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The sense of touch detects forces that bombard the body’s genetic screens in C. elegans and Drosophila melanogaster
surface. In metazoans, an assortment of morphologically have identified a plethora of molecules required for touch
sensation. Recent work has begun to uncover mechanisms
and functionally distinct mechanosensory cell types are
through which these molecules control force sensitivity. By
tuned to selectively respond to diverse mechanical stimuli, comparison, the analysis of touch reception in mammals is in its
such as vibration, stretch, and pressure. A comparative infancy. Here, we introduce commonly used model systems,
THE JOURNAL OF CELL BIOLOGY
evolutionary approach across mechanosensory cell types review emerging cell biological principles that govern touch
is detected by distinct neurons with elaborate sensory dendrites Drosophila mechanosensory neurons. Mechano-
that tile the body wall (Fig. 2 A, red; Way and Chalfie, 1989; sensory genes in Drosophila have been identified through for-
Chatzigeorgiou et al., 2010). Ciliated mechanosensory neurons ward genetic screens for insensitivity to gentle touch, noxious
innervating the nose mediate touch and physically sense food stimuli, and mating song (Kernan et al., 1994; Eberl et al., 1997;
particles, which impacts foraging behaviors (Fig. 2 A, green; Tracey et al., 2003). Like worms and mammals, flies have a siz-
Kaplan and Horvitz, 1993; Sawin et al., 2000; Li et al., 2006; able assortment of mechanosensory neurons essential for sur-
Kindt et al., 2007). Mating behaviors rely on ciliated male-specific vival (for review see Kernan, 2007; Smith and Lewin, 2009).
neurons called sensory rays (Fig. 2 A, orange; Liu and Sternberg, In addition to touch receptors, Drosophila require auditory recep-
1995; Barr and Sternberg, 1999; T. Liu et al., 2007). tors to distinguish mating songs, wing strain gauges to fly and
Thanks to a wealth of tools for analyzing the C. elegans proprioceptors to coordinate their six legs. Even in a cushy lab-
nervous system, mechanotransduction is best understood in this oratory setting, mechanosensory mutants perish because they
organism. Central to this success, Chalfie and colleagues de- are completely uncoordinated.
vised a simple behavioral assay for gentle-body touch and The fly’s body is studded with type I ciliated mechano
screened for genetic mutations that selectively caused mechano- sensory receptors such as chordotonal organs and bristles (Fig. 2;
sensory abnormality (mec) without impairing locomotion Kernan, 2007). Chordotonal organs, which are stretch receptors
(Sulston et al., 1975; Chalfie and Sulston, 1981; Chalfie and Au, attached to the skin, or cuticle, make up the fly’s ear and contribute
1989). Additional studies have dissected responses to nose touch to proprioception. Bristles serve as the principal proprioceptors
and harsh body touch (Way and Chalfie, 1989; Colbert et al., and touch receptors. These mechanosensory organs are generally
1997; Hart et al., 1999; Chatzigeorgiou et al., 2010). To delin- innervated by one sensory neuron, which extends a sensory cilium
eate the cellular basis of mechanosensation, laser ablation has into overlying structures (Fig. 2 C). Bristles act as levers to transmit
been used to pinpoint neurons required for touch-evoked behav- force to the cilium attached at its base by the dendritic cap. This
iors. Additionally, anatomical reconstructions have mapped compresses the ciliary membrane against the tubular bundle, an
connectivity to define neural networks that link touch sensation array of microtubules at the dendrite’s core. The resulting com-
to behaviors. Moreover, mechanosensory molecules can be pression is the putative gating stimulus for mechanotransduction
assigned to signaling pathways with physiological approaches, channels, which carry cation-selective, adapting currents remark-
including in vivo imaging and electrophysiology (Fig. 2 B). ably similar to those of vertebrate hair cells (Fig. 2 D).