Independent specialization of arthropod body segments has led to more than a century of debate on the homology of morphologically diverse segments1,2, each defined by a lateral appendage and a ganglion of the central nervous system. The plesiomorphic composition of the arthropod head remains enigmatic because variation in segments and corresponding appendages is extreme. Within extant arthropod classes (Chelicerata, Myriapoda, Crustacea and Hexapoda—including the insects), correspondences between the appendage-bearing second (deutocerebral) and third (tritocerebral) cephalic neuromeres have been recently resolved on the basis of immunohistochemistry1 and Hox gene expression patterns3,4. However, no appendage targets the first ganglion, the protocerebrum, and the corresponding segmental identity of this anterior region remains unclear5. Reconstructions of stem-group arthropods indicate that the anteriormost region originally might have borne an ocular apparatus and a frontal appendage innervated by the protocerebrum6. However, no study of the central nervous system in extant arthropods has been able to corroborate this idea directly, although recent analyses of cephalic gene expression patterns in insects suggest a segmental status for the protocerebral region7,8,9,10. Here we investigate the developmental neuroanatomy of a putative basal arthropod11, the pycnogonid sea spider, with immunohistochemical techniques. We show that the first pair of appendages, the chelifores, are innervated at an anterior position on the protocerebrum. This is the first true appendage shown to be innervated by the protocerebrum, and thus pycnogonid chelifores are not positionally homologous to appendages of extant arthropods but might, in fact, be homologous to the ‘great appendages’ of certain Cambrian stem-group arthropods.
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We thank W. Morrisey for diagram preparation; E. C. Seaver for assistance with immunohistochemistry; and J. Hanken, G. Das, G. Edgecombe and A. Hejnol for advice and discussion. We thank the Developmental Studies Hybridoma Bank for the anti-tubulin and Elav antibodies. This material is based on work supported by the National Science Foundation AToL program to G.G. and M.Q.M.
These images show an Anoplodactylus larva labelled with the neuronal marker, Elav (red). Anterior chelifores are directed up. The ocular nerves exiting the dorsal surface of the protocerebrum have been converted from red to yellow. A. Scale bar: 25 µm. In the upper row, ocular nerves are artificially coloured, the bottom row is raw data.
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Marine Biology (2007)