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AYÇİÇEĞİ (Helianthus Annuus L.) GENOTİPLERİNDE KURAKLIĞA DAYANIKLILIĞIN FİZYOLOJİK, BİYOKİMYASAL VE MOLEKÜLER DÜZEYDE İNCELENMESİ

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AYÇİÇEĞİ (Helianthus Annuus L.) GENOTİPLERİNDE KURAKLIĞA DAYANIKLILIĞIN FİZYOLOJİK, BİYOKİMYASAL VE MOLEKÜLER DÜZEYDE İNCELENMESİ AYÇİÇEĞİ (Helianthus annuus L.) GENOTİPLERİNDE KURAKLIĞA DAYANIKLILIĞIN FİZYOLOJİK, BİYOKİMYASAL VE MOLEKÜLER DÜZEYDE İNCELENMESİ INVESTIGATION OF DROUGHT TOLERANCE AT PHYSIOLOGICAL, BIOCHEMICAL AND MOLECULAR LEVELS IN SUNFLOWER (Helianthus annuus L.) GENOTYPES AYŞE SUNA BALKAN NALÇAİYİ PROF. DR. YASEMİN EKMEKÇİ Tez Danışmanı Hacettepe Üniversitesi Lisansüstü Eğitim-Öğretim ve Sınav Yönetmeliğinin Biyoloji Anabilim Dalı için Öngördüğü DOKTORA TEZİ olarak hazırlanmıştır. 2018 ÖZET AYÇİÇEĞİ (Helianthus annuus L.) GENOTİPLERİNDE KURAKLIĞA DAYANIKLILIĞIN FİZYOLOJİK, BİYOKİMYASAL VE MOLEKÜLER DÜZEYDE İNCELENMESİ Ayşe Suna BALKAN NALÇAİYİ Doktora, Biyoloji Bölümü Tez Danışmanı: Prof. Dr. Yasemin EKMEKÇİ Ocak 2018, 217 sayfa Bu çalışmada, ülkemizde yetiştirilen tescilli ayçiçeği çeşitleri ile atasal melezler ve kuraklığa toleranslı olduğu bilinen atasal Helianthus agrophyllus’un kuraklık ve iyileşme süreçlerinde oluşturdukları yanıtların morfolojik, fizyolojik, biyokimyasal ve moleküler (proteom) düzeyde incelenmesi amaçlanmıştır. Çalışmanın I. aşamasında ayçiçeği çeşitleri iki farklı şiddette (orta şiddetli-7 gün / şiddetli-9 gün) kuraklık ve sonrasında iyileşme (sulama- 5 gün) uygulamalarına maruz bırakılmıştır. Bu aşamada, çeşitlerin fotosentetik aktiviteleri (klorofil a fluoresans ölçümleri), membran hasarları (iyon sızıntısı), fotosentetik pigment ve su içerikleri ölçülmüştür. Fotosentetik performans indekslerinden, kuraklık ve iyileşme faktör indeksleri ile fizyolojik ve biyokimyasal ölçümlerden hasar indeksi ve iyileşme potansiyelleri hesaplanmıştır. Bu hesaplamalar doğrultusunda çeşitler; kuraklığa toleranslı (Şems, Sirena, Tarsan-1018), orta derecede toleranslı (Duna, Hornet, Tunca) ve duyarlı (Bosfora, Coral, Kaan) olmak üzere 3 tolerans düzeyinde sınıflandırılmıştır. Çalışmanın II. aşamasında, seçilen çeşitler ile birlikte ayçiçeğinin atasal melezleri (Helianthus annuus x Helianthus agrophyllus ve Helianthus annuus x Helianthus i deserticola) 9 gün kuraklık stresine ve devamında 5 gün iyileşme sürecine maruz bırakılarak genotiplerin kuraklığa oluşturduğu cevaplar bazı morfolojik, fotosentetik, fizyolojik ve biyokimyasal parametreler ile ortaya konulmuştur. Elde edilen verilerinden hasar indeks değerleri ve iyileşme potansiyelleri hesaplanarak genotipler kuraklık toleranslarına göre karakterize edilmiştir. Kuraklık koşullarında su içeriğini azalmasına bağlı olarak, tüm genotiplerde fenotipik karakterlerde (bitki boyu, yaprak sayısı ve taze/kuru ağırlık) değişimler saptanmıştır. Bununla birlikte, toleranslı genotipler duyarlı genotiplere göre; fotosentetik aktivitelerini ve pigment içeriklerini (klorofil/karotenoid) koruyup (eksitasyon enerjisini daha iyi yakalayıp etkin bir şekilde kullanabildiği), savunma kapasitesini [flavonoid, antosiyanin, glisin betain, süperoksit dismutaz (SOD), askorbat peroksidaz (APX), glutatyon redüktaz (GR), peroksidaz (POD) ve aldoz redüktaz (ALR)] arttırarak, stresle oluşan membran hasarı (MDA) ile H2O2 düzeyini düşük tutabilmiş ve strese dayanım sağlayabilmişlerdir. Çalışmanın III. aşamasında toleranslı (Tarsan-1018) ve duyarlı (Tunca) olarak karakterize edilen çeşitler ile atasal Helianthus agrophyllus’un protein profillerindeki değişimler proteomik analizler ile ortaya konulmuştur. MALDI-TOF/TOF MS/MS kütle spektrometresi ve MASCOT veri tabanı kullanılarak genotiplerde 63 adet protein tanımlanmış ve STRING veri tabanı ile protein-protein etkileşim örüntüleri oluşturulmuştur. Tanımlanan proteinlerin; fotosentez ve karbohidrat, enerji ve solunum, savunma, arjinin, nükleotid, yağ asidi ve glikolipid, protein ve sinyal metabolizması ile hücre çeper biyogenezinde görev alan proteinler olduğu belirlenmiştir. Duyarlı çeşitte, metabolizmalarda tanımlanan proteinlerin farklı yönde değişmesi ve kuraklıkla birlikte nükleotid ve protein metabolizmaları ile sinyal iletiminde yer alan protein ifadelerindeki azalmalar, Tunca’yı diğer genotiplere göre daha az başarılı kılmıştır. Toleranslı genotipler ise fotosentez ve karbon metabolizmaları ile birlikte enerji ve solunum ile yağ asiti ve glikolipid metabolizmalarındaki protein ifadelerini benzer yönde düzenleyerek ve sinyal iletim yolağında yer alan 14-3-3 benzeri protein ifadesini artırarak kuraklık koşullarına dayanımda daha iyi performans sergiledikleri belirlenmiştir. Anahtar Kelimeler: Fotosentetik aktivite, iyileşme, kuraklık, proteomik, savunma sistemleri, tolerans, yerel ve atasal ayçiçeği genotipleri ii ABSTRACT INVESTIGATION OF DROUGHT TOLERANCE AT PHYSIOLOGICAL, BIOCHEMICAL AND MOLECULAR LEVELS IN SUNFLOWER (Helianthus annuus L.) GENOTYPES Ayşe Suna BALKAN NALÇAİYİ Doctor of Philosophy, Department of Biology Supervisor: Prof. Dr. Yasemin EKMEKÇİ January 2018, 217 pages In this study, it was aimed to investigate the responses of registered sunflower cultivars grown in our country with their wild hybrids and wild Helianthus agrophyllus, identified to be drought tolerant, at morphological, physiological, biochemical and molecular (proteomic) levels during drought and recovery. In the first stage, sunflower cultivars were exposed to two different drought treatments (moderate -7 days / severe-9 days) and then recovery (irrigation-5 days). In this stage, photosynthetic activities (chlorophyll a fluorescence measurements), membrane damage (ion leakage), photosynthetic pigment and water contents of the cultivars were measured. Drought-recovery factor indexes and damage-recovery potential indexes were calculated from photosynthetic performance indexes, and physiological and biochemical measurements, respectively. These calculations allowed classification of the genotypes to three tolerance levels: drought-tolerant (Şems, Sirena, Tarsan-1018), moderately tolerant (Duna, Hornet, Tunca) and sensitive (Bosfora, Coral, Kaan). In the second stage of the study, the drought responses of selected cultivars with the wild hybrids of the sunflower (Helianthus annuus x Helianthus agrophyllus and Helianthus annuus x Helianthus deserticola) iii subjected to drought stress for 9 days and then 5-days recovery were proved by using the morphological, photosynthetic, physiological and biochemical parameters. Genotypes were characterized according to their drought tolerances by calculating damage indexes and recovery potentials from the obtained data. The changes in phenotypic characters (plant height, number of leaves and fresh / dry weight) were determined in all genotypes due to the reduced water content under drought conditions. However, tolerant genotypes compared to sensitive genotypes, were able to conserve photosynthetic activities (absorption/trapping of excitation energy and utilize efficiently), pigment contents (chlorophyll/carotenoid) and enhance defense capacity [flavonoids, anthocyanins, glycine betaine, superoxide dismutase (SOD), ascorbate peroxidase (APX), glutathione reductase (GR), peroxidase (POD) and aldose reductase (ALR)], thereby the level of H2O2 and stress-induced membrane damage (MDA) were kept low and were able to provide tolerance to stress. In the third stage, the alterations in the protein profiles of tolerant (Tarsan-1018) and sensitive (Tunca) cultivars and wild Helianthus agrophyllus were proved by proteomic analysis. 63 proteins were identified in genotypes with MALDI-TOF / TOF MS / MS mass spectrometry and MASCOT database, and the protein-protein interaction patterns were performed with STRING database. It has been determined that the identified proteins are involved in photosynthesis and carbohydrate, energy and respiration, defense, arginine, nucleotide, fatty acid and glycolipid, protein and signal metabolism and cell wall biogenesis. Different expression of proteins in metabolisms and the reductions in nucleotide and protein metabolisms, as well as protein involved in signal transduction of sensitive cultivar, made Tunca less successful in stress resistance compared to other genotypes. Tolerant genotypes were found to exhibit better performance in terms of photosynthesis and carbon metabolism, as well as protein expression in energy and respiration and fatty acid and glycolipid metabolisms in the same way, and increased the expression of 14-3- 3 like protein in signal transduction pathway increased resistance to drought conditions. Keywords: Photosynthetic activity, recovery, drought, proteomics, defense systems, tolerance, cultivated and wild sunflower genotypes. iv TEŞEKKÜR Tez çalışmamın her aşamasında büyük bir özveri ve sabır gösteren her zaman desteğini yanımda hissettiğim, bilgi ve tecrübelerini paylaşan, çalışmalarıma her zaman destek olan ve yol gösteren tez danışmanım değerli hocam Sayın Prof. Dr. Yasemin EKMEKÇİ’ye, Bilgileri ve deneyimleri ile bana yol göstererek destek olan tez izleme komitesi üyesi değerli hocalarım Sayın Prof. Dr. Füsun İNCİ EYİDOĞAN’a ve Sayın Doç. Dr. Nuran ÇİÇEK’e; Laboratuvar imkânlarını kullanmam için katkılarından dolayı Sayın Prof. Dr. Rukiye TIPIRDAMAZ’a; Prof. Dr. Zafir EKMEKÇİ’ye; Uzm. Yasemin ÖZTÜRK’e; Prof. Dr. Hatice MERGEN’e, Arş. Gör. Dr. Emel SAĞLAR’a, Arş. Gör. Tuğçe KARADUMAN’a ve Arş. Gör. Merve ÖZCAN’a; çalışma kapsamında kullanılan ayçiçeği tohumlarının temini için Dr. Veli PEKCAN’a; kütle spektrometresi analizlerindeki yardımlarından dolayı Prof. Dr. Murat KASAP’a ve Doç. Dr. Gürler AKPINAR’a; protein ekstraksiyonundaki ön çalışmalara yardımlarından dolayı Prof. Dr. Mustafa YILDIZ’a ve Yrd. Doç. Dr. Hakan TERZİ’ye; glisin betain ölçümündeki yardımlarından dolayı Doç. Dr. Evren YILDIZTUGAY’a; Çalışmalarımın her aşamasında yardımlarını esirgemeyen Arş. Gör. Dr. Handan SEVİM’e,
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