断营养影响青梗菜硝酸盐含量的转录组学分析

doi:10.7668/hbnxb.20193217

摘要/Abstract

摘要：

为降低青梗菜硝酸盐含量,在采收前对其进行断营养处理。生理分析发现,断营养5 d青梗菜产量无明显变化,但根、叶柄和叶片的硝酸盐含量分别降低了49.77%,23.90%,33.39%。转录组比较发现,断营养5 d青梗菜根、叶柄和叶片分别鉴定出301,2 270,2 271个差异表达基因(DEGs)。基因本体论(GO)分析表明,这些DEGs均主要富集在氮(N)代谢、碳(C)代谢和活性氧(ROS)代谢过程中。在N代谢过程中,根、叶柄和叶片硝酸盐摄取转运蛋白NPF7.2基因下调表达;叶柄和叶片NPF7.2的上游调控因子ERF104下调表达;叶片硝酸盐再转运蛋白NPF2.13和NPF1.1基因上调表达;根和叶片硝酸盐同化关键酶谷氨酰胺合成酶(GS)基因上调表达。在C代谢过程中,叶柄和叶片蔗糖合成关键酶磷酸蔗糖合酶(SPS)基因上调表达。在ROS代谢过程中,根的超氧化物歧化酶(SOD)基因和抗坏血酸过氧化物酶(APX)基因上调表达;叶柄和叶片细胞色素P450、过氧化物酶体和脂肪氧化酶下调表达;叶片过氧化氢酶(CAT)基因和过氧化物酶(POD)基因上调表达。综上,在青梗菜采收前5 d进行断营养处理,不仅可以保证产量不受影响,还可以降低硝酸盐含量,提高品质。

关键词: 青梗菜, 硝酸盐, 断营养, N代谢, C代谢, ROS代谢

Abstract:

In order to reduce the nitrate content of Brassica rapa L.ssp. chinensis L.,this study carried out nutrient-breaking treatment before the Brassica rapa L.ssp. chinensis L.was harvested.Physiological analysis of Brassica rapa L.ssp. chinensis L.nutrient-breaking treatment showed that Brassica rapa L.ssp.chinensis L.yield did not decrease significantly at 5 d,but the nitrate content of root,petiole and leaf decreased by 49.77%,23.90% and 33.39% respectively.Transcriptome comparison found that 301,2 270,and 2 271 differentially expressed genes(DEGs)were identified in Brassica rapa L.ssp.chinensis L.root,petiole,and leaf at 5 d,respectively.The Gene Ontology(GO)analysis revealed that these DEGs were mainly enriched in nitrogen(N),carbon(C)and reactive oxygen species(ROS)metabolism.In the process of N metabolism,nitrate uptake transporter NPF7.2 genes were down-regulated expression in root,petiole and leaf.The upstream regulator ERF104 of NPF7.2 were down-regulated expression in petiole and leaf.Nitrate retransporter NPF2.13 and NPF1.1 genes were up-regulated expression in leaf.Nitrate assimilation key enzyme glutamine synthetase(GS)genes were up-regulated expression in root and leaf.In the process of C metabolism,the sucrose phosphate synthase(SPS)genes,a key enzyme for sucrose synthesis in petiole and leaf,were up-regulated expression.In the process of ROS metabolism,superoxide dismutase(SOD)and ascorbate peroxidase(APX)genes were up-regulated expression in root.The expression of cytochrome P450,peroxisome and lipoxygenase genes were down-regulated expression in petiole and leaf.Catalase(CAT)and peroxidase(POD)genes were up-regulated expression in leaf.In summary,the nutrient-breaking treatment 5 d before harvesting can not only ensure that the yield of Brassica rapa L.ssp.chinensis L.was not affected,but also improve the quality of Brassica rapa L.ssp.chinensis L..

Key words: Brassica rapa L.ssp.chinensis L., Nitrate, Nutrient-breaking, N metabolism, C metabolism, ROS metabolism

宋丹华, 焦永刚, 石琳琪, 杨玉波, 郭敬华, 董灵迪. 断营养影响青梗菜硝酸盐含量的转录组学分析[J]. 华北农学报, 2023, 38(1): 74-83. doi: 10.7668/hbnxb.20193217.

SONG Danhua, JIAO Yonggang, SHI Linqi, YANG Yubo, GUO Jinghua, DONG Lingdi. Transcriptomics Analysis of the Influence of Nutrient-breaking on Nitrate Content in Brassica rapa L.ssp.chinensis L.[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(1): 74-83. doi: 10.7668/hbnxb.20193217.

http://www.hbnxb.net/CN/Y2023/V38/I1/74

图/表 12

表1 营养液中各离子浓度

Tab.1 The concentration of each ion in the nutrient solution

药品名称 Name of medicine	质量浓度/(mg/L) Concentration
$NO 3 -$	725.15
$SO 4 2 -$	721.54
K⁺	220.10
Mg²⁺	180.00
Ca²⁺	146.34
$PO 4 3 -$	125.74
EDTA^2-	21.67
Fe²⁺	3.33
$BO 3 3 -$	2.95
H⁺	2.80
Na⁺	1.72
Mn²⁺	0.87
Cl^-	0.49
Zn²⁺	0.27
$MoO 4 2 -$	0.19

表1 营养液中各离子浓度

Tab.1 The concentration of each ion in the nutrient solution

药品名称 Name of medicine	质量浓度/(mg/L) Concentration
$NO 3 -$	725.15
$SO 4 2 -$	721.54
K⁺	220.10
Mg²⁺	180.00
Ca²⁺	146.34
$PO 4 3 -$	125.74
EDTA^2-	21.67
Fe²⁺	3.33
$BO 3 3 -$	2.95
H⁺	2.80
Na⁺	1.72
Mn²⁺	0.87
Cl^-	0.49
Zn²⁺	0.27
$MoO 4 2 -$	0.19

表2 荧光定量PCR中所用差异表达基因及其引物序列

Tab.2 DEGs and their primer sequences in qRT-PCR

基因名称 Gene name	基因ID Gene ID	功能注释 Gene annotation	引物序列(5'—3') Primer sequence (5'—3')
PDE	BnaC05G0211900ZS	丙酮酸脱氢酶E1组分亚基α-2	F: ATGGTATGGATGTACTGGCTGTG
			R: CAATTTTCGCACCCTTTCAAT
NRT1	BnaA01G0117900ZS	蛋白质NRT1/ PTR家族7.2	F: GTGATTGCGGTAATGGCGAT
			R: GACGATGGAAACCAGCAGACTA
NADH	BnaC06G0258800ZS	硝酸还原酶	F: CCGTTGGGCATATCGACTT
			R: TGGCAAACTTAGGTTTACCGTT
GD3	BnaC01G0508900ZS	谷氨酸脱氢酶3	F: TCAACAGGGAGAATGCAAATG
			R: GAAGCCGCGAGTCATGTAGTT
PLTC	BnaA07G0357800ZS	质体蓝素次要异构体	F: ATCGTGTTCAAGAACAACGCAG
			R: ACGGTGAGCTTTCCAACCAT
PL18	BnaA03G0481700ZS	果胶裂解酶18	F: TGTTGCTTGGGCATAGTGATAC
			R: GGAGCCACAAATCTATTCCCTT
CCM	BnaA02G0349200ZS	细胞色素P450	F: CCCTCACATGGCGTCAGAA
			R: ACGAACCCAGAGCCAAAGTC
LPG3	BnaA08G0261700ZS	脂肪氧化酶3	F: ACCGTCCACTTGGACCGG
			R: CTCGTAAGGCAAAACTCCAGC
IP1	BnaA07G0336100ZS	无机焦磷酸酶1	F: TATTTAGGAGATGGAGCTGGTGAT
			R: GCTTTCACCATCGGTCCATT
β-Actin			F: GGAATTGCTGACCGTATGAGC
			R: ACCACGAACCAGAAGGCAGA

图1 断营养处理5 d青梗菜的生长状态

Fig.1 Growth status of Brassica rapa L.ssp.chinensis L. after 5 d of broken nutrition treatment

表3 不同时期单株质量

Tab.3 Weight per plant in different periodskg

处理 Treatment	断营养 Broken nutrition	对照 CK
T₀	0.068±0.006a	0.068±0.006a
T₁	0.070±0.009a	0.072±0.008a
T₂	0.075±0.007a	0.080±0.008a
T₃	0.079±0.007a	0.086±0.007a
T₄	0.084±0.006a	0.094±0.008a
T₅	0.087±0.007a	0.102±0.007a
T₆	0.093±0.009a	0.112±0.008b

图2 不同时期不同部位青梗菜的硝酸盐含量虚线表示硝酸盐含量3 100 mg/kg。

Fig.2 Nitrate content in different parts of Brassica rapa L.ssp.chinensis L.in different periods The dotted line indicates the nitrate content 3 100 mg/kg.

表4 转录组测序数据过滤后的质量统计

Tab.4 Quality statistics of filtered transcript group sequencing data

样品名称 Sample name	clean reads 数量 Number of clean reads	比对到参考基因组唯一位置的reads占 clean reads 的百分比 Percentage of unique mapped reads to clean reads	比对到参考基因组多处位置的reads 占clean reads 的百分比 Percentage of multiple mapped reads to clean reads	GC 含量 /% GC content	Q30 /%
根(对照-1)Root(CK-1)	26 835 645	62.73	2.80	47.56	92.31
根(对照-2)Root(CK-2)	23 523 067	81.04	4.17	47.71	92.51
根(对照-3)Root(CK-3)	22 008 544	81.30	3.98	47.49	93.10
根(断营养-1)Root(Broken nutrition-1)	24 949 367	81.56	4.07	47.57	91.95
根(断营养-2)Root(Broken nutrition-2)	24 268 227	81.41	3.99	47.61	92.73
根(断营养-3)Root(Broken nutrition-3)	22 717 607	81.62	4.18	47.62	92.16
叶柄(对照-1)Petiole(CK-1)	24 677 764	84.27	3.91	47.84	93.03
叶柄(对照-2)Petiole(CK-2)	25 146 080	84.17	4.17	47.73	93.36
叶柄(对照-3)Petiole(CK-3)	23 717 251	83.88	4.02	47.75	92.41
叶柄(断营养-1)Petiole(Broken nutrition-1)	23 112 395	84.08	3.92	47.62	92.85
叶柄(断营养-2)Petiole(Broken nutrition-2)	22 899 957	84.09	3.91	47.65	92.13
叶柄(断营养-3)Petiole(Broken nutrition-3)	23 787 469	84.63	3.80	47.88	93.10
叶片(对照-1)Leaf(CK-1)	21 507 242	80.93	4.44	47.48	91.87
叶片(对照-2)Leaf(CK-2)	23 149 355	83.73	4.49	47.66	93.02
叶片(对照-3)Leaf(CK-3)	21 890 795	83.46	4.53	47.91	92.36
叶片(断营养-1)Leaf(Broken nutrition-1)	24 079 426	83.57	5.23	48.32	93.19
叶片(断营养-2)Leaf(Broken nutrition-2)	22 826 628	83.86	4.60	47.82	92.99
叶片(断营养-3)Leaf(Broken nutrition-3)	22 926 956	83.10	5.31	48.07	92.80
平均Average	23 556 876	81.86	4.20	47.74	92.66

图3 基因表达相关性热图 Y415、Y416、Y417为根对照的3个重复;Y385、Y386、Y387为根断营养处理的3个重复;Y373、Y374、Y375为叶片断营养处理的3个重复;Y403、Y404、Y405为叶片对照的3个重复;Y409、Y410、Y411为叶柄对照的3个重复;Y379、Y378、Y379为叶柄断营养处理的3个重复。

Fig.3 Correlation heatmap of gene expression Y415,Y416,Y417 are three replicates of the root(CK);Y385,Y386,Y387 are three replicates of the root(broken nutrition);Y373,Y374,Y375 are three replicates of the leaf(broken nutrition);Y403,Y404,Y405 are three replicates of the leaf(CK);Y409,Y410,Y411 are three replicates of the petiole(CK);Y379,Y378,Y379 are three replicates of the petiole(broken nutrition).

图4 断营养处理下青梗菜根、叶柄和叶片差异表达基因

Fig.4 DEGs in root,petiole and leaf of Brassica rapa L.ssp. chinensis L.under nutrient-breaking treatment

图5 青梗菜不同部位间DEGs维恩图 G0.叶片的DEGs;G1.叶柄的DEGs;G2.根的DEGs。

Fig.5 Venn diagram of DEGs in different parts of Brassica rapa L.ssp.chinensis L. G0.DEGs of leaf ;G1.DEGs of petiole;G2.DEGs of root.

图6 断营养处理下青梗菜中生物过程GO显著富集term GO term中DEGs数与背景基因数的百分比 1.谷胱甘肽代谢过程;2.活性氮物质代谢过程;3.谷氨酸代谢过程;4.氮循环代谢过程;5.蛋白质代谢过程的调节;6.组氨酸代谢过程;7.蛋白质复合物亚基组织;8.纤维素生物合成过程;9.β-葡聚糖的生物合成过程;10.细胞葡萄糖醛酸化;11.糖鞘脂代谢过程;12.(+)-脱落酸D-吡喃葡萄糖基酯跨膜转运;13.木聚糖生物合成过程;14.含氨基葡萄糖的化合物代谢过程;15.几丁质分解代谢过程;16.氨基糖代谢过程;17.多糖分解代谢过程。

Fig.6 The term of biological function GO was significantly enriched in Brassica rapa L.ssp. chinensis L.under nutrient-breaking treatment The percentage of DEGs number 1.Glutathione metabolic process;2.Reactive nitrogen species metabolic process;3.Glutamate metabolic process;4.Nitrogen cycle metabolic process;5.Regulation of protein metabolic process;6.Histidine metabolic process;7.Protein complex subunit organization;8.Cellulose biosynthetic process;9.β-glucan biosynthetic process;10.Cellular glucuronidation;11.Glycosphingolipid metabolic process;12.(+)-abscisic acid D-glucopyranosyl ester transmembrane transport;13.Xylan biosynthetic process;14.Glucosamine-containing compound metabolic process;15.Chitin catabolic process;16. Amino sugar metabolic process;17.Polysaccharide catabolic process.

图7 断营养处理下青梗菜中分子功能GO显著富集term GO term中DEGs数与背景基因数的百分比 1.L-色氨酸氨基转移酶活性;2.谷胱甘肽结合;3.IAA-氨基酸共轭水解酶活性;4.受体丝氨酸/苏氨酸激酶结合;5.跨膜受体蛋白酪氨酸激酶活性;6.谷胱甘肽转移酶活性;7.高亲和力次级活性铵跨膜转运蛋白活性;8.氧化还原酶活性,氧化金属离子,NAD或NADP为受体;9.氧化还原酶活性,作为供体作用于二酚和相关物质;10.氧化还原酶活性,作用于供体的醛基或羰基上,氧为受体;11.氧化还原酶活性,作用于供体的CH-CH基团,NAD或NADP为受体;12.氧化还原酶活性,氧化金属离子;13.α-D-木糖苷木糖水解酶;14.葡甘露聚糖4-β-甘露糖基转移酶活性;15.半乳糖苷酶活性;16.甘露糖6-磷酸异构酶活性;17.(+)-脱落酸D-吡喃葡萄糖基酯跨膜转运蛋白活性;18.纤维二糖葡萄糖苷酶活性;19.糖苷跨膜转运蛋白活性;20.UDP-葡萄糖基转移酶活性;21.纤维素合酶(UDP形成)活性;22.β-葡聚糖酶活性;23.木葡聚糖特异性内切β-1,4-葡聚糖酶活性;24.几丁质结合。

Fig.7 The term of molecular function GO was significantly enriched in Brassica rapa L.ssp. chinensis L.under nutrient-breaking treatment The percentage of DEGs number 1.L-tryptophan aminotransferase activity;2.Glutathione binding;3.IAA-amino acid conjugate hydrolase activity;4.Receptor serine/threonine kinase binding;5.Transmembrane receptor protein tyrosine kinase activity;6.Glutathione transferase activity;7.High-affinity secondary active ammonium transmembrane transporter activity;8.Oxidoreductase activity,oxidizing metal ions,NAD or NADP as acceptor;9.Oxidoreductase activity,acting on diphenols and related substances as donors;10.Oxidoreductase activity,acting on the aldehyde or oxo group of donors,oxygen as acceptor;11.Oxidoreductase activity,acting on the CH-CH group of donors,NAD or NADP as acceptor;12.Oxidoreductase activity,oxidizing metal ions;13.α-D-xyloside xylohydrolase;14.Glucomannan 4-β-mannosyltransferase activity;15.Galactosidase activity;16.Mannose-6-phosphate isomerase activity;17.(+)-abscisic acid D-glucopyranosyl ester transmembrane transporter activity;18.Cellobiose glucosidase activity;19.Glucoside transmembrane transporter activity;20.UDP-glucosyltransferase activity;21.Cellulose synthase(UDP-forming)activity;22.β-glucanase activity;23.Xyloglucan-specific endo-β-1,4-glucanase activity;24.Chitin binding.

图8 转录组分析鉴定的DEGs的qRT-PCR验证 PDE、NRT1、NADH和GD3来自根;PLTC和PL18来自叶柄;CCM、LPG3和IP1来自叶片。

Fig.8 qRT-PCR verification of DEGs identified by transcriptome analysis PDE,NRT1,NADH and GD3 are derived from the root;PLTC and PL18 come from the petiole;CCM,LPG3 and IP1 come from the leaf.

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