結語與未來展望 - 水稻葉鞘在抽穗期間由儲存組織轉換成供源組織之分子調控機制

本論文為瞭解水稻葉鞘於抽穗期間儲存–供源轉換的分子調控機制，首先篩選出參與水稻葉鞘儲存-供源轉換之標識基因，包含醣類代謝相關基因 (AGP-L2、

GBSSII、SSSI、SBEI、SBEIII、SBEIV、sucrose synthase與 β-amylase)、蔗糖轉運蛋 白 (OsSUT1與OsSUT4) 及分子轉運蛋白 (putative sorbitol transporter、ammonium transporter與phosphate transporter) 等基因。藉由分析澱粉合成相關基因啟動子序 列及荷爾蒙實驗，得知ABA雖然並非是誘導葉鞘儲存-供源轉變之主因，但其可能於供源型葉鞘中促進澱粉分解及蔗糖轉運蛋白基因表現。我們也發現葡萄糖與蔗糖含量於抽穗後上升的趨勢與蔗糖轉運蛋白表現量相似，暗示其極可能影響蔗糖轉運蛋白在葉鞘中的表現，而醣類分子與ABA之間是否對蔗糖轉運蛋白有共同協調的關係存在，未來可以利用荷爾蒙抑制劑的處理來佐證。利用啟動子分析來了解水稻蔗糖轉運蛋白於供源型葉鞘中表現的分子調控機制，得到一段67 bp之啟動

子序列，其可能包含調控蔗糖轉運蛋白於抽穗後大量表現的調控區域。未來，尋找這段啟動子片段上之結合轉錄因子以及證實該轉錄因子之調控特性將是首要工作，預期能藉此更了解水稻葉鞘儲存-供源轉換之分子調控機制，在未來，可以應用於增強水稻葉鞘於抽穗前儲存醣類之能力或抽穗之後之供源強度，以提高水稻產量及降低環境逆境對水稻減產之衝擊。

參考資料

陳佳宜 (2007) 水稻種子發芽時期胚中蔗糖轉運蛋白基因之表現調控機制。國立臺灣大學生物資源暨農學院農藝系碩士論文。

Abe H, Yamaguchi-Shinozaki K, Urao T, Iwasaki T, Hosokawa D, Shinozaki K (1997) Role of ara-bidopsis MYC and MYB homologs in drought- and abscisic acid-regulated gene expression. Plant Cell 9:1859–1868

Abe H, Urao T, Ito T, Seki M, Shinozaki K, Yamaguchi–Shinozaki K (2003) Arabidopsis AtMYC2 (bHLH) and AtMYB2 (MYB) function as transcriptional activators in abscisic acid signaling. Plant Cell 15: 63–78

Akihiro T, Mizuno K, Fujimura T (2005) Gene expression of ADP–glucose pyrophosphorylase and starch contents in rice cultured cells are cooperatively regulated by sucrose and ABA. Plant Cell Phy-siol 46: 937–946

Aloni B, Daie J, Wyse RE (1986) Enhancement of ['^C]sucrose export from source leaves of Vicia faba by gibherelhc acid. Plant Physiol 82: 962-966

Aoki N, Hirose T, Scofield GN, Whitfeld PR, Furbank RT (2003) The sucrose transporter gene family in rice. Plant Cell Physiol 44: 223–232

Baba I, Kittaka A (1953) On the accumulation of starch grains at the shoot base of the rice plant. Proc Crop Sci Soc Jpn 22: 43–44

Barker L, Kuhn C, Weise A, Schulz A, Gebhardt C, Hirner B, Hellmann H, Schulze W, Ward JM, Frommer WB (2000) SUT2, a putative Suc sensor in sieve elements. Plant Cell 12: 1153–1164 Baroja-Fernandez E, Munoz FJ, Saikusa T, Rodriguez-Lopez M, Akazawa T, Pozueta-Romero J

(2003) Sucrose synthase catalyzes the de novo production of ADPglucose linked to starch biosynthesis in heterotrophic tissues of plants. Plant Cell Physiol 44: 500–509

Blakeney AG, Matheson TH (1984) Some properties of the stem and pollen starches of rice. Starch 36:

265–269

Blum A, Sinmena B, Mayer J, Golan G, Shpiler L (1994) Stem reserve mobilization supports wheat–grain filling under heat stress. Aust J Plant Physiol 21: 771–781

Bradford MM (1976) A rapid and sensitive method for the quantitation of microgram quantities of pro-tein utilizing the principle of propro-tein–dye binding. Anal Biochem 72: 248–254

Braun DM, Slewinski TL (2009) Genetic control of carbon partitioning in grasses: roles of sucrose transporters and tie–dyed loci in phloem loading. Plant Physiol 149: 71–81

Burton RA, Bewley JD, Smith AM, Bhattacharyya MK, Tatge H, Ring S, Bull V, Hamilton WDO, Martin C (1995) Starch branching enzymes belonging to distinct enzyme families are differentially expressed during pea embryo development. Plant J 7: 3–15

Chen C, Chen Z (2002) Potentiation of developmentally regulated plant defense response by AtWRKY18, a pathogen-induced Arabidopsis transcription factor. Plant Physiol 129: 706-16

Chen HJ, Wang SJ (2008) Molecular regulation of sink-source transition in rice leaf sheaths during the heading period. Acta Physiol Plant 30: 639-649

Chen WS, Liu HY, Yang ZH, Chen WH (1994) Gibberellin and temperature influence carbohydrate content and flowering in Phalaenopsis. Physiol Plant 90: 391–395

Chincinska IA, Liesche J, Krügel U, Michalska J, Geigenberger P, Grimm B, Kühn C (2008) Su-crose transporter StSUT4 from potato affects flowering, tuberization, and shade avoidance response.

Plant Physiol 146: 515–528

Chiou TJ, Bush DR (1998) Sucrose is a signal molecule in assimilate partitioning. Proc Natl Acad Sci USA 95: 4784–4788

Chu C and Lee TM (1989) The relationship between ethylene biosynthesis and chilling tolerance in seedlings of rice (Oryza sativa L.). Bot Bull Acad Sinica 30: 263–273

Cock JH, Yoshida S (1972) Accumulation of ¹⁴C–labelled carbohydrate before flowing and its subse-quent redistribution and respiration in the rice plant. Proc Crop Sci Soc Jpn 41: 226–234

Denyer K, Dunlap F, Thorbjornsen T, Keeling P, Smith AM (1996) The major form of ADP-glucose pyrophosphorylase in maize (Zea mays L.) endosperm is extra-plastidial. Plant Physiol 112: 779–785 Dian WM, Jiang HW, Chen QS, Liu FY, Wu P (2003) Cloning and characterization of the

gra-nule–bound starch synthase II gene in rice: gene expression is regulated by the nitrogen level, sugar and circadian rhythm. Planta 218: 261–268

Dian WM, Jiang HW, Wu P (2005) Evolution and expression analysis of starch synthase III and IV in rice. Journal of Experimental Botany 56: 623–632

Ding B, Parthasarathy MV, Niklas K, Turgeon R (1988) A morphometric analysis of the phloem-unloading pathway in developing tobacco leaves. Planta 176: 307–318

Edwards A, Borthakur A, Bornemann S, Venail J, Denyer K, Waite D, Fulton D, Smith A, Martin C (1999) Specificity of starch synthase isoforms from potato. Eur J Biochem 266: 724–736

Endler A, Meyer S, Schelbert S, Schneider T, Weschke W, Peters SW, Keller F, Baginsky S, Marti-noia E, Schmidt UG (2006) Identification of a vacuolar sucrose transporter in Hordeum vulgare and Arabidopsis thaliana by a tonoplast proteomic approach. Plant Physiol 141: 196–207

Eulgem T, Rushton PJ, Elmon Schmelzer E, Hahlbrock K, Somssich IE (1999) Early nuclear event in plant defence: rapid gene activation by WRKY transcription factors. EMBO J 18: 4689–4699

Eulgem T, Rushton PJ, Robatzek S, Somssich IE (2000) The WRKY superfamily of plant transcription factors. Trends Plant Sci 5: 199-206

Fried MG (1989) Measurement of protein-DNA interaction parameters by electrophoresis mobility shift assay. Electrophoresis 10: 366-376

Furbank RT, Scofield G, Hirose T, Wang XD, Patrick J, Offler C (2001) Cellular localisation and function of a sucrose transporter OsSUT1 in developing rice grains. Aust J Plant Physiol 28:

1187–1196

Gao Z, Maurousset L, Lemoine R, Yoo SD, Nocker S, Loescher W (2003) Cloning, expression, and 73

characterization of sorbitol transporters from developing sour cherry fruit and leaf sink tissues. Plant Physiol 131: 1–10

Ghosh HP, Preiss J (1966) Adenosine diphosphate glucose pyrophosphorylase. A regulatory enzyme in the biosynthesis of starch in spinach leaf chloroplasts. J Biol Chem 241: 4491–4504

Guan HP, Preiss J (1993) Differentiation of the properties of the starch branching isozymes from maize (Zea mays). Plant Physiol 102: 1269–1273

Hanggi E, Fleming AJ (2001) Sucrose synthase expression pattern in young maize leaves: implications for phloem transport. Planta 214: 326–329

Harms K, Wöhner RV, Schulz B, Frommer WB (1994) Isolation and characterization of P–type H(+)–ATPase genes from potato. Plant Mol Biol 26: 979–988

Hiei Y, Ohta S, Komari T, Kumashiro T (1994) Efficient transformation of rice (Oryza sativa L.) me-diated by Agrobacterium and sequence analysis of the boundaries of the T-DNA. Plant J 6: 271–282 Higo K, UgawaY, Iwamoto M, Korenaga T (1999) Plant cis–acting regulatory DNA elements (PLACE)

database. Nucleic Acids Res 27: 297–300

Hirose T, Endler A, Ohsugi R (1999) Gene expression of enzymes for starch and sucrose metabolism and transport in leaf sheaths of rice (Oryza sativa L.) during the heading period in relation to the sink to source transition. Plant Prod Sci 2: 178–183

Hirose T, Imaizumi N, Scofield GN, Furbank RT, Ohsugi R (1997) cDNA cloning and tissue–specific expression of a gene for sucrose transporter from rice (Oryza sativa L.). Plant Cell Physiol 38:

1389–1396

Hirose T, Ohdan T, Nakamura Y, Terao T (2006) Expression profiling of genes related to starch syn-thesis in rice leaf sheaths during the heading period. Physiol Plant 128: 425–435

Hu CY, Chee PP, Chesney RH, Zhou JH, Miller PD, O'Brien WT (1990) Intrinsic GUS–like activi-ties in seed plants. Plant Cell Rep 9: 1–5

Hurng WP, Lur HS, Liao CK, Kao CH (1994) Role of abscisic acid, ethylene and polyamines in flooding-promoted senescence of tobacco leaves. J Plant Physiol 143:102–105

Imlau A, Truernit E, Sauer N (1999) Cell-to-cell and long-distance trafficking of the green fluorescent protein in the phloem and symplastic unloading of the protein into sink tissues. Plant Cell 11: 309–322 Ishikawa T, Akita S, Li Q (1993) Relationship between contents of non–structural carbohydrates before

panicle initiation stage and grain yield in rice (Oryza sativa L.). Jpn J Crop Sci 62: 130–131

Ishimaru K, Hirose T, Aoki N, Takahashi S, Ono K, Yamamoto S, Wu J, Saji S, Baba T, Ugaki M, Matsumotoi T, Ohsugi R (2001) Antisense expression of a rice sucrose transporter OsSUT1 in rice (Oryza sativa L.). Plant Cell Physiol 42: 1181–1185

Ishimaru K, Kosone M, Sasaki H, Kashiwagi T (2004) Leaf contents differ depending on the position in a rice leaf sheath during sink–source transition. Plant Physiol Biochem 42: 855–860

Ishiwatari Y, Nemoto K, Fujiwara T, Chino M, Hayashi H (2000) In situ hybridization study of the rice phloem thioredoxin h mRNA accumulation– possible involvement in the differentiation of

vascu-74

lar tissues. Physiol Plant 109: 90–96

Itoh K, Yamaguchi J, Huang N, Rodriguez RL, Akazawa T, Shimamoto K (1995) Developmental hormonal regulation of rice -amylase (RAmy1A)-gusA fusion genes in transgenic rice seeds. Plan ysiol 107: 25–31

and t

Jefferson RA, Kavanagh T A, Bevan MW (1987) GUS fusions: β–glucuronidase as a sensitive and versatile gene fusion marker in higher plants. EMBO J 6: 3901–3907

Jones H, Eagles JE (1962) Translocation of ⁴carbon within and between leaves. Ann Bot N S 26:

505–510

Keppler D, Decker K (1974) Glycogen: Determination with amyloglucosidase. In” Bergmeyer HU (ed) Methods of enzymatic analysis” vol 3. Academic Press, New York, pp 1127–1131

Koch KE (1996) Carbohydrate–modulated gene expression in plants. Annu Rev Plant Physiol Plant Mol Biol 47: 509–540

Kossmann J, Abel GJ, Springer F, Lloyd JR, Willmitzer L (1999) Cloning and functional analysis of a cDNA encoding a starch synthase from potato (Solanum tuberosum L.) that is predominantly ex-pressed in leaf tissue. Planta 208: 503–11

Kumar A, Silim SN, Okamoto M, Siddiqi MY, Glass AD (2003) Differential expression of three mem-bers of the AMT1 gene family encoding putative high–affinity NH4+ transporters in roots of Oryza sa-tiva subspecies indica. Plant Cell Environ 26: 907–914

Kühn C, Hajirezaei MR, Fernie AR, Roessner-Tunali U, Czechowski T, Hirner B, Frommer WB (2003 ) The sucrose transporter StSUT1 localizes to sieve elements in potato tuber phloem and influ-ences tuber physiology and development. Plant Physiol 131: 102–13

Kühn C, Quick WP, Schulz A, Riesmeier JW, Sonnewald U, Frommer WB (1996) Companion cell-specific inhibition of the potato sucrose transporter SUT1. Plant Cell Environ 19: 1115–1123 Lam E, Chua NH (1989) ASF-2: A factor that binds to the cauliflower mosaic virus 35S promoter and a

conserved GATA motif in cab promoters. Plant Cell 1:1147–1156

Lao NT, Schoneveld O, Mould RM, Hibberd JM, Gray JC, Kavanagh TA (1999) An Arabidopsis gene encoding a chloroplast–targeted beta–amylase. Plant J 20: 519–527

Lazan H, Ng SY, Goh LY, Ali ZM (2004) Papaya beta–galactosidase/ galactanase isoforms in differen-tial cell wall hydrolysis and fruit softening during ripening. Plant Physiol Biochem 42: 847–853 Lee SK, Hwang SK, Han M, Eom JS, Kang HG, Han Y, Choi SB, Cho MH, Bhoo SH, An G, Hahn

TR, Okita TW, Jeon JS (2007) Identification of the ADP-glucose pyrophosphorylase isoforms essen-tial for starch synthesis in the leaf and seed endosperm of rice (Oryza sativa L.) Plant Mol Biol 65:

531–546

Lemoine R (2000) Sucrose transporters in plants: update on function and structure. Biochim Biophys Acta 1465: 246–262

Loescher WH, Marlow GG, Kennedy RA (1982) Sorbitol metabolism and sink-source interconversions in developing apple leaves. Plant Physiol 70: 335–339

Lu YK, Stemler AJ (2002) Extrinsic photosystem II carbonic anhydrase in maize mesophyll chloroplasts.

Plant Physiol 266: 16746–16754

Mae T, Ohira K (1981) The remobilization of nitrogen related to leaf growth and senescence in rice plants (Oryza sativa L.). Plant Cell Physiol 22: 1067–1074

Matsukura C, Saitoh T, Hirose T, Ohsugi R, Perata P, Yamaguchi J (2000) Sugar uptake and trans-port in rice embryo: expression of companion cell–specific sucrose transtrans-porter (OsSUT1) induced by sugar and light. Plant Physiol 124: 85–93

Matsuzaki A, Rutger JN (1991) Characteristics relating young panicle development and heading of closely related calrose rice genotypes. Jpn J Crop Sci 60: 298–305

Meyer S, Truernit E, Hu¨mmer C, Besenbeck R, Stadler R, Sauer N (2000) AtSUC3, a gene encod-ing a new Arabidopsis sucrose transporter, is expressed in cells adjacent to the vascular tissue and in a carpel cell layer. Plant J 24: 869–882

Miyazawa Y, Sakai A, Miyagishima S, Takano H, Kawano S, Kuroiwa T (1999) Auxin and cytokinin have opposite effects on amyloplast development and the expression of starch synthesis genes in cul-tured bright yellow–2 tobacco cells. Plant Physiol 121: 461–469

Muñoz FJ, Baroja–Fernandez E, Moran–Zorzano MT, Viale AM, Etxeberria E, Alonso–Casajus N, Pozueta–Romero J (2005) Sucrose synthase controls both intracellular ADP glucose levels and tran-sitory starch biosynthesis in source leaves. Plant Cell Physiol 46: 1366–1376

Nakamura M, Tsunoda T, Obokata J (2002) Photosynthesis nuclear genes generally lack TATA-boxes:

a tobacco photosystem I gene responds to light through an initiator. Plant J 29:1–10

Nakamura Y (1996) Some properties of starch debranching enzymes and their possible roles in amylo-pectin biosynthesis. Plant Sci 121: 1–18

Nakamura Y (2002) Towards a better understanding of the metabolic system for amylopectin biosynthe-sis in plants: rice endosperm as a model tissue. Plant Cell Physiol 43: 718–725

Nakamura Y, Yuki K, Park S, Ohya T (1989) Carbohydrate metabolism in the developing endosperm of rice grains. Plant cell Physiol 30: 833–839

Nakata PA, Okita TW (1995) Diferential regulation of ADP-glucose pyrophosphorylase in the sink and source tissues of potato. Plant Physiol 108: 361–368

Ngampanya B, Takeda T, Sonoda Y, Narangajavana J, Yamaguchi J (2002) Characterization of OsSUT2 cDNA expressed before flowering stage. Rice Genet News 19: 49–51

Nishi A, Nakamura Y, Tanaka N, Satoh H (2001) Biochemical and genetic effects of amy-lose–extender mutation in rice endosperm. Plant Physiol 127: 459–472

Nishiuchi T, Shishi H, Suzuki K (2004) Rapid and transient activation of transcription of the ERF3 gene by wounding in tobacco leaves: possible involvement of NtWRKYs and autorepression. J Biol Chem 279: 55355–55361

Nittylä T, Messerli G, Trevisan M, Chen J, Smith AM, Zeeman SC (2004) A novel maltose transporter is essential for starch degradation in leaves. Science 303: 87–89

Ohdan T, Francisco PB, Sawada T, Hirose T, Terao T, Satoh H, Nakamura Y (2005) Expression pro-filing of genes involved in starch synthesis in sink and source organs of rice. J Exp Bot 56: 3229–3244 Perez CM, Palmiano EP, Baun LC, Juliano BO (1971) Starch metabolism in the leaf sheaths and culm

of rice. Plant Physiol 47: 404–408

Rook F, Corke F, Card R, Munz G, Smith C, Bevan MW (2001) Impaired sucrose–induction mutants reveal the modulation of sugar–induced starch biosynthetic gene expression by abscisic acid signaling.

Plant J 26: 421–433

Ross EJH, Stone JM, Elowsky CG, Arredondo-Peter R, Klucas RV, Sarath G (2004) Activation of the Oryza sativa non-symbiotic haemoglobin-2 promoter by the cytokinin-regulated transcription fac-tor, ARR1. J Exp Bot 55:1721–1731

Salehuzzaman SNIM, Jacobsen E, Visser RGF (1994) Expression patterns of two starch biosynthetic genes in vitro cultured cassava plants and their induction by sugars. Plant Sci 98: 53–62

Samonte SOPB, Wilson LT, McClung AM, Tarpley L (2001) Seasonal dynamics of nonstructural car-bohydrate partitioning in fifteen diverse rice (Oryza sativa L.) genotypes. Crop Sci 41: 902–909 Sanford JC, Smith FD, Russell JA (1993) Optimizing the biolistic process for different biological

ap-plications. Meth Enzymol 217: 483–509

Sauer N (2007) Molecular physiology of higher plant sucrose transporters. FEBS Lett 581: 2309–2317 Scheible WR, González-Fontes A, Morcuende R, Lauerer M, Geiger M, Glaab J, Gojon A, Schulze

ED, Stitt M (1997) Tobacco mutants with a decreased number of functional nia genes compensates by modification the diurnal regulation of transcription, post-translational modification and turnover of ni-trate reductase. Planta 203: 304–319

Schneidereit A, Imlau A, Sauer N (2008) Conserved cis-regulatory elements for

DNA-binding-with-one-finger and homeo-domain-leucine-zipper transcription factors regulate com-panion cell-specific expression of the Arabidopsis thaliana SUCROSE TRANSPORTER 2 gene. Planta 228: 651–662

Scofield GN, Hirose T, Gaudron JA, Upadhyaya NM, Ohsugi R, Furbank RT (2002) Antisense sup-pression of the rice sucrose transporter gene, OsSUT1, leads to impaired grain filling and germination but does not affect photosynthesis. Funct Plant Biol 29: 815–826

Scofield GN, Aoki N, Hirose T, Takano M, Jenkins CL, Furbank RT (2007) The role of the sucrose transporter, OsSUT1, in germination and early seedling growth and development of rice plants. J Exp Bot 58: 483–495

Sheu JJ, Yu TS, Tong WF, Yu SM (1996) Carbohydrate starvation stimulates differential expression of rice a-amylase genes that is modulated through complicated transcriptional and post-transcriptional processes. J Biol Chem 271: 26998–27004

Sikka VK, Choi S, Kavakli IH, Sakulsingharoj C, Gupta S, Ito H, Okita TW (2001) Subcellular compartmentation and allosteric regulation of the rice endosperm ADP–glucose pyrophosphorylase.

Plant Sci 161: 461–468

Sivitz, AB, Reinders, A and Ward, JM (2008) Arabidopsis sucrose transporter AtSUC1 is important for pollen germination and sucrose–induced anthocyanin accumulation. Plant Physiol 147: 92–100 Smith AM, Denyer K, Martin C (1997) The synthesis of the starch granule. Annu Rev Plant Physiol

Plant Mol Biol 48: 67–87

Smith AM, Zeeman SC, Smith SM (2005) Starch degradation. Annu Rev Plant Biol 56: 73–98

Smith FW, Mudge SR, Rae AL, Glassop D (2003) Phosphate transport in plants. Plant Soil 248: 71–83 Soga Y, Nozaki M (1957) Studies on the relation between seasonal changes of carbohydrates

accumu-lated and the ripening at the stage of generative growth in the rice plant and the grain ripening. Proc Crop Sci Soc Jpn 26: 105–108

Sokolov LN, Déjardin A, Kleczkowski LA (1998) Sugars and light/dark exposure trigger differential regulation of ADP–glucose pyrophosphorylase genes in Arabidopsis thaliana (thale cress). Biochem J 336: 681–687

Sokolova SV,Balakshina NO,Krasavina MS (2002) Activation of Soluble Acid Invertase Accompa-nies the Cytokinin-Induced Source–Sink Leaf Transition. Russ J Plant Physiol 49:86-91

Sonoda Y, Ikeda A, Saiki S, Yamaya T, Yamaguchi J (2003) Feedback regulation of the ammonium transporter gene family AMT1 by glutamine in rice. Plant Cell Physiol 44: 1396–1402

Stead JA, Keen JN, McDowall KJ (2006) The identification of nucleic acid-interacting proteins using a simple proteomics-based approach that directly incorporates the electrophoretic mobility shift assay.

Mol Cell Proteomics 5: 1697–1702

Suenaga A, Moriya K, Sonoda Y, Ikeda A, Von Wiren N, Hayakawa T, Yamaguchi J, Yamaya T (2003) Constitutive expression of a novel–type ammonium transporter OsAMT2 in rice plants. Plant Cell Physiol 44: 206–211

Sun CX, Palmqvist S, Olsson H, Boren M, Ahlandsberg S, Jansson C (2003) A novel WRKY tran-scription factor, SUSIBA2, participates in sugar signaling in barley by binding to the sugarresponsive elements of the iso1 promoter. Plant Cell 15: 2076–2092

Takahashi S, Ishimaru K, Yazaki J, Fujii F, Shimbo K, Yamamoto K, Sakata K, Sasaki T, Kishimo-to N, Kikuchi S (2005) Microarray analysis of sink–source transition in rice leaf sheaths. Breed Sci 55:153–162

Takashi A, Mizuno K, Fujimura T (2005) Gene expression of ADP-glucose pyrophosphorylase and starch contents in rice cultured cells are cooperatively regulated by sucrose and ABA. Plant Cell Phy-siol 46: 937–946

Takeda T, Toyofuku K, Matsukura C, Yamaguchi J (2001) Sugar transporters involved in flowering and grain development of rice. J Plant Physiol 158: 465–470

Tanaka A (1961) Studies on the nutrio–physiology of leaves of rice. plant. J Fac Agric Hokkaido Univ 51:

449–550

Thomas BR, Rodriguez RL (1994) Metabolite signals regulate gene expression and source/sink rela-tions in cereal seedlings. Plant Physiol 106: 1235–1239

Thorsteinsson B, Tillberg E, Ericsson T (1990) Levels of IAA, ABA and carbohydrates in source and sink leaves of Betula pendula. Roth Scand J For Res 5:347–354

Toki S (1997) Rapid and efficient Agrobacterium-Mediated transformation in rice. Plant Mol Biol Rep 15:

16-21

Turgeon R (1989) The sink-source transition in leaves. Annu Rev Plant Physiol Plant Mol Biol 40:

119–38

Turgeon R, Webb JA (1975) Leaf development and phloem transport in Cucurbita pepo: carbon econo-my. Planta 123: 53–62

Urao T, Yamaguchi–Shinozaki K, Urao S, Shinozaki K (1993) An Arabidopsis myb homolog is

在文檔中水稻葉鞘在抽穗期間由儲存組織轉換成供源組織之分子調控機制 (頁 71-134)