Do chitons have a brain? New evidence for diversity and complexity in the polyplacophoran central nervous system.

Molluscs demonstrate astonishing morphological diversity, and the relationships among clades have been debated for more than a century. Molluscan nervous systems range from simple 'ladder-like' cords to the complex brains of cephalopods. Chitons (Polyplacophora) are assumed to retain many molluscan plesiomorphies, lacking neural condensation and ganglionic structure, and therefore a brain. We reconstructed three-dimensional anatomical models of the nervous system in eight species of chitons in an attempt to clarify chiton neuroarchitecture and its variability. We combined new data with digitised historic slide material originally used by malacologist Johannes Thiele (1860-1935). Reconstructions of whole nervous systems in Acanthochitona fascicularis, Callochiton septemvalvis, Chiton olivaceus, Hemiarthrum setulosum, Lepidochitona cinerea, Lepidopleurus cajetanus and Leptochiton asellus, and the anterior nervous system of Schizoplax brandtii, demonstrated consistent and substantial anterior neural concentration in the circumoesophageal nerve ring. This is further organised into three concentric tracts, corresponding to the lateral, ventral and cerebral nerve cords. These represent homologues to the three main pairs of ganglia in other molluscs. Their relative size, shape and organisation are highly variable among the examined taxa, but consistent with previous studies of select species, and we formulated a set of neuroanatomical characters for chitons. These support anatomical transitions at the ordinal and subordinal levels; the identification of robust homologies in neural architecture will be central to future comparisons across Mollusca and, more broadly, Lophotrochozoa. Contrary to almost all previous descriptions, the size and structure of the chiton anterior nerve ring unambiguously qualify it as a true brain with cordal substructure.