Biochemistry of Cork Formation as a Stress Responsesaibo/ADONIS/talks/CPinto... · Biochemistry of...
Transcript of Biochemistry of Cork Formation as a Stress Responsesaibo/ADONIS/talks/CPinto... · Biochemistry of...
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Biochemistry ofCork Formation as a Stress Response
Cândido Pinto RicardoInês Chaves March 5th, 2009
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Moore et al. 1998 Botany WCB/McGraw-Hill Companies
Secondary growth of dicot stems
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Moore et al. 1998 Botany WCB/McGraw-Hill Companies
Secondary growth of dicot stems Axial section of cork oak tree
Silva et al 2005 International Materials Reviews vol. 6 pp 345
In the cork oak tree, the phellogen forms a continuous layer of cells, enveloping the tree trunk. Each year is produced a 2-3 mm thick layer of cork that adheres to that of the previous year.
A- Cork (Phellem)B- Subero-phellogenic changeC- PhellogeniumD- Liber tissueE- Liber wood changeF- Wood (Xylem)G- BarkH- Lenticular channelI- Area for stopper productionJ- Annual growth rings
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www.winomagazine.com/blog2/?cat=8
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CorkApplications
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http://www.ccrc.uga.edu/~mao/intro/ouline.htm
Structure of plant cell wall
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http://www.ccrc.uga.edu/~mao/intro/ouline.htm
T- Tertiary wallS- Secondary wallW- Extractables
(waxes, terpenes, sterols, etc)P- Primary wallM- Middle lamellaPo- Pore
Structure of cork oak cell wall
Silva et al 2005 International Materials Reviews vol. 6 pp 345
Structure of plant cell wall
T SuberinS
W P M
Po
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Chemical composition (%):
Suberin 30-50Lignin 15-27Polysaccharides 12-25Extractables (waxes, terpenes, sterols, etc) 8-20Ash 2- 5Others 1- 5
Silva et al 2005 International Materials Reviews vol. 6:345
Cork from cork oak tree
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SuberinComplex polymerTwo distinct domains:Polyphenolic and polyaliphatic
Lulai 2007 Skin-set, wound-healing and related effectsin Dick Vreugdenhi Ed. Potato Biology and Biotechnology Advances and Prespectives Elsevier, Amsterdam
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Lignin structure
Phenolic precursors
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Triose-PCalvinCycle
Pi
CO2
Erythrose-4-P
Sucrose
PEP
DAHP
Shikimate
Triose-P
PEP
Pi
Cytosol
Quinic acid
Quercitol
○
○
Glucose-6-P myo- Inositol-1-P
3-DehydroquinateNADH
3-Dehydroshikimate
Plastid
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Triose-PCalvinCycle
Pi
CO2
Erythrose-4-P
Sucrose
PEP
DAHP
Phenylalanine
Shikimate
Triose-P
PEP
Pi
Chorismate
Cytosol
Quinic acid
Quercitol
○
○
Glucose-6-P myo- Inositol-1-P
3-DehydroquinateNADH
3-Dehydroshikimate
Arogenate
Tyrosine
Tryptophan
Plastid
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Triose-PCalvinCycle
Pi
CO2
Erythrose-4-P
Sucrose
PEP
DAHP
Phenylalanine
Shikimate
p-Coumaric acid
Triose-P
PEP
Pi
Chorismate
Phenylalanine
FlavonoidsCytosol
Quinic acid
Quercitol
○
○
Glucose-6-P myo- Inositol-1-P
3-DehydroquinateNADH
3-Dehydroshikimate
Arogenate
Tyrosine
Tryptophan
CoumarinsStilbenes
p-Coumaryl alcoolCaffeic acid
Ferulic acid
Sinapic acid
Coniferyl alcool
Sinapyl alcool
Plastid
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Triose-PCalvinCycle
Pi
CO2
Erythrose-4-P
Sucrose
PEP
DAHP
Phenylalanine
Shikimate
p-Coumaric acid
Triose-P
PEP
Pi
Chorismate
Phenylalanine
FlavonoidsCytosol
Quinic acid
Quercitol
○
○ LigninsLignans
Cork arom. domain
Glucose-6-P myo- Inositol-1-P
3-DehydroquinateNADH
3-Dehydroshikimate
Arogenate
Tyrosine
Tryptophan
CoumarinsStilbenes
p-Coumaryl alcoolCaffeic acid
Ferulic acid
Sinapic acid
Coniferyl alcool
Sinapyl alcool
Plastid
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Triose-PCalvinCycle
Pi
CO2
Erythrose-4-P
Sucrose
PEP
DAHP
Phenylalanine
Shikimate
p-Coumaric acid
Triose-P
PEP
Pi
Chorismate
Phenylalanine
FlavonoidsCytosol
Quinic acid
Quercitol
○
○ LigninsLignans
Cork arom. domain
Glucose-6-P myo- Inositol-1-P
3-DehydroquinateNADH
3-Dehydroshikimate
Arogenate
Tyrosine
Tryptophan
CoumarinsStilbenes
p-Coumaryl alcoolCaffeic acid
Ferulic acid
Sinapic acid
Coniferyl alcool
Sinapyl alcool
● DQSDQS
DQS-Dehydroquinate synthase; CM-Chorismate mutase; PAL-Phenylalanine ammonia lyase; CAD-Cinnamoyl alcohol dehydrogenase;CinOR-Cinnamoyl oxiredutase; ChS-Chalcone synthase
● CMCM
●PALPAL
CADCAD●
CinORCinOR●
ChSChS●
Plastid
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Aliphatic Precursors of Suberin
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KAS- β-Ketoacyl-ACP synthetaseAT - Acyl tranferaseTE - Trans-enoylaseDS - DesaturasePC - Phosphatidyl-cholineTG -Triacilglicerol
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Plastid
Endoplasmic reticulum
Plant Lipid Metabolism
1- Two dehydrogenase steps via ω-oxo acid interediate; 2- ω-Hydroxylation; 3- In-chain hydroxylation; 4- Epoxidation of the double bond, followed by hydration5- Fatty acid synthetase; 6- β-Ketoacyl-ACP synthetase II (KAS II) step of fatty acid synthetase; 7- Stearoyl-ACP Δ9-desaturase; 8- Fatty acid elongationACP-Acyl carrier protein
Harwood (1997) Plant Lipid Metabolism in Dey and Harborne Ed. Plant Biochemistry Academic Press, San Diego
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Comparative analysis:phellem versus xylem
GenesProteins
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Functional categories of genes contributing to cork formation
Soler et al. (2007) A Genomic Approach to Suberin Biosynthesis and Cork Differentiation. Plant Physiology 144:419–431
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Differentially Expressed Proteins 3 IEF 10
MM
Carbohydratemetabolism
Energy
Secondarymetabolism Membrane transport
Stress/Defence
Unkown
Regulation/Signalling
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10 ºC 28 ºC
15 day
30 day
Stress effects: Air Temperature (10 ºC and 28 ºC)
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Triose-PCalvinCycle
Pi
CO2
Erythrose-4-P
Sucrose
PEP
DAHP
Phenylalanine
Shikimate
p-Coumaric acid
Triose-P
PEP
Pi
Chorismate
Phenylalanine
FlavonoidsCytosol
Quinic acid
Quercitol
○
○ LigninsLignans
Cork arom. domain
Glucose-6-P myo- Inositol-1-P
3-DehydroquinateNADH
3-Dehydroshikimate
Arogenate
Tyrosine
Tryptophan
CoumarinsStilbenes
p-Coumaryl alcoolCaffeic acid
Ferulic acid
Sinapic acid
Coniferyl alcool
Sinapyl alcool
● DQSDQS
DQS-Dehydroquinate synthase; CM-Chorismate mutase; PAL-Phenylalanine ammonia lyase; CAD-Cinnamoyl alcohol dehydrogenase;CinOR-Cinnamoyl oxiredutase; ChS-Chalcone synthase
● CMCM
●PALPAL
CADCAD●
CinORCinOR●
ChSChS●
Plastid
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The suberized skin of potato tuberas a model to study cork metabolism
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Structure of Potato Periderm
Lulai 2007 Skin-set, wound-healing and related effectsin Dick Vreugdenhi Ed. Potato Biology and Biotechnology Advances and Prespectives Elsevier, Amsterdam
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Barel, G. et al. J. Exp. Bot. 2008 59:3347-3357
Potato tuber skin development
(A) Number of skin layers (suberized phellem cells) during tuber development
(B) Early stage in periderm development(C) Close-up of dividing phellogen cells(D) Mature skin following foliage removal Bar=200 µm
Cross-sections of tuber surface stained with Safranin O/Fast green and viewed by light (B–D, left panels) and UV (B–D, right panels) microscopy to examine tissue morphology and autofluorescence of suberized cells.
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Barel, G. et al. J. Exp. Bot. 2008 59:3347-3357
Representative 2-DE images of skin and tuber storage parenchyma (flesh) at the developmental
stage of 8 weeks post-sprout-emergence
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Barel, G. et al. J. Exp. Bot. 2008 59:3347-3357
Cell proliferation Oxidative stressActin (ACT) Ascorbate peroxidase 1 (APX1), cytosolicP23 tumor protein-like (P23/TCTP) Catalase isozyme 2 (CAT2)Proteasome {alpha}-7 subunit Catechol oxidase B, chloroplast precursorProteasome β-2A subunit Polyphenol oxidase (PPO)Translation init iation factor 5A-3Tubulin {alpha}-chain Plant defenceSignal transduction—cell wall Cysteine protease 1 (CYP1)Remorin (REM) Elicitor-inducible protein EIG-J7
Elicitor-inducible protein EIG-J7General metabolism Endochitinase 2 precursorUDP-glucose:protein transglucosylase (UPTG2) Endochitinase 2 precursorDisulphide-isomerase protein (PDI) PatatinTriosephosphate isomerase, (TPI) cytosolic isoform Patatin putative homologOxidative respiratory chain Patatin protein 07APFI (hypothetical protein F8G22.2) Pathogenesis-related protein 10 (PR-10)NADH-ubiquinone oxidoreductase 18 kDa subunit Pathogenesis-related protein 10 (PR-10)NADH:FMN oxidoreductase-like protein 2-Oxoglutarate-dependent dioxygenase (SPP2)
One-carbon (C1) metabolism Suberization/lignificationGlutamate-ammonia ligase (GS1) ACP-17 kDa β-hydroxyacyl-acyl carrier proteinSerine hydroxymethyltransferase 4 (SHMT4) Caffeoyl-CoA O-methyltransferase-5 (CCoAOMT-5)Methionine synthase (MS) Caffeoyl-CoA O-methyltransferase-6 (CCoAOMT-6)Abiotic and biotic stress Caffeoyl-CoA O-methyltransferase-3 (CCoAOMT-3)Plasma-membrane polypeptide (DREPP) Peroxidase (POD 18)
Peroxidase PER9-6 secretory (POD 20)Reference protein Peroxidase 136, class III , precursor (POD 9)Nascent polypeptide-associated complex NAC; UBA-like Peroxidase putative (POD 5)
Peroxidase, suberization-associated anionic peroxidase
List of proteins that accumulate differentially in potato tuber skin compared to tuber storage parenchyma
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EVALUATION OF WOUND-HEALING PROCESSES IN POTATO TUBER TISSUE
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Potato Slices
Day 0 Day 8Day 4
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Potato Slices
Day 0 Day 8Day 4
Cellular dediferentiationGene expressionOxidative stress response
Suberin deposition
Wounding - Wound response - Healing
Periderm formation
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Potato Slices
Day 0 Day 8Day 4
Cellular dediferentiationGene expressionOxidative stress response
Suberin deposition
Suberin detected
Wounding - Wound response - Healing
Periderm formation
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Time-course of peroxidase and oxidase activity in wound-healing potato tubers
NADPH-dependent O2.– generation
Peroxidase activity
Razem et al. J.Exp. Botany, Vol. 54:935-941
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Proteomics of the wound-healing process
3 10pI
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Clustered mean expression profiles of differentially expressed proteins
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Clustered mean expression profiles of differentially expressed proteins
Wound response
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Clustered mean expression profiles of differentially expressed proteins
Periderm reconstruction
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Clustered mean expression profiles of differentially expressed proteins
Periderm reinforcement
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Cork FormationCellular Processes• Phellogen proliferation • Phellogen derived cells
– Phellem commitment– Cell expansion– Cell senescence– Suberin biosynthesis and waxes deposition– Cell death
Biochemical Processes• Cork results from 4 main secondary metabolic pathways:
– Acyl-lipids (aliphatic suberin domain)– Phenylpropanoids (cork aromatic components)– Isoprenoids (wax terpenes and sterols)– Flavonoids (tannins)
• Peroxidase activity fundamental (presence of H2O2)• Integration of biochemical process almost unkown