Scientia Agricultura Sinica ›› 2022, Vol. 55 ›› Issue (7): 1399-1410.doi: 10.3864/j.issn.0578-1752.2022.07.011

• HORTICULTURE • Previous Articles     Next Articles

Effect of Exogenous Spermidine on Nitrogen Metabolism of Lettuce Under High-Temperature Stress

YU QiLong1(),HAN YingYan1,HAO JingHong1,QIN XiaoXiao1,LIU ChaoJie1(),FAN ShuangXi2()   

  1. 1College of Plant Science and Technology, Beijing University of Agriculture, Beijing 102206
    2Beijing Vocational College of Agriculture, Beijing 102442
  • Received:2021-07-02 Accepted:2021-09-09 Online:2022-04-01 Published:2022-04-18
  • Contact: ChaoJie LIU,ShuangXi FAN;;


【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

Table 1

Primer sequence information of genes of interest used in qPCR assay"

基因名称Gene name 上游引物 Forward primer (5′-3′) 下游引物 Reverse primer (5′-3′)

Table 2

Effects of exogenous Spd on the morphological indexes of lettuce under high-temperature stress"

处理Treatment 总鲜重
Total fresh weight (g/plant)
Total dry weight (g/plant)
Leaf length
Leaf width
Root length
Root volume
Root surface area
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

Fig. 1

The effect of exogenous Spd on the flow rate of NH4+ and NO3- ions in the roots of lettuce under high temperature stress The ion velocity value is 2.5 mm from the apex of the root tip. A positive value represents external discharge, and a negative value represents internal flow. Different lowercase letters indicate significant difference (P<0.05). The same as below"

Fig. 2

Effects of exogenous Spd on total nitrogen, nitrate nitrogen and ammonia nitrogen of lettuce under high-temperature stress"

Fig. 3

Effects of exogenous Spd on activities of key enzymes of nitrogen metabolism in lettuce under high-temperature stress"

Fig. 4

Effects of exogenous spermidine on gene expression of key enzymes in nitrogen metabolism of lettuce under high-temperature stress"

Table 3

Effects of exogenous Spd on the content of free amino acids in lettuce leaves under high temperature stress"

Free amino acid composition (μg·g-1)
处理 Treatment 含量均值
Average content (μg·g-1)
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

Table 4

Effects of exogenous Spd on the content of flavor amino acids in lettuce leaves under high temperature stress"

Flavor amino acid composition (μg·g-1)
处理 Treatment 含量均值
Average content (μg·g-1)
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
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