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Regulation of SPG4 and KANTB1 gene expressions by ELK1 transcription factor Skip to content Tez Künye Durumu Regulation of SPG4 and KANTB1 gene expressions by ELK1 transcription factor / SPG4 ve KATNB1 gen anlatımlarının ELK1 transkripsiyon faktörü ile düzenlenmesi Yazar:ECE SELÇUK Danışman: PROF. DR. ARZU KARABAY KORKMAZ Yer Bilgisi: İstanbul Teknik Üniversitesi / Fen Bilimleri Enstitüsü / İleri Teknolojiler Ana Bilim Dalı / Moleküler Biyoloji-Genetik ve Biyoteknoloji Bilim Dalı Konu:Biyoloji = Biology ; Genetik = Genetics Dizin:Blotting-western = Blotting-western ; Mikrotübüller = Microtubules ; Çeviriyazı = Transcription Onaylandı Yüksek Lisans İngilizce 2012 114 s. Hücre iskeleti tüm sitoplazmaya yayılan hücrenin şekil almasında, hareketinde, hücre içi molekül taşınması gibi ana mekanizmalarda görevli bir ağdır. Ökaryotik hücrelerde hücre iskeleti üç farklı yapıdan oluşmaktadır. Bunlar mikrofilamentler (aktin filamentler), ara filamentler ve mikrotübüllerdir.Mikrofilamentler, hücrede en fazla bulunan proteinlerden biridir ve evrimsel süreçte oldukça korunmuştur. Hücrenin şekil kazanmasında, hücre bölünmesinin sitokinez evresinde önemli roller üstlenen mikrofilamentler, globular aktinlerin ATP hidrolizi yardımıyla polimerizasyonu ile filament aktine dönüşür. Ara filamentler ise genellikle zar yapılarına yakın kısımlara yerleşmişlerdir. Örneğin çekirdek zarına çok yakın kısımlarda lamin adı verilen ara filamentler bulunur. Bunlar zara mekanik destek sağlarlar. Bunların haricinde hücreler arası bağların oluşumunda da görevleri vardır.Hücre iskeleti üçüncü elemanı olan mikrotübüller hücre bölünmesi sırasında kromozomların kutuplara çekilmesinde, hücrenin hareket edebilmesinde, hücrede vezikül taşınımında, nöronlarda akson oluşumunda ve nöronal şeklin kazanımında önemli rol oynar. Mikrotübüller hücrelerde stabil ve stabil olmayan iki farklı yapıda görülmektedir. Mikrotübüllerin ? ve ß tübülin adı verilen monomerlerinin polimerizasyonu ve depolimerizasyonu mikrotübüllere ?dinamik instabilite? diye adlandırılan özelliği kazandırır. Bu dinamizm, mikrotübüllerin uzayıp kısalmasını sağlar. Stabil mikrotübüller ise nöron gibi özelleşmiş hücrelerde bulunur. Çünkü nöronların, örneğin hücre gövdesinde üretilen proteinlerin akson uçlarına kadar taşınabilmesi için ray görevi görecek sabit bir yapıya ihtiyacı vardır. Bu stabilite mikrotübül ilişkili proteinler (MAP) tarafından kontrol edilir.Miktotübüller ile ilgili bilinmesi gereken diğer bir nokta da kesim mekanizmasıdır. Nöronların özel morfolojilerinin oluşmasında başı çeken mikrotübül kesimidir. Çünkü polimerizasyon ve depolimerizasyonun neden olduğu sadece uç kısımlarda meydana gelen uzalıp kısalma, nöron morfolojisinin oluşması için gerekli olan dallanma sürecinde zaman ve hız açısından yetersiz kalır. Uzun bir mikrotübülün daha kısa parçalara ayrılması daha avanajlıdır. Ayrıca mikrotübüllerin bir başka yere taşınabilmesi için de kesilmesi gerekir. Yani mikrotübül taşınabilirliği, mikrotübül boyu ile ters orantılıdır. Şu ana kadar spastin, katanin ve fidgetin adı verilen üç kesim enzimi tanımlanmıştır. Her üç enzim de ATP’yi parçalayarak işlev gören ATPaz ailesinin üyeleridir.Spastin, SPG4 geni tarafından kodlanırken, bir heterodimer olan katanin KATNA1 ve KATNB1 genlerinden kodlanan ve sırasıyla p60 ve de p80 diye isimlendirilen alt birimlerden oluşur. Kataninde enzimatik aktiviteyi gösteren asıl alt birim p60-katanindir. p80-katanin bu enzimatik aktiviteyi düzenleyendir ve bazı tanımlanmış bölgeleri ile p60-kataninin mikrotübülleri kesmesini arttırabilmektedir.Nöronlarda bu iki enzimin dağılımına bakıldığında, kataninin daha çok akson boyunca yoğunlaştığı, spastinin ise hücrenin kısa uzantılarının olduğu dallanma bölgelerinde fazla bulunduğu gözlemlenmiştir. Böylece spastinin mikrotübülleri daha kısa parçalara ayırdığı, kataninin ise daha uzun parçalar halinde kestiği gösterilmiştir.Bu iki enzimin çeşitli nörolojik hastalıklarla ilişkili olduğu bilinmektedir. Spastin genindeki delesyon, insersiyon gibi mutasyonların ya da alternatif kırpılmaların herediter spastik paraplejiye sebep olduğu gösterilmiştir. Spastin proteininin bu hastalıkla ilgisi tam olarak aydınlatılamamış olmakla beraber, mutant spastin proteininin mikrotübül kesme fonksiyonunu yitirmesinden dolayı akson uçlarında ihtiyaç duyulan kısa mikrotübüllerin sağlanamaması veya mutant formdaki spastinin akson uçlarındaki mikrotübüller üzerinde birikmesiyle veziküler transportun aksaması olası sebepler olarak bildirilmiştir. Mutasyonların haricinde gen anlatımının düzenlenmesinin de hastalığın ortaya çıkmasında etkili olacağı düşünülmüş ve spastin promotör karakterizasyonu yapılmıştır.Katanin ise Alzheimer hastalığı ile ilişkilidir. Çünkü mikrotübüllerin stabil kalmasını sağlayan ve normalde mikrotübüllerle etkileşim halinde olan tau proteininin hiperfosforilasyonu, mikrotübüllerin stabilitesini kaybetmesine neden olur. Böylece mikrotübüller katanin ile kesilmeye açık hale gelir. Bu sürecin Alzheimer hastalığı patolojisine neden olabileceği düşünülmektedir. Tüm bunlar göz önüne alındığında spastin ve kataninin hücre içindeki miktarları, dolayısıyla gen anlatım süreçleri ve düzenlenmesi oldukça önem kazanmaktadır.Bu genlerin anlatımlarının düzenlenmesinde etkili olan Elk1 transkripsiyon faktörünün rol oynadığı bulunmuştur. Elk1 hem gen anlatımını baskılayıcı hem de aktive edici bölgeler içermektedir. Elk1’in hedef gen anlatımını azaltması için aynı zamanda baskılanma bölgesinde SUMO adı verilen bir modifikasyon geçirmesi gerekmektedir. Ancak hücre dışı sinyallere bağlı olarak örneğin mitotik sinyallerle Elk1’in aktivatör bölgesi fosforilasyona uğradığında SUMO modifikasyonu ortadan kalkar ve Elk1 hedef gen anlatımını arttırır.Bu çalışmada spastin kodlayan SPG4 geninin ve p80-katanini kodlayan KATNB1 geninin anlatımlarının insan nöroblastoma hücrelerinde Elk1 transkripsiyon faktörü ile nasıl düzenlendiği araştırılmıştırElk1’in spastin promotörü üzerinde baskılayıcı etkisinin olduğu tespit edilmiş ve bu etkinin baskılayıcı bölgede gerçekleşen SUMO modifikasyonundan kaynaklandığı düşünülmüştür. Bu sebeple baskılayıcı bölge içermeyen yeni bir dizi oluşturulmuştur. Elk1 ve baskılayıcı bölge içermeyen Elk1’in spastin geni anlatımına olan etkileri karşılaştırıldığında, baskılayıcı bölge içermeyenin gen anlatımını arttırdığı bildirici gen (lusiferaz) deneyi ile saptanmıştır. Bu sonuç gen anlatım sürecinin kapsadığı transkripsiyon ve translasyon basamaklarında da sırasıyla mRNA ve protein düzeyinde de gerçek zamanlı polimeraz zincir reaksiyonu (PZR) ve immun boyama teknikleri ile doğrulanmıştır. Ardından SUMO modifikasyonunu elimine etmek ve promotör üzerindeki azaltıcı etkiyi tersine çevirebilmek için Elk1’in fosforile olması sağlanmıştır. Bunun için nöroblastoma hücrelerine mitotik bir ajan olan PMA bileşeni uygulanmış ve bildirici gen testi yeniden yapılmıştır. Buna göre PMA ile muamele edilmiş hücrelerde SPG4 gen anlatımının, PMA ile muamele edilmemiş hücrelere göre arttığı gözlemlenmiştir. Bu deney sonucu da protein düzeyinde doğrulanmıştır.Elk1’in KATNB1 promotörü üzerinde SPG4’den farklı olarak aktivatör rol oynadığı bildirici gen testi ile belirlenmiştir. Deney sonucu mRNA düzeyinde de gerçek zamanlı PZR yöntemiyle doğrulanmıştır. Elk1’in KCl kimyasalı ile SUMO modifikasyonuna uğraması tetiklenmiştir. KCl’ye maruz bırakılan hücrelerde diğerlerine göre daha az promotör aktivitesi belirlenmiştir. Elk1’in p80-katanin proteini seviyesini arttırdığı ve KCl uygulanmış hücrelerde p80-katanin protein üretiminin azaldığı immun boyama ve Western blotlama tekniği ile gösterilmiştir. Cytoskeleton is a network within the cell responsible for some cellular activities such as determining cell shape, providing cell movement and transport of some molecules. Microtubules are member of this network and have roles in chromosome segregation during mitosis, vesicular transport, axon formation and giving specific shapes of neurons. Microtubule severing is one of the major mechanisms that take role in formation of special neuron morphology. There are three microtubule severing enzymes called spastin, katanin and fidgetin. Spastin is encoded by SPG4 gene. However katanin is a heterodimer and its subunits p60 and p80 are encoded by two different genes KATNA1 and KATNB1, respectively. p60-katanin has the enzymatic activity for severing microtubules, while p80-katanin regulates role of p60-katanin in terms of stimulation or inhibition of p60-katanin activity.It is known that these enzymes are related with some neurological diseases. For instance, mutations in SPG4 gene lead to hereditary spastic paraplegia. Although exact mechanism has not been yet clear, it is thought that mutations in the SPG4 gene cause lack of short microtubules needed in axon ends, owing to loss of microtubule severing activity of spastin; or failure of vesicular transport due to accumulation of mutant spastin on microtubule in axon ends. Besides of mutations, regulation of SPG4 gene expression has also an effect on onset of disease. Katanin was found to be related with Alzheimer disease. In its unphosphorylated form, tau protein normally provides stabilization of microtubules and protects microtubules from the activity of severing enzymes. Hyperphosphorylation of tau results in dissociation of tau from microtubules and so microtubules become accessible for severing enzyme katanin. It is thought that this process may result in Alzheimer pathology.Elk1 is a transcription factor that has repressor and activator domains. The SUMO modification on repressor domain is necessary for inhibition of the target gene expression. However, when Elk1 is phosphorylated by extracellular signal such as a mitotic signal, Elk1 is de-SUMOylated and increases expression of target gene.In this study, the regulation of SPG4 and KATNB1 gene expressions by Elk1was investigated in human neuroblastoma cells.The repressive effect of Elk1 on spastin promoter was determined by using luciferase reporter assay and the reason for the repression is thought to be SUMO modification. Therefore new construct that did not include the repressor domain was constructed. According to the reporter assay data, Elk1 without repressor domain increased the promoter activity, whereas wt-Elk1 did not. This result was confirmed by analyzing mRNA and protein levels with real time PCR and immunostaning, respectively. Subsequently, PMA treatment was performed in order to de-SUMOylate Elk1 via phosphorylation. Following reporter gene assay, PMA treated and Elk1 transfected cells showed higher promoter activity compared to untreated and Elk1 transfected cell.Our observation indicated that, Elk1 acts as an activator on KATNB1 gene promoter differently than SPG4 promoter The results for KATNB1 gene were firstly confirmed in mRNA level by performing real time PCR. Then SUMOylation of Elk1 was triggered by using KCl. According to the reporter assay data, KCl treated and Elk1 transfected cells showed lower promoter activity compared to untreated and Elk1 transfected cells. Increase in p80-katanin protein level by Elk1 and decrease of p80-katanin protein level by KCl treatment were also shown with immunostaining and western blot analysis, respectively.
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