黃瓜重要農(nóng)藝性狀的分子生物學(xué)

張圣平，顧興芳

·導(dǎo)讀·

黃瓜重要農(nóng)藝性狀的分子生物學(xué)

張圣平，顧興芳

（中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所/農(nóng)業(yè)農(nóng)村部園藝作物生物學(xué)與種質(zhì)創(chuàng)制重點(diǎn)實(shí)驗(yàn)室，北京 100081）

黃瓜是我國(guó)的重要蔬菜作物，尤其在設(shè)施生產(chǎn)中占有舉足輕重的地位，2017年收獲面積為123.52萬(wàn)hm2，產(chǎn)量達(dá)到6 482.46萬(wàn)t[1]。豐產(chǎn)優(yōu)質(zhì)多抗新品種是黃瓜綠色高效生產(chǎn)的重要保障，而優(yōu)良新品種的選育離不開(kāi)準(zhǔn)確高效的育種技術(shù)。重要農(nóng)藝性狀分子生物學(xué)研究，如遺傳圖譜構(gòu)建、分子標(biāo)記開(kāi)發(fā)、優(yōu)異基因挖掘、功能基因組研究等，是高通量高效率分子設(shè)計(jì)育種技術(shù)研發(fā)的基礎(chǔ)。由于黃瓜遺傳背景狹窄，在全基因組序列信息破譯之前，相關(guān)分子生物學(xué)研究進(jìn)展緩慢。

2009年，黃瓜全基因組測(cè)序完成，數(shù)據(jù)量達(dá)到17.2 G，測(cè)序深度72.2×，注釋了26 682個(gè)基因[2]，使黃瓜成為第一個(gè)以二代測(cè)序技術(shù)為主完成測(cè)序的植物，也是世界上第一個(gè)完成測(cè)序的蔬菜作物，開(kāi)啟了黃瓜分子生物學(xué)和遺傳育種快速發(fā)展的新時(shí)期。10年來(lái)，基于黃瓜基因組測(cè)序的帶動(dòng)和功能基因組學(xué)的迅速發(fā)展，在黃瓜研究上凝聚了一批研究人員、謀劃了一批科研項(xiàng)目、取得了一批研究成果，特別是在高密度遺傳圖譜構(gòu)建、高質(zhì)量分子標(biāo)記開(kāi)發(fā)、重要農(nóng)藝性狀基因挖掘、基因調(diào)控網(wǎng)絡(luò)解析等方面取得了較大進(jìn)展，發(fā)表了影響因子大于6的SCI論文25篇，其中影響因子大于8的高水平論文12篇，申請(qǐng)國(guó)內(nèi)外專(zhuān)利超過(guò)200件，創(chuàng)建了高效的分子標(biāo)記聚合育種技術(shù)，實(shí)現(xiàn)了育種技術(shù)的更新?lián)Q代，國(guó)內(nèi)2項(xiàng)成果“黃瓜基因組和重要農(nóng)藝性狀基因研究”和“黃瓜優(yōu)質(zhì)多抗種質(zhì)資源創(chuàng)制與新品種選育”榮獲國(guó)家級(jí)獎(jiǎng)勵(lì)。

在我國(guó)完成華北密刺型黃瓜9930的全基因組測(cè)序之后，美國(guó)完成了加工型黃瓜GY14的全基因組測(cè)序[3]。最近中國(guó)農(nóng)業(yè)科學(xué)院蔬菜花卉研究所的科研人員進(jìn)一步組裝完成了9930黃瓜V3.0參考基因組，利用10×genomics技術(shù)平臺(tái)和Hi-C測(cè)序數(shù)據(jù)，組裝獲得了總長(zhǎng)度226.2 Mb，共包含174 contigs的參考基因組，包含1 374個(gè)全長(zhǎng)末端逆轉(zhuǎn)錄轉(zhuǎn)座子和1 078個(gè)新基因，可以更好的服務(wù)于黃瓜遺傳研究[4]。以9930基因組為參考，通過(guò)對(duì)115份黃瓜核心種質(zhì)進(jìn)行深度重測(cè)序，構(gòu)建了包含360多萬(wàn)個(gè)位點(diǎn)的全基因組遺傳變異圖譜，為全面了解黃瓜進(jìn)化及多樣性提供了新思路，并為全基因組設(shè)計(jì)育種奠定了良好基礎(chǔ)[5]。

黃瓜高密度遺傳圖譜構(gòu)建方面，第一張整合的高密度遺傳圖譜于2012年完成，使得定位在圖譜上的標(biāo)記位點(diǎn)達(dá)到1 369個(gè)[6]。隨后利用永久群體重組自交系（RILs）結(jié)合基因組測(cè)序，構(gòu)建了包含1 681個(gè)標(biāo)記的分子連鎖圖譜[7]。為重要農(nóng)藝性狀基因定位提供了有力的工具。

黃瓜抗病性狀分子生物學(xué)研究方面，將抗黑星病基因精細(xì)定位在Chr.2上0.29 cM的區(qū)域內(nèi)，獲得6個(gè)候選基因[8]；將抗西葫蘆黃化花葉病毒基因精細(xì)定位于Chr.6上50 kb的區(qū)段內(nèi)[9]；將枯萎病抗病主效QTL定位在Chr.2上2.4 cM內(nèi)[10]；檢測(cè)到PI197088黃瓜的11個(gè)與霜霉病抗性相關(guān)的QTL，4個(gè)與白粉病抗性相關(guān)的QTL[11]；檢測(cè)到TH118FLM黃瓜5個(gè)與霜霉病抗性相關(guān)的QTL[12]；檢測(cè)到PI183967黃瓜5個(gè)與莖部蔓枯病抗性相關(guān)的QTL[13]；在Chr.5上檢測(cè)到抗角斑病QTL位點(diǎn)psl5.1和psl5.2[14]。圖位克隆了GY14黃瓜抗炭疽病候選基因，該基因?qū)儆诩易宓闹参锟共』騕15]。黃瓜的感病丟失突變帶來(lái)了永久廣譜抗病性，為黃瓜抗病育種起到了重要作用，支撐了美國(guó)黃瓜產(chǎn)業(yè)的發(fā)展[16]。

黃瓜果實(shí)品質(zhì)性狀分子生物學(xué)研究方面，圖位克隆了短果基因[17]、小葉基因[18]、圓形葉片基因[19]、軟刺基因[20]、果瘤基因[21]、超高密度果刺新位點(diǎn)[22]，其中泛素化修飾并降解自身及其底物，進(jìn)而影響乙烯含量變化；編碼WD40重復(fù)蛋白并且與黃瓜器官大小的發(fā)育密切相關(guān)；編碼絲氨酸/蘇氨酸蛋白激酶。檢測(cè)到了8個(gè)與果實(shí)大小相關(guān)的QTL[23]；可以增強(qiáng)果實(shí)中乙烯的含量，進(jìn)而導(dǎo)致果實(shí)發(fā)生彎曲[24]；、和三個(gè)基因通過(guò)相互作用共同調(diào)節(jié)葉綠素含量的積累，從而影響嫩瓜果皮顏色[25]；將多效應(yīng)黑刺基因精細(xì)定位于Chr.4上50 kb的區(qū)段內(nèi)，獲得候選基因轉(zhuǎn)錄因子[26]；將心皮數(shù)基因精細(xì)定位在Chr.1上[27]，將果實(shí)多刺基因精細(xì)定位在Chr.2上[28]，將黃綠葉色基因精細(xì)定位在Chr.6上50.9 kb內(nèi)[29]。

黃瓜植株形態(tài)建成相關(guān)性狀分子生物學(xué)研究方面，解析了分枝、花打頂?shù)戎晷驼{(diào)控機(jī)制，是控制分枝的關(guān)鍵基因，通過(guò)抑制生長(zhǎng)素運(yùn)輸基因，導(dǎo)致側(cè)枝中生長(zhǎng)素過(guò)量積累，從而抑制側(cè)枝的生長(zhǎng)發(fā)育[30]?？寺×丝刂朴邢奚L(zhǎng)的關(guān)鍵基因，該基因通過(guò)與互作來(lái)影響花發(fā)育[31]。在性別表達(dá)和開(kāi)花時(shí)間研究上，將雄性不育基因精細(xì)定位到了76 kb區(qū)間，獲得候選基因[32]；檢測(cè)到3個(gè)調(diào)控開(kāi)花時(shí)間的QTL，其中FT6.2是主效QTL[33]。

基因調(diào)控網(wǎng)絡(luò)解析方面，明確了黃瓜苦味代謝和進(jìn)化源于兩個(gè)主轉(zhuǎn)錄因子直接調(diào)控的一個(gè)9基因模塊[34]；復(fù)合體調(diào)控果刺的其實(shí)發(fā)育[35]；復(fù)合體調(diào)控果瘤的形成[36]；在轉(zhuǎn)錄因子HD-ZIPIII和CsWUS 之間具有調(diào)配器的功能，從而調(diào)控花粉和胚珠的發(fā)育[38]；鑒定到一個(gè)控制果實(shí)長(zhǎng)度的關(guān)鍵基因，該基因通過(guò)抑制生長(zhǎng)素運(yùn)輸基因和的表達(dá)進(jìn)而減少生長(zhǎng)素積累[38]。

本專(zhuān)題所匯集的5篇論文，分別在黃瓜植株形態(tài)建成、雌花及果實(shí)發(fā)育、抗白粉病機(jī)制等方面取得了新進(jìn)展。在黃瓜Chr.1、Chr.2、Chr.3、Chr.4、Chr.5、Chr.6上，檢測(cè)到8個(gè)與下胚軸長(zhǎng)度密切關(guān)聯(lián)的SNP位點(diǎn)，挖掘到8個(gè)調(diào)控下胚軸長(zhǎng)度的候選基因[39]。從9930基因組中鑒定得到138個(gè)ERF基因家族成員，其中部分成員在不同性型材料中差異表達(dá)，可能參與雌花分化初期的基因表達(dá)調(diào)控[40]。克隆得到果實(shí)發(fā)育調(diào)控網(wǎng)絡(luò)中，參與胎座框的形成的重要基因，擬南芥異源過(guò)表達(dá)轉(zhuǎn)基因植株的果莢變短，花粉育性降低，且種子發(fā)育受到抑制[41]。分別在Chr.1、Chr.3、Chr.6上檢測(cè)到4個(gè)單性結(jié)實(shí)QTL，挖掘到4個(gè)與單性結(jié)實(shí)性狀相關(guān)的候選基因[42]。篩選獲得了抗白粉病新材料，在成熟期葉片的防御信號(hào)途徑相關(guān)基因表達(dá)上，抗病材料的表達(dá)高于感病材料[43]。這些僅僅是黃瓜重要農(nóng)藝性狀分子生物學(xué)研究領(lǐng)域中的一小部分內(nèi)容，謹(jǐn)希望以專(zhuān)題的形式呈現(xiàn)，促進(jìn)該學(xué)科領(lǐng)域的快速發(fā)展。

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Molecular Biology of Important Agronomic Traits in Cucumber

ZHANG ShengPing, GU XingFang

(Institute of Vegetables and Flowers, Chinese Academy of Agricultural Sciences/Key Laboratory of Horticultural Crop Biology and Germplasm Creation, Ministry of Agriculture and Rural Areas, Beijing 100081)

2019-12-14；

2019-12-30

中國(guó)農(nóng)業(yè)科學(xué)院創(chuàng)新工程（CAAS-ASTIP-2017-IVF）

張圣平，E-mail：zhangshengping@caas.cn。顧興芳，E-mail：guxingfang@caas.cn

10.3864/j.issn.0578-1752.2020.01.011

（責(zé)任編輯 趙伶俐）