| ·常規(guī)信息 最近更新：2024年10月4日 18:07:00 | |||||||||||||
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| 基因(座)名稱 | 顆粒結(jié)合型淀粉合酶; 蠟質(zhì)基因; 糯性基因 granule-bound starch synthase; waxy gene; glutinous endosperm | ||||||||||||
| 基因符號 | wx; OsGBSSI; qGC-6; Wx-mq; Wx-op | ||||||||||||
| 所在染色體 | 6 （已克隆） | ||||||||||||
水稻Wx 基因編碼顆粒淀粉合成酶(granule-bound starch synthase, GBSS)，是控制直鏈淀粉合成的主效基因，直接影響水稻胚乳和花粉中直鏈淀粉的含量(Amylose Content, AC)。非糯性基因(Wx)對糯性基因(wx)表現(xiàn)為不完全顯性，存在較為明顯的劑量效應(yīng)。 抗性淀粉的形成依賴于Waxya(Wxa)等位基因的高表達（Zhou et al. 2018）。 在非糯品種中，Wx 基因分化為Wxa 和Wxb 兩種等位基因，其中，野生稻全為Wxa ，秈稻以Wxa 為主，直鏈淀粉含量較高；粳稻基本為Wxb ，直鏈淀粉含量較低。這表明Wxb 基因是由Wxa 基因進化而來(Sano, 1984; Sano et al.1986)。Wx 基因位于第6 染色體的短臂，早在1990 年就已被克隆。粳型品種Hengfeng 的wx 基因由14 個外顯子和13 個內(nèi)含子組成，編碼由609個氨基酸組成的蛋白(Wang et al., 1990)。來自ARC6622 的等位基因Wxop 和來自Milky Queen 的等位基因Wx-mq 引起水稻胚乳的直鏈淀粉含量降低（Mikami et al., 1999; Sato et al. 2002）。 在灌漿期的籽粒中，Wx 基因有2 種mRNA 轉(zhuǎn)錄產(chǎn)物，長度分別為2.3 kb 和3.3 kb，糯稻品種和高AC 的品種分別只有3.3 kb 和2.3kb 的mRNA，而中等AC 的品種二者兼有之，Wx 蛋白與2.3 kb 的mRNA 的含量存在顯著相關(guān)性。研究發(fā)現(xiàn)，3.3 kb 的mRNA 實際上是Wx 轉(zhuǎn)錄的mRNA 前體，而2.3 kb RNA 為翻譯Wx 蛋白的mRNA，它由3.3 kb 的Wx mRNA 切除第一個1.0 kb 的內(nèi)含子后產(chǎn)生。因此，AC 的調(diào)控屬轉(zhuǎn)錄后調(diào)控，與切除內(nèi)含子的能力有關(guān)(Wang et al., 1995)。序列分析表明，與Wxa 相比，Wxb 的第1內(nèi)含子5'端剪切處發(fā)生由GT→TT的突變，導(dǎo)致第1 內(nèi)含子剪接效率降低及剪接不正常，粳型品種中成熟Wx 轉(zhuǎn)錄本含量降低，β-葡萄糖醛酸酶活性下降，從而引起直鏈淀粉含量下降(Hirano et al., 1998; Isshiki et al., 1998)。序列分析表明，Wxop 基因第1內(nèi)含子5'端的剪切位核苷酸與Wxa 一致(均是GT)，與Wxb 的TT不同，表明Wxop 由Wxa 進化而來，獨立于Wxb （Mikami et al., 1999）。 李家洋、錢前與顧銘洪研究組合作，通過關(guān)聯(lián)分析等手段對淀粉生物合成途徑中的18個基因進行系統(tǒng)研究，發(fā)現(xiàn)這些基因之間的關(guān)系，從而揭示了調(diào)控稻米食用和蒸煮品質(zhì)的精細調(diào)控網(wǎng)絡(luò)：通過影響直鏈淀粉含量(AC)、膠稠度(GC)和糊化溫度(GT)形成的復(fù)雜遺傳網(wǎng)絡(luò)來決定稻米蒸煮與食味品質(zhì)(ECQs)。其中，Wx和SSII-3是2個主要的通過影響AC、GC和GT來決定ECQs的基因，Wx是唯一對AC和GC具有主效作用的基因，而對GT具有微效性；SSII-3是唯一對GT具有主效作用，而對AC和GC具有微效性的基因。在消除主效基因的影響后，分別檢測到2個微效基因同時影響2個品質(zhì)性狀：ISA和SBE3同時對GC和GT有微效作用；也檢測到微效基因?qū)γ總�品質(zhì)性狀的特定作用：SSIII-2、AGPlar、PUL和SSI 4個基因只對AC有微效作用，AGPiso只對GC有微效影響，而SSIV-2只對GT有微效影響。同時本研究還表明，各個不同的淀粉合成相關(guān)基因(SSRGs)在淀粉合成的不同階段對品質(zhì)性狀產(chǎn)生影響。這一結(jié)論得到了遺傳轉(zhuǎn)化實驗的證實。此外，相關(guān)性分析和遺傳轉(zhuǎn)化為水稻育種提供了至關(guān)重要的信息，便于對水稻其它重要農(nóng)藝性狀的調(diào)查研究，也適用于對其它重要農(nóng)作物的研究（Tian et al., 2009）。 Wx 除了影響籽粒淀粉含量，還能夠調(diào)節(jié)膠稠度，圖位克隆證實其就是控制粳/秈雜交膠稠度的主效QTL qGC-6（Su et al. 2011） 用RT-PCR方法成功克隆了Wx-mq 基因，與Wx-b 基因cDNA全長序列相比，Wx-mq 的序列發(fā)生了2個位點的突變：一個是第497位核苷酸由G突變?yōu)锳，另一個是第595位核苷酸由T突變?yōu)镃。這兩個突變導(dǎo)致編碼產(chǎn)物也發(fā)生相應(yīng)變化：第158位氨基酸由精氨酸置換為組氨酸，第191位氨基酸由酪氨酸置換為組氨酸。推測Wx-mq 的這2個錯義突變是造成"Milky Queen"直鏈淀粉含量下降的原因（Sato et al., 2002）。 【W(wǎng)x 座上的微衛(wèi)星序列】Wx 基因序列存在一個(CT)n 微衛(wèi)星片段，在Wx 座位上具有微衛(wèi)星多態(tài)性，并鑒定出8 個不同的Wx 等位基因(Ayres et al., 1997)。舒慶堯等在74 個中國非糯秈稻和粳稻品種中也發(fā)現(xiàn)7 種(CT)n 多態(tài)。 【相關(guān)登錄號】
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| ·ONTOLOGY及相關(guān)基因 | |
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| 表型特征 | 粘性胚乳(TO:0000098), 蒸煮釀制品質(zhì)(TO:0000667), 膠稠度(TO:0000211), 種子生長發(fā)育特征(TO:0000653), 直鏈淀粉含量(TO:0000196), 支鏈淀粉含量(TO:0000097), 抗性淀粉含量(TO:9000696), 直鏈淀粉與支鏈淀粉比(TO:0000372) |
| 分子功能 | 淀粉合酶活性(GO:0009011) |
| 生物進程 | 淀粉生物合成(GO:0019252), 胚乳發(fā)育(GO:0009960) |
| 細胞結(jié)構(gòu) | 質(zhì)體(GO:0009536), 造粉體(GO:0009501), 造粉體淀粉粒(GO:0009568) |
| 形態(tài)構(gòu)造 | 胚乳(PO:0009089) |
| ·參考文獻 |
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