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Peter F. Surai
Selenium in
poultry nutrition
and health
Wageningen Academic
Publishers
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Selenium in poultry nutrition and health
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Selenium
in poultry nutrition
and health
Peter F. Surai
Wageningen Academic
Publishers
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EAN: 9789086863174
e-EAN: 9789086868650
ISBN: 978-90-8686-317-4
e-ISBN: 978-90-8686-865-0
DOI: 10.3920/978-90-8686-865-0
First published, 2018
© Wageningen Academic Publishers
e Netherlands, 2018
This work is subject to copyright. All
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Dedication
To my wife Helen, my daughter Katie, my son Anton,
my grandsons Oscar, Arthur and Henry
and my granddaughter Aiste
who gave me inspiration for writing this book.
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Selenium in poultry nutrition and health 7
Abouttheauthor
Dr. Peter Surai started his studies at Kharkov
University, Ukraine, where he obtained his
PhD and DSc in biochemistry studying
eects of antioxidants on poultry. Later he
became Professor of Human Physiology. In
1994 he moved to Scotland to continue his
antioxidant related research in poultry and
in 2000 he was promoted to a full Professor
of Nutritional Biochemistry at the Scottish
Agricultural College. Recently he was
awarded Honorary Professorships in 5
universities in various countries, including
UK, Hungary, Bulgaria and Ukraine. In 2010 he was elected to the Russian Academy of
Sciences as a foreign member. He has more than 750 research publications, including
150 papers in peer-reviewed journals and 13 books. In 1999 he received the prestigious
John Logie Baird Award for Innovation for the development of ‘super-eggs’ and, in
2000, e World’s Poultry Science Association Award for Research in recognition of
an outstanding contribution to the development of the poultry industry. In 2017 he
became a member of the team at the Moscow State Academy of Veterinary Medicine
and Biotechnology named aer K.I. Skryabin to conduct a research under a mega-
grant of the Government of Russian Federation (Contract No. 14.W03.31.0013).
For the last 15 years he has been lecturing all over the world visiting more than 70
countries.
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Selenium in poultry nutrition and health 9
Tableofcontents
Abouttheauthor 7
Preface 13
Abbreviations 15
Chapter 1
Antioxidant systems in animal body 19
1.1 Introduction 19
1.2 Free radicals and reactive oxygen and nitrogen species 19
1.3 ree levels of antioxidant defence 23
1.4 Superoxide dismutase in biological systems 26
1.5 Superoxide dismutase in avian biology 29
1.6 Other antioxidant mechanisms 30
1.7 Oxidative stress and transcription factors 46
1.8 Vitagene concept development 50
1.9 Conclusions 53
References 54
Chapter 2
Molecular mechanisms of selenium action: selenoproteins 67
2.1 Introduction 67
2.2 e selenoprotein family 67
2.3 Selenocysteine: the functional selenium 68
2.4 Glutathione peroxidases 70
2.5 Glutathione peroxidase activity eectors 83
2.6 GSH-Px and their biological roles 91
2.7 ioredoxin reductases as a major part of the thioredoxin system 93
2.8 Iodothyronine deiodinases 99
2.9 Other selenoproteins 103
2.10 General conclusions 119
References 122
Chapter 3
Selenium in feed: organic selenium concept 153
3.1 Introduction 153
3.2 Selenium in soils and plants 153
3.3 Selenium absorption and metabolism 160
3.4 Selenium status and bioavailability 169
3.5 Eectors of selenium absorption, metabolism and bioavailability 171
3.6 Selenium sources for poultry 172
3.7 Selenium-enriched yeast: pluses and minuses 175
3.8 SeMet and OH-SeMet 178
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10 Selenium in poultry nutrition and health
Table of contents
3.9 Chelated Se products 180
3.10 Nano-Se products 181
3.11 Conclusions 182
References 184
Chapter 4
Selenium deciency in poultry 195
4.1 Introduction 195
4.2 Exudative diathesis 196
4.3 Nutritional pancreatic atrophy 198
4.4 Nutritional encephalomalacia 199
4.5 Nutritional muscular dystrophy 205
4.6 Impaired immunocompetence 209
4.7 Impaired thyroid hormone metabolism 209
4.8 Reduced fertility 209
4.9 Reduced egg production and quality 209
4.10 Decreased hatchability and increased embryonic mortality 210
4.11 Conclusions 210
References 211
Chapter 5
Selenium in poultry nutrition 219
5.1 Introduction 219
5.2 Selenium for breeders 219
5.3 Selenium for commercial layers 240
5.4 Selenium for broilers 249
5.5 Conclusions 262
References 264
Сhapter 6
Selenium-enriched eggs and meat 279
6.1 Introduction 279
6.2 Selenium and human health 279
6.3 Strategies to deal with Se deciency in human diet 284
6.4 Addressing Se deciency in humans via Se-enriched eggs 287
6.5 Se-enriched eggs in a global context 293
6.6 Safety of Se-enriched eggs 293
6.7 Se-enriched meat 294
6.8 Optimal selenium forms in the diets for Se-egg and Se-meat production 296
6.9 Se-enriched eggs and meat as functional food 297
6.10 Conclusions 300
References 301
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Selenium in poultry nutrition and health 11
Table of contents
Chapter 7
Selenium and immunity 309
7.1 Introduction 309
7.2 Immune system and its evaluation 310
7.3 Phagocyte functions 322
7.4 Antibody production 325
7.5 Lymphocyte functions 327
7.6 In vitro eects of selenium on immune cells 331
7.7 Disease resistance 334
7.8 Immunoprotective eects of Se in stress conditions 335
7.9 Molecular mechanisms of immunomodulating properties of selenium 342
7.10 Immunocommunication, free radicals and selenium 346
7.11 Conclusions 352
References 355
Chapter 8
Antioxidant-prooxidant balance in the gut 369
8.1 Introduction 369
8.2 e gastrointestinal tract as a major site of antioxidant action 369
8.3 Prooxidants in the gastrointestinal tract 371
8.4 Antioxidant defences in the gastrointestinal tract 376
8.5 Specic place for Se-dependent enzymes in antioxidant defence of the
gastrointestinal tract 381
8.6 Role of vitagenes in the gut defence 383
8.7 Critical periods of the gut development 387
8.8 Conclusions 394
References 395
Chapter 9
Looking ahead 411
References 422
Index 425
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Selenium in poultry nutrition and health 13
Preface
Among many minerals selenium has a special place being the most controversial trace
element. Indeed a narrow gap between essentiality and toxicity and environmental
issues on the one hand and global selenium deciency on the other hand, fuel research
in this eld. ere were several breakthroughs in selenium research. e rst one was
the discovery of Se essentiality in early 1960s. e second one was the discovery in
1973 that glutathione peroxidase is a selenoprotein. e third one came almost 30 years
later with characterisation of main selenoproteins in human and animal body and
further understanding the role of selenium in nutrition and health. Indeed, this third
breakthrough is really a selenium revolution creating many hypotheses, stimulating
new research and providing practical applications in medicine and agriculture. New
insight in the role of free radicals as signalling molecules, understanding the role of
nutrients in gene expression and maternal programming, tremendous progress in
human and animal genome work created new demands for further research related
to biological roles of selenium.
Several comprehensive monographs and reviews have been recently published
addressing various Se-related issues. However, most of them were dealing with Se
roles in human health. Animal food-producing industry is developing very quickly
and a great body of information was accumulated indicating importance of Se in
maintenance of animal health, productive and reproductive performance. Our
previous comprehensive book ‘Selenium in Nutrition and Health’ was published in
2006 and a lot of important Se-related information has been accumulated for the last
10 years. erefore, the goal of this volume is to provide up to date information about
the roles of Se in poultry nutrition and health. In Chapter 1 a special emphasis is given
to the role of selenium as an essential part of the integrated antioxidant system of the
body with regulatory functions providing necessary connections between dierent
antioxidants. In fact selenium is called ‘the chief executive of the antioxidant defence’.
Chapter 2 is addressing molecular mechanisms of Se action describing major functions
of the selenoproteins. Indeed, the family of selenoproteins includes 25 members and
functions of many of them are still not well understood. Selenium in feed is described
in Chapter 3. e main idea of this chapter is to describe an organic Se concept.
Indeed, in grains and some other important food ingredients selenomethionine is the
main Se form. e idea was put forward that during evolution the digestive system
of human and animals was adapted to natural form of selenium consisting of SeMet
and other organic selenocompounds. erefore, this form of Se is more eciently
assimilated in the body than inorganic forms of selenium. In fact SeMet is considered
to be the storage form of selenium in the body. Accumulation of the Se reserves in
the body as a result of organic selenium consumption is considered as an adaptive
mechanism providing additional antioxidant defences in stress conditions. e three
generations of Se supplements for poultry are characterised. Chapter 4 is devoted to
Se-deciency diseases in poultry with a specic emphasis to new data on the eect of
Se deciency on the expression of various selenoproteins in dierent chicken tissues.
Indeed, oxidative stress is considered to be a driving force in the development of such
Se-deciency diseases as encephalomalacia, exudative diathesis, nutritional muscular
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14 Selenium in poultry nutrition and health
Preface
dystrophy, nutritional pancreatic atrophy, impaired immunocompetence and
decreased productive and reproductive performance of chickens. e data presented
in Chapter 5 indicate importance of Se in growth, development and reproduction of
poultry. e main idea of the chapter is to show benets of various forms of organic
Se on antioxidant defences in the body leading to improvement of productive and
reproductive performance of poultry and poultry product quality. Indeed, organic
selenium is proven to be the most eective form of Se supplementation for poultry
and farm animals. Chapter 6 is devoted to the link between animal industry and
human health and describing some features of new technologies for production of
Se-enriched eggs and meat. In fact, production of a range of Se-enriched products
is considered as an important solution for global Se deciency. Se-enriched eggs are
already on supermarket shelves in many countries worldwide with millions of such
eggs sold daily. Chapter 7 is devoted to the role of selenium in immunity. It is dicult
to overestimate immunomodulating properties of selenium and increased resistance
to various diseases of poultry/animals is a result of optimal Se status. e possibility
of virus mutation in the body of animals decient in selenium is of great importance
for understanding mechanisms of spreading such diseases as chicken inuenza, etc.
e last chapter is devoted to the antioxidant-prooxidant balance in the digestive
tract. It seems likely that this balance has been overlooked by scientists. However, the
specic roles of selenoproteins in such a balance need further investigation. Indeed,
chicken health starts from its gut. I understand that my views on the role of selenium
in poultry nutrition and health are sometimes dierent from those of other scientists
and therefore I would appreciate very much receiving any comments from readers
which will help me in my future research. I would like to thank my colleagues with
whom I have had the pleasure to collaborate and share my ideas related to natural
antioxidants and selenium in particular, who helped me at various stages of this
research by providing reprints of their recent publications. I am also indebted to
the World’s Poultry Science Association for the Research Award and a grant of the
Government of Russian Federation (Contract No. 14.W03.31.0013) supporting my
research.
Peter F. Surai
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Selenium in poultry nutrition and health 15
Abbreviations
5-LO 5-lipoxigenase
9-oxoODE 9-oxo-octadecadienoic acid
AA ascorbic acid
Ab antibody
AEC abdominal exudate cells
AFB1 aatoxin B1
ALS amyotrophic lateral sclerosis
AO anti-oxidant
APR acute phase response
AvBD avian beta-defensin
BD basal diet
BHA butylated hydroxyanisole
BHT butylated hydroxytoluene
C/EBP CCAAT-enhancer-binding protein
CAT catalase
CDS coding sequence
CNS central nervous system
ConA concanavalin A
CoQ coenzyme Q
COX-2 cyclooxygenase-2
CVB3 Coxsackie virus B3
DAA dehydroascorbic acid
DC dendritic cells
DDT dichlorodiphenyltrichloroethane
DHA docosahexaenoic acid
DHT dihydrotestosterone
Dio iodothyronine deiodinase
DON deoxynivalenol
DTH delayed-type hypersensitivity
EC-SOD extracellular superoxide dismutase
ED exudative diathesis
EFX enrooxacin
ER endoplasmic reticulum
ERO1 endoplamic reticulum oxidoreductin 1
FAK focal adhesion kinase
FB1 fumonisin B1
FCR feed conversion ratio
FcγR phagocytic Fcγ receptors
FDA US Food and Drug Administration
FO sh oil
FT3 free triiodothyronine
FT4 free thyroxine
GI-GSH-Px gastrointestinal glutathione peroxidase
GIT gastrointestinal tract
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16 Selenium in poultry nutrition and health
Abbreviations
GR glutathione reductase
GSH reduced glutathione
GSH-Px glutathione peroxidase
GSSG oxidised glutathione
GST glutathione S-transferase
H/L ratio heterophil to lymphocyte ratio
H2O2 hydrogen peroxide
HETE 15-hydroxyeicosatetraenoic acid
HI hemaglutination inhibition
HMSeBA selenomethionine hydroxyanalogue, 2-hydroxy-4-methylselenobutanoic
acid
HO-1 haeme oxygenase-1
HPETE 15-hydroperoxyeicosatetraenoic acid
HS heat stress
Hsf1 heat shock factor 1
HSP heat shock proteins
IBD Infectious bursal disease
ID iodothyronine deiodinase
IELs intraepithelial lymphocytes
IFN interferon
Ig immunoglobulin
IL-1 interleukin 1
IL-2R interleukin 2 receptor
IL-6 interleukin 6
iNOS inducible nitric oxide synthase
IκB inhibitor of kappa B
Keap1 Kelch-like-ECH-associated protein 1
LA linoleic acid
LAK lymphokine-activated killer
LDH lactate dehydrogenase
LOOH lipid hydroperoxide
LOX lipoxygenase
LP lipid peroxidation
LPS lipopolysaccharide
LTA lymphocyte transformation assay
LXA4 lipoxin A4
MAPK mitogen-activated protein kinase
MCP-1 monocyte chemoattractant protein-1
MD Marek’s disease
MDA malondialdehyde
Met methionine
MHC major histocompatibility complex
MIF macrophage inammatory protein 2
MLTC mixed lymphocyte/tumour cell cultures
Msr methionine sulfoxide reductase
NE nutritional encephalomalacia
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Abbreviations
Selenium in poultry nutrition and health 17
NecE necrotic enteritis
NF-κB nuclear factor-kappa B
NK cells natural killer cells
NKT natural killer T cells
NMD nutritional muscular dystrophy
NO nitric oxide
NPA nutritional pancreatic atrophy
NRC National Research Council
Nrf2 NF-E2-related factor 2
OCP organochlorine pesticides
ONOO- peroxynitrite
OTA ochratoxin A
PAMP pathogen-associated molecular patterns
PCB polychlorinated biphenyls
PCV2 porcine circovirus type 2
PFC plaque-forming cell
PGE2 prostaglandin E2
pGSH-Px plasma glutathione peroxidase
PHA phytohemagglutin
PH-GSH-Px phospholipid glutathione peroxidase
PI3K phosphatidylinositol 3-kinase
PLA2 phospholipase A2
PMN polymorphonuclear leukocytes
POP persistent organic pollutants
PPAR peroxisome proliferator-activated receptor
PPRE peroxisome proliferator response element
PRR pattern recognition receptors
Prx peroxiredoxin
PTGE prostaglandin E
PUFA polyunsaturated fatty acid
PWM pokeweed mitogen
RDA recommended daily allowance
RNS reactive nitrogen species
RXR retinoid-X receptor
SBP2 SECIS-binding protein
SECIS selenocysteine insertion sequence
SeCys selenocysteine
SelN selenoprotein N
SelP selenoprotein P
SelP-L long-form selenoprotein P
SelR selenoprotein R
SeMet selenomethionine
SeS selenoprotein S
SeW selenoprotein W
SM silymarin
SO Selisseo, OH-SeMet
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18 Selenium in poultry nutrition and health
Abbreviations
SOD superoxide dismutase
SP selenoproteins
SPS selenophosphate synthetase-2
SRBC sheep red blood cells
SS sodium selenite
SSC spermatogonial stem cells
SY selenium enriched yeast
T testosterone
T3 triiodothyronine
T4 thyroxine
T-AOC total antioxidant capacity
TBA thiobarbituric acid
TBARS thiobarbituric acid reactive substances
TCR T-cell receptor
cells T helper cells
TLR Toll-like receptors
TNF-α tumour necrosis factor alpha
Toc tocopherol
Trx thioredoxin
TrxR thioredoxin reductase
TSH thyroid-stimulating hormone
vMDV virulent Marek’s disease virus
VSMCs vascular smooth muscle cells
ZEA zearalenone
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