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Sultan Sazlığı Ekosisteminde Kanatlı Haemosporidialarının Moleküler Ekolojisi

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Sultan Sazlığı Ekosisteminde Kanatlı Haemosporidialarının Moleküler Ekolojisi Sultan Sazlığı Ekosisteminde Kanatlı Haemosporidialarının Moleküler Ekolojisi Program Kodu: 1001 Proje No: 114 O 646 Proje Yürütücüsü: Prof. Dr. Alparslan YILDIRIM Araştırmacılar: Prof. Dr. Bilal DİK Doç. Dr. Önder DÜZLÜ Dr. Arzu GÜRSOY ERGEN Bursiyerler: Biyolog Mübeccel OKUR Biyolog Zekiye KOÇER Haziran 2017 KAYSERİ ÖNSÖZ Apicomplexa anaç altında kanatlılarda enfeksiyona yol açan Haemosporidia takımındaki Hemoproteus, Leucocytozoon ve Plasmodium soylarındaki türler, özelleşmiş organellerden oluşan ve konak hücreye penetrasyonda önemli olan bir apikal kompleks yapıya sahip olup vektörlerle bulaştırılmaktadır. Vektör-kaynaklı hastalıkların bulaşma dinamiklerinde vektörlerin biyolojik ve ekolojik özellikleri önemli rol oynar. Vektör popülasyonları üzerine ekolojik çalışmalar, farklı kanatlı topluluklarında haemosporidian parazitlerin yayılışı ve bulaşma dinamikleri açısından kilit role sahiptir. Günümüze kadar yapılan çalışmalar kanatlı haemosporidian etkenlerinin konak spesifikliği düzeyinde geniş çeşitlilik gösterdiğini ortaya çıkarmıştır. Bazı haemosporidian nesilleri belirli kuş ailelerinde görülürken diğer bazıları bütün Aves sınıfında yayılış gösterebilmektedir. Geçmişte birçok Hemoproteus ve Leucocytozoon türünün rölatif olarak konak spesifikliği gösterdiği ve aynı ailedeki kuş türleri ile sınırlı olduğu, Plasmodium türlerinin ise daha geniş konak spekturumuna sahip olduğu ve karakterlerini değiştirerek çeşitli kanatlı familyalarını enfekte edebildiği kabul edilmiştir. Ancak günümüzde moleküler biyolojik tekniklerin kanatlı haemosporidanlarının teşhisinde ve karakterlerinin belirlenmesinde kullanılmaya başlanması ile birlikte bu etkenlerin konak özgün ya da generalist özellikte olmaları revizyon altına alınmıştır. Bu amaçla kurulan MalAvi (http://mbio- serv4.mbioekol.lu.se/avianmalaria/) veri tabanında Dünyanın farklı bölgelerinden kanatlı ve/veya vektör konaklardan izole edilmiş haemosporidian parazitler başta mitochondrial cytochrome b (mt-cytb) gen bölgesi karakterleri bazında kayıt altına alınmaya başlanmıştır. Bu şekilde farklı coğrafyalarda tespit edilmiş kanatlı haemosporidian parazitlerinin bir arada moleküler filogenetik klasifikasyonuna olanak sağlanmış ve şu ana kadar elde edilen veriler ile global tür çeşitliliğinin geçmişte bilinenden çok daha yüksek olduğu ve bu takım altında binlerce tür bulunabileceği gün yüzüne çıkmıştır. Bu açıdan başta mt-cytb sekans karakterizasyonu olmak üzere moleküler veriler temelinde, genetik farklılıklarla ortaya çıkarılan bu parazitler genel bir terimle kanatlı haemosporidian nesilleri (lineage) olarak tarif edilmiş ve her bir nesil izole edildiği kanatlı ve/veya insekt konak temelinde isimlendirilerek veri tabanlarına kayıtları sağlanmıştır. Günümüze kadar bu nesillerden çok az bir kısmının morfolojik ve gelişim özellikleri temelinde tür tanımlaması yapılabilmiştir. TÜBİTAK tarafından 114 O 646 kod numarası ile desteklenen bu araştırma projesiyle yabani kanatlılardan ötücü kuşlar (Passeriformes) ve diğer bazı non-passerin kanatlılarda enfeksiyona yol açan haemosporidian etkenlerinin (Hemoproteus, Leucocytozoon ve Plasmodium) Kayseri il sınırları içindeki Sultan Sazlığı ekosisteminde kanatlı konak ve vektörler (Culicidae ve Ceratopogonidae) bazında moleküler ekolojisi araştırılmıştır. ii İÇİNDEKİLER KAPAK ........................................................................................................................................ i ÖNSÖZ ....................................................................................................................................... ii İÇİNDEKİLER ........................................................................................................................... iii TABLO LİSTESİ ........................................................................................................................ v ŞEKİL LİSTESİ ........................................................................................................................ vii ÖZET ......................................................................................................................................... xi ABSTRACT .............................................................................................................................. xii 1. GİRİŞ ..................................................................................................................................... 1 2. LİTERATÜR BİLGİ................................................................................................................ 3 2.1 Kanatlı Haemosporidian Parazitlerin Taksonomi, Morfoloji ve Biyolojisi ...................... 3 2.2 Kanatlı Haemosporidian Parazitlerin Epidemiyolojisi .................................................... 7 2.3 Kanatlı Haemosporidian Parazitlerin Teşhisi............................................................... 11 2.4 Moleküler Veriler Işığında Kanatlı Haemosporidianlarında Global Çeşitlilik ............... 14 3. GEREÇ VE YÖNTEM ......................................................................................................... 17 3.1 Saha Çalışmaları .......................................................................................................... 17 3.1.1 Sultan Sazlığı Ekosisteminde Kuş Örneklemesi ve Kan Örneklerinin Toplanması .................................................................................................................. 17 3.1.2 Sultan Sazlığı Ekosisteminde İnsekt (Diptera) Örneklerinin Toplanması ...... 20 3.2 Laboratuvar Çalışmaları............................................................................................... 20 3.2.1 Perifer Kan Frotilerinin Boyanması ve Analizi ................................................ 20 3.2.2 İnsekt (Diptera) Örneklerinin İdentifikasyonu ................................................. 21 3.2.3 Kanatlı Kan Örnekleri ve İnsekt (Diptera) Örneklerinden Genomik DNA izolasyonu.................................................................................................................... 21 3.2.4 Kanatlı Kanları ve İnsektlerden Elde Edilen Genomik DNA İzolatlarında Haemosporidan Parazitlerin Mithocondrial Cytochrome b (Mt-cytb) Gen Bölgesinin Amplifikasyonu ............................................................................................................ 22 iii 3.2.5 Moleküler İdentifikasyon ve Genotipleme için Sivrisinek ve Culicoides Mitochondrial Cytochrome Oxidase Subunit I (mt-COI) Gen Bölgesinin Amplifikasyonu ........................................................................................................................ 22 3.2.6 Dişi Sivrisinek ve Culicoides Örneklerinde Kan Beslenme Analizlerinde Kanatlı Mt-cytb Gen Bölgesinin Amplifikasyonu ..................................................................... 23 3.2.7 Sekans ve Filogenetik Analizler ..................................................................... 24 3.2.8 Real Time PCR Analizleri ............................................................................... 27 3.3 İstatistiksel Analiz ......................................................................................................... 29 4. BULGULAR......................................................................................................................... 30 4.1 Sultan Sazlığı Yöresinde Örneklenen Kuş Türleri ve Özellikleri ................................. 30 4.2 Sultan Sazlığı Yöresinde İnsekt (Diptera) Türleri ........................................................ 35 4.2.1 Culicoides Örneklerinin Morfolojik ve Moleküler Analiz Sonuçları ................ 36 4.2.2 Sivrisinek Örneklerinin Morfolojik ve Moleküler Analiz Sonuçları .................. 43 4.3 Sivrisinek ve Culicoides Türlerine Ait Örneklerin Kan Beslenme Analiz Sonuçları ... 48 4.4 Kanatlı ve İnsekt Örneklerinde Haemosporidian Parazitlerin Analiz Sonuçları .......... 52 4.4.1 Kanatlı Türlerinde Haemosporidian Parazitlerin İdentifikasyonu, Dağılımı ve Moleküler Karakterizasyonu ........................................................................................ 52 4.4.2 Culicoides ve Sivrisinek Türlerinde Haemosporidian Parazitlerin İdentifikasyonu, Dağılımı ve Moleküler Karakterizasyonu .......................................... 98 4.4.3 Kanatlı ve insekt örneklerinde moleküler olarak karakterize edilen Haemosporidian parazitlerin filogenetik analizleri .................................................... 104 4.5 Haemosporidian Parazitlerin Real Time PCR Analiz Sonuçları ................................ 113 5. TARTIŞMA VE SONUÇ .................................................................................................... 119 6. KAYNAKLAR .................................................................................................................... 146 iv TABLO LİSTESİ Tablo 2.1 Günümüze kadar tür identifikasyonu sağlanan kanatlı haemosporidian türlerinin taksonomideki yeri...................................................................................................................... 3 Tablo 3.1 Haemosporidian mt-cytb gen bölgesinden dizayn edilen Sybergreen ve TaqMan real time PCR için primer ve prob sekansları ................................................................................. 28 Tablo 4.1 2015-2016 yılları arazi çalışmalarında halkalanan türler, sistematik yerleri, IUCN kategorileri ve statüleri ............................................................................................................. 30 Tablo 4.2 2015 yılı arazi çalışmasında yakalanan kuş türleri ve sayıları
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