CC BY
135
Естественные науки
Хохлова Л.П., Невмержицкая Ю.Ю. Роль цитоскелета в сигнальных системах растений // Учен. зап. Казан. ун-та. Сер. Естеств. науки. - 2011. - Т. 153, кн. 2. - С. 147-179.
УДК 581.1
РОЛЬ ЦИТОСКЕЛЕТА В СИГНАЛЬНЫХ СИСТЕМАХ РАСТЕНИЙ
Л.П. Хохлова, Ю.Ю. Невмержицкая Аннотация
В обзоре проведен анализ данных литературы и результатов собственных исследований об участии цитоскелета в сигнальных системах клеток растений. Основные компоненты цитоскелета - тубулиновые микротрубочки и актиновые микрофиламенты -обладают ярко выраженной динамической нестабильностью, постоянно находясь в состоянии сборки/разборки, изменяют свое структурное и полимерное состояние, локализацию, ориентацию, состав изотипов, контакты между собой и клеточными структурами. Именно эта реорганизация цитоскелета, происходящая под влиянием внешних и внутренних факторов, и определяет его сигнальные функции. Цитоскелет-зависимый сигналинг включает целую иерархию последовательных биохимических и физиологических изменений, влияющих на форму клеток и органов, микроструктуру тканей, деление, полярность, дифференцировку, различные типы подвижности клеток, а также эндо-, экзоцитоз и везикулярный транспорт веществ. Рассматриваются новые данные о возможности участия цитоскелета в межклеточных коммукациях через «синапсопо-добные» контакты.
Ключевые слова: растения, цитоскелет, динамическая нестабильность, сигналы, трансдукция, клеточный ответ.
Summary
L.P. Khokhlova, Yu.Yu. Nevmerzhitskaya. The Role of Cytoskeleton in Plant Signaling Systems.
This overview presents the analysis of literature and the author's own research results on the role of cytoskeleton in plant cells signaling systems. The main components of cytoskeleton - tubulin microtubules and actin microfilaments - are characterized by a crear dynamic instability being constantly in an assembly/disassembly state and changing in their structural and polymeric state, localization, orientation, isotope composition, and contacts between each other and cell structures. It is the reorganization of cytoskeleton occurring in response to external and internal factors that determines its signaling functions. Cytoskeleton-dependent signaling initiates a chain of biochemical and physiological reactions, which influence cells' and organs' form, microstructure of tissues, division, polarity, differentiation, different types of cells mobility, as well as exo- and endocytosis and vesicular transport of substances. The work also considers new data about the possible participation of cytoskeleton in intercellular communication via "synapse-like" contacts.
Key words: plants, cytoskeleton, dynamic instability, signals, transduction, cell response.
Литература
1. Lloyd C., Chan J. Microtubules and the Shape of Plants to Come // Nature Rev. Mol. Cell Biol. - 2004. - V. 5, No 1. - P. 13-23.
2. Клячко Н.Л. Динамика актина в растительной клетке // Материалы Всерос. конф., посв. 75-летию кафедры физиологии и биотехнологии растений. - Казань: Изд-во Казан. ун-та, 2007. - С. 52-58.
3. Baskin T.I. The Cytoskeleton // Biochemistry and Molecular Biology of Plant / Eds. B.B. Buchanan, W. Cruissem, R.L. Jones. - Rockville: Courier Companies, 2000. -V. 51. - P. 202-258.
4. Barlow W.P., Baluska F. Cytoskeletal perspectives on root growth and morphogenesis // Annu. Rev. Plant Physiol. Plant Mol. Biol. - 2000. - V. 51. - P. 289-322.
5. Azimzadeh J., Traas J., Pastuglia M. Molecular aspects of microtubule dynamics in plant // Curr. Opin. Plant Biol. - 2001. - V. 4, No 6. - P. 513-519.
6. Wasteneys G.O. Microtubule organization in the green kingdom: chaos or self-order? // J. Cell Sci. - 2002. - V. 115, No 7. - P. 1345-1354.
7. Hepler P.K., Vidali L., Cheung A.Y. Polarized Cell Growth in Higer Plants // Annu. Rev. Cell Dev. Biol. - 2001. - V. 17. - P. 159-187.
8. Staiger C.J., Lloyd C. W. The plant cytoskeleton // Curr. Opin. Cell Biol. - 1991. - V. 3, No 1. - P. 33-42.
9. Derksen I., Wilms F.H.A., Pierson E.S. The plant cytoskeleton: its significance in plant development // Acto Bot. Neerl. - 1990. - V. 39, No 1. - Р. 1-18.
10. Тарчевский И.А. Сигнальные системы клеток растений. - М.: Наука, 2002. - 293 с.
11. Nick P. Signaling to the microtubular cytoskeleton in plants // Int. Rev. Cytol. - 1998. -V. 184. - P. 33-79.
12. Gardiner J., Collings D.A., Harper J.D.I., Marc J. The effect of the phospholipase D-anta-gonist 1-butanol on seedling development and microtubule organisation in Arabidopsis // Plant Cell Physiol. - 2003. - V. 44, No 7. - P. 687-696
13. Nick P. Signals, motors, morphogenesis - the cytoskeleton in plant development // Plant Biol. - 1999. - V. 1, No 2. - P.169-179.
14. Tan Z., Boss W.F. Association of phosphatidylinositol kinase, phosphatidylinositol monophosphate kinase, and diacylglycerol kinase with the cytosceleton and F-actin fractions of carrot (Daucus carota L.) cells grown in suspension culture. Response to cell wall-degrading ensimes // Plant Physiol. - 1992. - V. 100, No 4. - P. 2116-2120.
15. Клячко Н.Л. Актин растений: множественные уровни регуляции // Физиол. раст. -2006. - Т. 53, № 5. - С. 790-798.
16. Клячко Н.Л. Актиновый цитоскелет и форма растительной клетки // Физиол. раст. -2004. - Т. 51, № 6. - С. 918-925.
17. Fu Y., Wu G., Yang Z. Rop GTPase-Dependent Dynamics of Tip-Localized F-Actin Controls Tip Growth in Pollen Tubes //J. Cell Biol. - 2001. - V. 152, No 5. - P. 1019-1032.
18. Новиков Г.В., Мошкова И.Е. Суперсемейство мономерных ГТФ-связывающих белков растений. 2. RAB-белки - регуляторы везикулярного транспорта и ответных реакций растений на стрессы // Физиол. раст. - 2008. - Т. 55, № 1. - С. 127-139.
19. Wu G., Gu Y., Li S., Yang Z. A Genome-Wide Analysis of Arabidopsis Rop-Interactive CRIB Motif-Containing Proteins That Act as Rop GTPase Targets // Plant Cell. - 2001. -V. 13, No 12. - P. 2841-2856.
20. Van Gestel K., Slegers H., von Witsch M., Samaj J., Baluska F., Verbelen J.P. Immuno-logical Evidence for the Presence of Plant Homologues of the Actin-Related Protein
Arp3 in Tobacco and Maize: Subcellular Localization to Actin-Enriched Pit Fields and Emerging Root Hairs // Protoplasma. - 2003. - V. 222, No 1-2. - P. 45-52.
21. Mathur J., Mathur N., KernebeckВ., HulskampM. Mutations in Actin-Related Proteins 2 and 3 Affect Cell Shape Development in Arabidopsis II // Plant Cell. - 2003. - V. 15, No 7. - P. 1632-1645.
22. Le J., El-Assal Salah El-Din, Basu D., Saad M.E., Szymanski D.B. Requirements for Arabidopsis AtARPl and AiARPJ during Epidermal Development // Curr. Biol. - 2003. -V. 13, No 8. - P. 1341-1347.
23. Li H., Lin Y., Heath R.M., Zhu M.X., Yang Z. Control of Pollen Tube Tip Growth by a Rop GTPase-Dependent Pathway That Leads to Tip-Localized Calcium Influx // Plant Cell. - 1999. - V. 11, No 9. - P. 1731-1742.
24. Mathur J. The Arp2/3 Complex: A Giving Plant Cells a Leading Edge // BioEssays. -2005. - V. 27, No 4. - P. 377-387.
25. Cheung A.Y., Wu H.M. Over expression of an Arabidopsis Formin Stimulates Supernumerary Actin Cable Formation from Pollen Tube Cell Membrane // The Plant Cell. -2004. - V. 16, No 1. - P. 257-269.
26. Baluska F., Salaj J., Mathur J., Braun M., Jasper F., Samaj J., Chua N.-H., Barlow P.W., Volkmann D. Root hair formation: F-actin-dependent tip growth is initiated by local assembly of profilin-supported F-actin meshworks accumulated within expansin-enriched bulges // Dev. Biol. - 2000. - V. 227, No 2. - P. 618-632.
27. Friml J., Wisniewska J., Benkova E., Mendgen K., Palme K. Lateral relocation of auxin efflux regulator PIN3 mediates tropism in Arabidopsis // Nature. - 2002. - V.415, No 2. -P. 806-809.
28. Orwar B.L., Sangwan V., Omann F., Dhindsa R. Early steps in cold sension by plant cells: of actin cytoskeleton and membrane fluidity // Plant J. - 2000. - V. 23, No 6. - P. 785-794.
29. Sangwan V., Orvar B.L., Beyerly J., Hirt H., Dhindsa R.S. Opposite changes in membrane fluidity mimic cold and heat stress activation of distinct plant MAP-kinase pathways // Plant J. - 2002. - V. 31, No 5. - P. 629-638.
30. Васильев А.Е. Сравнительная структурно-функциональная характеристика цитоске-лета животных и высших растений // Журн. общ. биол. - 1996. - Т. 57, № 3. -С. 293-325.
31. Heslop-Harrison J., Heslop-Harrison Y. Myosin Associated with the Surface of Organel-les, Vegetative Nuclei and Generative Cells in Angiosperm Pollen Grains and Tubes // J. Cell Sci. - 1989. - V. 94. - P. 319-325.
32. Reichelt S., Knight A.E., Hodge Т.Р., Baluska F., Samaj J., Volkmann D., Kendrick-Jones J. Characterization of Unconventional Myosin VIII in Plant Cells and Its Localization at the Postcytokinetic Cell Wall // Plant J. - 1999. - V. 19, No 5. - P. 555-567.
33. Lee Y.P.J., Liu B. Cytoskeletal Motors in Arabidopsis. Sixty-One Kinesins and Seventeen Miosins // Plant Physiol. - 2004. - V. 136, No 4. - P. 3877-3883.
34. Cyr R.J., Palevitz B.A. Organization of cortical microtubules in plant cells // Curr. Opin. Cell Biol. - 1995. - V. 7, No 1. - P. 65-71.
35. Bush D.S. Calcium regulation in plant cells and its role in signaling // Annu. Rev. Plant Phisiol. Plant Mol. Biol. - 1995. - V. 46. - P. 95-122.
36. Fisher D.D., Cyr R.J. Calcium levels affect the ability to immunolocalize calmodulin to cortical microtubules // Plant Physiol. - 1993. - V. 103, No 2. - P. 543-551.
37. Fisher D., Gilroy S., Cyr R. Evidence for opposing effects of calmodulin on cortical microtubules // Plant Physiol. - 1996. - V. 112, No 3. - P. 1079-1087.
38. Туркина M.B., Соколов О.И. Миозины - моторные белки актомиозиновой системы подвижности: связь с мембранами и путями передачи сигналов // Физиол. раст. -2001. - Т. 48, № 5. - С. 788-800.
39. Baluska F., Samay J., Wojtaszek P., Volkman D., Menzel D. Cytoskeleton-Plasma Membrane-Cell Wall Continuum in Plants. Emerging Links Revisited // Plant Physiol. - 2003. -V. 133, No 2. - P. 482-491.
40. Zhou J., Wang B., Li Y., Wang Y., Zhu L. Responses of Chrysanthemum Cells to Mechanical Stimulation Require Intact Microtubules and Plasma Membrane - Cell Wall Adhesion // J. Plant Growth Regul. - 2007. - V. 26, No 1. - P. 55-68.
41. Медведев С.С. Кальциевая сигнальная система растений // Физиол. раст. - 2005. -T. 52, № 2. - C. 282-305.
42. Abdrakhamanova A.F., Wang Q.Y., Khokhlova L.P., Nick P. Is Microtubule Disassembly a Trigger for Cold Acclimation? // Plant Cell Physiol. - 2003. - V.44, No 7. - P. 676-686.
43. Thomashow M.F. So what's new in the field of plant cold acclimation? Lots! // Plant Physiol. - 2001. - V. 125, No 1. - P. 89-93.
44. Monroy A.F., Castonguay Y., Laberge S., Sarhan F., Vezina L.P., Dhindsa R.R. A new cold-induced alfalfa gene is associated with enhanced hardening at subzero temperature // Plant Physiol. - 1993. - V. 102, No 3. - P. 873-879.
45. Nick P. Plant Microtubules. Development and Flexibility (Plant Cell Monogr. V. 11). -Berlin; Heidelberg; N. Y.: Springer-Verlag2008. - XIV + 269 p.
46. Sangwan V., Foulds I., Singh J., Dhindsa R.S. Cold-activation of Brassica napus BN115 promoter is mediated by structural changes in membranes and cytoskeleton, and requires Ca2+ influx // Plant J. - 2001. - V. 27, No 1. - P. 1-12.
47. Mazars C., Thion L., Thuleau P., Graziana A., Knight M.R., Moreau M., Ranjeva R. Organization of cytoskeleton controls the changes in cytosolic calcium of cold-shocked Nicotiana plumbaginifolia protoplasts // Cell Calcium. - 1997. - V. 22. - P. 413-420.
48. Oulett F., Carpenet E., Cope M J., Monroy A.F., Sarhan F. Regulation of a Wheat Actin-Depolymerizing factor during Cold Acclimation // Plant Physiol. - 2001. - V. 125, No 1. -P. 360-368.
49. Waser M., Mesaeli N., Spencer C., Michalak M. Regulation of Calreticulin Gene Expression by Calcium // J.Cell Biol. - 1997. - V. 138, No 3. - P. 547-557.
50. Denecke J., Carlsson L.E., Vidal S., Hoglund A., Ek B., Zeijl M.J., Sinjorgo K.M.C., Palva T. The Tobacco Homolog of Mammalian Calreticulin Is Present in Protein Complexes in vivo // Plant Cell. - 1995. - V. 7, No 4. - P. 391-406.
51. MichalakM., Mariani P., Opas M. Calreticulin, a Multifunctional Ca2+ Binding Chaperone of the Endoplasmic Reticulum // Biochem. Cell Biol. - 1998. - V.76, No 5. - P. 779-785.
52. Mery L., Mesaeli N., MichalakM., Opas M., Lew D.P., Krause K.H. Overexpresssion of Calreticulin Increases Intracellular Ca2+ Storage and Decreases Store-Operated Ca2+ Influx // J. Biol. Chem. - 1996. - V. 271, No 16. - P. 9332-9339.
53. Debhar S., Rennie P.S., Shago M., Leung-Hagesteijn C.Y., Filmus J., Hawley R.G., Bru-chovsky N., Cheng H., Matusik R.J., Giguere V. Inhibition of Nuclear Hormone Receptor Activity by Calreticulin // Nature. - 1994. - V.367, No 6462. - P. 480-483.
54. Sapiro R.G., Zhu Q., Bhoyroo V., Soling H.D. Definition of the Lectin-Like Properties of the Molecular Chaperone, Calreticulin, and Demonstration of Its Copurification with En-domannosidase from Rat Liver Golgi // J. Biol. Chem. - 1996. - V. 271, No 19. -P. 11588-11594.
55. Lievremont J.P., Rizzuto R., Hendershot L., Meldolesi J. Bip, a Major Chaperone Protein of the Endoplasmic Reticulum Lumen, Plays a Direct and Important Role in the Storage
of the Rapidly Exchanging Pool of Ca2+ // J. Biol. Chem. - 1997. - V. 272, No 49. -P. 30873-30879.
56. Anderson J.V., Li Q.B., Guy C.L. Structural Organization of the Spinach Endoplasmic Reticulum Luminal 70-kilodalton Heat-Shock Cognate Gene and Expression of 70-kilo-dalton Heat-Shock Genes during Cold Acclimation // Plant Physiol. - 1994. - V. 104, No 4. - P. 1359-1370.
57. Coppolino M.G., Dedhar S. Ligand-Specific, Transient Interaction between Integrins and Calreticulin during Cell Adhesion to Extracellular Matrix Proteins Is Dependent upon Phosphorylation/Dephosphorylation Events // Biochem. J. - 1999. - V. 340. - P. 41-50.
58. Mogelsvang S., Simpson P.J. Changes in the Levels of Seven Proteins Involved in Polypeptide Folding and Transport during Endosperm Development of Two Barley Genotypes Differing in Storage Protein Localization // Plant Mol. Biol. - 1998. - V. 36. - P. 541-552.
59. Олиневич О.В., Хохлова Л.П. Цитоскелет-зависимые изменения структурной организации ретикулоплазминов в клетках пшеницы при закаливании к холоду и действии абсцизовой кислоты // Физиол. раст. - 2002. - Т. 49, № 2. - С. 221-229.
60. Олиневич О.В., Хохлова Л.П., Раудаскоски М. Ретикулоплазмины - Бип и Калретику-лин при действии на проростки пшеницы Triticum aestivum L. низких температур и антимитотического агента оризалина // Докл. РАН. - 2001. - Т. 377, № 1. - С. 135-138.
61. Baluska F., Volkman D., Menzel D. Plant synapses: actin-based domains for cell-to-cell communication // Trend Plant Sci. - 2005. - V. 10, No 3. - P. 106-111.
62. Campanoni P., Blasius B., Nick P. Auxin Transport Synchronizes the Pattern of Cell Division in a Tobacco Cell Line1 // Plant Physiol. - 2003. - V. 133, No 3. - P. 1251-1260.
63. Schulze-Lefert P. Knocking on the heavens wall: pathogenesis of and resistance to bio-trophic fungi at the cell wall // Curr. Opin. Plant Biol. - 2004. - V. 7, No 1. - P. 1-7.
64. Клячко Н.Л. Цитоскелет и внутриклеточная подвижность у растений // Физиол. раст. - 2005. - Т. 52, № 5. - С. 786-795.
65. Reddy A.S.N. Molecular motors and their functions in plants // Int. Rev. Cytol. - 2001. -V. 204. - P. 97-178.
66. Coluccio L.M. Myosins: A Superfamily of Molecular Motors. - Dordrecht, The Netherlands: Springer, 2007. - 475 p.
67. Reddy A.S.H., Day I.S. Kinesins in the Arabidopsis Genome: A Comparative Analysis among Eukaryotes // BMC Genomics. - 2001. - V. 2, No 2. - URL: http://www.biomedcentral.com/content/pdf/1471-2164-2-2.pdf, свободный.
68. Marcus A.J., Li W., Ma H., Cyr R.J. A Kinesin Mutant with an Atypical Bipolar Spindle Undergoes Normal Mitosis // Mol. Biol. Cell. - 2003. - V. 14, No 4. - P. 1717-1726.
69. Preuss M.L., Delmer D.P., Liu B. The Cotton Kinesin-Like Calmodulin-Binding Protein Associates with Cortical Microtubules in Cotton Fibers // Plant Physiol. - 2003. - V. 132, No 1. - P. 154-160.
70. PreussM.L., Kovar D.P., Lee Y.-R. J., Starger С. J., Delmer D.P., Liu B. A Plant-Specific Kinesin Binds to Actin Microfilaments and Interacts with Cortical Мicrotubules in Cotton Fibers // Plant Physiol. - 2004. - V. 136, No 4. - P. 3945-3955.
71. Zhong P., Burk D.H., Morrison W.H., Ye Z.-H. A Kinesin-Like Protein Is Essential for Oriented Deposition of Cellulose Microfibrils and Cell Wall Strength // Cell. - 2002. -V. 14, No 12. - P. 3101-3117.
72. Moscatelli A., del Casino C., Lozzi L., Cai G., Scali Tiezzi A., Cresti M. High Molecular Weight Polypeptides Related to Dynein Heavy Chains in Nicotiana tabacum Pollen Tubes // J. Cell Sci. - 1995. - V. 1. - P. 1117-1125.
73. Romagnoli S., Cai G., Cresti M. In Vitro Assays Demonstrate That Pollen Tube Organelles Use Kinesin-Related Motor Proteins to Move along Microtubules // Plant Cell. - 2003. -V. 15, No 1. - P. 251-269.
74. Sakai Т., Kagawa Т., Kasahara M., Swartz Т.Е., Christie J. M., Briggs W.Ji., Wada M., Okada K. Arabidopsis Nphl and Np11: Blue Light Receptors That Mediate Both Photo-tropism and Chloroplast Relocation // Proc. Natl. Acad. Sci. USA. - 2001. - V. 98. -P. 6969-6974.
75. Sheahan M.B., Jose R.J., McCurdy D.W. Organelle Inheritance in Plant Cell Division: The Actin Cytoskeleton Is Required for Unbiased Inheritance of Chloroplasts, Mitochondria and Endoplasmic Reticulum in Dividing Protoplasts // Plant J. - 2004. - V. 37, No 3. -P. 379-390.
76. Wang Q.Y., Nick P. Cold acclimation can induce microtubular cold stability in a manner distinct from abscisic acid // Plant Cell Physiol. - 2001. - V. 42, No 9. - P. 999-1005.
77. Blancaflor E.B., Hasenstein K.H. Organization of cortical microtubules in graviresponding maize roots // Planta. - 1993. - V. 191, No 2. - P. 231-237.
78. Shibaoka H. Regulation by gibberellins of the orientation of cortical microtubules in plant cells // Austr. J. Plant Physiol. - 1993. - V. 20, No 5. - P. 461-470.
79. Shibaoka H. Involvement of wall microtubules in gibberellin promotion and kinetin inhibition of stem elongation // Plant Cell Physiol. - 1974. - V. 15, No 2. - P. 255-263.
80. Grabski S., Schindler M. Auxins and cytokinins as antipodal modulators of elasticity within the actin network of plant cells // Plant Physiol. - 1996. - V. 110, No 3. - P. 965-970.
81. Lang J.M., Eisinger W.R., Green P.B. Effects of ethylene on the orientation of microtubules and cellulose microfibrils of pea epicotyls with polylamellate cell walls // Proto-plasma. - 1982. - V. 110, No 1. - P. 5-14.
82. Sakiyama M., Shibaoka H. Effects of abscisic acid on the orientation and cold stability of cortical microtubules in epicotyl cells of the dwarf pea // Protoplasma. - 1990. - V. 157, No 1-3. - P. 165-171.
83. Клячко Н.Л. Фитогормоны и цитоскелет // Физиол. раст. - 2003. - Т. 50, № 3. -С. 475-480.
84. Хохлова Л.П., Олиневич О.В., Макарова М.В., Бочкарева М.А. Морфофизиологиче-ские изменения корней разных генотипов озимой пшеницы в связи с деструкцией цитоскелета // Физиол. раст. - 2006. - Т. 53, № 3. - C. 418-430.
85. Olinevich O.V., Khokhlova L.P., Raudaskoski M. The microtubule stability increases in abscisic acid-treated and cold-acclimated differentiating vascular root tissues of wheat // J. Plant Physiol. - 2002. - V. 159. - P. 465-472.
86. Олиневич О.В., Хохлова Л.П. Реорганизация тубулинового и актинового цитоскеле-та при закаливании растений Triticum aestivum L. к холоду и действии абсцизовой кислоты // Цитология. - 2002. - Т. 44, № 6. - С. 532-543.
87. Олиневич О.В., Хохлова Л.П. Влияние абсцизовой кислоты, низких температур и возраста растений на цитоскелетные и фосфорилированные белки // Биохимия. -2003. - Т. 68, Вып. 6. - С. 828-839.
88. Хохлова Л.П., Олиневич О.В. Реорганизация цитоскелета в клетках Triticum aestivum при закаливании растений к холоду и действии абсцизовой кислоты // Физиол. раст. - 2003. - Т. 50, № 4. - С. 528-540.
89. Veisz O., Galiba G., Sutka J. Effect of abscisic acid on the cold hardiness of wheat seedlings // J. Plant Physiol. - 1996. - V. 149. - P. 439-443.
90. Samaj J., Baluska F., Voigt B,. Schlicht M., Volkmann D., Menzel D. Endocytosis, Actin Cytoskeleton, and Signaling // Plant Physiol. - 2004. - V. 135, No 3. - P. 1150-1161.
91. Baluska F., Hlavacka A., Samaj J., Palme K., Robinson D.G., Matoh T., McCurdy D.W., Menzel D., Volkmann D. F-actin-dependent endocytosis of cell wall pectins in meristematic root cells: insights from brefeldin A-induced compartments // Plant Physiol. - 2002. -V. 130, No 1. - P. 422-431.
92. Geldner N., Friml J., Stierhof Y.-D., Jurgens G., Palme K. Auxin-transport inhibitors block PIN1 cycling and vesicle trafficking // Nature. - 2001. - V. 413, No 6854. - P. 425-428.
93. Geldner N., Anders N., Wolters H., Keicher J., Kornberger W., Muller P., Delbarre A., Ueda T., Nakano A., Jurgens G. The Arabidopsis GNOM ARF-GEF mediates endosomal recycling, auxin transport, and auxin-dependent plant growth // Cell. - 2003. - V. 112, No 2. - P. 219-230.
94. Grebe M., Xu J., Mobius W., Ueda T., Nakano A., Geuze H.J., Rook M.B., Scheres B. Arabidopsis sterol endocytosis involves actin-mediated trafficking via ARA6-positive early endosomes // Curr. Biol. - 2003. - V. 13, No 16. - P. 1378-1387.
95. Ueda T., Nakano A. Vesicular traffic: an integral part of plant life // Curr. Opin. Plant. Biol. - 2002. - V. 5, No 6. - P. 513-517.
96. Vernoud V., Horton A.C., Yang Z., Nielesen E. Analysis of the small GTPase gene super-family of Arabidopsis // Plant Physiol. - 2003. - V. 131, No 3. - P. 1191-1208.
97. Gu Y., Vernoud V., Fu Y., Yang Z. ROP GTPase Regulation of Pollen Tube Growth through the Dynamics of Tip-Localized F-Actin // J. Exp. Bot. - 2003. - V. 54, No 380. - P. 93-101.
98. Barth M., Holstein S.E.H. Identification and functional characterization of Arabidopsis AP180, a binding-partner of plant aC-adaptin // J. Cell Sci. - 2004. - V. 117. - P. 2051-2062.
99. XuX., Bittman R., Duportail G., Heissler D., Vilchezes C., London E. Effect of the structure of natural sterols and sphingolipids on the formation of ordered sphingolipid/sterol domains (rafts) // J. Biol. Chem. - 2001. - V. 276, No 36. - P. 33540-33546.
100. Lippincott-Schwartz J., Yuan L.C., Tipper C., AmherdtM., Orci L., Klausner R.D. Brefeldin A's effects on endosomes, lysosomes and the TGN suggest a general mechanism for regulating organelle structure and membrane traffic // Cell. - 1991. - V. 67, No 3. - P. 601-616.
101. Wood S.A., Brown W.J. The morphology but not the function of endosomes and lysosomes is altered by brefeldin A // J. Cell Biol. - 1992. - V. 119, No 2. - P. 273-285.
102. Baluska F., Samaj J., Hlavacka A., Kendrick-Jones J., Volkmann D. Actin-dependent fluid-phase endocytosis in inner cortex cells of maize root apices // J. Exp. Bot. - 2004. -V. 55, No 396. - P. 463-473.
103. Kang B.-H., Busse J.S., Bednarek S.Y. Members of the Arabidopsis dynamin-like gene family, ADL1, are essential for plant cytokinesis and polarized cell growth // Plant Cell. -2003. - V. 15, No 4. - P.899-913.
104. Макарова М.В. Морфофизиологические изменения корней озимой пшеницы в связи с деструкцией цитоскелета при действии индукторов морозоустойчивости: Авто-реф. канд. биол. наук. - Казань, 2007. - 25 с.
105. Pelkmans L., Puntener D., Helenius A. Local actin polymerization and dynamin recruitment in SV40-induced internalization of caveolae // Science. - 2002. - V. 296, No 5567. -P. 535-539.
106. Orth J.D., McNiven M.A. Dynamin at the actin-membrane interface // Curr. Opin. Cell Biol. - 2003. - V. 15, No 1. - P. 31-39.
107. Kang B.-H., Rancour D.M., Bednarek S. Y. The dynamin-like protein ADL1C is essential for plasma membrane maintenance during pollen maturation // Plant J. - 2003. - V. 35, No 1. - P. 1-15.
108. Hong Z., Geisler-Lee J., Zhang Z., Verma D.P.S. Phragmoplastin dynamics: multiple forms, microtubule association and their roles in cell plate formation in plants // Plant Mol. Biol. - 2003. - V. 53, No 3. - P. 297-312.
109. Samaj J., Ovecka M., Hlavacka A., Lecourieux F., Meskiene I., Lichtscheidl I., Lenart P., Salaj J., Volkmann D., Bogre L., Baluka F., Hirt H. Involvement of the mitogen-activated protein kinase SIMK in regulation of root hair tip growth // EMBO. - 2002. - V. 21, No 13. - P. 3296-3306.
110. Samaj J., Baluska F., Hirt H. From signal to cell polarity: mitogen-activated protein kinases as sensors and effectors of cytoskeleton dynamicity // J. Exp. Bot. - 2004. -V. 55, No 395. - P. 189-198.
111. Blancaflor E.B. Cortical actin filaments potentially interact with cortical microtubules inregulating polarity of cell expansion in primary roots of maize (Zea mays L.) // J. Plant Growth Regul. - 2000. - V. 19, No 4. - Р. 406-414.
112. Tominaga M., Morita K., Yokota E., Shimmen T. Microtubules regulate the organization of actin filaments at the cortical region in root hair cells of Hydrocharis // Protoplasma. -1997. - V. 199, No 1-2. - P. 83-92.
113. Lanerolle de P., Cole A.B. Cytoskeletal Proteins and Gene Regulation: Form, Function and Signal Transduction in the Nucleus // Sci. STKE. - 2002. - V. 139. - P. 30-35.
114. Giani S., Qin X., Faoro F., Bleviario D. In Rice, Oryzalin and Abscisic Acid Differentially Affect Tubulin mRNA and Protein Levels // Planta. - 1998. - V. 205, No 3. - P. 334-341.
115. Hoffman K.C., Vaughn K.C. Mitotic Disrupters Act by a Single Mechanism but Vary in Efficacy // Protoplasma. - 1994. - V. 179, No 1-2. - P. 16-25.
116. Кундельчук О.П., Тарасенко Л.В., Блюм Я.Б. Сравнение действия амипрофос-метила на структуру клеток корня у чувствительных и устойчивых к нему линий Nicotiana plumbaginifolia // Физиол. раст. - 2002. - Т. 49, № 3. - С. 425-430.
117. Volkmann D., Baluska F. Actin Cytoskeleton in Plants: From Transport Networks to Signaling Networks // Microsc. Res. Tech. - 1999. - V. 47, No 2. - P. 135-154.
118. Qiu J.-L., Jilk R., Marks M.D., Szymanski D.B. The Arabidopsis SPIKE 1 Gene Is Required for Normal Cell Shape Control and Tissue Development // Plant Cell. - 2002. -V. 14, No 1. - P. 101-118.
119. FrankM.J., Smith L.G. A Small, Novel Protein Highly Conserved in Plants and Animals Promotes the Polarized Growth and Division of Maize Leaf Epidermal Cells // Curr. Biol. - 2002. - V. 12, No 10. - P. 849-853.
Поступила в редакцию 10.10.10
Хохлова Людмила Петровна - доктор биологических наук, профессор кафедры физиологии и биотехнологии растений Казанского (Приволжского) федерального университета.
E-mail: [email protected]
Невмержицкая Юлия Юрьевна - кандидат биологический наук, старший преподаватель кафедры физиологии и биотехнологии растений Казанского (Приволжского) федерального университета.
E-mail: [email protected]
