Ear and Lateral Line of Vertebrates: Organization and Development

The mechano-electric transducers of vertebrates (the hair cell) form the basis for two distinct sensory systems, the lateral line and the inner ear. These two sensory systems share other features such as development and the central projections to the hindbrain and are therefore grouped together as the octavo-lateral system. This paper deals with the mechano-sensory hair cells, the organization of sensory organs in the lateral line and inner ear, their central projection, the development of the inner ear and, to a lesser extent, the lateral-line system as well as the system of efferent cells modifying their sensory responses.

Keywords: inner ear; development; lateral line; neuroanatomy; anatomy; vestibular; auditory

Figure 1. The mechano-sensory hair cell displays a marked asymmetry of the stereocilia (blue), which are always smaller the further they are away from the eccentric kinocilium (black). Mechano-electric transduction apparently requires the tip links between stereocilia (insert upper right). If deflection causes the tip links to relax, the mechanically gated ion channels will close. If the tip link tension is increased through appropriate deflection to the right, mechanically gated channels will be opened and the influx of potassium causes a proportional change in the resting potential of about 3 mV m–1 deflection (inset lower right).
Figure 2. This scheme shows the three octavo-lateral organs (ampullary electroreceptors, mechanosensory lateral line neuromasts, inner ear) and their primary connections in the hindbrain of a salamander, shown on a hemisection through the brainstem. In addition, the position of efferent cells is shown as well as their axonal trajectories to the two sets of mechanosensory organs. The inset on the right shows the approximate position of the dorso-lateral placodes that give rise to these organs and ganglia. Colour is identical for ganglia and their central nuclei in the hindbrain and the placodes of origin.
Figure 3. This drawing illustrates the presumed inductive interactions between the forming neural tube (n) and the mesoderm (represented by the notchord, not) to induce an otic placode (p) with a clear anterior/posterior axes (red and blue). Once induced and polarized through appropriate interactions with the neural tube and the mesoderm (indicated by arrows) the otic placode will invaginate into an otic pit (op). At this stage a medio-lateral and dorso-ventral axes will be added (red and green). Subsequently, the otic pit will close and separate from the ectoderm, fully polarized, (shades of red and green) to become the otocyst. Experimental evidence indicates that there is variation in the relative importance of the neural and mesodermal inductive actions (yellow and red arrows), probably over time (as shown here) and also between species. Several early genes that are important in the hindbrain or the otic placode have been characterized and to some extent tested for their alleged function. Both transplantation studies as well as gene deletion experiments indicate that these early interactions lead to the appropriate polarity formation of the otocyst and gene expression. Thus bone morphogenic protein 4 (BMP-4) and BDNF will be early on expressed in the future semicircular canal sensory epithelia (blue) and lunatic fringe, Fgf3 and NT-3 will be expressed in the segregating utricular, saccular and cochlear sensory epithelia (red). The receptors for the two neurotrophins expressed in the ear (trkB and trkC) will be expressed in the differentiating sensory neurons of the otic ganglion (green).
Figure 4. This image shows specific deletions achieved with certain genes (top) and the pattern of innervation and the loss in various single neurotrophin and neurotrophin receptor mutants (bottom). While Pax-2 appears to be specific for cochlea formation, Otx-1 and Hmx-3 seems to be specific for the horizontal canal formation. Note that in both BDNF and trkB mutants there is a complete loss of innervation to the semicircular canals, a severe reduction of the vestibular ganglion and a reduction of the apical innervation density in the cochlea. In contrast, in NT-3 and trkC mutations there is only a small reduction in the vestibular ganglion but a severe reduction in the cochlear innervation with an almost or complete loss in the basal turn.
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 Further Reading
    book Berlin CI (1999) The Efferent Auditory System:Basic Science and Clinical Applications. San Diego: Singular Publishing.
    book Coombs S, Görner P and Münz H (1989) The Mechanosensory Lateral Line, pp. 724. New York: Springer Verlag.
    book Ehret G and Romand R (1997) The Central Auditory System, p. 404. New York: Oxford University Press.
    Fritzsch B and Beisel K (2004) Keeping sensory cells and evolving neurons to connect them to the brain: molecular conservation and novelties in vertebrate ear development. Brain, Behavior and Evolution 64: 182–197.
    Guedry FE (1992) Perception of motion and position relative to the earth. An overview. Annals of the New York Academy of Sciences 656: 315–328.
    book Lewis ER, Leverenz EL and Bialek WS (1985) The Vertebrate Inner Ear, p. 248. Boca Raton: CRC Press.
    book Pickles JO (1994) An Introduction to the Physiology of Hearing, p. 367. London: Academic Press.
    Romand R and Varela-Nieto I (2003) Development of auditory and vestibular systems-3. Molecular development of the inner ear. Current Topics in Developmental Biology 57: 1–481.
    book Webster DB, Fay RR and Popper AN (1992) The Evolutionary Biology of Hearing, p. 859. New York: Springer.
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Fritzsch, Bernd(Jan 2006) Ear and Lateral Line of Vertebrates: Organization and Development. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1038/npg.els.0004187]