外源亚精胺对高温胁迫下生菜氮代谢的影响

doi:10.3864/j.issn.0578-1752.2022.07.011

摘要/Abstract

摘要：

【目的】在高温季节种植的生菜产量和营养品质不佳,而与生菜产量最密切的营养元素是氮。研究外源亚精胺（Spd）在生菜抗高温胁迫中的氮代谢调控机理,为降低高温对生菜产生的危害提供保护措施。【方法】试验材料为散叶生菜‘北散生3号',试验处理设置常温对照（昼/夜温度：22℃/17℃,CK）、高温胁迫处理（昼/夜温度：35℃/30℃,H）、高温喷施Spd处理（HS）。观察3个不同处理生菜的生长形态变化,研究生菜根系离子流速、含氮化合物（总氮、氨态氮、硝态氮）含量、氮代谢关键酶活性及基因相对表达水平的变化。此外,分析CK、H和HS处理对生菜叶片游离氨基酸含量的影响。【结果】与对照CK相比,生菜生长在H处理受到抑制;与H处理相比,HS处理的生菜总鲜重、总干重、叶长、叶宽、根长、根系体积和根系表面积均有所增加,分别增加了24.00%、24.62%、14.97%、11.83%、23.24%、29.47%和36.98%。CK、H和HS处理的生菜总氮含量变化与生菜的生长趋势一致。CK、H和HS处理的生菜根系NO₃^-净吸收量远小于NH₄⁺,即生菜根系主要以吸收NH₄⁺为主;H处理使根系NO₃^-外排而HS处理促进根系NO₃^-内流。HS处理有效抑制高温导致的叶片和根系氨态氮含量的积累。同时,与H处理相比,HS处理增加了叶片和根系的硝酸还原酶（NR）、谷氨酰胺合酶（GS）、谷氨酸合酶（GOGAT）、谷氨酸脱氢酶（GDH）的活性及上调LsNR、LsGS、LsGOGAT1、LsGDH的相对表达水平。与CK相比,H处理的总游离氨基酸（TAA）含量显著减少（P<0.05）;HS处理缓解了高温导致的TAA含量减少。处理CK、H和HS之间的鲜味氨基酸（UAA）、甜味氨基酸（SAA）和苦味氨基酸（BAA）的变化趋势与TAA一致,与其呈相反趋势的是芳香族氨基酸（AAA）。处理CK、H和HS之间的天冬氨酸（Asp）、苏氨酸（Thr）、谷氨酸（Glu）、甘氨酸（Gly）、缬氨酸（Val）、赖氨酸（Lys）、组氨酸（His）、精氨酸（Arg）和脯氨酸（Pro）这9种氨基酸与TAA的变化趋势一致。其中,检测的游离氨基酸中变异系数最大的是Arg。【结论】高温喷施Spd处理抑制了高温导致的生菜生物量积累的降低。与高温处理相比,高温喷施Spd处理缓解了高温胁迫导致的氮代谢紊乱,增加了叶片总游离氨基酸含量和呈味氨基酸中的甜味氨基酸、鲜味氨基酸、苦味氨基酸含量,有效地改善了生菜的耐高温能力,提升了高温胁迫下生菜的营养品质。

关键词: 高温, 生菜, 氮代谢, 亚精胺, 离子流速, 氨基酸

Abstract:

【Objective】 The lettuce yield and nutritional quality planted in high-temperature seasons are not good. Nitrogen is the most closely related nutrient element to lettuce yield. In this study, the regulation mechanism of exogenous spermidine (Spd) on nitrogen metabolism of lettuce under high-temperature stress was studied, and the changes of amino acid nutritional quality were observed, so as to provide some protective measures to reduce the harm of high temperature on lettuce. 【Method】 The test material was leaf lettuce Beisansheng 3. The test treatments were set as normal temperature (day/night: 22℃/17℃) control (CK), high-temperature stress (day/night: 35℃/30℃) treatment (H), and high temperature spraying Spd treatment (HS). The changes of lettuce morphology were observed, and the changes of the lettuce root ion flow rate, nitrogen compounds contents (total nitrogen, ammonia nitrogen, and nitrate nitrogen), and the activities of key enzymes in nitrogen metabolism as well as their relative expression levels were analyzed. In addition, the effects of the CK, H, and HS treatments on the lettuce leaf free amino acids content were studied. 【Result】 Compared with the CK, the lettuce growth of the H treatment was inhibited. Compared with the H treatment, the HS treatment increased total fresh weight, total dry weight, leaf length, leaf width, root length, root volume, and root surface area of lettuce by 24.00%, 24.62%, 14.97%, 11.83%, 23.24%, 29.47%, and 36.98%, respectively. The changes in the total nitrogen content of lettuce treated by CK, H, and HS were consistent with the growing trend of lettuce. The net absorption amount of NO₃^- in lettuce roots treated with CK, H and HS were much smaller than that of NH₄⁺, indicating that lettuce roots mainly absorbed NH₄⁺. The H treatment resulted in the outflow of NO₃^- from roots, while the HS treatment promoted the influx of NO₃^- from roots. The accumulation of ammonia nitrogen content in leaves under the HS treatment was effectively inhibited. Meanwhile, the activities of nitrate reductase (NR), glutamine synthase (GS), glutamate synthase (GOGAT), and glutamate dehydrogenase (GDH) under the HS treatment were increased and the relative gene expression levels of LsNR, LsGS, LsGOGAT1, and LsGDH in leaves and roots were up-regulated. Compared with the CK treatment, the H treatment reduced the total free amino acid (TAA) content (P<0.05); the HS treatment alleviated the reduction in TAA content caused by high temperature. The changes of umami amino acids (UAA), sweet amino acids (SAA), and bitter amino acids (BAA) among the CK, H, and HS treatment were consistent with those of TAA, while aromatic amino acids (AAA) were the opposite. The changes of aspartic acid (Asp), threonine (Thr), glutamic acid (Glu), glycine (Gly), valine (Val), lysine (Lys), histidine (His), arginine (Arg), and proline (Pro) among the CK, H, and HS treatment were consistent with those of TAA. Among the free amino acids tested, Arg had the largest coefficient of variation. 【Conclusion】Spraying Spd treatment inhibited the decrease of lettuce biomass accumulation under high temperature. For spraying Spd treatment, the nitrogen metabolism disorders under high-temperature stress was alleviated, and the leaf total free amino acid content, flavor amino acids of umami amino acids, sweet amino acids, and bitter amino acids were increased, while the high-temperature resistant capacity of lettuce was effectively enhanced and the nutritional quality of lettuce was improved.

Key words: high temperature, Lactuca sativa L., nitrogen metabolism, spermidine, ion flow rate, amino acid

余琦隆,韩莹琰,郝敬虹,秦晓晓,刘超杰,范双喜. 外源亚精胺对高温胁迫下生菜氮代谢的影响[J]. 中国农业科学, 2022, 55(7): 1399-1410.

YU QiLong,HAN YingYan,HAO JingHong,QIN XiaoXiao,LIU ChaoJie,FAN ShuangXi. Effect of Exogenous Spermidine on Nitrogen Metabolism of Lettuce Under High-Temperature Stress[J]. Scientia Agricultura Sinica, 2022, 55(7): 1399-1410.

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基因名称Gene name	上游引物 Forward primer (5′-3′)	下游引物 Reverse primer (5′-3′)
LsNR	CTCCGAGCCACCTTTGACTAAGC	CGCATATCTCCACCGTCCAATCC
LsGS	CTTCGCCACGCAGACCACATC	ATGGAGCAACCACGGTTAGCAAC
LsGOGAT	GCAGTTCGTGAGAAGCGTGAGAG	TGAAGCAGCGGATTCCACCTTTC
LsGOGAT1	CGTGAGGAAGATGTGCTGAAGGAG	GATGGTCGGGTTGCCTGTTCG
LsGDH	CCGTATGGTGGTGCGAAAGGTG	GAGCAGGAACATCGGTGTGGATTC
18S	GTGAGTGAAGAAGGGCAATG	CACTTTCAACCCGATTCACC

处理Treatment	总鲜重 Total fresh weight (g/plant)	总干重 Total dry weight (g/plant)	叶长 Leaf length (cm)	叶宽 Leaf width (cm)	根长 Root length (cm)	根系体积 Root volume (cm³)	根系表面积 Root surface area (cm²)
CK	15.60±0.49a	0.65±0.04b	13.78±0.25c	11.08±0.46a	24.30±0.98a	5.75±0.16a	281.96±10.49a
H	10.96±0.38c	0.65±0.73b	14.90±0.31b	7.86±0.24c	18.50±0.05c	3.19±0.35c	169.06±15.86c
HS	13.59±0.38b	0.81±0.11a	17.13±0.40a	8.79±0.28b	22.80±2.25b	4.13±0.40b	231.57±13.52b

游离氨基酸组成 Free amino acid composition (μg·g^-1)	处理 Treatment			含量均值 Average content (μg·g^-1)	变异系数 CV (%)
游离氨基酸组成 Free amino acid composition (μg·g^-1)	CK	H	HS	含量均值 Average content (μg·g^-1)	变异系数 CV (%)
天冬氨酸Asp	177.80±15.18a	100.46±10.14b	187.29±6.02a	155.18	30.69
苏氨酸Thr	528.62±6.69a	221.80±51.22c	332.73±24.36b	361.05	43.03
丝氨酸Ser	263.18±2.87a	195.94±5.54b	159.78±0.89c	206.30	25.44
谷氨酸Glu	949.11±46.19a	739.63±14.82c	839.33±32.91b	842.69	12.43
甘氨酸Gly	6.53±0.03a	4.81±0.47b	6.31±0.51a	5.88	15.91
丙氨酸Ala	128.55±1.59a	104.18±7.89b	85.38±0.93c	106.04	20.41
胱氨酸Cys	6.25±0.20a	6.39±0.16a	6.33±0.06a	6.32	1.11
缬氨酸Val	88.98±0.62b	68.14±3.20c	96.66±2.10a	84.59	17.45
蛋氨酸Met	2.90±0.04a	2.69±0.13a	2.88±0.13a	2.82	4.11
异亮氨酸Ile	17.32±0.05a	15.89±0.35b	15.41±0.6b	16.21	6.13
亮氨酸Leu	10.79±0.47a	6.77±0.22b	6.64±0.12b	8.07	29.25
酪氨酸Tyr	14.13±0.12b	16.28±0.52a	13.30±0.33c	14.57	10.56
苯丙氨酸Phe	23.63±0.08b	24.52±1.10ab	25.02±0.11a	24.39	2.89
赖氨酸Lys	14.10±0.24a	12.33±0.03b	11.55±0.36c	12.66	10.32
组氨酸His	9.50±0.21b	6.81±0.59c	12.60±1.76a	9.64	30.07
精氨酸Arg	60.55±0.71b	42.28±7.97c	101.88±7.04a	68.24	44.75
脯氨酸Pro	1085.58±30.69a	899.09±8.30c	943.91±4.74b	976.19	9.97
总氨基酸TAA	3387.54±44.14a	2468.01±63.91c	2847.01±37.26b	2900.85	15.93

呈味氨基酸组成 Flavor amino acid composition (μg·g^-1)	处理 Treatment			含量均值 Average content (μg·g^-1)	变异系数 CV (%)
呈味氨基酸组成 Flavor amino acid composition (μg·g^-1)	CK	H	HS	含量均值 Average content (μg·g^-1)	变异系数 CV (%)
鲜味氨基酸UAA	1276.09±34.67a	961.42±9.65c	1129.86±35.01b	1122.46	14.03
甜味氨基酸SAA	1886.89±31.66a	1323.64±55.74c	1449.03±28.59b	1553.19	19.04
芳香族氨基酸AAA	44.01±0.18b	47.19±1.56a	44.66±0.49b	45.29	3.71
苦味氨基酸BAA	180.54±1.36b	135.76±10.90c	223.47±9.43a	179.92	24.38