اخبار و اعلانات

 آمار

تعداد نشریات282
تعداد نشریات سازمان80
تعداد شماره‌ها3,272
تعداد مقالات36,149
تعداد مشاهده مقاله55,722,519
تعداد دریافت فایل اصل مقاله94,499,986
نادیان قمشه, حبیب اله. (1405). پاسخ‌های گیاه– قارچ میکوریز به تنش خشکی: واکاوی فرآیندهای زیستی و سازوکارهای دفاعی. سامانه مدیریت نشریات علمی, 14(1), 19-60. doi: 10.22092/sbj.2026.371633.290
حبیب اله نادیان قمشه. "پاسخ‌های گیاه– قارچ میکوریز به تنش خشکی: واکاوی فرآیندهای زیستی و سازوکارهای دفاعی". سامانه مدیریت نشریات علمی, 14, 1, 1405, 19-60. doi: 10.22092/sbj.2026.371633.290
نادیان قمشه, حبیب اله. (1405). 'پاسخ‌های گیاه– قارچ میکوریز به تنش خشکی: واکاوی فرآیندهای زیستی و سازوکارهای دفاعی', سامانه مدیریت نشریات علمی, 14(1), pp. 19-60. doi: 10.22092/sbj.2026.371633.290
نادیان قمشه, حبیب اله. پاسخ‌های گیاه– قارچ میکوریز به تنش خشکی: واکاوی فرآیندهای زیستی و سازوکارهای دفاعی. سامانه مدیریت نشریات علمی, 1405; 14(1): 19-60. doi: 10.22092/sbj.2026.371633.290

پاسخ‌های گیاه– قارچ میکوریز به تنش خشکی: واکاوی فرآیندهای زیستی و سازوکارهای دفاعی

زیست شناسی خاک
دوره 14، شماره 1، شهریور 1405، صفحه 19-60    اصل مقاله (2.04 M)
نوع مقاله: مقاله مروری
شناسه دیجیتال (DOI): 10.22092/sbj.2026.371633.290
نویسنده
حبیب اله نادیان قمشه*
(بازنشسته) گروه خاکشناسی-دانشگاه کشاورزی و منابع طبیعی رامین خوزستان
چکیده
تنش خشکی یکی از مهم‌ترین چالش‌های کشاورزی در مناطق خشک و نیمه‌خشک است که سبب آسیب‌های فراوان از جمله صدمات اکسیداتیو در گیاهان می‌شود. تحت تنش خشکی، بسیاری از فرایندهای زیستی گیاه عمدتا به سبب انباشت گونه‌های فعال اکسیژن مختل می‌شود. سازوکارهای متنوع دفاع آنتی‌اکسیدانی، گیاهان را قادر می‌سازد تا ضمن مقابله با تنش اکسیداتیو، توازن ردوکس و همستازی گونه‌های فعال اکسیژن را در شرایط تنش تا حدودی حفظ کنند. آنزیم‌های آنتی‌اکسیدان کلیدی مانند سوپراکسید دیسموتاز و کاتالاز نقش بسیار مهمی در خنثی کردن گونه‌های فعال اکسیژن دارند، حال آنکه آنتی‌اکسیدان‌های غیر آنزیمی بیشتر از طریق مهار و دفع گونه‌های فعال اکسیژن در کاهش تنش اکسیداتیو شرکت می‌کنند. سازوکارهای دفاعی گیاهان در همزیستی با قارچ‌های میکوریز آربسکولار در مقابله با تنش خشکی به صورت هم‌افزایی مشاهده می‌شود. قارچ‌های میکوریز علاوه بر فعالیت‌های آنتی‌اکسیدانی خود از طریق طیف وسیعی از متابولیت‌ها، قادرند با تنظیمات اسمزی، بیان ژن‌های کلیدی پاسخ‌دهنده به خشکی و نیز بهبود تغذیه گیاه آسیب‌های ناشی از تنش خشکی در گیاه میزبان خود را کاهش دهند. هر چند پیشرفت‌های اخیر با استفاده از زیست‌شناسی مولکولی و مهندسی ژنتیک تا حدودی توانسته است چگونگی سیگنال‌دهی گونه‌های فعال اکسیژن و سیستم‌های آنتی‌اکسیدان‌های آنزیمی و غیر آنزیمی و نیز برهمکنش‌های پیچیده هورمونی در پاسخ به تنش خشکی را روشن کنند، ولی هنوز بسیاری از این سازوکارها ناشناخته باقی مانده است. بدون شک پژوهش‌های آینده با تمرکز بر زوایای ناشناخته و پنهان تنظیم‌کننده‌های اصلی پاسخ‌های فیزیولوژیک و بیوشیمیایی گیاه در برابر تنش خشکی می‌توانند به توسعه کشاورزی پایدار و مقاوم به خشکی کمک کنند.
کلیدواژه‌ها
آنتی اکسیدان‌های آنزیمی و غیر آنزیمی؛ پراکسید هیدرژن؛ پرولین؛ تنش خشکی؛ قارچ‌های میکوریزا
عنوان مقاله [English]
Plant-mycorrhiza responses to drought stress: Analysis of vital processes and defense mechanisms
نویسندگان [English]
Habiballah Nadian Ghomsheh
Professor emeritus
چکیده [English]
Background and Objectives: Drought stress is one of the most critical challenges in agriculture, especially in arid and semi-arid regions. In light of ongoing climate change and prolonged drought periods, this issue has posed a serious threat to food security, particularly in countries located in Western Asia. Drought stress severely affects many physiological, enzymatic, biochemical, and nutritional processes in plants. Among the most important physiological and biochemical effects of drought stress are disturbances in photosynthesis and the photosynthetic electron transport chain, intercellular CO₂ concentration, photosynthetic rate, stomatal conductance, transpiration rate, excessive production of reactive oxygen species (ROS), disruptions in plant water potential, and alterations in the regulation and synthesis of enzymatic, non-enzymatic, and hormonal osmolytes. To cope with the damage caused by drought stress, plants—alongside appropriate morphological adjustments activate a set of protective mechanisms (such as increased synthesis of antioxidant metabolites) and regulatory mechanisms (such as enhancing osmotic regulators) in conjunction with molecular responses. In fact, upon perceiving stress signals through the roots, the plant increases the expression of regulatory protein genes such as calcium-dependent protein kinases and mitogen-activated protein kinases, as well as the expression of genes involved in the synthesis of osmolytes including proline, glycine betaine, LEA proteins, and soluble sugars. These responses not only mitigate oxidative stress but also help improve plant water relations. The defensive mechanisms of plants in symbiosis with arbuscular mycorrhizal fungi are activated synergistically to counteract drought stress.




Review Methods: Numerous studies have highlighted the pivotal role of soil organisms, particularly arbuscular mycorrhizal fungi, in alleviating the detrimental effects of drought stress and mediating the shared defense mechanisms between plants and fungi. These investigations encompass a wide range of plant responses under drought conditions, both in the presence and absence of mycorrhizal associations, and are derived from peer-reviewed journals and reputable scientific databases. This analytical review synthesizes the findings of researchers in the field, drawing on up-to-date studies on drought stress physiology, antioxidant metabolism, osmolyte accumulation, and molecular responses, accessed through major international databases including Web of Science and Scopus. Furthermore, Google Scholar provided a freely accessible platform for identifying recent publications, while specialized resources such as ScienceDirect, SpringerLink and Taylor & Francis offered valuable journal articles, book chapters, and authoritative reference works in the fields of mycorrhizal symbiosis and soil ecology.




Results: Under drought stress, plant water relations are disrupted due to a reduction in the moisture potential gradient between the soil solution and plant tissues. Stomatal closure, aimed at reducing water loss, is one of the plant’s earliest responses to drought conditions. This stress not only impairs photosynthetic processes but also leads to excessive production and accumulation of reactive oxygen species (ROS), resulting in oxidative damage at the cellular level. To cope with the damaging effects of drought stress, plants along with appropriate morphological adjustments activate a series of protective mechanisms (such as enhanced synthesis of antioxidant metabolites), regulatory mechanisms (such as increased osmotic adjustment), and molecular responses. Numerous studies have shown that mycorrhizal fungi can participate in these protective and regulatory mechanisms, thereby enhancing the host plant’s adaptive capacity through various pathways. Mycorrhizal colonization supports the plant by promoting the synthesis and accumulation of osmotic regulators and antioxidant compounds, thus providing effective protection under drought conditions. A reduction in osmotic metabolites is considered a drought avoidance mechanism, whereas increased accumulation of osmotic regulators and antioxidant compounds represents a drought tolerance strategy. In the interaction between mycorrhizal fungi and aquaporins in reducing drought stress in the host plant, several mechanisms are involved. Among these mechanisms is the upregulation or stabilization of key aquaporin genes such as PIP1;6, PIP2;1, and PIP2;5. Gene expression regulation—particularly when mediated by mycorrhizal fungi—constitutes one of the earliest and most important mechanisms enabling drought tolerance. The interaction between aquaporins and stomata under drought conditions is coordinated by mycorrhizal fungi in a manner that ensures optimal water relations depending on the intensity of the drought and the plant’s sensitivity to it. Upon sensing drought stress through their root system, plants transmit signals to various organs to adjust cellular processes accordingly. One such response is the synthesis of osmolytes such as proline, glycine betaine, and LEA proteins, which contribute to physiological, biochemical, and molecular adjustments that improve osmotic potential, enhance water relations, maintain cellular homeostasis, and ultimately increase drought tolerance—particularly in the presence of mycorrhizal fungi. Additionally, mycorrhizal plants exhibit enhanced synthesis of enzymatic antioxidants such as superoxide dismutase (SOD) and catalase (CAT), as well as antioxidant-related phytohormones including gibberellic acids, abscisic acid (ABA), indole-3-acetic acid (IAA), and jasmonic acid (JA). These compounds, each with distinct and sometimes contrasting physiological roles, act in a coordinated and synergistic manner to substantially mitigate the detrimental effects of drought stress on plant growth, nutrition, and physiological performance.




Conclusion: Plants employ complex antioxidant defense systems to mitigate the damaging effects of drought stress, relying on both enzymatic and non-enzymatic metabolites. Arbuscular mycorrhizal fungi substantially strengthen these defenses by enhancing ROS scavenging, stabilizing photosynthetic and redox processes, and improving osmotic adjustment, nutrient uptake, and the expression of drought-responsive genes. This symbiosis effectively contributes to ROS homeostasis and reduces oxidative damage, thereby improving plant resilience under water-limited conditions. Despite recent advances in molecular biology that have clarified elements of ROS signaling, antioxidant pathways, and hormonal crosstalk, many aspects of plant–mycorrhiza interactions under drought remain unresolved. Continued research using modern biotechnological approaches is essential for uncovering these mechanisms and developing more drought-tolerant crops that can support future global food security.
کلیدواژه‌ها [English]
drought stress, enzymatic and non-enzymatic antioxidants, mycorrhizal fungi, Hydrogen peroxide, Proline
مراجع
  1. Abdul-Aziz, M., Sabeem, M., Mullath, S.K., Brini, F. and Masmoudi, K. 2021. Plant group II LEA proteins: Intrinsically disordered structure for multiple functions in response to environmental stresses. Biomolecules, 11: 1662. doi: org/ 10.3390/biom11111662.
  2. Abo-Doma,, Edrees, S. and Abdel-Aziz, S.H. 2011. The effect of mycorrhiza growth and expression of some genes in barley. Egyptian Journal of Genetics and Cytology, 40:2, 301-313. doi: 10.21608/ejgc.2011.10794.
  3. Al-Amri, S.M. 2021. Application of bio-fertilizers for enhancing growth and yield of common bean plants grown under water stress conditions. Saudi Journal of Biological Sciences, vol. 28 (7) 3901–3908. doi: 10.1016/j.sjbs.2021.03.064.
  4. Alcazar, R., Cuevas, J.C., Patron, M., Altabella, T. and Tiburcio, A.F. 2006. Abscisic acid modulates polyamine metabolism under water stress in Arabidopsis thaliana. Physiologia plantarum,128, 448–455. doi: 10.1111/j.1399-3054.2006.00780.x.
  5. Alizadeh, O. and Nadian, H. 2010. Evaluation effect of water stress and nitrogen rates on amount of uptake some macro and micro elements in mycorrhizal and non-mycorrhizal corn plant. Research Journal of Biological Sciences, 5(5): 350-355, doi: 10.36478/rjbsci.2010.350.355 [In Persian].
  6. Anuradha, S. and Rao, S.S.R. 2008. The effect of brassinosteroids on radish (Raphanus sativus) seedlings growing under cadmium stress. Plant, Soil and Environment, 53, 465–472. doi: org/10.17221/2307-pse..
  7. Al-Karaki, G.N. 1998. Benefit, cost and water-use efficiency of arbuscular mycorrhizal durum wheat grown under drought stress, Mycorrhiza, 8 (1), 41–45. doi: org/10.1007/s005720050209.
  8. Aroca, R., Porcel, R., Ruiz-Lozano, J.M. 2006. How does arbuscular mycorrhizal symbiosis regulate root hydraulic properties and plasma membrane aquaporins in Phaseolus vulgarisunder drought, cold or salinity stresses? New Phytologist, 173(4), 808-816. doi: org/10.1111/j.1469 8137.2006.01961.x.
  9. Ashraf, M. and Foolad, M.R. 2007. Roles of glycine betaine and proline in improving plant abiotic stress resistance. Environmental and Experimental Botany, 59: 206–216. doi: org/10.1016/j.envexpbot.2005.12.006.
  10. Atkinson,E. 1977. Cellular energy metabolism and its regulation. Academic Press, London, UK. NLM Unique ID: 7708918, MMS ID 994946603406676; ISBN 9780120661503; ISBN 0120661500; NLM Unique ID 7708918. P 293.
  11. Auge, R.M., Toler, H.D. and Saxton, A.M. 2015. Arbuscular mycorrhizal symbiosis alters stomatal conductance of host plants more under drought than under amply watered conditions: a meta-analysis. Mycorrhiza, 25(1):13-24. doi:10.1007/s00572-014-0585-4. Epub 2014 May 16.
  12. Auge, R.M., Stodola, A.J.W., Ebel, R.C. and Duan, X. 1995. Leaf elongation and water relations of mycorrhizal sorghum in response to partial soil drying: two Glomus species at varying phosphorus fertilization. Journal of Experimental Botany, 46: 297-307. doi: org/10.1093/jxb/46.3.297.
  13. Barzana, G., Aroca, R., Paz, J.A., Chaumont, F., Martinez-Ballesta, M.C., Carvajal, M. and Ruiz-Lozano, J.M. 2012. Arbuscular mycorrhizal symbiosis increases relative apoplastic water flow in roots of the host plant under both well-watered and drought stress conditions. Annals Botany, 109: 1009-1017. doi: 1093/aob/mcs007.
  14. Barzana, G., Arco, R., Bienert, G.P. Chaumont, F. and Ruiz-Lozano, J.M. 2014. New Insights into the regulation of aquaporins by the arbuscular mycorrhizal symbiosis in maize plants under drought stress and possible implications for plant performance. Molecular Plant-Microbe Interactions, 27(4), 349-363, doi: org/10.1094/MPMI-09-13-0268-R.
  15. Begum, N., Ahanger, M.A., Su, Y., Lei, Y., Mustafa, N.S.A., Ahmad, P., Lixin Zhang, L. 2019. Improved drought tolerance by AMF inoculation in Maize (Zea mays) involves physiological and biochemical implications. Plants (Basel), 6;8(12):579. doi: 3390/plants8120579.
  16. Bendou, O., Gutierrez-Fernandez, I., Marcos-Barbero, E., Bueno-Ramos, N., Gonzalez-Hernandez, A., Morcuende, R. and  Arellano, J.B. 2021. Theoretical and experimental considerations for a rapid and high throughput measurement of catalase in vitro. Antioxidants, 11(1):21, doi: 3390/antiox11010021.
  17. Bethke, C., Badger, M.R., Russell, L. and Jonesa, R.L. 2004. Apoplastic synthesis of nitric oxide by plant tissues. The Plant Cell, 16: 332–341. doi: 10.1105/tpc.017822.
  18. Bhosale, K.S. and Shinde, B.P. 2011. Influence of arbuscular mycorrhizal fungi on proline and chlorophyll content in Zingiber Officinale Rosc grown under water stress. Indian Journal of Fundamental and Applied Life Sciences, 1 (3) :172-176. doi: http://www.cibtech.org/jls.htm
  19. Campomanes, P., Rothlisberger, U. Alfonso-Prieto, M. and Rovira, C. 2015. The Molecular mechanism of the catalase-like activity in horseradish peroxidase. Journal of the American Chemical Society, 137 (34), doi: org/10.1021/jacs.5b06796.
  20. Candar-Cakir, B.; Arican, E.; Zhang, B. 2016. Small RNA and degradome deep sequencing reveals drought-and tissue-specific micrornas and their important roles in drought-sensitive and drought-tolerant tomato Plant Biotechnology Journal, 14, 1727–1746. doi: 10.1111/pbi.12533.
  21. Chareesri, A., Deyn, G,B.D., Sergeeva, L., Polthanee, A. and Kuyper, T.W. 2020. Increased arbuscular mycorrhizal fungal colonization reduces yield loss of rice (Oryza sativa) under drought. Mycorrhiza, 30 (2-3), 315–328. doi: 10.1007/s00572-020-00953-z.
  22. Charpentier, M., Sun, J., Wen, J., Mysore, K.S. and Oldroyd, G. 2014. ABA promotion of arbuscular mycorrhizal colonization requires a component of the PP2A protein phosphatase complex. Plant Physiology,166, 2077–2090. doi:10.1104/pp.114.246371.
  23. Chandrasekaran, M. 20220. Arbuscular mycorrhizal fungi mediated enhanced biomass, root morphological traits and nutrient uptake under drought stress: A Meta-Analysis. Journal of Fungi, 8(7), 660; https://doi.org/10.3390/jof8070660.
  24. Chitarra, W., Maserti, B., Gambino, G., Guerrieri, E. and Balestrini, R. 2016a. Arbuscular mycorrhizal symbiosis-mediated tomato tolerance to drought. Plant Signaling and Behaviour, 11(7):00-00, doi:1080/15592324.2016.1197468.
  25. Chitarra, , Pagliarani, C., Maserti, B., Lumini, E., Siciliano, I., Cascone, P., Schubert, A., Gambino, G., Balestrini, R. and Guerrieri, E. 2016b. Insights on the impact of arbuscular mycorrhizal symbiosis on tomato tolerance to water stress. Plant Physiology, 171: 1009–1023. doi: 10.1104/pp.16.00307.
  26. Chen, S. and Schopfer, P. 1999. Hydroxyl‐radical production in physiological reactions: a novel function of peroxidase. European journal of biochemistry, 260, 726-735. doi: 10.1046/j.1432-1327.1999.00199.x.
  27. Chen, W., Men, P., Feng, H. and Wang, C. 2020. Effects of arbuscular mycorrhizal fungi on growth and physiological performance of Catalpa bungeiA.Mey. under drought stress. Forests, 11:10, 1117, doi: org/10.3390/f11101117.
  28. Choi, J., Lee, T., Cho, J., Servante, E.K., Pucker, B., Summers, W., Bowden, S., Rahimi, M., An, K., An, G., Bouwmeester, H.J., Wallington, E.J., Oldroyd, G. and Paszkowski, U. 2020. The negative regulator SMAX1 controls mycorrhizal symbiosis and strigolactone biosynthesis in rice. Nature Communications, 11:2114, doi: org/10.1038/s41467-020-16021-1.
  29. Chun, C., Paramasivan, M. and Chandrasekaran, M. 2018. Proline accumulation influenced by osmotic stress in arbuscular mycorrhizal symbiotic plants. Frontiers in Microbiology, 9:2525, doi: 10.3389/fmicb.2018.02525.
  30. Conner, A.C., Bill, R.M. and Conner, M.T. 2013. An emerging consensus on aquaporin translocation as a regulatory mechanism. Molecular Membrane Biology, 30:1–12. doi: 10.3109/09687688.2012.743194
  31. Corpas, F.J., Freschi, L., Rodriguez-Ruiz, M., Mioto, P.T. and Gonzalez-Gordo, S. 2018. Nitro-oxidative metabolism during fruit ripening. Journal of Experimental Botany, 69(14), 3449–3463. doi:10.1093/jxb/erx453.
  32. Das, D., Varshney, K., Ogawa, S., Torabi, S., Hüttl, R., Nelson, D.C. and Gutjahr, C. 2025. Ethylene promotes SMAX1 accumulation to inhibit arbuscular mycorrhiza symbiosis. Nature Communications, 16, Article No., doi: org/10.1038/s41467-025-57222-w.
  33. Demirel, U., Morris, W.L., Ducreux, L.J., Yavuz, C., Asim, A., Tindas, I., Campbell, R., Morris, J.A., Verrall, S.R., Hedley, P.E., Gokce, Z.N.O., Caliskan, S., Aksoy, E., Caliskan, M.E., Taylor, M.A. and Handcock, R.D. 2020, Physiological, biochemical, and transcriptional responses to single and combined abiotic stress in stress-tolerant and stress-sensitive potato genotypes. Frontiers in Plant Science, 11: 169, doi: org/10.3389/fpls.2020.00169.
  34. Delormel, T.Y. and Boudsocq, M. 2019. Properties and functions of calcium-dependent protein kinases and their relatives in Arabidopsis thaliana. New Phytologist, 224(2), 585-604. doi: org/10.1111/nph.16088.
  35. Fabregas, N., Lozano-Elena, F., Blasco-Escámez, D., Tohge, T., Martínez-Andújar, C., Albacete, A., Osorio, S., Bustamante, M., Riechmann, J.L., Nomura, T., Yokota, T., Conesa, A., Pérez Alfocea, F., Fernie, F.R., Cano-Delgado, A.I. 2018. Overexpression of the vascular brassinosteroid receptor BRL3 confers drought resistance without penalizing plant growth. Nature Communications, 9:4680, doi: 10.1038/s41467-018-06861-3.
  36. Floss, D.S., Levy, J.G., Lévesque-Tremblay, V., Pumplin, V.N. and Harrison, M.J. 2013. DELLA proteins regulate arbuscule formation in arbuscular mycorrhizal symbiosis. Proceedings of the National Academy of Sciences of the United States of America. 110(51), 5025-34.  doi: 10.1073/pnas.1308973110. 
  37. Foo, , McAdam, E.L., Weller, J.L. James, B. and Reid, J.B. 2016. Interactions between ethylene, gibberellins, and brassinosteroids in the development of rhizobial and mycorrhizal symbioses of pea. Journal of Experimental Botany, 67(8), 2413–2424, doi: org/10.1093/jxb/erw047.
  38. Foyer, C.H. and Hanke, G. 2022. ROS production and signalling in chloroplasts: cornerstones and evolving concepts. The Plant Journal, 111, 642–661. doi: 10.1111/tpj.15856.
  39. Fracetto, G.G.M., Peres, L.E.P. and Lambais, M.R. 2017. Gene expression analyses in tomato near isogenic lines provide evidence for ethylene and abscisic acid biosynthesis fine-tuning during arbuscular mycorrhiza development. Archives of Microbiology, Springer-Verlag Berlin Heidelberg 2017, doi: 10.1007/s00203-017-1354-5.
  40. Gavassi,A., Silva, G.S., Carolina de Marchi Santiago da Silva, C.M.S., Thompson, A.J., Macleod, K., Oliveira, P.M.R., Cavalheiro, M.F., Douglas Domingues, S. and Habermann, G. 2021, NCED expression is related to increased ABA biosynthesis and stomatal closure under aluminum stress. Environmental and Experimental Botany, 185: 104404, doi: org/10.1016/j.envexpbot.2021.104404.
  41. Ghosh, K., Islam, M.N., Siddiqui, M.N., Cao, X. and Khan, M.A.R. 2022. Proline, a multifaceted signalling molecule in plant responses to abiotic stress: understanding the physiological mechanisms. Plant Physiology, ISSN 1435-8603, doi:10.1111/plb.13363.
  42. Giel, R.D. and Guinel, F.C. 2002. Effects of elevated substrate-ethylene on colonization of leek (Allium porrum) by the arbuscular mycorrhizal fungus Glomus aggregatum. Biology Faculty Publications. 21, University of Guelp, hhttps://scholars.wlu.ca/biol_faculty/21.
  43. Gill, S.S. and Tuteja, N. 2010. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiology and Biochemistry, 48(12), 909-30.  doi: 10.1016/j.plaphy.2010.08.016. Epub 2010 Sep 15.
  44. Grumberg, B.C., Urcelay, C., Shroeder, M.A., Vargas-Gil, S. and Luna, C.M. 2015. The role of inoculum identity in drought stress mitigation by arbuscular mycorrhizal fungi in soybean. Biology and Fertility of Soils, 51, 1–10. doi: 10.1007/s00374-014-0942-7.
  45. Gui, L.X., Lu, S.S., Chen, Q., Yang, L. and Xiao, J.X. 2021. iTRAQ based proteomic analysis reveals positive impacts of arbuscular mycorrhizal fungi inoculation on photosynthesis and drought tolerance in blueberry. Trees, 35(1), 81– 92. doi: 10.1007/s00468-020-02015-5.
  46. Guo, N., Lu, W., Liu, C.Y. and Q.S. 2024. Regulation of arbuscular mycorrhizal fungi in citrus root hairs mediated by auxin efflux carrier protein PtPINs. Scientia Horticulturae, 337:113574, doi:org/10.1016/j.scienta.2024.113574.
  47. Gupta, R. and S.K. Chakrabarty, S.K. 2013. Gibberellic acid in plant, still a mystery unresolved. Plant Signaling and Behavior, 8:9, e25504. doi: 10.4161/psb.25504.
  48. Haghighania, H. Nadian, H. Ragali, F. Tavakoli, A.R. 2012. Effect of two species of arbuscular mycorrhizae on vegetative growth and P uptake by Mexican Lime Rootstock (Citrus aurantifolia) under drought stress. Journal of Seed and Plant Production, 28(4), 403-417.  doi: 22092/sppj.2017.110486. [In Persian].
  49. Halliwell, B., Gutteridge, J.M.C. 1999. Free radicals in biology and medicine. Oxford, UK, Oxford University Press. SBN (1999 edition): 0198500459 (10‑digit) / 9780198500452 (13‑digit) pp: 936
  50. Hasanuzzaman, , Borhannuddin-Bhuyan, M.H.M., Zulfiqar, F., Raza, A., Mohsin, S.M., Mahmud, J.A., Fujita, M. and Fotopoulos, V. 2020. Reactive oxygen species and antioxidant defense in plants under abiotic stress: Revisiting the crucial role of a universal defense regulator. Antioxidants, 29;9(8), 681. doi: 10.3390/antiox9080681.
  51. Hsu, P., Dubeaux, G., Takahashi, Y. and Schroeder, J.I. 2021. Signaling mechanisms in abscisic acid‐mediated stomatal closure. The Plant Journal, 105, 307–321. doi: 10.1111/tpj.15067.
  52. Huang, D., Ma, M., Wang, Q., Zang, M., Jing, G., Li, C. and Ma, F. 2020. Arbuscular mycorrhizal fungi enhanced drought resistance in apple by regulating genes in the MAPK pathway. Plant Physiology and Biochemistry, 149, 245–255. doi: org/10.1016/j.plaphy.2020.02.020.
  53. Huang, Y.M., Zou, Y.N. and Wu, Q.S. 2017. Alleviation of drought stress by mycorrhizas is related to increased root H2O2efflux in trifoliate orange. Scientific Reports, 7(42335), doi: 10.1038/srep42335.
  54. Hussain, M., Malik, M.A., Farooq, M., Ashraf, M.Y., Cheema, M.A. 2008. Improving drought tolerance by exogenous application of glycine betaine and salicylic acid in sunflower. Journal of Agronomy and Crop Science, 194: 193–199. doi: 10.1111/j.1439-037X.2008.00305.x.
  55. Irankhah, S., Chitarra, W., Mashreghi, M., Nerva, L., Ganjeali, A., Balestrini, R., Antoniou, C., Cheniany, M., and Lumini, E. 2020. Impact of an arbuscular mycorrhizal fungal inoculum and exogenous methyl jasmonate on fenugreek secondary metabolite production under water deficit. Environmental and Experimental Botany, 176, 104096. DOI: 10.1016/j.envexpbot.2020.104096.
  56. Jajic, , Sarna, T. and trzalka, K. 2015. Senescence, stress and reactive oxygen species. Plants, 4, 393–411.doi:10.3390/plants4030393. doi: 10.3390/plants4030393.
  57. Jamdhade, A.R., Sunkar, R. and Hivrale, V. 2017. Zymographic method for distinguishing different classes of superoxide dismutases in plants. Springer Nature, 221-227. doi: 10.1007/978-1-4939-7136-7_13.
  58. Jokar, N., Nadian. H. Khalilimoghdam, B., Haydari, M. and Gharineh, M. 2015. Effect of myorrhizal colonization on plant growth and proline content in some genotype of leeks (Allium ampeloprasum ssp. persicum L, Allium porrum). Plant production, 38(1), 15-26. doi: org/10.22055/ppd.2015.11115 [In Persian].
  59. Jokar, N., Nadian. H. Khalilimoghdam, B. and Haydari, M. 2013. Effects of arbuscular mycorrhizal fungus and drought stress on plant root growth, accumulation of proline and uptake some nutrients by three genotypes of leeks. Journal of Soil Biology. 1(2), doi:22092/sbj.2014.128410. [In Persian].
  60. Jakobsen, I., Abbott, L.K. and Robson, A.D. 1992. External hyphae of vesicular-arbuscular mycorrhizal fungi associated with Trifolium subterraneum 1. Spread of hyphae and phosphorus inflow into roots. New Phytologist 120: 371-380. doi: org/10.1111/j.1469-8137.1992.tb01077.x.
  61. Jomova, K., Raptova, R., Y. Alomar, S.Y., Alwasel, S.H., Nepovimova, E., Kuca, K. and Valko, M. 2023. Reactive oxygen species, toxicity, oxidative stress, and antioxidants: chronic diseases and aging, Archives of Toxicology, 97:2499–2574, doi: org/10.1007/s00204-023-03562-9.
  62. Ishii, T., Shrestha, Y. H., Matsumoto, I. and Kadoya, K. Effect of ethylene on the growth of vesicular-arbuscular mycorrhizal fungi and on the mycorrhizal formation of trifoliate orange roots. Journal of the Japanese Society for Horticultural Science. 65, 525–529. doi: 10.2503/jjshs.65.525.
  63. Karpinska, B. and Foyer, C.H. 2024. Superoxide signalling and antioxidant processing in the plant nucleus. Journal of Experimental Botany, 75(15), 4599–4610, doi: org/10.1093/jxb/erae090.
  64. Khamsuk, O., Sonjaroon, W., Suwanwong, S., Jutamanee, K. and Suksamrarn, A. 2018. Effects of 24-epibrassinolide and the synthetic brassinosteroid mimic on chili pepper under drought. Acta Physiologiae Plantarum, 40(6), 106. doi: 10.1007/s11738-018-2682-z.
  65. Khan, M.I., Nazir, F., Asgher, M., Per, T.S., Khan, N.A. 2015. Selenium and sulphur influence ethylene formation and alleviate cadmium-induced oxidative stress by improving proline and glutathione production in wheat. Journal of Plant Physiology, 173, 9–18. doi:org/10.1016/j.jplph.2014.09.011.
  66. Khan, M., Ali, S., Al Azzawi, T.N.I., Saqib, S. Ullah, F., Ayaz, A. and Zaman, W. 2023. The key roles of ROS and RNS as a signaling molecule in plant–microbe interactions. Antioxidants, 12, 268. doi: org/10.3390/ antiox12020268.
  67. Khan, W.U., Khan, L.U., Chen, D. and Chen, F. 2023. Comparative analyses of superoxide dismutase (SOD) gene family and expression profiling under multiple abiotic stresses in water lilies. Horticulturae, 9(7), 781, doi: org/10.3390/horticulturae9070781.
  68. Khan, P., Abdelbacki, A.M.M. Albaqami, M., Jan, R. and Kim, K.M. 2024. Proline promotes drought tolerance in maize. Biology, 14(1), 41, doi: org/10.3390/biology14010041.
  69. Khatri, K. and Rathore, M.S. 2022. Salt and osmotic stress-induced changes in physio-chemical responses, PSII photochemistry and chlorophyll a fluorescence in peanut. Plant Stress, 3:100063. doi: org/10.1016/j.stress.2022.100063.
  70. Khoyerdi, F.F., Shamshiri, M.H., Estaji, A. 1016. Changes in some physiological and osmotic parameters of several pistachio genotypes under drought stress. Science of Horticulture,198, 44–51. doi: 10.1016/j.scienta.2015.11.028.
  71. Kozlov, A.V., Javadov, S. and Sommer, N. Cellular ROS and antioxidants: Physiological and pathological role. Antioxidants, 13, 602. doi: org/ 10.3390/antiox13050602.
  72. Krajinski F, Biela A, Schubert D, Gianinazzi-Pearson V, Kaldenhoff R, Franken P (2000) Arbuscular mycorrhiza development regulates the mRNA abundance of Mtaqp1 encoding a mercury-insensitive aquaporin of Medicago truncatula. Planta, 211:85–90. doi: 1007/s004250000263.
  73. Kucera, B., Cohn M. A. and Leubner-Metzger, G. 2005. Plant hormone interactions during seed dormancy release and germination. Seed Science Research,15, 281–307. doi:1079/SSR2005218.
  74. Lanfranco, L., Mara-Novero, M. and Paola Bonfante, P. 2005. The mycorrhizal fungus Gigaspora margaritapossesses a CuZn superoxide dismutase that is up-regulated during symbiosis with legume hosts. Plant Physiology, 137 (4) 1319–30 doi: org/10.1104/pp.104.050435.
  75. Laxa, M., Liebthal, M., Telman, W., Chibani, K. and Dietz, K.J 2019. The role of the plant antioxidant system in drought tolerance. Antioxidants, 8, 94. doi: org/ 10.3390/antiox8040094.
  76. Lee, Y., Yamamoto, A.,  Ragaey, M., Hashiguchi, M., Akashi, R.  and Saeki, Y. 2024. Effect of drought stress and short-term recovery on proline and polyamine metabolism in Lotus japonicus. Agricultural Research, doi: org/10.1007/s40003-024-00832-5.
  77. Leon, J. and Costa-Broseta, A. 2020. Present knowledge and controversies, deficiencies, and misconceptions on nitric oxide synthesis, sensing, and signaling in plants. Plant Cell and Environment.43,1–15. doi: 10.1111/pce.13617.
  78. Li, J., Zhang, K., Meng, Y., Hu, J., Ding, M., Bian, J., Yan, M., Han, J. and Zhou, M. 2018. Jasmonic acid/ethylene signaling coordinates hydroxycinnamic acid amides biosynthesis through ORA59 transcription factor. Plant Journal,95, 444–457. doi: 10.1111/tpj.13960.
  79. Li, R., Shen, J., Wilkinson, S., Ford, B.G. and Davies, W.J. 2017. Stomatal and growth responses to hydraulic and chemical changes induced by progressive soil drying. Journal of Experimental Botany, 68, 21–22, 5883–5894. doi:10.1093/jxb/erx381.
  80. Li, J., Wang, Y., Pritchard, H.W. and Wang, X. 2014. The fluxes of H2O2 and O2 can be used to evaluate seed germination and Vigor of Caragana korshinskii. Planta, 239(6):1363-73. doi: 1007/s00425-014-2049-7.
  81. Li, Z., Zhang, Y., Liu, C., Gao, Y., Han, L. and Chu, H. 2022. Arbuscular mycorrhizal fungi contribute to reactive oxygen species homeostasis of Bombax ceiba under drought stress. Frontier Microbiology, 13,991781. doi: 10.3389/fmicb.2022.991781.
  82. Li, N., Li, J., Xie, J., Wenjing Rui, W., Pu, K., Gao, Y., Tiantian Wang, T. and Zhang, M. 2025. Glycine betaine and plant abiotic stresses: Unravelling physiological and molecular responses. Plant Science, 355, 112479, doi: org/10.1016/j.plantsci.2025.112479Get rights and content.
  83. Luo, C., Li, Z., Shi, Y., Gao, Y., Xu, Y., Zhang, Y. and Chu, H. 2024. Arbuscular mycorrhizal fungi enhance drought resistance in Bombax ceibaby regulating SOD family genes. PeerJ - (Peer Journal) 12:e17849, doi: 10.7717/peerj.17849.
  84. Luo, L.J. 2010. Breeding for water-saving and drought-resistance rice (WDR) in China. Journal of Experimental Botany, 61(13), 3509–3517, doi:10.1093/jxb/erq185.
  85. Ma, X.L., Wang, Y.J., Xie, S.L., Wang, C. and Wang, W. 2007. Glycine betaine application ameliorates negative effects of drought stress in tobacco. Russian Journal of Plant Physiology, 54, 472. doi: 10.1134/S1021443707040061.
  86. Mak, M., Babla, M., Xu, S.C., O’Carrigan, A., Liu, X.H., Gong, Y.M., Holford, P. and Chen, Z.H., 2014. Leaf mesophyll K+, H+ and Ca2+ fluxes are involved in drought-induced decrease in photosynthesis and stomatal closure in soybean. Environmental and Experimental Botany, 98, 1-12. Doi: org/10.1016/j.envexpbot.2013.10.003.
  87. Mansoor S, Ali Wani O, Lone JK, Manhas S, Kour N, Alam P, Ahmad A, Ahmad P 2022. Reactive oxygen species in plants: from source to sink. Antioxidants 11(2):225. Doi:org/10.3390/ antiox11020225.
  88. Martın-Rodrıguez, A., Morcillo, R.L., Vierheilig, H., Ocampo, J.A. Muller, J.L. and Jose Garrido, M.G. 2011. Ethylene-dependent⁄ethylene-independent ABA regulation of tomato plants colonized by arbuscular mycorrhiza fungi. New Phytologist, 190: 193–205, doi: 10.1111/j.1469-8137.2010.03610.x.
  89. Marschner, H. 2012. Marschner's mineral nutrition of higher plants, edited by: Petra Marschner, Academic Press/ Elsevie, third edition, SBN: 978‑0‑12‑384905‑2 (hardback).
  90. Martín-Rodríguez, , Huertas, R., Ho-Plagaro, T., Ocampo, J.A., Tureckova, V., Tarkowska, D., Ludwig-Muller, J., José, M. and García-Garrido, J.M. 2016, Arbuscular mycorrhizal symbiosis induces strigolactone biosynthesis under drought and improves drought tolerance in lettuce and tomato. Plant Cell and Environment. 39, 441–452. doi:10.1111/pce.1263.
  91. Mathur, N. and Yas. A.V. 1995. Influence of VA mycorrhizae on net photosynthesis and transpiration of Ziziphus mauritiana. Journal of Plant Physiology, 147, 328-330. doi: 10.1016/S0176-1617(11)82161-9.
  92. Maurya, A.K., 2020. Oxidative stress in crop plants, In: Agronomic Crops: Stress responses and tolerance; Hasanuzzaman, M., Ed.; Springer: Singapore, 2020; pp. 349–380. doi: 10.1007/978-981-15-0025-1.
  93. Maurel, C., Verdoucq, L., Luu, D.T. and Santoni, V. 2008. Plant aquaporins: membrane channels with multiple integrated functions. Annual Review of Plant Biology, 59: 595–624. doi:1146/annurev.arplant.59.032607.092734.
  94. McGuiness, P.N., Reid, J.B. and Foo, E. 2019. The role of gibberellins and brassinosteroids in nodulation and arbuscular mycorrhizal associations. Frontiers in Plant Science, 10, 443790. doi: org/10.3389/fpls.2019.00269.
  95. Mitra, D., Priyadarshini, A., Senapati, A. Behera, S., Chatterjee, I., Das Mohapatra, P.K. and Panneerselvam, P. 2022. Ecological importance of strigolactones hormone on arbuscular mycorrhizal fungi symbiosis in plant. Journal of Microbiology Research, 9, 160–163. doi: org/10.18231/j.ijmr.2022.029.
  96. Mittler, R. 2017. ROS are good. Trends in Plant Science, A Cell Press Journal, 22(1), 11-19. doi: 10.1016/j.tplants.2016.08.002.
  97. Mhamdi, A. and  Van Breusegem, F. 2018. Reactive oxygen species in plant development. Development, 145(15), dev164376. doi: 10.1242/dev.164376.
  98. Mohanta, T.K., Bae, H. 2017. Cloning and characterization of auxin efflux carrier genes EcPIN1a and EcPIN1b from finger millet. Eleusine coracana Biology, Environmental Science, 7:51. doi:10.1007/s13205-017-0689-6.
  99. Mosse, B. and Hyman. D.S. 1971. Plant growth responses to vesicular-arbuscular mycorrhizae, in unsterilized field soils. New Phytologist,70: 29-34. doi: 10.1111/j.1469-8137.1971.tb02505.x.
  • Mohamed, H.I. and Latif, H.H. 2017. Physiology and Molecular Biology of Plants. 23(3):545–556, doi:10.1007/s12298-017-0451-x.
  • Mosse, B. and Hyman. D. S. 1971. Plant growth responses to vesicular-arbuscular mycorrhizae. in unsterilized field soils. New Phytologist,70: 29-34. https://www.jstor.org/stable/2431048.
  • Nadian, H. 2025. Phosphorus uptake and transport mechanism in symbiotic plants with arbuscular mycorrhizal fungi (Knowns and unknowns). Journal of Soil Biology, 12(2), doi: 10.22092/SBJ.2024.36 6288.267.[In Persian].
  • Nadian H., Smith S.E., Alston, A.M. and Murray R.S. 1997. Effects of soil compaction on plant growth, phosphorus uptake and morphological characteristics of vesicular-arbuscular mycorrhizal colonisation of Trifolium subterraneum. New Phytologist, 135: 303-311. doi:1046/j.1469‑8137.1997.00653.x.
  • Nadian, H. 2011. The effect of drought stress and mycorrhizal symbiosis on growth and phosphorus uptake by two different sorghum cultivars in root morphology. Journal of Water and Soil Science, Esfahan University of Technology, 57: 25-30 doi: 20.1001.1.24763594.1390.15.57.10.2, [In Persian].
  • Nadian, H., Haydari, M., Gharineh, M.H., Daneshvar, M.H. 2013a. Effect of different levels of sodium chloride and mycorrhizal and uptake of P, Na and K by saffron (Crucus sativus, L.). Journal of Plant Production (Journal of Scientific Agriculture), 36(2), doi: pdf [In Persian].
  • Nadian, H. Fathi, G. and Abdollahi, 2013b. Phosphorus Inflow into Two Species of Clover Root with Different Morphology Colonized by AM Fungi. Iran Agricultural Research, 32(1), 41-54 Shiraz University. doi: 22099/iar.2013.1816.
  • Nath, M., Bhatt, D., Prasad, R., Gill, S.S., Anjum, N.A. and Tuteja, T. 2016. Reactive oxygen species generation-scavenging and signaling during plant-arbuscular mycorrhizal and piriformospora indica interaction under stress condition. Frontier in Plant Science, 7:1574. doi: 10.3389/fpls.2016.01574.
  • Nawaz, M. and Wang, Z. 2020. Abscisic acid and glycine betaine mediated tolerance mechanisms under drought stress and recovery in axonopus compressus: A New Insight. Scientific Reports, 10: 6942. doi: 10.1038/s41598‑020‑63447‑0.
  • Nazarli, H., Ahmadi, A. and Hadian, J. 2014. Salicylic acid and methyl jasmonate enhance drought tolerance in chamomile plants. Journal of HerbMed Pharmacology, 2014; 3(2), 87-92.  eISSN: 2345-5004. 
  • Nemeskeri, E., Molnar, K., Vigh, R., Nagy, J. and Dobos, A. 2015. Relationships between stomatal behaviour, spectral traits and water use and productivity of green peas (Pisum sativum) in dry seasons. Acta physiologiae plantarum, 37, 1–16. doi:10.1007/s11738-015-1776-0.
  • (2025). Development finance to combat desertification, land degradation and drought (OECD Publishing, Paris). https://doi.org/10.1787/23f62ad9-en.
  • Oh, E., Yamaguchi, S., Hu, J.H., Yusuke, J., Jung, B., Paik, I., Li, H.S., Sun, T.P., Kamia, Y. and Choi, G. 2007. PIL5, a phytochrome-interacting bHLH protein, regulates gibberellin responsiveness by binding directly to the GAI and RGA promoters in ArabidopsisPlant Cell, 19 (4), 1192–1208, doi: 10.1105/tpc.107.050153.
  • Orine, D., Defossez, E., Vergara, F., Uthe, H., van Dam, N.M. and Rasmann, S. 2022. Arbuscular mycorrhizal fungi prevent the negative effect of drought and modulate the growth-defence trade-off in tomato plants. Journal of Sustainable Agriculture and Environment. 1(3), 177-190. doi: org/10.1002/sae2.12018.
  • Paluch-Lubawa, E., Prosicka, B. and Polcyn, W. 2024. Expression patterns of maize PIP aquaporins in middle or upper leaves correlate with their different physiological responses to drought and mycorrhiza. Frontier in Plant Science, 13, doi: 10.3389/fpls.2024.1459016. eCollection 2024.
  • Paluch-Lubawa, E. and Polcyn, W. 2024. Tissue-specific accumulation of PIP aquaporins of a particular heteromeric composition is part of the maize response to mycorrhiza and drought. Scientific Reports, 17, 14(1):21712.  doi: 10.1038/s41598-024-72828-8.
  • Parent, B., Hachez, C., Redondo, E., Simonneau, T., Chaumont, F. and Tardieu, F. 2009. Drought and abscisic acid effects on aquaporin content translate into changes in hydraulic conductivity and leaf growth rate: a trans-scale approach. Plant Physiology, 149, 2000–2012. doi: org/10.1104/pp.108.130682.
  • Park, W., Scheffler, B.E., Bauer, P.J., Campbell, B.T. 2010. Identification of the family of aquaporin genes and their expression in upland cotton (Gossypium hirsutum). BMC Plant Biology, 10:142, http://www.biomedcentral.com/1471-2229/10/142.
  • Peng, Y.Y., Liu, C.Y. and Hao, 2025. Differential effects of arbuscular mycorrhizal fungi on rooting and physiology of Summer Black’Grape Cuttings. Horticulturae, 11, 825. Doi: org/10.3390/horticulturae11070825.
  • Pirasteh-Anosheh, H., Saed-Moucheshi, A., Pakniyat, H. and Pessarakli, M. 2016. Stomatal responses to drought stress. In: Ahmad, P. (Ed.), Water Stress and Crop Plants: A Sustainable approach. John Wiley & Sons, Ltd, Chichester, UK, 24–40, https:// doi.org/10.1002/9781119054450.ch3.
  • Pons, C. and Muller, C. 2022. Impacts of drought stress and mycorrhizal inoculation on the performance of two spring wheat cultivars. Plants, 11, 2187. doi:org/10.3390/plants11172187.
  • Porcel, R., Arco, R., Cano, C., Bago, A. and Ruiz-Lozano, J.M. 2006. Identification of a gene from the arbuscular mycorrhizal fungus Glomus intraradicesencoding for a 14-3-3 Protein that is up-regulated by drought stress during the AM symbiosis. Microbial Ecology, 52(3), 575-582, doi: 10.1007/s00248-006-9015-2. 
  • Porcel, R., Azcon, R. and Ruiz-Lozano, J.M. 2005. Evaluation of the role of genes encoding for dehydrin proteins (LEA D-11) during drought stress in arbuscular mycorrhizal Glycine max and Lactuca sativa plants. Journal of Experimental Botany, 56: 1933-1942. doi: org/10.1093/jxb/eri188.
  • Pozo, M.J., Cordier, C., Dumas-Gaudot, E., Gianinazzi S., Barea, J.M. and Azcón-Aguilar, C. 2002. Localized versus systemic effect of arbuscular mycorrhizal fungi on the defence responses to Phytophthorainfection in tomato plants. Journal of Experimental Botany53, 525–534. doi: org/10.1093/jexbot/53.368.525.
  • Quiroga, , Erice, G., Aroca, R., Chaumont, F., and Ruiz-Lozano, J.M. 2017. Enhanced drought stress tolerance by the arbuscular mycorrhizal symbiosis in a drought-sensitive Maize Cultivar Is Related to a Broader and Differential Regulation of Host Plant aquaporins than in a drought-tolerant cultivar. Frontier in Plant Science, 8,1056, Open access, doi: 10.3389/fpls.2017.01056.
  • Quigley, F., Rosenberg, J.M., Shachar-Hill, Y. and Bohnert, H.J. 2001. Genome Biology, 7;3(1):17. doi:1186/gb-2001-3-1-research0001.
  • Reis, A.D.P., Carvalhob, R.F., Costaa, I.B., Girioa, R.J.S., Gualbertoa, R., Spersa, R.C. and Gaiona, L.A. 2022. Hydrogen peroxide is involved in drought stress long-distance signaling controlling early stomatal closure in tomato plants. Brazilian Journal of Biology, 82, e267343, https://doi.org/10.1590/1519-6984.267343.
  • Reuscher, S., Akiyama, M., Mori, C., Aoki, K., Shibata, D., Shiratake, K. 2013. Genome-wide identification and expression analysis of aquaporins in tomato. PLoS One, 19, 8(11), e79052. doi: 10.1371/journal.pone.0079052. eCollection 2013.
  • Rodriguez-Gacio,, Matilla-Vázquez, M.A. and Matilla, A.J. 2009. Seed dormancy and ABA signaling. Plant Signaling and Behavior,4, 1035–1048, doi:10.4161/psb.4.11.9902.
  • Romera,R., Ruiz-Lozano, J.M., Zamarreño, A.M., Garcia-Mina, J.M. and Aroca-Ramal, R. 2016. Arbuscular mycorrhizal symbiosis and methyl jasmonate avoid the inhibition of root hydraulic conductivity caused by drought. Mycorrhiza, 26, 111–122. doi: 10.1007/s00572-016-0728-2.
  • Roussel, H., Bruns, S., Gianinazzi-Pearson, V., Hahlbrock, K. and Franken, P. 1997. Induction of a membrane intrinsic protein-encoding mRNA in arbuscular mycorrhiza and elicitor-stimulated cell suspension cultures of parsley. Plant Science, 126:203–210.
  • Ruiz-Sanchez, M., Armada, E., Munoz, Y., Garcia de Salamone, I.E,Aroca, R., Ruíz-Lozano, M. and Azcón, R. 2011. Azospirillum and arbuscular mycorrhizal colonization enhance rice growth and physiological traits under well-watered and drought conditions. Journal of Plant Physiology. 168(10), 1031–1037, doi: 10.1016/j.jplph.2010.12.019. Epub 2011 Mar 5. 
  • Ruiz-Sanchez, , Aroca, R., Munoz, Y., Polon, R. and Ruiz-Lozano, J.M. 2010. The arbuscular mycorrhizal symbiosis enhances the photosynthetic efficiency and the antioxidative response of rice plants subjected to drought stress. Journal of Plant Physiology, 167: 862-869.  https://doi.org/10.1016/j.jplph.2010.01.018.
  • Ruiz-Lozano, J.M., Collados, C., Barea, J.M., Azcón, R. 2001. Clonig of cDNAs encoding SODs from lettuce plants which show differential regulation by arbuscular mycorhizal symbiosis and by drought stress. Journal of Experimental Botany 52, 2241–2242.
  • Ruiz-Lozano, J.M., Perálvarez, M.C., Aroca, R. and Azcón, R. 2011. The application of a treated sugar beet waste residue to soil modifies the responses of mycorrhizal and non-mycorrhizal lettuce plants to drought stress. Plant Soil, 346, 153–166. doi: 10.1007/s11104-011-0805-z.
  • Ruiz-Lozano, J.M., Azcon, R., Gomez, M. 1995. Effects of arbuscular mycorrhizal Glomus species on drought tolerance: physiological and nutritional plant responses. Applied and Environmental Microbiology, 61, 456–60. doi: 1128/aem.61.2.456-460.1995.
  • Ruiz-Lozano, J.M., Azcón R. and Palma, J.M. 1996. Superoxide dismutase activity in arbuscular mycorrhizal Lactuca sativaplants subjected to drought stress. New Phytologist,134, 327–333.  doi: 1111/j.1469-8137.1996.tb04637.x 
  • Ruiz-Lozano, D., Porcel, R. and Aroca, R. Does the enhanced tolerance of arbuscular mycorrhizal plants to water deficit involve modulation of drought-induced plant genes? New Phytologist, 171 (4): 693-698. doi: org/10.1111/j.1469-8137.2006.01841.x.
  • Ruiz-Lozano, J.M., Porcel, R., Bárzana, G., Azcón, R. and Aroca, R., 2012. Contribution of arbuscular mycorrhizal symbiosis to plant drought tolerance: State of the art. In: Aroca, R. (ed.) Plant Responses to Drought Stress. Springer, Berlin, pp.335–362. doi: 10.1007/978-3-642-32653-0_13.
  • Safir, G.R., Boyer, J.S. and Gerdemann. J.W. 1971. Mycorrhizal enhancement of water transport in soybean. Science, 172, 581-583. doi: org/10.1126/science.172.3983.581.
  • Said, A.F., Alwutayd, K.M., KaAma, N., and Ghanem, O.M. 2025. Diversity and effectiveness of arbuscular mycorrhizal fungi species in alleviating drought stress in tomato. Notulae Botanicae Horti Agrobotanici Cluj-Napoca, 53(2), 14382. https://doi.org/10.15835/nbha53214382.
  • Saijo, Y. 2001. A Ca2+-dependent protein kinase that endows rice plants with cold- and salt-stress tolerance functions in vascular Plant Cell Physiology, 42(11), 1228–1233. doi: org/10.1093/pcp/pce158.
  • Salas-González, , Reyt, G., Flis, P., Custódio, V., Gopaulchan, D., Bakhoum, N., Dew, T.P.  Suresh , K., Franke, R.B., Dang, J.L., Salt, D.E. and Castrillo, G. 2021. Coordination between microbiota and root endodermis supports plant mineral nutrient homeostasis. Science, 8: 371 (6525) eabd0695, doi: 10.1126/science.abd0695. 
  • Seo, M., Hanada, A., Kuwahara, A., Endo, A., Okamoto, M., Yamauchi, Y., North, H., Marion-Poll, A., Sun, T.P., Koshiba, T., Kamiaya, Y., Yamaguchi, S. and Nambara, E. 2006. Regulation of hormone metabolism in Arabidopsisseeds: phytochrome regulation of abscisic acid metabolism and abscisic acid regulation of gibberellin metabolism. Plant Journal,48, 354–366. doi:10.1111/j.1365-313X.2006.02881.x.
  • Sepahvand, T., Etemad, V., Matinizadeh, M., Shirvany, A. 2021. Symbiosis of AMF with growth modulation and antioxidant capacity of Caucasian Hackberry (Celtis Caucasica ) seedlings under drought stress. Central Asian Journal of Environmental Science and Technology Innovation, 2(1), 20-35. doi: org/10.22034/CAJESTI.2021.01.03.
  • Shavrukov, Y., Kurishbayev, A., Jatayev, S., Shvidchenko, V., Zotova, L., Koekemoer, F., de Groot, S., Soole, K. and Peter Langridge, P. 2017. Early flowering as a drought escape mechanism in plants: How can it aid wheat production? Frontiers in Plant Science, 8:1950. doi: 10.3389/fpls.2017.01950.
  • Staszek, P. and Gniazdowska, A. 2020. Peroxynitrite induced signaling pathways in plant response to non‑proteinogenic amino acids. Planta, 252:5. https://doi.org/10.1007/s00425-020-03411-4.
  • Smirnoff, N. and Arnaud, D. 2019. Hydrogen peroxide metabolism and functions in plants. New Phytologist, 221, 1197–1214, doi: 10.1111/nph.15488.
  • Sood, M. 2025. Reactive oxygen species (ROS): plant perspectives on oxidative signalling and biotic stress response. Discover Plants, 2,187. doi: org/10.1007/s44372-025-00275-4.
  • Spinoso-Castillo,L., Moreno-Hernandez, M.D.R., Mancilla-Alvarez, E., Sanchez-Segura, L., Sanchez-Paez, R. and Bello-Bello, J.J. 2023. Arbuscular Mycorrhizal Symbiosis improves ex vitro acclimati-zation of sugarcane plantlets (Saccharum Spp.) under drought stress conditions. Plants, 12(3), 687. doi: org/10.3390/plants12030687.
  • Steudle E. 2000. Water uptake by roots: effects of water deficit. Journal of Experimental Botany 51, 1532–1542. doi:org/10.1093/jexbot/51.350.1531.
  • Steudle, E. and Peterson, C.A. 1998. How does water get through roots? Journal of Experimental Botany 49: 775–788. doi: org/10.1093/jxb/49.322.775.
  • Sun, , Li, S., Chen, W., Zhang, J., Zhang, L., Sun, W. and Wang, Z. 2021. Plant dehydrins: expression, regulatory networks, and protective roles in plants challenged by abiotic stress. International journal of Molecular Science, 23(22),12619. doi: 10.3390/ijms222312619.
  • Sun, , Liu, X., Kitagawa, Y., Calamita,G., Ding, X. 2024. Plant aquaporins: Their roles beyond water transport. The Crop Journal, 12(3), 641-655. https://doi.org/10.1016/j.cj.2024.04.00.
  • Tahiri, A. Meddich, A. Raklami, A., Alahmad, A., Bechtaoui, B., Anli, M., Göttfert, M., Heulin, T., Wafa Achouak, W. and Oufdou, K.  Assessing the potential role of compost, PGPR, and AMF in improving tomato plant growth, yield, fruit quality, and water stress tolerance. Journal of Soil Science and Plant Nutrition, 22(1), 743–764. doi: 10.1007/s42729‑021‑00684‑w.
  • Tominaga, T., Yamaguchi, K., Shigenobu, S., Yamato, M. and Kaminaka, H. 2020a. The effects of gibberellin on the expression of symbiosis-related genes in Paris-type arbuscular mycorrhizal symbio­sis in Eustoma grandiflorum. Plant Signaling & Behavior 15, 1784544. doi: org /10.1080/15592324.2020.1784544.
  • Tominaga, T., Miura, C., Takeda, N., Yuri Kanno, Y., Takemura, Y., Seo, M., Yamato, M. and Kaminaka, 2020b. Gibberellin promotes fungal entry and colonization during Paris-type arbuscular mycorrhizal symbiosis in Eustoma grandi­florum. Plant and Cell Physiology, 61, 565–575. doi: org/10.1093/pcp/pcz222.
  • Tyagi, S., Shumayla, Singh, S.P. and Upadhyay, S.K. 2019. Role of superoxide dismutases (SODs) in stress tolerance in plants. Springer Nature, In:  Molecular Approaches in Plant Biology and Environmental Challenges, 51-77. doi: org/10.1007/978-981-15-0690-1_3.
  • Uehlein, N, Fileschi, K., Eckert, M., Bienert, G.P., Bertl, A. and Kaldenhoff, R. 2007. Arbuscular mycorrhizal symbiosis and plant aquaporin expression. Phytochemistry, 68:122–129. doi: 1016/j.phytochem.2006.09.033.
  • UN Convention to Combat Desertification (UNCCD). UN Press Release, 9 December 2024.
  • Urao, T., Katagiri, T., Mizoguchi, T., Yamaguchi-Shinozaki, K., Hayashida, N., Shinozaki, K. Two genes that encode Ca2+-dependent protein kinases are induced by drought and high-salt stresses in Arabidopsis thaliana, Molecular Genetics and Genomics, 244(4), 331–340. doi: org/10.1007/BF00286684.
  • Volaire, F. 2018. A unified framework of plant adaptive strategies to drought: crossing scales and disciplines. Global Change Biology, 24(7), 2929–2938. doi: org/10.1111/gcb.14062.
  • Wang, C., El-Shetehy, M., Shine, M.B., Yu, K., Navarre, D., Wendehenne, D., Kachroo A. and Kachroo, P. 2014. Free radicals mediate systemic acquired Cell Reports 7(2), 3,48–355. doi: org/10.1016/j.celrep.2014.03.032
  • Wang, B.L., Tang, X.Y., Cheng, L.Y., Zhang, A.Z., Zhang, W.H., Zhang, F.S., Liu, J.Q., Cao, Y., Allan, D.L., Vance, C.P. and Shen, J.B. 2010. Nitric oxide is involved in phosphorus deficiency-induced cluster-root development and citrate exudation in white lupin. New Phytologist, 2010, 187:1112–1123. doi: 10.1111/j.1469-8137.2010.03323.x.
  • Wang, Q., Liang, X., Dong, Y., Xu, L., Zhang, X., Kong, J. and Liu, S. 2013. Effects of exogenous salicylic acid and nitric oxide on physiological characteristics of perennial ryegrass under cadmium stress. Journal of Plant Growth Regulation, 32, 721–731. doi: org/10. 1007/s00344-013-9339-3.
  • Wang, D., Ni, Y., Xi., K., Li, Y., Wu, W., Shan, H., Cheng, B. and Li, X. 2024. Aquaporin ZmTIP2;3 promotes drought resistance of maize through symbiosis with arbuscular mycorrhizal fungi. International Journal of Molecular Science,25(8), 4205. doi: org/10.3390/ijms25084205.
  • Wasternack, C., and Song, S. 2017. Jasmonates: biosynthesis, metabolism, and signaling by proteins activating and repressing transcription. Journal of Experimental Botany, 68(6), 1303–1321. doi: org/10.1093/jxb/erw443.
  • Wei,  S., Javed, T., Liu, T.T., Ahmad Ali, A. and Gao, S.J. 2025. Mechanisms of Abscisic acid (ABA)-mediated plant defense responses. Plant stress, 15, 100724. doi: org/10.1016/j.stress.2024.100724.
  • Wen, Y., Zhou, L.J., Xu, Y.J., Hashem, A., Abd_Allah, E.F. and Wu, Q.S. 2024. Growth performance and osmolyte regulation of drought-stressed walnut plants are improved by mycorrhiza. Agriculture, 14, 367. doi: org/10.3390/agriculture14030367.
  • Wilkinson,, Davies and W.J. 2010. Drought, ozone, ABA and ethylene: new insights from cell to plant to community. Plant, Cell & Environment, 33, 510–525. doi: org/10.1111/j.1365-3040.2009.02052.x
  • Wu, H.H., Zou, Y.N., Rahman, M.M., Ni, Q.D., and Wu, Q.S. 2017. Mycorrhizas alter sucrose and proline metabolism in trifoliate orange exposed to drought stress. Scientific Reports, 7, 42389. doi: 10.1038/srep42389.
  • Xie, W., Hao1, Z., Zhou, X. Jiang, X., Xu, L., Wu, S., Zhao, A., Xin Zhang, X. and Chen, B. 2018. Arbuscular mycorrhiza facilitates the accumulation of glycyrrhizin and liquiritin in Glycyrrhiza uralensis under drought stress. Mycorrhiza, Springer-Verlag GmbH Germany, part of Springer Nature 2018. doi: org/10.1007/s00572-018-0827-y.
  • Xu, L., Li, T., Wu, Z., Feng, H., Yu, M., Zhang, X. and Hen, B.C. 2018. Arbuscular mycorrhiza enhances drought tolerance of tomato plants by regulating the 14-3-3 genes in the ABA signaling pathway. Applied Soil Ecology, 125, 213-221, doi: org/10.1016/j.apsoil.2018.01.012.
  • Xu, D., Duan, X. Wang, B., Hong, B., Ho, T.H.D. and Wu, R. 1996. Expression of a late embryogenesis abundant protein gene, HVA7, from barley confers tolerance to water deficit and salt stress in transgenic rice. Plant Physiology, 110, 249-257. doi: 10.1104/pp.110.1.249.
  • Ye, Q., Wang, H. and Li, H. 2022. Arbuscular Mycorrhizal fungi improve growth, photosynthetic activity, and chlorophyll fluorescence of Vitis vinifera cv. ecolly under drought stress. Agronomy, 12(7), 1563, doi: org/10.3390/agronomy12071563.
  • Zhang, S.W., Li, C.H., Cao, J., Zhang, Y.C., Zhang, S.Q., Xia, Y.F., Sun, D.Y. and Sun, Y. 2009. Altered architecture and enhanced drought tolerance in rice via the down-regulation of indole-3-acetic acid byTLD1/OsGH3.13 Activation. Plant Physiology, 151, 1889–1901. doi: org/10.1104/pp.109.146803.
  • Zhang, Z., Zhang, J. and Huang, Y., 2014. Effect of arbuscular mycorrhizal fungi on the drought tolerance of Cyclobalanopsis glauca seedlings under greenhouse conditions. New Forests, 45(4), 545-556. doi:1007/s11056-014-9417-9.
  • Zhang, , Yu, Y., Zhang, J., Qian, X., Li, X. and Sun, X. 2024. Recent progress regarding jasmonates in tea plants: Biosynthesis, signaling, and function in stress responses. International Journal of Molecular Sciences, 25, 1079. doi: org/10.3390/ijms25021079.
  • Zhu, Y.X., Yang, L., Liu, N., Yang, J., Zhou, X.K., Xia, Y.C., He, Y., He, Y.Q., Gong, H.J. Ma, D.F. and Yin, J.L. 2019. Genome-wide identification, structure characterization, and expression pattern profiling of aquaporin gene family in cucumber. BMC Plant Biology, 7,19(1), 345. doi: 10.1186/s12870-019-1953-1.
  • Zhu, , Lu, P., Jiang, Y., Wang, M. and Zhang, L. 2014. Botany, Effects of brassinosteroid on antioxidant system in Salvia miltiorrhiza under drought stress. Journal of Research in Agriculture and Animal Science, 2(1), 01-06, www.questjournals.org, ISSN(Online): 2321-9459.
  • Zhao, H., Zhang, Y. and Zheng, Y. 2022. Integration of ABA, GA, and light signaling in seed germination through the regulation of ABI5. Frontier in Plant Science, 13, doi: org/10.3389/fpls.2022.1000803.
  1. Zou, N., Wan, Y.X., Zheng, F.L., Cheng, X.F.,  Hashem, A. and Wu, Q.S. 2025. Mycorrhizal trifoliate orange plants tolerate soil drought by enhancing photosynthetic physiological activities and reducing active GA3 levels. Tree Physiology, 45:8, tpaf073, doi: org/10.1093/treephys/tpaf073.
  2. Zou, Y.N., Huang, Y.M., Wu, Q.S. and He, X.H. 2015. Mycorrhiza-induced lower oxidative burst is related with higher antioxidant enzyme activities, net H2O2 effluxes, and Ca2+ influxes in trifoliate orange roots under drought stress. Mycorrhiza, 25(2),143-52, doi: 10.1007/s00572-014-0598-z. Epub 2014 Aug 3.
 

 

 

 

آمار
تعداد مشاهده مقاله: 216
تعداد دریافت فایل اصل مقاله: 123


counter
سامانه مدیریت نشریات علمی. طراحی و پیاده سازی از سیناوب