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Phylogeny and Character Evolution of the Genus Geum L. (Family Rosaceae) from Iran: Evidence from Analyses of Plastid and Nuclear DNA Sequences

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Phylogeny and Character Evolution of the Genus Geum L. (Family Rosaceae) from Iran: Evidence from Analyses of Plastid and Nuclear DNA Sequences Archive of SID Taxonomy and Biosystematics, 10th Year, No. 35, Summer 2018 1 Phylogeny and Character Evolution of the Genus Geum L. (Family Rosaceae) from Iran: Evidence from Analyses of Plastid and Nuclear DNA Sequences Marzieh Beygom Faghir 1*, Reyhaneh Pourmojib 2, Robabeh Shahi Shahvan 3 1 Assistant Professor, Department of Biology, Faculty of Science, University of Guilan, Rasht, Iran 2 M. S. Student of Ecological systematic_ plant biology, Faculty of Science, University of Guilan, Rasht, Iran 3 Ph. D. Student of Ecological systematic_ plant biology, Faculty of Biology, University of Tarbiat Modares, Tehran, Iran Abstract In the present study, molecular phylogeny of the genus Geum L. in Iran, including 5 species (G. kokanicum, G. iranicum G. heterocarpum, G. rivale and G. urbanum) from two subgenera (Orthostylus Fisch & Mey and Geum) were studied using nrDNA ITS and cpDNA rpl32-trnL(UAG) and combined sequence data. Phylogenetic analyses were performed using maximum parsimony approach as implemented in PAUP*, Bayesian method using MrBayes programm and Maximum Likeliood analysis using RaxmlGUI software. The parsimony analysis led to the formation of nrDNA ITS majority-rule consensus tree with a main clade (A), comprising Geum species and its related genera especially Coluria R. Br., Acomastylis Greene, Erythrocoma Greene and Novosieversia F. Bolle., arranged in two monophyletic groups. In all obtained trees, the two representatives of subgenus Geum completely were resolved and formed monophyletic group. While 3 species of the subgenus Orthostylus formed both tritomy (in nrDNA ITS sequence data analysis) and monophyletic group (in cpDNA rpl32-trnL (UAG) and combined sequence dataanalysis). The results of the present study showed that phylogenetic relationship of the genus is strongly under the influence of hybridization and polyploidy (allopolyploidy). The current result support circumscriptions of the genus presented in flora Iranica and flora of Iran. In this study, the evolutionary trends of fruits morphology, exin sculpturing, seed micromorphology and petiole anatomy were evaluated. Keywords: Phylogeny, Geum, Rosaceae, nrDNA ITS, cpDNA rpl32-trnL (UAG). * [email protected] Copyright©2019, University of Isfahan. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/BY-NC-ND/4.0), which permits others to download this work and share it with others as long as they credit it, but they cannot change it in any way or use it commercially. www.SID.ir Archive of SID تاکسونومی و بیوسیستماتیک، سال دهم، شماره سی و پنجم، تابستان 1397، صفحه -1 22 دريافت مقاله: 18/06/1396 پذيرش نهايی: 1397/10/23 فیلوژنی و تکامل صف تها در سردۀ .Geum L از تیرۀ گلسرخیان در ایران: شواهدی بر اساس تجزیه وتحلیلهای توالی DNA کلروپﻻستی و هستهای 3 2 *1 مرضیه بیگم فقیر ، ريحانه پورمجیب ، ربابه شاهی شاووان 1 استاديار گروه زيستشناسی، دانشکده علوم، دانشگاه گیﻻن، رشت ، ايران 2 دانشجوی کارشناسی ارشد بیوسیستماتیک اکولوژی، دانشکده علوم، دانشگاه گیﻻن، رشت، ايران 3 دانشجوی دکتری بیوسیستماتیک اکولوژی، دانشکده علوم، دانشگاه تربیت مدرس، تهران، ايران چکیده در پژوهش حاضر، فیلوژنی مولکولی سردۀ .Geum L از ايران شامل 5 گونۀ G. iranicum ،G. kokanicum، G. rivale ،G. heterocarpum و G. urbanum متعلق به دو زيرسردۀ )Orthostylus Fisch and Mey وGeum) با استفاده از دادههای حاصل از توالی cpDNA rpl32-trnL(UAG) ،nrDNA ITS و ترکیبی مطالعه شدند. * تجزيه وتحلیلهای فیلوژنی با استفاده از روشهای بیشینه صرفهجويی تعبیهشده در نرمافزار PAUP، بايزين با استفاده از نرمافزار MrBayes و بیشینه درستنمايی در نرمافزار RaxmlGUI اجرا شدند. نتايج تجزيهوتحلیل بیشینه صرفهجويی به تشکیل درخت مرکزی ITS ريبوزوم هستهای منجر شد که دارای شاخۀ اصلی A حامل گونههای Geum و سردههای مرتبطِ نزديک به آن )بهويژه Coluria R. Br. ،Acomastylis Greene ،Erythrocoma Greene و Novosieversia F. Bolle( در دو گروه تکنیاست. در تمام درختان حاصل، دو گونه از زيرسردۀ Geum گروهی تکنیا تشکیل دادند؛ در حالیکه گونههای زيرسردۀ Orthostylus بهشکل تریتومی (در تجزيهوتحلیل دادههای nrDNA ITS) و يا گروه تکنیا (در تجزيهوتحلیل دادههای کلروپﻻستی و ترکیبی) ظاهر شدند. نتايج بررسی حاضر نشان دادند فیلوژنی مولکولی اين سردهها بهشدت از دورگهگیری و پلیپلوئیدی )آلوپلیپلوئیدی( تأثیر میگیرد. يافتههای بررسی حاضر محدودۀ سیستماتیک گونههای اين سرده در فلورا ايرانیکا و فلور ايران را حمايت میکنند. در بررسی حاضر، تکامل صفتهای ر يختشناسی میوه، تزيینات اگزين دانۀ گرده، ريزريختشناسی بذر و صفتهای تشريحی دمبرگ ارزيابی شد. واژههای کلیدی: فیلوژنی،.cpDNA rpl32-trnL (UAG) ،nrDNA ITS ،Rosaceae ،Geum L. مقدمه. پیچیدگی آشکارا در مطالعههای تاکسونومیکی پیشین (Linnaeus, 1735; Bolle, 1933; Focke, 1894; Geum L. يکی از پیچیدهترين سردههای تیرۀ ;Rydberg, 1913; Juel, 1918; Hutchinson, 1967 (Robertson, 1974; Schulze-menz, 1964 گلسرخیان ازنظر سیستماتیکی محسوب میشود. اين بهويژه * [email protected] Copyright©2019, University of Isfahan. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/BY-NC-ND/4.0), which permits others to download this work and share it with others as long as they credit it, but they cannot change it in any way or use it commercially. www.SID.ir Archive of SID 2 تاکسونومی و بیوسیستماتیک، سال دهم، شماره سی و پنجم، تابستان 1397 در تعیین حد و مرز سرده و گونهها منعکس شده است. Coluria R. Br. ،Greene و Novosieversia F. Bolle( گونههای اين سرده در نواحی معتدله و قطبی دنیا و گیاهان علفی چندساله اند که روی درخت فیلوژنی عمدتاً در ارتفاعات و بیشتر در نیمکرۀ شمالی پراکنش تجزيهوتحلیل nrDNA ITS در درونگروه بزرگ دارند (Smedmark and Eriksson, 2002). پراکنش طايفۀ Colurieae و زيرطايفۀ Geinae قرار میگیرند اين گیاهان در ايران به مناطق شمال، شمالغرب و غرب (Smedmark and Eriksson, 2002). نتايج مطالعههای )با 4 گونه( و شمالشرق و مرکز )با 1 گونه( مربوط مولکولی پیشین محدودۀ سیستماتیکی Geum و میشود )جدول 1(. از میان اين گونهها، G. urbanum سردههای جداشده از آن )به غیراز Orthostylus .L در کمترين ارتفاع )200 متر( و Schlecht.) F. Bolle G. kokanicum) که در نیمکرۀ جنوبی انتشار .Regel and Schmalh ex Descer در بیشترين ارتفاع دارد( را حمايت نمیکنند Smedmark and) )3400 متر( از سطح دريا رشد میکند -Eriksson, 2002 (Schönbech(؛ اين پژوهشگران ابهامات Temsey, 1969; Khatamsaz, 1993(. تاکسونومیکی موجود را ناشی از پديدۀ آلوپلیپلوئیدی گونههایGeum ريزوم، برگهای طوقهای و بنساقۀ میدانند و نظريۀ تکامل مشبک را مطرح میکنند (Smedmark et al., 2005; Smedmark and ضخیم دارند، گلبرگها بیشتر زرد، سفید، نارنجی، Eriksson, 2006). تبارشناسی و تشخیص روابط بین دو قرمز تا ارغوانیاند، کاسبرگها و گلبرگ ها 5 عدد، زيرسرده و گونههای Geum در ايران با استفاده از پرچمها 20 عدد و میوه فندقه است. خامه مفصل دار، توالی های نوکلئوتیدی هسته ای و کلروپﻻستی ازجمله دارای رأس پايا و يا زودافت و قﻻبدار است. خامه در اهداف پژوهش حاضر است؛ همچنین بررسی شیوۀ گونههای .G. urbanum L و .G. rivale L مفصل تکامل صفت های ر يختشناسی میوه، تزيینات اگزين انتهايی و قﻻب افتان دارد که اصطﻻحاً آن را دانۀ گرده، ريزريخت شناسی بذر و صفتهای تشريحی Fish hook يا قﻻب ماهیگیری مینامند (Iltis, 1913) دمبرگ از ديگر اهداف مطالعۀ حاضرند. )جدول 2(. مطالعههای ياختهشناسی پیشین نشان میدهند آلوپلوئیدی، دورگه گیری بین گونهای و مواد و روشها. دوبرابرشدن کروموزومها نقش بسزايی در شکلگیری در پژوهش حاضر، روابط فیلوژنی مولکولی 5 گونۀ و تکامل گیاهان زيرطايفۀ Geinae دارند ,Gajewski) Geum شامل G. kokanicum ،G. heterocarpum، 1958). اعداد کروموزومی پايه در اين سرده بسیار G. rivale ،G. iranicum Khatamsaz و متغیرند )28، 42، 56، 70 و 84( و تعدادی از گونه های G. urbanum اين سرده هگزاپلوئید )مانند G. rivale و در ايران تجزيه وتحلیل شد. به اين منظور، ابتدا نمونههای گیاهی طی بهار و تابستان G. urbanum( و برخی تتراپلوئید G. heterocarpum) .Bioss( گزارش شدهاند Kalkman, 2004; Arens et) سالهای 93 و 94 از طبیعت جمعآوری و با مراجعه به فلورا ايرانیکا )Schönbech-Temsey, 1969( و فلور al., 2004(. گونههای Geum و خويشاوندان نزديک ايران )Khatamsaz, 1993( شناسايی شدند؛ نمونههای آن )شامل Acomastylis ،Erythrocoma Greene www.SID.ir Archive of SID فیلوژنی و تکامل صفتها در سردۀ .Geum L از تیرۀ گلسرخیان در ايران 3 جمع آوریشده در هرباريوم دانشکدۀ علوم پايۀ دانشگاه بهمنظور مقايسۀ موقعیت فیلوژنتیک گونههای ايرانی گیﻻن نگهداری میشوند. در اين تجزيهوتحلیل از Geum )جدول 1(، اطﻻعات توالی يابیشدۀ ناحیۀ نمونههای گیاهی موجود در هرباريومهای دانشکدۀ nrDNA ITS 21 تاکسون از بانک ژن (NCBI) نیز علوم پايۀ دانشگاه گیﻻن (GUH) و دانشکدۀ داروسازی استفاده شد )جدول 3(. دانشگاه علوم پزشکی تهران (THE) استفاده شد. جدول 1- ويژگیهای نمونههای مطالعهشده Accession. No / (nr DNA ITS/rpl32-trnL(UAG محل جمعآوری نام گونه نام زیرسرده خراسان: سرخچشمه، روئین، 2100 متر؛ Orthostylus LC336370/ LC336375 1.1. G. heterocarpum منصف (THE) 6738 خراسان: جادۀ بجنورد به اسفراين، گردنه اسدی، 1738 متر؛ 1.2. G. kokanicum LC336371/ LC336376 شاهی (GUH) 5751 خراسان: منطقۀ گرايل، امامزاده ذکريا، شیروان؛ 1.3. G. iranicum LC336372/ LC336377 منصف (THE) 6714 گیﻻن: 14 کیلومتری لنگرود به اطاقور؛ Geum .2 LC336373/ LC336378 2.1. G. rivale شاهی (GUH) 5753 گیﻻن: ﻻهیجان، سیاهکل، روستای لونک، 476 متر؛ 2.2.G. urbanum LC336374/ LC336379 فقیر و عاشوری )GUH(5749 مطالعۀ فیلوژنی مولکولی. (White, 1990) و به منظور انجام واکنش PCR برای مطالعۀ فیلوژنی مولکولی شامل مراحل استخراج توالی کلروپﻻستی (rpl32-trnL (UAG از آغازگرهای DNA، واکنش زنجیره ای پلیمراز rpl32 (Polymerase و (Shaw et al., 2007) trnL(UAG) استفاده Chain Reaction: PCR)، تعیین توالی و شد. توالی آغازگرهای استفاده شده در جدول )4(، هم رديف سازی توالی ها، تجزيه وتحلیل فیلوژنی داده ها مواد استفاده شده برای انجام واکنش PCR در جدول و ترسیم درخت فیلوژنتیک است. DNA کل ژنوم از )5( و برنامۀ PCR استفاده شده برای تکثیر قطعه های برگ های خشک شده و تازه با استفاده از کیت nrDNA ITS و (rpl32-trnL(UAG در جدول )6( استخراج DNA از نمونه های گیاهی Qiagene آورده شده است. استخراج شد. واکنش زنجیرهای پلیمراز (PCR). به منظور تکثیر توالی های nrDNA ITS از آغازگرهای Sang et al., 1995) ITS5m) و ITS4 www.SID.ir Archive of SID 4 تاکسونومی و بیوسیستماتیک، سال دهم، شماره سی و پنجم، تابستان 1397 جدول 2- رويشگاه و صفتهای کلیدی نمونههای مطالعهشده / صفتهای کلیدی متمایزکننده رویشگاه گونههای مطالعهشده نام گونه نام زیرسرده میوه دارای قﻻب، فقط قسمت باﻻی خامه پايا و Orthostylus آذربايجان و گلستان: 2000-1700 متر G.
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