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Antioxidants in plants
Rio December 2007
Antioxidants in plants
Institute of Plant Biotechnology
For Developing Countries
Rio, December 4, 2007
Mike May
IPBO
Website: http://www.ipbo.ugent.behttp://www.psb.ugent.be
http://www.efb-central.orghttp://www.pubresreg.org
E-mail : [email protected]
http://www.ipbo.ugent.be/http://www.psb.ugent.be/http://www.efb-central.org/http://www.pubresreg.org/http://www.pubresreg.org/http://www.efb-central.org/http://www.psb.ugent.be/http://www.ipbo.ugent.be/8/2/2019 Antioxidants in Plants
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Antioxidants in plants
Rio December 2007
Overview of seminar
Redox homeostasis:
Understanding functionthrough genetics:
Understanding thefunction of ascorbate:
Interactions with glutathioneand tocopherol:
Applications and future focus:
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Rio December 2007
New approaches for functional
analysis and crop improvement
1904, Agrobacteriumtumefaciens
1974, TiPlasmid, Gent
1990sgenomics
Overexpression& silencing
http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%26no=2%26tt=385%26oid=2be3f4eef1bb27cc%26ei=UTF-8http://rds.yahoo.com/_ylt=A9iby4GL3lJF7QcBxFaJzbkF;_ylu=X3oDMTBjdmNoOTVjBHBvcwMyBHNlYwNzcg--/SIG=1gd463pfr/EXP=1163145227/**http%3a//images.search.yahoo.com/search/images/view%3fback=http%253A%252F%252Fimages.search.yahoo.com%252Fsearch%252Fimages%253Fp%253DCROWN%252BGALL%2526ei%253DUTF-8%2526fr%253Dsfp%2526x%253Dwrt%26w=180%26h=143%26imgurl=soilplantlab.missouri.edu%252Fplant%252Fimages%252Fcrowngalleuonymus.jpg%26rurl=http%253A%252F%252Fsoilplantlab.missouri.edu%252Fplant%252Fdiseases%252Fcrowngall.htm%26size=12.9kB%26name=crowngalleuonymus.jpg%26p=CROWN%2bGALL%26type=jpeg%2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Antioxidants in Plants
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Antioxidants in plants
Rio December 2007
Redox homeostasis:
Reactive oxygen species
Antioxidants
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Antioxidants in plants
Rio December 2007
Aerobic life is based on the reduction of oxygen
via electron transfer systems: production in chloroplasts; consumption in mitochondria
This is an inefficient process; And results in the continuous production of
reactive oxygen species (ROS):
Hydrogen peroxide; Superoxide;
Hydroxyl radical
Reactive oxygen species
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Nucleus(not a source,
but ROS-mediated damageto DNA results inmutation)
Mitochondrion(ATP synthesis)
Chloroplast (water splitting, electrontransport)
PeroxisomeEndogenous sources of ROS in plants
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Pathogens(oxidative burst)
Mitochondrion(Herbicides, ozone)
Chloroplast (herbicides, highlight)
Drought, waterloggingpollutants, salinity
Stress related sources of ROS in plants
Temperatureextremes
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Unless ROS are removed from biological
systems, they cause damage to: Lipids (leakage of solutes);
Proteins (loss of function, aggregation);
Nucleic acids (mutation, nicks)
The evolution of aerobic life forms has gonehand in hand with the evolution of highlyconserved mechanisms for ROS removal andsensing of ROS.
ROS - consequences
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ROS-mediated chloroplast damage (rice)
Methyl
viologen
Salt
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Enzymatic processes: Superoxide dismutases peroxidases catalase thioredoxin/glutaredoxin
Major antioxidant compounds/cofactors: NAD(P)H ascorbate
tocopherols (vitamin E) glutathione pigments
polysaccharides
ROS - Removal
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ROS are consequently implicated in a wide variety ofdisorders:
In plants: photorespiratory damage; pathogen attack; Biotic and abiotic stresses
In animals: inflammatory disorders immune system Cancer Degenerative disorders such as Parkinsons and Alzheimer's genetic disorders eg Lou Gehrig syndrome, Downs syndrome
The study of ROS is therefore one of the most vibrant
areas of research in plant and animal science:
ROS pathology
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Fundamental: biotic and abiotic stresses; development; metabolic control; organelle dialogue;
Signalling, transcription factors, protein complexes Applied:
implication in human health (vitamin C/E);
functional foods; nutraceuticals; development of crops for climatic extremes
new growth regulators and herbicides
Study of ROS in plants
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The Ascorbate-Glutathione cycle
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Redox homeostasis
Diagram courtesyOf Christine Foyer
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Developmental control of synthesis
Environmental control of synthesis
Understanding the
function of ascorbate:
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Development-dependent and oxidative
stress-dependent Protein Carbonylformation
Source; Christine
Foyer, University ofNewcastle
A i id i l
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Ascorbate synthesis and accumulationdeclines with age: Maize
GLDH = Galactonolactone dehydrogenase, the lastenzyme in the biosynthetic pathway
Source; ChristineFoyer, University ofNewcastle
A ti id t i l t
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Ascorbate accumulates under high light inArabidopsis
LL IL HL
Source; ChristineFoyer, University of
Newcastle
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Antioxidants in plants
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Understanding function
through genetics:
vtcmutants of Arabidopsis
Pathogen-mediated signalling interactions with glutathione and tocopherol
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Two pathways for biosynthesis
Source; Wolucka &Van Montagu.Phytochemistry2007
L-Galactose pathway
L-Gulose pathway
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vtc 1
vtc 2/5
vtc 4
Source: Linster et al.,(2007) JBC 282, 18879-18885)
Mapping of Arabidopsismutants on the
L-galactose pathway
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Plants:
Galactolactonedehydrogenase(GLDH)
Animals:Gulonolactoneoxidase
(GLO)
Evolutionary divergence of pathways
Glucose 6-P
AscorbateHumans,higher primates
vtc 1
vtc 2/5
vtc 4
Gulose pathway
Galactose pathway
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Analysis of vtc1 & vtc2to study the rolesof ascorbate in plants
vtcmutants wereisolated in a screenfor ozone sensitivity
Source; ChristineFoyer, University ofNewcastle
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vtc1 mutants mature slower than wild type
Source; ChristineFoyer, University ofNewcastle
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Low ascorbate in the vtc1 mutants:Enhances resistance to biotrophic pathogens
vtc1
Wildtype
Source; ChristineFoyer, University ofNewcastle
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t o da ts p a ts
Rio December 2007
Low ascorbate in the vtc1 mutants:Creates a lesion-mimic phenotype
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p
Rio December 2007
Low ascorbate in the vtc1 mutants:Altered SAR signalling
NRP1 NRP1
NRP1 NRP1
Salicylic Acid
Pathogen
H2O2
Phytoalexins
NRP1 NRP1
NRP1 NRP1
GSH
NRP1SH
SH
WRKY gene expression
PR proteinsVtc1 flux
WT flux
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p
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Interactions with
glutathione and tocopherols:
Pathogen-mediated signalling
root development & cell cycle
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Biosynthesis of tocopherol
Porfivora et al(2002)PNAS 99, 12495-12500
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Tocopherol null mutants are not highly
stress-sensitive
Porfivora et al(2002)PNAS 99, 12495-12500
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Ascorbate, GSH and tocopherol
compensate for each other
Porfivora et al(2002)PNAS 99, 12495-12500
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Biosynthesis of GSH
Glu + Cys Glu-Cys Glu-Cys-Gly
Gly
GSH-1 GSH-2
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cad-2:(Heavy metal screen) cadmium sensitive
otherwise normal 15-30% GSH levels
pad-1:(pathogen screen) phytoalexin deficient
pathogen sensitive
otherwise normal 22% GSH levels
rml-1:(root meristem screen) root meristemless
GSH auxotroph almost undetectable GSH
gsh-1 mutants of Arabidopsis
demonstrate the importance of GSH
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GSH biosynthesis is stimulated in response topathogen attack;
pad-1 mutants are sensitive to pathogen attack
and do not accumulate phytoalexinscad-2is more resistant than pad-1
Lowering ascorbate enhances resistancewhilst lowering GSH enhances susceptibility
GSH and pathogen signalling
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NRP1 NRP1
NRP1 NRP1
Salicylic Acid
Pathogen
H2O2
Phytoalexins
NRP1 NRP1
NRP1 NRP1
GSH
NRP1SH
SH
WRKY gene expression
PR proteins
GSH and pathogen signalling
pad-1
Biosynthesis
X
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GSH and the cell cycle
Oxidative
Stress (e.g. P53)
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Quiescent
centre (G0)
Distribution of GSH in Arabidopsisroots
Root Hair
Source: MikeMay, Gent
University
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Role of GSH in cell cycle progress
in synchronised By-2 cell suspensions
GSH is needed for the G1 S transition
+ GSH
control
+ BSO
GSHlevels
DNASynthesis(S-phase)
Cell cyclephase
Source: MikeMay, GentUniversity
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Role of GSH in root but not shoot cell
cyclerml-1
rml-1+ GSH
Wild Type
Source: MikeMay, Gent
University
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Applications and future
focus:
Functional/super foods
Nutraceuticals Metabolomics
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Fruit or vegetable Ascorbic acid content
(mg per 100 g)
Rosehip 1000
Blackcurrant 200Kale 186Green pepper 128Horse radish 120Broccoli 113
Brussels sprouts 109Watercress 79Strawberry 59Spinach 51Oranges/lemons 50
Cabbage 47New potato 30Carrot 6
A. thalianasuspension cells 8
BY-2 suspension cells 0.2-2
Vitamin C content of some common foods
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Emblica officinalis: Amla or Indian Gooseberry
Euphorbiaceae;
Used in Ayurveda for over
2000 years; A component of a patented
formula for treatment of asthmaand allergic rhinitis in the USA;
Vitamin C (700mg per 100gfruit), flavonoids, polyphenols,alkaloids;
Immune system, heart, chroniclung problems, liver, circulation;
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Lycium chinense: Goji Berry
Solanaceae;
Used in TCM for over 2000
years 54 new product introductions in
2006
China produced 95,000 tonnesin 2004
$120 million in exports
Vitamin C (163% DRI),
polysaccharides, zeaxanthin,butyric acid
Immune system, eyesight, liver,circulation
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Acai berry; the amazonianwonderfruit