simple-spiking cells recorded in concurrent experiments were pyramidal cells. The cartwheel cell bodies reside in the lower part of layer 1 and the upper part of layer 2 of the nucleus. The cells are characterized by spiny dendrites penetrating the molecular layer, a lack of basal dendritic processes, and an axonal plexus invading layers 2 and 3, and the inner regions of layer 1. The cartwheel cell axons made putative synaptic contacts at the light microscopic level with pyramidal cells and small cells, including stellate cells, granule cells, and other cartwheel cells in layers 1 and 2. The axonal plexus of individual cartwheel cells suggests that they can inhibit cells receiving input from either the same or adjacent parallel fibers and that this inhibition is distributed along the isofrequency contours of the nucleus.

Primary mechanosensory receptors and interneurons in the cricket cercal sensory system are sensitive to the direction and frequency of air current stimuli. Receptors innervating long mechanoreceptor hairs (>1000 microm) are most sensitive to low-frequency air currents (<150 Hz); receptors innervating medium-length hairs (900-500 microm) are most sensitive to higher frequency ranges (150-400 Hz). Previous studies demonstrated that the projection pattern of the synaptic arborizations of long hair receptor afferents form a continuous map of air current direction within the terminal abdominal ganglion (). We demonstrate here that the projection pattern of the medium-length hair afferents also forms a continuous map of stimulus direction. However, the afferents from the long and medium-length hair afferents show very little spatial segregation with respect to their frequency sensitivity. The possible functional significance of this small degree of spatial segregation was investigated, by calculating the relative overlap between the long and medium-length hair afferents with the dendrites of two interneurons that are known to have different frequency sensitivities. Both interneurons were shown to have nearly equal anatomical overlap with long and medium hair afferents. Thus, the differential overlap of these interneurons with the two different classes of afferents was not adequate to explain the observed frequency selectivity of the interneurons. Other mechanisms such as selective connectivity between subsets of afferents and interneurons and/or differences in interneuron biophysical properties must play a role in establishing the frequency selectivities of these interneurons.