干旱胁迫下AMF对茶树光合特性及其基因表达的影响

doi:10.16420/j.issn.0513-353x.2023-0655

摘要/Abstract

摘要：

为探究干旱胁迫下丛枝菌根真菌（arbuscular mycorrhizal fungi，AMF）对茶树（Camellia sinensis）光合特性的影响，于温室盆栽条件下，对‘福鼎大白茶’实生苗进行水分和接种AMF试验。结果显示，干旱胁迫（田间最大持水量的55%）显著抑制了AMF对茶树根系的侵染，降低了叶片中叶绿素b含量、最大光量子效应（QY_max）值、胞间CO₂浓度（C_i）、气孔纵径和横径以及气孔开度，碳同化相关酶基因CsRbcL、CsTK、CsFBPase、CsPRK以及叶绿素合成酶基因CsHEME的表达下调；提高了叶绿素a/b、非光化学猝灭（NPQ_lss）参数，叶绿素合成酶CsHEMA1表达上调。正常供水（田间最大持水量的75%）和干旱胁迫条件下接种AMF，显著提高了叶绿素a和总叶绿素的含量、净光合速率（P_n）、气孔导度（G_s）、C_i、蒸腾速率（T_r），降低了NPQ_lss及气孔密度，不同程度地诱导了两种水分条件下碳同化相关酶基因CsRbcL、CsTK、CsFBPase、CsPRK以及叶绿素合成相关酶基因CsHEMA1、CsHEMC、CsHEME、CsHEMG和CsCHLE的表达。与此同时，接种AMF还显著增加了干旱胁迫下叶绿素b的含量、QY_max、气孔纵经、横径和气孔开度以及正常供水下叶绿素a/b。研究表明，接种AMF可促进茶树的光合作用，提高茶树对干旱的适应性，且在干旱胁迫下效果显著。

关键词: 茶树, 丛枝菌根真菌, 干旱胁迫, 气孔, 光合特性

Abstract:

In order to explore the effect of arbuscular mycorrhizal fungi（AMF）on the photosynthetic characteristics of tea plants under drought stress，the seedlings of Camellia sinensis. ‘Fuding Dabaicha’were subjected to water and AMF inoculation experiments under greenhouse pot conditions. The results showed that drought stress（55% of the maximum field water capacity）significantly inhibited the AMF infection on tea plant roots，reduced the chlorophyll b content，maximum quantum effect（QY_max）value，intercellular CO₂ concentration（C_i），stomatal vertical and horizontal diameter and stomatal opening，as well as down-regulated the expressions of carbon assimilation-related enzymes （CsRbcL，CsTK，CsFBPase，CsPRK）genes and chlorophyll synthesis-related enzyme gene CsHEME. Whereas，drought stress significantly increased the chlorophyll a/b and non-photochemical quenching （NPQ_lss）parameters，and up-regulated the expression of chlorophyll synthase enzymes gene CsHEMA1 in tea seedlings. Howbeit，under well-watered（75% of the maximum field water capacity）and drought stress conditions，inoculation with AMF significantly increased the content of chlorophyll a and total chlorophyll，net photosynthetic rate（P_n），stomatal conductance（G_s），C_i，and transpiration rate（T_r）but decreased NPQ_lss and stomatal density，accompany with the up-regulated expressions of carbon assimilation-related enzyme genes CsRbcL，CsTK，CsFBPase，CsPRK and chlorophyll synthesis-related enzyme genes CsHEMA1，CsHEMC，CsHEME，CsHEMG and CsCHLE in different degrees. Meanwhile，AMF inoculation also significantly increased the chlorophyll b content，QY_max，stomatal vertical，horizontal diameter and aperture under drought stress and chlorophyll a/b under well-watered. The results indicated that AMF can improve the adaptability of tea plant under drought stress by promoting photosynthesis，and the promotion effect was more significant under drought stress.

Key words: tea plant, arbuscular mycorrhizal fungi, drought stress, stomatal, photosynthetic characteristics

陈鑫, 邬晓龙, 刘升锐, 胡贤春, 刘春艳. 干旱胁迫下AMF对茶树光合特性及其基因表达的影响[J]. 园艺学报, 2024, 51(10): 2358-2370.

CHEN Xin, WU Xiaolong, LIU Shengrui, HU Xianchun, LIU Chunyan. Effects of AMF on Photosynthetic Characteristics and Gene Expressions of Tea Plants Under Drought Stress[J]. Acta Horticulturae Sinica, 2024, 51(10): 2358-2370.

图/表 10

表1 荧光定量引物序列

Table 1 Quantitative RT-PCR primer sequence

基因 Gene	名称 Name	正向引物（5′-3′） Forward primer sequence	反向引物（5′-3′） Reverse primer sequence
β-actin	肌动蛋白Actin	GGCGGATCAAGTGTTGGAAGGGAG	ACGCTTGGGATTGTATTCGGCATTA
CsCHLM	Mg-原卟啉Ⅸ甲基转移酶Magnesium protoporphyrin Ⅸ methyltransferase	CTCTATTGCCTCATTCCTC	ATTTAGTGTTTGGGTTGGT
CsCHLE	Mg-原卟啉Ⅸ单甲基酯环化酶Magnesium protoporphyrin Ⅸ monomethyl ester cyclase	CAATGACTGGAAGGCTAA	ATTCTTTGGTGTTGAGGC
CsHEMA1	谷氨酸-tRNA还原酶 Glutamyl-tRNA reductase	ATTCGTTGCGAGATTGTT	GCTGCTCCTTTCCTTTGT
CsHEMC	胆色素原脱氨酶 Porphobilinogen deaminase	TGACCGCCATTCTTTCTA	GCTAATCTTGTTTCCTCGT
CsHEMD	尿卟啉原Ⅲ合成酶 Uroporphyrinogen Ⅲ synthase	TGTCTGGGCTGTCTTCGA	CAAATCAGGCAACCGTGT
CsHEME	尿卟啉原Ⅲ脱羧酶 Uroporphyrinogen Ⅲ decarboxylase	ACATTCGCTTCTGTTCCC	TTTCTACTTCCAGCCCTC
CsHEMG	原卟啉原氧化酶Menaquinone dependent protoporphyrinogen oxidase	TCTGTGGAAGAAACGGAACT	CCGCAACGAAAGGGTCAA
CsPPOX	原卟啉Ⅳ原氧化酶 Protoporphyrinogen Ⅳ oxidase	GAAGCAGTTGACCGTGAC	AAAGCCCTCTGAACCCAC
CsRbcL	核酮糖-1,5-二磷酸羧化/加氧酶大亚基Ribulose- 1,5-bisphosphate carboxylase/ oxygenase large subunit	TGGCTGGAGATGGGACGA	CCTCTGGTAATCAGAACAGGGTT
CsTK	转酮醇酶 Transketolase	TGCCCAATGTTTTGATGCTACG	ATCCACCCTTTTCCACTCCCTC
CsFBPase	果糖-1,6-二磷酸酶 Furetose-1,6-bisphosphate phosphatase	GCAGATTGCTTCGTTGGTTC	TGCTATTATCCCTGTCCTCCC
CsPRK	核酮糖-5-磷酸激酶 Ribulose 5-phosphate kinase	AGTATTGGAGCCCGAAAGCC	CAAACAGGTAAACTGGGGTGAA

图1 茶树实生苗接种AMF后根系菌根发育状况（A、B）和茶根围土壤中的根外菌丝（C）

Fig. 1 Mycorrhizal development status（A，B）of tea seedlings after inoculation with AMF and external hyphae in rhizosphere soil（C）

图2 茶树实生苗在正常供水和干旱胁迫下接种AMF对根系菌根侵染率和土壤菌丝长度的影响图中的数值为平均数 ± 标准差（n = 6）。不同的小写字母表示不同处理间差异显著（P < 0.05）。

Fig. 2 Effects of AMF on mycorrhizal colonization and soil hyphae length of tea seedlings under well-watered and drought stress The data in the graph are mean ± SD（n = 6）. Different lowercase letters indicate significant differences （P < 0.05）between different treatments.

表2 茶实生苗在正常供水和干旱胁迫下接种AMF对叶片叶绿素含量的影响

Table 2 Effects of AMF on chlorophyll content of tea seedlings under well-watered and drought stress mg · g-1

处理 Treatment	叶绿素a Chlorophyll a	叶绿素b Chlorophyll b	叶绿素总含量 Total chlorophyll	叶绿素a/b Ratio of chlorophyll a/b
正常供水（对照）Well-watered（Control）	1.61 ± 0.07 c	0.65 ± 0.02 a	2.26 ± 0.10 c	2.48 ± 0.10 c
正常供水 + AMF Well-watered + AMF	1.85 ± 0.08 ab	0.68 ± 0.10 a	2.53 ± 0.17 a	2.72 ± 0.12 b
干旱胁迫 Drought stress	1.78 ± 0.03 b	0.46 ± 0.04 c	2.24 ± 0.04 c	3.87 ± 0.29 a
干旱胁迫 + AMF Drought stress + AMF	1.90 ± 0.07 a	0.55 ± 0.01 b	2.45 ± 0.08 b	3.45 ± 0.33 a

图3 茶树实生苗在正常供水和干旱胁迫下接种AMF对叶片叶绿素荧光参数QY_max、QY_lss和NPQ_lss的影响

Fig. 3 Effects of AMF on chlorophyll fluorescence parameters QY_max，QY_lss and NPQ_lss of tea seedlings under well-watered and drought stress

图4 茶树实生苗在正常供水和干旱胁迫下接种AMF对叶片光合作用及相关参数的影响图中的数值为平均数 ± 标准差（n = 6）。不同的小写字母表示不同处理间差异显著（P < 0.05）。

Fig. 4 Effects of AMF on photosynthetic parameters of tea seedlings under well-watered and drought stress The data in the graph are mean ± SD（n = 6）. Different lowercase letters indicate significant differences（P < 0.05）among different treatments.

表3 茶树实生苗在正常供水和干旱胁迫下接种AMF对气孔开度的影响

Table 3 Effects of AMF on stomatal aperture of tea seedlings under well-watered and drought stress

处理 Treatment	气孔密度/（个 · m^-2） Stomatal density	气孔横径/mm Stomatal vertical diameter	气孔纵径/mm Stomatal horizontal diameter	气孔开度/mm²Stomatal aperture
正常供水（对照）Well-watered	2.33 ± 0.21 a	0.175 ± 0.017 a	0.233 ± 0.015 a	0.032 ± 0.004 a
正常供水 + AMF Well watered + AMF	1.43 ± 0.12 c	0.184 ± 0.013 a	0.227 ± 0.014 ab	0.033 ± 0.004 a
干旱胁迫Drought stress	2.54 ± 0.19 a	0.147 ± 0.008 b	0.215 ± 0.010 b	0.025 ± 0.002 b
干旱胁迫 + AMF Drought stress + AMF	1.83 ± 0.19 b	0.172 ± 0.012 a	0.231 ± 0.009 a	0.031 ± 0.003 a

图5 茶树实生苗在正常供水和干旱胁迫下接种AMF后叶片气孔状况

Fig. 5 Stomatal status of tea leaves under well-watered and drought stress after AMF inoculation

图6 茶树实生苗在正常供水和干旱胁迫下接种AMF对叶片光合碳同化关键酶基因表达的影响图中的数值为平均数 ± 标准差（n = 6）。不同的小写字母表示不同处理间差异显著（P < 0.05）。

Fig. 6 Effects of AMF on gene expressions of key enzymes of photosynthetic carbon assimilation in tea seedlings under well-watered and drought stress The data in the graph are mean ± SD（n = 6）. Different lowercase letters indicate significant differences（P < 0.05） among different treatments.

图7 茶树实生苗在正常供水和干旱胁迫下接种AMF对叶片叶绿素合成相关酶基因表达量的影响图中的数值为平均数 ± 标准差（n = 6）。不同的小写字母表示不同处理间差异显著（P < 0.05）。

Fig. 7 Effects of AMF on gene expressions of chlorophyll synthesis-related enzymes in tea seedlings under well-watered and drought stress The data in the graph are mean ± SD（n = 6）. Different lowercase letters indicate significant differences （P < 0.05）among different treatments.

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