Gordon Fain

Gordon                         Fain                          Email: gfain@ucla.edu
Phone: (310) 206-4281
Address: 2121 TLSB
Research Interests: Physiology of mammalian photoreceptors


B.A. Biology Stanford University 1968
Ph.D. Biophysics Johns Hopkins University 1973

Research Interests:

A vertebrate photoreceptor uses a G-protein receptor (rhodopsin) and a G-protein cascade to produce the electrical response that signals a change in light intensity. Powerful new techniques have made it possible to understand the working of this cascade in extraordinary detail. The reason for this is that practically every protein involved in the cascade in a photoreceptor, from the pigment molecule rhodopsin to the G-protein and channels, but including also a large number of control proteins, are expressed only in the photoreceptors and nowhere else in the body. This makes it possible with genetic techniques to create mice in which these proteins have been knocked out, over or under expressed, or replaced with proteins of modified structure. We use electrical recording to study the effects of such genetic alterations on the light responses of mouse rods, in order to understand the role of these proteins in the visual cascade. We are especially interested in modulatory enzymes and their function in light and dark adaptation. We also have a long-standing interest in mechanisms of photoreceptor degeneration in genetically inherited disease. We are presently the only laboratory in the world making measurements of changes in Ca from mouse rods with fluorescent dyes, and our experiments have implicated both increases and decreases in Ca concentration as contributing factors in the triggering of apoptosis.


Li, X., McClellan, M.E., Tanito, M., Garteiser, P., Towner, R., Bissig, D., Berkowitz, B.A., Fliesler, S.J., Woodruff, M.L., Fain, G.L., Birch, D.G., Khan, M.S., Ash, J.D. and Elliot, M.H. (2012). Loss of caveolin-1 impairs retinal function due to disturbance of subretinal microenvironment Journal of Biological Chemistry 287: 16424-16434

Nymark, S., Frederiksen, R., Woodruff, M.L., Cornwall, M.C. and Fain, G.L. (2012). Bleaching of mouse rods: microspectrophotometry and suction-electrode recording Journal of Physiology 590: 2353-2364

Tsang, S.H., Woodruff, M.L., Lin, C.S., Jacobson, B.D., Naumann, M.C., Hsu, C.W., Davis, R.J., Cilluffo, M.C., Chen, J. and Fain, G.L. (2012). Effect of the ILE86TER mutation in the gamma subunit of cGMP phophodiesterase (PDE6) on rod photoreceptor signaling Cellular Signalling 24: 181-188

Fain, G.L. (2011). Adaption of mammalian photoreceptors to background light: putative role for direct modulation of phosphodiesterase Molecular Neurobiology 44: 374-382

Chen, J., Woodruff, M.L., Wang, T., Concepcion, F., Tranchina, D. and Fain, G.L. (2010). Channel modulation and the mechanism of light adaption in mouse rods J. Neuroscience 30: 16232-16240

Chen, C.K., Woodruff, M.L., Chen, F.S., Shim, H., Cilluffo, M.C. and Fain, G.L. (2010). Replacing the rod with the conde transducin alpha subunit decreases sensitivity and accelerates response decay J Physiol 588: 3231-3241

Kevalov, V.J., Cornwall, M.C. and Fain, G.L. (2010). Physiological studies of the interaction between opsin and chromophore in rod and cone visual pigments Retinoids: Methods and Protocols (Methods in Molecular Biology vol. 652) 95-114

Fain, G.L., Hardie, R. and Laughlin, S.B. (2010). Phototransduction and the evolution of photoreceptors Current Biology 20: R114-124

Chen, C.K., Woodruff, M.L., Chen, F.S., Chen, D. and Fain, G.L. (2010). Background light produces a recoverin-independent modulation of activated-rhodopsin lifetime in mouse rods J. Neuroscience 30: 1213-1220

Sampath, A.P. and Fain, G.L. (2009). Setting the absolute threshold vision F1000 Reports 2009 1:66: -

Okawa, H., Sampath, A.P., Laughlin, S.B. and Fain, G.F. (2008). ATP consumption by mammmalian rod photoreceptors in darkness and in light Current Biology 18: 1917-1921

Dizhoor, A.M., Woodruff, M.L., Olshevskaya, E.V., Cilluffo, M.C., Cornwall, M.C., Sieving, P.A. and Fain, G.L. (2008). Night blindness and the mechanism of constitutive signaling of mutant G90D Rhodosin Journal of Neuroscience 28: 11662-11672

Woodruff, M.L., Janisch, K.M., Peshenko, I.V., Dizhoor, A.M., Tsang, S.H., Fain, G.L. (2008). Modulation of Phosphodiesterase6 turnoff during background illumination in mouse rod photoreceptors Journal Neuroscience 28: 2064-2074

Woodruff, M.L., Olshevskaya, E.V., Savchenko, A.B., Peshenko, I.V., Barrett, R, Bush, R.A., Sieving, P,A, Fain, G.L., Diazoor, A.M. (2007). Constitutive excitation by Gly90Asp rhodopsin rescues rods from degeneration caused by elevated production of cyclic GMP in the dark J. Neurosci 27: 8805-8815

Leung, Y.-T., Fain, G.L., Matthews, H.R. (2007). Simultaneous measurement of current and calcium in the UV-sensitive cones of zebrafish J. Physiol 579: 15-27

Tsang, S.H., Woodruff, M.L., Janisch, K.M., Cilluffo, M.C, Farber, D.B., Fain, G.L. (2007). Removal of Phosphorylation Sites of Gamma Subunit of Phosphodiesterase6 Alters Rod Light Response J. Physiol 579: 303-312

Tsang, S.H., Woodruff, M.L., Chen, C.K., Yamashita, C.Y., Cilluffo, M.C., Rao, A.L., Farber, D.B., Fain, G.L. (2006). GAP-Independent termination of photoreceptor light response by excess gamma subunit of the c-GMP-phosphodiesterase J. Neurosci 26: 4472-4480

Fain, G.L. (2006). Why photoreceptors die (and why they don't) Bioessays 28: 344-354

Fan, J., Woodruff, M.L., Cilluffo, M.C., Crouch, R.K., Fain G.L. (2005). Opsin activation of transduction in the rods of dark-reared Rpe65 knockout mice J. Physiol 568: 83-95

Olshevskaya, E.V., Calvert, P.D., Woodruff, M.L., Peshenko, I.V., Savchenko, A.B., Makino, C.L., Ho, Y.-S., Fain, G.L. & Dizhoor, A.M. (2004). The Y99C mutation in guanylyl cyclase activating protein 1 increases intracellular free Ca2+ concentration and causes photoreceptor degeneration in transgenic mice Journal of Neuroscience 24: 6078-6085

Fain, G.L. (2003). Sensory Transduction Journal of Neuroscience -

Woodruff, M.L., Wang, Z., Chung, H.Y., Redmond, T.M., Fain, G.L., & Lem, J. (2003). Spontaneious activity of opsin apoprotein is a cause of Leber cogenital amaurosis Nature Genetics 35: 158-164

Matthews, H.R. and Fain, G.L. (2003). The effect of light on outer segment calcium in salamander rods Journal of Physiology 552: 763-776

Matthews, H.R. & Fain, G.L. (2001). A light-dependent increase in free Ca2+ concentration in salamander rod outer segment J. Physiol 532: 305-321

Matthews, H.R. & Fain, G.L. (2000). Laser spot confocal technique to measure cytoplasmic calcium concentration in photoreceptors Methods in Enzymol 316: 146-163

Cornwall, M.C., Jones, G.J., Kefalov, V.J., Fain, G.L. & Matthews, H.R. (2000). Electrophysiological methods for measurement of activation of phototransduction by bleached visual pigment in salamander photoreceptors Methods in Enzymol 316: 224-252

Fain, G. (1999). Cellular and Molecular Physiology of Neurons Methods in Enzymol -

Fain, G.L. & Lisman, J.E. (1999). Light, Ca 2+, and photoreceptor death: new evidence for the equivalent light hypothesis from arrestin knockout mice Invest. Ophthal. Vis. Sci 40: 2770-2772

Sampath, A.P., Matthews, H.R., Cornwall, M.C., Bandarchi, J. & Fain, G.L. (1999). Light-dependent changes in outer segment free Ca2+ concentration in salamander cone photoreceptors J. Gen. Physiol 113: 267-277

Fain, G.L. & Lisman, J.E. (1999). Visual Transduction Encyclopedia of Neuroscience -

Cilluffo, M.C.,Xia, S.-L., Farahbakhsh, N.A. & Fain, G.L. (1998). Synergistic receptor-activated calcium increases in single nonpigmented epithelial cells Invest. Ophthalmol. Vis. Sci 39: 1429-1435

Sampath, A.P., Matthews,H.R., Cornwall, M.C. & Fain, G.L. (1998). Bleached pigment produces a maintained decrease in outer segment Ca2+ in salamander rods J. Gen. Physiol 111: 53-64

Cilluffo, M.C., Farahbakhsh, N.A. & Fain, G.L. (1997). Functional and morphological differentiation of nonpigmented ciliary body epithelial cells grown on collagen rafts In Vitro Cell Dev Biol-Animal 33: 546-552

Xia, S.-L., Fain, G.L. & Farahbakhsh, N.A. (1997). Synergistic rise in Ca2+ produced by somatostatin and acetylcholine in ciliary body epithelial cells Exp. Eye Res 64: 627-635

Matthews, H.R., Cornwall, M.C. & Fain, G.L. (1996). Persistent activation of transducin by bleached rhodopsin in salamander rods J. Gen. Physiol 108: 557-563

Jones, G.J., M.C. Cornwall & G.L. Fain (1996). Equivalence of background and bleaching desensitization in isolated rod photoreceptors of the larval tiger salamander J. Gen. Physiol 108: 333-340

Zhou, Z.J. & Fain, G.L. (1996). Starburst amacrine cells change from spiking to non-spiking neurons during retinal development Proc.Natl. Acad. Sci 93: 8057-8062

Fain, G.L., Matthews, H.R. & Cornwall, M.C. (1996). Dark adaptation in vertebrate photoreceptors Trends in Neurosciences 19: 502-507

Matthews, H.R., Fain, G.L. & Cornwall, M.C. (1996). Role of cytoplasmic calcium concentration in the bleaching adaptation of salamander cone photoreceptors Journal of Physiology 490: 293-303

Lisman, J. & Fain, G. (1995). New support for the equivalent light hypothesis for retinitis pigmentosa Nature Medicine 1: 1254-1255

Zhou, Z.J. & Fain, G.L. (1995). Neurotransmitter receptors of starburst amacrine cells in rabbit retinal slices J. Neuroscience 15: 5334-5345

Cornwall, M.C., Matthews, H.R., Crouch, R.K. & Fain, G.L. (1995). Bleached pigment activates transduction in salamander cones J. Gen. Physiol 106: 543-557

Cohen, E.D., Zhou, Z.J. and Fain, G.L. (1994). Ligand-gated currents of alpha and beta ganglion cells in the cat retinal slice J. Neurophysiol 72: 1260-1269

Cornwall, M.C. and Fain, G.L. (1994). Bleached pigment activates transduction in isolated rods of the salamander retina J. Physiol 480: 261-279

Zhou, Z.J., Marshak, D.W. and Fain, G.L. (1994). Amino acid receptors of midget and parasol ganglion cells in primate retina Proc. Natl. Acad. Sci 91: 4907-4911

Cilluffo, M.C., Fain, M.J. and Fain, G.L. (1993). Tissue culture of rabbit ciliary body epithelial cells on permeable supports Exp. Eye Res 57: 513-526

Farahbakhsh, N.A., Cilluffo, M.C., Chronis, C. and Fain, G.L (1993). Dihydropyridine-sensitive Ca2+ spikes and Ca2+ currents in rabbit ciliary body epithelial cells Exp. Eye Res 58: 197-206

Fain, G.L. and Lisman, J.E. (1993). Photoreceptor degeneration in vitamin A deprivation and Retinitis Pigmentosa: The equivalent light hypothesis Exp. Eye Res 70: 335-340

Fain, G.L. and Cornwall, M.C. (1993). Light and dark adaptation in vertebrate photoreceptors In: Contrast Sensitivity: From Receptors to Clinic 3-32

Zhou, Z.J., Fain, G.L. and Dowling, J.E. (1992). Excitatory and inhibitory amino acid receptors on horizontal cells isolated from the white perch retina J. Neurophysiol 70: 8-19

Yazejian, B. and Fain, G.L. (1992). Whole-cell currents activated by nicotinic acetylcholine receptors on ganglion cells isolated from goldfish retina Visual Neurosci 10: 353-361

Yazejian, B. and Fain, G.L. (1992). Excitatory amino acid receptors on isolated retinal ganglion cells from the goldfish J. Neurophysiol 46: 94-107

Cilluffo, M.C., Cohen, B.N. and Fain, G.L. (1991). Nonpigmented cells of the rabbit ciliary body epithelium: tissue culture and voltage-gated currents Invest. Ophthalmol. Vis. Sci 32: 1619-1629

Fain, G.L. and Matthews, H.R. (1990). Calcium and the mechanism of light adaptation in vertebrate photoreceptors Trends in Neuroscience 13: 378-384

Frambach, D.A., Fain, G.L., Farber, D.B., and Bok, D. (1990). Beta adrenergic receptors on cultured human retinal pigment epithelium Invest. Ophthalmol. Vis. Sci 31: 1767-1772

Fain, G.L. and Schröder, W.H. (1990). Light-induced calcium release and re-uptake in toad rods J. Neurosci 10: 2238-2249

Matthews, H.R., Fain, G.L., Murphy, R.L.W. and Lamb, T.D (1990). Light adaptation in cone photoreceptors of the salamander: a role for cytoplasmic calcium J. Physiol 420: 447-469

Fain, G.L. and Farahbakhsh, N.A. (1989). Voltage-activated currents recorded from rabbit pigmented ciliary body epithelial cells in culture J. Physiol 418: 83-102

Cohen, B.N., Fain, G.L. and Fain, M.J. (1989). GABA and glycine channels in isolated ganglion cells from the goldfish retina J. Physiol 418: 53-82

Fain, G.L., Lamb, T.D., Matthews, H.R. and Murphy, R.L.W. (1989). Cytoplasmic calcium as the messenger for light adaptation in salamander rods J. Physiol 416: 215-243

Matthews, H.R., Murphy, R.L.W., Fain, G.L. and Lamb, T.D. (1988). Photoreceptor light adaptation is mediated by cytoplasmic calcium concentration Nature 334: 67-69

Fain, G. L., Cilluffo, M.C., Fain, M.J. and Lee, D.A. (1988). Isolation of non-pigmented epithelial cells from rabbit ciliary body Invest. Ophthalmol. Vis. Sci 29: 785-794

Fain, G.L., Smolka, A., Cilluffo, M.C., Fain, M.J., Lee, D.A., Brecha, N.C. and Sachs, G. (1988). Monoclonal antibodies to the H+-K+ ATPase of gastric mucosa selectively stain the non-pigmented cells of the rabbit ciliary body epithelium Invest. Ophthalmol. Vis. Sci 29: 817-821

Fain, G.L. and Schröder, W.H. (1988). Calcium in photoreceptors In: Ion Channel Modulation, A Symposium in Memory of Roger O. Eckert 103-118

Usukura, J., Fain, G.L. and Bok, D. (1988). [3H]ouabain localization of Na-K ATPase in the epithelium of rabbit ciliary body pars plicata Invest. Ophthalmol. Vis. Sci 29: 606-614

Fox, J.A., Pfeffer, B.A. and Fain, G.L. (1988). Single-channel recordings from cultured human retinal pigment epithelial cells J. Gen. Physiol 91: 193-222

Farahbakhsh, N.A. and Fain, G.L. (1987). Volume regulation of non-pigmented cells from ciliary epithelium Invest. Ophthalmol. Vis. Sci 28: 934-944

Fain, G.L. and Schröder, W.H. (1987). Calcium in dark-adapted toad rods: evidence for pooling and cyclic-guanosine-3=5=-monophosphate-dependent release J. Physiol 389: 361-384

Fain, G.L. and Hubbell, W.L. (1987). Visual transduction Encyclopedia of Neuroscience. Birkhauser, Boston 1297-1300

Fain, G.L. (1986). Evidence for a role of messenger substances in phototransduction Molecular Mechanism of Photoreception. Dahlem Konferenzen, Berlin 67-77

Fain, G.L. and Schröder, W.H. (1985). Calcium content and calcium exchange in dark-adapted toad rods J. Physiol 368: 641-665

Fain, G.L. and Schröder, W.H. (1985). Calcium content and light-induced release from photoreceptors: measurements with laser micro-mass analysis Contemporary Sensory Neurobiology 3-20

Schröder, W.H. and Fain, G.L. (1984). Light-dependent calcium release from photoreceptors measured by laser micro-mass analysis Nature 309: 268-270

Fain, G.L., Ishida, A.T. and Callery, S. (1983). Mechanisms of synaptic transmission in the retina Vision Res 23: 1239-1249

Woodruff, M.L. and Fain, G.L. (1982). Ca2+ -dependent changes in cyclic GMP levels are not correlated with opening and closing of the light-dependent permeability of toad photoreceptors J. Gen. Physiol 80: 537-555

Woodruff, M.L., Fain, G.L. and Bastian, B.L. (1982). Light-dependent ion influx into toad photoreceptors J. Gen. Physiol 80: 517-536

Bastian, B.L. and Fain, G.L. (1982). The effects of sodium replacement on the responses of toad rods J. Physiol 330: 331-347

Bastian, B.L. and Fain, G.L. (1982). The effects of low calcium and background light on the sensitivity of toad rods J. Physiol -

Lisman, J.E., Fain, G.L. and O?Day, P.M. (1982). Voltage-dependent conductances in Limulus ventral photoreceptors J. Gen. Physiol 79: 187-209

Bastian, B.L. and Fain, G.L. (1981). Calcium and the mechanism of light adaptation in rods J. Gen. Physiol 351-368

Ishida, A.T. and Fain, G.L. (1981). D-aspartate potentiates the effects of L-glutamate on horizontal cells in goldfish retina Proc. Natl. Acad. Sci. USA 78: 5890-5894

Fain, G.L. and Lisman, J.E. (1981). Membrane conductances of photoreceptors Prog. Biophys. Molec. Biol 37: 91-147

Fain, G.L. and Quandt, F.N. (1981). The effects of tetraethylammonium and cobalt ions on responses to extrinsic current in toad rods J. Physiol 303: 515-533

Fain, G.L., Gerschenfeld, H.M. and Quandt, F.N. (1980). Calcium spikes in toad rods J. Physiol 495-513

Fain, G.L. (1980). Integration by spikeless neurones in the retina Neurones Without Impulses: Their Significance in Vertebrate and Invertebrate Nervous Systems. Cambridge University Press, London 29-59

Bastian, B.L. and Fain, G.L. (1979). Light adaptation in toad rods: requirement for an internal messenger which is not calcium J. Physiol 297: 493-520

Fain, G.L. (1979). Le fonctionnement de la rétine La Recherche 10: 355-362

Fain, G.L., Quandt, F.N., Bastian, B.L. and Gerschenfeld, H.M. (1978). Contribution of a caesium-sensitive conductance increase to the rod photoresponse Nature 272: 467-469

Fain, G.L., Quandt, F.N. and Gerschenfeld, H.M. (1977). Calcium-dependent regenerative responses in rods Nature 269: 707-710

Fain, G.L. (1977). The threshold signal of photoreceptors Vertebrate Photoreception, a symposium of the Royal Society of London 305-323

Fain, G.L., Granda, A.M. & Maxell, J.H. (1977). Voltage signal of photoreceptors at visual threshold Nature 265: 181-183

Fain, G.L. (1976). Sensitivity of toad rods: dependence on wave-length and background illumination J. Physiol 261: 71-101

Fain, G.L., Gold, G.H. and Dowling, J.E. (1976). Receptor coupling in the toad retina Cold Spr. Harb. Symp. Quant. Biol 40: 547-561

Fain, G.L. (1975). Interactions of rod and cone signals in the mudpuppy retina J. Physiol 252: 735-769

Fain, G.L. (1975). Quantum sensitivity of rods in the toad retina Science 187: 838-841

Fain, G.L. (1973). Responses of rods and cones and the interactions of rods and cone signals in the mudpuppy retina The Johns Hopkins University -

Fain, G.L. and Dowling, J.E. (1973). Intracellular recordings from single rods and cones in the mudpuppy retina Science 180: 1178-1181