丛枝菌根真菌对盐胁迫下核桃幼苗生长及生理的影响

doi:10.16420/j.issn.0513-353x.2024-0348

摘要/Abstract

摘要：

为探究丛枝菌根真菌（AMF）对核桃幼苗耐盐性的影响和调控机理，以分别接种摩西斗管囊霉（Funneliformis mosseae）和异形根孢囊霉（Rhizophagus irregularis）的温室盆栽‘温185’核桃（Juglans regia‘Wen 185’）种子实生苗为试验材料，分析不同浓度NaCl胁迫下两种AMF对幼苗光合作用、叶绿素荧光参数、气孔特性、抗氧化酶活性、内源激素含量的影响。NaCl胁迫下，幼苗的生长发育受到抑制，光合性能下降，抗氧化酶（SOD、POD、APX）活性和脯氨酸（Pro）含量增加。总体上，接种AMF的幼苗表现出叶绿素含量、光合参数及内源激素（IAA、ABA、GA₃、ZR）水平提升，但对OJIP曲线的荧光强度（I-P）和电子传递效率（ABS/RC、TR_o/RC、ET_o/RC）的影响不显著。AMF接种降低了过氧化氢（H₂O₂）和丙二醛（MDA）的积累，减轻了氧化应激，并增加了100 mmol · L^-1 NaCl胁迫下的气孔宽度。接种F. mosseae增强了300 mmol · L^-1 NaCl胁迫时的抗氧化酶活性和Pro含量。主成分分析显示，AMF主要通过促进叶绿素合成，提高光合性能及增强POD活性和Pro含量来提高‘温185’核桃幼苗的耐盐性，其中接种F. mosseae的效果更好。

关键词: 核桃, 丛枝菌根真菌, 光合特性, 生理生化, 耐盐性

Abstract:

In order to investigate the effect and regulatory mechanisms of walnut seedlings’ salt tolerance influenced by arbuscular mycorrhizal fungi（AMF），in this study，Juglans regia‘Wen 185’ seedings were used as experimental material. Pot experiments were conducted in a greenhouse where seedlings were inoculated with Funneliformis mosseae and Rhizophagus irregularis. The effects of these two AMFs on photosynthetic parameters，chlorophyll fluorescence，stomatal characteristics，antioxidant enzyme activity，and endogenous hormone levels were analyzed under different NaCl concentrations （0，100，200，and 300 mmol · L^-1）. Under NaCl stress，seedling growth and development were inhibited，photosynthetic performance decreased，and the activities of antioxidant enzymes（SOD，POD，APX）and proline（Pro）content increased. Seedlings inoculated with AMF exhibited significant increases in chlorophyll content，photosynthetic parameters，and levels of endogenous hormones（IAA，ABA，GA₃，ZR），but the effects on the fluorescence intensity of the OJIP curve（I-P） and the efficiency of electron transport（ABS/RC，TR_o/RC，ET_o/RC）were not significant. AMF inoculation reduced the accumulation of hydrogen peroxide（H₂O₂）and malondialdehyde（MDA），mitigated oxidative stress，and improved stomatal width at 100 mmol · L^-1 NaCl. At 300 mmol · L^-1 NaCl，inoculation with F. mosseae enhanced antioxidant enzyme activity and Pro content. Principal component analysis showed that AMF mainly improved the salt tolerance of‘Wen 185’walnut seedlings by promoting chlorophyll synthesis，enhancing photosynthetic performance，and increasing POD activity and Pro content，with the inoculation of F. mosseae being more effective.

Key words: walnut, arbuscular mycorrhizal fungi, photosynthetic characteristics, physiology and biochemistry, salt tolerance

李敖, 郑旭, 吴承勖, 聂瑞宁, 姬新颖, 唐佳莉, 张俊佩. 丛枝菌根真菌对盐胁迫下核桃幼苗生长及生理的影响[J]. 园艺学报, 2025, 52(2): 423-438.

LI Ao, ZHENG Xu, WU Chengxu, NIE Ruining, JI Xinying, TANG Jiali, ZHANG Junpei. The Effect of Arbuscular Mycorrhizal Fungi on the Growth and Physiological Characteristics of Walnut Seedlings Under Salt Stress[J]. Acta Horticulturae Sinica, 2025, 52(2): 423-438.

图/表 12

图1 核桃幼苗接种AMF侵染情况（a、b）及其在不同浓度NaCl胁迫下的表型（c） NM：接种灭活菌剂，对照；F.m：接种Funneliformis mosseae；R.i：接种Rhizophagus irregularis。 ** 表示处理间差异极显著，t-test，α = 0.01

Fig. 1 AMF inoculation in walnut seedlings（a，b）and their phenotypes under various NaCl concentrations（c） NM：Inoculation with inactivated AMF inoculants，the control；F.m：Inoculation with Funneliformis mosseae；R.i：Inoculation with Rhizophagus irregularis. ** Indicates extremely significant differences between treatments，t-test，α = 0.01

表1 AMF对不同浓度NaCl胁迫下核桃幼苗生长的影响

Table 1 Effect of AMF on the growth of walnut seedlings under NaCl stress

NaCl/ （mmol · L^-1）	AMF接种 AMF inoculation	株高/cm Plant height	地径/mm Ground diameter	比叶面积/（cm² · g^-1） Specific leaf area	叶片含水量/％ Leaf water content
0	NM F.m	14.33 ± 0.98 abc 16.13 ± 0.96 a	5.97 ± 0.19 ab 6.40 ± 0.38 a	329.94 ± 7.41 bc 355.16 ± 8.16 a	73.12 ± 0.64 a 73.03 ± 0.80 a
0	R.i	16.25 ± 1.25 a	6.45 ± 0.61 a	350.28 ± 1.68 a	73.03 ± 0.34 a
100	NM F.m R.i	14.63 ± 1.29 abc 16.08 ± 1.10 a 14.10 ± 1.21 abc	5.70 ± 0.54 ab 6.45 ± 0.27 a 6.03 ± 0.20 ab	319.88 ± 5.52 cd 336.81 ± 4.05 b 319.96 ± 2.35 cd	71.11 ± 0.56 bc 71.97 ± 0.51 ab 71.97 ± 0.47 ab
200	NM F.m R.i	14.25 ± 1.60 abc 14.55 ± 1.43 abc 15.65 ± 0.93 ab	5.16 ± 0.13 b 5.75 ± 0.84 ab 5.48 ± 0.22 ab	289.68 ± 15.94 f 309.98 ± 2.23 de 303.88 ± 5.46 e	70.03 ± 0.26 cd 70.84 ± 0.90 bcd 69.61 ± 1.30 d
300	NM F.m R.i	13.63 ± 0.96 bc 15.13 ± 1.01 abc 13.25 ± 1.15 c	5.84 ± 1.01 ab 5.57 ± 0.41 ab 5.56 ± 0.29 ab	251.24 ± 8.10 h 299.84 ± 4.05 ef 266.31 ± 10.46 g	68.02 ± 1.25 e 70.31 ± 0.49 cd 69.53 ± 0.41 d

图2 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片叶绿素含量的变化 NM：接种灭活菌剂，对照；F.m：接种Funneliformis mosseae；R.i：接种Rhizophagus irregularis。不同小写字母表示处理间在P < 0.05水平差异显著。下同

Fig. 2 Changes in chlorophyll content in leaves of walnut seedlings under AMF inoculation and NaCl stress NM：Inoculation with inactivated AMF inoculants，the control；F.m：Inoculation with Funneliformis mosseae；R.i：Inoculation with Rhizophagus irregularis. Different small letters indicate a significant difference between the treatments at the P < 0.05 level. The same below

图3 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片光合参数的变化

Fig. 3 Changes of photosynthetic parameters in leaves of walnut seedlings under AMF inoculation and NaCl stress

图4 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片荧光强度变化

Fig. 4 Changes in fluorescence intensity in leaves of walnut seedlings under AMF inoculation and NaCl stress

图5 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片快速叶绿素荧光诱导动力学分析

Fig. 5 Fast chlorophyll fluorescence induction kinetics analysis in leaves of walnut seedlings under AMF inoculation and NaCl stress

图6 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片气孔扫描电镜观察

Fig. 6 Scanning electron microscopy observations of stomata in walnut seedling leaves under AMF inoculation and NaCl stress

图7 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片气孔特性的变化

Fig. 7 Changes of stomatal characteristics in leaves of walnut seedlings under AMF inoculation and NaCl stress

图8 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片抗氧化酶、脯氨酸、过氧化氢、丙二醛含量的变化

Fig. 8 Changes of antioxidant enzymes，proline，hydrogen peroxide and malondialdehyde contents in leaves of walnut seedlings under AMF inoculation and NaCl stress

图9 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片内源激素的变化

Fig. 9 Changes of endogenous hormones in leaves of walnut seedlings under AMF inoculation and NaCl stress

表2 AMF对不同浓度NaCl胁迫下各生理指标的双因素方差分析F值

Table 2 Two-way ANOVA F values with AMF for various physiological indicator under NaCl stress

指标Index	NaCl	AMF	NaCl × AMF
株高Plant height	2.798	2.023	1.68
地径Ground diameter	4.606	0.632	0.641
比叶面积Specific leaf area	143.643^***	34.491^***	3.846^**
叶片含水量Leaf water content	51.562^***	7.503^**	2.819^*
Chl. a	1129.989^***	84.332^***	12.145^***
Chl. b	523.973^***	53.622^***	20.335^***
Chl.（a+b）	988.983^***	80.524^***	14.573^***
P_n	840.754^***	73.736^***	9.115^***
G_s	1156.689^***	109.228^***	4.599^**
T_r	387.678^***	28.262^***	1.289
C_i	64.955^***	1.246	1.799
ABS/RC	7.164^***	1.565	0.965
TR_o/RC	6.997^**	0.836	0.712
ET_o/RC	3.296^*	4.882^*	0.855
DI_o/RC	5.196^**	3.395^*	1.398
F_v/F_m	3.07^*	4.687^*	1.419
PI_ABS	4.531^*	12.736^***	1.294
气孔长度Stomatal length	4.083^*	7.873^**	5.275^***
气孔宽度Stomatal width	12.215^***	6.167^**	1.657
气孔长宽比Stomatal length-width ratio	10.935^***	1.286	0.477
气孔密度Stomatal density	7.546^***	3.536^*	10.528^***
SOD	43.455^***	9.317^***	1.033
POD	114.64^***	1.555	5.271^***
APX	58.359^***	8.236^**	1.729
Pro	347.759^***	23.912^***	3.663^**
H₂O₂	369.106^***	72.425^***	7.615^***
MDA	39.018^***	79.852^***	19.385^***
IAA	6.973^**	196.747^***	5.551^***
ABA	41.951^***	960.551^***	41.726^***
GA₃	44.27^***	585.264^***	35.27^***
ZR	12.673^***	113.775^***	0.956

图10 AMF接种和不同浓度NaCl胁迫下核桃幼苗叶片各指标的主成分分析（A、B）和聚类热图（C） SL：气孔长度；D：地径；SW：气孔宽度；SLA：比叶面积；LWC：叶片含水量；H：株高；SD：气孔密度；SL/SW：气孔长宽比

Fig. 10 Principal component analysis（A，B） and cluster heatmap（C） of leaves of walnut seedlings under AMF inoculation and NaCl stress SL：Stomatal length；D：Ground diameter：SW：Stomatal width；SLA：Specific leaf area；LWC：Leaf water content；H：Plant height；SD：Stomatal density；SL/SW：Stomatal length-width ratio

参考文献 35

[1]	Ait-El-Mokhtar M, Laouane R B, Anli M, Boutasknit A, Wahbi S, Meddich A. 2019. Use of mycorrhizal fungi in improving tolerance of the date palm(Phoenix dactylifera L.) seedlings to salt stress. Scientia Horticulturae,253:429-438.
[2]	Amanifar S, Toghranegar Z. 2020. The efficiency of arbuscular mycorrhiza for improving tolerance of Valeriana officinalis L. and enhancing valerenic acid accumulation under salinity stress. Industrial Crops and Products,147:112234.
[3]	Cao Mingao, Zhang Fei, Huang Guangming, Liu Ruicheng, Liu Liping, Wu Qiangsheng, Xu Yongjie. 2023. The effect of arbuscular mycorrhizal fungi on the phosphorus absorption of walnut seedling roots under low phosphorus stress and its mechanism. Scientia Silvae Sinicae, 59 (12):117-124. (in Chinese)
	曹明奡, 张菲, 黄光明, 刘瑞成, 刘利平, 吴强盛, 徐永杰. 2023. 丛枝菌根真菌对低磷胁迫下核桃幼苗根系磷吸收的影响及机制. 林业科学, 59 (12):117-124.
[4]	Chen Xiaonan, Yilinuer Aili, Gao Wenli, Ma Xiaodong. 2022. The effects of arbuscular mycorrhizal fungi on the growth and physiology of Alhagi sparsifolia seedlings under salt stress. Pratacultural Science, 39 (9):1763-1772. (in Chinese)
	陈晓楠, 伊力努尔 · 艾力, 高文礼, 马晓东. 2022. 盐胁迫下丛枝菌根真菌对疏叶骆驼刺幼苗生长和生理的影响. 草业科学, 39 (9):1763-1772.
[5]	Chen J, Zhang H Q, Zhang X L, Tang M. 2020. Arbuscular mycorrhizal symbiosis mitigates oxidative injury in black locust under salt stress through modulating antioxidant defence of the plant. Environmental and Experimental Botany,175:104034.
[6]	Daszkowska-Golec A, Collin A, Sitko K, Janiak A, Kalaji H M, Szarejko I. 2019. Genetic and physiological dissection of photosynthesis in barley exposed to drought stress. International Journal of Molecular Sciences, 20 (24):6341.
[7]	Fattahi M, Mohammadkhani A, Shiran B, Baninasab B, Ravash R, Gogorcena Y. 2021. Beneficial effect of mycorrhiza on nutritional uptake and oxidative balance in pistachio(Pistacia spp.)rootstocks submitted to drought and salinity stress. Scientia Horticulturae,281:109937.
[8]	Fu Haiqi, Liu Xiao, Song Shu, Lü Wanjia, Yang Yongqing. 2023. Mechanisms of secondary metabolites in regulating plant resistance to salt-alkali stress. Plant Physiology Journal, 59 (4):727-740. (in Chinese)
	付海奇, 刘晓, 宋姝, 吕婉嘉, 杨永青. 2023. 次生代谢物调控植物抵抗盐碱胁迫的机制. 植物生理学报, 59 (4):727-740.
[9]	Ge Shibei, Jiang Xiaochun, Wang Lingyu, Yu Jingquan, Zhou Yanhong. 2020. Recent advances in the role and mechanism of arbuscular mycorrhiza-induced improvement of abiotic stress tolerance in horticultural plants. Acta Horticulturae Sinica, 47 (9):1752-1776. (in Chinese)
	葛诗蓓, 姜小春, 王羚羽, 喻景权, 周艳虹. 2020. 园艺植物丛枝菌根抗非生物胁迫的作用机制研究进展. 园艺学报, 47 (9):1752-1776.
[10]	Han S, Cheng Y, Wu G, He X W, Zhao G Z. 2024. Enhancing salt tolerance in poplar seedlings through arbuscular mycorrhizal fungi symbiosis. Plants, 13 (2):233.
[11]	Hao Han, Cao Lei, Chen Weinan, Hu Zenghui, Leng Pingsheng. 2020. Effects of salt stress on ion balance and physiological and biochemical characteristics of Quercus dentata seedlings. Acta Ecologica Sinica, 40 (19):6897-6904. (in Chinese)
	郝汉, 曹磊, 陈伟楠, 胡增辉, 冷平生. 2020. 盐胁迫对槲树(Quercus dentata)幼苗离子平衡及其生理生化特性的影响. 生态学报, 40 (19):6897-6904.
[12]	He Zhongqun, Li Huanxiu, Tang Haoru, He Chaoxing, Zhang Zhibin, Wang Huaisong. 2010. Effects of arbuscular mycorrhizal fungi on endogenous hormones in tomatoes under NaCl stress. Journal of Nuclear Agricultural Sciences, 24 (5):1099-1104. (in Chinese) doi: 10.11869/hnxb.2010.05.1099
	贺忠群, 李焕秀, 汤浩茹, 贺超兴, 张志斌, 王怀松. 2010. 丛枝菌根真菌对NaCl胁迫下番茄内源激素的影响. 核农学报, 24 (5):1099-1104. doi: 10.11869/hnxb.2010.05.1099
[13]	Huang P, Huang S, Ma Y, Danish S, Hareem M, Syed A, Elgorban A M, Eswaramoorthy R, Wong L S. 2024. Alleviation of salinity stress by EDTA chelated-biochar and arbuscular mycorrhizal fungi on maize via modulation of antioxidants activity and biochemical attributes. BMC Plant Biology, 24 (1):63. doi: 10.1186/s12870-024-04753-x pmid: 38262953
[14]	Huang Rongyan, Lu Chengwei, Xu Rong, Sun Zhengxuan, Fang Li, Chen Qinhua, Ma Tianyi, Liu Dianhui, Meng Weishan, Wang Yishun, Rao Huan, Liu Junchen. 2024. Effects of saline-alkali stress on the growth,physiology,and photosynthetic characteristics of‘Donghong’ sorbus. Acta Agretla Sinica, 32 (2):480-488. (in Chinese)
	黄荣雁, 路程伟, 许嵘, 孙正轩, 房莉, 陈钦华, 马天意, 刘殿辉, 孟维珊, 王一顺, 饶欢, 刘俊辰. 2024. 盐碱胁迫对‘冬红’花楸生长、生理及光合特性的影响. 草地学报, 32 (2):480-488. doi: 10.11733/j.issn.1007-0435.2024.02.015
[15]	Ji X Y, Tang J L, Zhang J P. 2022. Effects of salt stress on the morphology,growth and physiological parameters of Juglans microcarpa L. seedlings. Plants, 11 (18):2381.
[16]	Ji Xinying, Tang Jiali, Li Ao, Zheng Xu, Wang Hongxia, Zhang Junpei. 2024. Growth and physiological responses of different genotypes of walnut seedlings to salt stress. Scientia Silvae Sinicae,(2):65-77. (in Chinese)
	姬新颖, 唐佳莉, 李敖, 郑旭, 王红霞, 张俊佩. 2024. 盐胁迫下不同基因型核桃实生幼苗生长及生理响应. 林业科学,(2):65-77.
[17]	Jia Tingting, Chang Wei, Fan Xiaoxu, Song Fuqiang. 2018. Effects of AM fungi on photosynthesis and chlorophyll fluorescence characteristics of Elaeagnus angustifolia seedlings under salt stress. Acta Ecologica Sinica, 38 (4):1337-1347. (in Chinese)
	贾婷婷, 常伟, 范晓旭, 宋福强. 2018. 盐胁迫下AM真菌对沙枣苗木光合与叶绿素荧光特性的影响. 生态学报, 38 (4):1337-1347.
[18]	Khan T A, Saleem M, Fariduddin Q. 2023. Melatonin influences stomatal behavior,root morphology,cell viability,photosynthetic responses,fruit yield,and fruit quality of tomato plants exposed to salt stress. Journal of Plant Growth Regulation, 42 (4):2408-2432.
[19]	Li Pengmin, Gao Huiyuan, Reto J Strasser. 2005. Application of rapid chlorophyll fluorescence induction dynamics in photosynthesis research. Journal of Plant Physiology and Molecular Biology,(6):559-566. (in Chinese) pmid: 16361781
	李鹏民, 高辉远, Reto J Strasser. 2005. 快速叶绿素荧光诱导动力学分析在光合作用研究中的应用. 植物生理与分子生物学学报,(6):559-566. pmid: 16361781
[20]	Li Xiangyan. 2022. Physiological and ecological responses of three aquatic plants to Na⁺ and K⁺ salt stress[M. D. Dissertation]. Wuhan: Wuhan University. (in Chinese)
	李向彦. 2022. 三种水生植物对Na⁺盐和K⁺盐胁迫的生理生态响应研究[硕士论文]. 武汉: 武汉大学.
[21]	Lv Y, Xu N, Ha M R, Tan Z M, Guo S R, Wang J, Wang Y, Sang T, Shu S. 2024. Bacillus cereus enhances salt tolerance of cucumber seedlings by improving antioxidant metabolism and decreasing the ion toxicity. Scientia Horticulturae,328:112885.
[22]	Ma W Y, Qin Q Y, Zou Y N, Kuča K, Giri B, Wu Q S, Hashem A, AlArjani AlBandari F, Almutairi K F, Abd Allah E F, Xu Y J. 2022. Arbuscular mycorrhiza induces low oxidative burst in drought-stressed walnut through activating antioxidant defense systems and heat shock transcription factor expression. Frontiers in Plant Science,13:1089420.
[23]	Ma Z B, Zhao X C, He A, Cao Y, Han Q S, Lu Y J, Yong J W H, Huang J. 2022. Mycorrhizal symbiosis reprograms ion fluxes and fatty acid metabolism in wild jujube during salt stress. Plant Physiology, 189 (4):2481-2499. doi: 10.1093/plphys/kiac239 pmid: 35604107
[24]	Niu Linglei. 2016. Germination of walnut seeds and the effect of different salt concentrations on their growth characteristics. Xinjiang Agricultural Reclamation Science and Technology, 39 (12):19-24. (in Chinese)
	牛蛉磊. 2016. 核桃种子萌发及不同盐浓度对其生长特性的影响. 新疆农垦科技, 39 (12):19-24.
[25]	Shahvali R, Shiran B, Ravash R, Fallahi H, Banović Đ B. 2020. Effect of symbiosis with arbuscular mycorrhizal fungi on salt stress tolerance in GF677(peach × almond)rootstock. Scientia Horticulturae,272:109535.
[26]	Shang Yefei, Li Ming, Ding Bo, Niu Hao, Yang Zhenning, Chen Xiaoqiang, Cao Gaoyi, Xie Xiaodong. 2017. Advances in the study of auxin regulation of plant stomatal development. Chinese Bulletin of Botany, 52 (2):235-240. (in Chinese)
	商业绯, 李明, 丁博, 牛浩, 杨振宁, 陈小强, 曹高燚, 谢晓东. 2017. 生长素调控植物气孔发育的研究进展. 植物学报, 52 (2):235-240. doi: 10.11983/CBB16099
[27]	Testerink C, Munnik T. 2005. Phosphatidic acid:a multifunctional stress signaling lipid in plants. Trends in Plant Science, 10 (8):368-375. doi: 10.1016/j.tplants.2005.06.002 pmid: 16023886
[28]	Wang Baoqing. 2021. Study on the main traits and molecular basis of early-maturing walnut in Xinjiang[Ph. D. Dissertation]. Beijing: Chinese Academy of Forestry. (in Chinese)
	王宝庆. 2021. 新疆早实核桃主要性状及分子基础研究[博士论文]. 北京: 中国林业科学研究院.
[29]	Wang Jin. 2015. Study on the inoculation effects of arbuscular mycorrhizal fungi on walnut seedlings[M. D. Dissertation]. Jingzhou: Yangtze University. (in Chinese)
	王进. 2015. 丛枝菌根真菌对核桃幼苗接种效应研究[硕士论文]. 荆州: 长江大学.
[30]	Wang Wei, Mu Hongna, Yang Huimin, Sun Taoze. 2024. The influence of arbuscular mycorrhizal fungi(AMF)inoculation on the growth status and photosynthetic characteristics of Leucojum aestivum. Journal of Nanjing Forestry University(Natural Science Edition), https://link.cnki.net/urlid/32.1161.S.20240408.0944.002. in Chinese)
	王炜, 母洪娜, 杨慧敏, 孙陶泽. 2024. 接种丛枝菌根真菌(AMF)对夏雪片莲生长状况和光合特性的影响. 南京林业大学学报(自然科学版), https://link.cnki.net/urlid/32.1161.S.20240408.0944.002.
[31]	Wu Yuhan, Liu Wenhui, Liu Kaiqiang, Zhang Yongchao. 2022. Effects of drought stress on photosynthetic characteristics and reactive oxygen species scavenging system in oat seedlings. Acta Prataculturae Sinica, 31 (10):75-86. (in Chinese) doi: 10.11686/cyxb2021410
	吴雨涵, 刘文辉, 刘凯强, 张永超. 2022. 干旱胁迫对燕麦幼苗叶片光合特性及活性氧清除系统的影响. 草业学报, 31 (10):75-86. doi: 10.11686/cyxb2021410
[32]	Zhang D J, Tong C L, Wang Q S, Bie S. 2024. Mycorrhizas affect physiological performance,antioxidant system,photosynthesis,endogenous hormones,and water content in cotton under salt stress. Plants, 13 (6):805.
[33]	Zhao Xiuting, Wang Yanshuang, Duan Jie, Ma Lüyi, He Baohua, Jia Zhongkui, Sang Ziyang, Zhu Zhonglong. 2021. Effects of salt stress on the growth and photosynthetic characteristics of grafted Magnolia liliiflora seedlings. Scientia Silvae Sinicae, 57 (4):43-53. (in Chinese)
	赵秀婷, 王延双, 段劼, 马履一, 何宝华, 贾忠奎, 桑子阳, 朱仲龙. 2021. 盐胁迫对红花玉兰嫁接苗生长和光合特性的影响. 林业科学, 57 (4):43-53.
[34]	Zhu S P, Nong J F, Luo G T, Li Q P, Wang F S, Jiang D, Zhao X C. 2020. Varied tolerance and different responses of five citrus rootstocks to acid stress by principle component analysis and orthogonal analysis. Scientia Horticulturae,278:109853.
[35]	Zou Y N, Qin Q Y, Ma W Y, Zhou L J, Wu Q S, Xu Y J, Kuča K, Hashem A, AlArjani AlBandari F, Almutairi K F, AbdAllah E F. 2023. Metabolomics reveals arbuscular mycorrhizal fungi-mediated tolerance of walnut to soil drought. BMC Plant Biology, 23 (1):118.