两种磷素水平下小麦苗期对干旱胁迫的离子组和代谢组响应

doi:10.3864/j.issn.0578-1752.2022.02.004

摘要/Abstract

摘要：

【目的】探讨两种磷素供应下小麦植株对干旱胁迫的适应性机制及复水后的反应,为揭示小麦水磷互作机制及培育抗逆、磷高效的小麦品种提供理论基础。【方法】本研究采用新疆冬小麦主栽品种新冬23号为参试材料,采用离子组和代谢组学分析方法,研究了低磷（0.05 mmol·L^-1,LP）和常规磷（1.0 mmol·L^-1,CP）水平培养下,小麦植株对干旱胁迫（0、3、5、7 d）及复水3 d的反应。【结果】LP处理下小麦根系浓密。干旱胁迫0 d到7 d再到复水3 d,CP处理的地上部和整株干重呈先升后降的趋势,但LP处理则呈现先升高后降低再升高的趋势。复水3 d后,LP处理根系总体积上升率显著高于CP处理,而LP处理根系DNA含量下降率为21.7%,显著低于CP处理。离子组分析表明,与CP处理相比,LP处理中根系大部分元素含量下降,地上部大部分上升。代谢组分析表明,CP处理对干旱胁迫更加敏感,其中干旱胁迫对CP处理地上部影响大于根系。随着干旱胁迫时间的延长,对氨基酸、脂肪酸和萜类物质的代谢影响更大。【结论】低磷供应的小麦植株对干旱胁迫的敏感性降低,适应性更强,复水后恢复能力强于正常磷供应,其主要原因是根系形态的改变、离子平衡的重排以及氨基酸和脱落酸代谢的改变,以调节渗透平衡,维持细胞内离子稳态,增强了对干旱胁迫的耐性。

关键词: 小麦, 磷素, 干旱胁迫, 生长, 矿质元素, 代谢产物

Abstract:

【Objective】 This study was conducted to explore the adaptive mechanism of wheat plants to drought stress under two kinds of phosphorus supply conditions and its response after rehydration, so as to provide more information for revealing the interaction mechanism of water and phosphorus and breeding the wheat varieties with stress resistance and high phosphorus efficiency. 【Method】 The wheat cultivar Xindong23 were used. The response of wheat plants to drought stress (0, 3, 5, 7 d) and the reaction of rehydration 3 d under low phosphorus (LP: 0.05 mmol·L^-1) and conventional phosphorus (CP: 1.0 mmol·L^-1) were studied by means of ionomics and metabonomics. 【Result】The root system of wheat was dense under LP treatment. From 0 d to 7 d of drought stress and then to 3 d of re-watering, the dry weight of shoot and whole plant under CP treatment increased firstly and then decreased, but that under LP treatment increased firstly and then decreased and then increased. After re-watering for 3 days, the increase rate of total root volume under LP treatment was significantly higher than that under CP treatment. The decrease rate of root DNA content under LP treatment was 21.7%, which was significantly lower than that under CP treatment. Ion group analysis showed that compared with CP, the content of most elements in roots decreased and that in shoots increased under LP treatment. Metabonomics analysis showed that CP treatment was more sensitive to drought stress, and the effect of drought stress on shoot was greater than that of root. With the extension of drought stress time, the metabolism of amino acids, fatty acids and terpenoids was more affected. 【Conclusion】Wheat plants with low phosphorus supply were less sensitive and more adaptable to drought stress, and had stronger recovery ability after rewatering than those with normal phosphorus supply, the main reason is the change of root morphology, rearrangement of ion balance and the change of metabolism of amino acid and abscisic acid to regulate osmotic balance, maintain ion homeostasis and enhance tolerance to drought stress.

Key words: Triticum aestivum L., phosphoru s, drought stress, growth, mineral elements, metabolites

李刚, 白阳, 贾子颖, 马正阳, 张祥池, 李春艳, 李诚. 两种磷素水平下小麦苗期对干旱胁迫的离子组和代谢组响应[J]. 中国农业科学, 2022, 55(2): 280-294.

LI Gang, BAI Yang, JIA ZiYing, MA ZhengYang, ZHANG XiangChi, LI ChunYan, LI Cheng. Phosphorus Altered the Response of Ionomics and Metabolomics to Drought Stress in Wheat Seedlings[J]. Scientia Agricultura Sinica, 2022, 55(2): 280-294.

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https://www.chinaagrisci.com/CN/Y2022/V55/I2/280

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新冬23号小麦旱胁迫3 d和7 d对比下CP处理地上部差异代谢通路及差异代谢物"

编号 No.	代谢通路 Metabolic pathways	差异代谢物 Differential metabolite	干旱3 d/7 d Drought 3 d/7 d
1	氨基酸的生物合成 Biosynthesis of amino acids	活性腺苷甲硫胺酸S-Adenosylmethionine	2.013
		氧戊二酸Oxoglutaric acid	1.263
		2,6-二氨基庚二酸N6-Acetyl-LL-2,6-diaminoheptanedioate	1.648
		(6S,2S)-二氨基庚二酸LL-2,6-diaminoheptanedioate	1.554
		内消旋-2,6-二氨基庚酸Meso-2,6-diaminoheptanedioate	1.554
2	精氨酸生物合成Arginine biosynthesis	氧戊二酸Oxoglutaric acid	1.263
3	组氨酸Histidine metabolism	氧戊二酸Oxoglutaric acid	1.263
4	精氨酸和脯氨酸代谢 Arginine and proline metabolism	S-腺苷-L-蛋氨酸S-Adenosyl-L-methionine	2.013
		N2-琥珀酰-L-鸟氨酸N2-Succinyl-L-ornithine	1.648
		γ-谷氨酰氨基丁酸Gamma-Glutamyl-GABA	1.648
5	酪氨酸代谢Tyrosine metabolism	3-甲氧酪胺3-Methoxytyramine	1.430
6	色氨酸代谢Tryptophan metabolism	吲哚乙醛Indoleacetaldehyde	1.292
6	色氨酸代谢Tryptophan metabolism	N-乙酰血清素N-Acetylserotonin	1.406
7	赖氨酸生物合成 Lysine biosynthesis	氧戊二酸Oxoglutaric acid	1.263
		(2R,3R)-3-甲基谷氨酰-5-半醛-N6-赖氨酸 (2R,3R)-3-Methylglutamyl-5-semialdehyde-N6-lysine	1.480
		2,6-二氨基庚二酸N6-Acetyl-LL-2,6-diaminoheptanedioate	1.648
		LL-2,6-二氨基庚二酸酰胺LL-2,6-diaminoheptanedioate	1.554
		内消旋-2,6-二氨基庚酸Meso-2,6-diaminoheptanedioate	1.433
8	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	氧戊二酸Oxoglutaric acid	1.263
9	半胱氨酸和甲硫氨酸代谢 Cysteine and methionine metabolism	腺苷-L-甲硫氨酸 S-Adenosyl-L-methionine	2.013
		γ-L-谷氨酰胺基-L-2-氨基丁酸盐 Gamma-L-Glutamyl-L-2-aminobutyrate	1.648
		1.2-二羟基-5-（甲硫基）戊-1-烯-3-酮1,2-Dihydroxy-5-(methylthio)pent-1-en-3-one	1.548
		5-甲硫基-D-核糖5-Methylthio-D-ribose	1.548
10	玉米素生物合成Zeatin biosynthesis	S-腺苷-L-蛋氨酸S-Adenosyl-L-methionine,	2.012
11	苯丙氨酸代谢Phenylalanine metabolism	辣椒素Capsaicin	2.070
12	苯丙素生物合成 Phenylpropanoid biosynthesis	乙酸香豆醇酯 Coumaryl acetate	0.526
		甲基异丁香酚Methylisoeugenol	1.477
		阿魏酸Ferulic acid	0.721
		4-羟基-3-甲氧基肉桂酸4-Hydroxy-3-methoxycinnamic acid	0.721
		5-羟基松柏醛5-Hydroxyconiferaldehyde	0.721
13	碳代谢Carbon metabolism	氧戊二酸Oxoglutaric acid	1.263
14	戊糖和葡萄糖醛酸的相互转化 Pentose and glucuronate interconversions	氧戊二酸Oxoglutaric acid	1.263
15	光合作用Photosynthesis	质体醌醇Plastoquinol-1	1.606
16	TCA循环TCA cycle	氧戊二酸Oxoglutaric acid	1.263
17	丙酸代谢Propanoate metabolism	单磷酸盐Monophosphate	0.021
编号 No.	代谢通路 Metabolic pathways	差异代谢物 Differential metabolite	干旱3 d/7 d Drought 3 d/7 d
18	丁酸代谢Butanoate metabolism	氧戊二酸Oxoglutaric acid	1.263
19	抗坏血酸盐和醛酸盐代谢 Ascorbate and aldarate metabolism	氧戊二酸Oxoglutaric acid	1.263
20	α-亚麻酸代谢 Alpha-Linolenic acid metabolism	茉莉酸甲酯Methyl jasmonate	1.927
		(+)-7-异构乙基茉莉酮酸酯 (+)-7-Isomethyljasmonate	1.927
		硬脂酸Stearidonic acid	3.636
21	花生四烯酸代谢Arachidonic acid metabolism	16种16 metabolites	全部增加 All increase
22	类胡萝卜素生物合成Carotenoid biosynthesis	32种32 metabolites	18种减小 18 reduced
23	萜类骨架生物合成 Terpenoid backbone biosynthesis	反式,反式-金合欢醇(2E,6E)-Farnesol	2.203
		金合欢醇 2-trans,6-trans-Farnesal	1.976
		异戊二烯Isoprene	2.730
		全反式六异戊二烯基二磷酸酯 all-trans-Hexaprenyl diphosphate	0.564
24	单萜类生物合成 Monoterpenoid biosynthesis	去氧番木鳖苷 Deoxyloganin	1.581
		（-）-香芹酮 (-)-Carvone	1.554
		（-）-异戊烯酮 (-)-Isopiperitenone	1.554
		异戊烯酮 Ipsdienone	1.554
		（+）-（S）-香芹酮 (+)-(S)-Carvone	1.554
		（+）-门托呋喃 (+)-Menthofuran	1.554
		7-脱氧甘油酯7-Deoxyloganate	1.606
25	二萜类生物合成 Diterpenoid biosynthesis	2,3-脱氢赤霉素A9 2,3-Dehydro-gibberellin A9	1.572
		赤霉素A51分解代谢产物 Gibberellin A51-catabolite	1.599
		赤霉素A5 Gibberellin A5	1.599
		赤霉素A7 Gibberellin A7	1.599
		10β-14β-二羟紫杉-4（20） 10beta,14beta-Dihydroxytaxa-4(20)	1.447
		莫米拉克通 Momilactone A	1.676
		蚜虫素Aphidicolin	4.677
		11（12）-二烯-5α-乙酰氧基-10β-醇 11(12)-dien-5alpha-acetoxy-10beta-ol	1.574
		10-脱乙酰基巴卡汀 10-Deacetylbaccatin III	1.487
26	倍半萜类和三萜类生物合成 Sesquiterpenoid and triterpenoid biosynthesis	52种52 metabolites	51种增加 51 increased
表中只列出部分重要代谢通路及差异代谢物。下同 Only some important metabolic pathways and different metabolites were listed in the table. The same as below

表4

表5

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部位 Tissue	胁迫比较 Stress comparision		元素含量变化 Elements content (%)
部位 Tissue	胁迫比较 Stress comparision		K	Ca	P	Si	Na	Mg	Mn	Cu	Zn	Fe	S
根系 Root	D7	LP/CP	24.0	-78.5	275.7	301.9	-95.9	–	-87.7	-93.3	-87.1	-87.0	90.8
	R3	LP/CP	87.0	-48.9	138.6	282.9	-76.3	22.2	-97.1	-96.7	-91.9	-86.3	53
	CP	R3/D7	-26.3	-51.7	–	34.6	–	–	22.2	102.1	–	–	29.8
	LP	R3/D7	–	–	-33.7	28.2	405.3	–	-71.6	–	-41.9	–	–
地上部 Shoot	D7	LP/CP	-62.9	–	263.9	-88.8	297.3	-67.4	293.8	707.7	90.5	583.4	62.6
	R3	LP/CP	-42.7	–	262.9	-69.1	162.0	-62.7	992.8	442.4	120.8	607.6	41.2
	CP	R3/D7	-33.4	–	–	–	60.8	–	–	26.9	–	–	–
	LP	R3/D7	–	20.4	–	128.6	–	–	198.0	–	–	–	–

编号 No.	代谢通路 Metabolic pathways	差异代谢物 Differential metabolite	LP/CP
1	代谢途径Metabolic pathways	脱落酸ABA	0.162
2	植物信号转导Plant hormone signal transduction	脱落酸ABA	0.162
3	次级代谢物的生物合成Biosynthesis of secondary metabolites	脱落酸ABA	0.162
4	类胡萝卜素生物合成Carotenoid biosynthesis	脱落酸ABA	0.162

编号 No.	代谢通路Metabolic pathways	差异代谢物Differential metabolite	LP/CP
1	叶酸生物合成Folatebiosynthesis	环吡喃磷酸盐CPMP	0.058
2	硫代葡萄糖苷生物合成Glucosinolate biosynthesis	3-(甲基硫代)丙基硫代异氰酸酯3-Methylthiopropyl-desulfoglucosinolate	0.058
3	2-氧代羧酸代谢2-Oxocarboxylic acid metabolism	3-(甲基硫代)丙基硫代异氰酸酯3-Methylthiopropyl-desulfoglucosinolate	0.058
4	嘧啶代谢Pyrimidine metabolism	尿苷酸UMP	0.024
		尿苷5’-磷酸Pseudouridine 5'-phosphate	0.024
		3'-尿苷酸3'-UMP	0.024
5	代谢途径Metabolic pathways	尿苷酸UMP	0.024
		环吡喃磷酸酯 Cyclic pyranopterin monophosphate	0.058
		鸟苷酸GMP	0.058
6	嘌呤代谢Purine metabolism	鸟苷酸GMP	0.058
6	嘌呤代谢Purine metabolism	3'-鸟苷酸3'-GMP	0.058
7	次生代谢物的生物合成Biosynthesis of secondary metabolites	(甲硫基)丙基硫代葡萄糖苷3-Methylthiopropyl-desulfoglucosinolate	0.058
8	硫磺中继系统Sulfur relay system	环吡喃磷酸盐CPMP	0.058

编号 No.	代谢通路 Metabolic pathways	差异代谢物 Differential metabolite	干旱3 d/7 d Drought 3 d/7 d
1	色氨酸代谢Tryptophan metabolism	3-羟基邻氨基苯甲酸酯3-Hydroxyanthranilate	0.056
2	赖氨酸生物合成Lysine biosynthesis	(2R,3R)-3-甲基谷氨酰-5-半醛-N6-赖氨酸(2R,3R)-3-Methylglutamyl-5-semialdehyde-N6-lysine	8.775
3	氨基糖和核苷酸糖代谢 Amino sugar and nucleotide sugar metabolism	2,4-双(乙酰胺基)-2,4,6-三脱氧-β-L-醛缩吡喃糖 2,4-Bis(acetamido)-2,4,6-trideoxy-beta-L-altropyranose	3.135
3	氨基糖和核苷酸糖代谢 Amino sugar and nucleotide sugar metabolism	2,4-二乙酰氨基-2,4,6-三脱氧-α-D-甘露糖吡喃糖 2,4-Diacetamido-2,4,6-trideoxy-alpha-D-mannopyranose	3.135
4	α-亚麻酸代谢 Alpha-Linolenic acid metabolism	9-氧代壬酸9-Oxononanoic acid	1.496
4	α-亚麻酸代谢 Alpha-Linolenic acid metabolism	α-亚麻酸 Alpha-Linolenic acid	0.182
5	不饱和脂肪酸的生物合成 Biosynthesis of unsaturated fatty acids	(6Z,9Z,12Z)-十八碳三烯酸(6Z,9Z,12Z)-Octadecatrienoic acid	0.182
5	不饱和脂肪酸的生物合成 Biosynthesis of unsaturated fatty acids	α-亚麻酸alpha-Linolenate	0.182
6	二萜生物合成Diterpenoid biosynthesis	6β,7β-二羟基贝壳杉烯酸6beta,7beta-Dihydroxykaurenoic acid	0.270
7	单萜生物合成 Monoterpenoid biosynthesis	开联番木鳖苷Secologanin	0.320
8	亚油酸代谢Linoleic acid metabolism	12种12 metabolites	全部增加 All increase