Carotenoids Volume 5: Nutrition and Health pot

Carotenoids
Volume 5: Nutrition and Health
Edited by G. Britton
S. Liaaen-Jensen
H. Pfander

Birkhäuser Verlag
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Editors:
Dr. George Britton
53 Forest Road
Meols
Wirral
Merseyside
CH47 6AT
UK
Prof. Dr. Hanspeter Pfander
CaroteNature GmbH
Chief Operating Officer
Muristrasse 8e
3006 Bern
Switzerland
Professor Dr. Dr. h.c. Synnøve Liaaen-Jensen
Organic Chemistry Laboratories
Department of Chemistry
Norwegian University of Science and
Technology (NTNU)
7491 Trondheim
Norway
ISBN 978-3-7643-7500-3 e-ISBN 978-3-7643-7501-0
v
Dedication
NORMAN I. KRINSKY
29
th
June 1928 – 28
th
November 2008

For a period of more than 50 years from his Ph.D work on ‘Studies of Carotenoids and
Vitamin A Complexes with Proteins in Plasma and Tissues’, through his post-doctoral work
on vision, in the Harvard laboratory of Nobel laureate George Wald, to his long and
productive career at Tufts University, Boston, the name ‘Norman Krinsky’ has been
synonymous with ‘Carotenoids in Human Health and Nutrition’. A true giant in the field,
Norman pioneered so much of current thinking and understanding of the nutritional value and
health benefits of carotenoids, and this work is continued by the talented and dedicated group
of colleagues that he assembled at Tufts. It is no coincidence that so many chapters in this
Volume come from his associates, past and present, with him as a co-author on two of them.
A great teacher and communicator, Norman leaves us other legacies. The Gordon Research
Conferences on Carotenoids, initiated by him in 1992, continue to go from strength to
strength. He was co-chair of the 8th International Symposium on Carotenoids in Boston in
1987, and co-editor of the proceedings, published as Carotenoids: Chemistry and Biology
(editors N. I. Krinsky, M. M. Mathews-Roth and R. F. Taylor, 1990), and his more recent
book Carotenoids in Health and Disease (editors N. I. Krinsky, S. T. Mayne and H. Sies,
2004) is a most valuable collection of research reports on the subject.
Norman Krinsky’s contribution to carotenoid science is immense but he is also
remembered for his humanity and humour, his warmth, wit and wisdom. Always interested in
other people and their work, he was ever ready to guide and advise and has been a great
inspiration to so many. His perceptive comments and questions at conferences, always
constructive, never destructive, are legendary.
The editors all have our own memories of Norman, who was a good friend of all of us. He
will be very much missed; we will not see his like again.
That the field of Carotenoids in Human Nutrition and Health is as active and exciting as it
is today is due in no small part to our friend Norman Krinsky. It is most fitting and a great
honour for us to dedicate this volume to him and his memory.
vii
Contents
List of Contributors xxiii
Preface xxvii

Editors’ Notes on the Use of this Book xxix
In memoriam xxxi
Editors’ Acknowledgements xxxii
Abbreviations xxxiii


Chapter 1: Editors’ Introduction: A Healthy Debate
George Britton, Synnøve Liaaen-Jensen and Hanspeter Pfander

A. Introduction 1
B. Volume 5 2
1. Stra tegy 2
2. Relation to other volumes 2
3. Content of Volume 5 3
a) Nutrition 3
b) Carotenoids in health and disease 4
C. Conclusions 6
References 6


Chapter 2: Analysis of Carotenoids in Nutritional Studies
Frederick Khachik

A. Introduction 7
B.Isolation and Characterization 8
1.Strategy 8
2.Extraction 9
3. Sa ponification 10
4. Fractionation of carotenoids by thin-layer and column chromatography 11
viii

C. Identification and Structure Elucidation 12
D.HPLC of Carotenoids: General Aspects 12
1. Special features of carotenoids and HPLC 13
2. Stra tegy 14
3. Choice of system: Normal phase or reversed phase? 14
4. Normal phase 14
a)Silica columns 14
b) Silica-based bonded nitrile (CN) columns 16
5. Reversed phase 17
a) C
18
columns 17
b) C
30
columns 19
6. Temperature 20
7. Test chromatograms – standard mixture 20
8. Avoiding injection artefacts and peak distortion 21
E.Examples of Separations 23
1.Separation of carotenes 23
2. Separation of xanthophylls 25
3. E/Z Isomers 25
a) Carotenes 26
b) Xanthop hylls 26
4. Acyl esters 26
5. Optical isomers/enantiomers 28
F. Quantitative Analysis of Carotenoids by HPLC 28
1. Selection of an internal standard 28
a) Requirements of an internal standard 28
b) Examples of internal standards 29

c) Internal standard for carotenol esters 30
2. Use of an internal standard 30
3. Preparation of the internal standard calibration curves 31
G. HPLC of Carotenoids in Food 31
1. Green vegetables and fruits 31
2. Yellow/red fruits and vegetables containing mainly carotenes 33
3. Yellow/orange fruits and vegetables containing mainly xanthophylls and xanthophyll
esters 36
H.Analysis of Carotenoids in Human Serum, Milk, Major Organs, and Tissues 39
1. Human serum and milk 39
2. Major organs 41
I.Conclusions 43
References 43



ix
Chapter 3: Carotenoids in Food
George Britton and Frederick Khachik

A. Introduction 45
B. Distribution of Carotenoids in Vegetables and Fruits 47
1. Green vegetables and fruits 47
2. Yellow, orange and red fruits and vegetables 49
a) Fruits 49
b) Roots 52
c) Seeds 52
d) Flowers 53
e) Oils 53
3. Animal-derived food products 53

a) Eggs 54
b) Dairy produce 54
c) Seafood 54
4. Good sources 55
5. Additives, colourants 57
C. Effects of Environmental Conditions and Cultivation Practice 57
D.Effects of Storage, Processing and Cooking 58
1. Stability and loss or retention of carotenoids 58
2. Storage, cooking and processing 59
a) Transport and storage 59
b) Cooking and processing 60
3. Causes and mechanisms 61
a) Oxidation 61
b) Geometrical isomerization 62
c)Other changes 63
E.Conclusions and Recommendations 63
1. Analytical data 63
a)HPLC 63
b) Visual assessment 64
c) Instrumental 65
2. Some general conclusions 65
References 65





x
Chapter 4: Supplements
Alan Mortensen

A. Introduction 67
1. Market 67
2. Le gal 70
B. Carotenoids in Supplements 71
1. Which carotenoids? 71
2. Form ulations 73
a) Oil suspensions and oleoresins 74
b) Water-miscible formulations 74
3. A nalysis 75
C. Health Issues 76
1. Selling points 76
2. Bioavailability 77
3. Recommendations 79
References 80


Chapter 5: Microbial and Microalgal Carotenoids as Colourants and
Supplements
Laurent Dufossé

A. Introduction 83
B. Carotenoid Production by Microorganisms and Microalgae 84
1. β-Carotene 85
a) Dunaliella species 85
b) Blakeslea t rispora 86
c) Phycomyces bla kesleeanus 87
d) Mucor c ircinelloides 88
2. Lycopene 88
a) Blakeslea tris pora 88

b) Fusarium sporotrichioides 89
3. A staxanthin 89
a) Haematococcus pl uvialis 89
b) Xanthophyllomyces dendrorhous (formerly Phaffia rhodozyma) 90
c) Agrobacterium aurantiacum and other bacteria 91
4. Ze axanthin 91
5. C anthaxanthin 92
6. Torulene and torularhodin 93

xi
C. Prospects for Carotenoid Production by Genetically Modified Microorganisms 93
1. Escherichia coli and other hosts 93
2. Directed evolution and combinatorial biosynthesis 94
D. Concluding Comments 95
References 96


Chapter 6: Genetic Manipulation of Carotenoid Content and Composition
in Crop Plants

Paul D. Fraser and Peter M. Bramley

A. Introduction 99
B. Strategies for Enhancing Carotenoids in Crop Plants 101
1. General considerations 101
2. Experimental strategies 101
3. Optimizing conditions 104
a) Choice of crop 105
b) Choice of biosynthetic step(s) to target 105
c) Choice of promoter and gene/cDNA 106

d) Targeting of the transgenic protein 107
C. Examples of the Application of Metabolic Engineering to Carotenoid Formation in
Crop Plants 107
1. Tomato 107
2. Pota to 110
3. C arrot 110
4. R ice 111
5. Canola (rape seed) 111
D. Conclusions and Perspectives 111
References 112


Chapter 7: Absorption, Transport, Distribution in Tissues and
Bioavailability

Kirstie Canene-Adams and John W. Erdman Jr

A. Introduction 115
B. Absorption, Transport, and Storage in Tissues 116
1. Overview 116
2. Solubilization and incorporation into micelles 117
3. Intestinal absorption 119


xii
4. Transport in blood 121
a) Incorporation into chylomicrons 121
b) Other lipoproteins 121
5. Accumulation and distribution in tissues 122
a) General features 122

b) Blood 123
c) Liver 124
d) Adipose tissue 125
e) Eyes 125
f) Breast milk and colostrum 126
g) Breast 128
h) Male reproductive tissues 128
i) Skin 130
j) Adrenals 130
C. Bioavailability 131
1. Introduction 131
2. Effect of food matrix 132
a) Carotenoids in fruits and vegetables 132
b) Location of carotenoids 132
3. Effect of food processing 133
4. Structure and isomeric form of the carotenoid 134
a)
β
-Carotene 134
b) Lycopene 135
5. Effects of other dietary factors 135
a) Dietary fat 136
b) Inhibitors in the diet 136
c) Interactions between carotenoids 138
6. Human factors 139
a) ‘Non-responders’ 139
b) Age 139
c) Parasitic infections 140
D. Methods for Evaluating Carotenoid Bioavailability 140
1. Oral-faecal balance 141

2. Blood response 141
3. Triacylglycerol-rich fraction response 142
4. Digestion methods in vitro 142
5. Stable isotopes 143
6. Raman spectroscopy 143
E. The Future 144
References 144

xiii
Chapter 8: Carotenoids as Provitamin A
Guangwen Tang and Robert M. Russell

A.Introduction 149
B. Conversion into Vitamin A in vitro 151
C. The Conversion of Provitamin A Carotenoids into Vitamin A in vivo: Methods to
Determine Conversion Factors 153
1. Measuring radioactivity recovered in lymph and blood after feeding
radio-isotopically labelled β-carotene 153
2. Measuring the repletion doses of β-carotene and vitamin A needed to reverse
vitamin A deficiency in vitamin A depleted adults 154
3. Measuring changes of serum vitamin A levels after feeding synthetic β-carotene
or food rich in provitamin A carotenoids 155
4. Measuring changes in body stores of vitamin A after feeding dietary provitamin A
carotenoids (paired DRD test) 155
5. Measuring intestinal absorption by analysis of postprandial chylomicron fractions
after feeding synthetic β-carotene or food rich in provitamin A carotenoids 156
6. Measuring blood response kinetics after feeding β-carotene labelled with stable
isotopes 157
a)Single dose 157
b) Multiple doses 158

c) Use of labelled retinyl acetate as a reference 158
7. Feeding intrinsically labelled dietary provitamin A carotenoids in food 161
8. Conversion factors of β-carotene into retinol in humans: Summary 163
D. Factors that Affect the Bioabsorption and Conversion in vivo 165
1. Vitamin A status 165
2. Food matrix 165
3. Food preparation 166
4. Other carotenoids 166
5. Protein malnutrition 167
6. Intraluminal infections 167
7. Fat and fibre 167
E. Conversion in Tissues other than Intestine 168
F. Vitamin A Value of α
-Carotene and (cis)-β-Carotenes 168
G. Formation of Retinoic Acid from β-Carotene 168
H. Conclusion 169
References 169





xiv
Chapter 9: Vitamin A and Vitamin A Deficiency
George Britton

A. Introduction 173
B. Vitamin A 174
1. Basic biochemistry 174
2. Vitamin A status and requirements 176

3. Hypervitaminosis A: toxicity 176
C. Consequences of Vitamin A Deficiency 177
1. X erophthalmia 177
2. K eratinization 178
a) Eye tissues 178
b) Other epithelial tissues 178
3. Subclinical, systemic effects 179
a) Measles 179
b) Diarrhoea/dysentry 180
c) Respiratory infections 180
d) HIV and AIDS 180
e) Other infections 180
f) Immune response 180
E. Scale of Vitamin A Deficiency 181
1. Global distribution 181
2. Contributing factors 181
a) Age 181
b) Socioeconomic status 182
c) Seasonality 182
F. Strategies to Combat VAD 182
1. Supplements 183
a) Vitamin A 183
b) Provitamin carotenoids 183
2. Fortific ation 184
3. Dietary improvement 184
a) Home gardens 184
b) ‘Biofortification’ 184
c) Post-harvest treatment 185
4. Strategy overall 185
G. Underlying Causes 186

H. Conclusions 187
1. Place for carotenoid research 187
2. Political, educational, cultural 188
References 188
xv
Chapter 10: Epidemiology and Intervention Trials
Susan T. Mayne, Margaret E. Wright and Brenda Cartmel

A. Introduction to Epidemiology 191
B. Types of Epidemiological Studies 192
1. Observational study designs 192
a) Descriptive epidemiology 192
b) Analytical epidemiology 195
2. Intervention trials 196
a) Supplementation trials 196
b) Food-based interventions 197
3. Exposure assessment in epidemiological studies 198
a) Dietary assessment 198
b) Biomarker assessment 201
c) Assessment of multiple antioxidant nutrients: Antioxidant indices 203
C. Interpretation of Diet-Disease Associations Relevant to Carotenoids 203
1. Interpreting results of observational studies with carotenoid-containing foods 203
2. Interpreting results of intervention trials with carotenoid-containing foods 204
3. Interpreting results of carotenoid supplementation trials 205
4. Interpreting results of trials with intermediate endpoints 206
D. Future Directions 207
References 208


Chapter 11: Modulation of Intracellular Signalling Pathways by

Carotenoids

Paola Palozza, Simona Serini, Maria Ameruso and Sara Verdecchia

A. Introduction 211
B. Intercellular Communication and Signalling 212
1. Cell signalling pathways and mechanisms 213
2. Gap junction communication 213
3. The cell cycle and apoptosis 213
a) The cell cycle 213
b) Apoptosis 215
4. Reactive oxygen species as second messengers 215
5. Carotenoids as redox agents 216
C. Effects of Carotenoids on Cell Signalling and Communication 216
1. Modulation of cell cycle 216
2. Modulation of apoptosis 217
3. Modulation of the cell cycle and apoptosis via redox-sensitive proteins 218
xvi
4. Modulation of growth factors 220
5. Modulation of cell differentiation 221
6. Modulation of retinoid receptors 222
7. Redox-related modulation of transcription factors 223
a) NF-
κ
B 223
b) AP-1 224
c) Nrf2 and phase II enzymes 224
8. Modulation of hormone action 225
9. Modulation of peroxisome-proliferator activated receptors 226
10. Modulation of xenobiotic and other orphan nuclear receptors 227

11. Modulation of adhesion molecules and cytokines 228
12. Modulation of gap junction communication 229
D. Towards a Better Understanding of the Regulation of Cell Signalling
by Carotenoids 229
1. Delivery of carotenoids to cell cultures 229
2. Understanding effects and identifying biomarkers 230
References 230


Chapter 12: Antioxidant/Pro-oxidant Actions of Carotenoids
Kyung-Jin Yeum, Giancarlo Aldini, Robert M. Russell and Norman I. Krinsky

A. Introduction 235
B. Analytical Methods to Determine Antioxidant/Pro-oxidant Actions of
Carotenoids in Biological Samples 237
1. Total antioxidant capacity 237
2. Lipid peroxidation 241
3. Oxidation of low-density lipoprotein (LDL) 242
4. DNA damage 243
5. Other assays for biomarkers 243
a) Pulse radiolysis 243
b) HPLC/mass spectrometry 244
C. Studies of Antioxidant/Pro-oxidant Actions of Carotenoids 245
1. Studies in vitro 245
2. Studies ex vivo 247
3. Studies in vivo 248
D. Factors that Affect Antioxidant/Pro-oxidant Actions of Carotenoids 251
1. Concentration of carotenoids 252
2. Oxygen tension 253
3. Exposure to ultraviolet light 254

4. Oxidative stress 255
5. Interaction with membranes 25
xvii
6. Up-regulation of the receptor for advanced glycation endproducts (RAGE) 25
E. Interactions of Carotenoids 257
1. Interactions between carotenoids 257
2. Interactions of carotenoids with other antioxidants 258
F. Conclusions: Possible Biological Relevance of Antioxidant/Pro-oxidant Actions
of Carotenoids 259
References 262


Chapter 13: Carotenoids and Cancer
Cheryl L. Rock

A. Introduction 269
B. Lung Cancer 272
C. Breast and Ovarian Cancers 274
1. Breast cancer 274
2. Ovarian cancer 276
D. Prostate Cancer 276
E. Colorectal Cancer 278
F. Other Cancers 279
1. Cancer of the oral cavity, pharynx, and larynx (head and neck) 279
2. Cervical cancer 280
3. Other clinical trials with cancer outcomes 281
G. Conclusions 282
References 282

Chapter 14: Carotenoids and Coronary Heart Disease
Elizabeth J. Johnson and Norman I. Krinsky

A. Introduction 287
B. Observational Epidemiology 288
1. Case-control studies 288
2. Cohort studies 291
C. Randomized Control Trials 294
1. Carotenoids in the primary prevention of CHD 294
2. Carotenoids in the secondary prevention of CHD 295
3. Intervention trials and CHD biomarkers 296
D. Summary and Conclusions 297
References 298

xviii
Chapter 15: The Eye
Wolfgang Schalch, John T. Landrum and Richard A. Bone

A. Introduction 301
B. Anatomy of the Eye and Retina 302
C. Occurrence of Carotenoids in the Eye 304
1. Retina 304
2. Lens 307
3. Ciliary body and retinal pigment epithelium 307
D. The Macular Xanthophylls 307
E. ‘Classical’ Features of the Macular Pigment 310
1. General 310
a) Maxwell’s spot 310
b) Haidinger’s brushes 310
2. Effects of macular pigment on visual performance 312

a) Visual acuity and contrast sensitivity 312
b) Glare sensitivity and light scatter 313
F. Macular Pigment Optical Density (MPOD) and its Measurement 314
1. Analysis of carotenoids in retina and lens in vitro 314
2. Non-invasive determination of carotenoids in the retina in vivo 314
a) Quantitative estimation by psychophysical methods 314
b) Quantitative determination by physical methods 316
G. The Determinants of Macular Pigment Optical Density 317
1. Transport of carotenoids into the retina 317
2. D iet 318
3. Supple mentation 318
4. Other factors 320
H. The Role of Carotenoids in Risk Reduction of Macular Degeneration and Cataract
322
1. Mechanistic Basis 322
a) Absorption of blue light 322
b) Protection against photooxidation 322
c) Other properties 324
2. Evidence obtained from experiments with animals 324
3. Investigations in humans 325
a) Observational studies 325
b) Epidemiological studies 326
c) Supplementation studies (intervention trials) 327
I. Conclusions 328
References 330

xix
Chapter 16: Skin Photoprotection by Carotenoids
Regina Goralczyk and Karin Wertz

A. Introduction 335
B. Uptake and Metabolism of Carotenoids in Skin Cells 338
1. Humans and mouse models 338
2. Carotenoids in skin cell models 342
a) Culture conditions 342
b) Uptake and metabolism of carotenoids in skin cells 343
C. Photoprotection in vivo 345
1. Photosensitivity disorders 345
2. Photoc arcinogenesis 345
3. Sunburn 347
4. Photoa ging 348
5. Photoimmune modulation 350
D. Mechanistic Aspects of Photoprotection by Carotenoids 351
1. Inhibition of lipid peroxidation 351
2. Inhibition of UVA-induced expression of haem oxygenase 1 352
3. Prevention of mitochondrial DNA deletions 354
4. Metalloprotease inhibition 355
5. Use of microarray analysis to profile gene expression 357
E. Summary and Conclusion 359
References 359


Chapter 17: The Immune System
Boon P. Chew and Jean Soon Park

A. The Immune System and Disease 363
1. Introduction 363
2. Features of the immune system 364
a) The innate or antigen-non-specific immune system 364
b) The adaptive or antigen-specific specific immunity 365

c) Cell-mediated immune response 365
d) The humoural immune response 366
3. Nutritional intervention 366
4. Immunity and oxidative stress 366
B. Carotenoids and the Immune Response 367
1. Effects of carotenoids 367
a) Specific effects 3
b) Effects of carotenoid-rich foods and extracts 371
c) Model studies of health benefits 372
xx
C. Carotenoids and Disease 373
1. Age-related diseases 373
a) Age-related immunity decline 373
b) Neurodegenerative conditions 374
c) Rheumatoid arthritis 374
2. C ancer 374
3. Human immunodeficiency: HIV and AIDS 375
D. Mechanism of Action 376
E. Summary and Conclusions 378
References 379


Chapter 18: Biological Activities of Carotenoid Metabolites
Xiang-Dong Wang

A.Introduction 383
B. Carotenoid Metabolites 385
1. Enzymic central cleavage in vitro 385
a)
β

-Carotene 15,15
’
-oxygenase (BCO1) 385
b) Central cleavage of lycopene 386
2. Excentric enzymic cleavage in vitro 386
a)
β
-Carotene 9,10-oxygenase (BCO2) 386
b) Excentric cleavage of lycopene 387
3. Non-enzymic oxidative breakdown 388
4. Detection of central and excentric cleavage products in vivo 388
a) Metabolites of
β
-carotene 388
b) Metabolites of lycopene 389
C. Retinoids and the Retinoid Signalling Pathway 390
1. Retinoic acid and retinoic acid receptors 390
2. Effects of provitamin A carotenoids and their metabolites 391
a)
β
-Carotene and 14
’
-apo-
β
-caroten-14
’
-oic acid 391
b) Other provitamin A carotenoids 393
3. Effects of lycopene and its metabolites 393
a) Acycloretinoic acid 393

b) Other lycopene metabolites 394
c) Retinoid-dependent and retinoid-independent roles of carotenoid metabolites 395
D. Effects of Carotenoid Metabolites on Other Signalling and Communication Pathways
396
1. Nuclear factor-E2 related factor 2 (Nrf2) signalling pathway 396
a) Phase II enzymes and antioxidant-response elements 396
b) Effects of carotenoids and their metabolites 396
c) Lycopene metabolites 397
xxi
2. Carotenoid metabolites and the mitogen-activated protein kinase pathway 398
a)
β
-Carotene and metabolites 399
b) Lycopene and metabolites 399
3. Carotenoid metabolites and the insulin-like growth factor-1 (IGF-1) pathway 400
4. Carotenoid metabolites and gap-junction communication 401
E. Overview and Conclusions 402
References 404


Chapter 19: Editors’ Assessment
George Britton, Synnøve Liaaen-Jensen and Hanspeter Pfander

A. Introduction 409
B. From Food to Tissues 410
1. Sources, bioavailability and conversion 410
2. Variability between individuals 411
C. Carotenoids and Major Diseases: Practical Concerns and General Points 412
1. Human studies 412
a) Are effects due to carotenoids or to food? 413

b) Biomarkers 414
2. Cell cultures 414
3. Animal models 414
4. High dose, low dose and balance 415
5. Safety and toxicity 415
6. Geometrical isomers 416
7. Natural versus synthetic 417
D. How Might the Effects be Mediated? 418
1. Via antioxidant action 418
2. Via m etabolites 419
3. Via the immune system 419
E. Reports of Other Health Effects 419
1. Water-soluble carotenoids 420
2. Bone health 420
3. Metabolism and mitochondria 420
F. Final Comments: The Big Questions 421
References 422


Index 423
Postscript 431
xxii
Contents of Carotenoids Volume 4: Natural Functions

Chapter 1: Special Molecules, Special Properties
Chapter 2: Structure and Chirality
Chapter 3: E/Z Isomers and Isomerization
Chapter 4: Three-dimensional Structures of Carotenoids by X-ray Crystallography
Chapter 5: Aggregation and Interface Behaviour of Carotenoids
Chapter 6: Carotenoid-Protein Interactions

Chapter 7: Carotenoid Radicals and Radical Ions
Chapter 8: Structure and Properties of Carotenoid Cations
Chapter 9: Excited Electronic States, Photochemistry and Photophysics of
Carotenoids
Chapter 10: Functions of Intact Carotenoids
Chapter 11: Signal Functions of Carotenoid Colouration
Chapter 12: Carotenoids in Aquaculture: Fish and Crustaceans
Chapter 13: Xanthophylls in Poultry Feeding
Chapter 14: Carotenoids in Photosynthesis
Chapter 15: Functions of Carotenoid Metabolites and Breakdown Products
Chapter 16: Cleavage of β
-Carotene to Retinal
Chapter 17: Enzymic Pathways for Formation of Carotenoid Cleavage products
xxiii
List of Contributors
Giancarlo Aldini
Department of Pharmaceutical Sciences
Faculty of Pharmacy
University of Milan
Via Mangiagalli 25
20133 Milan
Italy
()

Maria Ameruso
Institute of General Pathology
Catholic University
School of Medicine
L. go F. Vito 1
00168 Rome

Italy

Richard A. Bone
Department of Physics
Florida International University
Miami
FL 33199
U.S.A.
()

Peter M. Bramley
Centre for Systems and Synthetic Biology
School of Biological Sciences
Royal Holloway University of London
Egham Hill
Egham
Surrey
TW20 0EX
U.K.
()


George Britton
53 Forest Road
Meols
Wirral
CH47 6AT
U.K.
()

Kirstie Canene-Adams
Department of Pathology
Johns Hopkins School of Medicine
The Bunting-Blaustein Cancer Research
Building
1650 Orleans Street
Baltimore
MD 21231-1000
U.S.A.
()

Brenda Cartmel
Yale School of Public Health
60 College Street
P. O. Box 208034
New Haven
CT 06520-8034
U.S.A.
()

Boon P. Chew
FSHN 110
School of Food Science
Washington State University
Pullman
WA 99164-6376
U.S.A.
()

xxiv

Laurent Dufossé
Université de la Réunion
ESIDAI
LCSNSA
Parc Technologique
2 Rue Joseph Wetzell
F-97490 Sainte-Clotilde
La Réunion
France
()

John W. Erdman Jr.
Department of Food Science and Human
Nutrition
University of Illinois,
451 Bevier Hall
905 S. Goodwin Avenue
Urbana
IL 61801
U.S.A.
()

Paul D. Fraser
Centre for Systems and Synthetic Biology
School of Biological Sciences
Royal Holloway University of London
Egham Hill
Egham
Surrey
TW20 0EX

U.K.
()

Regina Goralczyk
DSM Nutritional Products Ltd.
P.O. Box 2676
CH-4002 Basel
Switzerland
()

Elizabeth J. Johnson
Carotenoids and Health Laboratory
Jean Mayer USDA Human Nutrition Center
on Aging
Tufts University
711 Washington Street
Boston
MA 02111
U.S.A.
()
Norman I. Krinsky
Jean Mayer USDA Human Nutrition
Research Center on Aging
Department of Biochemistry
School of Medicine
Tufts University
711 Washington Street
Boston
MA 02111
U.S.A.

John T. Landrum
Department of Chemistry and Biochemistry
Florida International University
Miami
FL 33199
U.S.A.
()

Synnøve Liaaen-Jensen
Organic Chemistry Laboratories
Department of Chemistry
Norwegian University of Science and
Technology (NTNU)
7491 Trondheim
Norway
()

Susan T. Mayne
Yale School of Public Heath
60 College Street
P.O. Box 208034
New Haven
CT 06520-8034
U.S.A.
()

Alan Mortensen
Product Development
Color Division

Chr. Hansen
Bøge Allé 10-12
DK-2970 Hørsholm
Denmark
()





xxv
Paola Palozza
Institute of General Pathology
Catholic University
School of Medicine
L. go F. Vito 1
00168 Rome
Italy
()

Jean Soon Park
P & G Pet Care
Upstream R & D
6571 State Route 503 North
P.O. Box 189
Lewisburg
OH 45338
U.S.A.
()

Hanspeter Pfander
Muristrasse 8E
CH-3006 Bern
Switzerland
()

Cheryl L. Rock
Moores USCD Cancer Center
University of California San Diego
3855 Health Sciences Drive
La Jolla
CA 92093-0901
U.S.A.
()

Robert M. Russell
Office of Director
National Institutes of Health
Washington DC,
U.S.A.
()

Wolfgang Schalch
DSM Nutritional Products Ltd.
Wurmisweg 576
CH-4303 Kaiseraugst
Switzerland
()

Simona Serini

Institute of General Pathology
Catholic University
School of Medicine
L. go F. Vito 1
00168 Rome
Italy

Guangwen Tang
Carotenoids and Health Laboratory
Jean Mayer USDA Human Nutrition
Research Center on Aging
Tufts University
711 Washington Street
Boston
MA 02111
U.S.A.
()

Sara Verdecchia
Institute of General Pathology
Catholic University
School of Medicine
L. go F. Vito 1
00168 Rome
Italy

Xiang-Dong Wang
Nutrition and Cancer Biology Laboratory
Jean Mayer USDA Human Nutrition
Research Center on Aging

Tufts University
711 Washington Street
Boston
MA 02111
U.S.A.
()

Karin Wertz
DSM Nutritional Products Ltd.
P.O. Box 2676
CH-4002 Basel
Switzerland
()





xxvi
Margaret E. Wright
Department of Pathology
College of Medicine
University of Illinois at Chicago
840 South Wood Street
Chicago
IL 60612
U.S.A.
()

Kyung-Jin Yeum

Jean Mayer USDA Human Nutrition
Research Center on Aging
Tufts University
711 Washington Street
Boston
MA 02111
U.S.A.
()
xxvii
Preface
More than twenty years after the idea of this Carotenoids book series was first discussed, we
finally reach the end of the project with Volume 5, which covers the functions and actions of
carotenoids in human nutrition and health. In 1971, in Isler’s book Carotenoids, functions of
carotenoids and vitamin A were covered in just two chapters. Now, thanks to technical
developments and multidisciplinary approaches that make it possible to study functional
processes in great detail, this subject is the most rapidly expanding area of carotenoid research,
and occupies two full volumes, Volumes 4: Natural Functions and 5: Nutrition and Health.
Although Volume 5 can be used as a single stand-alone volume, the two were planned as
companion volumes to be used together. To understand the mechanisms of functions and
actions of carotenoids, including how carotenoids may be involved in maintaining human
health, requires understanding of the underlying principles, which are presented in the first
part of Volume 4.
The general philosophy and strategy of the series, to have expert authors review and
analyse critically a particular topic and present information and give guidance on practical
strategies and procedures is maintained in Volumes 4 and 5. It is also the aim that these
publications should be useful for both experienced carotenoid researchers and newcomers to
the field.
The material presented in the earlier volumes of the series is relevant to studies of
biological functions and actions. Biological studies must be supported by a rigorous analytical
base and carotenoids must be identified unequivocally. It is a common view that carotenoids

are difficult to work with. This may be daunting to newcomers to the field, especially if they
do not have a strong background in chemistry and analysis. There are difficulties; carotenoids
are less stable than most natural products, but ways to overcome the difficulties and to handle
these challenging compounds are well established and are described and discussed in Volume
1A which, together with Volume 1B, gives a comprehensive treatment of the isolation,
analysis and spectroscopic characterization of carotenoids as an essential foundation for all
carotenoid work. This is complemented by the Carotenoids Handbook (2004), which was
produced in association with this series and provides key analytical data for each of the 750 or
so known naturally-occurring carotenoids.
Volume 2 describes methods for the chemical synthesis of carotenoids, including those that
are needed as analytical standards and on a larger scale for biological trials. Functions and
actions are inextricably linked with biosynthesis and metabolism, covered in Volume 3.