低温循环诱导对日光温室黄瓜光合作用及产量的影响

doi:10.16420/j.issn.0513-353x.2024-0868

摘要/Abstract

摘要：

低温诱导的胁迫记忆作为植物应对环境变化的可塑性行为在提高植物对逆境的适应性中发挥重要作用。为探明低温循环诱导的胁迫记忆对日光温室黄瓜光合作用的环境适应性调控机制，以‘津优35号’黄瓜为试材，在人工气候室内进行处理。设未进行低温诱导（C0R0）、低温诱导1次（C1R1：昼/夜温度8 ℃/8 ℃处理24 h→25 ℃/18 ℃恢复48 h）、循环诱导2次（C2R2：低温和恢复条件同上）、循环诱导3次（C3R3：低温和恢复条件同上）共4个处理，诱导结束后均置于8 ℃/5 ℃下，48 ~ 72 h后取样测定光合作用相关指标；同时以常温（25 ℃/18 ℃）作为对照（CK）。在此基础上，选择低温循环诱导效果最佳的处理和C0R0的幼苗栽于日光温室中，研究低温循环诱导的胁迫记忆对冬季日光温室黄瓜生长及产量的影响。结果表明：低温诱导处理（C1R1、C2R2、C3R3）黄瓜幼苗的净光合速率（P_n）、气孔导度（G_s）、蒸腾速率（T_r）、核酮糖-1,5-二磷酸羧化酶（RuBPCase）、Rubisco活化酶（RCA）活性与mRNA表达，以及PSⅡ最大光化学效率（F_v/F_m）、PSⅡ实际光化学效率（Φ_PSⅡ）、PSⅡ电子传递速率[ETR(Ⅱ)]、光下PSⅡ实际光化学量子效率[Y(Ⅱ)]、PSⅠ电子传递速率[ETR(Ⅰ)]和PSⅠ有效光化学量子产量[Y(Ⅰ)]均显著高于C0R0，而胞间CO₂浓度（C_i）明显低于C0R0或与C0R0差异不显著，其中以C2R2处理的P_n、G_s、T_r、F_v/F_m、Φ_PSⅡ及RuBPCase和RCA活性与mRNA表达的增加幅度和C_i降低幅度最大，C1R1的最小；低温下C2R2处理的日光温室黄瓜P_n、F_v/F_m、Φ_PSⅡ也显著高于C0R0。可见，低温循环诱导通过维持低温下叶片较高的光合碳同化能力与光系统活性，减轻低温对黄瓜叶片光合机构的损伤，从而促进植株的生长发育，日光温室中产量比C0R0高10.25%。2次低温循环诱导的胁迫记忆时间为20 ~ 30 d。

关键词: 黄瓜, 低温循环诱导, 胁迫记忆, 光合作用, 产量

Abstract:

Cold-induced stress memory，as a flexible behavior in response to environmental changes，plays an important role in plant acclimation to adverse conditions. To elucidate the regulating mechanism of cold circulation-induced stress memory on the environmental adaptability of photosynthesis in cucumber in solar greenhouse.‘Jinyou 35’cucumber plants were used as experimental materials，and were treated in an artificial climate chamber. There were four treatments： no cold induction treatment（C0R0），single cold induction（C1R1：24 h treatment at a day/night temperature of 8 ℃/8 ℃ followed by a recovery period of 48 h at 25 ℃/18 ℃），repeated cold circulation induction twice（C2R2：same conditions as C1R1，cycle twice），and repeated cold circulation induction three times（C3R3：same conditions as C1R1 treatment，cycle three times）. Subsequently，the seedlings were exposed to chilling stress condition（8 ℃/5 ℃）. The C0R0 treatment at normal temperature（25 ℃/18 ℃）was used as the control（CK）. The relevant indexes of photosynthesis were measured after 48-72 h of low temperature. The seedlings exhibiting the most favorable cold circulation-induced effect and C0R0 treatments were then selected and cultivated within a solar greenhouse to investigate the effect of cold-induce memory on the photosynthetic rate（P_n），photochemical efficiency，growth and yield. The results showed that C1R1-，C2R2- and C3R3-treated seedlings exhibited significantly higher P_n，stomatal conductance（G_s），transpiration rate（T_r），as well as activities and mRNA expressions of ribulose-1,5-bisphosphate carboxylase（RuBPCase）and Rubisco activase（RCA）. Additionally，the C1R1，C2R2 and C3R3 treatments revealed an increase in maximum photochemical efficiency of PSⅡ（F_v/F_m），actual photochemical effect of PSⅡ（Φ_PSII），electron transfer rate of PSⅡ[ETR(II)]，quantum efficiency of PSⅡ photochemistry[Y(Ⅱ)]，electron transfer rate of PSⅠ[ETR(Ⅰ)]and quantum efficiency of PSⅠ photochemistry[Y(Ⅰ)]. Whereas，there were either lower or no significant differences observed in intercellular CO₂ concentration（C_i），compared with the C0R0 treatment under chilling stress. Among the four treatments，C2R2-treated plants exhibited the greatest increase in P_n，G_s，T_r，F_v/F_m，Φ_PSⅡ，activity and mRNA expression of RuBPCase and RCA，and decrease in C_i，while C1R1 treated seedlings showed the least changes in these parameters. The P_n，F_v/F_m and Φ_PSⅡof C2R2 treated cucumber grown in the solar greenhouse were significantly higher than those of the C0R0 at low temperatures. These findings indicate that the cold circulation induction maintain higher photosynthetic carbon assimilation capacity and photosystem activity，alleviate the damage to photosynthetic apparatus caused by chilling stress，thereby improve the growth and development of cucumber. The yield of C2R2 treatment was 10.25% higher than that of C0R0. The stress memory induced by two cycles of cold circulation in cucumber plants was between 20 and 30 days.

Key words: cucumber, cold circulation induction, stress memory, photosynthesis, yield

赵猛, 艾希珍, 戚成正, 毕焕改, 冯一清, 张晓伟. 低温循环诱导对日光温室黄瓜光合作用及产量的影响[J]. 园艺学报, 2025, 52(10): 2748-2760.

ZHAO Meng, AI Xizhen, QI Chengzheng, BI Huangai, FENG Yiqing, ZHANG Xiaowei. Effects of Cold Circulation Induction on Photosynthesis and Yield of Cucumber in Solar-Greenhouse[J]. Acta Horticulturae Sinica, 2025, 52(10): 2748-2760.

图/表 10

表1 引物序列

Table 1 Supplementary table primer sequences

基因 Gene	引物序列（5′-3′） Primer sequences
β-actin	F：CCACGAAACTACTTACAACTCCATC；R：GGGCTGTGATTTCCTTGCTC
rbcL	F：GCTATGGAATCGAGCCTGTTG；R：CCAAATACATTACCCACAATGGAAG
RCA	F：AAAGTGGGCTGTAGGCGTTG；R：CTTTTCTATTGTCATCTTCGGTTGG

图1 低温循环诱导次数对冷胁迫下黄瓜幼苗色素含量的影响 CK：常温对照；C0R0：未低温诱导；C1R1：低温诱导1次（8 ℃/8 ℃处理24 h→25 ℃/18 ℃恢复48 h）；C2R2：低温诱导2次；C3R3：低温诱导3次。图中数据为3次重复结果均值 ± 标准差，不同小写字母表示不同处理间差异显著（P < 0.05）；下同

Fig. 1 Effect of cold circulation induction times on the pigment contents in cucumber seedlings under chilling stress CK：25 ℃/18 ℃，the control；C0R0：No cold induction treatment；C1R1：Single cold induction（24 h treatment at a day/night temperature of 8 ℃/8 ℃ followed by a recovery period of 48 h at 25℃/18 ℃）；C2R2：Repeated cold circulation induction twice（same conditions as C1R1，cycle twice）；C3R3：Repeated cold circulation induction three times（same conditions as C1R1 treatment，cycle three times）. The present data are mean ± standard deviation of three biological replicates. Different lowercase letters indicate significant differences between treatments（P < 0.05），the same below

图2 低温循环诱导次数对冷胁迫下黄瓜幼苗净光合速率和气体交换参数的影响

Fig. 2 Effect of cold circulation induction times on Pn and gas exchange parameters of cucumber seedlings under chilling stress

图3 低温循环诱导次数对冷胁迫下黄瓜幼苗Fv/Fm、ΦPSⅡ的影响

Fig. 3 Effect of cold circulation induction times on Fv/Fm，ΦPSⅡ of cucumber seedlings under chilling stress

图4 低温循环诱导次数对冷胁迫下黄瓜幼苗ETR (Ⅰ)、ETR (Ⅱ)、Y (Ⅰ)、Y (Ⅱ)的影响

Fig. 4 Effect of cold circulation induction times on ETR (Ⅰ)，ETR (Ⅱ)，Y (Ⅰ)，Y (Ⅱ) of cucumber seedlings under chilling stress

图5 低温循环诱导次数对冷胁迫下黄瓜幼苗卡尔文循环关键酶活性与mRNA表达的影响

Fig. 5 Effect of cold circulation induction times on the activity and mRNA expression of key enzymes in Calvin cycle in cucumber seedlings under chilling stress

图6 日光温室光合光量子通量密度日变化（A）与气温昼夜变化（B）

Fig. 6 Diurnal variation of PPFD（A）and air temperature（B）in solar-greenhouse

图7 低温循环诱导对日光温室黄瓜叶片光合速率与光化学效率的影响

Fig. 7 Effect of cold circulation induction on photosynthetic rate and photochemical efficiency of cucumber leaves in solar greenhouse

表2 低温循环诱导对不同时期日光温室黄瓜生长量的影响

Table 2 Effect of cold circulation induction on the growth of cucumber in solar greenhouse

日期/（Y-M-D） Date	处理 Treatment	株高/cm Plant height	茎粗/mm Stem diameter	单株叶面积/cm² Leaf area per plant
2023-12-27	C0R0	17.09 ± 1.396 f	7.92 ± 0.30 c	616.07 ± 34.41 f
2023-12-27	C2R2	27.00 ± 1.04 e	8.31 ± 0.10 b	866.67 ± 54.76 e
2024-01-07	C0R0	50.20 ± 3.73 c	8.28 ± 0.48 b	1 696.57 ± 115.38 d
2024-01-07	C2R2	61.55 ± 3.62 b	9.64 ± 0.64 a	2 106.29 ± 170.80 c
2024-01-17	C0R0	104.63 ± 8.61 a	9.46 ± 0.55 a	3 269.93 ± 284.48 b
2024-01-17	C2R2	109.67 ± 9.14 a	9.86 ± 0.79 a	4 057.94 ± 474.06 a

表3 低温循环诱导对日光温室黄瓜产量的影响

Table 3 Effect of cold circulation induction on the yield of cucumber in solar greenhouse

处理 Treatment	单果质量/g Fruit weight	单株果数 Fruits per plant	产量/（kg · hm^-2） Yield	增产/% Yield increase
C0R0	206.10 ± 6.16 a	4.18 ± 0.25 b	15 502.25 ± 1 380.9 b	—
C2R2	200.04 ± 9.75 a	4.75 ± 0.14 a	17 091.00 ± 324.60 a	10.25

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