林业科学 ›› 2020, Vol. 56 ›› Issue (1): 38-53.doi: 10.11707/j.1001-7488.20200105
陆小雨1,2,陈竹1,唐菲1,傅松玲2,任杰1,*
收稿日期:
2019-02-01
出版日期:
2020-01-25
发布日期:
2020-02-24
通讯作者:
任杰
基金资助:
Xiaoyu Lu1,2,Zhu Chen1,Fei Tang1,Songling Fu2,Jie Ren1,*
Received:
2019-02-01
Online:
2020-01-25
Published:
2020-02-24
Contact:
Jie Ren
摘要:
目的: 红花槭秋彩叶的形成,与叶片中花青素苷的含量密切相关。本文旨在揭示红花槭中花青素苷的生物合成机理,为其叶色的定向改良提供理论依据。方法: 为解析花青素代谢物积累和基因表达水平的变化,以转色期同时具有绿叶、红叶和黄叶的红花槭特殊单株为材料,用超高效液相色谱串联质谱和高通量RNA测序的方法分别进行代谢组和转录组分析。结果: 1) 在红叶-绿叶、黄叶-绿叶、红叶-黄叶3个比较组中,代谢组正离子模式下分别检测出1 377、1 793、1 098个差异积累代谢物,负离子模式下分别检测出789、699、677个差异积累代谢物:红叶与绿叶相比,矢车菊素苷衍生物、天竺葵素苷元和飞燕草素苷元及其衍生物含量大幅上升;黄叶与绿叶相比,矢车菊素苷衍生物、飞燕草素苷及其衍生物含量增加,而天竺葵素苷及其衍生物减少。2)3个比较组中,转录组测序分别检测出28 536、43 017、27 110个差异表达基因:红叶与绿叶相比,花青素苷合成通路中89.5%的基因表达量增加;黄叶与绿叶相比,花青素苷合成通路中66.7%的基因表达量增加。3)红花槭花青素苷的生物合成中,有29个差异积累的相关代谢物和48个差异表达基因。4)差异代谢物和基因的网络互作分析显示,CHS2, CHS7, CHS8, F3H1, F3H5, F3H7, F3H8, F3H10, F3′H2, LAR, FLS1, FLS2, UFGT4逆向调控天竺葵素苷衍生物和飞燕草素苷衍生物的合成,UFGT5正向调控矢车菊素苷衍生物的合成。结论: 当红花槭叶片秋季变色时,花青素苷通路中大量基因的表达量上调,同时矢车菊素-3-(6″-乙酰半乳糖苷)和矢车菊素-3-阿拉伯糖苷含量大幅上升,此为红花槭叶片变色的主要驱动因子。
中图分类号:
陆小雨,陈竹,唐菲,傅松玲,任杰. 转录组与代谢组联合解析红花槭叶片中花青素苷变化机制[J]. 林业科学, 2020, 56(1): 38-53.
Xiaoyu Lu,Zhu Chen,Fei Tang,Songling Fu,Jie Ren. Combined Transcriptomic and Metabolomic Analysis Reveals Mechanism of Anthocyanin Changes in Red Maple(Acer rubrum) Leaves[J]. Scientia Silvae Sinicae, 2020, 56(1): 38-53.
表1
花青素苷衍生物名称"
代码Code | 中文名称Chinese name | 英文名称English name |
Pg-H-M-C-R-G | 天竺葵素-O(羟基-甲氧基-鼠李糖基-吡喃葡糖苷)-O-吡喃葡糖苷 | Pelargonidin-O-(hydroxy-methoxy-cinnamoyl-L-rhamnopyranosyl-glucopyranoside)-O-glucopyranoside |
Pg-G-H-G | 天竺葵素-O-(吡喃葡糖基-羟基-肉桂酸基-吡喃葡糖苷)-O-吡喃葡糖苷 | Pelargonidin-O-(glucopyranosyl-hydroxycinnamoyl-glucopyranoside)-O-glucopyranoside |
Pg-Gsy | 天竺葵素-3-(2-谷氨基-葡糖基芸香糖苷) | Pelargonidin 3-(2-glu-glucosylrutinoside) |
Pg-Ga | 天竺葵素-5-半乳糖苷 | Pelargonidin 5-galactoside |
Cy-Acet | 矢车菊素-3-(4-乙酰葡糖苷) | Cyanidin 3-(4-acetylglucoside) |
Cy-A-Ga | 矢车菊素-3-(6″-乙酰半乳糖苷) | Cyanidin 3-(6″-acetyl-galactoside) |
Cy-X-H-G | 矢车菊素-O-(吡喃木糖基-羟基肉桂基-吡喃葡糖苷) | Cyanidin-O-(xylopyranosyl-hydroxycinnamoyl-glucopyranosyl) |
Cy-S | 矢车菊素-3-桑布双糖苷 | Cyanidin 3-sambubioside |
Cy-Arab | 矢车菊素-3-阿拉伯糖苷 | Cyanidin 3-arabinoside |
Cy-L | 矢车菊素-3-昆布双糖苷 | Cyanidin 3-laminaribioside |
Del-M-Glu | 飞燕草素-3-(6″-丙二葡糖苷) | Delphinidin 3-(6″-malonyl-glucoside) |
Del-Acet | 飞燕草素-3-乙酰葡糖苷 | Delphinidin 3-acetylglucoside |
Del-di-M | 飞燕草素-3, 5-二-(6-O-丙二葡糖苷) | Delphinidin 3, 5-di(6-O-malonylglucoside) |
Del-S | 飞燕草素-3-桑布双糖苷 | Delphinidin 3-sambubioside |
图4
红叶、绿叶和黄叶中花青素苷合成途径上相关基因和代谢物的细节展示 CHS: 查尔酮合成酶;CHI: 查尔酮异构酶;F3H: 黄烷酮3-羟化酶;F3′H: 黄烷酮3′-羟化酶;F3′5′H: 黄烷酮3′5′-羟化酶;DFR: 二氢黄酮醇4-还原酶;ANS: 花青素苷合成酶;UFGT: 类黄酮3-O-葡糖基转移酶;FLS: 黄酮醇合成酶;LAR: 原花青素苷元还原酶;ANR: 花青素苷还原酶。P-Coumaroyl-CoA:对肉桂酸辅酶A;Naringenin chalcone:柚皮素查尔酮;Naringenin:柚皮素;DHK:二氢山奈酚;DHQ:二氢槲皮素;DHM:二氢杨梅素;Quercetin:槲皮素;Kaempferol:山奈酚;Myricetin:杨梅素;Leucocyanidin:原矢车菊素苷元;Leucopelargonidin:原天竺葵素苷元;Leucodelphnidin:原飞燕草素苷元;Catechin:儿茶素;Afzelechin:阿福豆素;Gallocatechin:没食子儿茶素;Cy: 矢车菊素苷;Pg: 天竺葵素苷;Del: 飞燕草素苷;Epicatechin:表儿茶素;Epiafzelechin:表阿福豆素;Epigallocatechin:表没食子儿茶素;Cy-3-glucoside:矢车菊素-3-葡糖苷;Pg-3-glucoside:天竺葵素-3-葡糖苷;Del-3-glucoside:飞燕草素-3-葡糖苷。"
表2
不同比较组间显著富集KEGG通路(修正P≤0.01)"
编号No. | 通路名称Pathway | 通路ID Pathway ID | 通路注释的差异表达基因DEGs with pathway annotation | 通路注释的所有基因All genes with pathway annotation | P | 修正P值Corrected P |
红叶-绿叶Red leaves-green leaves | ||||||
1 | 核糖体Ribosome | ko03010 | 286 | 441 | 2.67E-23 | 3.26E-21 |
2 | 卟啉和叶绿素代谢Porphyrin and chlorophyll metabolism | ko00860 | 120 | 193 | 7.66E-10 | 4.67E-08 |
3 | 类黄酮生物合成Flavonoid biosynthesis | ko00941 | 76 | 102 | 4.37E-09 | 1.78E-07 |
4 | 光合作用-天线蛋白质Photosynthesis-antenna proteins | ko00196 | 58 | 76 | 1.58E-07 | 4.82E-06 |
5 | 精氨酸和脯氨酸代谢Arginine and proline metabolism | ko00330 | 171 | 377 | 5.79E-06 | 0.000 137 722 |
6 | 半胱氨酸和蛋氨酸代谢Cysteine and methionine metabolism | ko00270 | 224 | 525 | 6.77E-06 | 0.000 137 722 |
7 | 苯丙氨酸代谢Phenylalanine metabolism | ko00360 | 115 | 231 | 8.66E-06 | 0.000 150 948 |
8 | 精氨酸生物合成Arginine biosynthesis | ko00220 | 86 | 162 | 1.94E-05 | 0.000 295 595 |
9 | 甘氨酸、丝氨酸和苏氨酸代谢Glycine, serine and threonine metabolism | ko00260 | 142 | 312 | 2.97E-05 | 0.000 403 202 |
10 | 乙醛酸和二羧酸代谢Glyoxylate and dicarboxylate metabolism | ko00630 | 246 | 612 | 6.04E-05 | 0.000 736 680 |
11 | 抗坏血酸代谢Ascorbate and aldarate metabolism | ko00053 | 124 | 274 | 0.000 109 140 | 0.001 210 459 |
12 | 泛醌和其他萜类化合物-醌生物合成Ubiquinone and other terpenoid-quinone biosynthesis | ko00130 | 88 | 182 | 0.000 194 958 | 0.001 982 071 |
13 | 磷酸戊糖途径Pentose phosphate pathway | ko00030 | 151 | 360 | 0.000 349 958 | 0.003 284 225 |
14 | 苯丙氨酸、酪氨酸和色氨酸生物合成Phenylalanine, tyrosine and tryptophan biosynthesis | ko00400 | 87 | 186 | 0.000 479 922 | 0.004 182 176 |
15 | 植物-病原体相互作用Plant-pathogen interaction | ko04626 | 230 | 595 | 0.000 584 893 | 0.004 757 129 |
16 | 光合生物中的碳固定Carbon fixation in photosynthetic organisms | ko00710 | 229 | 593 | 0.000 625 676 | 0.004 770 781 |
17 | 谷胱甘肽代谢Glutathione metabolism | ko00480 | 158 | 388 | 0.000 727 198 | 0.005 094 233 |
18 | 苯丙烷类生物合成Phenylpropanoid biosynthesis | ko00940 | 96 | 214 | 0.000 751 608 | 0.005 094 233 |
19 | 氰氨基酸代谢Cyanoamino acid metabolism | ko00460 | 50 | 94 | 0.000 929 480 | 0.005 968 242 |
20 | 泛酸和CoA生物合成Pantothenate and CoA biosynthesis | ko00770 | 65 | 133 | 0.000 998 095 | 0.006 088 377 |
21 | 类胡萝卜素生物合成Carotenoid biosynthesis | ko00906 | 67 | 140 | 0.001 276 839 | 0.007 417 825 |
22 | 缬氨酸、亮氨酸和异亮氨酸的生物合成Valine, leucine and isoleucine biosynthesis | ko00290 | 55 | 110 | 0.001 581 266 | 0.008 768 837 |
23 | 果糖和甘露糖代谢Fructose and mannose metabolism | ko00051 | 125 | 304 | 0.001 850 311 | 0.009 814 695 |
24 | 二苯乙烯类、二芳基庚烷类和姜辣素类的生物合成Stilbenoid, diarylheptanoid and gingerol biosynthesis | ko00945 | 27 | 42 | 0.001 955 262 | 0.009 939 251 |
绿叶-黄叶Green leaves-yellow leaves | ||||||
1 | 核糖体Ribosome | ko03010 | 322 | 441 | 1.91E-13 | 2.33E-11 |
2 | 卟啉和叶绿素代谢Porphyrin and chlorophyll metabolism | ko00860 | 136 | 193 | 5.56E-06 | 0.000 339 316 |
红叶-黄叶Red leaves-yellow leaves | ||||||
1 | 类黄酮生物合成Flavonoid biosynthesis | ko00941 | 73 | 102 | 1.82E-10 | 2.22E-08 |
2 | 苯丙烷类生物合成Phenylpropanoid biosynthesis | ko00940 | 108 | 214 | 3.92E-08 | 2.39E-06 |
3 | 甘油脂代谢Glycerolipid metabolism | ko00561 | 202 | 520 | 1.16E-06 | 4.73E-05 |
4 | 调节自噬Regulation of autophagy | ko04140 | 118 | 287 | 2.67E-05 | 0.000 812 852 |
5 | 核糖体Ribosome | ko03010 | 164 | 441 | 6.85E-05 | 0.001 671 012 |
6 | 淀粉和蔗糖代谢Starch and sucrose metabolism | ko00500 | 370 | 1 149 | 0.000 168 590 | 0.003 427 987 |
7 | 苯丙氨酸、酪氨酸和色氨酸生物合成Phenylalanine, tyrosine and tryptophan biosynthesis | ko00400 | 78 | 186 | 0.000 350 292 | 0.005 966 480 |
8 | 糖胺聚糖降解Glycosaminoglycan degradation | ko00531 | 23 | 33 | 0.000 391 245 | 0.005 966 480 |
9 | N-聚糖生物合成N-glycan biosynthesis | ko00510 | 75 | 182 | 0.000 664 722 | 0.009 010 676 |
10 | 果糖和甘露糖代谢Fructose and mannose metabolism | ko00051 | 113 | 304 | 0.000 842 426 | 0.009 555 408 |
11 | 萜类骨架生物合成Terpenoid backbone biosynthesis | ko00900 | 67 | 160 | 0.000 896 156 | 0.009 555 408 |
12 | 脂肪酸延伸率Fatty acid elongation | ko00062 | 25 | 41 | 0.000 939 876 | 0.009 555 408 |
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