Neuroanatomy of Mud dragons: a comprehensive view of the nervous system in Echinoderes (Kinorhyncha) by confocal laser scanning microscopy.
Background: The Scalidophora (Kinorhyncha, Loricifera and Priapulida) have an important phylogenetic position as early branching ecdysozoans, yet the architecture of their nervous organ systems is notably underinvestigated. Without such information, and in the absence of a stable phylogenetic context, we are inhibited from producing adequate hypotheses about the evolution and diversification of ecdysozoan nervous systems. Here, we utilize confocal laser scanning microscopy to characterize serotonergic, tubulinergic and FMRFamidergic immunoreactivity patterns in a comparative neuroanatomical study with three species of Echinoderes, the most speciose, abundant and diverse genus within Kinorhyncha. Results: Neuroanatomy in Echinoderes as revealed by acetylated α-tubulin immunoreactivity includes a circumpharyngeal brain and ten neurite bundles in the head region that converge into five longitudinal nerves within the trunk. The ventral nerve cord is ganglionated, emerging from the brain with two connectives that converge in trunk segments 2–3, and diverge again within segment 8. The longitudinal nerves and ventral nerve cord are connected by two transverse neurites in segments 2–9. Differences among species correlate with the number, position and innervation of cuticular structures along the body. Patterns of serotoninergic and FMRFamidergic immunoreactivity correlate with the position of the brain neuropil and the ventral nerve cord. Distinct serotonergic and FMRFamidergic somata are associated with the brain neuropil and specific trunk segments along the ventral nerve cord. Conclusions: Neural architecture is highly conserved across all three species, suggesting that our results reveal a pattern that is common to more than 40%of the species within Kinorhyncha. The nervous system of Echinoderes is segmented along most of the trunk; however, posterior trunk segments exhibit modifications that are likely associated with sensorial, motor or reproductive functions. Although all kinorhynchs show some evidence of an externally segmented trunk, it is unclear whether external segmentation matches internal segmentation of nervous and muscular organ systems across Kinorhyncha, as we observed in Echinoderes. The neuroanatomical data provided in this study not only expand the limited knowledge on kinorhynch nervous systems but also establish a comparative morphological framework within Scalidophora that will support broader inferences about the evolution of neural architecture among the deepest branching lineages of the Ecdysozoa.
This work was supported by infrastructure and operating grants to BSL from the Tula Foundation (Centre for Microbial Diversity and Evolution), the National Sciences and Engineering Research Council of Canada (NSERC 2014–05258), and the Canadian Institute for Advanced Research Program in Integrated Microbial Biodiversity. These resources supported collections in British Columbia, provided access to materials and infrastructure, and salary for the first author. Field collections, imaging and research in Florida (USA) were funded by a Smithsonian Institution/Link Foundation Graduate Fellowship in Marine Science to MH. The funding entities had no role in the design, analyses, interpretation of data or writing of the manuscript.
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