SummaryThe inner ear and the epibranchial ganglia constitute much of the sensory system in the caudal vertebrate head. The inner ear consists of mechanosensory hair cells, their neurons, and structures necessary for sound and balance sensation. The epibranchial ganglia are knots of neurons that innervate and relay sensory signals from several visceral organs and the taste buds. Their development was once thought to be independent, in line with their independent functions. However, recent studies indicate that both systems arise from a morphologically distinct common precursor domain: the posterior placodal area. This review summarises recent studies into the induction, morphogenesis and innervation of these systems and discusses lineage restriction and cell specification in the context of their common origin.
Key words: Epibranchial, Inner ear, Neurogenesis, Placode, Signalling
IntroductionCranial placodes, found in all vertebrates, are transient thickenings of ectoderm that contribute extensively to the sensory component of the cephalic peripheral nervous system (see Box 1 and Glossary, Box 2). Individual placodes give rise to characteristic cell types, although the diversity of placodal derivatives varies (Box 1). Some placodes, such as the olfactory, otic and lateral line placodes, can form the receptive cell that responds to a stimulus, as well as the sensory neurons that transmit this information (Box 1). Others, such as the epibranchial and trigeminal placodes, only give rise to sensory neurons. The lens and adenohypophyseal placodes generate no sensory derivatives (Baker and Bronner-Fraser, 2001;Webb and Noden, 1993;Begbie and Graham, 2001b). In this review, we focus on the inner ear (or otic) placode and the epibranchial series of placodes and discuss their origins from a common progenitor domain: the posterior placodal area (PPA) (Fig. 1).The otic placode forms the complex inner ear structure that detects sound and balance, as well as the neurons that convey this information to the auditory hindbrain. The otic placodes form distinctive paired depressions adjacent to the caudal hindbrain and progressively deepen to form otocysts (see Glossary, Box 2). Transcriptional networks, influenced by extrinsic signals, drive the regional differentiation of the otic placode to generate mechanosensory hair cells, supporting cells and neurons (see Box 1 and Glossary, Box 2). This progressive differentiation results in a remarkable convolution of the simple spherical otocyst into an intricate structure that is dedicated to receiving information on balance, angular velocity and sound. These later morphogenetic events have been well reviewed (Bok et al., 2007;Fritzsch et al., 2006;Torres and Giráldez, 1998) and will not be covered here.The epibranchial placodes give rise to the geniculate, petrosal and nodose ganglia, which contribute sensory neurons to cranial nerves VII (facial), IX (glossopharyngeal) and X (vagus), in that order (see Box 1 and Glossary, Box 2; Fig. 1). Epibranchial placodes are located ventral to the...