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Information on EC 2.5.1.115 - homogentisate phytyltransferase
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EC Tree
2.5.1.115 homogentisate phytyltransferaseIUBMB Comments
Requires Mg2+ for activity . Involved in the biosynthesis of the vitamin E tocopherols. While the enzyme from the cyanobacterium Synechocystis PCC 6803 has an appreciable activity with geranylgeranyl diphosphate (EC 2.5.1.116, homogentisate geranylgeranyltransferase), the enzyme from the plant Arabidopsis thaliana has only a low activity with that substrate [1,3,4].
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Word Map
- 2.5.1.115
- tocopherol
-
tocotrienols
- alpha-tocopherol
-
geranylgeranyl
- gamma-tocopherol
-
phytylation
-
prenylation
- synechocystis
-
tocochromanols
- prenyltransferase
-
lipid-soluble
-
tocopherol-deficient
-
seed-specific
- agriculture
- nutrition
- synthesis
The expected taxonomic range for this enzyme is: Bacteria, Eukaryota, Archaea
Reaction Schemes
Synonyms
homogentisate phytyltransferase, vte2-1, slr1736, vte2-2, vte2.1, 
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SYNONYM 
ORGANISM
UNIPROT
COMMENTARY 
LITERATURE
HPT
HPT1
VTE2
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REACTION
REACTION DIAGRAM
COMMENTARY
ORGANISM
UNIPROT
LITERATURE
phytyl diphosphate + homogentisate = diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
-
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PATHWAY SOURCE
PATHWAYS
BRENDA
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SYSTEMATIC NAME
IUBMB Comments
phytyl diphosphate:homogentisate phytyltransferase
Requires Mg2+ for activity [3]. Involved in the biosynthesis of the vitamin E tocopherols. While the enzyme from the cyanobacterium Synechocystis PCC 6803 has an appreciable activity with geranylgeranyl diphosphate (EC 2.5.1.116, homogentisate geranylgeranyltransferase), the enzyme from the plant Arabidopsis thaliana has only a low activity with that substrate [1,3,4].
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SUBSTRATE
PRODUCT
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate)
(Substrate)
LITERATURE
(Substrate)
(Substrate)
COMMENTARY
(Product)
(Product)
LITERATURE
(Product)
(Product)
Reversibility
r=reversible
ir=irreversible
?=not specified
r=reversible
ir=irreversible
?=not specified
geranylgeranyl diphosphate + homogentisate
diphosphate + 2-methyl-6-geranylgeranylbenzene-1,4-diol + CO2
geranylgeranyl diphosphate + homogentisate
diphosphate + 2-methyl-6-geranylgeranylbenzoquinol + CO2
-
-
-
?
geranylgeranyl diphosphate + homogentisate
diphosphate + 6-geranylgeranyl-2-methylbenzene-1,4-diol + CO2
about 18% of the activity with phythyldiphosphate
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzoquinol + CO2
-
-
-
?
geranylgeranyl diphosphate + homogentisate
diphosphate + 2-methyl-6-geranylgeranylbenzene-1,4-diol + CO2
-
-
-
?
geranylgeranyl diphosphate + homogentisate
diphosphate + 2-methyl-6-geranylgeranylbenzene-1,4-diol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
the enzyme is involved in the biosynthesis of the vitamin E tocopherols
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
no activity with geranylgeranyl diphosphate, no activity with solanesyl diphosphate
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
preferred substrate
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
the enzyme is involved in the biosynthesis of the vitamin E tocopherols
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
phytyl diphosphate and homogentisate are the preferred substrates in vito, no activity with solanesyl diphosphate
-
-
?
additional information
?
-
enzyme is more active with phythyl diphosphate than with geranylgeranyl diphosphate but inactive with solanesyl diphosphate or farnesyl diphosphate
-
-
?
additional information
?
-
recombinant Arabidopsis thaliana HPT preferentially catalyzes in vitro production of the tocotrienol precursor geranylgeranyl benzoquinol only when presented with high molar ratios of geranylgeranyl diphosphate:phytyl diphosphate
-
-
?
additional information
?
-
enzyme is more active with phythyl diphosphate than with geranylgeranyl diphosphate but inactive with solanesyl diphosphate or farnesyl diphosphate
-
-
?
additional information
?
-
no substrate: solanesyl diphosphate
-
-
?
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NATURAL SUBSTRATE
NATURAL PRODUCT
REACTION DIAGRAM
ORGANISM
UNIPROT
COMMENTARY
(Substrate)
(Substrate)
LITERATURE
(Substrate)
(Substrate)
COMMENTARY
(Product)
(Product)
LITERATURE
(Product)
(Product)
REVERSIBILITY
r=reversible
ir=irreversible
?=not specified
r=reversible
ir=irreversible
?=not specified
geranylgeranyl diphosphate + homogentisate
diphosphate + 2-methyl-6-geranylgeranylbenzoquinol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzoquinol + CO2
-
-
-
?
additional information
?
-
recombinant Arabidopsis thaliana HPT preferentially catalyzes in vitro production of the tocotrienol precursor geranylgeranyl benzoquinol only when presented with high molar ratios of geranylgeranyl diphosphate:phytyl diphosphate
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
the enzyme is involved in the biosynthesis of the vitamin E tocopherols
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
-
-
-
-
?
phytyl diphosphate + homogentisate
diphosphate + 2-methyl-6-phytylbenzene-1,4-diol + CO2
the enzyme is involved in the biosynthesis of the vitamin E tocopherols
-
-
?
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METALS and IONS
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
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INHIBITOR
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
IMAGE
n-dodecyl beta-D-maltoside
stimulation at low concentration, strong inhibition above 0.1 mM
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ACTIVATING COMPOUND
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
IMAGE
n-dodecyl beta-D-maltoside
stimulation at low concentration, strong inhibition above 0.1 mM
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KM VALUE [mM]
SUBSTRATE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
IMAGE
0.001
(2E,7R,11R)-3,7,11,15-tetramethylhexadec-2-en-1-yl trihydrogen diphosphate
pH 7.5, 28°C
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pH OPTIMUM
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
7.5
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pI VALUE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
9.5
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ORGANISM
COMMENTARY
LITERATURE
UNIPROT
SEQUENCE DB
SOURCE
-
SwissProt
brenda
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SOURCE TISSUE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
SOURCE
-
gene expression level gradually decreases during fruit ripening, reaching a background level in ripened apple fruits
brenda
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LOCALIZATION
ORGANISM
UNIPROT
COMMENTARY
GeneOntology No.
LITERATURE
SOURCE
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GENERAL INFORMATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
malfunction
disruption of Synechocystis sp. PCC 6803 HPT function causes an absence of tocopherols without affecting plastoquinone-9 levels, indicating that separate polyprenyltransferases exist for tocopherol and plastoquinone synthesis in Synechocystis sp. PCC 6803
physiological function
physiological function
the enzyme is involved in the biosynthesis of the vitamin E tocopherols. Tocopherol functions to limit lipid oxidation during seed desiccation, quiescence and germination and early seedling development. Protects thylakoid membrane lipids from photooxidation and is required for low-temperature adaptation
physiological function
the enzyme is involved in the biosynthesis of the vitamin E tocopherols. Tocopherol functions to limit lipid oxidation during seed desiccation, quiescence and germination and early seedling development. Protects thylakoid membrane lipids from photooxidation and is required for low-temperature adaptation
physiological function
a null mutant obtained by insertional inactivation does not accumulate tocopherols, and is rescued by the Arabidopsis HPT1 ortholog. The membrane fraction of wild-type Synechocystis sp. PCC 6803 is capable of catalyzing the phytylation of homogentisate, whereas the membrane fraction from the slr1736 null mutant is not
physiological function
a Synechocystis sp. PCC 6803 homogentisate phytyltransferase null mutant is rescued by the Arabidopsis HPT1 ortholog. Antisense expression of HPT1 in Arabidopsis results in reduced seed tocopherol levels, whereas seed-specific sense expression results in increased seed tocopherol levels
physiological function
abiotic stress results in an 18 and 8fold increase in total tocopherol content in wild-type and isoform HPT1 overexpressing leaves, respectively, with tocopherol levels in the ovrexpressing strain being 2- to 4fold higher than wild type at all experimental time points. Increased total tocopherol levels correlate with elevated HPT mRNA levels and HPT specific activity, suggesting that HPT activity limits total tocopherol synthesis during abiotic stress. Overexpression of gamma-tocopherol methyltransferase in the HPT1 overexpression strain results in nearly complete conversion of gamma- and delta-tocopherols to alpha- and beta-tocopherols
physiological function
in leaves, isoform HPT1 constitutive overexpression results in a 10fold increase in HPT specific activity and a 4.4fold increase in total tocopherol content relative to wild type. In seeds, HPT1 overexpression results in a 4fold increase in HPT specific activity and a total seed tocopherol content that is 40% higher than wild type, primarily because of an increase in gamma-tocopherol content. This enlarged pool of gamma-tocopherol is almost entirely converted to alpha-tocopherol by crossing HPT1 overexpressing plants with lines constitutively overexpressing gamma-tocopherol methyltransferase. Seed of the resulting double-overexpressing lines displays a 12fold increase in vitamin E activity relative to wild type
physiological function
seed-specific expression of isoform VTE2-2 in Arabidopsis results in increased seed-tocopherol levels
physiological function
SLR1736 gene disruption does not lead to significant differences in cellular plastoquinone, phylloquinone, and chlorophyll levels, but causes absence of tocopherols without affecting plastoquinone-9 levels
physiological function
-
T1 leaves of transgenic tomato plants overexpressing the apple enzyme contain1.8- to 3.6fold and1.6- to 2.9fold higher levels of alpha-tocopherol and gamma-tocopherol, respectively, than those in control plants. The levels of alpha-tocopherol and gamma-tocopherol in T1 fruits increases up to 1.7fold and 3.1fold, respectively
physiological function
an Arabidopsis thalina that accumulates tocotrienols and homogentisate by co-expression of Arabidopsis thaliana hydroxyphenylpyruvate dioxygenase and Escherichia coli bi-functional chorismate mutase/prephenate dehydrogenase loses tocotrienol production when crossed into the vte2-1 HPT null mutant. Hydroxyphenylpyruvate dioxygenase/TyrA-induced tocotrienol production requires HPT and occurs upon enrichment of geranylgeranyl diphosphaterelative to phytyl diphosphate in prenyl diphosphate pools
physiological function
-
expression of homogentisate phytyltransferase, tocopherol cyclase, and gamma-tocopherol methyltransferase on a synthetic operon to synthesize vitamin E in leaves of Nicotiana tabacum and Lycopersicum esculentum. Homogentisate phytyltransferase is the limiting enzymatic step in the pathway. An up to tenfold increase in total tocochromanol accumulation is achieved. A light-dependent regulatory link exosts between tocochromanol metabolism and the pathways of photosynthetic pigment biosynthesis
physiological function
-
transgenic tobacco plants expressing VTE2 under the control of stress-inducible promoters show increased levels of alpha-tocopherol when exposed to drought conditions. The accumulation of alpha-tocopherol correlates with higher water content and increased photosynthetic performance and less oxidative stress damage
physiological function
-
the enzyme catalyzes a key step in rice tocopherol biosynthesis, confers cold tolerance and regulates rice development by affecting the accumulation of DELLA protein SLENDER RICE1
Show AA Sequence and Transmembrane Helices (246 entries)
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MOLECULAR WEIGHT
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
30974
functional protein, after processing a transit peptide with 304 residues
44400
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SUBUNIT
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
?
x * 32744, including transit peptide, x * 30974, without transit peptide, calculated
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POSTTRANSLATIONAL MODIFICATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
proteolytic modification
sequence contains a 19 amin acid plastidic transit peptide
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CLONED (Commentary)
ORGANISM
UNIPROT
LITERATURE
co-expressed with tocopherol cyclase in Nicotiana benthamiana leaves
expression in Escherichia coli
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EXPRESSION
ORGANISM
UNIPROT
LITERATURE
abiotic stress results in an 18 and 8fold increase in total tocopherol content in wild-type and isoform HPT1 overexpressing leaves, respectively, with tocopherol levels in the ovrexpressing strain being 2- to 4fold higher than wild type at all experimental time points. Increased total tocopherol levels correlate with elevated HPT mRNA levels and HPT specific activity
enzyme is constitutively expressed in all analyzed tissues except in kernel 15 weeks after anthesis. Enzyme is highly expressed at the fruit ripening stage of 17 weeks after anthesis
isoform Hpt1 is strongly regulated by dark and low-temperature. Under the same treatments, alpha-tocopherol increases in cultures exposed to darkness or heat, whereas gamma-tocopherol does it in low temperature
the expression patterns of the five tocopherol biosynthetic genes vary in different plant tissues and under different stress conditions. All five genes are highly expressed in leaf tissues, whereas 4-hydroxyphenylpyruvate dioxygenase and homogentisate phytyltransferase are highly expressed in the thick roots. The expression patterns of these five genes significantly differ in response to PEG, NaCl and H2O2-mediated oxidative stress. Homogentisate phytyltransferase is strongly induced following PEG treatment
enzyme is constitutively expressed in all analyzed tissues except in kernel 15 weeks after anthesis. Enzyme is highly expressed at the fruit ripening stage of 17 weeks after anthesis
enzyme is constitutively expressed in all analyzed tissues except in kernel 15 weeks after anthesis. Enzyme is highly expressed at the fruit ripening stage of 17 weeks after anthesis
the expression patterns of the five tocopherol biosynthetic genes vary in different plant tissues and under different stress conditions. All five genes are highly expressed in leaf tissues, whereas 4-hydroxyphenylpyruvate dioxygenase and homogentisate phytyltransferase are highly expressed in the thick roots. The expression patterns of these five genes significantly differ in response to PEG, NaCl and H2O2-mediated oxidative stress. Homogentisate phytyltransferase is strongly induced following PEG treatment
-
the expression patterns of the five tocopherol biosynthetic genes vary in different plant tissues and under different stress conditions. All five genes are highly expressed in leaf tissues, whereas 4-hydroxyphenylpyruvate dioxygenase and homogentisate phytyltransferase are highly expressed in the thick roots. The expression patterns of these five genes significantly differ in response to PEG, NaCl and H2O2-mediated oxidative stress. Homogentisate phytyltransferase is strongly induced following PEG treatment
-
-
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APPLICATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
agriculture
nutrition
in leaves, isoform HPT1 constitutive overexpression results in a 10fold increase in HPT specific activity and a 4.4fold increase in total tocopherol content relative to wild type. In seeds, HPT1 overexpression results in a 4fold increase in HPT specific activity and a total seed tocopherol content that is 40% higher than wild type, primarily because of an increase in gamma-tocopherol content. This enlarged pool of gamma-tocopherol is almost entirely converted to alpha-tocopherol by crossing HPT1 overexpressing plants with lines constitutively overexpressing gamma-tocopherol methyltransferase. Seed of the resulting double-overexpressing lines displays a 12fold increase in vitamin E activity relative to wild type
synthesis
constitutive expression of genes encoding homogentisate phytyltransferase HPT and tocopherol cyclase TC individually and in combination in tobacco. The alpha-tocopherol content in transgenic tobacco plants expressing HPT, TC, and both HPT and TC is increased by 5.4-, 4.0-, and 7.1fold, respectively, when compared to the wild type
agriculture
-
stress-induced expression of VTE2 can be used to increase the vitamin E content and to diminish detrimental effects of environmental stress in plants
agriculture
the transient expression of the enzyme gene enhances the vitamin E levels in transgenic Nicotiana tabacum leaves. The stable expression of the enzyme gene is an efficient strategy to enhance vitamin E biosynthesis in agricultural crop breeding
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REF.
AUTHORS
TITLE
JOURNAL
VOL.
PAGES
YEAR
ORGANISM (UNIPROT)
PUBMED ID
SOURCE
Sadre, R.; Gruber, J.; Frentzen, M.
Characterization of homogentisate prenyltransferases involved in plastoquinone-9 and tocochromanol biosynthesis
FEBS Lett.
580
5357-5362
2006
Synechocystis sp. (P73726), Arabidopsis thaliana (Q8VWJ1)
brenda
Collakova, E.; DellaPenna, D.
Isolation and functional analysis of homogentisate phytyltransferase from Synechocystis sp. PCC 6803 and Arabidopsis
Plant Physiol.
127
1113-1124
2001
Synechocystis sp. (P73726), Synechocystis sp., Arabidopsis thaliana (Q8VWJ1)
brenda
Harish, M.; Dachinamoorthy, P.; Balamurugan, S.; Bala Murugan, S.; Sathishkumar, R.
Overexpression of homogentisate phytyltransferase (HPT) and tocopherol cyclase (TC) enhances alpha-tocopherol content in transgenic tobacco
Biol. Plant.
57
395-400
2013
Arabidopsis thaliana (Q8VWJ1)
-
brenda
Seo, Y.S.; Kim, S.J.; Harn, C.H.; Kim, W.T.
Ectopic expression of apple fruit homogentisate phytyltransferase gene (MdHPT1) increases tocopherol in transgenic tomato (Solanum lycopersicum cv. Micro-Tom) leaves and fruits
Phytochemistry
72
321-329
2011
Malus domestica
brenda
Zhang, C.; Cahoon, R.E.; Hunter, S.C.; Chen, M.; Han, J.; Cahoon, E.B.
Genetic and biochemical basis for alternative routes of tocotrienol biosynthesis for enhanced vitamin E antioxidant production
Plant J.
73
628-639
2013
Arabidopsis thaliana (Q8VWJ1)
brenda
Savidge, B.; Weiss, J.D.; Wong, Y.H.; Lassner, M.W.; Mitsky, T.A.; Shewmaker, C.K.; Post-Beittenmiller, D.; Valentin, H.E.
Isolation and characterization of homogentisate phytyltransferase genes from Synechocystis sp. PCC 6803 and Arabidopsis
Plant Physiol.
129
321-332
2002
Synechocystis sp. (P73726), Synechocystis sp., Arabidopsis thaliana (Q8VWJ1)
brenda
Collakova, E.; DellaPenna, D.
Homogentisate phytyltransferase activity is limiting for tocopherol biosynthesis in Arabidopsis
Plant Physiol.
131
632-642
2003
Arabidopsis thaliana (Q8VWJ1)
brenda
Collakova, E.; DellaPenna, D.
The role of homogentisate phytyltransferase and other tocopherol pathway enzymes in the regulation of tocopherol synthesis during abiotic stress
Plant Physiol.
133
930-940
2003
Arabidopsis thaliana (Q8VWJ1)
brenda
Venkatesh, T.V.; Karunanandaa, B.; Free, D.L.; Rottnek, J.M.; Baszis, S.R.; Valentin, H.E.
Identification and characterization of an Arabidopsis homogentisate phytyltransferase paralog
Planta
223
1134-1144
2006
Arabidopsis thaliana (Q1ACB3)
brenda
Espinoza, A.; San Martin, A.; Lopez-Climent, M.; Ruiz-Lara, S.; Gomez-Cadenas, A.; Casaretto, J.A.
Engineered drought-induced biosynthesis of alpha-tocopherol alleviates stress-induced leaf damage in tobacco
J. Plant Physiol.
170
1285-1294
2013
Solanum chilense
brenda
Galvez-Valdivieso, G.; Cardenosa, R.; Pineda, M.; Aguilar, M.
Homogentisate phytyltransferase from the unicellular green alga Chlamydomonas reinhardtii
J. Plant Physiol.
188
80-88
2015
Chlamydomonas reinhardtii (A0A078LN07), Chlamydomonas reinhardtii
brenda
Kong, S.; Abdullah, S.; Ho, C.; Amiruddin, M.
Molecular cloning, gene expression profiling and in silico sequence analysis of Vitamin E biosynthetic genes from the oil palm
Plant Gene
5
100-108
2016
Elaeis oleifera (W8NZU5), Elaeis guineensis (W8P019)
-
brenda
Ji, C.Y.; Kim, Y.H.; Kim, H.S.; Ke, Q.; Kim, G.W.; Park, S.C.; Lee, H.S.; Jeong, J.C.; Kwak, S.S.
Molecular characterization of tocopherol biosynthetic genes in sweetpotato that respond to stress and activate the tocopherol production in tobacco
Plant Physiol. Biochem.
106
118-128
2016
Ipomoea batatas, Ipomoea batatas Lam
brenda
Lu, Y.; Rijzaani, H.; Karcher, D.; Ruf, S.; Bock, R.
Efficient metabolic pathway engineering in transgenic tobacco and tomato plastids with synthetic multigene operons
Proc. Natl. Acad. Sci. USA
110
E623-E632
2013
Synechocystis sp.
brenda
Sathish, S.; Preethy, K.S.; Venkatesh, R.; Sathishkumar, R.
Rapid enhancement of alpha-tocopherol content in Nicotiana benthamiana by transient expression of Arabidopsis thaliana tocopherol cyclase and homogentisate phytyl transferase genes
3 Biotech
8
485
2018
Arabidopsis thaliana (Q8VWJ1), Arabidopsis thaliana
brenda
Wunnakup, T.; Vimolmangkang, S.; De-Eknamkul, W.
Transient expression of the homogentisate phytyltransferase gene from Clitoria ternatea causes metabolic enhancement of alpha-tocopherol biosynthesis and chlorophyll degradation in tomato leaves
J. Plant Biochem. Biotechnol.
27
55-67
2018
Clitoria ternatea (A0A1L2BI34)
-
brenda
Zhang, Y.; Liu, K.; Zhu, X.; Wu, Y.; Zhang, S.; Chen, H.; Ling, J.; Wang, Y.; Fang, X.
Rice tocopherol deficiency 1 encodes a homogentisate phytyltransferase essential for tocopherol biosynthesis and plant development in rice
Plant Cell Rep.
37
775-787
2018
Oryza sativa
brenda
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