Segmental variation in identified neurons may provide an opportunity to examine extrinsic influences on neuronal phenotype, since segmentally homologous neurons must contain much the same intrinsic information, having arisen from very similar or identical precursors. Two large serotonergic Retzius (Rz) cells are found in each segmental ganglion of the leech Hirudo medicinalis. While most Rz cells innervate the body wall in their own segment and, by way of axons in the interganglionic connectives, the body wall of adjacent segments, the Rz cells in ganglia 5 and 6 [Rz(5,6)] lack interganglionic axons and innervate only the reproductive tissue (Glover and Mason, 1986). Here we describe and quantify the development of differences between Rz(5,6) and other Rz cells in peripheral innervation, neuropilar arborization, and soma size. We filled individual Rz cells with Lucifer yellow or HRP in adults and in staged embryos. During the first 72 hr of outgrowth of Rz cell processes, the morphology of Rz(5,6) was indistinguishable from that of other Rz cells. Only after the processes of Rz(5,6) reached the reproductive tissue did they begin to differ from their segmental homologs. This temporal correlation suggests that these morphological differences arise because of some interaction between Rz(5,6) and their target tissue.
Planarians are free-living aquatic flatworms that possess a well-documented photophobic response to light. With a true central nervous system and simple cerebral eyes (ocelli), planarians are an emerging model for regenerative eye research. However, comparatively little is known about the physiology of their photoreception or how their behavior is affected by various wavelengths. Most phototactic studies have examined planarian behavior using white light. Here, we describe a novel planarian behavioral assay to test responses to small ranges of visible wavelengths (red, blue, green), as well as ultraviolet (UV) and infrared (IR) which have not previously been examined. Our data show that planarians display behavioral responses across a range of wavelengths. These responses occur in a hierarchy, with the shortest wavelengths (UV) causing the most intense photophobic responses while longer wavelengths produce no effect (red) or an apparent attraction (IR). In addition, our data reveals that planarian photophobia is comprised of both a general photophobic response (that drives planarians to escape the light source regardless of wavelength) and wavelength-specific responses that encompass specific behavioral reactions to individual wavelengths. Our results serve to improve the understanding of planarian phototaxis and suggest that behavioral studies performed with white light mask a complex behavioral interaction with the environment.
Invertebrates have proved to be important experimental systems for examining questions related to growth cone navigation and nerve formation, in large part because of their simpler nervous systems. However, such apparent simplicity can be deceiving because the final stereotyped patterns may be the result of multiple developmental mechanisms and not necessarily the sole consequence of the pathway choices of individual growth cones. We have examined the normal sequence of events that are involved in the formation of the major peripheral nerves in leech embryos by employing (1) an antibody directed against acetylated tubulin to label neurons growing out from the central nervous system, (2) the Lan3-2 antibody to label a specific population of peripheral neurons growing into the central nervous system, and (3) intracellular dye filling of single cells. We found that the mature pattern of nerves was characterized by a pair of large nerve roots, each of which branched into two major tracts. The earliest axonal projections did not, however, establish this pattern definitively. Rather, each of the four nerves initially formed as discrete, roughly parallel tracts without bifurcation, with the final branching pattern of the nerve roots being generated by a secondary condensation. In addition, we found that some of the nerves were pioneered in different ways and by different groups of neurons. One of the nerves was established by central neurons growing peripherally, another by peripheral neurons growing centrally. These results suggest that the formation of common nerves and neuronal pathfinding in the leech involves multiple sets of growth cone guidance strategies and morphogenetic mechanisms that belie its apparent simplicity.
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