David Copenhagen, PhD
Professor
Ophthalmology
School of Medicine

Grants and Projects

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• P30 - Core Grant for Vision Research, NIH, 1977-2024
• Melanopsin-mediated light responses in the embryonic retina, NIH, 2015-2018
• Light regulated vascular development of the eye, NIH, 2013-2017
• Training Program for the Visual Sciences, NIH, 1990-2017
• Synaptic Interactions and Mechanisms in the Retina, NIH, 1977-2015
• Function and development in the synapse, NIH, 1980-2009
• Synaptic Interactions and Mechanisms in the Retina, NIH, 1977-2004

Publications (103)

Top publication keywords:
Photoreceptor Cells2-Amino-5-phosphonovalerateRetinal Rod Photoreceptor CellsRod OpsinsSynapsesAmbystomaTrigeminal GanglionRetinal Ganglion CellsSynaptic TransmissionVision, OcularRetinaLight Signal TransductionVisual PathwaysCalcium-Transporting ATPasesLight

• Light responses of melanopsin-expressing ganglion cells in the foetal mammalian retina.
  bioRxiv 2019 Jan Verweij, Shawnta Y. Chaney, Derek Bredl, Shruti Vemaraju, Gabriele M. König, Evi Kostenis, Richard A. Lang, David R. Copenhagen
• Fetal stage melanopsin (OPN4) and GNAQ (Gaq) signaling regulates vascular development of the eye.
  bioRxiv 2019 Shruti Vemaraju, Gowri Nayak, William E Miller, David R Copenhagen, Richard A Lang
• Current understanding of photophobia, visual networks and headaches.
  Cephalalgia : an international journal of headache 2018 Noseda R, Copenhagen D, Burstein R
• Melanopsin expression in the cornea.
  Visual neuroscience 2018 Delwig A, Chaney SY, Bertke AS, Verweij J, Quirce S, Larsen DD, Yang C, Buhr E, VAN Gelder R, Gallar J, Margolis T, Copenhagen DR
• Retinofugal Projections from Melanopsin-Expressing Retinal Ganglion Cells Revealed by Intraocular Injections of Cre-Dependent Virus.
  PloS one 2016 Delwig A, Larsen DD, Yasumura D, Yang CF, Shah NM, Copenhagen DR

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• Glutamatergic neurotransmission from melanopsin retinal ganglion cells is required for neonatal photoaversion but not adult pupillary light reflex.
  PloS one 2013 Delwig A, Majumdar S, Ahern K, LaVail MM, Edwards R, Hnasko TS, Copenhagen DR
• Length of day during early gestation as a predictor of risk for severe retinopathy of prematurity.
  Ophthalmology 2013 Yang MB, Rao S, Copenhagen DR, Lang RA
• A direct and melanopsin-dependent fetal light response regulates mouse eye development.
  Nature 2013 Rao S, Chun C, Fan J, Kofron JM, Yang MB, Hegde RS, Ferrara N, Copenhagen DR, Lang RA
• Light evokes melanopsin-dependent vocalization and neural activation associated with aversive experience in neonatal mice.
  PloS one 2012 Delwig A, Logan AM, Copenhagen DR, Ahn AH
• Overexpression of neurotrophin-3 stimulates a second wave of dopaminergic amacrine cell genesis after birth in the mouse retina.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2011 Yoshida M, Feng L, Grimbert F, Rangarajan KV, Buggele W, Copenhagen DR, Cang J, Liu X
• Light as a regulator of vascular regression.
  Developmental Biology 2011 Sujata Rao, Christina Chun, Samer Hattar, Napoleone Ferrara, David Copenhagen, Richard Lang
• TrkB (tropomyosin-related kinase B) controls the assembly and maintenance of GABAergic synapses in the cerebellar cortex.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2011 Chen AI, Nguyen CN, Copenhagen DR, Badurek S, Minichiello L, Ranscht B, Reichardt LF
• Melanopsin-dependent light avoidance in neonatal mice.
  Proceedings of the National Academy of Sciences of the United States of America 2010 Johnson J, Wu V, Donovan M, Majumdar S, Rentería RC, Porco T, Van Gelder RN, Copenhagen DR
• Regulation of neonatal development of retinal ganglion cell dendrites by neurotrophin-3 overexpression.
  The Journal of comparative neurology 2009 Liu X, Robinson ML, Schreiber AM, Wu V, Lavail MM, Cang J, Copenhagen DR
• Synaptic and extrasynaptic factors governing glutamatergic retinal waves.
  Neuron 2009 Blankenship AG, Ford KJ, Johnson J, Seal RP, Edwards RH, Copenhagen DR, Feller MB
• Retinal TrkB receptors regulate neural development in the inner, but not outer, retina.
  Molecular and cellular neurosciences 2008 Grishanin RN, Yang H, Liu X, Donohue-Rolfe K, Nune GC, Zang K, Xu B, Duncan JL, Lavail MM, Copenhagen DR, Reichardt LF
• 1.24 Plasticity of Retinal Circuitry.
  The Senses: A Comprehensive Reference 2008 N. Tian, D. Copenhagen
• Brain-derived neurotrophic factor and TrkB modulate visual experience-dependent refinement of neuronal pathways in retina.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2007 Liu X, Grishanin RN, Tolwani RJ, Rentería RC, Xu B, Reichardt LF, Copenhagen DR
• Vesicular glutamate transporter 1 is required for photoreceptor synaptic signaling but not for intrinsic visual functions.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2007 Johnson J, Fremeau RT, Duncan JL, Rentería RC, Yang H, Hua Z, Liu X, LaVail MM, Edwards RH, Copenhagen DR
• Intrinsic ON responses of the retinal OFF pathway are suppressed by the ON pathway.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2006 Rentería RC, Tian N, Cang J, Nakanishi S, Stryker MP, Copenhagen DR
• Scotopic visual signaling in the mouse retina is modulated by high-affinity plasma membrane calcium extrusion.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2006 Duncan JL, Yang H, Doan T, Silverstein RS, Murphy GJ, Nune G, Liu X, Copenhagen D, Tempel BL, Rieke F, Krizaj D
• Neuronal pentraxins mediate synaptic refinement in the developing visual system.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2006 Bjartmar L, Huberman AD, Ullian EM, Rentería RC, Liu X, Xu W, Prezioso J, Susman MW, Stellwagen D, Stokes CC, Cho R, Worley P, Malenka RC, Ball S, Peachey NS, Copenhagen D, Chapman B, Nakamoto M, …
• Development of precise maps in visual cortex requires patterned spontaneous activity in the retina.
  Neuron 2005 Cang J, Rentería RC, Kaneko M, Liu X, Copenhagen DR, Stryker MP
• Ephrin-As and neural activity are required for eye-specific patterning during retinogeniculate mapping.
  Nature neuroscience 2005 Pfeiffenberger C, Cutforth T, Woods G, Yamada J, Rentería RC, Copenhagen DR, Flanagan JG, Feldheim DA
• Ontogeny of plasma membrane Ca2+ ATPase isoforms in the neural retina of the postnatal rat.
  Visual neuroscience 2005 Rentería RC, Strehler EE, Copenhagen DR, Krizaj D
• Vesicular glutamate transporter 3 expression identifies glutamatergic amacrine cells in the rodent retina.
  The Journal of comparative neurology 2004 Johnson J, Sherry DM, Liu X, Fremeau RT, Seal RP, Edwards RH, Copenhagen DR
• Expression of calcium transporters in the retina of the tiger salamander (Ambystoma tigrinum).
  The Journal of comparative neurology 2004 Krizaj D, Liu X, Copenhagen DR
• N-type and L-type calcium channels mediate glycinergic synaptic inputs to retinal ganglion cells of tiger salamanders.
  Visual neuroscience 2004 Bieda MC, Copenhagen DR
• Vesicular glutamate transporters 1 and 2 target to functionally distinct synaptic release sites.
  Science (New York, N.Y.) 2004 Fremeau RT, Kam K, Qureshi T, Johnson J, Copenhagen DR, Storm-Mathisen J, Chaudhry FA, Nicoll RA, Edwards RH
• Need rods? Get glycine receptors and taurine.
  Neuron 2004 Rentería RC, Johnson J, Copenhagen DR
• Visual stimulation is required for refinement of ON and OFF pathways in postnatal retina.
  Neuron 2003 Tian N, Copenhagen DR
• The H+-coupled electrogenic lysosomal amino acid transporter LYAAT1 localizes to the axon and plasma membrane of hippocampal neurons.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2003 Wreden CC, Johnson J, Tran C, Seal RP, Copenhagen DR, Reimer RJ, Edwards RH
• Ryanodine stores and calcium regulation in the inner segments of salamander rods and cones.
  The Journal of physiology 2003 Krizaj D, Lai FA, Copenhagen DR
• Vesicular neurotransmitter transporter expression in developing postnatal rodent retina: GABA and glycine precede glutamate.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2003 Johnson J, Tian N, Caywood MS, Reimer RJ, Edwards RH, Copenhagen DR
• Development and Plasticity in the Postnatal Mouse Retina.
  The Neural Basis of Early Vision 2003 David R. Copenhagen, Ning Tian
• SN1 Catalyzes Transport of Glutamine Across Müller Cells in Retina.
  The Neural Basis of Early Vision 2003 Kyoh-Ichi Takahashi, David Krizaj, David R. Copenhagen
• The identification of vesicular glutamate transporter 3 suggests novel modes of signaling by glutamate.
  Proceedings of the National Academy of Sciences of the United States of America 2002 Fremeau RT, Burman J, Qureshi T, Tran CH, Proctor J, Johnson J, Zhang H, Sulzer D, Copenhagen DR, Storm-Mathisen J, Reimer RJ, Chaudhry FA, Edwards RH
• Cell-specific expression of plasma membrane calcium ATPase isoforms in retinal neurons.
  The Journal of comparative neurology 2002 Krizaj D, Demarco SJ, Johnson J, Strehler EE, Copenhagen DR
• Calcium regulation in photoreceptors.
  Frontiers in bioscience : a journal and virtual library 2002 Krizaj D, Copenhagen DR
• Coupled and uncoupled proton movement by amino acid transport system N.
  The EMBO journal 2001 Chaudhry FA, Krizaj D, Larsson P, Reimer RJ, Wreden C, Storm-Mathisen J, Copenhagen D, Kavanaugh M, Edwards RH
• Visual deprivation alters development of synaptic function in inner retina after eye opening.
  Neuron 2001 Tian N, Copenhagen DR
• Is the retina going digital?
  Neuron 2001 Copenhagen D
• BDNF-Induced potentiation of spontaneous twitching in innervated myocytes requires calcium release from intracellular stores.
  Journal of neurophysiology 2000 Kleiman RJ, Tian N, Krizaj D, Hwang TN, Copenhagen DR, Reichardt LF
• Combining physiology and genetics in the zebrafish retina.
  The Journal of physiology 2000 Baier H, Copenhagen D
• Inhibition is not required for the production of transient spiking responses from retinal ganglion cells.
  Visual neuroscience 2000 Bieda MC, Copenhagen DR
• Localization and developmental expression patterns of the neuronal K-Cl cotransporter (KCC2) in the rat retina.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 2000 Vu TQ, Payne JA, Copenhagen DR
• Molecular analysis of system N suggests novel physiological roles in nitrogen metabolism and synaptic transmission.
  Cell 1999 Chaudhry FA, Reimer RJ, Krizaj D, Barber D, Storm-Mathisen J, Copenhagen DR, Edwards RH
• The role of the synthetic enzyme GAD65 in the control of neuronal gamma-aminobutyric acid release.
  Proceedings of the National Academy of Sciences of the United States of America 1999 Tian N, Petersen C, Kash S, Baekkeskov S, Copenhagen D, Nicoll R
• Automatic detection, characterization, and discrimination of kinetically distinct spontaneous synaptic events.
  Journal of neuroscience methods 1999 Hwang TN, Copenhagen DR
• Caffeine-sensitive calcium stores regulate synaptic transmission from retinal rod photoreceptors.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 1999 Krizaj D, Bao JX, Schmitz Y, Witkovsky P, Copenhagen DR
• Sodium action potentials are not required for light-evoked release of GABA or glycine from retinal amacrine cells.
  Journal of neurophysiology 1999 Bieda MC, Copenhagen DR
• Analysis of excitatory and inhibitory spontaneous synaptic activity in mouse retinal ganglion cells.
  Journal of neurophysiology 1998 Tian N, Hwang TN, Copenhagen DR
• Compartmentalization of calcium extrusion mechanisms in the outer and inner segments of photoreceptors.
  Neuron 1998 Krizaj D, Copenhagen DR
• Metabotropic glutamate receptor-mediated suppression of an inward rectifier current is linked via a cGMP cascade.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 1997 Dixon DB, Copenhagen DR
• Quinine, intracellular pH and modulation of hemi-gap junctions in catfish horizontal cells.
  Vision research 1996 Dixon DB, Takahashi K, Bieda M, Copenhagen DR
• Retinal development: on the crest of an exciting wave.
  Current biology : CB 1996 Copenhagen DR
• Passive electrical cable properties and synaptic excitation of tiger salamander retinal ganglion cells.
  Visual neuroscience 1996 Taylor WR, Mittman S, Copenhagen DR
• Modulation of neuronal function by intracellular pH.
  Neuroscience research 1996 Takahashi KI, Copenhagen DR
• The relationship between light-evoked synaptic excitation and spiking behaviour of salamander retinal ganglion cells.
  The Journal of physiology 1995 Diamond JS, Copenhagen DR
• Characterization of spontaneous excitatory synaptic currents in salamander retinal ganglion cells.
  The Journal of physiology 1995 Taylor WR, Chen E, Copenhagen DR
• Intracellular alkalinization enhances inward rectifier K+ current in retinal horizontal cells of catfish.
  Zoological science 1995 Takahashi K, Copenhagen DR
• Actions of nipecotic acid and SKF89976A on GABA transporter in cone-driven horizontal cells dissociated from the catfish retina.
  The Japanese journal of physiology 1995 Takahashi K, Miyoshi S, Kaneko A, Copenhagen DR
• Nitric oxide synthase in Müller cells and neurons of salamander and fish retina.
  The Journal of neuroscience : the official journal of the Society for Neuroscience 1994 Liepe BA, Stone C, Koistinaho J, Copenhagen DR
• GABA transporter mediated-current in cone-driven horizontal cells of catfish retina.
  Neuroscience Research Supplements 1994 Kyoh-Ichi Takahashi, Shun-Ichiro Miyoshi, Akimichi Kaneko, David R. Copenhagen
• The contribution of NMDA and non-NMDA receptors to the light-evoked input-output characteristics of retinal ganglion cells.
  Neuron 1993 Diamond JS, Copenhagen DR
• L-glutamate suppresses HVA calcium current in catfish horizontal cells by raising intracellular proton concentration.
  Neuron 1993 Dixon DB, Takahashi K, Copenhagen DR
• Modulation of a sustained calcium current by intracellular pH in horizontal cells of fish retina.
  The Journal of general physiology 1993 Takahashi K, Dixon DB, Copenhagen DR
• Suppression of a calcium current by CNQX and kynurenate.
  Neuroscience letters 1992 Dixon DB, Copenhagen DR
• APB suppresses synaptic input to retinal horizontal cells in fish: a direct action on horizontal cells modulated by intracellular pH.
  Journal of neurophysiology 1992 Takahashi K, Copenhagen DR
• Two types of glutamate receptors differentially excite amacrine cells in the tiger salamander retina.
  The Journal of physiology 1992 Dixon DB, Copenhagen DR
• Synaptic transmission in the retina.
  Current opinion in neurobiology 1991 Copenhagen DR
• Application of a fluorometric method to measure glutamate release from single retinal photoreceptors.
  Journal of neuroscience methods 1991 Ayoub GS, Copenhagen DR
• Concomitant activation of two types of glutamate receptor mediates excitation of salamander retinal ganglion cells.
  The Journal of physiology 1990 Mittman S, Taylor WR, Copenhagen DR
• Signal transmission through the dark-adapted retina of the toad (Bufo marinus). Gain, convergence, and signal/noise.
  The Journal of general physiology 1990 Copenhagen DR, Hemilä S, Reuter T
• Weber and noise adaptation in the retina of the toad Bufo marinus.
  The Journal of general physiology 1990 Donner K, Copenhagen DR, Reuter T
• Intracellular cesium separates two glutamate conductances in retinal bipolar cells of goldfish.
  Vision research 1990 Nawy S, Copenhagen DR
• Release of endogenous excitatory amino acids from turtle photoreceptors.
  Nature 1989 Copenhagen DR, Jahr CE
• The glutamate analog 2-amino-4-phosphonobutyrate antagonizes synaptic transmission from cones to horizontal cells in the goldfish retina.
  Proceedings of the National Academy of Sciences of the United States of America 1989 Nawy S, Sie A, Copenhagen DR
• Release of endogenous glutamate from isolated cone photoreceptors of the lizard.
  Neuroscience research. Supplement : the official journal of the Japan Neuroscience Society 1989 Ayoub GS, Korenbrot JI, Copenhagen DR
• Synaptic transfer of rod signals to horizontal and bipolar cells in the retina of the toad (Bufo marinus).
  The Journal of physiology 1988 Belgum JH, Copenhagen DR
• Bubble pressure measurement of micropipet tip outer diameter.
  Journal of neuroscience methods 1987 Mittman S, Flaming DG, Copenhagen DR, Belgum JH
• Ganglion cell performance at absolute threshold in toad retina: effects of dark events in rods.
  The Journal of physiology 1987 Copenhagen DR, Donner K, Reuter T
• Spatial spread of adaptation within the cone network of turtle retina.
  The Journal of physiology 1987 Copenhagen DR, Green DG
• Development and characterization of a polymer gel with an immobilized enzyme to measure L-glutamate.
  Analytical biochemistry 1987 Korenbrot JI, Perry R, Copenhagen DR
• Multiple classes of glutamate receptor on depolarizing bipolar cells in retina.
  Nature 1987 Nawy S, Copenhagen DR
• Does the random distribution of discrete photoreceptor events limit the sensitivity of the retina?
  Neuroscience research. Supplement : the official journal of the Japan Neuroscience Society 1986 Reuter T, Donner K, Copenhagen DR
• Does the random distribution of discrete photoreceptor events limit the sensitivity of the retina?.
  Neuroscience Research Supplements 1986 Tom Reuter, Kristian Donner, David R. Copenhagen
• The absence of spread of adaptation between rod photoreceptors in turtle retina.
  The Journal of physiology 1985 Copenhagen DR, Green DG
• The focal nature of retinal illumination from the operating microscope.
  Archives of ophthalmology (Chicago, Ill. : 1960) 1985 Irvine AR, Copenhagen DR
• An analysis of transmission from cones to hyperpolarizing bipolar cells in the retina of the turtle.
  The Journal of physiology 1983 Ashmore JF, Copenhagen DR
• Kinetics of synaptic transmission from photoreceptors to horizontal and bipolar cells in turtle retina.
  Vision research 1983 Copenhagen DR, Ashmore JF, Schnapf JK
• Differences in the kinetics of rod and cone synaptic transmission.
  Nature 1982 Schnapf JL, Copenhagen DR
• Horizontal cells in turtle retina: structure, synaptic connections, and visual processing.
  Progress in clinical and biological research 1982 Leeper HF, Copenhagen DR
• Different postsynaptic events in two types of retinal bipolar cell.
  Nature 1980 Ashmore JF, Copenhagen DR
• Current-voltage relations in the rod photoreceptor network of the turtle retina.
  The Journal of physiology 1980 Copenhagen DR, Owen WG
• Mixed rod-cone responses in horizontal cells of snapping turtle retina.
  Vision Research 1979 D.R. Copenhagen, H.F. Leeper
• Mixed rod-cone responses in horizontal cells of snapping turtle retina.
  Vision research 1979 Leeper HF, Copenhagen DR
• Evidence for passive electrotonic interactions in red rods of toad retina.
  Nature 1978 Leeper HF, Normann RA, Copenhagen DR
• Functional characteristics of lateral interactions between rods in the retina of the snapping turtle.
  The Journal of physiology 1976 Copenhagen DR, Owen WG
• Coupling between rod photoreceptors in a vertebrate retina.
  Nature 1976 Copenhagen DR, Owen WG
• Time course of threshold elevation in on-off ganglion cells of Necturus retina: effects of lateral interactions.
  Vision research 1975 Copenhagen DR
• Control of retinal sensitivity. 3. Lateral interactions at the inner plexiform layer.
  The Journal of general physiology 1974 Werblin FS, Copenhagen DR
• Study of valveless, nonocclusive pump.
  JAAMI : journal of the Association for the Advancement of Medical Instrumentation 1969 Copenhagen D, Vogelhut P