Mutasyonlar genelde iki tiptir: nokta mutasyonu ve kromozom bozuklukları. Nokta mutasyonlarında sadece bir baz çifti değişime uğrar. İnsana göz atacak olursak; insan genomu 3.1 milyar DNA bazı taşır ve her baz, hücre bölünmesinin gerçekleşebilmesi için tam olarak eşlenmelidir. Bu durumda nadir de olsa hatalar meydana gelir. Yaklaşık olarak her 10 milyar baz çiftinden biri değişime uğrar. Hatalardan en sık rastlananı “nokta yer değiştirmesi”dir (point substitution). Nokta yer değiştirmesine göre daha az rastlanan 3 hata ise şunlardır: Bazlardan birinin kopyalanamaması (silme), tek bir baz için iki kopya üretilmesi, yeni bir bazın veya yeni birçok bazın eklenmesi (ekleme). Kromozom bozuklukları, nokta mutasyonlarına göre daha büyük çaplı mutasyonlardır. Kromozom bozuklukları; mayoz bölünme sürecinde kromozomlar-arası eşit olmayan parça değişimi* esnasında, DNA rekombinasyonunda meydana gelen kaymalarla ya da ters-çevrilebilir (transposable) durumlar esnasında ortaya çıkabilir. Genler ve hatta tüm kromozomlar yerdeğiştirebilir, eşlenebilir ya da bahsedilen hatalar nedeniyle silinebilir. (Görsel 1)
Görsel 1: Birbiriyle akraba olan 7 tropik balık türünün DNA dizilimindeki mutasyon (Veriler, epinepheine serranidlere ait intron 6 LDH-A geninin dizilenmesinden elde edilmiştir.) Orijinal DNA bilgisi üstte, dizilemeden sonra DNA bilgisi ise altta gösterilmiştir. Nokta yer değiştirmeleri kırmızıyla belirtilmiştir. Sarı kutucuk içerisindeki kesik çizgiler ise 12 bazın silinme işlemini göstermektedir.
Mutasyonlar, farklı etkilere sahiptir. Her mutasyon aynı özelliği göstermez. Mutasyonların çoğu zararlı mutasyonlardır. Zararlı olmayan mutasyonlar ise ya çok küçük etkilere sahip ya da hiçbir etkiye sahip olmayan mutasyonlardır. Ama çok nadir de olsa, DNA dizilimindeki değişiklik organizma için yararlı bir etkiye sahip olabilir.
Vücut hücresinde meydana gelip sonraki nesle aktarılmayan mutasyonlar somatik mutasyonlardır. Üreme (gamet) hücresinde veya üreme hücresinin oluşmasına sebep olan bir hücrede meydana gelen mutasyonlar ise farklı ve eşsizdir. Çünkü bu mutasyonlar, yetişkin (gamet üreten) bireyi neredeyse hiç etkilemezken; sonraki nesle aktarılıp sonraki nesli etkiler. Üremede kullanılan bir hücrede meydana gelen değişimlere germ-line** mutasyonu adı verilir ve burada meydana gelen bir mutasyon değişimin zamanla çoğalmasına olanak tanır. Eğer bu mutasyon, yavru bireyin fenotipi üzerinde zararlı bir etki yaratırsa, bu mutasyona “genetik bozukluk” denir. Öte yandan, eğer bu mutasyon yavru bireyin uyum başarısını artırıyorsa bu mutasyona “adaptasyon” denir. Bu yüzden, sonraki nesillerin uyum başarısını etkileyen her türlü mutasyon evrimin yapı taşıdır.
Mutasyon, evrimin bir şartıdır. Her bir organizmadaki her bir genetik özellik, başlangıçta, mutasyondan meydana gelmiştir. Yeni genetik varyant (allel), üremeyle yayılır ve farklılaşan üreme evrimi tanımlayıcı bir özelliktir. Organizmaların daha verimli beslenmesini, büyümesini ve çoğalmasını sağlayan mutasyonlar; mutant gen çifti (allel) sayısının zamanla artmasını sağlar. Bu, anlaması gayet kolay bir durumdur. Az bir zaman sonra, adaptif özelliği gelişmiş olan popülasyon adaptif özelliği daha az gelişmiş olan orijinal popülasyondan ekolojik ve/veya fiziksel olarak çok daha farklı bir hale gelebilir. Hatta, özellikle küçük popülasyonlarda, zararlı mutasyonlar bazı genlerinde adaptif alleller taşıyan bireyleri öldürerek evrimsel değişikliklere bile yol açabilir.
Mutasyonların çoğu genin üzerindeki tek bir noktada gerçekleşir. Genellikle tek bir proteini değiştirir ve bu yüzden önemsiz olarak görülebilir. Örneğin, genler tükürük bezlerindeki sindirim enzimlerinin yapısını ve verimliliğini kontrol eder. İlk bakışta, tükürük enzimleri üzerindeki mutasyonların hayatta kalmaya etkisinin az olduğu sanılabilir. Oysa ki yılan salyalarındaki zehrin küçük mutasyonların birbirini izlemesi ile oluştuğu ve bu yüzden bu mutasyonların yılanın evrimini çok önemli bir şekilde etkilediği kesinlikle söylenebilir. Yılanların ataları üzerindeki doğal seçilim, onların daha saldırgan ve daha zehirli enzim üretmelerini sağlamıştır. Ama, tabii ki bu mutasyonlar rastgele gerçekleşip, farklı yılan gruplarında farklı zehirler üretmiştir. Yılan zehri farklı etkilere sahip farklı proteinlerin karışımıdır. Bu yüzden birbirlerine genetik yakınlığı olan türler, yine zehirli olan diğer yılan ailelerine göre daha farklı bir zehir karışımına sahiptir. Su yılanlarının, mercan yılanlarının ve kobraların (Elapidae ailesinden olanların) ataları sinir sistemine saldıran zehir evrimleştirmiş iken Engerek yılanlarının (Viperidae ailesi; çıngıraklı yılan ve bushmaster yılanlarını da kapsar) zehri kardiyovasküler sisteme saldırır. İki aile de atalarından zehrin gücü ilgili küçük avantajlar almış birçok türe sahiptir. Ve mutasyonlar zaman içerisinde biriktikçe zehir ve tür çeşitliliği de artmıştır.
Pek çok türün evrimsel geçmişi küçük nokta mutasyonlarının birikimleri sonucu şekillenmiş olsa da bazı durumlarda evrim çok daha çabuk gerçekleşebilir. Organizma tiplerinden birkaçı, eşeyli üreme öncesinde hatalı mayoz geçiren atalara sahiptir. Bu hatalı mayoz sonucunda her kromozom çifti tam olarak eşlenir ve kromozom sayısı 2 katına çıkar. Bu durum Kuzey Amerika’daki gri ağaç kurbağalarında “anlık türleşme (instant speciation)” olarak gözlemlenir. (Görsel 2)
Görsel 2: Gri ağaç kurbağalarının (Hyla versicolor) evrimsel geçmişi mutasyon ve mutasyonun potansiyel etkilerini göstermek açısından yerinde bir örnek. Gri ağaç kurbağalarının atalarından biri olan Hyla chrysocelis’in mayoz esnasında 24 kromozomunu yanlış sıralaması sonucunda Hyla Versicolor meydana geldi. Hylo versicolor boyut, şekil ve renk açısından Hyla chrysocelis’in tıpatıp aynısı. Ama bu yeni tür 48 kromozoma sahip ve bu yeni türün çiftleşme çağrılarından biri orijinal Hyla chrysocelis’e göre daha farklı.
Genom boyutunun iki katına çıkması, eğer tohumlu bir bitkiyseniz, meyvelerinizin veya tohumlarınızın anormal bir şekilde büyümesini sağladığı için elbette bir avantaj olacaktır! Yediğimiz tahılların çoğu diğer otlarla karşılaştırdığımızda çok daha büyük tohumlara sahiptir. Bu durum, modern pirincin ve modern buğdayın atalarında gerçekleşmiş olan genomik eşlenmeden kaynaklanmaktadır. Yani, modern pirincin ve modern buğdayın atalarının üreme organlarında meydana gelen bir hata gelecek nesillere başarıyla aktarılmış ve bu özellik günümüze kadar ulaşmıştır. İnsanlar, bu işlemi yapay seçilim ile taklit edip, en büyük tohumlara ve en büyük meyvelere sahip farklı bitkileri melezlemişlerdir ve bu yolla modern birçok zirai mahsul türü üretilmiştir. İlk olarak Charles Darwin ve Alfred Russell Wallace tarafından tanımlanan ‘doğal seçilimle meydana gelen evrim’ fikri, evrimsel uyum başarısı daha gelişmiş olan bireylerin hayatta kalması ilkesine dayanıyordu. Uyum başarısı; genetik bozukluklar, zehirli tükürük, daha büyük boyuta sahip yavru birey ya da herhangi başka bir etmenden dolayı gerçekleşmiş olsa da, her durumda, tüm kalıtsal varyasyonlar sadece mutasyon ile var olabilir. Evrimin hammaddesi rastgele genetik değişimlerdir ve evrim bu değişimler olmaksızın düşünülemez.
* Kromozomların eşit oranda crossing over'a (mayoz bölünme sırasında eşlenik kromozomların çaprazlanması) maruz kalmaması durumu
** Eşey hücre öncülleri
Yazar: Joel L. Carlin (Department of Biology, Gustavus Adolphus)
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