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Oxford Textbook of
Cancer Biology
Oxford Textbook of
Cancer Biology
EDITED BY
Francesco Pezzella
Nuffield Division of Clinical Laboratory Sciences,
Radcliffe Department of Medicine,
University of Oxford, Oxford, UK
Mahvash Tavassoli
Department Mucosal and Salivary Biology,
King’s College London,
London, UK
David J. Kerr
Nuffield Division of Clinical Laboratory Sciences,
Radcliffe Department of Medicine,
University of Oxford, Oxford, UK;
Weill Cornell College of Medicine,
New York, USA
1
3
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© Oxford University Press 2019
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First Edition published in 2019
Impression: 1
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Preface
The textbook is dead. Long live the textbook! With increased output understand that without novel basic science and the generation of
of rapidly published new data and availability of teaching material new knowledge, there cannot be sustainable innovations in cancer
on the web, it has often been predicted that the textbook will be- diagnosis and therapy. We have structured this book logically and
come extinct. However, in our experience, it has also become in- trust that the inquisitive reader will select which chapters to explore
creasingly difficult to find a comprehensive text which enables us in greater depth.
to catch up with the current state of art in multiple fields, within There is a difference between the textbooks of today and yes-
a wider contextual framework. While the high number of research terday: before, publication was the terminus or end of the work for
and review papers provide a continuous update on increasingly its authors; now, because of the integration between the printed
narrow and specialized topics in cancer biology, we think there will book and online resources, this is no longer the case. This will allow
be always a need for concise, coherent descriptions of the funda- us to annually review, revise, and update the chapters on the online
mentals on areas like cell cycle or cell death. This is particularly im- version of the book to reflect recent developments in the field.
portant for students who require a platform of basic information Finally, as this is a cancer textbook, we would like to remember
before venturing more deeply into the literature. We have assembled our parents, relatives, friends and, of course, patients whose lives
a fantastic cast of authors, each of whom are outstanding in their have been affected and in many cases, ended too soon by this dis-
field, and have attempted, when relevant, to make the translational ease. We hope this book is another small step forward in the right
link to the application of cancer biology for patient benefit. We must direction.
Acknowledgements
We would like to thank our friends Sandor Paku, Balazs Dome, and of creating this book: Andrea, Caroline, Janine, Sree, and Anya.
Andrew Reynolds for granting us permission to use the picture on We also would like to thank all the authors for their work and their
the cover of the book, illustrating a non-angiogenic tumour growing willingness and commitment to write.
in a mouse model.
We would also like to acknowledge the help and support by
Oxford University Press staff that guided us through the process
Contents
4. Genetics and genetic instability in cancer 43 15. Telomerase and immortalization 209
Mark A. Glaire and David N. Church Laura Collopy and Kazunori Tomita
7. Chemical carcinogens 79 18. Oxygen and cancer: The response to hypoxia 255
David H. Phillips Adrian L. Harris and Margaret Ashcroft
SECTION IV SECTION VI
Cancer microenvironment The biology of cancer treatment
21. Cancer-associated stroma 303 28. Principles of chemotherapy 413
Wilma Mesker and Rob Tollenaar David J. Kerr, Daniel Haller, and Jaap Verweij
22. Blood vessels and cancer 314 29. Immunotherapy and tumour resistance to
Francesco Pezzella and Robert Kerbel immune-mediated control and elimination 423
Gwennaëlle C. Monnot and Pedro Romero
23. Cancer immunology 330
Herman Waldmann 30. Biological effect of radiotherapy
on cancer cells 438
Anna Dubrovska, Mechthild Krause, and Michael Baumann
SECTION V
Global vision of cancer
SECTION VII
24. Molecular profiling in cancer research and Conclusions
personalized medicine 347
Pieter-Jan van Dam and Steven Van Laere 31. Benign tumours: The forgotten neoplasms 453
Francesco Pezzella, Adrian L. Harris, and Mahvash Tavassoli
25. Proteomics and metabolomics applications in
cancer biology 363 32. Conclusions: Cancer biology, a moveable
Pedro Cutillas and Benedikt M. Kessler feast 463
David J. Kerr, Francesco Pezzella, and Mahvash Tavassoli
26. Cancer systems biology: From molecular
profiles to pathways, signalling networks, and
therapeutic vulnerabilities 375 Index 469
Lieven Verbeke and Steven Van Laere
FRS2 fibroblast growth factor receptor substrate 2 mTOR mammalian target of rapamycin
GAB2 GRB2 associated binding protein MYC MYC proto-oncogene
GADD45 growth arrest and DNA damage 45 MYCN MYCN protooncogene, neuroblastoma derived
GATA4 GATA-binding protein 4 MYH9 non-muscle heavy chain
GC germinal centre NAD nicotinamide adenine dinucleotide
GF growth factor NBS Nijmegen breakage syndrome
GFR growth factor receptors NER nucleotide excision repair
GG-NER global genome nucleotide excision repair NF-kB nuclear factor kappa-light-chain-enhancer of
GGR global genome repair activated B cells
GRB2 growth factor-receptor-bound protein 2 NHEJ non-homologous end-joining
HAV hepatitis A virus NPC nasopharyngeal carcinoma
HBV hepatitis B virus NPM nucleophosmin
HCV hepatitis C virus NPM-ALK nucleophosmin-anaplastic lymphoma kinase
HDAC histone deacetylase NSCLC non-small cell lung cancer
HER2 human epidermal growth factor receptor 2 p53BP1 p53 binding protein 1
HIV Human Immunodeficiency Virus p130CAS breast cancer anti-estrogen resistance 1
HIF-1 hypoxia-inducible factor p130Cas breast cancer anti-oestrogen resistance protein 1
HNSCC head and neck squamous cell carcinoma pAMPK phosphorylated 5’ adenosine monophosphate-
HPV human papilloma virus activated protein kinase
HR homologous recombination PAR poly-ADP-ribose
HRP horseradish peroxidase PARG poly (ADP-ribose) glycohydrolase
HSC haematopoietic stem cells PARP poly (ADP-ribose) polymerase
HTLV human T-lymphotropic retrovirus PCAF P300/CBP-associated factor
HVS herpesvirus saimiri PCNA proliferating cell nuclear antigen
IARC International Agency for Research on Cancer PDGF platelet-derived growth factor
ICL interstrand DNA crosslink PI3K phosphoinositide 3-kinase
IDL insertion and deletion loop PKB protein kinase B
IMT inflammatory myofibroblastic tumours PLC-g phospholipase C-gamma
IR ionizing radiation PLK1 polo kinase 1
IR Insulin receptor PLWHA people living with HIV/AIDS
IRF-4 interferon regulatory factor 4 PML promyelocytic leukaemia
IRS-1 insulin receptor substrate 1 PR progesterone receptor
JAK3 Janus kinase 3 PTEN phosphatase and tensin homologue
JNK-C Jun N-terminal kinase p16 cyclin-dependent kinase inhibitor 2A
KAP-1 KRAB-associated protein-1 p53 TP53 or tumour protein
KEGG Kyoto Encyclopedia of Genes and Genomes p56 phosphoglycerate kinase
KIF5B kinesin/family member 5B P53R2 p53-inducible ribonucleotide reductase small
KRAS Kirsten rat sarcoma viral oncogene homologue subunit 2-like protein
KS Kaposi’s sarcoma RAD51, radiation repair 51 and 52
KSHV Kaposi’s sarcoma-associated herpesvirus RAD52
LC3 microtubule-associated protein 1A/1B-light RANBP2 Ran-binding protein 2
chain 3 KRAS rat sarcoma viral oncogene homologue
LFS Li-Fraumeni syndrome Rb retinoblastoma protein
LUCA last unknown common ancestor Redd1 regulated in development and DNA damage
MAP MUTYH-associated polyposis response 1
MAPK mitogen-activated protein kinase RISC RNA-induced silencing complex
MCD multicentric Castleman disease RNA ribonucleic acid
MCL mantle cell lymphoma ROS reactive oxygen species
MCPV Merkel cell polyomavirus RPA replication protein A
MDC1 mediator of DNA-damage checkpoint 1 RSV rouse sarcoma virus
MDM2 mouse double minute 2 homologue RTK receptor tyrosine kinase
MIN microsatellite instability SAC spindle assembly checkpoint
MLH1 MutL homologue 1 SASP senescence-associated secretory phenotype
MMR mismatch repair SEC31L1 SEC31 homologue A
MMP matrix metalloproteinase SH2 src homology 2
MRN Mre11/Rad50/Nbs1 complex SHM somatic hypermutation
MSH2 MutS homologue 2 SHP2 protein-tyrosine phosphatase 1D or protein-
MSN moesin tyrosine phosphatase 2C
Abbreviations xiii
Balkees Abderrahman, Department of Breast Carus, Technische Universität Dresden; and Mechthild Krause, Department of Radiotherapy
Medical Oncology, University of Texas, MD Helmholtz-Zentrum Dresden-Rossendorf, and Radiation Oncology, Faculty of Medicine
Anderson Cancer Center, Houston, USA Institute of Radiooncology-OncoRay; Cancer and University Hospital Carl Gustav Carus,
Margaret Ashcroft, Department of Medicine, Consortium (DKTK), partner site Dresden, Technische Universität Dresden; German Cancer
University of Cambridge, Cambridge, UK and German Cancer Research Center (DKFZ), Consortium (DKTK), partner site Dresden,
Germany and German Cancer Research Center (DKFZ);
Nicholas Athanasou, Nuffield Department
Nadège Gaborit, Institut de Recherche en OncoRay – National Center for Radiation
of Orthopaedics, Rheumatology, and Research in Oncology, Faculty of Medicine and
Musculoskeletal Science, University of Oxford, Cancérologie de Montpellier, INSERM U1194,
Université de Montpellier, Institut régional du University Hospital Carl Gustav Carus, Technische
Oxford, UK
Cancer de Montpellier, Montpellier, France Universität Dresden, Helmholtz-Zentrum
Michael Baumann, German Cancer Research Dresden - Rossendorf; Helmholtz-Zentrum
Center (DKFZ); and Department of Radiotherapy Kevin Gatter†, Nuffield Division of Clinical
Dresden - Rossendorf, Institute of Radiooncology
and Radiation Oncology, Faculty of Medicine Laboratory Sciences, Radcliffe Department of – OncoRay, Dresden, Germany; National Center
and University Hospital Carl Gustav Carus, Medicine, University of Oxford, Oxford, UK for Tumor Diseases (NCT), Partner Site Dresden;
Technische Universität Dresden, Germany Mark A. Glaire, Cancer Genomics and German Cancer Research Center (DKFZ); Faculty
Karim Bensaad, Department of Oncology, Immunology Group and NIHR Comprehensive of Medicine and University Hospital Carl Gustav
University of Oxford, Oxford, UK Biomedical Research Centre, The Wellcome Carus, Technische Universität Dresden; and
Centre for Human Genetics, University of Helmholtz Association / Helmholtz-Zentrum
Jessica Bullenkamp, Molecular and Clinical
Oxford, Oxford, UK Dresden - Rossendorf (HZDR), Germany
Sciences Research Institute, St. George’s
University London, London, UK Betty Gration, MRC Molecular Haematology Unit, Nicholas La Thangue, Department of Oncology,
Radcliffe Department of Medicine, Weatherall University of Oxford, Oxford, UK
Giacomo Buscemi, Department of Biosciences,
Institute of Molecular Medicine, University of Andrew P. Mazar, Monopar Therapeutics,
University of Milan, Milan, Italy Oxford, Oxford, UK Wilmette, USA
Simon Carr, Department of Oncology, University
Daniel Haller, Department of Medicine, Perelman Wilma Mesker, Department of Surgery, Leiden
of Oxford, Oxford, UK School of Medicine, University of Pennsylvania, University Medical Center, Leiden, the Netherlands
David N. Church, Cancer Genomics and Philadelphia, USA
Kingsley Micklem, Nuffield Division Clinical
Immunology Group and NIHR Comprehensive Adrian L. Harris, Department of Oncology, Laboratory Science, Radcliffe Department of
Biomedical Research Centre, The Wellcome University of Oxford, Oxford, UK Medicine, University of Oxford, Oxford, UK
Centre for Human Genetics, University of
Oxford, Oxford, UK Edward Hookway, Nuffield Department Gwennaëlle C. Monnot, Ludwig Cancer Research
of Orthopaedics, Rheumatology and Center, Department of Fundamental Oncology,
Laura C. Collopy, Cancer Institute, Faculty of
Musculoskeletal Sciences, University of Oxford, Faculty of Biology and Medicine, University of
Medical Sciences, University College London, Oxford, UK Lausanne, Lausanne, Switzerland
London, UK
Jiangting Hu, Radcliffe Department of Medicine, Udo Oppermann, Nuffield Department of
Pedro Cutillas, Cell Signalling and Proteomics
University of Oxford, Oxford, UK Orthopaedics, Rheumatology, and Musculoskeletal
Group, Barts Cancer Institute (CRUK Centre),
V. Craig Jordan, Department of Breast Medical Science, University of Oxford, Oxford, UK
Queen Mary University of London, London, UK
Oncology, University of Texas MD Anderson Francesco Pezzella, Nuffield Division of Clinical
Blossom Damania, Lineberger Comprehensive
Cancer Center, Houston, USA Laboratory Sciences, Radcliffe Department of
Cancer Center and Department of Microbiology
Robert Kerbel, Biological Sciences Platform, Medicine, University of Oxford; and Cellular
and Immunology, School of Medicine, University
Sunnybrook Research Institute, Department Pathology Clinical Service Unit, Oxford
of North Carolina, Chapel Hill, USA
of Medical Biophysics, University of Toronto, University Hospitals, Oxford, UK
Dirk P. Dittmer, Lineberger Comprehensive Cancer
Toronto, Canada David H. Phillips, Department of Analytical,
Center and Department of Microbiology and
David J. Kerr, Nuffield Division of Clinical Environmental and Forensic Sciences, School of
Immunology, School of Medicine, University of
Laboratory Sciences, Radcliffe Department of Population Health and Environmental Sciences,
North Carolina, Chapel Hill, USA
Medicine, University of Oxford, Oxford, UK; and King’s College London, London, UK
Tom Donnem, Department of Oncology,
Weill Cornell College of Medicine, New York, Karen Pulford, Emeritus Reader in
University Hospital of North Norway and the USA Immunodiagnostics, Nuffield Division
Arctic University of Norway, Tromso, Norway
Benedikt M. Kessler, Target Discovery Institute, of Clinical Laboratory Sciences, Radcliffe
Anna Dubrovska, OncoRay-National Center for Department of Medicine, University of Oxford,
Nuffield Department of Medicine, University of
Radiation Research in Oncology, Faculty of Oxford, UK
Oxford, Oxford, UK
Medicine and University Hospital Carl Gustav
† It is with regret we report the death of Kevin Gatter during the preparation of this textbook.
xvi Contributors
Chao-Nan Qian, Department of Nasopharyngeal Weatherall Institute of Molecular Medicine, Lieven Verbeke, Department of Information
Carcinoma, State Key Laboratory of Oncology University of Oxford, Oxford, UK Technology, Ghent University, Ghent, Belgium
South China, Sun Yat-Sen University Cancer Mahvash Tavassoli, Department Mucosal and Jaap Verweij, Department of Medical Oncology,
Center, Guangzhou, China Salivary Biology, King’s College London, Erasmus University Medical Centre, Rotterdam,
Lynn Quek, MRC Molecular Haematology Unit, London, UK the Netherlands
Radcliffe Department of Medicine, Weatherall Rob Tollenaar, Department of Surgery, Leiden Paresh Vyas, MRC Molecular Haematology Unit,
Institute of Molecular Medicine, University of University Medical Center, Leiden, the Radcliffe Department of Medicine, Weatherall
Oxford, Oxford, UK Netherlands Institute of Molecular Medicine, University of
Andrea Rasola, Department of Biomedical Kazunori Tomita, Cancer Institute, Faculty of Oxford, Oxford, UK
Sciences, University of Padova, Padova, Italy Medical Sciences, University College London, Herman Waldmann, Sir William Dunn School of
Pedro Romero, Ludwig Cancer Research Center, London, UK Pathology, University of Oxford, Oxford, UK
Department of Fundamental Oncology, Faculty Andrey Ugolkov, Division of Hematology and Yan-Qun Xiang, Department of Nasopharyngeal
of Biology and Medicine, University of Lausanne, Oncology, Feinberg School of Medicine, Carcinoma, Sun Yat-Sen University Cancer
Lausanne, Switzerland Northwestern University, Chicago, USA Center, Guangzhou, China
Almut Schulze, Department of Biochemistry and Pieter-Jan van Dam, Faculty of Medicine and Yosef Yarden, Department of Biological
Molecular Biology, Biocenter, University of Health Sciences, University Antwerp, Antwerp, Regulation, Weizmann Institute of Science,
Würzburg, Würzburg, Germany Belgium Rehovot, Israel
Connor Sweeney, MRC Molecular Haematology Steven Van Laere, HistoGeneX NV, Antwerp,
Unit, Radcliffe Department of Medicine, Belgium
SECTION I
The multicellular organism