مطالعه الگوی بیان ژن bZIP transcription factor 1 و فعالیت برخی آنزیم‌های آنتی-اکسیدانت و تجمع پرولین در ذرت (Zea mays L.) تحت تنش شوری

نوع مقاله : مقاله پژوهشی

نویسندگان

گروه تولید و ژنتیک گیاهی، دانشکده کشاورزی، دانشگاه ارومیه، اومیه، ایران.

10.22126/cbb.2026.13714.1136

چکیده

مقدمه: ذرت (Zea mays L.)، گیاهی دیپلوئید از خانواده گندمیان بوده و از لحاظ فتوسنتز جزو گیاهان C4 می­باشد. تنش شوری سبب افزایش تولید گونه­های فعال اکسیژن (ROS) می­شود که این خود باعث ایجاد تنش ثانویه به نام تنش اکسیداتیو در گیاهان می­شود. افزایش سطح ROS به‌ویژه موجب پراکسیداسیون اسیدهای چرب غیراشباع شده و در نتیجه، فرآورده‌های ثانویه‌ای مانند کتون‌ها، مالون‌دی‌آلدئید و سایر ترکیبات مشتق از آن‌ها تولید می‌شود. پراکسیداسیون لیپیدها و تولید مالون‌دی‌آلدئید به‌عنوان یکی از شاخص‌های مهم آسیب اکسیداتیو، به‌طور گسترده برای ارزیابی میزان حساسیت یا تحمل ژنوتیپ‌های گیاهی به تنش شوری مورد استفاده قرار می‌گیرد. گیاهان برای مقابله با تنش اکسیداتیو ناشی از شوری، مجموعه‌ای از سازوکارهای دفاعی آنزیمی و غیرآنزیمی را فعال می‌کنند. آنزیم‌های آنتی‌اکسیدانت نقش اساسی در سم‌زدایی ROS و حفظ هموستاز اکسیداتیو ایفا می‌کنند. علاوه بر این، در تنش شوری، افزایش سنتز پرولین به‌عنوان یک اسمولیت فعال موجب حفظ تعادل اسمزی و محافظت سلول‌ها در برابر کم‌آبی می‌شود. خانواده‌ی bZIP یکی از گسترده‌ترین گروه‌های عوامل رونویسی در گیاهان محسوب می‌شود که نقشی محوری در پاسخ به تنش‌های زیستی و غیرزیستی ایفا می‌کند. این پژوهش با هدف شناخت پاسخ‌های فیزیولوژیکی، بیوشیمیایی و مولکولی ذرت به تنش شوری، با تمرکز بر فعالیت آنزیم‌های آنتی‌اکسیدانت، تجمع پرولین، پراکسیداسیون لیپیدی و الگوی بیان ژن bZIP transcription factor1  در دو لاین متحمل و حساس تحت تنش شوری ۸ دسی‌زیمنس بر متر انجام شد.
مواد و روش‌ها: دو لاین ذرت شامل لاین متحمل (R10; P14L2) و حساس (S46; MO17) به شوری بر اساس نتایج مطالعات پیشین انتخاب شدند (Ebrahimipour et al., 2020). آزمایش به‌صورت فاکتوریل در قالب طرح پایه کاملاً تصادفی با سه تکرار اجرا گردید. فاکتور اول شامل دو ژنوتیپ ذرت و فاکتور دوم شامل دو سطح شوری (شاهد و 8 دسی‌زیمنس بر متر) بود. گیاهان در اتاق رشد کنترل‌شده دانشکده کشاورزی دانشگاه ارومیه تحت شرایط دمایی 2±25 درجه سانتی‌گراد، رطوبت نسبی 65–60 درصد و دوره نوری 16 ساعت روشنایی و 8 ساعت تاریکی پرورش یافتند. آبیاری گیاهان از مرحله دو برگی با محلول غذایی هوگلند انجام شد. اعمال تنش شوری از مرحله هشت‌برگی به‌مدت 20 روز با استفاده از محلول غذایی حاوی NaCl، 8 دسی‌زیمنس بر متر، انجام گرفت. پس از پایان دوره تنش، نمونه‌برداری از برگ‌های جوان به‌منظور انجام آزمایش‌های بیوشیمیایی و مولکولی انجام و نمونه‌ها بلافاصله در دمای 80- درجه سانتی‌گراد نگهداری شدند. برای بررسی الگوی بیان ژن bZIP transcription factor 1، آغازگرهای اختصاصی ژن هدف و ژن مرجع (اکتین) بر اساس توالی‌های موجود در پایگاه داده NCBI طراحی شدند.
یافته‌ها: نتایج نشان داد که تنش شوری موجب افزایش معنی‌دار پراکسیداسیون لیپیدها (مالون‌دی‌آلدئید) و تجمع پرولین در هر دو لاین شد، به‌طوری که شدت این افزایش در لاین متحمل بیشتر بود. فعالیت آنزیم‌های سوپراکسید دیسموتاز و آسکوربات پراکسیداز و کاتالاز تحت تنش شوری افزایش یافت، در حالی که فعالیت گایاکول پراکسیداز کاهش نشان داد. کاتالاز و آسکوربات نیز پاسخ وابسته به ژنوتیپ از خود بروز دادند و فعالیت آن­ها در لاین حساس نسبت به لاین متحمل افزایش بیشتری داشت. تحلیل همبستگی صفات، ارتباط مثبت و معنی‌دار بین پرولین، مالون‌دی‌آلدئید و فعالیت آنزیم‌های آنتی‌اکسیدانت کلیدی را تأیید کرد. در سطح مولکولی، بیان ژن bZIP transcription factor1 به‌طور معنی‌داری تحت تأثیر زمان، لاین و اثر متقابل آن‌ها قرار گرفت؛ به‌طوری که لاین حساس افزایش بیان زودهنگام و گذرا و لاین متحمل افزایش بیان تأخیری اما پایدار این ژن را نشان داد.
نتیجه‌گیری: در مجموع، نتایج این پژوهش نشان می‌دهد که تحمل شوری در ذرت حاصل تعامل هماهنگ میان تنظیم سیستم آنتی‌اکسیدانت، تجمع اسمولیت‌ها و کنترل زمانی بیان ژن‌های تنظیم‌کننده پاسخ به تنش است و ژنbZIP transcription factor1  می‌تواند به‌عنوان یک نشانگر مولکولی کلیدی در بهبود تحمل شوری مورد توجه قرار گیرد.

کلیدواژه‌ها

موضوعات

عنوان مقاله [English]

Studying bZIP transcription factor 1 gene expression pattern and the activity of some antioxidant enzymes and proline accumulation in maize (Zea mays L.) under salinity stress

نویسندگان [English]

  • Marjan Jannatdoust
  • Reza Darvishzadeh
Department of Plant Production and Genetics, Faculty of Agriculture, Urmia University, Urmia, Iran.
چکیده [English]

Introduction: Maize is a diploid species belonging to the Poaceae family and is classified as a C4 plant in terms of its photosynthetic pathway. Salinity stress results in the increased production of ROS, which in turn triggers a secondary stress known as oxidative stress in plants. Elevated ROS levels specifically induce the peroxidation of unsaturated fatty acids, resulting in the production of secondary products such as ketones, MDA, and other derived compounds. Lipid peroxidation and MDA production, as critical indicators of oxidative damage, are widely utilized to evaluate the degree of sensitivity or tolerance of plant genotypes to salinity stress. To combat salinity-induced oxidative stress, plants activate a suite of enzymatic and non-enzymatic defense mechanisms. Antioxidant enzymes play a fundamental role in ROS detoxification and the maintenance of oxidative homeostasis. Additionally, under salinity stress, the increased synthesis of proline as an active osmolyte facilitates the maintenance of osmotic balance and protects cells against dehydration. The bZIP family represents one of the most extensive groups of transcription factors in plants, playing a crucial role in responding to both biotic and abiotic stresses. This research was conducted to investigate the physiological, biochemical, and molecular responses of maize to salinity stress, focusing on antioxidant enzyme activity, proline accumulation, lipid peroxidation, and the expression pattern of the bZIP transcription factor 1 gene in two tolerant and sensitive lines under a salinity stress.
Materials and methods: Two maize lines, a tolerant line (R10; P14L2) and a sensitive line (S46; MO17), were selected based on the results of previous studies. The experiment was implemented as a factorial based on a CRD with three replications. The first factor consisted of the two maize genotypes, and the second factor included two salinity levels (control and 8 dS/m). Plants were grown in a controlled growth chamber at Urmia University, under temperature conditions of 25°C, a relative humidity of 60–65%, and a photoperiod of 16 hours of light and 8 hours of darkness. Irrigation was performed using Hoagland nutrient solution starting from the two-leaf stage. Salinity stress was applied from the eight-leaf stage for a duration of 20 days using a nutrient solution containing NaCl at a concentration of 8dS/m. Following the stress period, young leaves were sampled for biochemical and molecular analyses, and samples were immediately stored at -80°C. To investigate the expression pattern of the bZIP transcription factor 1 gene, specific primers for the target gene and the reference gene (actin) were designed based on sequences available in the NCBI.
Results: The results demonstrated that salinity stress significantly increased lipid peroxidation, as indicated by malondialdehyde accumulation, as well as proline content in both lines, with a greater magnitude of increase observed in the tolerant line. The activities of superoxide dismutase, ascorbate peroxidase, and catalase were enhanced under salinity stress, whereas guaiacol peroxidase activity showed a decline. Catalase and ascorbate peroxidase exhibited genotype-dependent responses, with higher activity levels recorded in the sensitive line compared to the tolerant line. Correlation analysis revealed significant positive relation among proline content, malondialdehyde levels, and the activities of key antioxidant enzymes. At the molecular level, the expression of the bZIP transcription factor1 gene was significantly affected by time, genotype, and their interaction; the sensitive line displayed an early and transient upregulation of gene expression, while the tolerant line exhibited a delayed but sustained induction of this gene.
Conclusion: Overall, these findings indicate that salinity tolerance in maize is achieved through a coordinated interaction between antioxidant defense regulation, osmolyte accumulation, and temporal control of stress-responsive regulatory gene expression, and that bZIP may serve as a key molecular marker for improving salinity tolerance in maize.

کلیدواژه‌ها [English]

  • : Abiotic stress
  • Antioxidant enzymes
  • Gene expression
  • Maize
  • Proline and Real-time PCR

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