Harvard University - Department of Molecular & Cellular Biology


Gordon and Llura Gund Professor of Neurosciences, Emeritus

Email: dowling@mcb.harvard.edu
Phone: 617-495-2245

Mail: NW 347.20
Northwest Building
52 Oxford St
Cambridge, MA  02138

Members of the Dowling Lab
List of Publications from PubMed


MCB 365. Neurobiology
Catalog Number: 8349  View Course Website
Term: Fall Term And Spring Term 2014-2015.
Instructor: John Dowling
Course Level: Exclusively for Graduates
BIOPHYS 317. Biophysical Aspects of the Visual System
Catalog Number: 4770  View Course Website
Term: Fall Term And Spring Term 2014-2015.
Instructor: John Dowling
Course Level: Exclusively for Graduates
(View all MCB Courses)


The vertebrate retina is an accessible part of the central nervous system; an understanding of retinal mechanisms should provide clues concerning neural mechanisms throughout the brain. Over the years, our group has been concerned with the cells of the retina, their structure, function and synaptic interactions.

Our current interests are focused in two directions; both employ the zebrafish as a model system First, we have developed behavioral tests to isolate visual system specific mutations from chemically-mutagenized zebrafish. Both recessive and dominant mutations that affect the retina have been isolated and are currently being analyzed histologically, electrophysiologically and biochemically. Ongoing screening is continually revealing new mutants.

A second interest is the molecular basis of retinal development. In particular, the effects of retinoic acid on candidate genes involved in early eye and photoreceptor development are being explored as well as detailed examination of early retinal development in zebrafish.


Schmitt, E. A and Dowling, J. E. (l994) Early eye morphogenesis in the zebrafish, Brachydanio rerio, J. Comp. Neurol., 344, 532-542.

Brockerhoff, S. E., Hurley, J. B., Janssen-Bienhold, U., Neuhauss, S. C., Driever, W. and Dowling, J. E. (l995) A behavioral screen for isolating zebrafish mutants. Proc. Natl. Acad. Sci., 92, 10545-10549.

Schmitt, E. A. and Dowling, J. E. (1996) A comparison of topographical patterns of ganglion and photoreceptor cell differentiation in the retina of the zebrafish, Danio rerio, J. Comp. Neurol., 371, 222-234.

Hyatt, G. A., Schmitt, E. A., Fadool, J. M. and Dowling, J. E. (1996) Retinoic acid alters photoreceptor development in vivo. Proc. Natl. Acad. Sci., 93, 13298-13303.

Brockerhoff, S. E., Hurley, J. B., Niemi, G. A. and Dowling, J. E. (1997) A new form of inherited red-blindness in zebrafish. J. Neurosci., 17, 4236-4242.

Fadool, J. M., Brockerhoff, S. E., Hyatt, G. A. and Dowling, J. E. (1997) Mutations affecting eye morphology in the developing zebrafish (Danio rerio). Develop. Genetics, 20, 288-295.

Li, L. and Dowling, J. E. (1997) A dominant form of inherited retinal degeneration caused by a non-photoreceptor cell-specific mutation, Proc. Natl. Acad. Sci., 94, 11645-11650.

Fadool, J.M., Hartl, D.L. and Dowling, J.E. (1998) Transposition of the mariner element Drosophila mauritiana in zebrafish. Proc. Natl. Acad. Sci., 95, 5182-5186.

Li, L. and Dowling, J.E. (1998) Zebrafish visual sensitivity is regulated by a circadian clock. Visual Neuroscience, 15:851-857.

Qian, H., Dowling, J. E. and Ripps, H. (1999) A single amino acid in the second transmembrane domain of GABA rho subunits is a principal determinant of the response kinetics of GABAc receptors.  J. Neurobiol. 40, 67-76.

Li, L. and Dowling, J. E. (2000) Disruption of the olfactoretinal centrifugal pathway may relate to the visual system defect in night blindness b mutant zebrafish. J. Neurosci. 20:1883-1892.

Li. L. and Dowling, J. E. (2000) Effects of dopamine depletion on visual sensitivity of zebrafish. J. Neurosci. 20:1893-1903.

Link, B. A., Fadool, J. M., Malicki, J. and Dowling, J. E. (2000) The zebrafish young mutation acts non-cell autonomously to uncouple differentiation from specifications for all retinal cells. Development, 127, 2177-2188.

Peterson, R.T., Link, B. A., Dowling, J. E. and Schreiber, S. L. (2000) Small molecule developmental screens reveal the logic and timing of vertebrate development, Proc. Natl. Acad. Sci., 97, 12965-12969.

Allwardt, B., Lall, A. B., Brockerhoff, S. E. and Dowling, J. E. (2001) Synapse formation is arrested in retinal photoreceptors of the zebrafish nrc mutant. J. Neurosci., 21, 2330-2342.

Link, B. A., Kainz, P. M., Ryou, T. and Dowling, J. E. (2001) The perplexed and confused mutations affect distinct stages during the transition from proliferating to post-mitotic cells within the zebrafish retina. Develop. Biol., 236, 436-453.

Allwardt, B. A. and Dowling, J. E. (2001) The pineal gland in wild-type and two mutant zebrafish with retinal defects. J. Neurocytol., 130, 493-501.
Darland, T. and Dowling, J. E.  (2001) Behavioral screening for cocaine sensitivity in mutagenized zebrafish.  Proc. Natl. Acad. Sci., 98, 11691-11696.

Perkins, B. D., Kainz, P. M., O’Malley, D. M. and Dowling, J. E.  (2002) Transgenic expression of a GFP-rhodopsin COOH-terminal fusion protein in zebrafish rod photoreceptors. Visual Neurosci., 19, 257-264.

Mangrum, W. I., Dowling, J. E. and Cohen, E. D. (2002) Ganglion cell types in the zebrafish retina.  Vis. Neurosci., 19, 767-779.

Gregg, R. G., Willer, G. B., Fadool, J. M., Dowling, J. E. and Link, B. A. (2003) Positional cloning of the young mutation identified an essential role for the Brahma chromatin remodeling complex in mediating retinal cell differentiation, Proc. Natl. Acad. Sci., 100, 6535-6540.

Kainz, P. M., Adolph, A. R., Wong, K. Y.  and Dowling, J. E. (2003) The lazy eye zebrafish mutation affects Müller glial cells, compromising photoreceptor function and causing partial blindness, J. Comp. Neurol., 463, 265-280.

Wong, K. Y., Gray, J., Hayward, C. J. C., Adolph, A. R. and Dowling, J. E. (2004) Glutamatergic mechanisms in the outer retina of larval zebrafish:  Analysis of electroretinogram b- and d-waves using a novel preparation, Zebrafish, 1 (2).      
Perkins, B. D., Fadool, J. M. and Dowling, J. E.  (2004) Photoreceptor Structure and Development Analyses using GFP Transgenes in Zebrafish: Cellular & Developmental Biology, 2nd edition, (Eds. H. W. Detrich, M. Westerfield, and L. I. Zon), (Vol. 76 of Methods in Cell Biology).
Wong, K. Y., Adolph, A. R. and Dowling, J. E. (2005) Retinal bipolar cells input mechanisms in giant danio: I. Electroretinographic analysis. J. Neurophysiol. 93, 84-93.       
Wong, K. Y., Cohen, E. D. and Dowling, J. E.  (2005) Retinal bipolar cell input mechanisms in giant danio:  II.  Patch-clamp analysis of ON bipolar cells.  J. Neurophysiol. 93, 94-107.
Wong, K. Y. and Dowling, J. E.  (2005) Retinal Bipoolar Cell Input Mechanisms in Giant Danio:  III.  ON-OFF Bipolar Cells and their Color Opponent,  J. Neurophysiol., 94, 265-272.
Gross, J. M., Perkins, B. D., Amsterdam, A., Egana, A., Darland, T., Matsui, J. I., Sciascia, S., Hopkins, N. and Dowling, J. E.  (2005) Identification of zebrafish insertional mutants with defects in visual system development and function, 170, 245-261, Genetics.
Perkins, B. D., Nicholas, C. S., Baye, L. M., Link, B. A. and Dowling, J. E.  (2005) The dazed  gene is necessary for late cell type development and retinal cell maintenance in the zebrafish retina.  Developmental Dynamics, 233, 680-694. 
Gross, J. M. and Dowling, J. E.  (2005) Tbx2b is essential for neuronal differentiation along the dorsal/ventral axis of the zebrafish retina.   Proc. Nat. Acad. Sci., 102, 4371-4376.          
Fadool, J. M. and Dowling, J. E.  (2005) Genetic analysis of retinal development using  zebrafish mutants, in “Retinal Development”, edited by Everlyne Sernagor,  Stephen Eglen, William Harris and Rachel Wong, Cambridge University Press,  pp. 342-370.
Qian, H., Zhu, Y, Ramsey, D. J., Chappell, R. L., Dowling, J. E., and Ripps, H.  (2005) Directional asymmetries in the optokinetic response of larval zebrafish (Danio rerio). Zebrafish, 2, 189-196.
Leung, Y. F. and Dowling, J. E.  (2005) Gene expression profiling of zebrafish embryonic retina.   Zebrafish, 2, 269-285.
Matsui, J. I., Egana, A. L., Sponholtz, T. R., Adolph, A. R. and Dowling, J. E. (2006) Effects of ethanol on photoreceptors and visual function in developing zebrafish.  Invest. Ophthal. Vis. Sci.,47, 4589-4597.

Leung Y.F., Ma, P., Dowling J.E. (2006) Gene expression profiling of zebrafish embryonic retinal pigment epithelium in vivo. Invest. Ophthalmol. Vis. Sci. 48, 881-890.

Emran, F. Rihel, J., Adolph, A., Wong, K. Y., Kraves, S. and Dowling, J. E. (2007) OFF- ganglion cells cannot drive the optokinetic reflex in zebrafish. Proc. Natl. Acad. Sci., 104, 19126-19131.

Song, P., Matsui, J. and Dowling, J. E. (2007) The morphological types and connectivity of horizontal cells found in the adult zebrafish (Danio rerio) retina. J. Comp. Neurol., 506, 328-338.

Fadool, J. M. and Dowling, J. E. (2008) Zebrafish: A model system for the study of eye genetics. Progress in Retinal and Eye Research. 27, 89-110.

Kojima, D., Torii, M. Fukada, Y. and Dowling, J. E. (2008) Differential expression of duplicated VAL-opsin genes in the developing zebrafish. J. Neurochem. 104, 1364-1371.

Leung, Y. F.. P. Ma, Link, B. A. and Dowling, J. E. (2008) Factorial microarray analysis of zebrafish retinal development, Proc. Natl. Acad. Sci., 105, 12909-12914./p>

Kojima, D., Dowling, J. E. and Fukada, Y. (2008) Probing pineal-specific gene expression with transgenic zebrafish. In: Photochemistry and Photobiology, Wiley- Blackwell, Inc., 84, 1011-1015.

Yong, N. L., Matsui, J. I. and Dowling, J. E. (2009) Specificity of the horizontal cell-photoreceptor connections in the zebrafish (Danio rerio) retina, J. Comp. Neurol. In press.

updated: 02/24/2015