不同盐碱胁迫对棉花营养元素吸收转运以及代谢的影响

鲁晓宇; 郭家鑫; 陶一凡; 叶扬; 桂诚浩; 郭慧娟; 闵伟

doi:10.12357/cjea.20220581

不同盐碱胁迫对棉花营养元素吸收转运以及代谢的影响

doi: 10.12357/cjea.20220581

石河子大学农学院资源与环境科学系　石河子　832003

基金项目: 兵团中青年科技创新领军人才计划(2020CB020)、石河子大学青年创新人才培养计划(CXPY202111)和农业农村部西北绿洲农业环境重点实验室开放基金(XBLZ-20214)资助

详细信息

作者简介:
鲁晓宇, 主要从事植物逆境生理研究。E-mail: 1804221194@qq.com

通讯作者:
闵伟, 主要从事土壤水盐运移、土壤肥力与调控及土壤微生物分子生态研究。E-mail: minwei555@126.com

中图分类号: S562
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出版历程
- 收稿日期: 2022-07-26
- 录用日期: 2022-09-19
- 网络出版日期: 2022-11-07
- 刊出日期: 2023-03-10

Effects of different salt and alkali stress on absorption, transportation, and metabolism of nutrient elements in cotton

Department of Resources and Environmental Sciences, Shihezi University, Shihezi 832003, China

Funds: This study was supported by the Program of Youth Science and Technology Innovation Leader of the Xinjiang Production and Construction Corps (2020CB020), the Youth Innovation Talent Cultivation Program of Shihezi University (CXPY202111), and the Open Fund of Key Laboratory of Northwest Oasis Agro-Environment, Ministry of Agriculture of China (XBLZ-20214).

More Information

Corresponding author: E-mail: minwei555@126.com

摘要

摘要: 新疆盐碱地类型多且积盐严重, 因此, 探讨不同盐碱胁迫对棉花养分吸收转运和代谢的影响, 揭示棉花对不同盐碱胁迫的耐受机制, 可为新疆不同盐碱地类型棉花栽培提供一定的理论基础。本研究设置对照(CK)、氯化钠(NaCl)盐胁迫(CS)、硫酸钠(Na₂SO₄)盐胁迫(SS)和碱(NaHCO₃+Na₂CO₃)胁迫(AS) 4个处理, 通过离子组和代谢组学的方法, 探究不同盐碱胁迫对棉花根和叶中营养元素含量以及代谢的影响。结果表明, 盐碱胁迫显著抑制棉花生长, 与CK相比, CS、SS和AS处理的棉花总生物量分别显著(P<0.05, 下同)降低51.7%、47.8%和52.3%, CS处理叶片N含量显著增加, P、K、Ca、Mg和S含量显著降低, 茎中N、P、K、Ca、Mg和S含量均显著降低, 根中N、Ca和Mg含量显著降低, P含量显著增加; SS处理叶中P、Ca和Mg含量显著降低, S含量显著增加, 茎中P、Ca和Mg含量显著降低, S含量显著增加, 根中P和Ca含量显著降低, Mg和S含量显著增加; AS处理叶中P、K、Ca、Mg和S含量显著降低, 茎中N、P、Ca、Mg和S含量显著降低, 根中N、P和S含量显著降低, Mg含量显著增加。CS处理下棉花叶片和根系中分别筛选出7条差异代谢通路, SS处理下棉花叶片和根系中分别筛选出16和29条差异代谢通路, AS处理下棉花叶片和根系中分别筛选出8条和18条差异代谢通路。氯化钠胁迫抑制棉花P、Ca、Mg和S的转运能力, 但是对代谢的影响相对较小; 硫酸钠胁迫下棉花体内积累的S促进了氨基酸类代谢; 碱胁迫抑制了P、K、Ca、Mg和S的转运能力, 根部有机酸代谢增强且有机酸显著积累, 叶片中亚油酸显著积累。
- 棉花 /
- NaCl胁迫 /
- Na₂SO₄胁迫 /
- 碱胁迫 /
- 营养元素 /
- 代谢组
Abstract: There are many types of saline-alkali land in Xinjiang, and salt accumulation is a serious problem. Therefore, this study aimed to explore the effects of different saline-alkali stresses on the absorption, transportation, and metabolism of cotton nutrients and to reveal the tolerance mechanism of cotton to different salt and alkali stresses to provide a theoretical basis for cotton cultivation on different types of saline-alkali land in Xinjiang. Four treatments were used in this study: control (CK), NaCl stress (CS), Na₂SO₄ stress (SS), and alkali (NaHCO₃+Na₂CO₃) stress (AS). The effects of different salt and alkali stresses on the content and metabolism of nutrient elements in cotton roots and leaves were explored using ionomics and metabonomics. Compared with CK, the total biomass of CS, SS, and AS treatments decreased significantly (P<0.05) by 51.7%, 47.8%, and 52.3%, respectively. Compared with CK, CS treatment significantly (P<0.05) increased the content of N in leaves and the content of P in roots; significantly (P<0.05) decreased the contents of P, K, Ca, Mg, and S in leaves; the contents of N, P, K, Ca, Mg, and S in stems; the content of N, Ca, and Mg in roots. Under AS treatment, the contents of P, K, Ca, Mg and S in leaves; the contents of N, P, Ca, Mg and S in stems; and the contents of N, P and S in roots significantly (P<0.05) decreased, while Mg content in roots significantly (P<0.05) increased over the CK treatment. Seven differential metabolic pathways were screened from cotton leaves and roots under the CS treatment; 16 and 29 differential metabolic pathways were screened from cotton leaves and roots under the SS treatment; and eight and 18 differential metabolic pathways were screened from cotton leaves and roots under the AS treatment. NaCl stress inhibited the transport of P, Ca, Mg, S, and N uptake in cotton but promoted the transport of N and K; NaCl stress had relatively little effect on metabolism, only the accumulation of amino acids and organic acids. Na₂SO₄ stress did not significantly inhibit the absorption and transport of N and K but promoted the absorption of Mg and S. However, it inhibited the absorption and transport of Ca and Mg in cotton. Na₂SO₄ stress had a significant impact on metabolism. The significant accumulation of S incotton promotes the metabolism of amino acids, and the enhancement of amino acid metabolism also indirectly promotes the intensity of other metabolic pathways, making cotton more tolerant to Na₂SO₄. Alkali (NaHCO₃+Na₂CO₃) stress inhibited N and S uptake and P, K, Ca, Mg, and S transport but increased Mg uptake, which significantly changed the metabolism of organic acids in cotton, enhanced the metabolism of organic acids in roots, significantly accumulated organic acids, and significantly accumulated linoleic acid in leaves.
- Cotton /
- NaCl stress /
- Na₂SO₄ stress /
- Alkali stress /
- Nutrient elements /
- Metabolomics

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表 1 不同处理土壤盐碱类型及盐碱化程度

Table 1. Type and degree of saline and alkaline in soil of different treatments

处理 Treatment	盐碱类型 Saline alkali type	含盐量 Salt content (g∙kg⁻¹)	电导率 Electrical conductivity (EC_1:5, dS∙m⁻¹)	pH (1∶2.5)
CK	非盐(碱)化 Control (non salting/alkalization)	0.53	0.17	8.16
CS	氯化钠盐胁迫 NaCl stress	4.43	1.39	8.43
SS	硫酸钠盐胁迫 Na₂SO₄ stress	6.43	2.01	8.19
AS	碱胁迫 Na₂CO₃+NaHCO₃ stress	2.03	0.63	9.92

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表 2 不同盐碱胁迫对棉花生物量的影响

Table 2. Effects of different salt and alkaline stresses on cotton biomass

处理 Treatment	生物量 Biomass (g∙plant⁻¹)
	叶 Leaf	茎 Stem	根 Root	总 Total
CK	1.44±0.01a	1.69±0.02a	0.93±0.01a	4.05±0.04a
CS	0.75±0.00b	0.58±0.02d	0.63±0.01b	1.96±0.02c
SS	0.77±0.01b	0.83±0.01b	0.52±0.02c	2.11±0.02b
AS	0.57±0.01c	0.72±0.03c	0.65±0.02b	1.93±0.03c
各处理说明见表1。同列不同小写字母表示不同处理间差异显著(P<0.05)。The description of each treatment is shown in Table 1. Different lowercase letters in the same column indicate significant differences among treatments (P<0.05).

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表 3 不同盐碱胁迫对棉花养分吸收和分配的影响

Table 3. Effects of different saline alkali stresses on nutrient uptake and distribution in cotton

处理 Treatment		含量 Content (g∙kg⁻¹)						器官分配比 Distribution ratio of organ
		N	P	K	Ca	Mg	S	N	P	K	Ca	Mg	S
叶 Leaves	CK	38.91±0.82b	2.77±0.08a	26.73±0.56a	56.67±1.23a	11.72±0.40a	21.02±0.33b	0.55b	0.45a	0.45b	0.75c	0.59b	0.84b
	CS	44.33±0.58a	2.22±0.09c	22.25±0.72c	44.54±0.58b	9.48±0.23c	18.02±0.64c	0.64a	0.44ab	0.51a	0.80a	0.65a	0.86a
	SS	38.19±0.11b	2.06±0.03d	27.50±0.26a	41.91±0.14c	10.14±0.06b	27.82±0.46a	0.55b	0.46a	0.47b	0.77b	0.60b	0.86a
	AS	37.86±0.44b	2.38±0.08b	23.88±0.56b	33.99±0.41d	8.52±0.15d	17.33±0.36c	0.55b	0.43b	0.38c	0.67d	0.48d	0.81c
茎 Stem	CK	19.86±0.41a	2.09±0.01a	21.70±0.11a	14.15±0.16a	5.93±0.03a	2.83±0.02b	0.33a	0.40a	0.44a	0.22b	0.35b	0.13a
	CS	18.63±0.43b	1.92±0.04b	16.62±0.23b	10.41±0.21b	4.70±0.09d	2.56±0.04c	0.21c	0.30c	0.30c	0.15d	0.25d	0.10d
	SS	20.29±0.59a	1.57±0.04d	21.75±0.27a	9.36±0.32d	4.91±0.02c	3.33±0.07a	0.32b	0.38b	0.40b	0.19c	0.31c	0.11b
	AS	18.02±0.19b	1.74±0.01c	21.53±0.15a	9.91±0.07c	5.34±0.08b	2.43±0.05d	0.33ab	0.39ab	0.43a	0.25a	0.38a	0.14a
根 Roots	CK	12.88±0.15a	1.36±0.04b	10.20±0.43ab	3.80±0.08a	1.69±0.03d	1.18±0.04b	0.12c	0.14c	0.11b	0.03d	0.06d	0.03b
	CS	12.38±0.25b	1.52±0.04a	9.82±0.06b	3.26±0.06b	1.78±0.01c	1.21±0.05b	0.15a	0.26a	0.19a	0.05b	0.10b	0.05a
	SS	13.16±0.22a	1.06±0.02c	10.63±0.25a	3.08±0.04c	2.11±0.03b	1.42±0.05a	0.13b	0.16b	0.13b	0.04c	0.09c	0.03b
	AS	7.73±0.31c	0.90±0.01d	10.65±0.03a	3.76±0.08a	2.30±0.04a	0.96±0.01c	0.12bc	0.18b	0.19a	0.08a	0.14a	0.05a
各处理说明见表1。同列不同小写字母表示不同处理间差异显著(P<0.05)。The description of each treatment is shown in Table 1. Different lowercase letters in the same column indicate significant differences among treatments (P<0.05).

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表 4 NaCl胁迫下棉花根差异代谢通路和差异代谢物

Table 4. Differential metabolic pathways and metabolites in cotton roots under NaCl stress

编号 No.	代谢通路 Metabolic pathway	差异代谢物 Differential metabolite	变化倍数 Fold change
1	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	L-谷氨酰胺 L-Glutamine	1.26
1	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	琥珀酸半醛 Succinic semialdehyde	1.28
2	丁酸代谢 Butanoate metabolism	琥珀酸半醛 Succinic semialdehyde	1.28
3	泛酸和辅酶A生物合成 Pantothenate and CoA biosynthesis	泛酸 Pantothenate	1.35
3	泛酸和辅酶A生物合成 Pantothenate and CoA biosynthesis	二氢尿嘧啶 Dihydrouracil	1.32
4	精氨酸生物合成 Arginine biosynthesis	L-谷氨酰胺 L-Glutamine	1.26
5	半乳糖代谢 Galactose metabolism	肌醇 myo-Inositol	1.39
		蔗糖 Sucrose	1.40
		UDP-D-半乳糖 UDP-D-Galactose	1.13
6	精氨酸和脯氨酸代谢 Arginine and proline metabolism	4-乙酰氨基丁酸酯 4-acetamidobutanoate	0.74
7	磷酸肌醇代谢 Inositol phosphate metabolism	肌醇 myo-Inositol	1.39

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表 5 Na₂SO₄胁迫下棉花根差异代谢通路和差异代谢物

Table 5. Differential metabolic pathways and metabolites in cotton roots under Na₂SO₄ stress

编号 No.	代谢通路 Metabolic pathway	差异代谢物 Differential metabolite	变化倍数 Fold change
1	异喹啉生物碱的生物合成 Isoquinoline alkaloid biosynthesis	L-酪氨酸 L-Tyrosine	1.35
1	异喹啉生物碱的生物合成 Isoquinoline alkaloid biosynthesis	多巴胺 Dopamine	1.36
2	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	L-丙氨酸 L-Alanine	2.85
		L-天冬酰胺 L-Asparagine	1.27
		丙酮酸 Pyruvate	1.49
		L-谷氨酰胺 L-Glutamine	1.37
		L-谷氨酸 L-Glutamate	1.79
		琥珀酸 Succinate	2.10
		琥珀酸半醛 Succinic semialdehyde	1.74
3	牛磺酸和低牛磺酸代谢 Taurine and hypotaurine metabolism	牛磺酸 Taurine	1.57
4	C5支化二元酸代谢 C5-Branched dibasic acid metabolism	柠檬酸 Citraconic acid	1.34
4	C5支化二元酸代谢 C5-Branched dibasic acid metabolism	丙酮酸 Pyruvate	1.49
5	苯丙氨酸代谢 Phenylalanine metabolism	苯丙氨酸 L-Phenylalanine	1.54
6	甘油磷脂代谢 Glycerophospholipid metabolism	甘油磷酸胆碱 Glycerophosphocholine	0.62
6	甘油磷脂代谢 Glycerophospholipid metabolism	磷酰胆碱 Phosphorylcholine	1.33
7	精氨酸生物合成 Arginine biosynthesis	L-谷氨酸 L-Glutamate	1.79
		N-乙酰-L-谷氨酸 N-Acetyl-L-glutamate	0.75
		L-瓜氨酸 L-Citrulline	3.58
		L-谷氨酰胺 L-Glutamine	1.37
		N2-乙酰-L-鸟氨酸 N2-Acetyl-L-ornithine	0.47
8	淀粉和蔗糖代谢 Starch and sucrose metabolism	蔗糖 Sucrose	1.49
		D-麦芽糖 D-Maltose	2.33
		D-葡萄糖6-磷酸 D-Glucose 6-phosphate	1.81
		α, α-海藻糖 Alpha, alpha-Trehalose	1.80
9	甘氨酸、丝氨酸和苏氨酸代谢 Glycine, serine and threonine metabolism	L-苏氨酸 L-Threonine	0.79
		L-丝氨酸 L-Serine	1.57
		丙酮酸 Pyruvate	1.49
10	丙酮酸代谢 Pyruvate metabolism	丙酮酸 Pyruvate	1.49
10	丙酮酸代谢 Pyruvate metabolism	L-苹果酸 L-Malic acid	2.76
11	乙醛酸和二羧酸代谢 Glyoxylate and dicarboxylate metabolism	D-核酮糖1,5-二磷酸 D-Ribulose 1,5-bisphosphate	1.46
		顺乌头酸 Cis-Aconitate	2.10
		L-苹果酸 L-Malic acid	2.76
		琥珀酸 Succinate	2.10
		磷酸乙醇酸 Phosphoglycolic acid	0.68
		L-谷氨酸 L-Glutamate	1.79
		L-丝氨酸 L-Serine	1.57
		L-谷氨酰胺 L-Glutamine	1.37
		甘油酸 Glyceric acid	1.44
12	半乳糖代谢 Galactose metabolism	UDP-D-半乳糖 UDP-D-Galactose	1.22
		水苏糖 Stachyose	2.92
		棉子糖 Raffinose	2.32
		肌醇 Myo-Inositol	1.61
		蔗糖 Sucrose	1.49
		甘油 Glycerol	5.50
13	戊糖磷酸途径 Pentose phosphate pathway	D-核糖 D-Ribose	2.39
13	戊糖磷酸途径 Pentose phosphate pathway	D-核糖5-磷酸 D-Ribose 5-phosphate	0.38
14	丁酸代谢 Butanoate metabolism	L-谷氨酸 L-Glutamate	1.79
		丙酮酸 Pyruvate	1.49
		琥珀酸半醛 Succinic semialdehyde	1.74
		琥珀酸 Succinate	2.10
15	柠檬酸循环(TCA循环) Citrate cycle (TCA cycle)	L-苹果酸 L-Malic acid	2.76
		琥珀酸 Succinate	2.10
				顺乌头酸 Cis-Aconitate	2.10
				丙酮酸 Pyruvate	1.49
16	光合生物的固碳作用 Carbon fixation in photosynthetic organisms	L-苹果酸 L-Malic acid	2.76
		丙酮酸 Pyruvate	1.49
		L-丙氨酸 L-Alanine	2.85
		D-核糖5-磷酸 D-Ribose 5-phosphate	0.38
		D-核酮糖1,5-二磷酸 D-Ribulose 1,5-bisphosphate	1.46
17	谷胱甘肽代谢 Glutathione metabolism	谷胱甘肽二硫化物 Glutathione disulfide	1.93
		L-谷氨酸 L-Glutamate	1.79
		γ-谷氨酰半胱氨酸 Gamma-Glutamylcysteine	1.85
		L-焦谷氨酸 L-Pyroglutamic acid	2.46
18	精氨酸和脯氨酸代谢 Arginine and proline metabolism	脯氨酸 Proline	1.43
		4-胍基丁酸 4-Guanidinobutyric acid	1.22
		4-乙酰氨基丁酸酯 4-acetamidobutanoate	0.70
		L-谷氨酸 L-Glutamate	1.79
19	氨基糖和核苷酸糖代谢 Amino sugar and nucleotide sugar metabolism	UDP-D-半乳糖 UDP-D-Galactose	1.22
19	氨基糖和核苷酸糖代谢 Amino sugar and nucleotide sugar metabolism	UDP-N-乙酰葡糖胺 UDP-N-acetylglucosamine	1.57
20	缬氨酸、亮氨酸和异亮氨酸生物合成 Valine, leucine and isoleucine biosynthesis	L-亮氨酸 L-Leucine	0.83
		柠檬酸 Citraconic acid	1.34
		L-苏氨酸 L-Threonine	0.79
		丙酮酸 Pyruvate	1.49
		L-异亮氨酸 L-Isoleucine	0.44
21	泛酸和辅酶A生物合成 Pantothenate and CoA biosynthesis	丙酮酸 Pyruvate	1.49
		尿嘧啶 Uracil	1.83
		泛酸 Pantothenate	1.50
22	角质、木栓碱和蜡的生物合成 Cutin, suberine and wax biosynthesis	油酸 Oleic acid	0.68
22	角质、木栓碱和蜡的生物合成 Cutin, suberine and wax biosynthesis	苯甲酸 Behenic acid	1.96
23	类黄酮生物合成 Flavonoid biosynthesis	柚皮素 Naringenin	0.80
24	糖酵解/糖异生 Glycolysis / Gluconeogenesis	丙酮酸 Pyruvate	1.49
25	酪氨酸代谢 Tyrosine metabolism	L-酪氨酸 L-Tyrosine	1.35
		多巴胺 Dopamine	1.36
		丙酮酸 Pyruvate	1.49
26	氨酰tRNA生物合成 Aminoacyl-tRNA biosynthesis	L-天冬酰胺 L-Asparagine	1.27
		L-组氨酸 L-Histidine	0.75
		苯丙氨酸 L-Phenylalanine	1.54
		L-谷氨酰胺 L-Glutamine	1.37
		L-丝氨酸 L-Serine	1.57
		L-丙氨酸 L-Alanine	2.85
		L-异亮氨酸 L-Isoleucine	0.44
		L-亮氨酸 L-Leucine	0.83
		L-苏氨酸 L-Threonine	0.79
		L-酪氨酸 L-Tyrosine	1.35
		L-脯氨酸 L-Proline	1.44
		L-谷氨酸 L-Glutamate	1.79
27	硫代谢 Sulfur metabolism	L-丝氨酸 L-Serine	1.57
		O-乙酰-L-丝氨酸 O-Acetyl-L-serine	3.34
		琥珀酸 Succinate	2.10
28	苯丙氨酸、酪氨酸和色氨酸生物合成 Phenylalanine, tyrosine and tryptophan biosynthesis	预苯酸 Prephenate	1.69
		L-苯丙氨酸 L-Phenylalanine	1.54
		L-酪氨酸 L-Tyrosine	1.35
29	肌醇磷酸代谢 Inositol phosphate metabolism	肌醇 Myo-Inositol	1.61
29	肌醇磷酸代谢 Inositol phosphate metabolism	D-葡萄糖6-磷酸 D-Glucose 6-phosphate	1.81

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表 6 Na₂CO₃+NaHCO₃胁迫下棉花根差异代谢通路和差异代谢物

Table 6. Differential metabolic pathways and metabolites in cotton roots under Na₂CO₃+NaHCO₃ stress

编号 No.	代谢通路 Metabolic pathway	差异代谢物 Differential metabolite	变化倍数 Fold change
1	异喹啉生物碱的生物合成 Isoquinoline alkaloid biosynthesis	L-酪氨酸 L-Tyrosine	1.30
1	异喹啉生物碱的生物合成 Isoquinoline alkaloid biosynthesis	多巴胺 Dopamine	1.33
2	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	L-天冬酰胺 L-Asparagine	1.23
		L-丙氨酸 L-Alanine	3.29
		L-谷氨酰胺 L-Glutamine	0.72
		L-谷氨酸 L-Glutamate	2.00
		琥珀酸 Succinate	2.91
		4-氨基丁酸 4-Aminobutyric acid	1.16
		琥珀酸半醛 Succinic semialdehyde	1.58
		琥珀酸 Succinate	2.91
3	维生素B6代谢 Vitamin B6 metabolism	吡哆醇 Pyridoxine	2.62
3	维生素B6代谢 Vitamin B6 metabolism	5’-磷酸吡哆醛 Pyridoxal 5’-phosphate	0.76
4	精氨酸生物合成 Arginine biosynthesis	L-谷氨酸 L-Glutamate	2.00
		N-乙酰-L-谷氨酸 N-Acetyl-L-glutamate	0.28
		L-瓜氨酸 L-Citrulline	2.86
		L-谷氨酰胺 L-Glutamine	0.72
5	丁酸代谢 Butanoate metabolism	L-谷氨酸 L-Glutamate	2.00
		琥珀酸 Succinate	2.91
		4-氨基丁酸 4-Aminobutyric acid	1.16
		琥珀酸半醛 Succinic semialdehyde	1.58
6	乙醛酸和二羧酸代谢 Glyoxylate and dicarboxylate metabolism	顺乌头酸 Cis-Aconitate	1.27
		琥珀酸 Succinate	2.91
		L-谷氨酸 L-Glutamate	2.00
		L-谷氨酰胺 L-Glutamine	0.72
		磷酸乙醇酸 Phosphoglycolic acid	0.47
		L-苹果酸 L-Malic acid	4.20
7	戊糖磷酸途径 Pentose phosphate pathway	D-核糖 D-Ribose	2.49
7	戊糖磷酸途径 Pentose phosphate pathway	D-核糖5-磷酸 D-Ribose 5-phosphate	0.15
8	淀粉和蔗糖代谢 Starch and sucrose metabolism	蔗糖 Sucrose	1.34
8	淀粉和蔗糖代谢 Starch and sucrose metabolism	麦芽糖 Maltose	2.45
9	柠檬酸循环(TCA循环) Citrate cycle (TCA cycle)	琥珀酸 Succinate	2.91
		顺乌头酸 Cis-Aconitate	1.27
		L-苹果酸 L-Malic acid	4.20
10	甘油磷脂代谢 Glycerophospholipid metabolism	胆碱 Choline	1.27
10	甘油磷脂代谢 Glycerophospholipid metabolism	甘油磷酸胆碱 Glycerophosphocholine	0.45
11	丙酮酸代谢 Pyruvate metabolism	L-苹果酸 L-Malic acid	4.20
12	半乳糖代谢 Galactose metabolism	水苏糖 Stachyose	4.37
		棉子糖 Raffinose	2.29
		肌醇 Myo-Inositol	1.59
		蔗糖 Sucrose	1.34
13	谷胱甘肽代谢 Glutathione metabolism	L-谷氨酸盐 L-Glutamate	2.00
		谷胱甘肽二硫化物 Glutathione disulfide	1.62
		γ-谷氨酰半胱氨酸 Gamma-Glutamylcysteine	2.73
14	甘氨酸、丝氨酸和苏氨酸代谢 Glycine, serine and threonine metabolism	胆碱 Choline	1.27
		L-苏氨酸 L-Threonine	0.66
		甘油酸 Glyceric acid	2.23
15	泛酸和辅酶A生物合成 Pantothenate and CoA biosynthesis	泛酸 Pantothenate	2.03
16	角质、木栓碱和蜡的生物合成 Cutin, suberine and wax biosynthesis	油酸 Oleic acid	0.47
17	酪氨酸代谢 Tyrosine metabolism	L-酪氨酸 L-Tyrosine	1.30
17	酪氨酸代谢 Tyrosine metabolism	多巴胺 Dopamine	1.33
18	肌醇磷酸代谢 Inositol phosphate metabolism	肌醇 Myo-Inositol	1.59

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表 7 NaCl胁迫下棉花叶差异代谢通路和差异代谢物

Table 7. Differential metabolic pathways and metabolites in cotton leaves under NaCl stress

编号 No.	代谢通路 Metabolic pathway	差异代谢物 Differential metabolite	变化倍数 Fold change
1	异喹啉生物碱生物合成 Isoquinoline alkaloid biosynthesis	L-酪氨酸 L-Tyrosine	1.53
2	丙酮酸代谢 Pyruvate metabolism	L-苹果酸 L-Malic acid	1.43
3	芪类、二芳基庚烷类和姜酚的生物合成 Stilbenoid, diarylheptanoid and gingerol biosynthesis	绿原酸 Chlorogenic acid	1.64
4	氰基氨基酸代谢 Cyanoamino acid metabolism	野樱皮甙 Prunasin	1.26
5	磷酸戊糖途径 Pentose phosphate pathway	6-磷酸-D-葡萄糖酸 6-Phospho-D-gluconate	1.22
6	酪氨酸代谢 Tyrosine metabolism	L-酪氨酸 L-Tyrosine	1.54
7	萜类骨架生物合成 Terpenoid backbone biosynthesis	1-脱氧-D-木酮糖5-磷酸 1-Deoxy-D-xylulose 5-phosphate	1.82
7	萜类骨架生物合成 Terpenoid backbone biosynthesis	(R)-5-磷酸甲羟戊酸 (R)-mevalonic acid 5-Phosphate	1.75

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表 8 Na₂SO₄胁迫下棉花叶差异代谢通路和差异代谢物

Table 8. Differential metabolic pathways and metabolites in cotton leaves under Na₂SO₄ stress

编号 No.	代谢通路 Metabolic pathway	差异代谢物 Differential metabolite	变化倍数 Fold change
1	亚油酸代谢 Linoleic acid metabolism	亚油酸 Linoleic acid	4.24
2	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	L-天冬氨酸 L-Aspartate	2.17
		L-天门冬酰胺 L-Asparagine	0.65
		丙酮酸 Pyruvate	1.34
		氧戊二酸 Oxoglutaric acid	1.48
		L-谷氨酰胺 L-Glutamine	0.79
		L-谷氨酸 L-Glutamate	1.79
		琥珀酸 Succinate	1.42
3	黄酮和黄酮醇的生物合成 Flavone and flavonol biosynthesis	山奈酚 Kaempferol	1.47
4	戊糖磷酸途径 Pentose phosphate pathway	6-磷酸-D-葡萄糖酸 6-Phospho-D-gluconate	1.23
4	戊糖磷酸途径 Pentose phosphate pathway	D-核糖5-磷酸 D-Ribose 5-phosphate	1.52
5	色氨酸代谢 Tryptophan metabolism	2-氧代辛酸 2-Oxoadipic acid	0.51
		L-色氨酸 L-Tryptophan	0.27
		N-乙酰血清素 N-Acetylserotonin	3.14
		吲哚乙酸 Indoleacetic acid	0.4
6	柠檬酸循环(TCA 循环) Citrate cycle (TCA cycle)	琥珀酸 Succinate	1.42
		柠檬酸 Citrate	6.14
		顺乌头酸 Cis-Aconitate	1.39
		丙酮酸 Pyruvate	1.34
		2-氧代辛酸 2-Oxoadipic acid	0.51
7	赖氨酸降解 Lysine degradation	2-氧代辛酸 2-Oxoadipic acid	0.51
7	赖氨酸降解 Lysine degradation	糖苷 Saccharopine	0.52
8	苯丙氨酸代谢 Phenylalanine metabolism	苯乙酸 Phenylacetic acid	1.15
		苯乙胺 Phenylethylamine	0.68
		反式肉桂酸酯 Trans-cinnamate	1.20
9	精氨酸生物合成 Arginine biosynthesis	2-氧代辛酸 2-Oxoadipic acid	0.51
		L-谷氨酸 L-Glutamate	1.79
		L-谷氨酰胺 L-Glutamine	0.79
		L-天冬氨酸 L-Aspartate	2.17
		N2-乙酰-L-鸟氨酸 N2-Acetyl-L-ornithine	0.57
10	半乳糖代谢 Galactose metabolism	α-D-葡萄糖 Alpha-D-Glucose	0.64
		水苏糖 Stachyose	0.74
		棉子糖 Raffinose	2.67
		蔗糖 Sucrose	1.21
11	乙醛酸和二羧酸代谢 Glyoxylate and dicarboxylate metabolism	顺乌头酸 Cis-Aconitate	1.39
		柠檬酸 Citrate	6.14
		琥珀酸 Succinate	1.42
		L-谷氨酸 L-Glutamate	1.79
		L-谷氨酰胺 L-Glutamine	0.79
		2-氧代辛酸 2-Oxoadipic acid	0.51
		甘油酸 Glyceric acid	1.28
12	丙酮酸代谢 Pyruvate metabolism	丙酮酸 Pyruvate	1.34
13	甘氨酸、丝氨酸和苏氨酸代谢 Glycine, serine and threonine metabolism	L-天冬氨酸 L-Aspartate	2.17
		甘油酸 Glyceric acid	1.28
		L-苏氨酸 L-Threonine	1.52
		L-色氨酸 L-Tryptophan	0.27
		丙酮酸 Pyruvate	1.34
14	芪类、二芳基庚烷类和姜辣素生物合成 Stilbenoid, diarylheptanoid and gingerol biosynthesis	绿原酸 Chlorogenic acid	1.79
15	糖酵解/糖异生 Glycolysis / Gluconeogenesis	α-D-葡萄糖 Alpha-D-Glucose	0.64
15	糖酵解/糖异生 Glycolysis / Gluconeogenesis	丙酮酸 Pyruvate	1.34
16	缬氨酸、亮氨酸和异亮氨酸的生物合成 Valine, leucine and isoleucine biosynthesis	L-苏氨酸 L-Threonine	1.52
16		丙酮酸 Pyruvate	1.34

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表 9 Na₂CO₃+NaHCO₃胁迫下棉花叶差异代谢通路和差异代谢物

Table 9. Differential metabolic pathways and metabolites in cotton leaves under Na₂CO₃+NaHCO₃ stress

编号 No.	代谢通路 Metabolic pathway	差异代谢物 Differential metabolite	变化倍数 Fold change
1	亚油酸代谢 Linoleic acid metabolism	亚油酸 Linoleic acid	3.47
2	黄酮和黄酮醇生物合成 Flavone and flavonol biosynthesis	山奈酚 Kaempferol	2.87
3	戊糖磷酸途径 Pentose phosphate pathway	6-磷酸-D-葡萄糖酸 6-Phospho-D-gluconate	0.58
3	戊糖磷酸途径 Pentose phosphate pathway	D-核糖5-磷酸 D-Ribose 5-phosphate	1.50
4	丙氨酸、天冬氨酸和谷氨酸代谢 Alanine, aspartate and glutamate metabolism	L-天冬氨酸 L-Aspartate	1.95
		L-谷氨酰胺 L-Glutamine	0.57
		琥珀酸 Succinate	0.35
5	硫胺素代谢 Thiamine metabolism	一磷酸硫胺素 Thiamine monophosphate	0.36
6	赖氨酸降解 Lysine degradation	2-氧代辛酸 2-Oxoadipic acid	0.35
6	赖氨酸降解 Lysine degradation	糖苷 Saccharopine	0.59
7	精氨酸和脯氨酸代谢 Arginine and proline metabolism	L-精氨酸 L-Arginine	2.59
		羟脯氨酸 Hydroxyproline	1.83
		腐胺 Putrescine	1.58
8	色氨酸代谢 Tryptophan metabolism	2-氧代辛酸 2-Oxoadipic acid	0.35
		N-乙酰血清素 N-Acetylserotonin	8.23
		吲哚乙酸 Indoleacetic acid	0.36

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