英漢對照分子生物學導論(第三版)（簡體書）

王勇，江蘇大學食品與生物工程學院生物技術系副教授，2002-2016年承擔本科生和碩士研究生分子生物學雙語教學任務，2004年至今承擔本科生分子生物學雙語教學任務，2007-2011年承擔本科生分子生物學英語教學任務。

Foreword
前言
With the incoming of the third edition of An Introduction to Molecular Biology with Chinese Translation, we have to tell everybody with a heavy heart that: we have lost contact with Dr. Lapan since June 2019. After failure of many contacting attempts ever since, we were connected to the corresponding author of his previously published paper. Unfortunately, the answer is also being out of contact. As such, we could only wish everything of Dr. Lapan be alright!
在《英漢對照分子生物學導論》第三版即將出版之際，我們不得不懷著沉重的心情告訴大家：自2019年6月以來我們與勒潘博士失去了聯繫。在後續多次聯繫嘗試失敗後，我們聯繫到了他先前發表論文的通訊作者，很遺憾得知也是失聯。如此，惟願勒潘博士一切安好！
In this new edition,“Molecular Identification” and “Gene Editing” sections are added into chapters 2 and 10 respectively. The former introduces application of molecular biology knowledge in social practice using nucleic acid testing and DNA fingerprinting as examples. The latter briefly outlines development stages of gene editing technology and functioning mechanism of the latest gene editing system. Due to absence of Dr. Lapan in this revision, no audio files are available for these two sections. Please make allowance for this.
本次再版分別在第2章、第10章增加了“分子鑒定”與“基因編輯”兩節內容。前者以核酸檢測、DNA指紋分析為實例介紹分子生物學知識在社會實踐中的應用；後者簡要概述基因編輯技術的發展階段及最新基因編輯系統的作用機理。由於勒潘博士缺席本次修訂工作，因此這兩節內容沒有音頻文件，希望大家諒解。
Additionally, the following revisions have been made, including: addition of ten figures into chapters 5 to 7 and chapter 9, corrections/revisions to over sixty words/sentences, supplement of glossary into the book, and updating of the accessory teaching courseware. We sincerely hope that this new edition could help everybody better understand and learn knowledge in basic theory and practical application of molecular biology.
此外，還完成了下面幾項修訂工作，包括：增加了十幅插圖到第5～7章及第9章中，修正了六十多處字句，增補了名詞解釋到書中，更新了配套教學課件。我們誠摯希望本書的再版能幫助大家更好地理解與學習分子生物學基礎理論及其實際應用知識。

Best wishes,
Yong Wang
Xianghui Qi
March 2024
致以良好祝願！
王勇齊
向輝
2024年3月

Foreword to the First Edition
第一版前言
One of the wonderful things about molecular biology is that the field depends on constant communicationImportant questions can only be answered through the collaboration of researchers from all over the world, who each contribute their piece of the puzzle and build on each others workBut this lively interaction also poses challenges, particularly the problem of languageHow can scientists collaborate if they speak and publish in different languages?
關於分子生物學的美好事物之一就是這一領域的發展依賴於不斷的交流。重大問題只能通過全世界研究人員的合作才能找到答案，每個研究小組為解答難題貢獻出他們的研究成果，並在他人研究成果的基礎上開展更深入的研究。但是要開展這種密切合作也面臨著許多挑戰，特別是語言問題。如果科學家們說著不同的語言、用不同的語言發表文章，那他們之間怎麼去開展合作呢？
At current stage, most molecular biological researches are published in English, as are most international conferencesThe language has become so central because a large number of scientists and research centers that have made important contributions to molecular biology were, and still are, based in England and the United StatesThe predominance of one language is not necessarily fair, but it is a reality that most professionals working in the field must face.
現階段，大多數分子生物學研究報告用英文發表，大多數國際會議也用英語發言。由於對分子生物學做出過重大貢獻的許多科學家和研究中心都在英國和美國，現在也仍然是這些國家在發揮著主導作用，因此英語已經成為首要的語言。一種語言起主導作用不一定是公平的，但這是大多數在這一領域工作的專業人員必須面對的現實。
For Chinese students, learning biology in English can be very exciting, but also quite challenging. Chinese and English are very different languages, with completely different phonetics and writing systemsOn top of this, biology textbooks and articles in English are especially difficult to understand, even for native English speakers. There is a lot of unique vocabularies, and there may be sentence structures particular to scientific writing.
對中國學生來說，用英語學習生物學會是很令人興奮的，但也會是頗具挑戰性的。漢語和英語是很不一樣的語言，具有完全不同的語音和語法體系。此外，生物學的英語教材和文章特別難以理解，即使對母語為英語的讀者來說也是如此。書中有許多特殊的詞匯，還會有一些特殊的科技寫作方面的慣用句子結構。
In many cases, Chinese students are presented with a giant textbook only in English and instructors hope for the bestOur experience as teachers of molecular biology in China is that this approach is unrealistic. Most students in China at this time, especially those who are on a course of scientific study, do not have enough training in English to understand textbooks written for native English speakersStruggling with the language often comes at the expense of actually learning the concepts and principles of molecular biology.
許多時候，中國學生拿到手的是一本厚厚的英文教材，教師希望學生從中獲得最大的收益。我們在中國講授分子生物學的體會是：這樣的做法是不切合實際的。這一階段的大多數中國學生，特別是那些還處於科學研究課程學習階段的學生，並不具備足夠的英語能力去理解為母語是英語的讀者編寫的教材。在語言上的些許進步實際上是以犧牲理解分子生物學概念與原理為代價的。
We have put together this book with the hope of creating a more realistic, more sympathetic, and more specialized approach to learn molecular biology in English. Advanced students can learn by reading the English portion of the book, and refer to the Chinese translation to check their understanding, or for reference on individual words. Beginners in English can learn from the Chinese portion, and refer to the English for an introduction to vocabularies.With both languages present, students and instructors have more options for how to learn!
我們編寫此書的目的是想為用英語學習分子生物學提供一種更實際、更讓人喜歡和更專業的方法。程度好的學生可以閱讀本書的英文部分，並參考中文譯文以檢查他們的理解情況，或參考個別單詞的中文含義。英語程度差一點的學生可以從本書的中文部分學習，並參考英文而開始掌握專業詞匯。使用這樣的英漢對照教材，在如何學習上學生和教師都可以有更多的選擇！
An added feature of this book is that all the English texts have been accompanied by audio files.They are especially helpful for those who intend to improve their listening comprehension of academic EnglishMeanwhile,the elaborately designed figures will enable readers to understand concepts and principles in molecular terms much more easily.
本書的另一個特點是：所有的英語課文都有配套的音頻文件。它們對那些希望提高自身專業英語聽力水平的人特別有幫助。同時，精心設計的插圖能使讀者更容易地從分子水平上理解相關的概念與原理。
The book covers the material that we feel an advanced, undergraduate student of molecular biology should knowUnlike many textbooks used for molecular biology, this is not meant as a reference book, a large volume where one can look up any facts about the cellThose kinds of volumes have their use, but we feel that it is unproductive to present students with such piles of informationIf students can master all of the information present even in this relatively brief book, they will already be more informed than most students of the field, and even some professionals!
本書包括的內容是我們認為一個分子生物學高年級本科生應該掌握的。與許多分子生物學教材不同，它不是一本參考書，不是一本讓人查閱關於細胞各方面內容的大部頭著作。那些大部頭著作自然有它們的用武之地，而我們覺得向學生提供這麼一大堆信息並不是很有成效的做法。如果學生能掌握這本相對來說比較簡單的書中的所有內容，他們就已經是這一領域學生中出類拔萃的了，他們甚至可能比一些專業人員還要強！
It is with great excitement that we invite you to learn molecular biology with us in EnglishThis is an important age for China and the rest of the world!As China becomes every day more prosperous, its research centers and companies are improving and have much to offer scientific communities around the worldAt the same time, the continued development of China depends on the knowledge and skills that have been developed in the West and continue to flourish thereIn molecular biology, there is much to gain if we can share and communicate with each other!
能邀請你與我們一道用英語來學習分子生物學讓我們覺得很興奮。中國和世界的其他國家正處於一個重要的時代！隨著中國的日益進步與繁榮，中國的研究中心和公司正在取得更大的進展，它們正在為世界科學共同體提供更多的研究成果。同時，中國的持續發展需要了解西方已經擁有的和即將涌現的知識和技能。在分子生物學領域，如果我們能夠共享知識與技能並互相交流，我們就都可以取得更大的成就！

Sincerely,
Sylvain W. Lapan
Yong Wang
October 2007
西爾維恩 W. 勒
潘王勇
謹識
2007年10月

Foreword to the Second Edition
第二版前言
Dear Readers,
With this new edition of An Introduction to Molecular Biology with Chinese Translation we continue our mission that has captivated and motivated us from the beginningFirstly, to convey fundamental concepts of how genetic information is replicated, expressed and modifiedSecondly, to assist Chinese readers in learning words and phrases in the English language used to describe these processesWe have been honored by the positive feedback of students over the past decade, and used these comments to maintain and improve the qualities of this book that students value the most: the native English main text, accurate translation, clear figures and audio files narrated in American English
親愛的讀者：
在新版《英漢對照分子生物學導論》中，我們繼續努力達成促使與激勵我們編寫本書的初衷——首先，講述遺傳信息是如何進行復制、表達和改變的基本概念；其次，幫助中國讀者學習描述這些生命過程的英語詞匯與短語。我們很榮幸在過去十年裡收到不少學生對本書的積極反饋，這些反饋表明了學生最關心本書的以下質量：地道的英語課文，準確的中文翻譯，清晰的圖解以及美式英語音頻文件。在新版《英漢對照分子生物學導論》中，這些質量都得到了維護與提升。
The past decade has witnessed remarkable advancements in biotechnology — the reduction in cost of DNA sequencing and synthesis, the application of genome engineering techniques such as CRISPR, the use of transposases for generating DNA libraries and profiling chromatin, to name just a fewIn each case, these advances would not have been made if not for basic research into genome structure, stability, and expressionWe have an enormous amount still to gain from the basic study of molecular biology in a variety of organisms, as well as the innovative application of discoveries to solve challenges in biotechnologyIt is our sincere hope that this new edition will advance the goal of introducing Chinese and Englishspeaking students to this exciting field and fostering global collaboration.
過去十年裡生物技術領域取得了令人矚目的成就——DNA測序與合成成本顯著降低、CRISPR等基因編輯技術的應用、使用轉座酶生成DNA文庫及分析染色質等等，不勝枚舉。具體到某一方面，所有成就的取得都是建立在對基因組結構與穩定性以及基因表達特性的基礎研究之上的。現今，分子生物學基礎研究仍有待在許多不同生物中展開，生物技術領域所面臨的挑戰也需要創造性地應用分子生物學基礎研究成果來應對。我們誠摯希望新版《英漢對照分子生物學導論》能有助於引導中國學生與英語國家學生進入這一令人興奮的領域並培養全球合作精神。

Best wishes,
Sylvain W. Lapan
Yong WangJan 2018致以良好祝願！
西爾維恩·W. 勒潘王勇2018年1月

Introduction 緒論1

Chapter1Amino Acids to Proteins 第1章從氨基酸到蛋白質5
1.1Protein Composition 1.1蛋白質的組成5
1.2Protein Conformations 1.2蛋白質的構象8
1.2.1Describing Protein Structure 1.2.1描述蛋白質的結構9
1.2.2Chemical and Physical Basis for Protein Folding 1.2.2蛋白質折疊的化學和物理基礎13
1.3Protein Structure and Function：A Few Examples 1.3蛋白質的結構與功能：幾個例子16
1.4The Dynamics of Proteins 1.4蛋白質動力學18
1.5Experiments 1.5實驗研究21
Summary 小結21
Vocabulary 詞匯22
Review Questions 習題23
Exploration Questions 思考題23

Chapter2Nucleic Acids 第2章核酸24
2.1Properties of a Genetic Material 2.1遺傳物質的性質24
2.2Nucleic Acids and DNA structure 2.2核酸與DNA結構25
2.2.1Nucleotides 2.2.1核苷酸25
2.2.2General Structure of Nucleic Acids 2.2.2核酸的一般結構27
2.2.3Structure of DNA 2.2.3DNA的結構30
2.3Molecular Identification 2.3分子鑒定33
2.3.1Identification of DNA Molecules 2.3.1DNA分子鑒定33
2.3.2Identification of RNA Molecules 2.3.2RNA分子鑒定35
2.4DNA as the Genetic Material 2.4DNA作為遺傳物質37
2.5DNA in the Cell 2.5細胞中的DNA38
2.6RNA(Ribonucleic Acid) 2.6RNA（核糖核酸）40
2.7Experiments 2.7實驗研究42
Summary 小結44
Vocabulary 詞匯44
Review Questions 習題45
Exploration Questions 思考題46

Chapter3Transcription in Prokaryotes:Mechanism and Regulation 第3章原核生物轉錄：機理與調控47
3.1Why Use an RNA Intermediate 3.1為什麼使用RNA作為中間物47
3.2Mechanism of Transcription 3.2轉錄機理49
3.2.1Promoters 3.2.1啟動子49
3.2.2RNA Polymerase 3.2.2RNA聚合酶51
3.2.3Transcription Mechanism in Three Phases 3.2.3轉錄機理的三個階段53
3.3Regulation of Gene Expression in Prokaryotes 3.3原核生物基因表達調控58
3.3.1Coordinate Regulation 3.3.1協同調控59
3.3.2The Lac Operon 3.3.2乳糖操縱子61
3.3.3The Trp Operon 3.3.3色氨酸操縱子64
3.3.4Ara and Gal Operons 3.3.4阿拉伯糖與半乳糖操縱子70
3.4Experiments 3.4實驗研究72
Summary 小結76
Vocabulary 詞匯77
Review Questions 習題78
Exploration Questions 思考題79

Chapter4Transcription in Eukaryotes:Mechanism and Regulation 第4章真核生物轉錄：機理與調控80
4.1Eukaryotic RNA Polymerases 4.1真核生物RNA聚合酶80
4.2Eukaryotic Promoters 4.2真核生物啟動子82
4.3General Transcription Factors and Initiation 4.3通用轉錄因子與轉錄起始83
4.3.1TFⅡD 4.3.1TFⅡD84
4.3.2Other TFⅡs 4.3.2其他TFⅡs86
4.3.3General Transcription Factors for RNA Polymerase Ⅰ and Ⅲ 4.3.3RNA聚合酶Ⅰ和Ⅲ的通用轉錄因子86
4.4Specific Transcription Factors and Transcriptional Regulation 4.4特異轉錄因子與轉錄調控88
4.4.1Activators 4.4.1激活蛋白89
4.4.2Repressors 4.4.2阻遏蛋白91
4.4.3Enhancers and Silencers 4.4.3增強子和沉默子91
4.5Structures of Specific Transcription Factors 4.5特異轉錄因子的結構93
4.5.1DNA-Binding Motifs in Prokaryotes 4.5.1原核生物DNA結合基序94
4.5.2DNA-Binding Motifs in Eukaryotes 4.5.2真核生物DNA結合基序95
4.6Experiments 4.6實驗研究97
4.6.1RNA Polymerase Targets 4.6.1RNA聚合酶的目標97
4.6.2Modularity of Specific Transcription Factors 4.6.2特異轉錄因子的模塊化98
Summary 小結99
Vocabulary 詞匯100
Review Questions 習題100
Exploration Questions 思考題101

Chapter5mRNA Modifications in Eukaryotes 第5章真核生物mRNA的修飾102
5.1Capping 5.1加帽102
5.2Polyadenylation 5.2聚腺苷酸化104
5.3Splicing 5.3剪接106
5.3.1The Basic Splicing Reaction 5.3.1基本的剪接反應106
5.3.2Proteins Involved in Splicing 5.3.2在剪接中發揮作用的蛋白質108
5.3.3Self-Splicing 5.3.3自我剪接110
5.3.4Trans-Splicing 5.3.4反式剪接111
5.3.5Reasons for Introns 5.3.5內含子存在的原因113
5.4mRNA Editing 5.4mRNA編輯115
5.5Experiments 5.5實驗研究118
Summary 小結120
Vocabulary 詞匯120
Review Questions 習題120
Exploration Questions 思考題121

Chapter6Translation 第6章翻譯122
6.1The Genetic Code 6.1遺傳密碼122
6.2Mechanism of Translation in Prokaryotes 6.2原核生物翻譯機理125
6.2.1Initiation 6.2.1起始125
6.2.2Elongation 6.2.2延伸128
6.2.3Termination 6.2.3終止132
6.3Translation in Eukaryotes 6.3真核生物翻譯132
6.4tRNA Structure and Wobble 6.4tRNA的結構與搖擺136
6.4.1Anti-codons 6.4.1反密碼子137
6.4.2Wobble 6.4.2搖擺139
6.5Experiments 6.5實驗研究140
6.5.1Deciphering the Genetic Code 6.5.1破譯遺傳密碼140
6.5.2Direction of Translation 6.5.2翻譯的方向141
Summary 小結142
Vocabulary 詞匯143
Review Questions 習題143
Exploration Questions 思考題144

Chapter7Regulation of Gene Expression in Eukaryotes 第7章真核生物基因表達調控145
7.1Histones and Transcriptional Regulation 7.1組蛋白與轉錄調控146
7.1.1Histones and DNA Organization 7.1.1組蛋白與DNA組織146
7.1.2Histones and Transcription 7.1.2組蛋白與轉錄149
7.1.3Covalent Modification of Histones 7.1.3組蛋白的共價修飾151
7.1.4Proteins that Recognize and Modify Histones 7.1.4識別和修飾組蛋白的蛋白質153
7.2Post-Transcriptional Regulation 7.2轉錄後調控156
7.3Nuclear Export 7.3細胞核輸出158
7.4RNA Stability 7.4RNA穩定性160
7.4.1mRNA Stability Regulation by Proteins 7.4.1蛋白質調控mRNA穩定性160
7.4.2mRNA Stability Regulation by Small RNAs 7.4.2小RNA調控mRNA穩定性163
7.5Translational Control 7.5翻譯調控165
7.5.1Global Control 7.5.1全局控制166
7.5.2mRNA-Specific Control 7.5.2mRNA特異性控制167
7.6mRNA Localization 7.6mRNA定位170
7.7Protein Regulation 7.7蛋白質調控172
7.8Experiments 7.8實驗研究172
7.8.1Beads-on-a-string Structure 7.8.1線珠結構172
7.8.2Repression of Gene Expression in Heterochromatin 7.8.2異染色質中基因表達的抑制174
Summary 小結174
Vocabulary 詞匯175
Review Questions 習題176
Exploration Questions 思考題177

Chapter8DNA Replication 第8章DNA復制178
8.1Semi-Conservative Replication 8.1半保留復制178
8.2Initiation of Replication 8.2復制的起始179
8.3Semi-Discontinuous Replication 8.3半不連續復制181
8.4Elongation of Replication and its Proteins 8.4復制延伸及其相關蛋白184
8.4.1Helicase and SSBs 8.4.1解旋酶與SSB184
8.4.2DNA Polymerases 8.4.2DNA聚合酶185
8.4.3Explanation for 3′→5′Synthesis 8.4.3關於3′→5′合成188
8.4.4Primers 8.4.4引物191
8.5DNA Topology 8.5DNA拓撲學193
8.6DNA Replication in Eukaryotes 8.6真核生物DNA復制195
8.6.1Initiation of DNA Replication in Eukaryotes 8.6.1真核生物DNA復制起始197
8.6.2Telomeres 8.6.2端粒199
8.7Experiments 8.7實驗研究200
Summary 小結203
Vocabulary 詞匯203
Review Questions 習題203
Exploration Questions 思考題204

Chapter9Mutations and Mutation Repair 第9章突變與突變修復205
9.1DNA Damage and Mutations 9.1DNA損傷與突變205
9.2Point Mutations 9.2點突變206
9.2.1Mismatched Base 9.2.1錯配的堿基206
9.2.2Spontaneous Mutations 9.2.2自發突變207
9.2.3Induced Mutations 9.2.3誘發突變209
9.3Insertions and Deletions 9.3插入和缺失213
9.3.1Strand Slippage 9.3.1鏈滑動213
9.3.2Transposons 9.3.2轉座子214
9.3.3Intercalating Agents 9.3.3嵌入劑215
9.4Large-Scale DNA Changes 9.4大規模DNA變化216
9.5Consequences of DNA Mutations 9.5DNA突變的後果217
9.5.1Consequences of Point Mutations 9.5.1點突變的後果217
9.5.2Consequences of Insertions and Deletions 9.5.2插入和缺失的後果219
9.5.3Consequences of Translocations 9.5.3易位的後果220
9.5.4Mutation Hot Spots 9.5.4突變熱點221
9.6Mutation Repair 9.6突變修復222
9.6.1Direct Reversal 9.6.1直接回復223
9.6.2Mismatch Repair 9.6.2錯配修復223
9.6.3Nucleotide Excision Repair 9.6.3核苷酸切除修復225
9.6.4Base Excision Repair 9.6.4堿基切除修復227
9.6.5Double-Stranded Break Repair 9.6.5雙鏈斷裂修復228
9.7Experiments 9.7實驗研究230
9.7.1Nucleotide Excision Repair and Human Disease 9.7.1核苷酸切除修復與人類疾病230
9.7.2The Ames Test 9.7.2埃姆斯測驗法231
Summary 小結232
Vocabulary 詞匯232
Review Questions 習題233
Exploration Questions 思考題234

Chapter10Recombination 第10章重組235
10.1Homologous Recombination 10.1同源重組235
10.1.1Mechanism for Crossing-Over 10.1.1交換機理236
10.1.2Mechanism for Double-Stranded Break Repair 10.1.2雙鏈斷裂修復機理240
10.1.3The RecBCD Pathway 10.1.3RecBCD途徑241
10.1.4Gene Conversion 10.1.4基因轉換241
10.2Non-homologous Recombination 10.2非同源重組243
10.2.1Transposons 10.2.1轉座子243
10.2.2Retrotransposons 10.2.2反轉錄轉座子249
10.2.3Bacteriophage λ Integration 10.2.3λ噬菌體的整合250
10.3Gene Editing 10.3基因編輯251
10.3.1ZFNs System 10.3.1ZFN系統251
10.3.2TALENs System 10.3.2TALEN系統253
10.3.3CRISPR/Cas System 10.3.3CRISPR/Cas系統254
Summary 小結257
Vocabulary 詞匯257
Review Questions 習題258
Exploration Questions 思考題258

Answers to Review Questions 習題答案259

Index(Chinese) 中文索引262

Index(English) 英文索引268

Glossary 名詞解釋274

References 參考文獻302