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8113-Yasham Neden Var-Nick Lane-Ebru Qilic-2015-318S.Pdf KOÇ ÜNiVERSiTESi YAYINLARI: 87 BiYOLOJi Yaşam Neden Var? Nick Lane lngilizceden çeviren: Ebru Kılıç Yayına hazırlayan: Hülya Haripoğlu Düzelti: Elvan Özkaya iç rasarım: Kamuran Ok Kapak rasarımı: James Jones The Vital Question © Nick Lane, 2015 ©Koç Üniversiresi Yayınları, 2015 1. Baskı: lsranbul, Nisan 2016 Bu kitabın yazarı, eserin kendi orijinal yararımı olduğunu ve eserde dile getirilen rüm görüşlerin kendisine air olduğunu, bunlardan dolayı kendisinden başka kimsenin sorumlu rurulamayacağını; eserde üçüncü şahısların haklarını ihlal edebilecek kısımlar olmadığını kabul eder. Baskı: 12.marbaa Sertifika no: 33094 Naro Caddesi 14/1 Seyranrepe Kağırhane/lsranbul +90 212 284 0226 Koç Üniversiresi Yayınları lsriklal Caddesi No:181 Merkez Han Beyoğlu/lsranbul +90 212 393 6000 [email protected] • www.kocuniversirypress.com • www.kocuniversiresiyayinlari.com Koç Universiry Suna Kıraç Library Caraloging-in-Publicarion Dara Lane, Nick, 1967- Yaşam neden var?/ Nick Lane; lngilizceden çeviren Ebru Kılıç; yayına hazırlayan Hülya Haripoğlu. pages; cm. lncludes bibliographical references and index. ISBN 978-605-5250-94-2 ı. Life--Origin--Popular works. 2. Cells. 1. Kılıç, Ebru. il. Haripoğlu, Hülya. 111. Tirle. QH325.L3520 2016 Yaşam Neden Var? NICKLANE lngilizceden Çeviren: Ebru Kılıç ffi1KÜY İçindeki le� Resim Listesi 7 TEŞEKKÜR 11 GiRİŞ 17 Yaşam Neden Olduğu Gibidir? BiRİNCi BÖLÜM 31 Yaşam Nedir? Yaşamın ilk 2 Milyar Yılının Kısa Ta rihi 35 Genler ve Doğal Ortamla ilgili Sorun 39 Biyolojinin Kalbindeki Kara Delik 43 Karmaşıklık Yo lunda Kayıp Adımlar 52 Ya nlış Soru 57 iKİNCİ BÖLÜM 61 Yaşamak Nedir? Enerji, Entropi ve Yapı 63 Tu haf.Dar Biyolojik Enerji Yelpazesi 69 Biyolojinin Başlıca Bilmecelerinden Biri 79 Ya şam Tü müyle Elektronlarla ilgilidir 81 Yaşam Protonlarla İlgilidir 85 İKiNCİ KISIM 89 Yaşamın Kökeni ÜÇÜNCÜ BÖLÜM 91 Yaşamın Kökeninde Enerji Bir Hücre Nasıl Ya pılır 96 Akış Reaktörleri Olarak Hidrotermal Menfezler 101 Alkalin Olmanın Önemi 107 Proton Gücü 112 DÖRDÜNCÜ BÖLÜM 126 Hücrelerin Ortaya Çıkışı LUCA'ya Uzanan Taşlı Yol Zar Geçirgenliği Sorunu 132 Bakteriler ve Arkeler Neden Te melden Farklıdır? 140 ÜÇÜNCÜ KISIM 147 Karmaşıklık BEŞiNCİ BÖLÜM 149 Karmaşık Hücrelerin Kökeni 149 Karmaşıklığın Kimerik Kökeni 152 Bakteriler Neden Bakteridir? 157 Gen Başına Enerji 161 Ökaryotlar Nasıl Kaçmıştır? 167 Mirokondriler, Karmaşıklığın Anahtarları 173 ALTINCI BÖLÜM 179 Eşeyli Üreme ve Ölümün Kökenleri Genlerimizin Ya pısındaki Sır 185 lnrronlar ve Hücre Çekirdeğinin Kökeni 190 Eşeyli Üremenin Kökeni 195 iki Cinsiyet 202 Ölümsüz Germ Hattı, Ölümlü Beden 208 YEDiNCi BÖLÜM 215 Güç ve Şan Tü rlerin Kökeni Üzerine 221 Cinsiyet Kararlılığı ve Haldane Kuralı 227 Ölüm Eşiği 235 Yaşlanmayla ilgili Serbest Radikal Kuramı 241 SON SÖZ 251 Derinliklerden Sözlük 261 Notlar 267 Kaynakça 281 Dizin 309 Resim Listesi Resim 1 Karmaşık hücrelerin kimerik kökenlerini gösteren bir yaşam ağacı. İzniyle yeniden basılmıştır: Martin, W. , "Mosaic bacterial chromosomes: a cha llenge en route to a tree of genomes", BioEssays 21: 99-104, 1999. 27 Resim 2 Yaşamın zaman çizelgesi. 37 Resim 3 Ökaryotların karmaşıklığı. İzniyle yeniden basılmıştır: (A) Fawcett O. Th e Celi. WB Saunders, Philadelphia, 1981. (B) Mark Farmer, University of Georgia. (C) Newcastle University Biomedicine Scientific Facilities; (O) Peter Letcher, Alabama University. 45 Resim 4 Arkezoa - meşhur (ama yanlış) kayıp halka. İzniyle yeniden basılmıştır: (A) Katz L.A., "Changing perspectives on the origin of eukaryotes", Tr endsin Ecology and Evolution 13: 493-497, 1998. (B) Adam RD, "Biology of Giardia la mblia", Clinical Reviews in Microbiology 14: 447-75, 2001. 47 Resim 5 Ökaryor "süper grupları." İzniyle yeniden basılmıştır: Koonin, E.,V., "The origin and early evolution of eukaryores in rhe lighr of phylogenomics", Genome Biology 11:209, 2010. 50 Resim 6 Biyolojinin kalbindeki kara delik. Fotomikrograf, izniyle yeniden basılmıştır: Soh, E. Y., Shin H.J., Im K., "The prorecrive effecrs of monoclonal amibodies in mice from Na egleria fowleri infecrion", Korean ]ournalof Parasitology 30: 113-123, 1992. 53 Resim 7 Bir lipid zarının yapısı. İzniyle yeniden basılmıştır: Singer, S. ], Nicolson, G. L., "The fluid mosaic model of rhe strucrure of celi membranes", Science 175: 720-31, 1972. 65 Resim 8 Solunum zincirinde birinci kompleks. İzniyle yeniden basılmıştır: (A) Sazanov L. A, Hinchliffe, P. , "Srrucrure of rhe hydrophi lic domain of respiratory complex I from Th ermus thermophiles': Science 311: 1430-1436, 2006. (B) Baradaran, R., BerrisfordJ. M., Minhas, G.S., Sazanov, L. A., "Crysral strucrure of rhe entire respiratory complex I", Na ture 494: 443-48, 2013. (C) Vinorhkumar K. R., Zhu, ]., Hirst, ]., ''Architecrure of mammalian respiratory complex I", Na ture 515: 80-84, 2014. 73 Resim 9 Mitokondriler nasıl çalışır? Fotomikrograf, izniyle yeniden basılmıştır: Fawcett, O., The Celi, W. B. Saunders, Philadelphia, 1981. 76 8 YAŞAM NEDEN VAR' Resim 10 ATP semazının yapısı. İzniyle yeniden basılmıştır: David S. Goodsell, The Machinery of Life,Spri nger, New York, 2009. 78 Resim 11 Demir sülfür mineralleri ve demir sülfür toplulukları. İzniyle değiştirilmiştir: Russell, M. J., Martin, W. , "The rocky roots of the acetyl CoA pathway", Tr ends in Biochemical Sciences 29 358063, 2004. 104 Resim 12 Denizlerin derinindeki hidrotermal menfezler. Fotoğraflar Deborah S. Kelley ve Oceanography Society'nin izniyle yeniden basılmıştır; Oceanography 18, Eylül 2005 106 Resim 13 Te rmoforezle organik maddelerin aşırı yoğunlaşması. İzniyle yeniden basılmıştır: (a-c) Baaske, P., Weinert, F. M., Duhr, S., vd. "Extreme accumulation of nucleotides in simulated hydrothermal pore systems", Proceedings Na tiona!Academy Sciences USA 104: 9346-9351, 2007. (d) Herschy, B., Whicher, A., Camprubi, E., Wa tson, C., Dartnell, L., Wa rd, J., Evans, J. R. G., Lane, N., ''An origin-of-life reactor to simulate alkaline hydrothermal vents", jo urnal of Molecular Evolution 79: 213-27, 2014. 109 Resim 14 H2 ve C02'den nasıl organik madde yapılır? İzniyle yeniden basılmıştır: Herschy, B., Whicher, A., Camprubi, E., Watson, C., Dartnell, L., Wa rd, J., Evans, J. R. G., Lane, N., "An origin-of-lifereactor to simulate alkaline hydrothermal vents", jo urnal of Mo lecular Evolution 79: 213-27, 2014. 115 Resim 15 Ya şamın üç alemini gösteren meşhur ama yanıltıcı ağaç. İzniyle değiştirilmiştir: Woese, C. R., Kandler, O., Wheelis, M. L., "Towards a natura! system of organisms: proposal forthe domains Archaea, Bacteria, and Eucarya", Proceedings Na tiona!Academy Sciences USA 87: 4576-4579, 1990. 121 Resim 16 "Hayret verici kaybolan ağaç." İzniyle yeniden basılmıştır: Sousa, F. L., Thiergart, T. , Landan, G., Nelson-Sathi, S., Pereira, I. A. C., Ailen, J. F., Lane, N., Martin, W. F., "Early bioenergetic evolution", Ph ilosophical Transactions Royal Society B 368: 20130088, 2013. 123 Resim 17 Doğal bir proton basamağının enerji sağladığı bir hücre. İzniyle deği ştirilmiştir: Sojo, V. , Pomiankowski, A., Lane, N., "A bioenergetic hasis formembrane divergence in archaea and bacteria", PL OS Biology 12 (8): el001926, 2014. 133 Resim 18 Metan yaparak enerji üretmek. 135 Resim 19 Bakteriler ve arkenin kökeni. İzniyle değiştirilmiştir: Sojo, V., Pomiankowski, A., Lane, N., ''A bioenergetic hasis for membrane divergence in archaea and bacteria " PL OS Biology 12(8): el001926, 2014. 141 Resim 20 Etkin pompalamanın olası evrimi. 144 RESİM LİSTESi 1 9 Resim 21 Ökaryotların dikkat çekici kimerikliği. İzniyle yeniden basılmıştır: Thiergart, T. , Landan, G., Schrenk, M., Dagan, T. , Martin, W. F., "A n evolutionary network of genes present in the eukaryote common ancestor polis genomes on eukaryotic and mitochondrial origin", Genome Biology and Evolution 4: 466-485, 2012. 155 Resim 22 Ya şa mın üç değil, iki alemi. İzniyle yeniden basılmıştır: Williams, T. A., Poster, P. G., Cox, C. J., Embley, T. M., ''.An archaeal origin of eukaryotes supports only two primary domains of li fe", Nature 504: 231-236, 2013. 158 Resim 23 ''.Aşırıpoliploidlik" gösteren dev bakteriler. (A) ve (B) Cornell Üniversitesi'n den Esther Angert'in izniyle yeniden basılmıştır: (C) ve (O) Rostock, Leibnitz Baltık Denizi Araştırmaları Enstitüsü'nden Heide Schulz­ Vo gt'un izniyle Lane, N., Martin, W. 'nin yazdığı "The energetics of genome complexity"de yeniden basılmıştır, Nature 467: 929-934 2010; ve Schulz H. N.,"The genus Thiomargarita", Prokaryotes 6: 1156-1163, 2006. 166 Resim 24 Bakteriler ve ökaryotlarda gen başına enerji. Lane, N., Martin, W. "Energy per gene in bacteria and eukaryotes", Lane, N., Martin, W. , "The energetics of genome complexity" den orijinal veriler, Nature 467: 929-934, 2010; Lane, N., "Bioenergetic constraints on the evolution of complex life" da değiştirilerek kullanılmıştır, Cold Spring Harbor Perspectives in Biology, doi : 10.1101/cshperspect.a015982 CSHP, 2014. 168 Resim 25 Başka bakterilerin içinde yaşayan bakteriler. İzniyle yeniden basılmıştır: (Yukarıda) Wujek, O. E., "l mracellular bacteria in the blue-green-alga Pleurocapsa minor", Transactions ofthe American Microscopical Society 98: 143-145, 1979. (Alrra) Gatehouse, L. N., Sutherland, P., Forgie, S.A., Kaji, R., Christellera, J. T. , "Molecular and histological characterization of primary (beta-proteobacteria) and secondary (gammaproteobacteria) endosymbioms of three mealybug species", Applied Environmental Microbiology 78: 1187, 2012. 171 Resim 26 Çekirdeksel gözenekler. İzniyle yeniden basılmıştır: Fawcett O., The Cet!, W. B. Saunders, Philadelphia, 1981. 182 Resim 27 Kendi kendisini dilimleyen
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