The authors and publishers are pleased to present the twenty-ninth
edition of Harper's Illustrated Biochemistry.
The first edition of
this text, entitled Harper's Biochemistry,
was published in 1939
under the sole authorship of Dr Harold Harper, University of California, San
Francisco. Subsequently, various authors have contributed to the text.
Cover Illustration for the Twenty-Ninth Edition
The cover illustration for the 29th edition commemorates Elizabeth H.
Blackburn, Carol W. Greider, and Jack W. Szostak, who shared the 2009 Nobel
Prize in Physiology or Medicine for their seminal work on telomeres and the
enzyme telomerase. Telomeres comprise up to 200 copies of a repeating DNA
sequence called a G-quadruplex, a structure named for the unique cyclic
arrangement of four sets of four guanine bases hydrogen-bonded in
head-to-tail fashion that stabilize this structure. In the illustration, the
phosphodiester backbone of the DNA is represented by a ribbon and the
guanine bases by filled hexagons fused to filled pentagons. The spectral
color gradation from purple to red facilitates tracing the progression of
the polynucleotide chain. The four sets of cyclic tetra-guanine units can be
seen in center stacked from top to bottom and tilted roughly 45° from
left to right (Adapted from Protein Data Bank ID no. 2KKA).
As a consequence of the unidirectional nature of DNA
replication, each time a chromosome is replicated, the number of
G-quadruplex units is reduced. When the supply of telomere units is
completely exhausted, replication ceases and the cell transitions to a
senescent state. Scientists speculate that the telomere serves as a
countdown clock that limits the number of times a somatic cell can divide,
and hence its lifespan.
Changes in the Twenty-Ninth Edition
Consistent with our goal of providing students with a text that describes
and illustrates biochemistry in a medically relevant, up-to-date,
comprehensive, and yet relatively concise manner, in addition to updating
every chapter, significant new material appears in this edition.
Each chapter now begins with a brief statement of its
objectives followed by a brief account of its biomedical importance. A major
addition is the inclusion of over 250 multiple-choice exam questions with
answers given in an answer bank.
Major Additional Changes Include Three Entirely New Chapters:
"Biochemistry of Aging"
"Biochemistry of Cancer"
Additional Significant Changes Include:
• Inclusion of aspects of epidemiology in the chapter on
"Bioinformatics and Computational Biology".
• New figures that
illustrate key approaches for identifying possible active sites,
ligand-binding sites, and other interaction sites (Section I), and various
aspects of metabolism (Section II).
• New tables that summarize
aspects of metabolic diseases, including those of purine, pyrimidine, and
amino acid metabolism (Section III).
• Expanded discussion of
non-coding RNAs, DNA damage repair and human diseases, epigenetic factors
that control eukaryotic gene expression, the activities of miRNAs, and
powerful new assays to monitor and characterize transcription genome-wide
• New tables that address vitamin and mineral
requirements and a greatly expanded discussion of iron metabolism in health
and disease (Section VI).
Organization of the Book
Following two introductory chapters, the text is divided into six main
sections. All sections and chapters emphasize the medical relevance of
structures and functions of proteins and enzymes. This section also contains
a chapter on Bioinformatics and Computational Biology, reflecting the
increasing importance of these topics in modern biochemistry, biology, and
explains how various
cellular reactions either utilize or release energy, and traces the pathways
by which carbohydrates and lipids are synthesized and degraded. Also
described are the many functions of these molecules.Section III
deals with the amino acids, their metabolic fates,
certain features of protein catabolism, and the biochemistry of the
porphyrins and bile pigments.Section IV
describes the structure and function of nucleotides and nucleic acids, DNA
replication and repair, RNA synthesis and modification, protein synthesis,
the principles of recombinant DNA technology, and new understanding of how
gene expression is regulated.Section V
deals with aspects of extracellular and intracellular communication. Topics
include membrane structure and function, the molecular bases of the actions
of hormones, and the field of signal transduction.Section VI
includes fifteen special topics: nutrition, digestion, and
and minerals; free radicals and antioxidants;
intracellular trafficking and sorting of proteins; glycoproteins; the
extracellular matrix; muscle and the cytoskeleton; plasma proteins and
immunoglobulins; hemostasis and thrombosis; red and white blood cells; the
metabolism of xenobiotics; the biochemistry of aging; the biochemistry of
cancer; clinical chemistry; and sixteen biochemically oriented case
histories. The latter chapter concludes with a brief epilog indicating some
major challenges for medicine for which biochemistry and related disciplines
will play important roles in finding solutions.Appendix
lists useful web sites and biochemical journals and others with significant
The authors thank Michael Weitz for his role in the planning of this
edition, and Brian Kearns for his key role in getting this edition ready for
publication. We also thank Mala Arora and her colleagues at Thomson Digital
for their efforts in editing, typesetting, and artwork, and Calvin "Nic"
Steussy of Purdue University for his assistance in generating the cover
Suggestions from students and
colleagues around the world have been most helpful in the formulation of
this edition. We look forward to receiving similar input in the future.
Rob Murray acknowledges with thanks Joe Varghese and Molly
Jacob as co-authors of Chapters 50, 55, and 56, Fred Keeley for his many
contributions to Chapter 48, Peter Gross for co-authorship of Chapters 51
and 57, and Margaret Rand for co-authorship of Chapter 51. Special thanks
are extended to Reinhart Reithmeier, Alan Volchuk, and David Williams for
reviewing and making invaluable suggestions for the revision of Chapters 40
Robert K. Murray
David A. Bender
Kathleen M. Botham
Peter J. Kennelly
Victor W. Rodwell