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莲的分子生物学研究进展

摘要:

莲(Nelumbo)作为真双子叶基部类群植物,对植物系统进化研究具有重要意义,同时,作为重要的水生园艺植物,还具有观赏、食用及药用等经济价值。近年来,随着莲基因组测序、组装及注释工作的不断完善,莲分子生物学研究得到了极大的发展。本文综述了莲分子生物学研究的最新进展,主要包括3个方面:(1)莲基因组研究的最新进展及其意义;(2)莲各个园艺性状及不同生理过程的遗传机理及多组学研究状况;(3)调控莲重要园艺性状关键基因的挖掘及其功能验证的相关进展。最后,对目前莲的相关研究所面临的问题及今后的发展趋势进行了展望。

关键词: 莲  /  基因组  /  组学  /  功能研究  /  分子生物学  

Abstract:

Lotus (Nelumbo), as a basal eudicotyledon, is of great significance for studying the evolution and phylogeny of plants. It is also an economically important aquatic horticultural plant with ornamental, edible, and medicinal value. With the completion of genome sequencing, assembly, and annotation in the last decade, molecular level studies have greatly driven and expanded our understanding of all biological aspects of this species. In this review, we systematically discuss the latest advances in molecular biology studies of the lotus, with a focus on: (1) Recent progress in genomic research; (2) Genetic and multiomics studies of different horticultural traits and physiological processes; and (3) Characterization of key genes controlling important horticultural traits and their functions. Based on these, we also discuss current research challenges and future prospects.

图  1   莲地下茎不同发育时期

S1:莲地下茎膨大早期;S2: 莲地下茎膨大中期;S3:莲地下茎膨大后期。

Figure  1.   Different developmental stages of lotus rhizome

S1: Early stage of lotus rhizome enlargement; S2: Middle stage of lotus rhizome enlargement; S3: Late stage of lotus rhizome enlargement.

图  2   莲花瓣的花青素生物合成途径

PAL:苯丙氨酸解氨酶;C4H:肉桂酸-4-羟化酶;4CL:4-香豆酸辅酶A连接酶;CHS:查耳酮合酶;CHI:查尔酮异构酶;F3H:黄烷酮3-羟化酶;F3'H:黄酮类3'-羟化酶;F3'5'H:类黄酮3',5'-羟化酶;DFR:二氢黄酮醇4-还原酶;ANS:花色素合成酶;UFGT:花青素3-O-葡萄糖基转移酶;FLS:黄酮醇合成酶;OMT:氧甲基转移酶;RT:鼠李糖基转移酶。红色框为粉红色、红色莲、嵌色莲(红色部分)品种涉及的途径;黄色框为白色、黄色、嵌色莲(白色部分)品种涉及的途径。方框中颜色为对应的化合物颜色。

Figure  2.   Anthocyanin biosynthesis pathway in lotus petals

PAL: Phenylalanine ammonia lyase; C4H: Cinnamate 4-hydroxylase; 4CL: 4-Coumaroyl CoA ligase; CHS: Chalcone synthase; CHI: Chalcone isomerase; F3H: Flavanone 3-hydroxylase; F3'H: Flavonoid 3'-hydroxylase; F3'5'H: Flavonoid 3'5'-hydroxylase; DFR: Dihydroflavonol reductase; ANS: Anthocyanidin synthase; UFGT: Anthocyanin 3-O-glucosyltransferase; FLS: Flavonol synthase; OMT: O-Methyltransferase; RT: Rhamnosyltransferase. Red frame shows anthocyanin biosynthesis pathway involved in cultivars of pink, red, and pied lotus (red part); yellow frame shows anthocyanin biosynthesis pathway involved in cultivars of white, yellow, and pied lotus (white part). Colors in boxes are corresponding chemical compound colors.

图  3   莲花发育的ABCE模型

a:萼片;b:正常花瓣;c:瓣化雄蕊,形态趋近正常花瓣;d:瓣化雄蕊,形态趋近正常雄蕊;e:正常雄蕊;f: 瓣化心皮;g: 正常心皮。

Figure  3.   ABCE model of floral development in lotus

a: Sepal; b: Petal; c: Petaloid stamen, shape approximates normal petal; d: Petaloid stamen, shape approximates normal stamen; e: Stamen; f: Petaloid carpel; g: Carpel.

表  1   莲基因组信息

Table  1   Genomic information of Nelumbo

莲品种
Lotus variety基因组大小
Genome size / MbGC含量
GC content / %测序深度
Genome coverage组装水平
Assembly level参考文献
Reference 亚洲莲
Nelumbo nucifera Gaertn.‘中国古代莲’
804.6539.50100 × Scaffold[2]797.6839.16100 × 染色体[4]813.238.97129.2 × 染色体[6]‘太子莲’
79239.0090 × Scaffold[5]782.7639.1290 × 染色体[4]80738.81−染色体[8]美洲黄莲
Nelumbo lutea Pers.美洲黄莲843−79 × 染色体[9]

表  2   莲地下茎膨大过程中涉及的信号通路及其基因或蛋白

Table  2   Signaling pathway genes and proteins involved in rhizome enlargement in lotus

信号通路
Signaling pathway基因或蛋白
Gene or protein参考文献
Reference 光周期
PHYB, GI, COL, FT, PHYA, FYPP, HY5, EFL4, FRIGIDA, COP9[21, 22]激素
ABF, PP2C, PYL, PYR, snRK2, AUX/IAA, ARF, GH3, SAUR, EIN, EBF, ERF, CRE, AHP, ARR, JAR1,
COI1, JAZ, PCNT115, GRF, SKP1, cullin, F-box protein, CUL4, LHB1B2, LOX6[21, 22, 25]钙离子
Calcium-transporting ATPase, CHERP, cbpP, CALM1, CAM, CML, Calreticulin, Calcineurin,
Caltractin, Calnexin, Annexin, CPK[21] [1]

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