بررسی تغییرات در بیان ژن P5CS، و برخی صفات فیزیولوژیک و بیوشیمایی گندم در پاسخ به باکتری آزوسپیریلوم و قارچ پیریفورمواسپورا ایندیکا تحت تنش شوری

نوع مقاله : مقاله پژوهشی

نویسندگان

1 دانشیار گروه زراعت و اصلاح نباتات، دانشکده کشاورزی، دانشگاه ایلام، ایلام، ایران.

2 دانش آموخته دکتری آگروتکنولوژی، گروه زراعت و اصلاح نباتات، دانکشده کشاورزی، دانشگاه ایلام، ایلام، ایران.

چکیده

مقدمه: تنش‌های محیطی از بزرگ‌ترین تهدیدها برای ثبات در تولید محصولات کشاورزی محسوب می‌گردند. تغییرات آب و هوایی شدت تنش‌های محیطی را در مناطق مختلف جهان افزایش داده است. خشکی و شوری مهمترین تنش‌های محیطی مناطق خشک و نیمه خشک جهان هستند. در دهه‌های اخیر راهکارهای مختلفی جهت کاهش اثرات تنش شوری بر گیاهان زراعی معرفی شده است. استفاده از میکروارگانیسم‌های مفید خاک از جمله روش‌های  توصیه شده می‌باشد.
مواد و روش‌ها: در مطالعه حاضر اثر باکتری آزوسپیریلوم و قارچ پیریفورمواسپورا ایندیکا در تلقیح جداگانه و توامان آنها به‌منظور بررسی تغییرات مولکولی، بیوشیمایی و فیزیولوژی گندم تحت اعمال و عدم اعمال تنش شوری بررسی گردید. آزمایش به‌صورت فاکتوریل سه عاملی هر کدام با دو سطح بر پایه طرح کاملاٌ تصادفی و در سه تکرار در گلخانه دانشکده کشاورزی دانشگاه ایلام انجام گرفت. سه تیمار آزمایش شامل تنش شوری (0 و 12 دسی زیمنس بر متر)، تلقیح با باکتری آزوسپیریلوم (108 × 1.8 سلول زنده در هر میلی‌لیتر محیط کشت)، و کاربرد قارچ  پیریفورمواسپورا ایندیکا (10 گرم حاوی کلامیدوسپور در یک کیلوگرم خاک) بود. جهت اعمال تنش شوری 12 دسی زیمنس بر متر از مخلوط دو نمک کلرید کلسیم (65 میلی گرم) و کلرید سدیم (375/0 گرم) استفاده گردید. 21 و 28 روز پس از سبز شدن گندم شوری اعمال گردید.  از گندم رقم پیشگام جهت این آزمایش استفاده گردید. 24 ساعت پس از اعمال دومین آبیاری با آب شور نمونه‌گیری برگ به جهت استخراج RNA کل و مطالعات مرتبط با بیان ژن دلتا-پیرولین-5 کربوکسیلات سینتاز که آنزیم کلیدی در مسیر سنتز پرولین است انجام گرفت. میزان تغییرات پرولین برگ، میزان تولید مالون دی آلدهید، نشت یونی، محتوای آب نسبی برگ، محتوای نیتروژن، میزان رنگیزههای کلروفیلی در پاسخ به میکروارگانیسم‌های مذکور مورد بررسی قرار گرفت. همچنین میزان کلونیزاسیون قارچ و  سوکسینات دهیدروژناز که معیاری از میزان فعالیت زیستی قارچ در ریشه گیاه است نیز مورد سنجش قرار گرفت.
یافته‌ها: درصد کلونیزاسیون قارچ با ریشه گندم رقم پیشگام  9/24 درصد بود. تلقیح با آزوسپیریلوم میزان کلونیزاسیون ریشه با قارچ را 17/3 درصد افزایش داد و به مقدار 29 درصد رساند. شوری تاثیر معنی‌داری بر سوکسینات دهیدروژناز نداشت اما تلقیح با باکتری میزان آن را به صورت معنی‌داری افزایش داد. گیاهان تلقیح شده با قارچ و باکتری از محتوای آب نسبی بیشتر اما نشت یونی و مالون دی آلدهید  کمتری تحت شوری برخوردار بودند. شوری موجب تغییر معنی‌دار در میزان رنگیزه‌های کلروفیلی نگردید؛ هر چند تلقیح با قارچ و باکتری میزان کلروفیل a را افزایش داد. تاثیر شوری، قارچ و باکتری و اثر متقابل آنها بر میزان بیان ژن دلتا-پیرولین-5 کربوکسیلات سینتاز معنی‌دار بود. گیاهان شاهد سطوح بیشتری از بیان ژن را نشان دادند. در حالیکه گیاهان تلقیح شده با قارچ و باکتری از میزان بیان ژن کمتری برخوردار بودند. همچنین کاربرد همزمان باکتری و قارچ بر میزان بیان ژن دلتا-پیرولین-5 کربوکسیلات سینتاز معنی‌دار بود و موجب کاهش بیشتر آن در مقایسه با گیاهانی که فقط با قارچ تلقیح شده بودند گردید. اثر شوری، قارچ و باکتری و نیز اثرات متقابل آنها بر میزان غلظت پرولین برگ معنی‌دار بود. تحت تنش شوری میزان پرولین در گیاهان تلقیح شده با قارچ کمتر از گیاهان شاهد بود. چنین روندی نیز در گیاهان تلقیح شده با باکتری مشاهده گردید به‌طوری که گیاهان تلقیح شده سطوح کمتری از پرولین را تحت تنش شوری نشان دادند.
نتیجه‌گیری: گیاهان تلقیح شده با قارچ و باکتری نسبت به گیاهان شاهد کمتر تحت تاثیر تنش شوری قرار گرفتند که میزان تولید کمتر مالون دی آلدهید و نشت الکترولیت و  بالاتر بودن میزان رنگریزه کلروفیلی و محتوای آب نسبی برگ تایید کننده این نتایج بود. همچنین میزان بیان ژن دلتا-پیرولین-5 کربوکسیلات سینتاز و تجمع پرولین در گیاهان تحت تنش شوری در مقایسه با گیاهان کنترل نشان داد که میکروارگانیسم‌های استفاده شده از طریق دیگری شدت تنش را بر گیاه میزبان کاهش داده‌اند.

کلیدواژه‌ها

موضوعات

عنوان مقاله [English]

Investigation of modulation in P5CS gene expression and some physiological and biochemical traits of wheat in response to Azospirillum and Piriformospora indica inoculation under salinity stress

نویسندگان [English]

  • Mohammad Javad Zarae 1
  • Zohreh Karimi 2
1 Associate Professor, Department of Agronomy and Crop Breeding, Faculty of Agriculture, Ilam University, Ilam, Iran.
2 Ph.D. Graduated in Agrotechnology, Department of Agronomy and Crop Breeding, Faculty of Agriculture, Ilam University, Ilam, Iran.
چکیده [English]

Introduction: Environmental stresses such as salinity are one of the major threats to the sustainability of crop production. Climate change has increased the intensity of these environmental tensions in different regions of the world. Drought and salinity are the critical environmental stresses in arid and semi-arid regions. In recent decades, various strategies have been introduced to reduce the effects of salinity stress on crops. The use of beneficial soil microorganisms is one of these methods.
Materials and methods: In the present study, the effect ofAzospirillum bacteria and Piriformospora indica fungus in separate and combined inoculation was investigated to identify the role of these above-mentioned plant growth-promoting microorganisms in the modulation of the molecular, biochemical, and physiological aspects in wheat under salt stress conditions. A 2×2×2 factorial experimental design was implemented in a completely randomized block design with three replications under greenhouse conditions in 2022. The experiment was conducted in the pot. The main treatments included salt stress imposition, inoculation with and without Azospirillum zaea, and endophytic fungus Piriformospora indica application. Salinity was induced using calcium chloride (65 mg) and sodium chloride (0.375 g) to achieve a salinity level of 12 dS/m. Saline water irrigation was performed 21 and 28 days after wheat germination. Normal water with a salinity of 0.3 decisiemens per meter was used for the control treatment. The wheat cultivar Pishgam was used for this experiment. Twenty-four hours after applying the second salinity treatment, leaf sampling was conducted to extract total RNA and study the expression of the delta-pyrroline-5-carboxylate synthase gene. The changes in leaf proline content and the expression of the gene responsible for proline synthesis (pyrroline-5-carboxylate synthase), malondialdehyde production, ion leakage, leaf relative water content, nitrogen content, and chlorophyll pigment levels in response to applied microorganisms under both salinity and non-salinity conditions were investigated. Additionally, succinate dehydrogenase activity and root colonization rate with fungus were measured under the influence of salinity and inoculation with Azospirillum.
Results: The percentage of fungal colonization in wheat roots of cultivar Pishgam was 24.9%. Inoculation with Azospirillum increased the root colonization rate by 3.17%. Salinity had no significant effect on succinate dehydrogenase, but inoculation with Azospirillum significantly increased succinate dehydrogenase. The plants inoculated with P. indica and Azospirillumhad a higher relative water content in leaves under salt stress. Plants inoculated with bacteria and fungus also had higher nitrogen content. Salinity increased the amount of ion leakage, but inoculation with fungus and bacteria prevented it from increasing. Salinity did not cause a significant change in the amount of chlorophyll pigments, although inoculation with P. indicaand Azospirillum increased the amount of chlorophyll a. The effect of salinity, P. indica, Azospirillum, and their interaction on the expression level of the delta-pyrroline-5 carboxylate synthase gene was significant. Control plants showed higher levels of gene expression, while the inoculated plants with P. indica and Azospirillum had lower gene expression levels. Additionally, the interaction between Azospirillum and P. indica on the expression level of the delta-pyrroline-5 carboxylate synthase gene was significant and it caused a greater decrease compared to the plants that were only inoculated with P. indica. The effect of salinity, P. indica, and bacteria, as well as their mutual effects on leaf proline concentration, was significant. Under salt stress, the amount of proline in the plants inoculated with P. indica was lower than that of the control plants. A similar trend was observed in plants inoculated with bacteria, showing lower levels of proline under salt stress. The expression trend of the delta-pyrroline-5-carboxylate synthase gene was consistent with the trend of proline accumulation under salt stress, indicating the effect of salt induction on the expression of this gene.
Conclusion: The inoculated plants with P. indica and Azospirillum were less affected by induced salt stress than the control plants. The inoculated plants exhibited a lower rate of malondialdehyde and electrolyte leakage, as well as a higher level of chlorophyll pigment and leaf-relative water content. Additionally, P5CS gene expression and proline accumulation in inoculated plants were lower than in control plants.

کلیدواژه‌ها [English]

  • Ion leakage
  • malondialdehyde
  • plants-microbe interaction
  • proline.succinate dehydrogenase

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