PubMed · 16881080
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Sheri J Y Mizumori. 2006. Hippocampal place fields: a neural code for episodic memory?. https://doi.org/10.1002/hipo.20209
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During action potential conduction, the axonal specializations at the node, together with the adjacent paranodal terminations of the myelin sheath, interact with glial processes that invest the nodal gap. The nature of the mutual signals between axons and myelinating glia, however, are not well understood. Here we have characterized the distribution of inositol 1,4,5-trisphosphate receptors (IP(3)Rs) in the axoglial apparatus by immunohistochemistry, using known myelin domain-specific markers. While IP(3)R1 is not expressed in the Schwann cells or the axon, IP(3)R2 and IP(3)R3 are expressed in distinct cellular domains, suggesting distinct signaling roles for the two receptors. IP(3)R3 is the most predominant isoform in Schwann cells, and is expressed in particularly dense patches in the paranodal region. In addition to IP(3)Rs, two other members of the metabotropic Ca(2+) signaling pathway, G(alpha)q, and P(2)Y1 type of purinoceptors were also found in Schwann cells. Their pattern of expression matches the expression of their signaling partners, the IP(3)Rs. One interesting finding to emerge from this study is the expression of connexin 32 (Cx32) in close proximity with IP(3)R3. Although IP(3)R3 and Cx32 are not colocalized, their expression in the same membrane areas raises the question whether Schwann cell Ca(2+) signals either control the function of the gap junctions, or whether the gap junctional channels serve as conduits for rapid radial spread of Ca(2+) signals initiated during action potential propagation.
Voltage-gated sodium (Na(v)) channels accumulate at the axon initial segment (IS), where their high density supports spike initiation. Maintenance of this high density of Na(v) channels involves a macromolecular complex that includes the cytoskeletal linker protein ankyrin-G, the only protein known to bind Na(v) channels and localize them at the IS. We found previously that Na(v)1.6 is the predominant Na(v) channel isoform at IS of adult rodent retinal ganglion cells. However, here we report that Na(v)1.6 immunostaining is consistently reduced or absent in short regions of the IS proximal to the soma, although both ankyrin-G and pan-Na(v) antibodies stain this region. We show that this proximal IS subregion is a unique axonal microdomain, containing an accumulation of Na(v)1.1 channels that are spatially segregated from the Na(v)1.6 channels of the distal IS. Additionally, we find that axonal K(v)1.2 potassium channels are present within the distal IS, but are also excluded from the Na(v)1.1-enriched proximal IS microdomain. Because ankyrin-G was prominent in both proximal and distal subcompartments of the IS, where it colocalized with either Na(v)1.1 or Na(v)1.6, respectively, mechanisms other than association with ankyrin-G must mediate differential targeting of Na(v) channel subtypes to achieve the spatial precision observed within the IS. This precise arrangement of ion channels within the axon initial segment is likely an important determinant of the firing properties of ganglion cells and other mammalian neurons.
We present evidence for parallel auditory thalamocortical pathways that serve two different behaviors. The pallid bat listens for prey-generated noise (5-35 kHz) to localize prey, while reserving echolocation [downward frequency-modulated (FM) sweeps, 60-30 kHz] for obstacle avoidance. Its auditory cortex contains a tonotopic map representing frequencies from 6 to 70 kHz. The high-frequency (BF > 30 kHz) representation is dominated by FM sweep-selective neurons, whereas most neurons tuned to lower frequencies prefer noise. Retrograde tracer injections into these physiologically distinct cortical regions revealed that the high-frequency region receives input from the suprageniculate (SG) nucleus, but not the ventral division of the medial geniculate body (MGBv), in all experiments (n = 9). In contrast, the low-frequency region receives tonotopically organized input from the MGBv in all experiments (n = 16). Labeling in the SG was observed in only two of these experiments. Both cortical regions also receive sparse inputs from medial (MGBm) and parts of the dorsal division (MGBd) outside the SG. These results show that the low- and high-frequency regions of a single tonotopic map receive dominant inputs from different thalamic divisions. Within the low-frequency region, most neurons are binaurally inhibited, and an orderly map of interaural intensity difference (IID) sensitivity is present. We show that the input to the IID map arises from topographically organized projections from the MGBv. As observed in other species, a frequency-dependent organization is observed in the lateromedial direction in the MGBv. These data demonstrate that MGBv-to-auditory cortex connections are organized with respect to both frequency and binaural selectivity.