Development and Disease



William Pak



Pak WL. 2010. Why Drosophila to study phototransduction. J. Neurogenet. 24(2): 55-66.

Watson GB, Chouinard SW, Cook KR, Geng C, , Gifford JM, Gustafson GD, Hasler JM, Larrinua IM, Letherer TJ, Mitchell JC, Pak WK, Salgado VL, Sparks TC, Stilwell GE. 2010. A spinosyn-sensitive Drosophila melanogaster nicotinic acetylcholine receptor identified through chemically induced target site resistance, resistance gene identification, and heterologous expression. Insect Biochem Mol Biol. 405(5): 376-384.

Lu, H., Leung,H-T., Wang, N., Pak, W.L., Shieh, B-H. 2009. Role of Ca²+/Calmodulin-dependent protein kinase II in Drosophila photoreceptors. J. Biol Chem. 284: 11100-11109.

Shahrestani P, Leung H-T, Le PK, Pak WL, Tse S, Ocorr K, Huang T. 2009. Heterozygous mutation of Drosophila Opal causes the development of multiple organ abnormalities in an age-dependent and organ-specific manner. PLos One. 4(8): e6867.

Wang, N., Leung, H-T., Pak, W. L., Carl, Y. T., Wadzinski, B. E., Shieh, B-H. 2008. Role of protein phosphatase 2A in regulating the visual signaling in Drosophila. J Neurosci. 28: 1444-1451.

Leung, H-T., Tseng-Crank, J., Kim, E., Mahapatra, C., Shino, S., Zhou, Y., An, L., Doerge, R. W., Pak, W. L. 2008. DAG lipase activity is necessary for TRP channel regulation in Drosophila photoreceptors. Neuron. 58: 884-896.

Long, A. A., Kim, E., Leung, H-T, Woodruff, E. III, An, L., Doerge, R. W., Pak, W. L., Broadie, K. 2008. Presynaptic calcium channel localization and calcium dependent synaptic vesicle exocytosis regulated by the Fuseless protein. Journal of Neuroscience. 28: 3668-3682.

Lee, S., Leung, H-T., Kim, E., Jang, J., Lee, E., Baek, K., Pak, W. L., Yoon, J. 2007. Effects of a mutation in the Drosophila porin gene encoding mitochondrial voltage-dependent anion channel protein on phototransduction. Develop Neurobiol. 67: 1533-1545.

Geng, C., Pellegrino, A., Bowman, J., Zhu, L., Pak, W. L. 2004. Complete RNAi rescue of neuronal degeneration in a constitutively active Drosophila TRP channel mutant. Biochim Biophys Acta. 1674: 91-97.

Yoon, J., Leung, H-T., Lee S., Geng, C., Kim, Y., Baek, K., Pak, W.L. 2004. Specific molecular alterations in the norpA-encoded phospholipase C of Drosophila and their effects on electrophysiological responses in vivo. J. Neurochem. 89: 998-1008.

Ben-Shahar, Y., Leung, H-T., Pak, W. L., Sokolowski, M. B., Robinson, G. E. 2003. cGMP-dependent changes in phototaxis: a possible role for the foraging gene in honey bee division of labor. J Exp Biol. 206: 2507-2515.

Pak, W. L., Leung, H-T. 2003. Genetic approaches to visual transduction in Drosophila melanogaster. Receptors and Channels. 9: 149-167.

Geng, C., Pak, W. L. 2002. Photoreceptor degeneration and Ca2+ influx through light-activated channels of Drosophila. Adv Exp Med Biol. 514: 585-599.

Geng, C., Leung, H-T., Skingsley, D. R., Iovchev, M. I., Yin, Z., Semenov, E. P., Burg, M. G., Hardie, R. C., Pak, W. L. 2002. Target of Drosophila photoreceptor synaptic transmission is a histamine-gated chloride channel encoded by ort (hclA). J Biol Chem. 277: 42113-42120.

Hong, Y. S., Park, S., Geng, C., Baek, K., Bowman, J. D., Yoon, J., Pak, W. L. 2002. Single amino acid change in the fifth transmembrane segment of the TRP Ca2+ channel causes massive degeneration of photoreceptors. J Biol Chem. 277: 33884-33889.

Iovchev, M., Kodrov, P., Wolstenholme, A. J., Pak, W. L., Semenov, E. P. 2002. Altered drug resistance and recovery from paralysis in Drosophila melanogaster with a deficient histamine-gated chloride channel. J Neurogenet. 16: 249-261.

Georgiev, P. G., Wolstenholme, A.J., Pak, W.L., Semenov, E. P. 2002. Differential responses to avermectins in ort mutants of Drosophila melanogaster. Pesticide Biochem. and Physiol. 72: 65-71.

Leung, H-T., Geng, C., Pak, W. L. 2000. Phenotypes of trpl mutants and interactions between the transient receptor potential (TRP) and TRP-like channels in Drosophila. J Neurosci. 20: 6797-6803.

Yoon, J., Ben-Ami, H. C., Hong, Y. S., Park, S., Strong, L. L. R., Bowman, J., Geng, C., Baek, K., Minke, B., Pak, W. L. 2000. Novel mechanism of massive photoreceptor degeneration caused by mutations in the trp gene of Drosophila. J Neurosci. 20: 649-659.

Li, C., Geng, C., Leung, H-T., Hong, Y. S., Strong, L. L. R., Schneuwly, S., Pak, W. L. 1999. INAF, a protein required for transient receptor potential Ca2+ channel function. Proc Natl Acad Sci U S A. 96: 13474-13479.

Semenov, E. P., Pak, W. L. 1999. Diversification of Drosophila chloride channel gene by multiple posttranscriptional mRNA modifications. J Neurochem. 72: 66-72.

Melzig, J., Burg, M., Gruhn, M., Pak, W. L., Buchner, E. 1998. Selective histamine uptake rescues photo- and mechanoreceptor function of histidine decarboxylase-deficient Drosophila mutant. J Neurosci. 18: 7160-7166.

Bentrop, J., Schwab, K., Pak, W. L., Paulsen, R. 1997. Site-directed mutagenesis of highly conserved amino acids in the first cytoplasmic loop of Drosophila Rh1 opsin blocks rhodopsin synthesis in the nascent state. EMBO J. 16: 1600-1609.

Burg, M. G., Geng, C., Guan, Y., Koliantz, G., Pak, W. L. 1996. Drosophila rosA gene, which when mutant causes aberrant photoreceptor oscillation, encodes a novel neurotransmitter transporter homologue. J Neurogenetics. 11: 59-79.

Ferreira, P. A., Nakayama, T. A., Pak, W. L., Travis, G. H. 1996. Cyclophilin-related protein RanBP2 acts as chaperone for red/green opsin. Nature. 383: 637-640.

Melzig, J., Buchner, S., Wiebel, F., Wolf, R., Burg, M., Pak, W. L., Buchner, E. 1996. Genetic depletion of histamine from the nervous system of Drosophila eliminates specific visual and mechanosensory behavior. J Comp Physiol A. 179: 763-773.

Pearn, M. T., Randall, L. L., Shortridge, R. D., Burg, M. G., Pak, W. L. 1996. Molecular, biochemical, and electrophysiological characterization of Drosophila norpA mutants. J Biol Chem. 271: 4937-4945.

Pak, W.L. 1996. Use of Drosophila mutants in vision research. Mol. Cells. 6: 117-124.

Ferreira, P. A., Hom, J. T., Pak, W. L. 1995. Retina-specifically expressed novel subtypes of bovine cyclophilin. J Biol Chem. 270: 23179-23188.

Liu, W., Yoon, J., Burg, M., Chen, L., Pak, W. L. 1995. Molecular characterization of two Drosophila guanylate cyclases expressed in the nervous system. J Biol Chem. 270: 12418-12427.

Pak, W. L. 1995. Drosophila in vision research: The Friedenwald Lecture. Invest Ophthalmol Vis Sci. 36: 2340-2357.

Ferreira, P. A., Pak, W. L. 1994. Bovine phospholipase C highly homologous to the NorpA protein of Drosophila is expressed specifically in cones. J Biol Chem. 269: 3129-3131.

Matsumoto, H., Kurien, B. T., Takagi, Y., Kahn, E. S., Kinumi, T., Komori, N., Yamada, T., Hayashi, F., Isono, K., Pak, W. L., Jackson, K. W., Tobin, S. 1994. Phosrestin I undergoes the earliest light-induced phosphorylation by a calcium/calmodulin-dependent protein kinase in Drosophila photoreceptors. Neuron. 12: 997-1010.

Buchner, E., Buchner, S., Burg, M. G., Hofbauer, A., Pak, W. L., Pollack, I. 1993. Histamine is a major mechanosensory neurotransmitter in Drosophila. Cell Tissue Res. 273: 119-125.

Burg, M. G., Sarthy, P. V., Koliantz, G., Pak, W. L. 1993. Genetic identification and cDNA cloning of a Drosophila histidine decarboxylase gene and its requirement for photoreceptor transmitter synthesis. EMBO J. 12: 911-919.

Ferreira, P. A., Shortridge, R. D., Pak, W. L. 1993. Distinctive subtypes of bovine phospholipase C that have preferential expression in the retina and high homology to the norpA gene product of Drosophila. Proc Natl Acad Sci U S A. 90: 6042-6056.

Leonard, D. S., Bowman, V. D., Ready, D. F., Pak, W. L. 1992. Degeneration of photoreceptors in rhodopsin mutants of Drosophila. J Neurobiol. 23: 605-626.

Pak, W. L. 1991. Molecular genetic studies of photoreceptor function using Drosophila mutants. Prog Clin Biol Res. 362: 1-32.

Schneuwly, S., Burg, M. G., Lending, C.R., Perdew, M. H., Pak, W. L. 1991. Properties of photoreceptor-specific phospholipase C encoded by the norpA gene of Drosophila melanogaster. J Biol Chem. 266: 24314-24319.

Shortridge, R. D., Yoon, J., Lending, C. R., Bloomquist, B. T., Perdew, M. H., Pak, W. L. 1991. A Drosophila phospholipase C gene that is expressed in the central nervous system. J Biol Chem. 266: 12474-12480.

Pak, W.L., Shortridge, R.D. 1991. Inositol phospholipid and invertebrate photoreceptors. Photochem Photobiol. 53: 871-875.

Pflugfelder, G. O., Schwarz, H., Roth, H., Poeck, B., Sigl, A., Kerscher, S., Jonschker, B., Pak, W. L., Heisenberg, M. 1990. Genetic and molecular characterization of the optomotor-blind gene locus in Drosophila melanogaster. Genetics. 126: 91-104.

O'Tousa, J. E., Leonard, D. S., Pak, W. L. 1989. Morphological defects in oraJK84 photoreceptors caused by mutation in R1-6 opsin gene of Drosophila. J Neurogenet. 6: 41-52.

Schneuwly, S., Shortridge, R. D., Larrivee, D. C., Ono, T., Ozaki, M., Pak, W. L. 1989. Drosophila ninaA gene encodes an eye-specific cyclophilin (cyclosporine A binding protein). Proc Natl Acad Sci U S A. 86: 5390-5394.

Yoon, J., Shortridge, R. D., Bloomquist, B. T., Schneuwly, S., Perdew, M. H., Pak, W. L. 1989. Molecular characterization of Drosophila gene encoding G0 alpha subunit homolog. J Biol Chem. 264: 18536-18543.

Bloomquist, B. T., Shortridge, R. D., Schneuwly, S., Perdew, M., Montell, C., Steller, H., Rubin, G., Pak, W. L. 1988. Isolation of a putative phospholipase C gene of Drosophila, norpA, and its role in phototransduction. Cell. 54: 723-733.

Pak, W. L., O'Tousa, J. E. 1988. Molecular analysis of visual pigment genes. Photochem Photobiol. 47: 877-882.

Matsumoto, H., Isono, K., Pye, Q. N., Pak, W. L. 1987. Gene encoding cytoskeletal proteins in Drosophila rhabdomeres. Proc Natl Acad Sci U S A. 84: 985-989.

Banga, S. S., Bloomquist, B. T., Brodberg, R. K., Pye, Q. N., Larrivee, D. C., Mason, J. M., Boyd, J. B., Pak, W. L. 1986. Cytogenetic characterization of the 4BC region on the X chromosome of Drosophila melanogaster: Localization of the mei-9, norpA and omb genes. Chromosoma. 93: 341-346.

Johnson, E. C., Pak, W. L. 1986. Electrophysiological study of Drosophila rhodopsin mutants. J Gen Physiol. 88: 651-673.

Pak, W. L. 1986. Genes encoding visual pigments. Photobiochemistry and Photobiophysics. 13: 229-244.

Homyk, T., Jr., Isono, K., Pak, W. L. 1985. Developmental and physiological analysis of a conditional mutation affecting photoreceptor and optic lobe development in Drosophila melanogaster. J Neurogenet. 2: 309-324.

Nichols, R., Pak, W. L. 1985. Characterization of Drosophila melanogaster rhodopsin. J Biol Chem. 260: 12670-12674.

O'Tousa, J. E., Baehr, W., Martin, R. L., Hirsh, J., Pak, W. L., Applebury, M. L. 1985. The Drosophila ninaE gene encodes an opsin. Cell. 40: 839-850.

Matsumoto, H., Pak, W. L. 1984. Light-induced phosphorylation of retina-specific polypeptides of Drosophila in vivo. Science. 223: 184-186.

Scavarda, N. J., O'Tousa, J., Pak, W. L. 1983. Drosophila locus with gene dosage effects on rhodopsin. Proc Natl Acad Sci U S A. 80: 4441-4445.

Stephenson, R. S., O'Tousa, J., Scavarda, N. J., Randall, L. L., Pak, W. L. 1983. Drosophila mutants with reduced rhodopsin content. Symp Soc Exp Biol. 36: 477-501.

Matsumoto, H., O'Tousa, J., Pak, W. L. 1982. Light-induced modification of Drosophila retinal polypeptides in vivo. Science. 217: 839-841.

Pak, W. L. 1982. Relation of bleaching to sensitivity of vertebrate photoreceptors. Methods Enzymol. 81: 423-431.

Schinz, R. H., Lo, M-VC., Larrivee, D. C., Pak, W. L. 1982. Freeze-fracture study of the Drosophila photoreceptor membrane: Mutations affecting membrane particle density. J Cell Biol. 93: 961-969.

Larrivee, D. C., Conrad, S. K., Stephenson, R. S., Pak, W. L. 1981. Mutations that selectively affect rhodopsin concentration in the peripheral photoreceptors of Drosophila melanogaster. J Gen Physiol. 78: 521-545.

Lo, M-VC., Pak, W. L. 1981. Light-induced pigment granule migration in the retinular cells of Drosophila melanogaster. Comparison of wild-type with ERG-defective mutants. J Gen Physiol. 77: 155-175.

Lo, M-VC., Wong, F., Pak, W. L. 1980. Increase in intracellular free calcium concentration of Limulus photoreceptors caused by a metabolic inhibitor. Vision Res. 20: 539-544.

Pak, W. L., Conrad, S. K., Kremer, N. E., Larrivee, D. C., Schinz, R. H., Wong, F. 1980. Photoreceptor function. Basic Life Sci. 16: 331-346.

Stephenson, R. S., Pak, W. L. 1980. Heterogenic components of a fast electrical potential in Drosophila compound eye and their relation to visual pigment photoconversion. J Gen Physiol. 75: 353-379.

Lo, M-VC., Pak, W. L. 1978. Desensitization of peripheral photoreceptors shown by blue-induced decrease in transmittance of Drosophila rhabdomeres. Nature. 273: 772-774.

Waloga, G., Pak, W. L. 1978. Ionic mechanism for the generation of horizontal cell potentials in isolated axolotl retina. J Gen Physiol. 71: 69-92.

Wu, C-F., Pak, W. L. 1978. Light-induced voltage noise in the photoreceptor of Drosophila melanogaster. J Gen Physiol. 71: 249-268.

Pak, W. L., Pinto, L. H. 1976. Genetic approach to the study of the nervous system. Annu Rev Biophys Bioeng. 5: 397-448.

Waloga, G., Pak, W. L. 1976. Horizontal cell potentials: Dependence on external sodium ion concentration. Science. 191: 964-966.

Wong, F., Wu, C-F., Mauro, A., Pak, W. L. 1976. Persistence of prolonged light-induced conductance change in arthropod photoreceptors on recovery from anoxia. Nature. 264: 661-664.

Pak, W.L., Ostroy, S.E., Deland, M.C., Wu C-F. 1976. Photoreceptor mutant of Drosophila: Is protein involved in intermediate steps of phototransduction. Science. 194: 956-959.

Grabowski, S. R., Pak, W. L. 1975. Intracellular recordings of rod responses during dark-adaptation. J Physiol. 247: 363-391.

Minke, B., Wu, C-F., Pak, W. L. 1975. Induction of photoreceptor voltage noise in the dark in Drosophila mutant. Nature. 258: 84-87.

Wu, C-F., Pak, W. L. 1975. Quantal basis of photoreceptor spectral sensitivity of Drosophila melanogaster. J Gen Physiol. 66: 149-168.

Minke, B., Wu, C-F., Pak, W.L. 1975. Isolation of light-induced response of the central retinula cells from the electroretinogram of Drosophila. J. Comp. Physiol.. 98: 345-355.

Ostroy, S. E., Pak, W. L. 1974. Protein and electroretinogram changes in the alleles of the norpAP12 Drosophila phototransduction mutant. Biochim Biophys Acta. 368: 259-268.

Ostroy, S. E., Wilson, M., Pak, W. L. 1974. Drosophila rhodopsin: Photochemistry, extraction and differences in the norpAP12 phototransduction mutant. Biochem Biophys Res Commun. 59: 960-966.

Pak, W. L., Lidington, K. J. 1974. Fast electrical potential from a long-lived, long-wavelength photoproduct of fly visual pigment. J Gen Physiol. 63: 740-756.

Pinto, L. H., Pak, W. L. 1974. Light-induced changes in photoreceptor membrane resistance and potential in Gecko retinas. II. Preparations with active lateral interactions. J Gen Physiol. 64: 49-69.

Pinto, L. H., Pak, W. L. 1974. Light-induced changes in photoreceptor membrane resistance and potential in Gecko retinas. I. Preparations treated to reduce lateral interactions. J Gen Physiol. 64: 26-48.

Deland, M. C., Pak, W. L. 1973. Reversibly temperature sensitive phototransduction mutant of Drosophila melanogaster. Nat New Biol. 224: 184-186.

Ostroy, S. E., Pak, W. L. 1973. Protein differences associated with a phototransduction mutant of Drosophila. Nat New Biol. 243: 120-121.

Alawi, A. A., Jennings, V., Grossfield, J., Pak, W. L. 1972. Phototransduction mutants of Drosophila melanogaster. Adv Exp Med Biol. 24: 1-21.

Grabowski, S. R., Pinto, L. H., Pak, W. L. 1972. Adaptation in retinal rods of axolotl: Intracellular recordings. Science. 176: 1240-1243.

Alawi, A. A., Pak, W. L. 1971. On-transient of insect electroretinogram: Its cellular origin. Science. 172: 1055-1057.

Murakami, M., Pak, W. L. 1970. Intracellularly recorded early receptor potential of vertebrate photoreceptors. Vision Res. 10: 965-975.

Pak, W. L., Grossfield, J., Arnold, K. S. 1970. Mutants of the visual pathway of Drosophila melanogaster. Nature. 227: 518-520.

Pak, W. L., Grossfield, J., White, N. V. 1969. Nonphototactic mutants in a study of vision in Drosophila. Nature. 222: 351-354.

Pak, W. L. 1968. Rapid photoresponses in the retina and their relevance to vision research. Photochem Photobiol. 8: 495-503.

Pak, W. L., Helmrich, H. G. 1968. Absence of photodichroism in the retinal receptors. Vision Res. 8: 585-589.

Pak, W. L., Boes, R. J. 1967. Rhodopsin: Responses from transient intermediates formed during its bleaching. Science. 155: 1131-1133.

Pak, W. L., Ebrey, T. G., Rozzi, V.P. 1967. Effect of changes in the chemical environment of the retina on the two components of the early receptor potential. Nature. 214: 109-110.

Pak, W. L., Ebrey, T. G. 1966. Early receptor potentials of rods and cones in rodents. J Gen Physiol. 49: 1199-1208.

Pak, W. Y. 1966. [Military and civil nurses project report. Nursing care of schizophrenia]. Taehan Kanho. 5: 45-50.

Pak, W. L. 1965. Some properties of the early electrical response in the vertebrate retina. Cold Spring Harb Symp Quant Biol. 30: 493-499.

Pak, W. L., Ebrey, T. G. 1965. Visual receptor potential observed at sub-zero temperatures. Nature. 205: 484-486.

Pak, W. L., Cone, R. A. 1964. Isolation and identification of the initial peak of the early receptor potential. Nature. 204: 836-838.

Pak, WL;, Greisen, KI. 1962. Nuclear interactions of high energy cosmic rays. Phys. Rev. 126: 1668-1670.

Pak, WL, Ozaki, S, Roe, BP, Greisen, KI. 1961. Momentum spectrum of muons. Physical Review. 121: 905-907.


Long AA, Mahapatra CT, Woodruff EA III, Rohrbough J, Leung H-T, Shino S, An L, Doerge RW, Metzstein MM, Pak WL, Broadie K. 2010. The nonsense-mediated decay pathway maintains synapse architecture and synaptic vesicle cycle efficacy. J Cell Sci. 123: 3303-3315.