Carsten Carlberg
Eunike Velleuer
Cancer
Biology
How Science Works
Second Edition
,Cancer Biology
,Carsten Carlberg • Eunike Velleuer
Cancer Biology
How Science Works
Second Edition
,Carsten Carlberg Eunike Velleuer
InLife, Institute of Animal Reproduction Department for Cytopathology
Polish Academy of Sciences Heinrich-Heine University Düsseldorf
Olsztyn, Poland Düsseldorf, Germany
School of Medicine, Institute of Department for Pediatric Hemato-Oncology
Biomedicine Helios Children’s Hospital
University of Eastern Finland Krefeld, Germany
Kuopio, Finland
ISBN 978-3-032-18588-4 ISBN 978-3-032-18589-1 (eBook)
https://doi.org/10.1007/978-3-032-18589-1
1st Edition: Cancer Biology: How Science Works, © Springer Nature Switzerland AG 2021
© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland
AG 2021, 2026
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether
the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and trans-
mission or information storage and retrieval, electronic adaptation, computer software, or by similar or
dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publica-
tion does not imply, even in the absence of a specific statement, that such names are exempt from the
relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
If disposing of this product, please recycle the paper.
,Preface
This book begins with sobering news: approximately one in two individuals in high-
income countries will receive a cancer diagnosis during their lifetime, meaning a
malignant tumor will be detected in their body. Yet there is encouraging news as
well: fewer than half of all cancer patients die from the disease, and every second
cancer-related death is preventable. Cancer is not a single disease but a heteroge-
neous group of disorders that can arise in virtually any organ. What they all share is
uncontrolled cellular growth.
Every newly diagnosed malignant tumor in adulthood reflects an individual his-
tory of two or more decades of tumorigenesis. Cancer is traditionally viewed as a
disease of the genome, driven by accumulated DNA point mutations, as well as
chromosomal translocations, deletions, and amplifications. Tumorigenesis also
involves profound alterations in cellular identity, responsiveness to internal and
external signals, and major shifts in the transcriptome. These changes are rooted in
epigenomic dysregulation. In fact, most cancers harbor mutations and alterations
not only in their genomes but also in their epigenomes.
Cells in our body have the capacity to grow when needed: during wound healing
or in the continuous renewal of blood, skin, and intestinal tissues. Because malig-
nant transformation is typically slow, cancer is largely an age-related disease, one
that seems inevitable as we grow older. Yet tumorigenesis depends on numerous
environmental influences, including the ability of the immune system to detect and
eliminate cancer cells. Many environmental factors that either promote or protect
against cancer lie within our own control: avoiding smoking, choosing a healthy
diet, and staying physically active are key examples. Thus, cancer prevention
through lifestyle choices remains the most powerful strategy for reducing can-
cer risk.
Understanding cancer requires insight into ourselves, both in detail and within a
broader context. Basic biology explains the cellular mechanisms of growth, differ-
entiation, and cell death that make life possible. Every human being represents an
intricate interplay of hundreds of cell types, each forming specialized tissues and
organs. These processes must be precisely coordinated during embryonic develop-
ment and maintained throughout adult life to preserve homeostasis. Studying the
cellular and molecular foundations of these processes is deeply fascinating, yet also
demanding. Much of what we know about normal biology has arisen from investi-
gating what happens when these mechanisms go awry in disease.
v
, vi Preface
Accordingly, this book not only describes the fundamental mechanisms underly-
ing cancer but also aims to offer a more holistic perspective placing molecular
insights into the wider context of the personal and societal implications of lifestyle
choices.
The content of this book is linked to the lecture course “Cancer Biology,” taught
by one of us (C. Carlberg) at the University of Eastern Finland in Kuopio since
2005, alongside related courses in Molecular Medicine and Genetics, Molecular
Immunology, and Nutrigenomics. The biological concepts introduced here are com-
plemented by clinical insights drawn from daily oncology practice, contributed by
one of us (E. Velleuer). This book also relates to the books Gene Regulation and
Epigenetics: How Science Works (ISBN 978-3-031-68729-7), Aging: How Science
Works (ISBN 978-3-031-61256-5), Molecular Immunology: How Science Works
(ISBN 978-3-031-04024-5), and Nutrigenomics: How Science Works (ISBN
978-3-031-85880-2), which may interest readers seeking more detailed information.
Chapters 1, 2, and 3 provide a general overview of cancer. Chapters 4, 5, 6, 7, and
8 examine the molecular basis of the disease, Chaps. 9, 10, and 11 discuss its cel-
lular aspects, and Chaps. 12 and 13 present the principles of effective cancer ther-
apy. By integrating fundamental biological mechanisms with clinical examples, we
aim to make this book not only informative but also personally engaging. A glossary
in the appendix defines key specialized terms; gene and protein name abbreviations
are explained in the abbreviations list.
We hope readers will enjoy this visually rich book and become as enthusiastic as
we are about understanding life, and its malfunctioning counterpart, cancer biology.
Kuopio, Finland Carsten Carlberg
Düsseldorf, Germany Eunike Velleuer
December 2025
Eunike Velleuer
Cancer
Biology
How Science Works
Second Edition
,Cancer Biology
,Carsten Carlberg • Eunike Velleuer
Cancer Biology
How Science Works
Second Edition
,Carsten Carlberg Eunike Velleuer
InLife, Institute of Animal Reproduction Department for Cytopathology
Polish Academy of Sciences Heinrich-Heine University Düsseldorf
Olsztyn, Poland Düsseldorf, Germany
School of Medicine, Institute of Department for Pediatric Hemato-Oncology
Biomedicine Helios Children’s Hospital
University of Eastern Finland Krefeld, Germany
Kuopio, Finland
ISBN 978-3-032-18588-4 ISBN 978-3-032-18589-1 (eBook)
https://doi.org/10.1007/978-3-032-18589-1
1st Edition: Cancer Biology: How Science Works, © Springer Nature Switzerland AG 2021
© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland
AG 2021, 2026
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether
the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and trans-
mission or information storage and retrieval, electronic adaptation, computer software, or by similar or
dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publica-
tion does not imply, even in the absence of a specific statement, that such names are exempt from the
relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
If disposing of this product, please recycle the paper.
,Preface
This book begins with sobering news: approximately one in two individuals in high-
income countries will receive a cancer diagnosis during their lifetime, meaning a
malignant tumor will be detected in their body. Yet there is encouraging news as
well: fewer than half of all cancer patients die from the disease, and every second
cancer-related death is preventable. Cancer is not a single disease but a heteroge-
neous group of disorders that can arise in virtually any organ. What they all share is
uncontrolled cellular growth.
Every newly diagnosed malignant tumor in adulthood reflects an individual his-
tory of two or more decades of tumorigenesis. Cancer is traditionally viewed as a
disease of the genome, driven by accumulated DNA point mutations, as well as
chromosomal translocations, deletions, and amplifications. Tumorigenesis also
involves profound alterations in cellular identity, responsiveness to internal and
external signals, and major shifts in the transcriptome. These changes are rooted in
epigenomic dysregulation. In fact, most cancers harbor mutations and alterations
not only in their genomes but also in their epigenomes.
Cells in our body have the capacity to grow when needed: during wound healing
or in the continuous renewal of blood, skin, and intestinal tissues. Because malig-
nant transformation is typically slow, cancer is largely an age-related disease, one
that seems inevitable as we grow older. Yet tumorigenesis depends on numerous
environmental influences, including the ability of the immune system to detect and
eliminate cancer cells. Many environmental factors that either promote or protect
against cancer lie within our own control: avoiding smoking, choosing a healthy
diet, and staying physically active are key examples. Thus, cancer prevention
through lifestyle choices remains the most powerful strategy for reducing can-
cer risk.
Understanding cancer requires insight into ourselves, both in detail and within a
broader context. Basic biology explains the cellular mechanisms of growth, differ-
entiation, and cell death that make life possible. Every human being represents an
intricate interplay of hundreds of cell types, each forming specialized tissues and
organs. These processes must be precisely coordinated during embryonic develop-
ment and maintained throughout adult life to preserve homeostasis. Studying the
cellular and molecular foundations of these processes is deeply fascinating, yet also
demanding. Much of what we know about normal biology has arisen from investi-
gating what happens when these mechanisms go awry in disease.
v
, vi Preface
Accordingly, this book not only describes the fundamental mechanisms underly-
ing cancer but also aims to offer a more holistic perspective placing molecular
insights into the wider context of the personal and societal implications of lifestyle
choices.
The content of this book is linked to the lecture course “Cancer Biology,” taught
by one of us (C. Carlberg) at the University of Eastern Finland in Kuopio since
2005, alongside related courses in Molecular Medicine and Genetics, Molecular
Immunology, and Nutrigenomics. The biological concepts introduced here are com-
plemented by clinical insights drawn from daily oncology practice, contributed by
one of us (E. Velleuer). This book also relates to the books Gene Regulation and
Epigenetics: How Science Works (ISBN 978-3-031-68729-7), Aging: How Science
Works (ISBN 978-3-031-61256-5), Molecular Immunology: How Science Works
(ISBN 978-3-031-04024-5), and Nutrigenomics: How Science Works (ISBN
978-3-031-85880-2), which may interest readers seeking more detailed information.
Chapters 1, 2, and 3 provide a general overview of cancer. Chapters 4, 5, 6, 7, and
8 examine the molecular basis of the disease, Chaps. 9, 10, and 11 discuss its cel-
lular aspects, and Chaps. 12 and 13 present the principles of effective cancer ther-
apy. By integrating fundamental biological mechanisms with clinical examples, we
aim to make this book not only informative but also personally engaging. A glossary
in the appendix defines key specialized terms; gene and protein name abbreviations
are explained in the abbreviations list.
We hope readers will enjoy this visually rich book and become as enthusiastic as
we are about understanding life, and its malfunctioning counterpart, cancer biology.
Kuopio, Finland Carsten Carlberg
Düsseldorf, Germany Eunike Velleuer
December 2025
