Cell-intrinsic regulators of proliferation in vertebrate retinal progenitors

Ohnuma, 2001, Cell cycle and cell fate in the nervous system, Curr. Opin. Neurobiol., 11, 66, 10.1016/S0959-4388(00)00175-6 Pardee, 1974, A restriction point for control of normal animal cell proliferation, Proc. Natl. Acad. Sci. U.S.A., 71, 1286, 10.1073/pnas.71.4.1286 Pardee, 1989, G1 events and regulation of cell proliferation, Science, 246, 603, 10.1126/science.2683075 Planas-Silva, 1997, The restriction point and control of cell proliferation, Curr. Opin. Cell. Biol., 9, 768, 10.1016/S0955-0674(97)80076-2 Reh, 1998, Multipotential stem cells and progenitors in the vertebrate retina, J. Neurobiol., 36, 206, 10.1002/(SICI)1097-4695(199808)36:2<206::AID-NEU8>3.0.CO;2-5 Cameron, 1998, Regulation of neurogenesis by growth factors and neurotransmitters, J. Neurobiol., 36, 287, 10.1002/(SICI)1097-4695(199808)36:2<287::AID-NEU13>3.0.CO;2-B Levine, 2000, Soluble factors and the development of rod photoreceptors, Cell. Mol. Life Sci., 57, 224, 10.1007/PL00000686 Megason, 2002, A mitogen gradient of dorsal midline Wnts organizes growth in the CNS, Development, 129, 2087, 10.1242/dev.129.9.2087 Knoblich, 1994, Cyclin E controls S phase progression and its down-regulation during Drosophila embryogenesis is required for the arrest of cell proliferation, Cell, 77, 107, 10.1016/0092-8674(94)90239-9 Lane, 1996, Dacapo, a cyclin-dependent kinase inhibitor, stops cell proliferation during Drosophila development, Cell, 87, 1225, 10.1016/S0092-8674(00)81818-8 Brook, 1996, Requirements for dE2F function in proliferating cells and in post-mitotic differentiating cells, EMBO J., 15, 3676, 10.1002/j.1460-2075.1996.tb00737.x de Nooij, 1996, A cyclin-dependent kinase inhibitor, Dacapo, is necessary for timely exit from the cell cycle during Drosophila embryogenesis, Cell, 87, 1237, 10.1016/S0092-8674(00)81819-X Du, 1996, RBF, a novel RB-related gene that regulates E2F activity and interacts with cyclin E in Drosophila, Genes Dev., 10, 1206, 10.1101/gad.10.10.1206 Du, 1996, Ectopic expression of dE2F and dDP induces cell proliferation and death in the Drosophila eye, EMBO J., 15, 3684, 10.1002/j.1460-2075.1996.tb00738.x Finley, 1996, Isolation of Drosophila cyclin D, a protein expressed in the morphogenetic furrow before entry into S phase, Proc. Natl. Acad. Sci. U.S.A., 93, 3011, 10.1073/pnas.93.7.3011 Secombe, 1998, Analysis of a Drosophila cyclin E hypomorphic mutation suggests a novel role for cyclin E in cell proliferation control during eye imaginal disc development, Genetics, 149, 1867, 10.1093/genetics/149.4.1867 Du, 1999, The role of RBF in the introduction of G1 regulation during Drosophila embryogenesis, EMBO J., 18, 916, 10.1093/emboj/18.4.916 Du, 2000, Suppression of the rbf null mutants by a de2f1 allele that lacks transactivation domain, Development, 127, 367, 10.1242/dev.127.2.367 Classon, 2002, The retinoblastoma tumour suppressor in development and cancer, Nat. Rev. Cancer, 2, 910, 10.1038/nrc950 Jacks, 1992, Effects of an Rb mutation in the mouse, Nature, 359, 295, 10.1038/359295a0 Lee, 1992, Mice deficient for Rb are nonviable and show defects in neurogenesis and haematopoiesis, Nature, 359, 288, 10.1038/359288a0 Lee, 1994, Dual roles of the retinoblastoma protein in cell cycle regulation and neuron differentiation, Genes Dev., 8, 2008, 10.1101/gad.8.17.2008 Maandag, 1994, Developmental rescue of an embryonic-lethal mutation in the retinoblastoma gene in chimeric mice, EMBO J., 13, 4260, 10.1002/j.1460-2075.1994.tb06746.x Robanus-Maandag, 1998, p107 is a suppressor of retinoblastoma development in pRb-deficient mice, Genes Dev., 12, 1599, 10.1101/gad.12.11.1599 Griep, 1998, Multiple genetic loci modify risk for retinoblastoma in transgenic mice, Invest. Ophthalmol. Vis. Sci., 39, 2723 Sage, 2003, Acute mutation of retinoblastoma gene function is sufficient for cell cycle re-entry, Nature, 424, 223, 10.1038/nature01764 Trimarchi, 2002, Sibling rivalry in the e2f family, Nat. Rev. Mol. Cell. Biol., 3, 11, 10.1038/nrm714 Wu, 2001, The E2F1-3 transcription factors are essential for cellular proliferation, Nature, 414, 457, 10.1038/35106593 Cooper-Kuhn, 2002, Impaired adult neurogenesis in mice lacking the transcription factor E2F1, Mol. Cell. Neurosci., 21, 312, 10.1006/mcne.2002.1176 Dagnino, 1997, Expression patterns of the E2F family of transcription factors during mouse nervous system development, Mech. Dev., 66, 13, 10.1016/S0925-4773(97)00083-X Espanel, 1998, Regulation of E2F-1 gene expression in avian cells, Oncogene, 17, 585, 10.1038/sj.onc.1201962 Kastner, 1998, Transient accumulation of retinoblastoma/E2F-1 protein complexes correlates with the onset of neuronal differentiation in the developing quail neural retina, Cell. Growth Differ., 9, 857 Lin, 2001, The proliferative and apoptotic activities of E2F1 in the mouse retina, Oncogene, 20, 7073, 10.1038/sj.onc.1204932 Fantl, 1995, Mice lacking cyclin D1 are small and show defects in eye and mammary gland development, Genes Dev., 9, 2364, 10.1101/gad.9.19.2364 Ma, 1998, A unique pattern of photoreceptor degeneration in cyclin D1 mutant mice, Proc. Natl. Acad. Sci. U.S.A., 95, 9938, 10.1073/pnas.95.17.9938 Sicinski, 1995, Cyclin D1 provides a link between development and oncogenesis in the retina and breast, Cell, 82, 621, 10.1016/0092-8674(95)90034-9 Geng, 1999, Rescue of cyclin D1 deficiency by knockin cyclin E, Cell, 97, 767, 10.1016/S0092-8674(00)80788-6 Geng, 2001, Deletion of p27(Kip1) gene restores normal development in cyclin D1-deficient mice, Proc. Natl. Acad. Sci. U.S.A., 98, 194, 10.1073/pnas.011522998 Tong, 2001, Genetic evidence for the interactions of cyclin D1 and p27(Kip1) in mice, Mol. Cell. Biol., 21, 1319, 10.1128/MCB.21.4.1319-1328.2001 Green, 2003, Genetic rescue of cell number in a mouse model of microphthalmia: interactions between Chx10 and G1-phase cell cycle regulators, Development, 130, 539, 10.1242/dev.00275 Ciemerych, 2002, Development of mice expressing a single D-type cyclin, Genes Dev., 16, 3277, 10.1101/gad.1023602 Mitsuhashi, 2001, Overexpression of p27Kip1 lengthens the G1 phase in a mouse model that targets inducible gene expression to central nervous system progenitor cells, Proc. Natl. Acad. Sci. U.S.A., 98, 6435, 10.1073/pnas.111051398 Fero, 1996, A syndrome of multiorgan hyperplasia with features of gigantism, tumorigenesis, and female sterility in p27(Kip1)-deficient mice, Cell, 85, 733, 10.1016/S0092-8674(00)81239-8 Kiyokawa, 1996, Enhanced growth of mice lacking the cyclin-dependent kinase inhibitor function of p27(Kip1), Cell, 85, 721, 10.1016/S0092-8674(00)81238-6 Nakayama, 1996, Mice lacking p27(Kip1) display increased body size, multiple organ hyperplasia, retinal dysplasia, and pituitary tumors, Cell, 85, 707, 10.1016/S0092-8674(00)81237-4 Durand, 1998, p27Kip1 alters the response of cells to mitogen and is part of a cell-intrinsic timer that arrests the cell cycle and initiates differentiation, Curr. Biol., 8, 431, 10.1016/S0960-9822(98)70177-0 Ohnuma, 1999, p27Xic1, a Cdk inhibitor, promotes the determination of glial cells in Xenopus retina, Cell, 99, 499, 10.1016/S0092-8674(00)81538-X Levine, 2000, p27(Kip1) regulates cell cycle withdrawal of late multipotent progenitor cells in the mammalian retina, Dev. Biol., 219, 299, 10.1006/dbio.2000.9622 Dyer, 2001, p27Kip1 and p57Kip2 regulate proliferation in distinct retinal progenitor cell populations, J. Neurosci., 21, 4259, 10.1523/JNEUROSCI.21-12-04259.2001 Dyer, 2000, Control of Muller glial cell proliferation and activation following retinal injury, Nat. Neurosci., 3, 873, 10.1038/78774 Dyer, 2000, p57(Kip2) regulates progenitor cell proliferation and amacrine interneuron development in the mouse retina, Development, 127, 3593, 10.1242/dev.127.16.3593 Zindy, 1997, Expression of INK4 inhibitors of cyclin D-dependent kinases during mouse brain development, Cell Growth Differ., 8, 1139 Cunningham, 2002, The cyclin-dependent kinase inhibitors p19(Ink4d) and p27(Kip1) are coexpressed in select retinal cells and act cooperatively to control cell cycle exit, Mol. Cell. Neurosci., 19, 359, 10.1006/mcne.2001.1090 Defoe W, Levine EM. Expression of the cyclin-dependent kinase inhibitor p27Kip1 by developing retinal pigment epithelium. Gene Expression Patterns 2003;3:615–9. Acampora, 1999, Otx genes in corticogenesis and brain development, Cereb. Cortex, 9, 533, 10.1093/cercor/9.6.533 Estivill-Torrus, 2002, Pax6 is required to regulate the cell cycle and the rate of progression from symmetrical to asymmetrical division in mammalian cortical progenitors, Development, 129, 455, 10.1242/dev.129.2.455 Galli, 2002, Emx2 regulates the proliferation of stem cells of the adult mammalian central nervous system, Development, 129, 1633, 10.1242/dev.129.7.1633 Hanashima, 2002, Brain factor-1 controls the proliferation and differentiation of neocortical progenitor cells through independent mechanisms, J. Neurosci., 22, 6526, 10.1523/JNEUROSCI.22-15-06526.2002 Tole, 2000, Emx2 is required for growth of the hippocampus but not for hippocampal field specification, J. Neurosci., 20, 2618, 10.1523/JNEUROSCI.20-07-02618.2000 Warren, 1999, The transcription factor, Pax6 is required for cell proliferation and differentiation in the developing cerebral cortex, Cereb. Cortex, 9, 627, 10.1093/cercor/9.6.627 Xuan, 1995, Winged helix transcription factor BF-1 is essential for the development of the cerebral hemispheres, Neuron, 14, 1141, 10.1016/0896-6273(95)90262-7 Mathers, 1997, The Rx homeobox gene is essential for vertebrate eye development, Nature, 387, 603, 10.1038/42475 Casarosa, 1997, Xrx1, a novel Xenopus homeobox gene expressed during eye and pineal gland development, Mech. Dev., 61, 187, 10.1016/S0925-4773(96)00640-5 Furukawa, 1997, rax, a novel paired-type homeobox gene, shows expression in the anterior neural fold and developing retina, Proc. Natl. Acad. Sci. U.S.A., 94, 3088, 10.1073/pnas.94.7.3088 Chuang, 1999, Expression of three Rx homeobox genes in embryonic and adult zebrafish, Mech. Dev., 84, 195, 10.1016/S0925-4773(99)00077-5 Deschet, 1999, Expression of the medaka (Oryzias latipes) Ol-Rx3 paired-like gene in two diencephalic derivatives, the eye and the hypothalamus, Mech. Dev., 83, 179, 10.1016/S0925-4773(99)00037-4 Ohuchi, 1999, Identification of chick rax/rx genes with overlapping patterns of expression during early eye and brain development, Mech. Dev., 85, 193, 10.1016/S0925-4773(99)00094-5 Chen, 2002, The chicken RaxL gene plays a role in the initiation of photoreceptor differentiation, Development, 129, 5363, 10.1242/dev.00114 Andreazzoli, 1999, Role of Xrx1 in Xenopus eye and anterior brain development, Development, 126, 2451, 10.1242/dev.126.11.2451 Chuang, 2001, Zebrafish genes rx1 and rx2 help define the region of forebrain that gives rise to retina, Dev. Biol., 231, 13, 10.1006/dbio.2000.0125 Zhang, 2000, Function of Rx, but not Pax6, is essential for the formation of retinal progenitor cells in mice, Genesis, 28, 135, 10.1002/1526-968X(200011/12)28:3/4<135::AID-GENE70>3.0.CO;2-P Winkler, 2000, The conditional medaka mutation eyeless uncouples patterning and morphogenesis of the eye, Development, 127, 1911, 10.1242/dev.127.9.1911 Loosli, 2001, Medaka eyeless is the key factor linking retinal determination and eye growth, Development, 128, 4035, 10.1242/dev.128.20.4035 Casarosa, 2003, Xrx1 controls proliferation and multipotency of retinal progenitors, Mol. Cell. Neurosci., 22, 25, 10.1016/S1044-7431(02)00025-8 Ashery-Padan, 2001, Pax6 lights-up the way for eye development, Curr. Opin. Cell. Biol., 13, 706, 10.1016/S0955-0674(00)00274-X Marquardt, 2001, Pax6 is required for the multipotent state of retinal progenitor cells, Cell, 105, 43, 10.1016/S0092-8674(01)00295-1 Bernier, 2001, Isolation and characterization of a downstream target of Pax6 in the mammalian retinal primordium, Development, 128, 3987, 10.1242/dev.128.20.3987 Gotz, 1998, Pax6 controls radial glia differentiation in the cerebral cortex, Neuron, 21, 1031, 10.1016/S0896-6273(00)80621-2 Warren, 1997, Roles of Pax-6 in murine diencephalic development, Development, 124, 1573, 10.1242/dev.124.8.1573 Kawakami, 2000, Six family genes—structure and function as transcription factors and their roles in development, Bioessays, 22, 616, 10.1002/1521-1878(200007)22:7<616::AID-BIES4>3.0.CO;2-R Zhou, 2000, Cloning and expression of xSix3, the Xenopus homologue of murine Six3, Mech. Dev., 91, 327, 10.1016/S0925-4773(99)00270-1 Ghanbari, 2001, Molecular cloning and embryonic expression of Xenopus six homeobox genes, Mech. Dev., 101, 271, 10.1016/S0925-4773(00)00572-4 Loosli, 1999, Six3 overexpression initiates the formation of ectopic retina, Genes Dev., 13, 649, 10.1101/gad.13.6.649 Kobayashi, 1998, Overexpression of the forebrain-specific homeobox gene six3 induces rostral forebrain enlargement in zebrafish, Development, 125, 2973, 10.1242/dev.125.15.2973 Bernier, 2000, Expanded retina territory by midbrain transformation upon overexpression of Six6 (Optx2) in Xenopus embryos, Mech. Dev., 93, 59, 10.1016/S0925-4773(00)00271-9 Lagutin, 2001, Six3 promotes the formation of ectopic optic vesicle-like structures in mouse embryos, Dev. Dyn., 221, 342, 10.1002/dvdy.1148 Carl, 2002, Six3 inactivation reveals its essential role for the formation and patterning of the vertebrate eye, Development, 129, 4057, 10.1242/dev.129.17.4057 Wallis, 1999, Mutations in the homeodomain of the human SIX3 gene cause holoprosencephaly, Nat. Genet., 22, 196, 10.1038/9718 Toy, 1998, The optx2 homeobox gene is expressed in early precursors of the eye and activates retina-specific genes, Proc. Natl. Acad. Sci. U.S.A., 95, 10643, 10.1073/pnas.95.18.10643 Toy, 1999, Expression of the optx2 homeobox gene during mouse development, Mech. Dev., 83, 183, 10.1016/S0925-4773(99)00049-0 Lopez-Rios, 1999, Six9 (Optx2), a new member of the six gene family of transcription factors, is expressed at early stages of vertebrate ocular and pituitary development, Mech. Dev., 83, 155, 10.1016/S0925-4773(99)00017-9 Zuber, 1999, Giant eyes in Xenopus laevis by overexpression of XOptx2, Cell, 98, 341, 10.1016/S0092-8674(00)81963-7 Jean, 1999, Six6 (Optx2) is a novel murine Six3-related homeobox gene that demarcates the presumptive pituitary/hypothalamic axis and the ventral optic stalk, Mech. Dev., 84, 31, 10.1016/S0925-4773(99)00068-4 Kobayashi, 2001, The homeobox protein Six3 interacts with the Groucho corepressor and acts as a transcriptional repressor in eye and forebrain formation, Dev. Biol., 232, 315, 10.1006/dbio.2001.0185 Zhu, 2002, Six3-mediated auto repression and eye development requires its interaction with members of the Groucho-related family of co-repressors, Development, 129, 2835, 10.1242/dev.129.12.2835 Lopez-Rios, 2003, Six3 and Six6 activity is modulated by members of the groucho family, Development, 130, 185, 10.1242/dev.00185 Li, 2002, Tissue-specific regulation of retinal and pituitary precursor cell proliferation, Science, 297, 1180, 10.1126/science.1073263 Galliot, 1999, Evolution of homeobox genes: Q50 paired-like genes founded the paired class, Dev. Genes E, 209, 186, 10.1007/s004270050243 Liu, 1994, Developmental expression of a novel murine homeobox gene (Chx10): evidence for roles in determination of the neuroretina and inner nuclear layer, Neuron, 13, 377, 10.1016/0896-6273(94)90354-9 Levine, 1997, Vsx-1 and Vsx-2: two Chx10-like homeobox genes expressed in overlapping domains in the adult goldfish retina, J. Comp. Neurol., 387, 439, 10.1002/(SICI)1096-9861(19971027)387:3<439::AID-CNE9>3.0.CO;2-1 Passini, 1997, Vsx-1 and Vsx-2: differential expression of two paired-like homeobox genes during zebrafish and goldfish retinogenesis, J. Comp. Neurol., 388, 495, 10.1002/(SICI)1096-9861(19971124)388:3<495::AID-CNE11>3.0.CO;2-L Passini, 1998, Vsx-2, a gene encoding a paired-type homeodomain, is expressed in the retina, hindbrain, and spinal cord during goldfish embryogenesis, Brain Res. Dev. Brain. Res., 109, 129, 10.1016/S0165-3806(98)00069-8 Barabino, 1997, Inactivation of the zebrafish homologue of Chx10 by antisense oligonucleotides causes eye malformations similar to the ocular retardation phenotype, Mech. Dev., 63, 133, 10.1016/S0925-4773(97)00036-1 Belecky-Adams, 1997, Pax-6, Prox 1, and Chx10 homeobox gene expression correlates with phenotypic fate of retinal precursor cells, Invest. Ophthalmol. Vis. Sci., 38, 1293 Chen, 2000, Expression of Chx10 and Chx10-1 in the developing chicken retina, Mech. Dev., 90, 293, 10.1016/S0925-4773(99)00251-8 Fuhrmann, 2000, Extraocular mesenchyme patterns the optic vesicle during early eye development in the embryonic chick, Development, 127, 4599, 10.1242/dev.127.21.4599 Theiler, 1976, A new allele of ocular retardation: early development and morphogenetic cell death, Anat. Embryol. (Berl.), 150, 85, 10.1007/BF00346288 Truslove, 1962, A gene causing ocular retardation in the mouse, J. Embryol. Exp. Morph., 10, 652 Burmeister, 1996, Ocular retardation mouse caused by Chx10 homeobox null allele: impaired retinal progenitor proliferation and bipolar cell differentiation, Nat. Genet., 12, 376, 10.1038/ng0496-376 Ferda Percin, 2000, Human microphthalmia associated with mutations in the retinal homeobox gene CHX10, Nat. Genet., 25, 397, 10.1038/78071 Robb, 1978, Ocular retardation (or) in the mouse, Invest. Ophthalmol. Vis. Sci., 17, 468 Silver, 1979, Studies on the development of the eye cup and optic nerve in normal mice and in mutants with congenital optic nerve aplasia, Dev. Biol., 68, 175, 10.1016/0012-1606(79)90252-5 Konyukhov, 1971, Genetic control over the duration of G 1 phase, Experientia, 27, 970, 10.1007/BF02135780 Bone-Larson, 2000, Partial rescue of the ocular retardation phenotype by genetic modifiers, J. Neurobiol., 42, 232, 10.1002/(SICI)1097-4695(20000205)42:2<232::AID-NEU7>3.0.CO;2-4 Tomita, 1996, Mammalian hairy and enhancer of split homolog 1 regulates differentiation of retinal neurons and is essential for eye morphogenesis, Neuron, 16, 723, 10.1016/S0896-6273(00)80093-8 Jan, 2001, Asymmetric cell division in the Drosophila nervous system, Nat. Rev. Neurosci., 2, 772, 10.1038/35097516 Li, 2000, Pan-neural prospero terminates cell proliferation during Drosophila neurogenesis, Genes Dev., 14, 147, 10.1101/gad.14.2.147 Oliver, 1993, Prox 1 a prospero-related homeobox gene expressed during mouse development, Mech. Dev., 44, 3, 10.1016/0925-4773(93)90012-M Tomarev, 1996, Chicken homeobox gene Prox 1 related to Drosophila prospero is expressed in the developing lens and retina, Dev. Dyn., 206, 354, 10.1002/(SICI)1097-0177(199608)206:4<354::AID-AJA2>3.0.CO;2-H Glasgow, 1998, Restricted expression of the homeobox gene prox 1 in developing zebrafish, Mech. Dev., 76, 175, 10.1016/S0925-4773(98)00121-X Dyer, 2003, Prox1 function controls progenitor cell proliferation and horizontal cell genesis in the mammalian retina, Nat. Genet., 34, 53, 10.1038/ng1144 Wawersik, 2000, Vertebrate eye development as modeled in Drosophila, Hum. Mol. Genet., 9, 917, 10.1093/hmg/9.6.917 Hanson, 2001, Mammalian homologues of the Drosophila eye specification genes, Semin. Cell. Dev. Biol., 12, 475, 10.1006/scdb.2001.0271 Alexiades, 1996, Quantitative analysis of proliferation and cell cycle length during development of the rat retina, Dev. Dyn., 205, 293, 10.1002/(SICI)1097-0177(199603)205:3<293::AID-AJA9>3.0.CO;2-D