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Biochemistry: The Molecular Basis of Cell Structure and Function (1977 г.)

  • Издателство: Worth Publishers, Inc.
  • ISBN / UPC: 0879010479

Biochemistry: The Molecular Basis of Cell Structure and Function (1977 г.)

  • Издателство: Worth Publishers, Inc.
  • ISBN / UPC: 0879010479

Биохимия: Молекулярната основа на клетъчната структура и функция (второ издание, книга от Алберт Ленинджър на английски език)


Albert L. Lehninger  (автор)   |   биохимия  (етикет)


Издателство:   Worth Publishers, Inc.
Език: английски език
Раздел: Химия и химични технологии


Твърда корица, голям формат  |  1104 стр.  |  2301 гр.

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Introduction: The Molecular Logic of Living Organisms 1
The Molecular Components of Cells 15
1. Biomolecules and Cells 17
2. Water 39
3. Proteins and Their Biological Functions: A Bird's-Eye View 57
4. The Amino Acid Building Blocks of Proteins 71
5. Proteins: Covalent Backbone and Amino Acid Sequence 95
6. Proteins: Three-Dimensional Conformation 125
7. Proteins: Purification and Characterization 157
8. Enzymes: Kinetics and Inhibition 183
9. Enzymes: Mechanism, Structure, and Regulation 217
10. Sugars, Storage Polysaccharides, and Cell Walls 249
11. Lipids, Lipoproteins, and Membranes 279
12. Nucleotides and the Covalent Structure of Nucleic Acids 309
13. Vitamins and Coenzymes 335
Catabolism and the Generation of Phosphate-Bond Energy 361
14. Metabolic and Energy-Transfer Pathways: A Survey of Intermediary Metabolism 363
15. Bioenergetic Principles and the ATP Cycle 387
16. Glycolysis 417
17. The Tricarboxylic Acid Cycle and the Phosphogluconate Pathway 443
18. Oxidation-Reduction Enzymes and Electron Transport 477
19. Oxidative Phosphorylation, Mitochondrial Structure, and the Compartmentation of Respiratory Metabolism 509
20. Oxidation of Fatty Acids 543
21. Oxidative Degradation of Amino Acids 559
22. Photosynthetic Electron Transport and Phosphorylation 587
Biosynthesis and the Utilization of Phosphate-Bond Energy 617
23. The Biosynthesis of Carbohydrates 623
24. The Biosynthesis of Lipids 659
25. The Biosynthesis of Amino Acids and Some Derivatives; Metabolism of Inorganic Nitrogen 693
26. The Biosynthesis of Nucleotides 729
27. The Biochemistry of Muscle and Motile Systems 749
28. Active Transport across Membranes 779
29. Biochemical Aspects of Hormone Action 807
30. Organ Interrelationships in the Metabolism of Mammals 829
Replication, Transcription, and Translation of Genetic Information 853
31. DNA and the Structure of the Genetic Material 859
32. Replication and Transcription of DNA 891
33. Translation: The Biosynthesis of Proteins 929
34. The Genetic Code 957
35. Regulation of Gene Expression 977
36. The Molecular Basis of Morphogenesis 1011
37. The Origin of Life 1031
Appendix A: A Chronology of Biochemistry 1058
Appendix B: The Research Literature of Biochemistry 1064
Appendix C: Abbreviations Common in Biochemical Research Literature 1066
Appendix D: Unit Abbreviation and Prefixes and Physical Constants 1068
Appendix E: International Atomic Weights 1069
Appendix F: Logarithms 1070
Appendix G: Solutions to Problems 1072
Index 1077






Someone long ago said that to be happy at one's work, one must be fit for it, have a sense of success in it, and not do too much of it. In the rush to finish the second edition, there was often too much to do, but even in the lowest moments my spirits were buoyed by the hundreds of heartening letters I received from students who used the first edition.


I am tempted to call this second edition a brand new text, since nearly every paragraph was rewritten and there are so many additions. But I have kept the organization and style that students and teachers found useful the first time around. I have also tried to hold the focus on central bioche­mical concepts and on the impact of new discoveries in this exciting field, rather than dwell on encyclopedic detail.


I continue to believe that biochemistry is best approached by starting with a set of organizing principles rather than scattered facts and hypotheses. Biochemistry has a structure, a set of unifying themes, all the wonderful consequences of genetic control over the shape and function of proteins. The compelling sense of this pattern of organizing principles and their implications is set forth in an Introduction (page 3), which I call "The Molecular Logic of Living Organisms." As with the first edition, the rest of the book builds upon this introductory essay, beginning with the structures and prop­erties of biomolecules, moving to a study of energy-yielding processes and energy-requiring processes, and ending with the culmination of our journey —the replication, transcrip­tion, and translation of genetic information.


The book is now cross-referenced with page numbers, not just chapter numbers. All enzymatic reactions are balanced, the metabolic pathway illustrations are essentially complete, and all enzyme names are included. (I am using the recom­mended enzyme names of the 1972 report of the Enzyme Commission.) The problems have been completely re­worked and there are more of them than before.


There is so much other new material in the second edi­tion that I can only take room here to mention the most salient. There are many more applications of biochemical knowledge to problems of human health and disease. There are two new chapters particularly relevant to human bio­chemistry—"Biochemical Aspects of Hormone Action" and "Organ Interrelationships in the Metabolism of Mammals." Among other things, the hormone chapter deals with the organization of the endocrine system, the relationship of the hypothalamus and pituitary, and the role of hormone re­ceptors and intracellular messengers, such as cyclic AMP. The chapter on integration of metabolism includes an ac­count of the metabolic characteristics of each organ as well as the metabolic interplay between organs, metabolic rate, transport of nutrients and gases by the blood, the function of the kidneys in adjustment of urine composition, and the special role of the liver and adipose tissue in processing and distributing fuel supplies. I have also added a discussion of starvation and diabetes mellitus to show how the human organism adjusts its metabolism in response to stress.


In the area of proteins and enzymes, I have included de­tailed discussion of the conformation of proteins, the kinet­ics and thermodynamics of protein folding and unfolding, and the sequential and all-or-none models of allosteric tran­sitions. There is also a new chapter on the art and science of protein purification and characterization. The enzyme chapters are expanded with new material on enzyme reac­tion mechanisms and kinetics, including bisubstrate reac­tions. And I have written a new chapter on vitamins and coenzymes.


The most completely rewritten portion of the book is Part 4. The chapter on DNA includes new material on the struc­ture of bacterial and viral genomes as well as much new in­formation on eukaryotic chromosomes, including repetititve sequences and inverted repetitions. The chapter on DNA replication and transcription has also been revised and up­dated to include the important new advances of the last few years, as have the chapters on protein synthesis and the amino acid code. The chapter on regulation of gene expres­sion has received particular attention because of the great strides made in this important area of cell biology. The mate­rial on morphogenesis and self-assembly, as well as that on the origin of life, has also been substantially revised.


All told, I found the writing of this second edition to be a renewed biochemical education for myself. I can only hope I may have made the way a little easier for others.


Many of the chapters have been reviewed by experts in the fields concerned: Bruce M. Alberts, Daniel I. Arnon, William Beranek, Jr., Clanton C. Black, Jr., Ralph A. Brad-shaw, Alexander C. Brownie, Richard M. Caprioli, Waldo E. Cohn, Stanley Dagley, Julian E. Davies, Richard E. Dickerson, John T. Edsall, Irving Geis, Govindjee, Franklin M. Harold, William F. Harrington, Jens G. Hauge, Bernard L. Horecker, William P. Jencks, Thomas H. Jukes, Walter Kauzmann, Daniel E. Koshland, Jr., Robert Lehman, John M. Lowenstein, Alan H. Mehler, Alton Meister, Daniel J. O'Kane, Peter L. Pedersen, Norman S. Radin, William J. Rutter, Carl Sagan, Norman G. Sansing, Irwin H. Segel, Thomas P. Singer, David B. Sprinson, Donald F. Steiner, Jack L. Strominger, Charles Sweeley, Thomas E. Thompson, Gordon M. Tomkins, John L. Westley, and William B. Wood. I owe them much for their critical advice and encouragement. Moreover, many teachers and students took the time to send me useful suggestions based on their experience with the first edition. As before, I will greatly appreciate comments from those who use this new book.


For help in preparing this second edition I must also express my deepest thanks to two fine teachers of biochemis­try: Herbert Friedmann of the University of Chicago, who read every chapter at least twice, examined each statement with microscopic attention, and ruthlessly skewered many bad phrases, and Richard Mintel, who enthusiastically devel­oped many new problems and their solutions, made useful suggestions for teaching certain topics, and, not least, con­tributed many hours to patient proofreading, always with cheerful ebullience. I also wish to thank Daniel DiMaio, a Johns Hopkins medical student, for introducing me to his "Cell Game" and for formulating many of the problems in Part 4. In addition, I thank Linda Hansford, who laid out the pages and backstopped our proofreading. And I must thank my secretary Peggy Ford for again keeping every­thing flying while typing manuscript against impossible deadlines. My thanks to Chris Mello, too.


Again, it was a pleasure to work with all the good people of Worth Publishers, Inc.; they produced this book with their usual care and high standards. But above all, I thank my wife, who has, on top of everything else, endured my writing six books —she is an everlasting source of encouragement.


Albert L. Lehninger

Sparks, Maryland May, 1975






This book is written for students who are taking their first and perhaps their only course in biochemistry, whether as undergraduates or as graduate or medical students. I under­took this task because I want to convey to students my pic­ture of what this science has recently become. Biochemistry is no longer a mere catalog of the biological occurrence and enzymatic reactions of a large number of organic com­pounds. In the last few years it has acquired, along with many new facts, a set of organizing principles which have made it a much simpler field to comprehend, and, at the same time, a more powerful way of analyzing many impor­tant problems in biology.


How has this come about? Each field of scientific study at some time in its evolution undergoes a profound transition in which a collection of widely scattered facts and hypothe­ses crystallizes into a logical pattern, unified by a few basic concepts. Biochemistry has been undergoing such a transi­tion, stimulated by new experimental findings and new in­sights. Among these are the recognition of the principles of energy transfer in cells, the mechanisms by which the major metabolic pathways are regulated, the importance of mem­branes, ribosomes, and other ultrastructural elements of cells in their molecular activities, and the far-reaching conclusion that the amino acid sequence determines the three-dimen­sional conformation of protein molecules and thus their bio­logical functions. The new knowledge of the molecular basis of genetics, which has transformed all of biology, has had the most profound influence. Because of these developments biochemistry now has a central story, a leitmotiv, which I have tried to express in simple terms in the Introduction.


This book is concerned primarily with biochemistry at the cell level, where its organizing principles are most clearly evident. Central concepts are emphasized rather than an en­cyclopedic treatment of biochemical details. There are four major parts in the book:

  1. Biomolecules
  2. Energy-yielding processes
  3. Energy-requiring processes
  4. Transfer of genetic information


These are subdivided into what I believe is a logical progres­sion of chapters, each of which is a manageable "package" for both students and teachers, equivalent to the content of one lecture or discussion period. I agree with many teachers that the structure and properties of some biomolecules may best be taught together with their metabolism. This approach is quite feasible using this book, although for student conve­nience I have chosen to collect most of the material on the structure, chemistry, and occurrence of the various types of biomolecules into one section. I believe this makes for easy reference, while still allowing for flexibility of approach.


Biochemistry has many new frontiers today. I have tried to sketch out some of the most promising in chapters on the regulation of protein synthesis and its role in cell differentia­tion, the molecular basis of self-assembly and mor­phogenesis, and the origin of life. These chapters may well be out-of-date soon, but I hope they will serve to acquaint students with some of the biochemistry of the future.




Many may think it foolhardy for a single author to attempt to write a comprehensive textbook of biochemistry. However, my publishers have made it possible to enlist the criticism and advice of a number of chemists, biochemists, and bio­logists expert in research and/or teaching in the areas cov­ered by the book. Each chapter has been read and criticized by at least one and often several authorities. To them I owe a great deal, not only for kind encouragement and sometimes deservedly blunt criticism, but also for the insight and per­spective that only the real expert can convey. It is perhaps inevitable that some errors of fact, interpretation, or empha­sis will be found, but I trust no one will attribute these to anyone but me. I will greatly appreciate receiving from stu­dents and teachers alike their comments, criticisms, notice of errors, and advice about improvements that can be made in later printings or editions.


To the following reviewers I give my most sincere thanks: Jay Martin Anderson, Christian B. Anfinsen, Robert E. Beyer, R. G. S. Bidwell, Rodney L. Biltonen, Konrad E. Bloch, Ben­jamin Bouck, Daniel Branton, Robert H. Burris, Melvin Calvin, Roderick K. Clayton, Helena Curtis, Robert E. Davies, Bernard D. Davis, John T. Edsall, Paul T. Englund, Allan H. Fenselau, J. Lawrence Fox, Richard Goldsby, Ursula Johnson Goodenough, Guido Guidotti, Gordon G. Hammes, William F. Harrington, Edward C. Heath, Harold G. Hempling, Donald P. Hollis, Lloyd L. Ingraham, Andre T. Jagendorf, William P. Jencks, Daniel E. Koshland, Jr., Sir Hans A. Krebs, Myron Ledbetter, William J. Lennarz, Richard C. Le-wontin, Julius Marmur, Daniel Nathans, Leslie Orgel, Peter L. Pedersen, Keith R. Porter, David Prescott, John Sinclair,

Gunther Stent, Jack L. Strominger, Maurice Sussman, Serge N. Timasheff, and William B. Wood. My thanks go to many others, acknowledged at the end of the book, who gen­erously gave me permission to use drawings, electron micro­graphs, and other illustrative material.


I am also grateful to the officers and staff of Worth Pub­lishers for their genuine interest in the needs of both stu­dents and teachers, their appreciation of the struggles of a university author, and above all, their desire to produce an educationally useful book.


My colleagues in the Department of Physiological Chem­istry of The Johns Hopkins School of Medicine furnished much advice and also took on many responsibilities which the gestation of this book had forced me to neglect. I also owe a great deal to Johns Hopkins medical students, who taught me whatever I have learned about teaching. Two of them, Bill Scott and Penny Pate, gave me much help in the early stages of preparation of the manuscript; one happy out­come is that they are now Mr. and Mrs. William Wallace Scott, Jr. To Linda Hansford I am particularly indebted for invaluable help with proof-reading, indexing, checking of problems, and collection of data and references. Thanks also to Ronald Garrett, who photographed the molecular models and to my secretary Peggy Ford, who not only effectively marshalled my time and attention among teaching, research, departmental administration, and book-writing, but also typed many chapters of the manuscript.


Finally, I want to express my deep appreciation to my family, who patiently endured the many weekends and eve­nings that were devoted to writing and who gave encourage­ment when it was most needed.


Albert L. Lehninger

Sparks, Maryland March, 1970

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Albert L. Lehninger
Worth Publishers, Inc.
New York
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