两个马铃薯品种冬、春种植模式下叶片糖苷生物碱合成代谢差异分析

doi:10.16420/j.issn.0513-353x.2024-0667

摘要/Abstract

摘要： 为探究云南省冬（W）、春（S）种植模式下生境差异对不同基因型马铃薯叶片糖苷生物碱（steroidal glycoalkaoids，SGA）合成代谢的影响，以青薯9号（Q9）和丽薯6号（L6）为材料，通过比较蛋白质组学和代谢组学联合分析技术，分析了这两个品种在冬春季节种植期间，处于块茎关键生长时期时叶片中的SGA代谢变化。结果表明：（1）SGA的含量在叶片中随着块茎的发育成熟总体呈上升趋势，但在块茎膨大期中的合成差异最为明显，因此，选择这一时期的叶片开展了基于两种比较模式（Q9-leafW vs. Q9-leafS；Q9-leafW vs. L6-leafW）的组学分析；（2）因品种基因型差异和种植模式引起的生境条件改变，从整体蛋白质表达调控及其导致的代谢物重分配水平上，对马铃薯叶片在两种比较模式下的全局代谢分化造成了初生和次生层面的显著影响；（3）在两种比较模式中分别鉴定到了4个与SGA生物合成高度相关的差异表达蛋白质和差异积累代谢物，它们在SGA生物合成路径的初级和次级代谢阶段中发挥了调控SGA合成积累的不同作用，这可能是引起叶片总SGA在含量和组分上显著差异的关键原因。

关键词: 马铃薯, 种植模式, 叶片, 糖苷生物碱, 组学

Abstract:

To investigate the impacts of habitat discrepancy on the steroidal glycoalkaoids（SGA）anabolism in the leaves of different potato genotypes under the winter（W）and spring（S）cropping patterns in Yunnan Province，the Qingshu 9（Q9）and Lishu 6（L6）potato varieties were chosen as the major experimental materials for the joint comparative proteomic and metabolomic analysis. Alterations occurred on the leaf SGA metabolism have been preliminarily characterized at the key tuber growing stages，when the two potato varieties were cultivated under the winter and spring cropping patterns，respectively. Results indicated that：（1）Total contents of SGA in the potato leaves displayed an uprising trend with the tuber maturation，but had the most obvious significant differences at the tuber bulking stage，which was thereby selected for omics research based on the two comparison modes（Q9-leafW vs. Q9-leafS；Q9-leafW vs. L6-leafW）（2）It was found that the genotypic difference and cropping pattern induced habitat discrepancy had greatly disturbed the overall protein expression regulation and the resulted metabolites repartitioning in potato leaves on both primary and secondary metabolic levels.（3）There were a total of four differentially expressed proteins and accumulated metabolites，which are highly associated with the SGA biosynthesis，identified in the two comparison modes，respectively. Their diversified functions exhibited distinct regulatory effects on the composition and content variations in the potato leaf SGA across the primary and secondary metabolic phases of the SGA biosynthesis.

Key words: potato, cropping pattern, leaf, SGA, omics

祝兴喆, 苏仙月, 苏旺, 张红林, 潘哲超, 刘涛, 徐笑宇. 两个马铃薯品种冬、春种植模式下叶片糖苷生物碱合成代谢差异分析[J]. 园艺学报, 2025, 52(7): 1883-1900.

ZHU Xingzhe, SU Xianyue, SU Wang, ZHANG Honglin, PAN Zhechao, LIU Tao, and XU Xiaoyu. Understanding the Anabolic Differences of Steroidal Glycoalkaloids in Leaf Between Two Potato Varieties Grown at the Winter and Spring Cropping Patterns[J]. Acta Horticulturae Sinica, 2025, 52(7): 1883-1900.

图/表 8

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类别 Category	可溶性糖/ （mg · g^-1DW） Soluble sugar	还原性总糖/ （mg · g^-1DW） Total reducing sugar	水溶性蛋白质/ （mg · g^-1DW） Water-soluble protein	总淀粉含量/ （% DW） Total starch content	直链淀粉含量/ （%DW） Amylose content	维生素C含量/ （nmol · g^-1FW） Vitamin C content
Q9（S）	4.50 b	1.51 b	3.59 c	19.62 a	7.02 a	673.87 b
Q9（W）	3.86 a	0.70 c	6.49 a	18.81 a	6.27 a	936.83 a
L6（S）	8.31 a	2.66 a	5.38 b	12.16 c	3.92 b	808.02 b
L6（W）	8.26 a	2.06 b	4.12 c	14.12 b	4.71 b	1104.46 a

类别 Category	可溶性糖/ （mg · g^-1DW） Soluble sugar	还原性总糖/ （mg · g^-1DW） Total reducing sugar	水溶性蛋白质/ （mg · g^-1DW） Water-soluble protein	总淀粉含量/ （% DW） Total starch content	直链淀粉含量/ （%DW） Amylose content	维生素C含量/ （nmol · g^-1FW） Vitamin C content
Q9（S）	4.50 b	1.51 b	3.59 c	19.62 a	7.02 a	673.87 b
Q9（W）	3.86 a	0.70 c	6.49 a	18.81 a	6.27 a	936.83 a
L6（S）	8.31 a	2.66 a	5.38 b	12.16 c	3.92 b	808.02 b
L6（W）	8.26 a	2.06 b	4.12 c	14.12 b	4.71 b	1104.46 a

亚细胞定位Subcellular localization	蛋白数量Number of proteins		亚细胞定位Subcellular localization	蛋白数量Number of proteins
亚细胞定位Subcellular localization	Q9-leafW vs. Q9-leafS	Q9-leafW vs. L6-leafW	亚细胞定位Subcellular localization	Q9-leafW vs. Q9-leafS	Q9-leafW vs. L6-leafW
液泡Vacuole	14	13	膜Membrane	94	45
单代膜蛋白Single-pass membrane protein	3	1	多通道膜蛋白Multi-pass membrane protein	79	35
分泌蛋白Secreted protein	27	17	细胞质Cytoplasm	17	8
质体Plastid	48	28	基质侧Stromal side	1	-
细胞核Nucleus	10	11	高尔基体膜Golgi apparatus membrane	2	-
线粒体内膜Mitochondrion inner membrane	3	2	单通道 II 型膜蛋白Single-pass type II membrane protein	1	-
管腔侧Lumenal side	1	1	质体膜Plastid membrane	3	-
脂锚定蛋白Lipid-anchor	2	4	核仁Nucleolus	2	-
膜间隙Intermembrane side	1	2	基质蛋白侧Matrix side	1	-
糖基磷脂酰肌醇锚定蛋白GPI-anchor	2	4	单通道IV 型膜蛋白Single-pass type IV membrane protein	2	-
高尔基体Golgi apparatus	5	2	乙醛酸循环体Glyoxysome	1	-
细胞外空间Extracellular space	8	4	内生体膜Endosome membrane	1	-
内质网膜Extracellular space	2	1	衣被凹陷Coated pit	1	-
内质网Endoplasmic reticulum	1	1	胞质小泡Cytoplasmic vesicle	1	-
胞质Cytosol	2	1	胞质侧Cytoplasmic side	2	-
染色体Chromosome	2	2	细胞质Cytoplasm	17	-
叶绿体类囊体膜Chloroplast thylakoid membrane	27	13	包衣小泡膜COPII-coated vesicle membrane	1	-
叶绿体类囊体管腔Chloroplast Thylakoidlumen	3	3	胞质小泡膜Cytoplasmic vesicle membrane	1	-
叶绿体类囊体Chloroplast thylakoid	1	1	顺面高尔基网Cis-Golgi network	1	-
叶绿体基质Chloroplast stroma	1	1	叶绿体膜Chloroplast membrane	3	-
叶绿体被膜Chloroplast envelope	1	1	叶绿体内膜Chloroplast inner membran	1	-
叶绿体Chloroplast	12	9	细胞膜Cell membrane	7	9
细胞壁Cell wall	10	5	胞间连丝Plasmodesma	-	1
单通道 I 型膜蛋白Single-pass type I membrane protein	-	2	线粒体外模Mitochondrion outer membrane	-	1
质外体Apoplast	7	4	线粒体Mitochondrion	-	1
造粉体Amyloplast	2	2	细胞骨架Cytoskeleton	-	1
过氧物酶体Peroxisome	3	2	细胞连接Cell junction	-	1
外周膜蛋白Peripheral membrane protein	9	6	单通道II型膜蛋白Single-pass type II membrane protein	1	-

亚细胞定位Subcellular localization	蛋白数量Number of proteins		亚细胞定位Subcellular localization	蛋白数量Number of proteins
亚细胞定位Subcellular localization	Q9-leafW vs. Q9-leafS	Q9-leafW vs. L6-leafW	亚细胞定位Subcellular localization	Q9-leafW vs. Q9-leafS	Q9-leafW vs. L6-leafW
液泡Vacuole	14	13	膜Membrane	94	45
单代膜蛋白Single-pass membrane protein	3	1	多通道膜蛋白Multi-pass membrane protein	79	35
分泌蛋白Secreted protein	27	17	细胞质Cytoplasm	17	8
质体Plastid	48	28	基质侧Stromal side	1	-
细胞核Nucleus	10	11	高尔基体膜Golgi apparatus membrane	2	-
线粒体内膜Mitochondrion inner membrane	3	2	单通道 II 型膜蛋白Single-pass type II membrane protein	1	-
管腔侧Lumenal side	1	1	质体膜Plastid membrane	3	-
脂锚定蛋白Lipid-anchor	2	4	核仁Nucleolus	2	-
膜间隙Intermembrane side	1	2	基质蛋白侧Matrix side	1	-
糖基磷脂酰肌醇锚定蛋白GPI-anchor	2	4	单通道IV 型膜蛋白Single-pass type IV membrane protein	2	-
高尔基体Golgi apparatus	5	2	乙醛酸循环体Glyoxysome	1	-
细胞外空间Extracellular space	8	4	内生体膜Endosome membrane	1	-
内质网膜Extracellular space	2	1	衣被凹陷Coated pit	1	-
内质网Endoplasmic reticulum	1	1	胞质小泡Cytoplasmic vesicle	1	-
胞质Cytosol	2	1	胞质侧Cytoplasmic side	2	-
染色体Chromosome	2	2	细胞质Cytoplasm	17	-
叶绿体类囊体膜Chloroplast thylakoid membrane	27	13	包衣小泡膜COPII-coated vesicle membrane	1	-
叶绿体类囊体管腔Chloroplast Thylakoidlumen	3	3	胞质小泡膜Cytoplasmic vesicle membrane	1	-
叶绿体类囊体Chloroplast thylakoid	1	1	顺面高尔基网Cis-Golgi network	1	-
叶绿体基质Chloroplast stroma	1	1	叶绿体膜Chloroplast membrane	3	-
叶绿体被膜Chloroplast envelope	1	1	叶绿体内膜Chloroplast inner membran	1	-
叶绿体Chloroplast	12	9	细胞膜Cell membrane	7	9
细胞壁Cell wall	10	5	胞间连丝Plasmodesma	-	1
单通道 I 型膜蛋白Single-pass type I membrane protein	-	2	线粒体外模Mitochondrion outer membrane	-	1
质外体Apoplast	7	4	线粒体Mitochondrion	-	1
造粉体Amyloplast	2	2	细胞骨架Cytoskeleton	-	1
过氧物酶体Peroxisome	3	2	细胞连接Cell junction	-	1
外周膜蛋白Peripheral membrane protein	9	6	单通道II型膜蛋白Single-pass type II membrane protein	1	-

代谢物的名称与分类The nomenclature and classification of metabolites	表达量倍数差异Fold difference in expression
代谢物的名称与分类The nomenclature and classification of metabolites	Q9-leafW vs. Q9-leafS	Q9-leafW vs. L6-leafW
SGAs生物合成相关的物质（6） Substances related to biosynthesis of SGAs（6）
7-酮基胆固醇 7-Ketocholesterol	0.19	1.24
番茄碱 Tomatidine	8.07	5.98
茄啶 Solanidine	-0.69	-1.47
β-茄碱 beta-Solanine	3.44	4.44
α-茄碱 alpha-Solanine	-5.06	-4.06
α-卡茄碱 alpha-Chaconine	-0.34	-1.03
脂类物质（18） lipid（18）
单硬脂酸甘油酯 Glyceryl monostearate	-1.60	-
2-棕榈酰甘油 2-Palmitoylglycerol	-1.29	-
(1R,2S,4R,5R)-5-羟基-1,7,7-三甲基双环[2.2.1]庚-2-基6-O-[(2R,3R,4R)-四氢-3,4-二羟基-4-(羟甲基)-2-呋喃基]-β-D-吡喃葡萄糖苷 (1R,2S,4R,5R)-5-Hydroxy-1,7,7-trimethylbicyclo[2.2.1]hept-2-yl6-O-[（2R,3R,4R）-tetrahydro-3,4-dihydroxy-4-(hydroxymethyl)-2-furanyl]-β-D-glucopyranoside	-2.84	-
4,4,8,10,14-五甲基-17-(4,5,6-三羟基-6-甲基庚烷-2-基)-2，5，6，7，9，15-六氢-1H-环戊二烯[a]菲-3,16-二酮 4，4，8，10，14-Pentamethyl-17-(4,5,6-trihydroxy-6-methylheptan-2-yl)-2,5,6,7,9,15-hexahydro-1H-cyclopenta[a]phenanthrene-3，16-dione	-2.74	-
芥酸酰胺 Erucamide	-2.18	-
叶黄素 Lutein	-1.40	-
贝母甲素 Peimine	-	-1.18
3,5,5-三甲基-4-[3-[(6-O-β-D-木吡喃糖基-β-D-吡喃葡萄糖基)氧]丁基]-2-环己烯-1-酮 3,5,5-Trimethyl-4-[3-[(6-O-β-D-xylopyranosyl-β-D-glucopyranosyl)oxy]butyl]-2-cyclohexen-1-one	-	1.33
9S-羟基-10E,12Z,15Z-十八碳三烯酸 9-HOTrE	3.43	-
十八碳四烯酸 FA 18︰4	3.17	-
9,12,13-三羟基十八碳-10,15-二烯酸9,12,13-Trihydroxyoctadeca-10,15-dienoic acid	1.32	-
乳糖醇 Lactitol	4.40	-
脂肪酸 Sebacic acid	3.47	-
16-羟基棕榈酸 FA 16︰0；2O	4.49	-
(S)-2-羟基-2-甲基琥珀酸 (S)-2-Hydroxy-2-methylsuccinic acid	-	3.17
12,13-二羟基十八烷基-9-烯酸 12,13-Dihydroxyoctadec-9-enoic acid	-	2.26
(Z)-5,8,11-三羟基十八烷基-9-烯酸 (Z)-5,8,11-Trihydroxyoctadec-9-enoic acid	-	1.24
十八碳二烯四氧酸 FA 18：2；4O	-	-1.00
有机氧化物（11） Organic oxide（11）
蔗糖 Sucrose	-	-1.69
苯乙基-2-O-(β-D-吡喃木糖基)-β-D-吡喃葡萄糖苷Phenethyl2-O-(β-D-xylopyranosyl)-β-D-glucopyranoside	-	-1.22
苯甲醇 + 己烷 Benzyl alcohol + Hex-Pen	-1.84	-
决奈达隆 Dronedarone	3.86	-
苯甲醇 + 己烷 Benzyl alcohol + Hex-Hex	-2.00	-
(2S,3S,4S,5R,6R）-6-(3-苯甲酰氧基-2-羟基丙氧基)-3,4,5-三羟基氧烷-2-羧酸 (2S,3S,4S,5R,6R）-6-(3-benzoyloxy-2-hydroxypropoxy）-3,4,5-trihydroxyoxane-2-carboxylic acid	-	3.97
3-O-阿魏酰奎尼酸 3-O-Feruloylquinic acid	-	1.58
红景天苷 Salidroside	-	3.16
山梨醇 Sorbo	-	4.64
乳酮糖 Lactulose	-	-1.79
红霉素 Erythromycin	-	-2.18
苯丙类和聚酮类物质（9） Phenylpropyl and polyketone substances（9）
8,8-二甲基-2-苯基-4H，8-H-苯并[1，2-b：3,4-b’]二吡喃-4-酮 8,8-Dimethyl-2-phenyl-4H，8H-benzo[1，2-b：3,4-b’]dipyran-4-one	1.60	-
黄芩苷II Scutellarioside II	-1.74	-
山奈酚 Kaempferol	-3.64	-
阿魏酰-O-甲氧基酪胺 Feruloyl O-methyldopamine	4.21	-
(S)-7-(2-O-6-脱氧-α-L-甘露糖基)-β-D-吡喃葡萄糖基)-2,3-二氢-5-羟基-2-(3-羟基-4-甲氧基苯基)-4H-1-苯并吡喃-4-酮 (S)-7-(2-O-6-Deoxy-α-L-mannopyranosyl)-β-D-glucopyranosyl) oxy)-2,3-dihydro-5-hydroxy-2-(3-hydroxy-4-methoxyphenyl)-4H-1-benzopyran-4-one	-	-1.15
3-（4-羟基-3-甲氧基苯基）-2-丙烯酸 3-（4-Hydroxy-3-methoxyphenyl）-2-propenoic acid	-	4.45
芦丁 Rutin	-	1.58
（+/-）-木脂酰胺（+/-）-Grossamide	-	3.62
秦皮甲素 Esculin	-	-1.12
有机杂环化合物（6） Organic Heterocyclic Compounds（6）
（3R,4R,5R）-3-[（11E）-11,15-十六碳二烯-9-炔-1-基]-4-羟基-5-甲基二氢-2（3H）-呋喃酮（3R,4R,5R）-3-[（11E）-11，15-Hexadecadien-9-yn-1-yl]-4-hydroxy-5-methyldihydro-2（3H）-furanone	-3.32	-
培高利特 Pergolide	0.88	-
14,15-脱氧喜沃林 Deoxykhivorin	1.72	-
3-[(7S,8R)-5-羟基-2,2,7,8-四甲基-6-氧代-7,8-二氢吡喃[3,2-g]铬-10-基]-3-苯基丙酸 3-[(7S,8R)-5-Hydroxy-2,2,7,8-tetramethyl-6-oxo-7,8-dihydropyrano[3,2-g]chromen-10-yl]-3-phenylpropanoic acid	-2.56	-
脱镁叶绿甲酯酸A Pheophorbide A	2.58	-
升麻素苷 prim-O-b-D-Glucosylcimifugin	-	3.27
有机酸及其衍生物（3） Organic acids and their derivatives（3）
1-氨基-2-羧基吡咯烷 Pyrrolidin-1-ium-2-carboxylate	3.92	-
波纳丁 Bonactin	-	1.94
氢离子；2-羟基丙烷-1,2,3-三羧酸盐 Hydron；2-hydroxypropane-1,2,3-tricarboxylate	-2.12	-
苯环型化合物（1） Benzene ring compound（1）
甜菜安 Desmedipham	-	2.18
核苷、核苷酸及其类似物（1） Nucleosides，nucleotides and their analogs（1）
阿糖尿苷 Arabinofuranosyluracil	2.79	-