Biochemical Basis of Touch Perception: Mechanosensory Function of Degenerin/Epithelial Na+ Channels (2002)

degenerin/epithelial Na+ channel dorsal root ganglion rapidly adapting slowly adapting A-fiber mechano-nociceptors Personal experience informs us that receptors in the skin transmit a wealth of tactile information. We can detect the caress of a cheek, the texture of fabric, a fly alighting on our hand, and a pinch of our arm. Yet of all the vertebrate senses, touch is the least understood at the molecular level. For many years it has been postulated that the core components of mechanosensors are ion channels (for a review and references see Ref. 1Hamill O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). Such channels could convert mechanical energy directly into an electrical signal; this could account for the very high speed response of mechanosensors. In contrast, sensory receptors that detect light, odors, and most tastes initiate second messenger cascades, which then activate ion channels to generate electrical activity.One of the first clues identifying ion channel mechanosensors came from Chalfie and colleagues (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar). To identify components of the mechanosensory apparatus in Caenorhabditis elegans, they developed a genetic screen, assaying the response to light touch. The screen led to the first members of the degenerin/epithelial Na+ channel (DEG/ENaC)1 family. The DEG/ENaC proteins discovered, including MEC-4 and MEC-10, may comprise the core of a multiprotein ion channel complex that opens in response to mechanical stimulation (4Gu G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar, 5Tavernarakis N. Driscoll M. Annu. Rev. Physiol. 1997; 59: 659-689Crossref PubMed Scopus (183) Google Scholar). Here we review insights into how DEG/ENaC channels may contribute to mechanosensation.Anatomical and Physiological Components of MechanosensationPrevious research has taught us much about the bases for the sense of touch. At the behavioral level, psychometric studies in humans have distinguished several distinct cutaneous sensory inputs (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar). At the anatomic level, investigators have identified a variety of specialized cutaneous sensory structures; some examples include Meissner corpuscles, Merkel cell-neurite complexes, lanceolate and pilo-Ruffini fibers surrounding hair follicles, and free nerve endings (Fig.1A) (7Munger B.L. Ide PubMed Scopus Google Scholar). The for in the dorsal root ganglion At the level, that distinct mechanosensory are of sensory include rapidly adapting and slowly adapting A-fiber mechano-nociceptors and mechano-nociceptors (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar, M. 1997; PubMed Scopus Google Scholar). of distinct For fibers and to very light touch. a is they of the skin in the of the include nerve endings in Meissner and lanceolate In contrast, fibers to high mechanical a and slowly to a fibers include some free nerve this of has been into the that may convert mechanical into electrical DEG/ENaC proteins a a of have and and and an Driscoll M. PubMed Scopus Google Scholar). The a of the which PubMed Scopus Google a this to that of some mechanosensors. For the of a is a that mechanical to a a is at the in mechanosensory DEG/ENaC of DEG/ENaC is that of the is to a it in some DEG/ENaC to on the of that a channel that are PubMed Scopus Google Scholar, S. M. G.A. PubMed Scopus Google Scholar). a of the and contribute to the channel Annu. Rev. Physiol. PubMed Scopus Google Scholar). The channels are a of Na+ some channels may have a the very high are to some and may to a of this in some DEG/ENaC members generate an is the channel of and G. B. Nature. 1994; 367: PubMed Scopus Google Scholar). members of examples are the Na+ channel from G. M. Nature. PubMed Scopus Google and and 1996; PubMed Scopus Google Scholar, G. M. Nature. 1997; PubMed Scopus Google Scholar, G. M. 1997; PubMed Scopus Google members PubMed Scopus Google have been to We proteins could ion channels the that the DEG/ENaC the and have have been in DEG/ENaC and of G. B. Nature. 1994; 367: PubMed Scopus Google Scholar, G. N. M. PubMed Scopus Google Na+ channel and 1996; PubMed Scopus Google Scholar, M. PubMed Scopus Google Scholar, G. N. M. 1996; PubMed Scopus Google Scholar, B. Proc. Natl. Acad. Sci. U. S. A. 1997; PubMed Scopus Google ion G. M. Nature. 1997; PubMed Scopus Google Scholar, M. PubMed Scopus Google Scholar, B. Proc. Natl. Acad. Sci. U. S. A. 1997; PubMed Scopus Google Scholar, S. Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google root ion G. M. 1997; PubMed Scopus Google Scholar, M. PubMed Scopus Google G. M. Physiol. Scopus Google ion channel S. PubMed Scopus Google Scholar, PubMed Scopus Google and have is in a DEG/ENaC at the of DEG/ENaC channels in mechanosensory elegans, MEC-4 and (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar, G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar, 5Tavernarakis N. Driscoll M. Annu. Rev. Physiol. 1997; 59: 659-689Crossref PubMed Scopus (183) Google Scholar). In and PubMed Scopus Google Scholar). The sensory of the they sense touch and in to the of specialized and and the of studies have of and and in mechanosensory to the and to specialized cutaneous mechanosensory including the Merkel cell-neurite complex PubMed Scopus Google Scholar, B. G. G. Driscoll M. M. Google Scholar). in several cutaneous mechanosensory including corpuscles, Meissner corpuscles, Merkel cell-neurite complexes, and lanceolate nerve endings surrounding the hair B. Nature. PubMed Scopus Google Scholar, 2001; PubMed Google Scholar). DEG/ENaC channels they can detect tactile of DEG/ENaC in a Merkel cell-neurite of and nerve in lanceolate fibers surrounding hair PubMed Scopus Google Scholar, PubMed Scopus Google Scholar, and B. Nature. PubMed Scopus Google of DEG/ENaC a the a from the in the MEC-4 and this that channels contribute to (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar). DEG/ENaC and may have mechanosensory in in B. 1996; PubMed Scopus Google Scholar, N. S. Driscoll M. 1997; PubMed Scopus Google the mechanosensory of a DEG/ENaC the B. Nature. PubMed Scopus Google Scholar). the response of sensory fibers to mechanical an in In the of a mechanical the of in In to mechanical the a The of a to and for the of The of is for the and of touch (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google of from and Ref. B. Nature. PubMed Scopus Google and that activate some DEG/ENaC channels and that they contribute to M. PubMed Scopus Google Scholar). may this in Proc. Natl. Acad. Sci. U. S. A. 2001; PubMed Scopus Google and DEG/ENaC channels may contribute to in some cutaneous and are in the and of mechanosensors. DEG/ENaC channels generate in in M. and it that to from the of they are to detect tactile PubMed Google Scholar, of New Scholar). is We that in mechanosensory nerve DEG/ENaC channels are to proteins that touch We that is a of DEG/ENaC in are the this is the then the the response of channels mechanosensors. For DEG/ENaC channels of a Proc. Natl. Acad. Sci. U. S. A. 2001; PubMed Scopus Google Scholar). the the of to a mechanical We of DEG/ENaC in and at least in to a tactile of DEG/ENaC to in channels have been for the ion channels in hair studies of that and are for A. PubMed Scopus Google channels are to channels in Nature. 2001; PubMed Scopus Google in detect that the and and in PubMed Scopus Google Scholar). of nerve an DEG/ENaC detect and to generate a that and have a in they in DEG/ENaC the is on this we the genetic the and the in that the is it is that the of in B. Nature. PubMed Scopus Google Scholar). that are and a mechanosensory has been in B. 1996; PubMed Scopus Google Scholar, N. S. Driscoll M. 1997; PubMed Scopus Google Scholar). We that in DEG/ENaC contribute to in for the DEG/ENaC channels are at the of in and in in response to the of the nerve endings this a to DEG/ENaC In it to the all the to a in in a the can it may to about how the channels DEG/ENaC are about how mechanosensors O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). could directly activate the channel channels Martinac B. Nature. 1994; PubMed Scopus Google Scholar, G. PubMed Scopus Google Scholar). is is proteins are channels that are may directly O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). are about can this at how this in in is which a channel to a tactile could of an that a channel O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). for G. PubMed Google from a an that a to account for the very high speed of for a tactile of PubMed Scopus Google Scholar). and Martinac O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google that the response of mechanosensors may to this some mechanosensory nerve including and Merkel to to nerve endings PubMed Scopus Google Scholar, PubMed Scopus Google Scholar). endings proteins G. PubMed Scopus Google Scholar). Yet some that of the Merkel the nerve to M. PubMed Scopus Google Scholar). We the mechanosensory that mechanosensory channel the mechanosensory a O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google is the we for DEG/ENaC In this the channel the the O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar, G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Driscoll M. PubMed Scopus Google Scholar, Nature. 2001; PubMed Scopus Google Scholar). of this complex and to the genetic identified several for tactile (4Gu G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar). The of MEC-4 and channel may and an At the the and the may to the mechanosensory complex a that a B. 1996; PubMed Scopus Google DEG/ENaC proteins have been and of Annu. Rev. Physiol. PubMed Scopus Google and and Google the for are may to channel members to channels Nature. 2001; PubMed Scopus Google Scholar). proteins that DEG/ENaC proteins in us how mechanical that mechanical of sensory nerve endings an ion channel complex that DEG/ENaC the core a that The of molecular including DEG/ENaC and and the to sensory receptors may of this channel to in the we experience in our sense of touch. Personal experience informs us that receptors in the skin transmit a wealth of tactile information. We can detect the caress of a cheek, the texture of fabric, a fly alighting on our hand, and a pinch of our arm. Yet of all the vertebrate senses, touch is the least understood at the molecular level. For many years it has been postulated that the core components of mechanosensors are ion channels (for a review and references see Ref. 1Hamill O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). Such channels could convert mechanical energy directly into an electrical signal; this could account for the very high speed response of mechanosensors. In contrast, sensory receptors that detect light, odors, and most tastes initiate second messenger cascades, which then activate ion channels to generate electrical of the first clues identifying ion channel mechanosensors came from Chalfie and colleagues (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar). To identify components of the mechanosensory apparatus in Caenorhabditis elegans, they developed a genetic screen, assaying the response to light touch. The screen led to the first members of the degenerin/epithelial Na+ channel (DEG/ENaC)1 family. The DEG/ENaC proteins discovered, including MEC-4 and MEC-10, may comprise the core of a multiprotein ion channel complex that opens in response to mechanical stimulation (4Gu G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar, 5Tavernarakis N. Driscoll M. Annu. Rev. Physiol. 1997; 59: 659-689Crossref PubMed Scopus (183) Google Scholar). Here we review insights into how DEG/ENaC channels may contribute to and Physiological Components of MechanosensationPrevious research has taught us much about the bases for the sense of touch. At the behavioral level, psychometric studies in humans have distinguished several distinct cutaneous sensory inputs (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar). At the anatomic level, investigators have identified a variety of specialized cutaneous sensory structures; some examples include Meissner corpuscles, Merkel cell-neurite complexes, lanceolate and pilo-Ruffini fibers surrounding hair follicles, and free nerve endings (Fig.1A) (7Munger B.L. Ide PubMed Scopus Google Scholar). The for in the dorsal root ganglion At the level, that distinct mechanosensory are of sensory include rapidly adapting and slowly adapting A-fiber mechano-nociceptors and mechano-nociceptors (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar, M. 1997; PubMed Scopus Google Scholar). of distinct For fibers and to very light touch. a is they of the skin in the of the include nerve endings in Meissner and lanceolate In contrast, fibers to high mechanical a and slowly to a fibers include some free nerve this of has been into the that may convert mechanical into electrical research has taught us much about the bases for the sense of touch. At the behavioral level, psychometric studies in humans have distinguished several distinct cutaneous sensory inputs (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar). At the anatomic level, investigators have identified a variety of specialized cutaneous sensory structures; some examples include Meissner corpuscles, Merkel cell-neurite complexes, lanceolate and pilo-Ruffini fibers surrounding hair follicles, and free nerve endings (Fig.1A) (7Munger B.L. Ide PubMed Scopus Google Scholar). The for in the dorsal root ganglion At the level, that distinct mechanosensory are of sensory include rapidly adapting and slowly adapting A-fiber mechano-nociceptors and mechano-nociceptors (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar, M. 1997; PubMed Scopus Google Scholar). of distinct For fibers and to very light touch. a is they of the skin in the of the include nerve endings in Meissner and lanceolate In contrast, fibers to high mechanical a and slowly to a fibers include some free nerve Yet this of has been into the that may convert mechanical into electrical The DEG/ENaC proteins a a of have and and and an Driscoll M. PubMed Scopus Google Scholar). The a of the which PubMed Scopus Google a this to that of some mechanosensors. For the of a is a that mechanical to a a is at the in mechanosensory DEG/ENaC DEG/ENaC to on the of that a channel that are PubMed Scopus Google Scholar, S. M. G.A. PubMed Scopus Google Scholar). a of the and contribute to the channel Annu. Rev. Physiol. PubMed Scopus Google Scholar). The channels are a of Na+ some channels may have a the very high are to some and may to a of this in some DEG/ENaC members generate an is the channel of and G. B. Nature. 1994; 367: PubMed Scopus Google Scholar). members of examples are the Na+ channel from G. M. Nature. PubMed Scopus Google and and 1996; PubMed Scopus Google Scholar, G. M. Nature. 1997; PubMed Scopus Google Scholar, G. M. 1997; PubMed Scopus Google members PubMed Scopus Google have been to We proteins could ion channels the that the DEG/ENaC the and have have been in DEG/ENaC and of G. B. Nature. 1994; 367: PubMed Scopus Google Scholar, G. N. M. PubMed Scopus Google Na+ channel and 1996; PubMed Scopus Google Scholar, M. PubMed Scopus Google Scholar, G. N. M. 1996; PubMed Scopus Google Scholar, B. Proc. Natl. Acad. Sci. U. S. A. 1997; PubMed Scopus Google ion G. M. Nature. 1997; PubMed Scopus Google Scholar, M. PubMed Scopus Google Scholar, B. Proc. Natl. Acad. Sci. U. S. A. 1997; PubMed Scopus Google Scholar, S. Proc. Natl. Acad. Sci. U. S. A. PubMed Scopus Google root ion G. M. 1997; PubMed Scopus Google Scholar, M. PubMed Scopus Google G. M. Physiol. Scopus Google ion channel S. PubMed Scopus Google Scholar, PubMed Scopus Google and have is in a DEG/ENaC proteins a a of have and and and an Driscoll M. PubMed Scopus Google Scholar). The a of the which PubMed Scopus Google a this to that of some mechanosensors. For the of a is a that mechanical to a a is at the in mechanosensory DEG/ENaC DEG/ENaC to on the of that a channel that are PubMed Scopus Google Scholar, S. M. G.A. PubMed Scopus Google Scholar). a of the and contribute to the channel Annu. Rev. Physiol. PubMed Scopus Google Scholar). The channels are a of Na+ some channels may have a the very high are to some and may to a of this family. in some DEG/ENaC members generate an is the channel of and G. B. Nature. 1994; 367: PubMed Scopus Google Scholar). members of examples are the Na+ channel from G. M. Nature. PubMed Scopus Google and and 1996; PubMed Scopus Google Scholar, G. M. Nature. 1997; PubMed Scopus Google Scholar, G. M. 1997; PubMed Scopus Google members PubMed Scopus Google have been to We proteins could ion channels the that the DEG/ENaC the and have have been in and have is DEG/ENaC at the of DEG/ENaC channels in mechanosensory elegans, MEC-4 and (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar, G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar, 5Tavernarakis N. Driscoll M. Annu. Rev. Physiol. 1997; 59: 659-689Crossref PubMed Scopus (183) Google Scholar). In and PubMed Scopus Google Scholar). The sensory of the they sense touch and in to the of specialized and and the of studies have of and and in mechanosensory to the and to specialized cutaneous mechanosensory including the Merkel cell-neurite complex PubMed Scopus Google Scholar, B. G. G. Driscoll M. M. Google Scholar). in several cutaneous mechanosensory including corpuscles, Meissner corpuscles, Merkel cell-neurite complexes, and lanceolate nerve endings surrounding the hair B. Nature. PubMed Scopus Google Scholar, 2001; PubMed Google Scholar). DEG/ENaC channels they can detect tactile DEG/ENaC channels in mechanosensory elegans, MEC-4 and (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar, G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar, 5Tavernarakis N. Driscoll M. Annu. Rev. Physiol. 1997; 59: 659-689Crossref PubMed Scopus (183) Google Scholar). In and PubMed Scopus Google Scholar). The sensory of the they sense touch and in to the of specialized and and the of studies have of and and in mechanosensory to the and to specialized cutaneous mechanosensory including the Merkel cell-neurite complex PubMed Scopus Google Scholar, B. G. G. Driscoll M. M. Google Scholar). in several cutaneous mechanosensory including corpuscles, Meissner corpuscles, Merkel cell-neurite complexes, and lanceolate nerve endings surrounding the hair B. Nature. PubMed Scopus Google Scholar, 2001; PubMed Google Scholar). DEG/ENaC channels they can detect tactile of DEG/ENaC a the a from the in the MEC-4 and this that channels contribute to (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar). DEG/ENaC and may have mechanosensory in in B. 1996; PubMed Scopus Google Scholar, N. S. Driscoll M. 1997; PubMed Scopus Google the mechanosensory of a DEG/ENaC the B. Nature. PubMed Scopus Google Scholar). the response of sensory fibers to mechanical an in In the of a mechanical the of in In to mechanical the a The of a to and for the of The of is for the and of touch (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar). a the a from the in the MEC-4 and this that channels contribute to (2Driscoll M. Chalfie M. Nature. 1991; 349: 588-593Crossref PubMed Scopus (452) Google Scholar, 3Huang M. Chalfie M. Nature. 1994; 367: 467-470Crossref PubMed Scopus (343) Google Scholar). DEG/ENaC and may have mechanosensory in in B. 1996; PubMed Scopus Google Scholar, N. S. Driscoll M. 1997; PubMed Scopus Google Scholar). To the mechanosensory of a DEG/ENaC the B. Nature. PubMed Scopus Google Scholar). the response of sensory fibers to mechanical an in In the of a mechanical the of in In to mechanical the a The of a to and for the of The of is for the and of touch (6Perl E.R. Scott S.A. Sensory Neuron: Diversity, Development, and Plasticity. Oxford University Press, New York1992: 3-23Google Scholar). and that activate some DEG/ENaC channels and that they contribute to M. PubMed Scopus Google Scholar). may this in Proc. Natl. Acad. Sci. U. S. A. 2001; PubMed Scopus Google and DEG/ENaC channels may contribute to in some cutaneous and are in the and of mechanosensors. DEG/ENaC channels generate in in M. and it that to from the of they are to detect tactile PubMed Google Scholar, of New Scholar). is We that in mechanosensory nerve DEG/ENaC channels are to proteins that touch We that is a of DEG/ENaC in are the this is the then the the response of channels mechanosensors. For DEG/ENaC channels of a Proc. Natl. Acad. Sci. U. S. A. 2001; PubMed Scopus Google Scholar). the the of to a mechanical We of DEG/ENaC in and at least in to a tactile The that activate some DEG/ENaC channels and that they contribute to M. PubMed Scopus Google Scholar). may this in Proc. Natl. Acad. Sci. U. S. A. 2001; PubMed Scopus Google and DEG/ENaC channels may contribute to in some cutaneous and are in the and of mechanosensors. DEG/ENaC channels generate in in M. and it that to from the of they are to detect tactile PubMed Google Scholar, of New Scholar). is We that in mechanosensory nerve DEG/ENaC channels are to proteins that touch We that is a of DEG/ENaC in are the this is the then the the response of channels mechanosensors. For DEG/ENaC channels of a Proc. Natl. Acad. Sci. U. S. A. 2001; PubMed Scopus Google Scholar). the the of to a mechanical We of DEG/ENaC in and at least in to a tactile of DEG/ENaC to in channels have been for the ion channels in hair studies of that and are for A. 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Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). could directly activate the channel channels Martinac B. Nature. 1994; PubMed Scopus Google Scholar, G. PubMed Scopus Google Scholar). is is proteins are channels that are may directly O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). are about can this at how this in in is could of an that a channel O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar). for G. PubMed Google from a an that a to account for the very high speed of for a tactile of PubMed Scopus Google Scholar). and Martinac O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google that the response of mechanosensors may to this some mechanosensory nerve including and Merkel to to nerve endings PubMed Scopus Google Scholar, PubMed Scopus Google Scholar). endings proteins G. PubMed Scopus Google Scholar). Yet some that of the Merkel the nerve to M. PubMed Scopus Google Scholar). We the mechanosensory that mechanosensory channel the mechanosensory The a O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google is the we for DEG/ENaC In this the channel the the O.P. Martinac B. Physiol. Rev. 2001; 81: 685-740Crossref PubMed Scopus (914) Google Scholar, G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Driscoll M. PubMed Scopus Google Scholar, Nature. 2001; PubMed Scopus Google Scholar). of this complex and to the genetic identified several for tactile (4Gu G. Caldwell G.A. Chalfie M. Proc. Natl. Acad. Sci. U. S. A. 1996; 93: 6577-6582Crossref PubMed Scopus (90) Google Scholar). The of MEC-4 and channel may and an At the the and the may to the mechanosensory complex a that a B. 1996; PubMed Scopus Google Scholar). For DEG/ENaC proteins have been and of Annu. Rev. Physiol. 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