臺灣博碩士論文加值系統

OsMADS45是屬於MADS box中的E群基因，參與水稻稻穗發育過程中的調控。雖然Jeon等人於2000年的研究顯示，水稻中大量表現OsMADS45會導致其轉殖株提早開花並導致殖株矮化，但影響此現象的分子機制仍然不明瞭。因此我們構築以Ubi啟動子持續表現的OsMADS45，並轉殖到對光週期較不敏感的台農67號水稻品系，進而探討大量表現OsMADS45所造成的分子機轉。大量表現OsMADS45的台農67號水稻轉殖株，其開花時間明顯較野生型提早，所提早的開花時間是為Jeon轉殖株的8倍(提早40天抽穗)，此外轉殖株外觀呈現矮化並有較低的穀粒產量和多數的空穗。大量表現OsMADS45並不會影響開花調控因子的日夜週期韻律，然而在轉殖株中，Heading Date 3a (Hd3a)以及RICE FlOWERING LOCUS T1 (RFT1)兩開花素均提早了20天開始表現。此外大量表現OsMADS45亦會影響Heading Date 1 (Hd1)，促使其在幼苗期表現量降低而於開花後提高表現。再者OsMADS45轉殖株中，OsMADS14與OsMADS18兩基因的表現時間點皆與提早表現的Hd3a與RFT1相符。OsMADS45的大量表現並不會影響Hd1以及Early heading date 1 (Ehd1)上游基因的表現，例如OsGI、Ehd2/Osld1/RID1以及OsMADS50。本實驗結果證實大量表現OsMADS45時，會誘導Hd3a與RFT1基因提早表現，導致轉殖株發育初期即有Hd3a與RFT1的存在，並促使OsMADS14與OsMADS18提高表現，進而造成轉殖株提早進入生殖生長週期，導致提早開花。

The rice gene, OsMADS45, which belongs to the MADS-box E class gene, participates in the regulation of floral development. Previous studies have revealed that ectopic expression of OsMADS45 induces early flowering and reduces plant height under short-day (SD) conditions. However, the regulation mechanism of OsMADS45 overexpression remains unknown. We introduce an OsMADS45 overexpression construct Ubi:OsMADS45 into TNG67 plants (an Hd1 (Heading date 1) and Ehd1 (Early heading date 1) defective rice cultivar grown in Taiwan), and we analyzed the expression patterns of various floral regulators to understand the regulation pathways affected by OsMADS45 expression. The transgenic rice exhibit a heading date approximately 40 days earlier than that observed in TNG67 plants, and transgenic rice display small plant size and low grain yield. OsMADS45 overexpression did not alter the oscillating rhythm of the examined floral regulatory genes, Hd3a (Heading Date 3a) and RFT1 (RICE FLOWERING LOCUS T1), but advanced (by approximately 20 days) the up-regulation of the two florigens and suppressed the expression of Hd1 at the juvenile stage. The expression levels of OsMADS14 and OsMADS18, which are two well-known reproductive phase transition markers, were also increased at early developmental stages and are believed to be the major regulators responsible for early flowering in OsMADS45-overexpressing transgenic rice. OsMADS45 overexpression did not influence other floral regulator genes upstream of Hd1 and Ehd1, such as OsGI (OsGIGANTEA), Ehd2/Osld1/RID1 and OsMADS50. These results indicate that in transgenic rice, OsMADS45 overexpressing ectopically activates the upstream genes Hd3a and RFT1 at early developmental stage and up-regulates the expression of OsMADS14 and OsMADS18, which promote early flowering.

中文摘要 VII
ABSTRACT VIII
1. INTRODUCTION 1
2. LITERATURE REVIEW 3
Flowering regulation in rice 3
Rice floral development genes 4
The function and interaction of OsMADS45 6
3. MATERIALS AND METHODS 8
Plant material and growth conditions 8
Ubi:OsMADS45 transgenic rice 8
Analyses of agronomic traits 9
Measurement of photosynthesis rate 9
Genomic DNA extraction and gene sequencing 9
Southern blot analysis 11
RNA isolation and semi-quantitative RT-PCR 12
4. RESULTS 14
Ectopic expression of OsMADS45 causes early flowering in TNG67 rice variety 14
Ectopic expression of OsMADS45 does not alter the oscillation rhythm of flowering time regulators 15
Expression analysis of various floral regulatory genes in 45OX transgenic rice 16
The expression of RFT1 is activated at early developmental stages in 45OX transgenic rice 16
The up-regulation of Hd3a and down-regulation of Hd1 at early development stages were observed in 45OX transgenic rice 17
The expression of OsMADS14 and OsMADS18 is up-regulated in 45OX transgenic rice 18
Genes Hd1 and Ehd1 are defective and an alternative spliced Hd1 mRNA was identified in TNG67 rice 20
5. DISCUSSION 22
6. LITERATURE CITED 29
7. TABLES AND FIGURES 37
8. SUPPLEMETARY DATA 44

APPENDIX: Analyses of PEPC:daao T-DNA insertion mutant. 49
1. INTRODUCTION 49
2. LITERATURE REVIEW 50
3. MATERIALS AND METHODS 53
Growth conditions 53
Southern blot analysis 53
Inverse PCR (iPCR) 53
The breeding of TK9/PEPC:daao hybrid Rice 54
The construction of Os02g0743200 antisense vector and Agrobacteria mediated gene transformation 54
4. RESULTS 55
The Os02g0743200 gene was the T-DNA inserted site in PEPC:daao transgenic rice 55
Os02g0743200 is similar to Sorghum bicolor SORBIDRAFT_04g028190 55
The uncorrelated phenotypes of PEPC:daao/TK9 hybridized rice 56
Construction of Os02g0743200 antisense plasmid 56
5. DISCUSSION 58
6. FUTURE WORKS 61
7. LITERATURE CITED 62
8. TABLES AND FIGURES 65
9. SUPPLEMENTAL DATA OF APPENDIX 76

 
Contents of tables

Table 1. Agronomic traits of TNG67 and Ubi:OsMADS45 transgenic rice plants. 37


APPENDIX

Table 1. The primers used in this test. 65
Table 2. The genotypes and agronomic traits of the F2 generation of PEPC:daao/TK9 hybridized rice 66

 
Contents of figures

Figure 1. The genome type and genetic expression of Ubi:OsMADS45 T3 transgenic rice. 38
Figure 2. The phenotypes of Ubi:OsMADS45 transgenic lines.. 39
Figure 3. The diurnal expression patterns of the floral regulatory genes of the Ehd2-OsMADS50-Ehd1-RFT1 and OsGI-Hd1-Hd3a flowering pathways. 40
Figure 4. Expression analysis of the floral regulatory genes Ehd2, OsMADS50, Ehd1, RFT1, OsGI, Hd1, Hd3a, OsMADS14 and OsMADS18 in TNG67 and Ubi:OsMADS45 transgenic rice at various growth stages 41
Figure 5. A schematic diagram showing the sequence variations and cDNA of Hd1 of TNG67 rice compare with Nipponbare plants 42
Figure 6. A model of the early flowering signal pathway in 45OX transgenic rice 43


APPENDIX

Figure 1. Southern blot of PEPC:daao transgenic rice 67
Figure 2. The amplification of flanking inserted sequence of PEPC:daao by iPCR analysis. 68
Figure 3. Os02g0743200 gene was inserted in PEPC:daao transgenic rice. 69
Figure 4. The RT-PCR analysis of TK9 and PEPC:daao inserted rice 70
Figure 5. The gene location and alignment of Os02g0743200 71
Figure 6. The genotypes (A) and phenotypes (B) of F2 generation of PEPC:daao/TK9 hybridized rice. 72
Figure 7. The construction of Os02g0743200 antisense plasmid 73
Figure 8. Calli and plants of antisense Os02g0743200 resulting from Agrobacterium mediated transformation 74
Figure 9. The comparison of TK9 (left) and Os02g0743200 overexpresssing (right) seeds. 75

 
Contents of supplemental data

Supplemental Table 1. The primer pairs used in this study. 44
Supplemental Figure 1. The flowering pathway. 48
Supplemental Figure 2. The transformation vector of the Ubi:OsMADS45 transgenic rice 46
Supplemental Figure 3. The genomic DNA sequence of the Hd1 gene of TNG67 rice 47
Supplemental Figure 4. A schematic diagram showing the location of sequence variation and the detailed cDNA sequences of Ehd1 of TNG67 rice 48



APPENDIX.

Supplemental Table 1. The agronomic traits of T2 transgenic rice 76
Supplemental Table 2. The material and programs of iPCR 77
Supplemental Table 3. Media used in this test. 79
Supplemental Figure 1. Morphological characteristics of TK9 rice (left) and PEPC:daao transgenic plant (right). 84
Supplemental Figure 2. The strategy of antisense gene construction. 85
Supplemental Figure 3. RT-PCR analysis of Os02g0743200 antisense (an-1, 2, 3 and 4) and overexpression (ov-2) lines 86
Supplemental Figure 4. The pollen viability test of TK9 and Os02g0743200 antisense lines (an-1, 2 and 3) 87

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