青藏高原高山嵩草光合功能对模拟夜间低温的响应

doi:10.17521/cjpe.2021.0446

植物生态学报 ›› 2023, Vol. 47 ›› Issue (3): 361-373.DOI: 10.17521/cjpe.2021.0446 cstr: 32100.14.cjpe.2021.0446

所属专题：青藏高原植物生态学：生理生态学；光合作用

青藏高原高山嵩草光合功能对模拟夜间低温的响应

师生波¹^,³^,^*(), 周党卫¹, 李天才¹, 德科加², 杲秀珍¹, 马家麟¹, 孙涛³, 王方琳³

¹中国科学院西北高原生物研究所高原生物适应与进化重点实验室, 西宁 810001
²青海大学畜牧兽医科学院, 西宁 810016
³甘肃省治沙研究所甘肃省荒漠化与风沙灾害重点实验室, 兰州 730070

收稿日期:2021-12-02 接受日期:2022-05-21 出版日期:2023-03-20 发布日期:2022-10-11
作者简介:* E-mail: sbshi@nwipb.cas.cn
基金资助:
青海省自然科学基金(2019-ZJ-7016);青海省创新平台建设专项(2017-ZJ-Y20);青海省创新平台建设专项(2021-ZJ-Y05);国家自然科学基金(31660237)

Responses of photosynthetic function of Kobresia pygmaea to simulated nocturnal low temperature on the Qingzang Plateau

SHI Sheng-Bo¹^,³^,^*(), ZHOU Dang-Wei¹, LI Tian-Cai¹, DE Ke-Jia², GAO Xiu-Zhen¹, MA Jia-Lin¹, SUN Tao³, WANG Fang-Lin³

¹Key Laboratory of Adaptation and Evolution of Plateau Biology, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining 810001, China
²Qinghai Academy of Animal Science and Veterinary Medicine, Qinghai University, Xining 810016, China
³State Key Laboratory Breeding Base of Desertification and Aeolian Sand Disaster Combating, Gansu Desert Control Research Institute, Lanzhou 730070, China

Received:2021-12-02 Accepted:2022-05-21 Online:2023-03-20 Published:2022-10-11
Supported by:
Qinghai Province Naturnal Science Foundation(2019-ZJ-7016);Construction Project for Innovation Platform of Qinghai Province(2017-ZJ-Y20);Construction Project for Innovation Platform of Qinghai Province(2021-ZJ-Y05);National Natural Science Foundation of China(31660237)

摘要/Abstract

摘要：

昼夜温差大是青藏高原的典型气候特征, 夜间低温作为植物生长季内非常频繁的非生物胁迫因子, 对典型高山植物日间光合生理功能的影响如何, 尚缺乏研究。该研究以采自青海大学-清华大学三江源高寒草地生态系统野外观测站的高山嵩草(Kobresia pygmaea)为材料, 应用叶绿素荧光图像分析手段, 研究了模拟夜间低温对叶片光系统II (PSII)非光化学猝灭中光诱导和非光诱导的量子产量, 及慢弛豫相和快弛豫相组分的影响。结果表明: 0 ℃夜间低温对日间PSII相对电子传递速率、PSII反应中心开放比率(q_L)和PSII非光化学猝灭系数(q_NP)的快速光响应曲线影响较小; 400和1 500 µmol·m^-2·s^-1稳态作用光强下的比较证实, 夜间低温并没有影响到光合机构活性及非光化学能量耗散过程。PSII反应中心激发能分配的量子通量分析表明, PSII实际光化学量子效率、PSII非光化学猝灭中非调节性和调节性能量耗散量子产量的相对比率在第3天高光强下, 对照组和夜间低温组分别为: 36:19:45和38:19:43; 较低光强下为66:22:12和66:23:11。非光化学猝灭(NPQ)中快弛豫相(NPQ_f)为主要组分, 而慢弛豫相(NPQ_s)所占份额(NPQ_s/NPQ)在对照植株的第1天和第3天分别为11%和10%, 夜间低温组则为13%和12%。因此, 0 ℃夜间低温后, 高山嵩草PSII反应中心发生光抑制的几率增大, 较低光强和夜间低温能导致光合诱导时间的延长; 但光化学能量转换和保护性的调节机制尚能有效分配吸收的光能, 夜间低温没有加剧过剩激发能难以调节耗散的趋势。

关键词: 夜间低温, 光系统II非光化学猝灭, 高山嵩草, 青藏高原, 叶绿素荧光

Abstract:

Aims Kobresia pygmaea is a perennial cushion herb from the Cyperaceae family with a height of 1-3 cm and small linear leaves about 1 mm wide. It is mainly distributed on the low slopes of the high mountains ranging from 3 000 m to 5 960 m on the Qingzang Plateau. Its habitat is harsh, and extreme climate conditions such as low temperature, strong wind, and high sunlight intensity are the main abiotic stresses during plants growing season. The objectives of this study were to analyze the photochemical and non-photochemical energy distribution of the photosystem II (PSII) reaction center in K. pygmaea leaves, and their quenching protection mechanism after nocturnal low-temperature (NLT) treatment.

Methods Kobresia pygmaea meadow turfs (30 cm × 15 cm) were collected from the Alpine Grassland Ecosystem Research Station of the Resource of Three Rivers. The turf blocks were separated into two groups, one group was kept in a culture room with a temperature of 24/18 °C (day/night) as a control treatment, and another was kept in an artificial climate chamber with 0 °C in the evening as an NLT treatment. During the daytime, the NLT group was moved back to the culture room and irradiated together with the control group. On day 0, day 1, day 3, and day 5 after NLT treatment, the chlorophyll fluorescence of K. pygmaea leaves including, the light-response curve, PSII photochemical efficiency at 400 and 1 500 μmol·m⁻²·s⁻¹ steady-state light intensities, and dark relaxation were monitored using CF imager. Then, based on the “Lake Model”, the variation of the PSII actual photochemical efficiency (Φ_PSII), the quantum yield of non-regulated energy dissipation (Φ_NO) and regulated energy dissipation (Φ_NPQ) were explored. Additionally, the fast and slow relaxation components of PSII non-photochemical quenching were determined.

Important findings Nocturnal low temperature had limited effects on the rapid light-response curves of PSII relative electron transfer rate through PSII (rETR), the fraction of open PSII centers (q_L), and PSII non-photochemical quenching coefficient (q_NP). The comparison of chlorophyll fluorescence between 400 and 1 500 μmol·m⁻²·s⁻¹ steady-state light intensities confirmed that NLT treatment did not affect the activity of the PSII reaction center and the process of non-photochemical quenching of K. pygmaea. On the third day after NLT treatment, under high light intensity, the ratios of Φ_PSII:Φ_NO:Φ_NPQ were 36:19:45 and 38:19:43 in the control and NLT groups, respectively; while under lower light intensity, they were 66:22:12 and 66:23:11, respectively. The fast relaxation component (NPQ_f) was the main component in non-photochemical quenching (NPQ); the proportion of the slow relaxation component in non-photochemical quenching was 11% and 10% on day 1 and day 3 in control group, and 13% and 12% in NLT group, respectively. Our results indicated that the probability of photoinhibition of the PSII reaction center in K. pygmaeawas increased after NLT treatment; low light intensity and NLT led to the prolongation of photosynthetic induction time. Overall, the NLT treatment did not increase the tendency of excess excitation energy to be difficult to regulate and dissipate in K. pygmaea leaves, since PSII photochemical energy dissipation and protective regulation mechanism still effectively distributed the absorbed light energy.

Key words: nocturnal low temperature, photosystem II non-photochemical quenching, Kobresia pygmaea, Qingzang Plateau, chlorophyll fluorescence

师生波, 周党卫, 李天才, 德科加, 杲秀珍, 马家麟, 孙涛, 王方琳. 青藏高原高山嵩草光合功能对模拟夜间低温的响应. 植物生态学报, 2023, 47(3): 361-373. DOI: 10.17521/cjpe.2021.0446

SHI Sheng-Bo, ZHOU Dang-Wei, LI Tian-Cai, DE Ke-Jia, GAO Xiu-Zhen, MA Jia-Lin, SUN Tao, WANG Fang-Lin. Responses of photosynthetic function of Kobresia pygmaea to simulated nocturnal low temperature on the Qingzang Plateau. Chinese Journal of Plant Ecology, 2023, 47(3): 361-373. DOI: 10.17521/cjpe.2021.0446

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0446

https://www.plant-ecology.com/CN/Y2023/V47/I3/361

图/表 8

图1 高山嵩草叶片光系统II (PSII)反应中心开放比率(qL) (A)、PSII非光化学猝灭系数(qNP) (B)和PSII相对表观电子传递速率(rETR) (C)的快速光响应曲线。PPFD, 光合有效辐射通量密度。CK, 对照; NLT, 夜间低温。

Fig. 1 Rapid light-response curves of the fraction of open PSII centers (qL) (A), PSII non-photochemical quenching coefficient (qNP) (B), and relative electron transfer rate through PSII (rETR) (C) of Kobresia pygmaea leaves. PPFD, photosynthetical active photon flux density. CK, control; NLT, nocturnal low temperature treatment.

图2 稳态作用光强下高山嵩草叶片光系统II相对表观电子传递速率(rETR)的变化及对夜间低温(NLT)的响应(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照组和夜间低温组在不同时长间rETR的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05); *, 对照(CK)组和低温处理组间差异显著(p < 0.05)。

Fig. 2 Relative electron transfer rate through photosystem II (rETR) of Kobresia pygmaea leaves under nocturnal low-temperature (NLT) treatment at steady-state light intensities (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among different days of the treatment in the control and NLT groups, respectively (α = 0.05). ns, no significant difference between the control (CK) and NLT groups (p > 0.05); *, significant difference between the control and NLT groups (p < 0.05).

图3 夜间低温(NLT)对不同稳态光强下高山嵩草叶片光系统II (PSII)反应中心开放比率(qL) (A、B)和PSII非光化学猝灭系数(qNP) (C、D)的影响及光诱导平衡(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照(CK)组和夜间低温组在15 min的光照平衡期间qL和qNP的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05); *, 对照组和低温处理组间差异显著(p < 0.05); **和***, 对照组和低温处理组间差异极显著(p < 0.01和p < 0.001)。

Fig. 3 Fraction of open photosystem II (PSII) centers (qL) (A, B) and PSII non-photochemical quenching coefficient (qNP) (C, D) of Kobresia pygmaea leaves under necturnal low-temperature (NLT) treatment at steady-state light intensities (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among different irradiation times in the control (CK) and NLT groups, respectively (α = 0.05). ns, no significant difference between the control and NLT groups (p > 0.05); *, significant difference between the control and NLT groups (p < 0.05); ** and ***, highly significant difference between the control and NLT groups (p < 0.01 and p < 0.001).

图4 不同时长高山嵩草叶片光系统II (PSII)反应中心开放比率(qL) (A、B)和PSII非光化学猝灭系数(qNP) (C、D)的变化及夜间低温(NLT)的影响(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照(CK)组和夜间低温组在不同处理时长间qL和qNP的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05); *, 对照组和低温处理组间差异显著(p < 0.05); **, 对照组和低温处理组间差异极显著(p < 0.01)。

Fig. 4 Fraction of open photosystem II (PSII) centers (qL) (A, B) and PSII non-photochemical quenching coefficient (qNP) (C, D) of Kobresia pygmaea leaves under different days of nocturnal low-temperature (NLT) treatment (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among different days of the treatment in the control (CK) and NLT groups, respectively (α = 0.05). ns, no significant difference between the control and NLT groups (p > 0.05); *, significant difference between the control group and NLT groups (p < 0.05); **, highly significant difference between the control and NLT groups (p < 0.01).

图5 稳态作用光强下夜间低温(NLT)处理对高山嵩草叶片光系统II (PSII)实际光化学量子效率(ΦPSII) (A、B)、非调节性能量耗散量子产量(ΦNO) (C、D)和调节性能量耗散量子产量(ΦNPQ) (E、F)的影响及随光诱导时间的变化(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照(CK)组和夜间低温组在15 min的光照平衡期间ΦPSII、ΦNO和ΦNPQ的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05); *, 对照组和低温处理组间差异显著(p < 0.05); **和***, 对照组和低温处理组间差异极显著(p < 0.01和p < 0.001)。

Fig. 5 Photosystem II (PSII) actual photochemical efficiency (ΦPSII) (A, B), and the quantum yield of non-regulated energy dissipation (ΦNO) (C, D) and regulated energy dissipation (ΦNPQ) (E, F) of Kobresia pygmaea leaves under different days of nocturnal low-temperature (NLT) treatment (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among different days of the treatment in the control (CK) and NLT groups, respectively (α = 0.05). ns, no significant difference between the control and NLT groups (p > 0.05); *, significant difference between the control and NLT groups (p < 0.05); **, highly significant difference between the control group and NLT groups (p < 0.01).

图6 不同时长夜间低温(NLT)处理高山嵩草叶片光系统II (PSII)实际光化学量子效率(ΦPSII) (A、B)、非调节性能量耗散量子产量(ΦNO) (C、D)和调节性能量耗散量子产量(ΦNPQ) (E、F)的变化及对夜间低温的响应(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照(CK)组和夜间低温组在不同处理时长间ΦPSII、ΦNO和ΦNPQ的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05); *, 对照组和低温处理组间差异显著(p < 0.05); **, 对照组和低温处理组间差异极显著(p < 0.01)。

Fig. 6 Photosystem II (PSII) actual photochemical efficiency (ΦPSII) (A, B), and the quantum yield of non-regulated energy dissipation (ΦNO) (C, D) and regulated energy dissipation (ΦNPQ) (E, F) of Kobresia pygmaea leaves under nocturnal low-temperature (NLT) treatment at steady-state light intensities (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among irradiation times in the control (CK) and NLT groups, respectively (α = 0.05). ns, no significant difference between the control and NLT groups (p > 0.05); *, significant difference between the control and NLT groups (p < 0.05); ** and ***, highly significant difference between the control and NLT groups (p < 0.01 and p < 0.001).

图7 夜间低温(NLT)对高山嵩草叶片光系统II (PSII)非光化学猝灭(NPQ) (A)和快弛豫与慢弛豫组分(NPQf和NPQs) (B、C)的影响及随暗驰豫时间的变化(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照(CK)组和夜间低温组在15 min的光照平衡期间NPQ、NPQf和NPQs的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05); *, 对照组和低温处理组间差异显著(p < 0.05)。

Fig. 7 Photosystem II (PSII) non-photochemical quenching (NPQ) (A) and its fast and slow components (NPQf and NPQs) (B, C) of Kobresia pygmaea leaves under nocturnal low-temperature (NLT) treatment and their variation with dark relaxation time (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among different dark relaxation times in the control (CK) and NLT groups, respectively (α = 0.05). ns, no significant difference between the control and NLT groups (p > 0.05); *, significant difference between the control and NLT groups (p < 0.05).

图8 不同时长夜间低温(NLT)处理高山嵩草叶片光系统II (PSII)非光化学猝灭(NPQ) (A)和快弛豫与慢弛豫组分(NPQf和NPQs) (B、C)的变化及对夜间低温的响应(平均值±标准差, n = 30)。不同小写字母和大写字母分别表示对照(CK)组和夜间低温组在不同处理时长间NPQ、NPQf和NPQs的差异显著性(α = 0.05)。ns, 对照组和低温处理组间差异不显著(p > 0.05)。

Fig. 8 Photosystem II (PSII) non-photochemical quenching (NPQ) (A) and its fast and slow components (NPQf and NPQs) (B, C) of Kobresia pygmaea leaves under different days of nocturnal low-temperature (NLT) treatment (mean ± SD, n = 30). Different lowercase and uppercase letters indicate significant differences among different days of the treatment in the control (CK) and NLT groups, respectively (α = 0.05). ns, no significant difference between the control and NLT groups (p > 0.05).

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青藏高原高山嵩草光合功能对模拟夜间低温的响应

Responses of photosynthetic function of Kobresia pygmaea to simulated nocturnal low temperature on the Qingzang Plateau

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