Development of tyrosine hydroxylase-immunoreactive cell populations and fiber pathways in the brain of the dogfish Scyliorhinus canicula: new perspectives on the evolution of the vertebrate catecholaminergic system.

Developmental studies of the central catecholaminergic (CA) system are essential for understanding its evolution. To obtain knowledge about the CA system in chondrichthyans, an ancient gnathostome group, we used immunohistochemical techniques for detecting tyrosine hydroxylase (TH), the initial rate-limiting enzyme of the CA synthesis, to study: 1) the neuromery of developing TH-immunoreactive (ir) neuronal populations, 2) the development of TH-ir innervation, and 3) the organization of TH-ir cells and fibers in the brain of postembryonic stages of the shark Scyliorhinus canicula. The first TH-ir cells appeared in the hypothalamus and rostral diencephalon (suprachiasmatic, posterior recess and posterior tubercle nuclei at embryonic stage 26, and dorsomedial hypothalamus at stage 28); then in more caudal basal regions of the diencephalon and rostral mesencephalon (substantia nigra/ventral tegmental area); and later on in the anterior (locus coeruleus/nucleus subcoeruleus) and posterior (vagal lobe and reticular formation) rhombencephalon. The appearance of TH-ir cells in the telencephalon (pallium) was rather late (stage [S]31) with respect to the other TH-ir prosencephalic populations. The first TH-ir fibers arose from cells of the posterior tubercle (S30) and formed recognizable ascending (toward dorsal and rostral territories) and descending pathways at S31. When the second half of embryonic development started (S32), TH-ir fibers innervated most brain areas, and nearly all TH-ir cell groups of the postembryonic brain were already established. This study provides key information about the evolution of the developmental patterns of central CA systems in fishes and thus may help in understanding how the vertebrate CA systems have evolved.