丛枝菌根真菌对苹果品种耐盐碱的影响及缓解机理

doi:10.16420/j.issn.0513-353x.2024-0858

摘要/Abstract

摘要：

为探究接种丛枝菌根真菌（AMF）对盐碱胁迫下苹果嫁接苗生长的影响及其作用机理，以1年生‘秦脆’和‘瑞雪’嫁接盆苗（砧木M26）为试材，设置对照（清水灌溉）、盐碱胁迫（0.2 mol · L^-1 NaHCO₃︰NaCl = 1︰1，SA）、接种AMF（接种根内球囊霉，AMF）和SA + AMF（提前60 d接种AMF，再浇灌SA）处理。SA处理显著降低了‘秦脆’和‘瑞雪’的株高、茎粗、新梢生长量、根面积、根长、根体积以及叶绿素含量和光合速率，而接种AMF缓解了盐碱胁迫对苹果幼苗造成的伤害；与SA处理相比，SA + AMF处理‘秦脆’和‘瑞雪’的抗氧化酶活性显著升高，‘秦脆’叶片的POD、SOD和CAT活性分别升高了28.27%、48.72%和37.01%，而‘瑞雪’叶片POD、SOD和CAT活性分别升高了12.75%、23.37%和16.10%，并且叶片和根系的MDA含量均显著下降；‘秦脆’和‘瑞雪’的TTC还原强度分别升高71.40%和34.29%，根系柠檬酸含量分别增长了105.26%和122.22%，AMF通过促进柠檬酸的分泌来降低根系pH，增强了‘秦脆’和‘瑞雪’的耐盐碱性。盐碱条件下AMF共生降低了叶片中Na⁺含量，根系和叶片中的K⁺含量进一步上升，Na⁺/K⁺比值显著下降，其中‘秦脆’根系的Na⁺/K⁺比值恢复到了对照水平。可见，接种AMF能够促进苗木树势的恢复，增加根系活力、叶绿素含量和光合速率，调节Na⁺/K⁺比率来维持离子平衡，减少盐碱胁迫对植株造成的离子毒害，通过增强植株的SOD、POD和CAT活性来减少活性氧的积累，减少植株所受到的氧化损伤。此外，盐碱胁迫下AMF通过增加柠檬酸分泌来平衡根系pH，减少根系受到的pH胁迫，以此来缓解盐碱胁迫对植株造成的生长抑制。

关键词: 苹果, 丛枝菌根真菌, 盐碱胁迫, 离子稳态, 氧化损伤

Abstract:

To investigate the effects and mechanisms of the arbuscular mycorrhizal fungi（AMF）on the growth of apple grafted under salt-alkali stress，‘Qincui’and‘Ruixue’as experimental materials was grafted on M26. The experiment included four treatment groups：the control（CK，Irrigated with water），the salinity-alkality（0.2 mol · L^-1 NaHCO₃︰NaCl = 1︰1，SA），inoculation with AMF（Inoculated with Glomus intraradices，AMF），and SA + AMF（inoculation with AMF 60 d in advance，followed by watering with SA）. SA treatment significantly reduced plant height，stem diameter，shoot growth，root area，root length，root volume，chlorophyll content，and photosynthetic rate in‘Qincui’ and‘Ruixue’apple seedlings，however，AMF inoculation alleviated the growth inhibition caused by salt-alkali stress. Compared with SA treatment，the SA + AMF treatment significantly enhanced the activities of antioxidant enzymes（POD，SOD，and CAT）in‘Qincui’and‘Ruixue’. In‘Qincui’leaves，POD，SOD，and CAT activities increased by 28.27%，48.72%，and 37.01%，respectively，while in‘Ruixue’，these activities increased by 12.75%，23.37%，and 16.10%，and MDA content in leaves and roots significantly decreased. Compared with SA treatment，the SA + AMF treatment significantly enhanced TTC reduction intensity of‘Qincui’and‘Ruixue’increased by 71.40% and 34.29%，the root citrate content increased by 105.26% and 122.22%，respectively. AMF facilitated citrate secretion，thereby reducing root pH and enhancing the salt-alkali tolerance of‘Qincui’and‘Ruixue’. Under salt-alkali stress，AMF symbiosis decreased Na⁺ content in the leaves，while K⁺ content in both roots and leaves increased，leading to a significant reduction in the Na⁺/K⁺ ratio. Notably，the Na⁺/K⁺ ratio in the roots of‘Qincui’returned to control levels. These findings indicate that AMF inoculation promotes seedling vigor，root activity，chlorophyll content，and photosynthetic rate. Inaddition，AMF regulates the Na⁺/K⁺ ratio to maintain ion homeostasis，thereby mitigating ion toxicity caused by salt-alkali stress. Moreover，AMF enhanced the activities of antioxidant enzymes（SOD，POD and CAT），thereby reducing reactive oxygen species（ROS）accumulation and oxidative damage. Under salt-alkali stress，AMF also increases citrate secretion，which maintain balance of root pH and alleviates pH stress，further mitigating the growth inhibition caused by salt-alkali conditions.

Key words: apple, arbuscular mycorrhizal fungi, saline-alkalini stress, ion homeostasis, oxidative damage

杜丽君, 刘英材, 史妍妍, 汪洋, 李春晖, 何银涛, 王旭, 边传杰, 张满让. 丛枝菌根真菌对苹果品种耐盐碱的影响及缓解机理[J]. 园艺学报, 2025, 52(9): 2410-2424.

DU Lijun, LIU Yingcai, SHI Yanyan, WANG Yang, LI Chunhui, HE Yintao, WANG Xu, BIAN Chuanjie, ZHANG Manrang. Effect of Arbuscular Mycorrhizal Fungi on the Growth and Mechanism of Different Apple Cultivar Under Salt-Alkali Stress[J]. Acta Horticulturae Sinica, 2025, 52(9): 2410-2424.

图/表 11

图1 盐碱胁迫下AMF对‘秦脆’和‘瑞雪’苹果嫁接苗根系的侵染情况 SA：0.2 mol · L-1 NaHCO3 + NaCl；AMF：接种根内球囊霉；SA + AMF：AMF接种60 d后再进行SA处理。不同小写字母表示在0.05水平差异显著。下同

Fig. 1 The Phenotype and mycorrhizal infection rate of‘Qincui’and‘Ruixue’apples SA：0.2 mol · L-1 NaHCO3 + NaCl；AMF：Inoculation with Rhizophagus intraradices；SA + AMF：SA treatment applied 60 days after AMF inoculation. Different lowercase letters indicate significant difference at the 0.05 probability level. The same below

图2 盐碱胁迫20 d后‘秦脆’和‘瑞雪’苹果嫁接苗表型

Fig. 2 The phenotypes of‘Qincui’and‘Ruixue’apple grafted seedings after 20 days under salt-alkali stress

表1 盐碱胁迫20 d后‘秦脆’和‘瑞雪’苹果嫁接苗的生理指标

Table 1 Physiological indexes of‘Qincui’and‘Ruixue’apple grafted seedings after 20 days under salt-alkali treatment

品种 Cultivar	处理 Treatment	株高/cm Height	茎粗/mm Stem diameter	新梢生长量/cm Shoot growth	根面积/cm² Root area	根长/cm Root length	根体积/cm³ Root volume
‘秦脆’ ‘Qincui’	对照Control	106.0 ± 2.6 b	7.3 ± 0.3 b	82.3 ± 2.5 b	96.1 ± 3.5 b	3 294.6 ± 74.8 b	2 630.6 ± 187.5 b
	SA	83.0 ± 4.4 d	6.4 ± 0.2 c	71.3 ± 2.3 c	77.1 ± 1.3 d	2 672.1 ± 180.5 d	1 924.0 ± 155.8 c
	AMF	120.7 ± 5.6 a	8.1 ± 0.5 a	100.7 ± 2.1 a	117.1 ± 5.3 a	3 509.4 ± 110.3 a	3 061.3 ± 161.4 a
	SA + AMF	97.0 ± 5.2 c	7.2 ± 0.3 b	80.3 ± 2.5 b	83.6 ± 2.5 c	3 056.6 ± 19.1 c	2 437.3 ± 231.1 b
‘瑞雪’ ‘Ruixue’	对照Control	81.0 ± 1.7 b	7.1 ± 0.2 c	73.9 ± 2.0 b	91.4 ± 0.6 b	3 060.0 ± 122.6 b	2 592.3 ± 133.2 b
	SA	70.0 ± 1.7 d	6.3 ± 0.2 d	66.6 ± 1.4 c	72.7 ± 2.2 c	2 551.1 ± 174.8 d	1 914.3 ± 124.1 c
	AMF	89.0 ± 3.6 a	8.1 ± 0.2 a	98.2 ± 0.2 a	102.9 ± 3.0 a	3 441.2 ± 136.5 a	3 030.0 ± 123.1 a
	SA + AMF	77.7 ± 1.5 c	7.6 ± 0.2 b	71.3 ± 0.6 b	91.7 ± 8.2 b	2 706.4 ± 272.3 c	2 347.7 ± 186.9 b

图3 盐碱胁迫20 d后‘秦脆’和‘瑞雪’苹果的叶片叶绿素相对含量（A）和光合速率（B）

Fig. 3 Chlorophyll content（A）and photosynthetic rate（B）of‘Qincui’and‘Ruixue’apple after 20 days under salt-alkali treatment

图4 盐碱胁迫下AMF处理苹果嫁接苗超氧阴离子和H2O2染色

Fig. 4 Effect of AMF treatment on and H2O2 staing in apple grafted under saline-alkali

图5 盐碱胁迫下AMF处理对苹果嫁接苗超氧阴离子−和H2O2含量的影响

Fig. 5 Effect of AMF treatment on and H2O2 contents in apple grafted under saline-alkali

图6 盐碱处理20 d后‘秦脆’和‘瑞雪’苹果POD、SOD、CAT活性和MDA含量

Fig. 6 POD，SOD，CAT activities and MDA content in‘Qincui’and‘Ruixue’after 20 days under salt-alkali treatment

图7 盐碱胁迫后苹果嫁接苗根系TTC染色根尖颜色越红表示根系越有活力

Fig. 7 TTC staining of apple seedlings under saline-alkali stress The redder the root tip color indicates the more active of root

图8 盐碱胁迫后苹果嫁接苗根系TTC活性

Fig. 8 TTC activity of apple seedlings under saline-alkali stress

图9 盐碱胁迫后苹果嫁接苗苹果酸含量和柠檬酸含量

Fig. 9 Malic acid and citric acid content of apple seedlings under saline-alkali stress

图10 盐碱胁迫下AMF对‘秦脆’和‘瑞雪’苹果Na+含量、K+含量、Na+︰K+比率的影响

Fig. 10 Effects of saline-alkali stress and AMF on the Na+ content，K+ content and Na+︰K+ ratio of‘Qincui’and‘Ruixue’

参考文献 49

[1]	Altaf M A, Hao Y Y, Shu H H, Jin W H, Chen C H, Li L, Zhang Y, Mumtaz M A, Fu H H, Cheng S H, Zhu G P, Wang Z W. 2024. Melatonin mitigates cold-induced damage to pepper seedlings by promoting redox homeostasis and regulating antioxidant profiling. Horticultural Plant Journal,10:532-544.
[2]	Bahadur A, Batool A, Nasir F, Jiang S J, Qin M S, Zhang Q, Pan J B, Liu Y J, Feng H Y. 2019. Mechanistic insights into arbuscular mycorrhizal fungi-mediated drought stress tolerance in plants. International Journal Molecular Science, 20 (17):4199.
[3]	Chen Z J, Yu L, Liu W J, Zhang J, Wang N, Chen X S. 2021. Research progress of fruit color development in apple(Malus domestica Borkh.). Plant Physiology and Biochemistry,162:267-279.
[4]	Cook C, Huskey D, Mazzola M, Somera T. 2024. Effect of rootstock genotype and Arbuscular Mycorrhizal Fungal(AMF)apecies on early colonization of apple. Plants, 13 (10):1388.
[5]	Gao Hua, Zhao Zhenyang. 2016. A new apple variety-Rui Xue. China Fruit Industry Information, 33 (4):57. (in Chinese)
	高华, 赵政阳. 2016. 苹果新品种——瑞雪. 中国果业信息, 33 (4):57.
[6]	Gao X P, Guo H H, Zhang Q, Guo H X, Zhang L, Zhang C Y, Gou Z Y, Liu Y, Wei J M, Chen A Y, Chu Z H, Zeng F C. 2020. Arbuscular Mycorrhizal Fungi(AMF)enhanced the growth,yield,fiber quality and phosphorus regulation in upland cotton(Gossypium hirsutum L.). Science Reports,10:2084.
[7]	Gong X Q, Shi S T, Dou F F, Song Y, Ma F W. 2017. Exogenous melatonin alleviates alkaline stress in Malus hupehensis Rehd. by regulating the biosynthesis of polyamines. Molecules,22:1542.
[8]	Guo R, Shi L X, Yan C G, Zhong X L, Gu F X, Liu Q, Xia X. 2017. Ionomic and metabolic responses to neutral salt or alkaline salt stresses in maize(Zea mays L.) seedlings. BMC Plant Biology,17:41.
[9]	Javid M, Ford R, Nicolas M E. 2012. Tolerance responses of Brassica juncea to salinity,alkalinity and alkaline salinity. Functional Plant Biology,39:699-707.
[10]	Jin X Q, Liu T, Xu J J, Gao Z X, Hu X H. 2019. Exogenous GABA enhances muskmelon tolerance to salinity-alkalinity stress by regulating redox balance and chlorophyll biosynthesis. BMC Plant Biology,19:48.
[11]	Li Ao, Zheng Xu, Wu Chengxu, Nie Ruining, Ji Xinying, Tang Jiali, Zhang Junpei. 2025. The effect of arbuscular mycorrhizal fungi on the growth and physiological characteristics of walnut seedlings under salt stress. Acta Horticulturae Sinica, 52 (2):423-438. (in Chinese) doi: 10.16420/j.issn.0513-353x.2024-0348
	李敖, 郑旭, 吴承勖, 聂瑞宁, 姬新颖, 唐佳莉, 张俊佩. 2025. 丛枝菌根真菌对盐胁迫下核桃幼苗生长及生理的影响. 园艺学报, 52 (2):423-438. doi: 10.16420/j.issn.0513-353x.2024-0348
[12]	Li N, Zhang Z H, Gao S, Lv Y, Chen Z J, Cao B L, Xu K. 2021. Different responses of two Chinese cabbage(Brassica rapa L. ssp. pekinensis)cultivars in photosynthetic characteristics and chloroplast ultrastructure to salt and alkali stress. Planta, 254 (5):102.
[13]	Li W, Zhu C S, Song Y L, Yuan Y F, Li M, Sun Y K. 2024. Arbuscular mycorrhizal fungi by inducing watermelon roots secretion phthalates, altering soil enzyme activity and bacterial community composition to alleviate the watermelon wilt. BMC Plant Biology,24:593.
[14]	Li W Q, Zheng W J, Peng Y, Shao Y, Liu C T, Li J, Hu Y Y, Zhao B R, Mao B G. 2023. OsPMS1 Mutation enhances salt tolerance by suppressing ROS accumulation, maintaining Na⁺/K⁺ homeostasis,and promoting ABA biosynthesis. Genes(Basel), 14 (8):1621.
[15]	Li Z, Wu N, Meng S, Wu F, Liu T. 2020. Arbuscular mycorrhizal fungi(AMF)enhance the tolerance of Euonymus maackii Rupr. at a moderate level of salinity. PLoS ONE, 15 (4):e0231497.
[16]	Liang Shengmin, Zhang Fei, Wu Qiangsheng. 2023. Arbuscular mycorrhizal fungi improve drought tolerance of trifoliate orange seedlings by regulating root polyamines. Acta Horticulturae Sinica, 50 (12):2680-2688. (in Chinese) doi: 10.16420/j.issn.0513-353x.2022-1036
	梁圣敏, 张菲, 吴强盛. 2023. 丛枝菌根真菌通过调节枳根系多胺提高抗旱性. 园艺学报, 50 (12):2680-2688. doi: 10.16420/j.issn.0513-353x.2022-1036
[17]	Lin J X, Wang Y N, Sun S N, Mu C S, Yan X F. 2017. Effects of arbuscular mycorrhizal fungi on the growth,photosynthesis and photosynthetic pigments of Leymus chinensis seedlings under salt-alkali stress and nitrogen deposition. Science Total Environment,576:234-241.
[18]	Liu D, Zheng K Y, Wang Y, Zhang Y, Lao R M, Qin Z Y, Li T, Zhao Z W. 2022. Harnessing an arbuscular mycorrhizal fungus to improve the adaptability of a facultative metallophytic poplar(Populus yunnanensis)to cadmium stress:Physiological and molecular responses. Journal of Hazardous Materials,424:127430.
[19]	Liu R C, Yang L, Zou Y N, Wu Q S. 2023. Root-associated endophytic fungi modulate endogenous auxin and cytokinin levels to improve plant biomass and root morphology of trifoliate orange. Horticultural Plant Journal,9:463-472.
[20]	Liu Yanxiang, Yang Haimeng, Lai Yanfen, Zheng Jiena, Zhang Fuping. 2013. Research on catalase activity of Averrhoa carambola fruit. Journal of southern Agriculture, 44 (2):304-307. (in Chinese)
	刘燕湘, 杨海萌, 赖燕芬, 郑杰娜, 张福平. 2013. 杨桃果实过氧化氢酶活性研究. 南方农业学报, 44 (2):304-307.
[21]	Lu Jianzeng, Zhou Liyan, Huang Xinyi, Wu Fengzhi, Gao Danmei. 2024. Effects of carbon pool strength on plant growth and potassium uptake of tomato intercropped with potato-onion. Acta Horticulturae Sinica, 51 (11):1-13. (in Chinese)
	陆建增, 周丽颜, 黄馨仪, 吴凤芝, 高丹美. 2024. 番茄碳库强度对其与伴生分蘖洋葱和AMF的关系及钾吸收的影响. 园艺学报, 51 (11):1-13.
[22]	Ma C Q, Bian C J, Liu W J, Sun Z J, Xi X L, Guo D M, Liu X T, Tian Y K, Wang C H, Zheng X D. 2022. Strigolactone alleviates the salinity-alkalinity stress of Malus hupehensis seedlings. Frontiers in Plant Science,13:901782.
[23]	Mathur S, Sharma M P, Jajoo A. 2018. Improved photosynthetic efficacy of maize(Zea mays)plants with arbuscular mycorrhizal fungi(AMF)under high temperature stress. Journal of Photochem and Photobiology B-Biology,180:149-154.
[24]	Min Z, Li R Y, Chen L, Zhang Y, Li Z Y, Liu M, Ju Y L, Fang Y L. 2018. Alleviation of drought stress in grapevine by foliar-applied strigolactones. Plant Physiology Biochemistry,135:99-110.
[25]	Nacoon S, Jogloy S, Riddech N, Mongkolthanaruk W, Ekprasert J, Cooper J, Boonlue S. 2021. Combination of arbuscular mycorrhizal fungi and phosphate solubilizing bacteria on growth and production of Helianthus tuberosus under field condition. Science Reports, 11 (1):6501.
[26]	Peng Z C, Zulfiqar T, Yang H C, Wang M, Zhang F H. 2024. Effect of Arbuscular Mycorrhizal Fungi(AMF)on photosynthetic characteristics of cotton seedlings under saline-alkali stress. Science Reports, 14 (1):8633.
[27]	Phillips J M, Hayman D S. 1970. Improved procedures for clearing roots and staining parasitic and vesicular-Arbuscular Mycorrhizal Fungi for rapid assessment of infection. Transactions of the British Mycological Society,55:158-161.
[28]	Porcel R, Aroca R, Azcon R, Ruiz-Lozano J. 2016. Regulation of cation transporter genes by the arbuscular mycorrhizal symbiosis in rice plants subjected to salinity suggests improved salt tolerance due to reduced Na⁺ root-to-shoot distribution. Mycorrhiza,26:673-684.
[29]	Shang C Y, Liu X Y, Chen G, Li G B, Hu S S, Zheng H, Ge L, Long Y H, Wang Q M, Hu X H. 2024. SlWRKY 81 regulates Spd synthesis and Na⁺/K⁺ homeostasis through interaction with SlJAZ1 mediated JA pathway to improve tomato saline-alkali resistance. Plant Journal, 118 (6):1774-1792.
[30]	Shi C C, Yang F, Liu Z H, Li Y M, Di X L, Wang J H, Lin J X. 2021. Uniform water potential induced by salt,alkali, and drought stresses has different impacts on the seedling of hordeum jubatum:from growth,photosynthesis,and chlorophyll fluorescence. Frontiers in Plant Science,12:733236.
[31]	Song L L, Yu Y L, Chen H Z, Feng Y W, Chen S, Zhang H H, Zhou H J, Meng L, Wang Y. 2024. Response of photosynthetic characteristics and antioxidant system in the leaves of safflower to NaCl and NaHCO₃. Plant Cell Reports, 43 (6):146. doi: 10.1007/s00299-024-03234-7 pmid: 38764051
[32]	Sun Z J, Li J Y, Guo D M, Wang T C, Tian Y K, Ma C Q, Liu X L, Wang C H, Zheng X D. 2023. Melatonin enhances KCl salinity tolerance by maintaining K⁺ homeostasis in Malus hupehensis. Plant Biotechnol Journal, 21 (11):2273-2290.
[33]	Sun Zhijuan, Liu Wenjie, Zheng Xiaodong, Xi Xiangli, Ma Changqing, Liu Xiaoli, Tian Yike. 2023. Effects and functional mechanism of melatonin on the growth of Malus hupehensis seedlings under saline-alkali stress. Acta Horticulturae Sinica, 50 (8):1697-1710. (in Chinese)
	孙志娟, 刘文杰, 郑晓东, 袭祥利, 马长青, 刘晓丽, 王彩虹, 田义轲. 2023. 褪黑素对盐碱胁迫下平邑甜茶幼苗生长的影响及其机制. 园艺学报, 50 (8):1697-1710.
[34]	Tawaraya K, Tokairin K, Wagatuma T. 2001. Dependence of Alliun fistulosum cultivars on the arbuscular mycorrhizal funguf, Glomus fasciculatum. Applied Soil Ecology,17:119-124.
[35]	Wei H J, He W Y, Mao X J, Liao S K, Wang Q, Wang Z H, Tang M, Xu T Y, Chen H. 2024. Arbuscular mycorrhizal fungi and exogenous Ca²⁺ application synergistically enhance salt and alkali resistance in perennial ryegrass through diverse adaptive strategies. Microbiol Research,289:127906.
[36]	Wu C, Bi Y L, Zhu W B. 2024a. Is the amount of water transported by arbuscular mycorrhizal fungal hyphae negligible? Insights from a compartmentalized experimental study. Plant Soil,499:537-552.
[37]	Wu Y J, He H, Ren J Y, Shen H C, Sahito Z A, Li B, Tang X Y, Tao Q, Huang R, Wang C Q. 2024b. Assembly patterns and key taxa of bacterial communities in the rhizosphere soil of moso bamboo(Phyllostachys pubescens)under different Cd and Pb pollution. International Journal of Phytoremediation, 26 (11):1776-1786.
[38]	Xu H N, Sun X D, Wang X F, Shi Q H, Yang X Y, Yang F J. 2011. Involvement of a cucumber MAPK gene(CsNMAPK)in positive regulation of ROS scavengence and osmotic adjustment under salt stress. Scientia Horticulturae,127:488-493.
[39]	Xu J W, Yang J Y, Xu Z J, Zhao D K, Hu X H. 2020. Exogenous spermine-induced expression of SlSPMS gene improves salinity-alkalinity stress tolerance by regulating the antioxidant enzyme system and ion homeostasis in tomato. Plant Physiology Biochemistry,157:79-92.
[40]	Yang C X, Zhao W N, Wang Y N, Zhang L, Huang S C, Lin J X. 2020. Metabolomics analysis reveals the alkali tolerance mechanism in puccinellia tenuiflora plants inoculated with arbuscular mycorrhizal fungi. Microorganisms, 8 (3):327.
[41]	Yang Y Q, Guo Y. 2018. Elucidating the molecular mechanisms mediating plant salt-stress responses. New Phytology,217:523-539.
[42]	Yin X L, Zhang W, Feng Z W, Feng G D, Zhu H H, Yao Q. 2024. Improved observation of colonized roots reveals the regulation of arbuscule development and senescence by drought stress in the arbuscular mycorrhizae of citrus. Horticultural Plant Journal,10:425-436.
[43]	Zhai Shulin, Liu Xiaolin, Yao Luying, Zhan Wenjing, Gao Wenjun. 2023. Effect of AMF inoculation on root configuration of alfalfa under mixed saline-alkali stress. Feed Researh, 46 (21):90-94. (in Chinese)
	翟书林, 刘晓琳, 姚璐莹, 詹文晶, 高文俊. 2023. 混合盐碱下接种丛枝菌根真菌(AMF)对紫花苜蓿根系构型的影响. 饲料研究, 46 (21):90-94.
[44]	Zhang B, Shi F, Zheng X, Pan H Y, Wen Y Q, Song F Q. 2023a. Effects of AMF compound inoculants on growth,ion homeostasis,and salt tolerance-related gene expression in oryza sativa L. under salt treatments. Rice, 16 (1):18.
[45]	Zhang J X, Diao F W, Hao B H, Xu L, Jia B B, Hou Y Z, Ding S L, Guo W. 2023b. Multiomics reveals Claroideoglomus etunicatum regulates plant hormone signal transduction,photosynthesis and La compartmentalization in maize to promote growth under La stress. Ecotoxicology and Environmental Safety,262:115128.
[46]	Zhang M H, Wu M K, Xu T, Cao J F, Zhang Z H, Zhang T X, Xie Q Y, Wang J, Sun S W, Zhang Q Z, Ma R Y, Xie L N. 2024. A putative Na⁺/H⁺ antiporter BpSOS1 contributes to salt tolerance in birch. Plant Science,346:112181.
[47]	Zheng X D, Li Y Q, Xi X L, Ma C Q, Sun Z J, Yang X Q, Li X Y, Tian Y K, Wang C H. 2020. Exogenous strigolactones alleviate KCl stress by regulating photosynthesis,ROS migration and ion transport in Malus hupehensis Rehd. Plant Physiology Biochemistry,159:113-122.
[48]	Zheng Xiaodong, Xi Xiangli, Li Yuqi, Sun Zhijuan, Ma Changqing, Han Mingsan, Li Shaoxuan, Tian Yike, Wang Caihong. 2022. Effects and regulating mechanism of exogenous brassinosteroids on the growth of Malus hupehensis under saline-alkali stress. Acta Horticulturae Sinica, 49 (7):1401-1414. (in Chinese) doi: 10.16420/j.issn.0513-353x.2021-0499
	郑晓东, 袭祥利, 李玉琪, 孙志娟, 马长青, 韩明三, 李少旋, 田义轲, 王彩虹. 2022. 油菜素内酯对盐碱胁迫下平邑甜茶幼苗生长的影响及调控机理研究. 园艺学报, 49 (7):1401-1414. doi: 10.16420/j.issn.0513-353x.2021-0499
[49]	Zou Yangjun, Ma Fengwang, Fu Xuanchang, Li Cuiying, An Guiyang, Li Mingjun, Li Chao, Dang Zhiming. 2019. A new late ripening apple cultivar‘Qincui’. Acta Horticulturae Sinica, 46 (5):1011-1012. (in Chinese) doi: 10.16420/j.issn.0513-353x.2018-0368
	邹养军, 马锋旺, 符轩畅, 李翠英, 安贵阳, 李明军, 李超, 党志明. 2019. 晚熟苹果新品种‘秦脆’. 园艺学报, 46 (5):1011-1012. doi: 10.16420/j.issn.0513-353x.2018-0368