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Bioactive Foods
and Extracts
Cancer Treatment and Prevention

Bioactive Foods
and Extracts
Cancer Treatment and Prevention
Edited by
Ronald Ross Watson
Victor R. Preedy
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v
Contents
Preface ix
Acknowledgments xi
Editors xiii
Contributors xv
SECTION I Herbal Medicines and Bioactive Foods
and Cancer Treatment
1Chapter Glucosinolates in Brassica and Cancer 3
Pablo Velasco, Marta Francisco, and María Elena Cartea
2Chapter MK615, an Extract of the Japanese Apricot (ume): A Promising
Anticancer and Anti-In ammatory Compound 31
Tokihiko Sawada and Keiichi Kubota
3Chapter Rhodiola and Related Plants: A Role in Cancer Prevention and Therapy 37
Kelly J. Gauger and Sallie Smith Schneider
4Chapter Greco-Arab and Islamic Herbal Medicine and Cancer Treatment/Prevention 49
Omar Said, Hilal Zaid, and Bashar Saad

5Chapter Indian Vegetarian Diet and Cancer Prevention 67
Manjeshwar Shrinath Baliga, Jason Jerome D’Souza,
Raghavendra Haniadka, and Rajesh Arora
6Chapter Isothiocyanate-Modi ed Pathways in Cancer Prevention and Treatment 87
Jana Jakubikova and Jan Sedlak
7Chapter Is Tulsi a Panacea for Cancer Prevention and/or Therapy? An Evidence-
Based Revisit 105
Manjeshwar Shrinath Baliga, Farhan Fazal, Rajesh Wilson,
and Rajesh Arora
8Chapter Culinary Spices in Cancer Chemoprevention 123
John R. Endres, Irfan Qureshi, Amy Clewell, and Alexander G. Schauss
vi Contents
SECTION II Bioactive Foods and Supplements
in Cancer Prevention
9Chapter Bioactive Foods in Cancer Prevention 143
Rakesh Sharma and Jose Katz
1Chapter 0 Curcumin-Mediated Cellular Responses in Chemical Carcinogenesis:
In Vivo Studies 181
Girish B. Maru, Asha G. Ramchandani, Gaurav Kumar,
and Rachana Garg
1Chapter 1 Prunes and Plums in Health Promotion 205
Felina Marie Cordova and Ronald Ross Watson
1Chapter 2 Cancer Preventive Phytochemicals from Southeast Asian Countries:
In Vivo Activities and Underlying Molecular Mechanisms 211
Akira Murakami
1Chapter 3 Mushroom-Derived Substances for Cancer Prevention and Treatment 223
Nili Ruimi, Nesly Dotan, Yaniv Menachem, Judy Bartuv-Tal,
and Jamal Mahajna
1Chapter 4 Dietary Phytochemicals in Prevention and during Cancer Treatment 237
Ali-Reza Waladkhani

1Chapter 5 Effect of Phytochemicals on Stress Management and Mental Health 257
Ali-Reza Waladkhani
1Chapter 6 Fruits, Vegetables, and Their Extracts in Health of the
Upper Gastrointestinal Tract 269
Simin Bolourchi-Vaghe
1Chapter 7 Signi cance of Processing for the Chemopreventive Potential
of Tomato-Based Products 279
Valeri V. Mossine and Thomas P. Mawhinney
1Chapter 8 Glycemic Index, Glycemic Load, and Cancer Risk 301
Helen G. Mulholland, Liam J. Murray, and Marie M. Cantwell
Contents vii
SECTION III Cancers Specifi c, Targeted Therapies with
Bioactive Foods and Their Products
1Chapter 9 Bioactive Foods and Extracts in Prostate Cancer Prevention 319
Faysal A. Ya and Wassim Kassouf
2Chapter 0 Control of Prostate Cancer Proliferation and Gene Expression
Using Herbal Supplements 329
Joseph M. Wu and Tze-chen Hsieh
2Chapter 1 Plant-Derived Antioxidants and Use in Prevention and
Treatment of Prostate Cancer 353
Carlie Margaret Abersold and Simin Bolourchi-Vaghe
2Chapter 2 Cruciferous Vegetables and Their Components in the
Prevention of Breast Cancer 363
Susan E. Steck
2Chapter 3 The Role of Flavonoids in Fruits and Vegetables Related to
Breast Cancer Prevention 379
Catherine Christie, Nancy Correa-Matos, Lauri Wright,
Julia A. Watkins, and Alex Bouziane
2Chapter 4 Chemopreventive Properties of Ginseng Compounds on Colon Cancer 393
Chong-Zhi Wang and Chun-Su Yuan

2Chapter 5 Effects of Pentacyclic Triterpenes from Olives on Colon Cancer 403
M. Emília Juan and Joana M. Planas
2Chapter 6 Fruit Phenolics in Colon Cancer Prevention and Treatment 415
Michael A. Lea
2Chapter 7 Diet, Epigenetics, and Colonic Fermentation and Their Role in
Colorectal Cancer 427
Ramesh P. Arasaradnam and K.D. Bardhan
2Chapter 8 Alcohol Consumption and Risk of Colorectal Cancer 441
Brenda W.C. Bongaerts and Matty P. Weijenberg
2Chapter 9 Plants Antioxidants and Lung Cancer Risk 455
Alberto Ruano-Raviña, Mónica Pérez-Ríos, and Juan Miguel Barros-Dios
viii Contents
3Chapter 0 Epidemiologic Review of Head and Neck Cancers, Oral Precancers,
and Dietary Risk Factors in India 471
Mia Hashibe
IV SECTION Nonbotanical Dietary Components
3Chapter 1 Vitamin D and Cancer 481
Arnaud Teichert and Daniel D. Bikle
3Chapter 2 Dietary Selenium and Liver Cancer 497
Divinia N. Stemm and Howard P. Glauert
3Chapter 3 Cancer, Probiotics, and Clinical Practice 505
Gian Vincenzo Zuccotti, Fabio Meneghin, and Dario Dilillo
3Chapter 4 Indian Herbal Medicine for Cancer Therapy and Prevention: Bench to Bedside 519
Rajesh Arora, Poonam Malhotra, R. Chawla, D. Gupta, R.K. Sharma, and
Manjeshwar Shrinath Baliga
3Chapter 5 Dietary Intake and the Development of Lung Cancer 545
Lauri Wright, Catherine Christie, and Julia A. Watkins
3Chapter 6 Herbs and Bioactive Compounds in Prevention and Treatment of
Hepatocellular Carcinoma 555
Anuradha Sehrawat and Vijay Kumar

3Chapter 7 New Zealand Christmas Tree: Historic Uses and Cancer Prevention 583
Felina Marie Cordova and Ronald Ross Watson
3Chapter 8 Nutrition and Colorectal Cancer 587
Mitra Rangarajan and Gerard E. Mullin
3Chapter 9 Feijoa (Pineapple Guava) Fruit: A Role in Health Promotion? 603
Mercedes C. Argüelles and Ronald Ross Watson
4Chapter 0 Indian Bael (Aegle marmelos) for the Prevention/Treatment of Cancer:
Weighing Cost versus Bene t 609
Manjeshwar Shrinath Baliga, Harshith P. Bhat, Ponemone Venkatesh,
R.K. Sharma, and Rajesh Arora
Index 623
ix
Preface
There is a considerable historical record for the cancer preventative effects of consumption of
vegetables, fruits, and herbs. Indeed, the usage of foods and their extracts as therapeutic tools
appears in ancient and modern cultures. Recently, however, there has been an increase in the amount
of scienti c research relating to the effects of plant products in cancer treatment. Clearly informa-
tion is vital for the researcher, physician, and government regulator with increased availability and
media evidence that such agents may have ef cacy. Especially in the United States, the use of
botanicals and their extracts is widely available and unregulated. Therefore, information from sci-
enti c research is critical in helping researchers and healthcare professionals make decisions on the
bene ts, risks, or value of botanicals and their extracts in the prevention and treatment of cancers.
Ronald Ross Watson

xi
Acknowledgments
The work of the editorial assistant, Bethany L. Stevens, in communicating with authors, working
with the manuscripts and the publisher was critical to the successful completion of the book and is
much appreciated. Her daily responses to queries and collection of manuscripts and documents
were extremely helpful. Support for her work was graciously provided by Elwood Richard, treasurer

of the National Health Research Institute in support of its mission to communicate to scientists
about bioactive foods and dietary supplements was vital ( />nhri/). This was part of their efforts to educate scientists and the lay public on the health and eco-
nomic bene ts of nutrients in the diet as well as supplements. Finally, Nguyen T. Nga and Mari
Stoddard of the Arizona Health Sciences Library were instrumental in  nding the authors and their
addresses in the early stages of the book’s preparation.

xiii
Editors
Ronald Ross Watson, PhD, attended the University of Idaho but graduated from Brigham Young
University in Provo, Utah, with a degree in chemistry in 1966. He earned his PhD in biochemistry
from Michigan State University in 1971. His postdoctoral schooling in nutrition and microbiology
was completed at the Harvard School of Public Health, where he gained two years of postdoctoral
research experience in immunology and nutrition.
From 1973 to 1974 Dr. Watson was assistant professor of immunology and performed research
at the University of Mississippi Medical Center in Jackson. He was assistant professor of microbiol-
ogy and immunology at the Indiana University Medical School from 1974 to 1978 and associate
professor at Purdue University in the Department of Food and Nutrition from 1978 to 1982. In 1982,
Dr. Watson joined the faculty at the University of Arizona Health Sciences Center in the Department
of Family and Community Medicine of the School of Medicine. He is currently professor of health
promotion sciences in the Mel and Enid Zuckerman Arizona College of Public Health.
Dr. Watson is a member of several national and international nutrition, immunology, cancer, and
alcoholism research societies. He is presently funded by the National Heart Blood and Lung Institute
to study heart diseases. In addition, he has an NIH grant from NCCAM to study dietary supple-
ments in modulating immune function and thus the heart structure and function. For 30 years he
was funded by Wallace Research Foundation to study dietary supplements in health promotion.
Dr. Watson has edited more than 35 books on nutrition and 53 other scienti c books. He has pub-
lished more than 500 research and review articles. His major research emphasis have been in the
 eld of dietary supplements in health promotion in senior citizens, diabetic, arthritic, hypertensive,
and heart failure patients and development of new dietary supplements and immunomodulators to
change the structure of heart and thus the function in old or obese mice and humans with metabolic

syndrome.
Professor Victor R. Preedy, BSc, DSc, FIBiol, FRCPath, FRSPH, is currently professor of nutri-
tional biochemistry in the Department of Nutrition and Dietetics, King’s College London and hon-
orary professor of clinical biochemistry in the Department of Clinical Biochemistry, King’s College
Hospital. He is also director of the Genomics Centre, King’s College London. He is presently a
member of the School of Medicine, King’s College London. Professor Preedy obtained his PhD in
1981 and in 1992 he received a membership to the Royal College of Pathologists, based on his pub-
lished works. He was elected as a Fellow of the Royal College of Pathologists in 2000. In 1993 he
gained a DSc degree for his outstanding contribution to protein metabolism. Professor Preedy was
elected as a Fellow to the Royal Society for the Promotion of Health (2004) and The Royal Institute
of Public Health (2004). In 2009 he was elected a Fellow of the Royal Society for Public Health
(RSPH). The RSPH is governed by the Royal Charter and Her Majesty The Queen is its Patron.
Professor Preedy has written or edited more than 550 articles, which includes over 160 peer-reviewed
manuscripts based on original research and 85 reviews and 30 books. His interests pertain to mat-
ters concerning nutrition and health at the individual and societal levels.

xv
Carlie Margaret Abersold
Chief Clinical Dietitian
River Garden Hebrew Home
Jacksonville, Florida
Ramesh P. Arasaradnam
Clinical Sciences Research Institute
Warwick Medical School,
University Hospital Coventry and
Warwick
Coventry, United Kingdom
Mercedes C. Argüelles
University of Arizona
Tucson, Arizona

Rajesh Arora
Division of Radiation Biology and
Radioprotection
Institute of Nuclear Medicine and
Allied Sciences
New Delhi, India
and
Department of Chemistry and
Biochemistry
Trakia University
Stara Zagora, Bulgaria
Manjeshwar Shrinath Baliga
Department of Research and
Development
Father Muller Medical College
Mangalore, Karnataka, India
K.D. Bardhan
Department of General Medicine
Rotherham General Hospital
NHS Trust
Rotherham, United Kingdom
Juan Miguel Barros-Dios
Department of Preventive Medicine
and Public Health
University of Santiago de Compostela
and
Service of Preventive Medicine
Clinic University Hospital of Santiago
de Compostela
Santiago de Compostela, Spain

and
CIBER de Epidemiología y
Salud Pública
CIBERESP
Santiago de Compostela, Spain
Judy Bartuv-Tal
Cancer Drug Discover Program
MIGAL—Galilee Technology Center
Kiryat Shimona, Israel
Harshith P. Bhat
Research Centre
Maharani Lakshmi Ammani
Women’s College
Bangalore, Karnataka, India
Daniel D. Bikle
Veterinary Affairs Medical
Center
University of California
San Francisco, California
Simin Bolourchi-Vaghe
Department of Nutrition and
Dietetics
University of North Florida
Jacksonville, Florida
Contributors
xvi Contributors
Brenda W.C. Bongaerts
Department of Pathology
Maastricht University
Maastricht, The Netherlands

Alex Bouziane
Department of Nutrition and Dietetics
University of North Florida
Jacksonville, Florida
Marie M. Cantwell
Centre for Public Health
Queen’s University Belfast
Belfast, Northern Ireland, United Kingdom
María Elena Cartea
Department of Plant Genetics
Spanish Council for Scienti c Research
(CSIC)
Pontevedra, Spain
R. Chawla
Division of Radiation Biology and
Radioprotection
Institute of Nuclear Medicine and
Allied Sciences
New Delhi, India
Catherine Christie
Department of Nutrition and Dietetics
University of North Florida
Jacksonville, Florida
Amy Clewell
Natural and Medicinal Products Research
AIBMR Life Sciences
Puyallup, Washington
Felina Marie Cordova
University of Arizona
Tucson, Arizona

Nancy Correa-Matos
Department of Nutrition and Dietetics
University of North Florida
Jacksonville, Florida
Jason Jerome D’Souza
Father Muller Medical College
Mangalore, Karnataka, India
Dario Dilillo
Department of Pediatrics
University of Milan
Milan, Italy
Nesly Dotan
Cancer Drug Discover Program
MIGAL—Galilee Technology Center
Kiryat Shimona, Israel
John R. Endres
Natural and Medicinal Products Research
AIBMR Life Sciences
Puyallup, Washington
Farhan Fazal
Father Muller Medical College
Mangalore, Karnataka, India
Marta Francisco
Department of Plant Genetics
Spanish Council for Scienti c Research (CSIC)
Pontevedra, Spain
Rachana Garg
Advanced Centre for Treatment, Research
and Education in Cancer (ACTREC)
Tata Memorial Centre

Navi Mumbai, India
Kelly J. Gauger
Pioneer Valley Life Sciences Institute
Center for Excellence in Apoptosis Research
Spring eld, Massachusetts
Howard P. Glauert
Graduate Center for Toxicology
and
Graduate Center for Nutritional Sciences
University of Kentucky
Lexington, Kentucky
D. Gupta
Division of Radiation Biology and
Radioprotection
Institute of Nuclear Medicine and Allied Sciences
New Delhi, India
Raghavendra Haniaddka
Father Muller Medical College
Mangalore, Karnataka, India
Contributors xvii
Mia Hashibe
International Agency for Research
on Cancer
Lyon, France
Tze-chen Hsieh
Department of Biochemistry and
Molecular Biology
New York Medical College
Valhalla, New York
Jana Jakubikova

Department of Tumor Immunology
Cancer Research Institute
Bratislava, Slovak Republic
and
Department of Medical Oncology
Dana Farber Cancer Institute
and
Department of Medicine
Harvard Medical School
Boston, Massachusetts
M. Emília Juan
Departament de Fisiologia
Universitat de Barcelona
Barcelona, Spain
Wassim Kassouf
Department of Surgery (Urology)
McGill University
Montreal, Québec, Canada
Jose Katz
Department of Medicine
Columbia University
and
Katz Cardiomedical Centers
New York, New York
Keiichi Kubota
Second Department of Surgery
Dokkyo Medical University
Tochigi, Japan
Gaurav Kumar
Advanced Centre for Treatment, Research

and Education in Cancer (ACTREC)
Tata Memorial Centre
Navi Mumbai, India
Vijay Kumar
Virology Group
International Centre for Genetic Engineering
and Biotechnology
New Delhi, India
Michael A. Lea
Department of Biochemistry and
Molecular Biology
UMDNJ—New Jersey Medical School
Newark, New Jersey
Jamal Mahajna
Cancer Drug Discover Program
MIGAL—Galilee Technology Center
Kiryat Shimona, Israel
Poonam Malhotra
Division of Radiation Biology and
Radioprotection
Institute of Nuclear Medicine and
Allied Sciences
New Delhi, India
Girish B. Maru
Advanced Centre for Treatment, Research
and Education in Cancer (ACTREC)
Tata Memorial Centre
Navi Mumbai, India
Thomas P. Mawhinney
Department of Biochemistry

University of Missouri
Columbia, Missouri
Yaniv Menachem
Cancer Drug Discover Program
MIGAL—Galilee Technology Center
Kiryat Shimona, Israel
Fabio Meneghin
Department of Pediatrics
University of Milan
Milan, Italy
xviii Contributors
Valeri V. Mossine
Department of Biochemistry
University of Missouri
Columbia, Missouri
Helen G. Mulholland
Centre for Public Health
Queen’s University Belfast
Belfast, Northern Ireland, United Kingdom
Gerard E. Mullin
Division of Gastroenterology
The Johns Hopkins Hospital
Baltimore, Maryland
Akira Murakami
Division of Food Science and
Biotechnology
Kyoto University
Kyoto, Japan
Liam J. Murray
Centre for Public Health

Queen’s University Belfast
Belfast, Northern Ireland, United Kingdom
Mónica Pérez-Ríos
Department of Preventive Medicine and
Public Health
University of Santiago de Compostela
and
Galician Department of Health
General Directorate of Public Health
Santiago de Compostela, Spain
Joana M. Planas
Departament de Fisiologia
Universitat de Barcelona
Barcelona, Spain
Irfan Qureshi
Natural and Medicinal Products Research
AIBMR Life Sciences
Puyallup, Washington
Asha G. Ramchandani
Advanced Centre for Treatment, Research and
Education in Cancer (ACTREC)
Tata Memorial Centre
Navi Mumbai, India
Mitra Rangarajan
Department of Gastroenterology and Hepatology
Johns Hopkins University School of Medicine
Baltimore, Maryland
Alberto Ruano-Raviña
Department of Preventive Medicine
and Public Health

University of Santiago de Compostela
and
Galician Department of Health
General Directorate of Public Health and
Plani cation
Santiago de Compostela, Spain
and
CIBER de Epidemiología y Salud Pública
CIBERESP
Santiago de Compostela, Spain
Nili Ruimi
Cancer Drug Discover Program
MIGAL—Galilee Technology Center
Kiryat Shimona, Israel
Bashar Saad
Qasemi Research Center
Al-Qasemi Academic College
Baga Algharbiya, Israel
and
Arab American University
Jenin, Palestine
Omar Said
Qasemi Research Center
Al-Qasemi Academic College
Baga Algharbiya, Israel
and
Antaki Center for Herbal Medicine
Kufur kana, Israel
and
Research and Development Regional

Center
The Galilee Society
Shefa Amr, Israel
Contributors xix
Tokihiko Sawada
Second Department of Surgery
Dokkyo Medical University
Tochigi, Japan
Alexander G. Schauss
Natural and Medicinal Products Research
AIBMR Life Sciences
Puyallup, Washington
Sallie Smith Schneider
Pioneer Valley Life Sciences Institute
Center for Excellence in Apoptosis Research
Spring eld, Massachusetts
Jan Sedlak
Department of Tumor Immunology
Cancer Research Institute
Bratislava, Slovak Republic
Anuradha Sehrawat
Virology Group
International Centre for Genetic Engineering
and Biotechnology
New Delhi, India
Rakesh Sharma
Department of Medicine
Columbia University
New York, New York
R.K. Sharma

Division of Radiation Biology and
Radioprotection
Institute of Nuclear Medicine and Allied Sciences
Delhi, India
Susan E. Steck
Department of Epidemiology and Biostatistics
University of South Carolina
Columbia, South Carolina
Divinia N. Stemm
Graduate Center for Toxicology
University of Kentucky
Lexington, Kentucky
and
Department of Environmental Quality
Remediation and Redevelopment Division,
Michigan
Lansing, Michigan
Arnaud Teichert
Endocrine Unit
University of California
San Francisco, California
Pablo Velasco
Department of Plant Genetics
Spanish Council for Scienti c
Research (CSIC)
Pontevedra, Spain
Ponemone Venkatesh
Department of Radiobiology
Kasturba Medical College
Manipal, Karnataka, India

Ali-Reza Waladkhani
Krankenanstalt Mutterhaus Borromaerinnen
Klinikum Mutterhaus der Borromäerinnen
Trier, Germany
Chong-Zhi Wang
Tang Center for Herbal Medicine Research
and
Department of Anesthesia and Critical Care
University of Chicago
Chicago, Illinois
Julia A. Watkins
Department of Nutrition and Dietetics
University of North Florida
Jacksonville, Florida
Ronald Ross Watson
University of Arizona
Tucson, Arizona
Matty P. Weijenberg
Department of Pathology
Maastricht University
Maastricht, The Netherlands
Rajesh Wilson
Father Muller Medical College
Mangalore, Karnataka, India
Lauri Wright
Department of Nutrition and Dietetics
University of North Florida
Jacksonville, Florida
xx Contributors
Joseph M. Wu

Department of Biochemistry and
Molecular Biology
New York Medical College
Valhalla, New York
Faysal A. Ya
Department of Surgery (Urology)
McGill University
Montreal, Québec, Canada
Chun-Su Yuan
Tang Center for Herbal Medicine Research
and
Department of Anesthesia and Critical Care
University of Chicago
Chicago, Illinois
Hilal Zaid
Qasemi Research Center
Al-Qasemi Academic College
Baga Algharbiya, Israel
Gian Vincenzo Zuccotti
Department of Pediatrics
University of Milan
Milan, Italy
ISection
Herbal Medicines and Bioactive Foods
and Cancer Treatment

3
1
Glucosinolates in Brassica
and Cancer

Pablo Velasco, Marta Francisco, and María Elena Cartea
INTRODUCTION
In the last 20 years, a number of epidemiological studies have shown that tumor formation and
incidence of cardiovascular diseases are inversely related to the intake of fruit and vegetables
(Verhoeven et al., 1996). One group of vegetables that has been widely approved for its bene cial
effects on human health are the vegetables from the family Brassicaceae (= Cruciferae). The family
Brassicaceae is a large group, having about 3000 species in 350 genera, including several types of
edible plants, which are sometimes referred to as “the cabbage family.” The petals of plants of this
family have a distinctive cruciform arrangement, which is the origin of the initial term Cruciferae.
These plants can be annuals, biennials or perennials. They are well adapted to average temperatures
of 16–18°C and are thus grown during the cool season in temperate areas. Crops of Brassicaceae
are mainly distributed in temperate regions of the northern hemisphere: in the areas of Southwestern
and Central Asia, China and Japan, Europe, the Mediterranean region and North America. Brassica
production has grown steadily and vegetables represent a major part of the human diet worldwide.
Despite the great diversity among the Brassicaceae, members of only a few genera are used in
human diet (JARC, 2004).
The Brassica genus belongs to the Brassicaceae family and, economically speaking, it is the
most important genus within the tribe Brassiceae, containing 37 different species. The taxonomy of
this genus is complex. Gómez-Campo (1999) presented a complete classi cation of the genus
CONTENTS
Introduction 3
Glucosinolates and Hydrolysis Products 8
Glucosinolate-Derived Isothiocyanates 9
Glucosinolate-Derived Indoles 14
Glucosinolate-Derived Nitriles 15
Mechanisms of Cancer Protection by Glucosinolate Degradation Products 16
Glucosinolate-Derived Isothiocyanates 16
Blocking Mechanisms: Phase I and Phase II Enzymes 16
Suppression via Antiproliferative Mechanisms 17
Apoptosis 17

Glucosinolate-Derived Indoles 19
Glucosinolate-Derived Nitriles 19
Detoxi cation 19
Tumor Growth Inhibition 20
Enhancing Glucosinolates in Horticultural Crucifers: Breeding and Biotechnology 20
References 22
4 Bioactive Foods and Extracts
Brassica and its allied genera, indicating subgenera, sections, species and subspecies, later updated
by the same author (Gómez-Campo, 2003). The genus includes a group of six interrelated species of
worldwide economic importance. U (1935) studied the cytology of the genus and established the
relationships among the genomes of the six species. The three diploid Brassica species, Brassica
nigra (L.) Koch (2n = 16), Brassica oleracea L. (2n = 18) and Brassica rapa L. (2n = 20), form the
classic Triangle of U (Figure 1.1). In nature, these species have hybridized in different combinations
to give rise to the three amphidiploid species, namely Brassica carinata A. Braun (2n = 4x = 34),
Brassica juncea (L.) Czern. (2n = 4x = 36) and Brassica napus L. (2n = 4x = 8).
The genus is categorized into oilseed, forage, condiment, and vegetable crops by using their
buds, in orescences, leaves, roots, seeds, and stems. The same species can be utilized for several
uses according to different forms or types. Four species, B. oleracea, B. rapa, B. napus, and B. jun-
cea, contain crops that have a horticultural use. The principal vegetable species is B. oleracea,
which includes vegetable and forage forms, such as kale, cabbage, broccoli, Brussels sprouts,
cauli ower, and others; B. rapa includes vegetable forms, such as turnip, Chinese cabbage, and pak
choi, along with forage and oilseed types; B. napus crops are mainly used as oilseed (rapeseed),
although forage and vegetable types like leaf rape and “nabicol” are also included;  nally, the
mustard group, which is formed by three species, B. carinata, B. nigra, and B. juncea, is mainly
used as a condiment because of their seeds, although leaves of B. juncea are also consumed as
vegetables in Asian countries.
Other cruciferous vegetables that are used in human diet belong to other genera of the Brassicaceae
family. Some of them are used as condiments, as is the case of Sinapis alba (white mustard), but
most of them are used as salad crops, such as Nasturtium of cinale (watercress) and Eruca sativa,
E. vesicaria, Diplotaxis tenuifolia (rocket salads), D. muralis (wall rocket), which is reported to be

a component of mixed soups in Italy and D. erucoides (white wall rocket), which is used as a raw
salad plant in Spain, Sicily, and Malta. The use of edible D. catholica  owers has been reported in
Spain. Diplotaxis harra, an important fodder plant in North Africa, is used as a medicinal plant in
Tunisia, but, in Sicily, it is also consumed after being cooked. Diplotaxis acris, grazed by animals
and used raw in salads in North Africa, is reported to be a component of green salads in Iraq and
Jordan as well. Diplotaxis simplex is reported to be either a medium-value pasture or an edible
species in Egypt (D’Antuono et al., 2009). Finally, other crops can be used because of their roots, as
it is the case of Raphanus sativus (radish), which is an important vegetable crop worldwide,
B
B. nigra
n = 8
BC
B. carinata
n = 17
C
B. oleracea
n = 9
A
B. rapa
n = 10
AC
B. napus
n = 19
AB
B. juncea
n = 18
FIGURE 1.1 Triangle of U (Adapted from U, N. 1935. Jpn. J. Bot. 7:389–452.)

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