Physiological Reviews, Vol 76, 651-685, Copyright © 1996 by American Physiological Society

JOURNAL ARTICLE

Regulation of protein transport to the nucleus: central role of phosphorylation

D. A. Jans and S. Hubner
Nuclear Signalling Laboratory, Division for Biochemistry and Molecular Biology, John Curtin School of Medical Research, Australian National University, Canberra, Australia.

Nuclear protein transport is integral to eukaryotic cell processes such as differentiation, transformation, and the control of gene expression. Although the targeting role of nuclear localization signals (NLSs) has been known for some time, more recent results indicate that NLS-dependent nuclear protein import is precisely regulated. Phosphorylation appears to be the main mechanism controlling the nuclear transport of a number of proteins, including transcription factors such as NFkappaB, c-rel, dorsal, and SWI5 from yeast. Cytoplasmic retention factors, intra- and intermolecular NLS masking, and NLS masking by phosphorylation are some of the mechanisms by which phosphorylation specifically regulates nuclear transport. Even nuclear localization of the archetypal NLS-containing simian virus 40 large tumor antigen (T-ag) is regulated, namely by the "CcN motif," which comprises the T-ag NLS ("N") determining ultimate subcellular destination, a casein kinase II site ("C") 13 amino acids NH2-terminal to the NLS modulating the rate of nuclear import, and a cyclin-dependent kinase site ("c") adjacent to the NLS regulating the maximal level of nuclear accumulation. The CcN motif appears to be a special form of phosphorylation-regulated NLS (prNLS), where phosphorylation at site(s) close to the NLS specifically regulates NLS function. The regulation of nuclear transport through phosphorylation and prNLSs appears to be common in eukaryotic cells from yeast and plants to higher mammals.

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				K. Welch, J. Franke, M. Kohler, and I. G. Macara RanBP3 Contains an Unusual Nuclear Localization Signal That Is Imported Preferentially by Importin-alpha 3 Mol. Cell. Biol., December 1, 1999; 19(12): 8400 - 8411. [Abstract] [Full Text] [PDF]

				M. L. Kingsley-Kallesen, D. Kelly, and A. Rizzino Transcriptional Regulation of the Transforming Growth Factor-beta 2 Promoter by cAMP-responsive Element-binding Protein (CREB) and Activating Transcription Factor-1 (ATF-1) Is Modulated by Protein Kinases and the Coactivators p300 and CREB-binding Protein J. Biol. Chem., November 26, 1999; 274(48): 34020 - 34028. [Abstract] [Full Text] [PDF]

				T. G. Canty Jr, E. M. Boyle Jr, A. Farr, E. N. Morgan, E. D. Verrier, and T. H. Pohlman Oxidative Stress Induces NF-{kappa}B Nuclear Translocation Without Degradation of I{kappa}B{alpha} Circulation, November 9, 1999; 100 (2009): II-361 - II-364. [Abstract] [Full Text] [PDF]

				A. M. Healy, M. Peters-Golden, J. P. Yao, and T. G. Brock Identification of a Bipartite Nuclear Localization Sequence Necessary for Nuclear Import of 5-Lipoxygenase J. Biol. Chem., October 15, 1999; 274(42): 29812 - 29818. [Abstract] [Full Text] [PDF]

				F. Wellmer, S. Kircher, A. Rugner, H. Frohnmeyer, E. Schafer, and K. Harter Phosphorylation of the Parsley bZIP Transcription Factor CPRF2 Is Regulated by Light J. Biol. Chem., October 8, 1999; 274(41): 29476 - 29482. [Abstract] [Full Text] [PDF]

				S. Hubner, H. M. S. Smith, W. Hu, C. K. Chan, H.-P. Rihs, B. M. Paschal, N. V. Raikhel, and D. A. Jans Plant Importin alpha Binds Nuclear Localization Sequences with High Affinity and Can Mediate Nuclear Import Independent of Importin beta J. Biol. Chem., August 6, 1999; 274(32): 22610 - 22617. [Abstract] [Full Text] [PDF]

				I. J. Gonzalez-Robayna, T. N. Alliston, P. Buse, G. L. Firestone, and J. S. Richards Functional and Subcellular Changes in the A-Kinase-Signaling Pathway: Relation to Aromatase and Sgk Expression during the Transition of Granulosa Cells to Luteal Cells Mol. Endocrinol., August 1, 1999; 13(8): 1318 - 1337. [Abstract] [Full Text]

				W. Ding, M. M. Witte, and R. E. Scott Transformation Blocks Differentiation-induced Inhibition of Serum Response Factor Interactions with Serum Response Elements Cancer Res., August 1, 1999; 59(15): 3795 - 3802. [Abstract] [Full Text] [PDF]

				K. L. Loveland, D. Herszfeld, B. Chu, E. Rames, E. Christy, L. J. Briggs, R. Shakri, D. M. de Kretser, and D. A. Jans Novel Low Molecular Weight Microtubule-associated Protein-2 Isoforms Contain a Functional Nuclear Localization Sequence J. Biol. Chem., July 2, 1999; 274(27): 19261 - 19268. [Abstract] [Full Text] [PDF]

				V. Lang, M. Semichon, F. Michel, C. Brossard, H. Gary-Gouy, and G. Bismuth Fyn Membrane Localization Is Necessary to Induce the Constitutive Tyrosine Phosphorylation of Sam68 in the Nucleus of T Lymphocytes J. Immunol., June 15, 1999; 162(12): 7224 - 7232. [Abstract] [Full Text] [PDF]

				J.-L. Liu, Y. Ye, Z. Qian, Y. Qian, D. J. Templeton, L. F. Lee, and H.-J. Kung Functional Interactions between Herpesvirus Oncoprotein MEQ and Cell Cycle Regulator CDK2 J. Virol., May 1, 1999; 73(5): 4208 - 4219. [Abstract] [Full Text]

				S. E. Perry, M. D. Lehti, and D. E. Fernandez The MADS-Domain Protein AGAMOUS-Like 15 Accumulates in Embryonic Tissues with Diverse Origins Plant Physiology, May 1, 1999; 120(1): 121 - 130. [Abstract] [Full Text]

				K. Okamoto, H. Tanaka, H. Ogawa, Y. Makino, H. Eguchi, S.-i. Hayashi, N. Yoshikawa, L. Poellinger, K. Umesono, and I. Makino Redox-dependent Regulation of Nuclear Import of the Glucocorticoid Receptor J. Biol. Chem., April 9, 1999; 274(15): 10363 - 10371. [Abstract] [Full Text] [PDF]

				M. Kann, B. Sodeik, A. Vlachou, W. H. Gerlich, and A. Helenius Phosphorylation-dependent Binding of Hepatitis B Virus Core Particles to the Nuclear Pore Complex J. Cell Biol., April 5, 1999; 145(1): 45 - 55. [Abstract] [Full Text] [PDF]

				A. Stathopoulos-Gerontides, J. J. Guo, and M. S. Cyert Yeast calcineurin regulates nuclear localization of the Crz1p transcription factor through dephosphorylation Genes & Dev., April 1, 1999; 13(7): 798 - 803. [Abstract] [Full Text]

				H. M.S. Smith and N. V. Raikhel Protein Targeting to the Nuclear Pore. What Can We Learn from Plants? Plant Physiology, April 1, 1999; 119(4): 1157 - 1164. [Full Text]

				N. Ohad, R. Yadegari, L. Margossian, M. Hannon, D. Michaeli, J. J. Harada, R. B. Goldberg, and R. L. Fischer Mutations in FIE, a WD Polycomb Group Gene, Allow Endosperm Development without Fertilization PLANT CELL, March 1, 1999; 11(3): 407 - 416. [Abstract] [Full Text] [PDF]

				J. Herrington, L. Rui, G. Luo, L.-y. Yu-Lee, and C. Carter-Su A Functional DNA Binding Domain Is Required for Growth Hormone-induced Nuclear Accumulation of Stat5B J. Biol. Chem., February 19, 1999; 274(8): 5138 - 5145. [Abstract] [Full Text] [PDF]

				I. Ben-Dor, O. Bern, and T. Tennenbaum Cell Cycle-dependent Nuclear Accumulation of the p94fer Tyrosine Kinase Is Regulated by Its NH2 Terminus and Is Affected by Kinase Domain Integrity and ATP Binding Cell Growth Differ., February 1, 1999; 10(2): 113 - 129. [Abstract] [Full Text]

				S. Kircher, F. Wellmer, P. Nick, A. Rugner, E. Schafer, and K. Harter Nuclear Import of the Parsley bZIP Transcription Factor CPRF2 Is Regulated by Phytochrome Photoreceptors J. Cell Biol., January 25, 1999; 144(2): 201 - 211. [Abstract] [Full Text] [PDF]