园艺学报 ›› 2025, Vol. 52 ›› Issue (12): 3401-3411.doi: 10.16420/j.issn.0513-353x.2024-1060
齐晓庆, 高澄玥, 李浙浙, 杨秀莲, 王良桂, 岳远征*(
)
收稿日期:2025-06-16
修回日期:2025-10-30
出版日期:2025-12-25
发布日期:2025-12-20
通讯作者:
基金资助:
QI Xiaoqing, GAO Chengyue, LI Zhezhe, YANG Xiulian, WANG Lianggui, YUE Yuanzheng*(
)
Received:2025-06-16
Revised:2025-10-30
Published:2025-12-25
Online:2025-12-20
摘要:
叶色嵌合体独特的斑块或条纹状表型,在观赏植物育种中具有重要价值。因此,研究叶色嵌合体形成的分子调控机制对于定向改良观赏植物的叶色具有重要意义。本文中系统综述了叶色嵌合体的分类、分子调控机制及育种方法,重点从分子机制层面探讨了叶色嵌合体的形成原因以及人工定向培育叶色嵌合体的技术方法。并进一步指出目前该领域在质体发育、层间信号传导及其生存机制等核心问题上仍存在空白,未来需深入探索以指导其稳定创制与应用。
齐晓庆, 高澄玥, 李浙浙, 杨秀莲, 王良桂, 岳远征. 观赏植物叶色嵌合体的研究进展[J]. 园艺学报, 2025, 52(12): 3401-3411.
QI Xiaoqing, GAO Chengyue, LI Zhezhe, YANG Xiulian, WANG Lianggui, YUE Yuanzheng. Research Progress on Leaf Variegation Chimeras of Ornamental Plants[J]. Acta Horticulturae Sinica, 2025, 52(12): 3401-3411.
图2 3种叶色嵌合体茎尖分生组织的细胞层横截面示意图 图中圆环代表茎尖分生组织不同的细胞层,颜色代表细胞层中基因型不同的细胞
Fig. 2 Schematic diagram of cross-section of the cell layer of the apical meristem of three leaf variegation chimeras The rings in the figure represent the different cell layers of the apical meristem,and the colours represent cells with different genotypes in the cell layers
图3 叶色嵌合体的质体分选过程示意图 a:正常的茎尖分生组织细胞;b:L2层的一个细胞中的单个质体(黄色)突变形成“异质性的”细胞;c:当L2层细胞和质体分裂时,逐渐形成只有发育正常或突变质体“同质体”细胞和两者混合的“异质体”细胞;d:L2层具有突变质体的细胞入侵L3层。虚线表示细胞分裂的平面,绿点表示发育正常的质体,黄点表示突变质体
Fig. 3 Schematic diagram of plasmid sorting process for leaf variegation chimeras a:Normal apical meristem cells;b:A single plastid (yellow) in a cell of the L2 layer mutates to form a‘heteroplasmic’cell;c:As the cells and plastids of the L2 layer divide;they gradually form a‘homoplasmic’cell with only normal or mutant plastids and a mixture of both;d:Cells with mutant plastids in the L2 layer invade the L3 layer. The dotted lines indicate the planes of cell division;green dots indicate normal plasmodesmata and yellow dots indicate mutant plasmodesmata
图4 叶色嵌合体中叶绿体发育和质体基因转录调控示意图 野生型的细胞核和质体基因控制叶绿体的良好发育及突变型的细胞核和质体基因使叶绿体发育不良
Fig. 4 Schematic diagram of regulatory map of chloroplast development and plastid gene transcription in leaf variegation chimeras The nucleus and plastid genes in the wild-type control the good development of chloroplasts,and the mutant nuclear and plastid gene cause poor development of chloroplasts
| 序号No. | 名称 Name | 分子机制 Molecular mechanism | 参考文献 Reference |
|---|---|---|---|
| 1 | 蝴蝶兰 Phalaenopsis aphrodite | VAR2单核苷酸插入 VAR2 single nucleotide insertion | Kurniawan et al., |
| 2 | 斑叶君子兰 Clivia miniata | rpoC2的长片段缺失突变 Long fragment deletion mutation in rpoC2 | Wu et al., |
| 3 | 向日葵 Helianthus annuus | psaA点突变 Point mutation in psaA | Azarin et al., |
| 4 | 黄金葛 Epipremnum aureum | EaZIP转录水平差异或早期EaZIP转录物无法有效积累 EaZIP transcript level differences or ineffective accumulation of early EaZIP transcripts | Hung et al., |
| 5 | 尖萼报春苣苔 Primulina pungentisepala | CHL-2上调 CHL-2 upregulation | Chen et al., |
| 6 | 小果榕 Ficus microcarpa | psba、psbb、psbc和psbd均下调;clh1 和clh2 上调;mTERF缺陷 psba,psbb,psbc and psbd are all downregulated;clh1 and clh2 are upregulated;mTERF deficiency | Shih et al., |
| 7 | 日本鸢尾 Iris japonica | GLK下调 GLK downregulation | Yu et al., |
| 8 | 绣球花 Hydrangea macrophylla | HmAP2/ERFs下调 HmAP2/ERFs downregulation | Qi et al., |
| 9 | 虎杖 Reynoutria japonica | rpoC2点突变 rpoC2 point mutation | Park et al., |
| 10 | 垂叶榕 Ficus benjamina | rpoC1短片段缺失 Short fragment deletion in rpoC1 | Park et al., |
| 11 | 粉花绣线菊 Spiraea japonica | rpoB短片段插入 Short fragment insertion in rpoB | Park et al., |
| 12 | 南美榕 Ficus natalensis | rpoB短片段插入 Short fragment insertion in rpoB | Park et al., |
| 13 | 木槿 Hibiscus syriacus | rpoB短片段插入Short fragment insertion in rpoB | Park et al., |
| 14 | 洋常春藤 Hedera helix | psaA短片段缺失 Short fragment deletion in psaA | Park et al., |
| 15 | 合果芋 Syngonium podophyllum | rpoB短片段缺失Short fragment deletion in rpoB | Park et al., |
| 16 | 日本柳杉 Cryptomeria japonica | matK短片段插入突变 Short fragment insertion mutation in matK | Hirao et al., |
| 17 | 大花惠兰 Cymbidium hybrid | ycf3点突变 ycf3 point mutation | Park et al., |
| 18 | 墨兰 Cymbidium sinense | atpI短片段插入 Short fragment insertion in atpI | Park et al., |
| 19 | 长柄玉簪 Hosta longipes | atpE短片段插入 Short fragment insertion in atpE | Park et al., |
| 20 | 吊兰 Chlorophytum comosum | rpoC2短片段缺失 Short fragment deletion in rpoC2 | Park et al., |
| 21 | 富贵竹 Dracaena sanderiana | ycf3内含子点突变 Intronic point mutation in ycf3 | Park et al., |
| 22 | 红苞凤梨 Ananas comosus | VDE、PSY、PDS上调;GS和GDH下调; GLK1、PLA2、FTIP4、AS特异性表达 VDE,PSY,PDS upregulated;GS and GDH downregulated;GLK1,PLA2,FTIP4,AS specifically expressed | Xue et al., Feng et al., |
| 23 | 长果山菅 Dinaella tasmanica | psbA、matK、rps16和trnK的消除 Elimination of psbA,matK,rps16 and trnK | Zhou et al., |
| 24 | 台湾蝴蝶兰 Phalaenopsis aphrodite | PsbO和PsbP下调 PsbO and PsbP downregulated | Tsai et al., |
| 25 | 常春藤叶仙客来 Cyclamen hederifolium | 基因间区域rps4-trnT-UGU的插入/缺失突变 Insertion/Deletion mutation in the intergenic region rps4-trnT-UGU | Mao et al., |
| 26 | 楸树 Catalpa fargesii | PPOX、LHCB1、PsbO、PsbP、PsbQ和PsbR磷酸化差异;CfALKBH5调控m6A甲基化来调控基因表达 Phosphorylation differences in PPOX,LHCB1,PsbO,PsbP,PsbQ,and PsbR;CfALKBH5 regulates gene expression by modulating m6A methylation | Wang et al., |
表1 叶色嵌合体的分子形成机制
Table 1 Molecular mechanism of leaf variegation chimeras
| 序号No. | 名称 Name | 分子机制 Molecular mechanism | 参考文献 Reference |
|---|---|---|---|
| 1 | 蝴蝶兰 Phalaenopsis aphrodite | VAR2单核苷酸插入 VAR2 single nucleotide insertion | Kurniawan et al., |
| 2 | 斑叶君子兰 Clivia miniata | rpoC2的长片段缺失突变 Long fragment deletion mutation in rpoC2 | Wu et al., |
| 3 | 向日葵 Helianthus annuus | psaA点突变 Point mutation in psaA | Azarin et al., |
| 4 | 黄金葛 Epipremnum aureum | EaZIP转录水平差异或早期EaZIP转录物无法有效积累 EaZIP transcript level differences or ineffective accumulation of early EaZIP transcripts | Hung et al., |
| 5 | 尖萼报春苣苔 Primulina pungentisepala | CHL-2上调 CHL-2 upregulation | Chen et al., |
| 6 | 小果榕 Ficus microcarpa | psba、psbb、psbc和psbd均下调;clh1 和clh2 上调;mTERF缺陷 psba,psbb,psbc and psbd are all downregulated;clh1 and clh2 are upregulated;mTERF deficiency | Shih et al., |
| 7 | 日本鸢尾 Iris japonica | GLK下调 GLK downregulation | Yu et al., |
| 8 | 绣球花 Hydrangea macrophylla | HmAP2/ERFs下调 HmAP2/ERFs downregulation | Qi et al., |
| 9 | 虎杖 Reynoutria japonica | rpoC2点突变 rpoC2 point mutation | Park et al., |
| 10 | 垂叶榕 Ficus benjamina | rpoC1短片段缺失 Short fragment deletion in rpoC1 | Park et al., |
| 11 | 粉花绣线菊 Spiraea japonica | rpoB短片段插入 Short fragment insertion in rpoB | Park et al., |
| 12 | 南美榕 Ficus natalensis | rpoB短片段插入 Short fragment insertion in rpoB | Park et al., |
| 13 | 木槿 Hibiscus syriacus | rpoB短片段插入Short fragment insertion in rpoB | Park et al., |
| 14 | 洋常春藤 Hedera helix | psaA短片段缺失 Short fragment deletion in psaA | Park et al., |
| 15 | 合果芋 Syngonium podophyllum | rpoB短片段缺失Short fragment deletion in rpoB | Park et al., |
| 16 | 日本柳杉 Cryptomeria japonica | matK短片段插入突变 Short fragment insertion mutation in matK | Hirao et al., |
| 17 | 大花惠兰 Cymbidium hybrid | ycf3点突变 ycf3 point mutation | Park et al., |
| 18 | 墨兰 Cymbidium sinense | atpI短片段插入 Short fragment insertion in atpI | Park et al., |
| 19 | 长柄玉簪 Hosta longipes | atpE短片段插入 Short fragment insertion in atpE | Park et al., |
| 20 | 吊兰 Chlorophytum comosum | rpoC2短片段缺失 Short fragment deletion in rpoC2 | Park et al., |
| 21 | 富贵竹 Dracaena sanderiana | ycf3内含子点突变 Intronic point mutation in ycf3 | Park et al., |
| 22 | 红苞凤梨 Ananas comosus | VDE、PSY、PDS上调;GS和GDH下调; GLK1、PLA2、FTIP4、AS特异性表达 VDE,PSY,PDS upregulated;GS and GDH downregulated;GLK1,PLA2,FTIP4,AS specifically expressed | Xue et al., Feng et al., |
| 23 | 长果山菅 Dinaella tasmanica | psbA、matK、rps16和trnK的消除 Elimination of psbA,matK,rps16 and trnK | Zhou et al., |
| 24 | 台湾蝴蝶兰 Phalaenopsis aphrodite | PsbO和PsbP下调 PsbO and PsbP downregulated | Tsai et al., |
| 25 | 常春藤叶仙客来 Cyclamen hederifolium | 基因间区域rps4-trnT-UGU的插入/缺失突变 Insertion/Deletion mutation in the intergenic region rps4-trnT-UGU | Mao et al., |
| 26 | 楸树 Catalpa fargesii | PPOX、LHCB1、PsbO、PsbP、PsbQ和PsbR磷酸化差异;CfALKBH5调控m6A甲基化来调控基因表达 Phosphorylation differences in PPOX,LHCB1,PsbO,PsbP,PsbQ,and PsbR;CfALKBH5 regulates gene expression by modulating m6A methylation | Wang et al., |
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