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    BIOLOGI CAMPBELL REECE - MITCHELL eee iT ee SS [ed BIOLOGI EDIS! KELIMA JILID Il Neil A. Campbell University of California, Riverside Jane B. Reece Palo Alto, California Lawrence G. Mitchell University of Montana > lily XC4W-ECT- PENERBIT ERLANGGA EGET aL LJ. H. Baping Raya No, 100 Cirecas, Jakarta 13740 e-mail: mahameru@rad netid (Anggota IKAP!) ‘Campbell, Neil A. Biologivell A. Campbel, Lawrence G. Mitchell; alin Manaiu ; editor, Amalia Safi, Lemeda Simarmata, Hilerius W. Hardani, —Jakarta: Erlangga, 2003. 472 him. ; 240 280 em, Judul aati Biology. ISBN 979-688-467-4 (no. jl. lengkap) ISBN 979-666-408-2 (i. 1) ISBN 979-€88-469-0 (il. 2) ISBN 979-668-470-4 (ji. 3) 1. Biolog. |. Judul, I. Reece, Jane B IL Michel, Lawrence . IV. Sef, Amaia. V, Simarmata, Lemeda. Vi Haran, Hiarius W. ara BUKU INI DITERBITKAN ATAS KERJA SAMA PENERBIT FRLANGGA DENGAN PUSAT PERBUKUAN DEPDIKNAS Gambar Sampul Depan: Magnolia Blosom, Tower of Jeveb, 1925 Foro oleh Imogen Cunningham, © 1974 ‘The Imogen Cunningham Trast. Judul Astiz BIOLOGY, Fifth Edition Campbell, Reece, Mitchell Hak cipta © 1987, 1990, 1993, 1996, 1999 pada Beyjaminn Gemmings, sebuah imprint dati Addison Wesley Longman, In¢. Hak terjemahan he dalam Bahasa Indonesia pada Fenerbir Erlangea, betdasarkan pesjanjian pada 19 April 2000. Alb Bahass + Prof, Ds. Ir, Wasmen Manalu Institut Pertanian Bogor leah Tim Juruan Biolog, PMIPA — ITB Dr. Ahmad Ridwan De. Ayd Trsnawary Yosuf De. Erly Marwani De. Iiawati Dr. Madita Ramdani Moc De. Taufkurahman De Riskita Rachmi E. De Sony Hem Sumarsono Bditor + Amalia Safe, TR M.Si Lemeda Simarmata, S. Hilarius W. Hardani, §.T. Buku ini disee dan dilayour olch Bagian Produksi Penerbie Erlangga dengan PoweMac G4 (Agaramond 10). 7 06 876543 Dicesk olch + PT Gelora Aksara Pratams All righs reserved. No part of this publication may be reproduced. stored in rericval stem, or ransmitied in any form er by any means electron nechanical, photocopying, recording, or ary other media embediments row known or hereafer to become known, without the prion written permision of the publiber: Dilarang keras mengutip, menjiplak, memperbanyak, memotokopi, baik sebagian maupun keseluruban isi buku ini serta memperjualbelikannya ‘anpa izin terule dasi Poserhir Flanges © HAK CIPTA DILINDUNGI OLEH UNDANG-UNDANG Ekosistem bergantung pada fungi ebagsi pengutal dan simbion 197 Sejumtah fangi adalah patogen 198 Banyak hewan, termasukt manusia, memakan fungi 198 Hubungan Filogenetk Fungi Fung’ dan hewan kemungkinon berevolasi dari neck moyang protista yang sama 198 32 Pengantar Evolusi Hewan 202 ‘Apa yang Dimaksud dengan Hewan? 202 Gambaran Umum mengenai Filogeni dan Keanckaragaman Hewan 203 Parazoa tidak memilik jaringansejat 205 Radista dan bibieris adalah cabang.utama eumetazon 205 Evolusi rongga tubuh menghailkan hewan yang lebih kompleks 206 Selomata bercabang menjadi protostoms dan deurerostoma 207 Asal Mula Keanckaragaman Hewan 209 Sebagian besar flu hevan muncul dalam suatu rentang waktu geologis yang rlatf sngkat 209 Genetika perkembangan memperjélas pemahaman kita mengenai diversifikasi di masa Kambrium 210 33 tnverebrata 213 Parzzea 213 Frum Porifera: Spons adalah stil dan memibkieubuh berpori seta koanosit 213 Radiasa 25 Filum Cnidaria: Anggora flu Cnidaria memilki simert radial, rongga gastrovaskule, dan enidosit 215 Fillum Ctenophora: Ubue-ubue ssi memilik barban lempeng sia din koloblas yang lengket 218 Asclomata 21K Filum Platyhelminthes: Cacing pipih adalah hewan sselomata yang pipihsecara dersoventral 218 Preudosclomara 220 Filum Rovfera: Anggota filum Rotifera memliki shang dan rmahkora sia 21 Filum Nematoda: Cacing gig tidak beregmen dan bereubuh silndsis dengan ujung yang meruncing 21 Sclomata: Protostoma 22 Filum Nemertex: Pos ccing proboscis di pohon filogenetik masih behum jelas 222 Filurnfilum Lophophoraa: Bryozoa, Phoronida, dan Brachiopoda memiliki tentakel bers di selling rmuluinya 223 XX DAFTAR IS RINCI J1LID Ut Filum Mollusca: Anggota fikim Mollusca memiliki kak berocor, mass vsera, dan suatu mantel 204 Filum Annelida: Anggotafilum Annelida adalah cacing Ierscgmen 27 Filum Arthropoda: Anggotafilum Arthropoda memilki segmentsi regional, tungkai bersendi, dan eksoskeleon 250 Selomata: Deuterostoma 240 Filum Echinodermata: Anggota filum Echinodermata :memiliki sistem pembuluh air dan simersi dial sekunder 240, Filum Chordata: Anggota filam Chordata melipati dua subilum invertebrta dan semua verebrata 242 3A tvolusi dan Keancharagaman Vertebrata 247 ChosdaeaInvenebrta dan Asal Mila Vreebrata 247 [Empat ciri anatomis yang merupakan karaluersik ‘Blum Ghowdag nar Chordata invertcbrata memberikan petunjule mengenai ‘sal mula verebrata 248 Pengancar Verebrata 250 Pia neural (neural rest sealissi(chephalzation) yang nyata, tulang punggung, dan sistem srkulast ertucup merupakan karakternik khas Subfilam Vertebrata 250 Gambaran umum keanekaragaman vertebrata 251 Superkelas Agnatha: Verebraa Tak Berhang 251 Lamprey dan Hagfsh adalah hewan agratha yang mash hhidup sae ini 23 Superkelas Gnathoscomata I than Rahang vertebrataberevlusi dari kerangka penyokong cela faring 254 Endoskeleon bertulangrawan yang diperkuat ol butiran berkalsium merupakan ciri dagnestik untuk Kels Chondrichthyes 255 Endoskeleton bertulang, operkulum, dan kancung renang ‘merupakan citi kas Kelas Osteichthyes 256 Superkelas Gnathosomata 1 Terapoda 258 Amphibia adalah Kelas retrapoda yang paling cua 258 vols tclur amniotik memperbesar keberhasilan vertebrau di darat 261 Sema hewan amniota memiliki cst yang dinarankan dari reptia 20 Burung berawal sebagai reptiiatebang 26 Keanckargaman mamalia meningkat jam serelh terjadinya kepanahan di masa Kretaseus 20 Primata dan Filogeni Homo sapiens 275 Byolusiprimata memberikan pandangan bagi pemahaman asal mula manusia 25 Manusia merupakan cabang yang sangae muda pada ppohon sila vertebrata 276 aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page that is unavailable for viewing or reached your viewing limit for this book. aa You have either reached a page 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Lexy, SB. “The Challenge of Antibiotic Rsisance” Sciewfic American, Mat 1998, Rineian mergenai ancaman, penebabnya, dan kerungkinan slus sae meen ana Madigan, ML, dan BL. Nas. “Exemophiles” Scns American, Ape 1997, Sorte pongnte pokes yung era bidap len Knghemgen yang tidak berabubar dan encimensim yang. memmungkinkan mereka ‘akan hal erat. Mier LV, “Bacterial Gene Swapping in Natu” Scenic American, Jnuat 1998, Nenjcskan kondist yang mengplang atau mencxgah perkaren fe ananpeses Monaseri R. "Deep Dwolers” Sconce Mewn, 29 Maret 1997. Meng pokarve yang hidup sngat dalam i alarm lark Bur ‘The Race Aunts Lethal Mivobar. Chery Chase, MD: The Howard Hughes Metical Inte. 1996 Sebuah buku dengan. banyak pen, dengan ders yang bik yang menyelidi pony infest yang mengancam ‘misyaraat manus moder. Tortory GJ, BR. Fake, din CL, Cue, Mobily: An feducton, eh ‘Redwood City, CA: Bejamin/Cummings, 1998. Bu tts ureum. Tavis J. “Spying Diseases From the Sy” Scere Nrox. 2 Agustus 1997 ‘Menjelakanpenggnaan data stl unca mengidenifks pemandangan tempat cgay ako penyi 124 UNIT LIMA * SEJARAH EVOLUSI DARI KEAN Kunjungi edisi khusus The Biology Place for BIOLOGY, Fith Edition, di heep:/iwowebiology.com/campbell, Kunjungilah Bab 27 untuk memperoleh sumber online, rermasuk aktivtas belajar,ujian praktek, dan hubungan ke situs web (web ste) bertkut: “American Society for Microbiology (ASM)” Inilah home page ASM. Dengan 40,000 anggota di seluruh dunia, inilah organises eunggal imu-lmu hayai yang tetua dan erbesse. Situs ini meliputi proyek Biofilm Imaging dan the National Col- lkction for Micobiclogy Teaching and Learning, “Antoni van Leeuwenhoek (1632-1723)" Biognfi online penemu mikroskop dan penemu baker “TIGR Microbial Database” Situs ini mengandung, suaru dafar genom mikroba yang telsh éipublikascan aru yang sedang dalam proses pengururan. Kunjungi The Biology Plice untuk meacobs aktvias beljar yang discbut “Investigating the Bucteral Genome” oleh Tom Terry, Universiy of Connecticut, yang membuat penggunaan ekstensif dari database TIGR ini “Tntreduction to the Archaea: Lifts Extremists” Suatu pengantar Domain Arkhaea dengan link ke sumberdaya arkhaca di seluruh dunia. EKARAGAMAN BIOLOGIS idak ada lagi pemandangan yang lebib menyenangkan dibandingkan dengan melibarsekian vibu makhluk hidup dalam sects keci ai,” tis Anton van Leewwenbuek stead menemulean dunia milroba lebih dari tiga abad slams, Dunia iru itemuckan kerbali ole setiap mahasiswa biologi dengan cara me- Bhat melalui mikzoskop, setetes air kolam yang penuh dengan berancka raga makhluk yang kita scbue pretista (lihat foro dé sebelah). Protisa bers: eukarioik, dan babkan provisa yang paling sederhana sekalipun jauh lebih kompleks dibandingkan dengan prokariota, Eulariota pertama yang berevolusi dari nenck moyang ‘prokariotit kemungkinan bersift uniseller dan oleh sebab itu disebut protsta. Kata ita mengandung arti sesuatu yang angat tua (Bahasa Yunani provos, ‘pertama’). Eukariota pertome itu bukan saja merupakan pendahulu protita modem yang sangat berancka ragam, tetap juga nenck moyang bagi semua eukariote—tumbuhan, fungi, dan hewan, Dua di ancara bagian-bagian yang paling ber- ‘makna dalam sejarah kehidupan—asal mula sel ewkaritik dan hemunculan eukariota multseluler berikurmya—terjadi selama evolu protisa. Bab ini akan menelusuriasal mula selseleuavitik: menghej eanckaragamen, evolui, dan ekologi prota; dan membaas asal ula evolsioner dari organisme muiseller PENGANTAR PROTISTA. Paling tidak satu miliar tahun sebelum munculnya cumbuhan, fangi, dan hewan, terdapat procista, turunan eukariorik paling awal dari prokariota. Fesil tertua yang diduga foil protista adalah cobjek dari zaman Prakambrium yang disebut acritarch (Bahas YYanani, “tidak jelas asa usulnya’) yang berumur 2,1 miliar hun, Beberapa di antaranya memiliki ukuran dan struktur yang sesuai dengan kalie sista (lapsul pelindung) yang lepas, mirip dengan kkulit pelindung yang dibuat oleh protista tertentu saat ini Miliaran tahun pertama evolusi eukariota tampaknya menghasi- Jaan beberapa garis keturunan yang berlinan. Protista adalah cukariota yang paling berancka ragam Semua protsts adalah cukariota, akan tetapi protsta sangat ber aneka ragam, sehingga hanya sedikit korakteristik umum lain yang dapat diseburkan tanpa perkecualian, Sesungguhnya, varias protista dalam hal struktur dan fangsi, melebihi kelompok orgs- nisme lainnya. Sebagian besar dari sekicar 60.000 speses protsta yang dikerahui hidup saat ini bersifat uniseluler,tetapi ada bebe- ‘apa spesies berkoloni dan bersifac multiseluler. Karens sebagian bhesar prossta hersifae uniselule, maka protista dapar dianggap sebagai organisme eukariotik yang paling sederhana. Tetapi pads ‘ingkat seluler, kebanyakan protista luar biasa kompleksnya— paling rumic di ancara semua sel. Kita seharusnya dapat menduge hal itu, dari organisme yang dalam batasannya sebagai suatu sel tunggal harus melakukan semua fungsi dasir yang dilakukan olch kumpulan scl-sel cerspesiaisasi yang menyusun cubuk ASAL MULA KEANEKARAGAMAN EUKARIOTIK Tenganear Protita Prosista adalah euariow yang paling berancka ragam Simbiosisteribac dalam munculnya cakariota dari prokariora Sistematika dan Flogeni Protsta Taksa monofiletik muncul dari peneliian modern dala siste- smatika prot AAnggou calon kingdom Arkinacos tidak meni ddan dapat mowakili gris kerurunan awal eukatiotik Calon kingdom Euglenozoa mencakup flagela autotrofik dan hetcroerofil Rongga (alveoli) subpermukaan adalah citi khas calon kingdom Alveolata Suat kumpulin eukasiots mubiseluker yang berancki gam bergerak dengan menggunakan pseudopodia (kaki semu) Jamar lendir memilki adapeasi strakrural dan siklus hidup yang meningkatkan peran ekologisnya sebagai pengucal Diatom, alga pirang, alga cokelat, dan jamur ait adalah anggora clon kingdom Seamenopila Adaptasi serukrural dan biokimiawi membancu rampet laut bet- tahan hidup dan bereprodulsi di per laut Beberapa alga menilii siklus hidup dengan pexilran generasi muliscller haploid dan diploid Alga menth (cal kinglom Rhodophyta) vidal memiliki Nagel Alga hija dan tumbuhan kemungkian memiliki nenck moyang foroautowofik yang sama Mukiseluaritas muncul secara independen beberapa kali ivokonde Bahan dengan hak cipta tumbuhan dan hewan, Setiap protista uniseluler sama sekali tidak analog dengan sebuah sel cunggal dari manusia atau organisme multiseluler lainnya, tetapi protista uniseluler itu sendici adalah suate organisme, seiempuena cumbuhan atau hewan yang utuh, Protista sangat beraneka ragam dalam hal metabolisme, dan scbagai kelompok, protista merapakan organisme yang paling berancka ragam dalam hal nucrisi di antara seluruh eukarioca Sebagian besar protista memiliki metabolisme yang bersifat aerobik, yang menggunakan mitokondria uncuk respirasi selulernya (beberapa protista tidak memiliki mitokondria dan dapat hidup dalam lingkungan anaerobik atau mengandung, bakteri mucualistik yang melakukan respirasi). Beberapa procista adalah foroaucorrof dengan kloroplas, beberaps lagi adalah heterotrof yang meayerap molekul organik atau menelan partikel makanan yang lebih besar, dan yang lainnya adalah miksotrof, chpat melikukan forosintesis dan nutisi heterotrofik (contohnys adalah Euglena, yang ditunjukkan pada GaMar 28.1). Sangat bermanfaat dalam konteks ekologis uncuk mengelompokkan seanekaragaman nutrisitersebut ke dalam tiga kelompok: protista, yang menelan makanannya (seperti hewan), atau protozoa (cunggal, protozoan); protista yang melakukan absorbsi (seperti fungi: kelompok ini tidak memiliki nama umum lain); dan protista forosintetik (seperti tumbuhan), algse (canggal, alga) Meskipun sangat umum digunskan, istlah protozoa dan algae tidak memiliki dasar dalam filogeni dan tidak ada nilai pentingnya dalam taksonomi. (Isilah age mengec pada foroautotiof akuatik yang relatif sedeshana, yang meliputi beberapa organisme yang kadang-kadang dikbsifilasikan dalam kingdom tumbuban.) Meskipun semua alga memiliki klorofil, 4. pigmen “primer” yang sima dengan yang ditemukan pada ‘tumbuhan, alga sangat berbeda dalam hal pigmen asesorisnya— pigmen yang menangkap panjang gelombang cahaya yang tidak dapat ditangkap Klorofil 2. Pigmen pigmen tersebut meliputi bentuk lain klorofil (kehijauan), karoten (kuning-oranye), xan- tofil (Kecokelatan), dan fikobilin (variasi merah dan bira). Cam- puran pigmen dalam Moroplas akan memberikan warna khas pada alga. Banyak di antara nama ilmiah dan nama umum untuk alga didasarkan pada warna (misalnya, Chlorophyta adalah alga hijau). Sebagian besar protista hersifae motil, memiliki flagela atau sila pada suacu saat dalam siklus hidup mereka. Penting uncuk, dipahami bahva flagela pada prokariota dan eukariota bukanlah sruleur yang homolog. Flagela prokariota melekat pada per- ‘mukaan sel (lihat cawaaR 27.5). Kebalikannya, flagela dan silia eukariota merupakan perpanjangan sitoplasma, dengan berkas rikrotubul yang tertutup olch membran plasma (lihac Gasman 7.24). Silia dan flagela eukariota memiliki ultastrukcur dasar yang sama (pengaruran mikrotubul 9 + 2), akan terapi silia lebih pendek dan lebih banyak. Sila dan flagela menggerakkan sel dengan kekuatan denyutan berirama, yang analog dengan dayung pada sampan (lihat Gavia 7.23b). Reproduksi dan siklus hidup sangat bervariasi pada protista Pembelahan mitosis erjadi pada sebagian besar protsta, tetapi terdapat banyak variasi dalam proses yang belum diketahui pada, cukariotalainnya. Beberapa protista secaraeksklusif adalah asek- sual yang lain juga dapat bereprodulesi ecara seksual atau paling 126 Flag Binele mata — Deck Flags kode haps ‘Vdeule é enero S-—_—— \orog Cadangin ——= =o ‘makina > but if oe ratte paramilon) E Membean 2 phsma, GAMBAR 28.1 * Euglena: contoh eukariota yang sengat kompleks. PProtsta ini merupakan salah satu organisme yang unum menempati air kolam yang Keruh. Gamba ini menunjakkan struktur yang lebih kkompleks pada mikroorganisme eukariotk dibandingkan dengan pro- kariota. Euglena menggunakan flagela “bertinsel” yang panjang untuk ‘menarik dirinya sendiri melalui er. Bink mata, suatu organel ber- pigmen, berfungs! sebagai pelindung cahaya. Bergantung pada possi ‘organise itu, bint maia itu memungkinkan hanya eahaya der arch tertentu saja yang dapat mencapai detektor cahaya, pembengkaken dekat dasarflagela panjang ity, Hasinye adslah pergerakan orgenisime tersebut menyu cahaya dengan intensitas yang tepat, suatu adaptasi yang penting untuk meningkatkan fotosintesis. Euglena tidak memil dinding sel, akan tetapi memiiki lempengan fleksbet kuat yang ferbuat dari protein di bawah membran plasmanya. Vakuola Kontrakil berfungsi ‘sebagai ‘pompa lambung.” yang mengeluarkan kelabihan air yang ‘masuk ke dalam sel melalui proses osmosis dar ingkungan hipotonik. Kolesihan makanan yang dibuat di kloroplas col akan disimpan dalam

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