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Information on EC 2.5.1.79 - thermospermine synthase
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2.5.1.79 thermospermine synthaseIUBMB Comments
This plant enzyme is crucial for the proper functioning of xylem vessel elements in the vascular tissues of plants .
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Word Map
- 2.5.1.79
- polyamine
- xylem
- auxin
-
dwarf
- putrescine
-
uorfs
- dwarfism
-
tetraamine
-
internode
-
scots
- aminopropyltransferases
-
procambial
-
conifer
- analysis
The expected taxonomic range for this enzyme is: Eukaryota, Bacteria, Archaea
Reaction Schemes
Synonyms
sac51, acaulis5, thermospermine synthase, tspms, popacaulis5, ghacl5, t-spm synthase, 
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SYNONYM 
ORGANISM
UNIPROT
COMMENTARY 
LITERATURE
ACAULIS5
ACL5
AtACL5
GhACL5
T-Spm synthase
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REACTION
REACTION DIAGRAM
COMMENTARY
ORGANISM
UNIPROT
LITERATURE
S-adenosyl 3-(methylsulfanyl)propylamine + spermidine = S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
-
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SYSTEMATIC NAME
IUBMB Comments
S-adenosyl 3-(methylthio)propylamine:spermidine 3-aminopropyltransferase (thermospermine synthesizing)
This plant enzyme is crucial for the proper functioning of xylem vessel elements in the vascular tissues of plants [3].
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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
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl-L-methioninamine + spermidine
5'-methylthioadenosine + thermospermine
-
-
-
?
S-adenosyl-L-methioninamine + spermidine
5'-methylthioadenosine + thermospermine
-
SAC51 is one of the key transcription factors controlling stem elongation. Plants acquire the ability to synthesize thermospermine at an early stage of evolution by horizontal gene transfer from a prokaryote
-
-
?
S-adenosyl-L-methioninamine + spermidine
5'-methylthioadenosine + thermospermine
-
-
sole product
-
?
S-adenosyl-L-methioninamine + spermidine
5'-methylthioadenosine + thermospermine
-
-
sole product, no synthesis of spermine
-
?
additional information
?
-
isoform Acl5 is required for stem elongation
-
-
?
additional information
?
-
-
isoform Acl5 is required for stem elongation
-
-
?
additional information
?
-
-
ACL5 is required for correct xylem specification through regulation of the lifetime of the xylem elements
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-
?
additional information
?
-
the enzyme GhACL5 contains potential S-adenosyl-L-methionine binding sites
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-
?
additional information
?
-
-
the enzyme GhACL5 contains potential S-adenosyl-L-methionine binding sites
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-
?
additional information
?
-
the enzyme GhACL5 contains potential S-adenosyl-L-methionine binding sites
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-
?
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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
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
S-adenosyl-L-methioninamine + spermidine
5'-methylthioadenosine + thermospermine
-
SAC51 is one of the key transcription factors controlling stem elongation. Plants acquire the ability to synthesize thermospermine at an early stage of evolution by horizontal gene transfer from a prokaryote
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
S-adenosyl 3-(methylthio)propylamine + spermidine
S-methyl-5'-thioadenosine + thermospermine + H+
-
-
-
?
additional information
?
-
isoform Acl5 is required for stem elongation
-
-
?
additional information
?
-
-
isoform Acl5 is required for stem elongation
-
-
?
additional information
?
-
-
ACL5 is required for correct xylem specification through regulation of the lifetime of the xylem elements
-
-
?
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DISEASE
TITLE OF PUBLICATION
LINK TO PUBMED
Dwarfism
Complexity and Conservation of Thermospermine-Responsive uORFs of SAC51 Family Genes in Angiosperms.
Dwarfism
Mutations in Ribosomal Proteins, RPL4 and RACK1, Suppress the Phenotype of a Thermospermine-Deficient Mutant of Arabidopsis thaliana.
Gallstones
Structural analysis of gallstones with thin-section petrographic microscopy: a study of 100 gallstones from Taiwanese patients.
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KM VALUE [mM]
SUBSTRATE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
IMAGE
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SPECIFIC ACTIVITY [µmol/min/mg]
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
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pH OPTIMUM
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
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TEMPERATURE OPTIMUM
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
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ORGANISM
COMMENTARY
LITERATURE
UNIPROT
SEQUENCE DB
SOURCE
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SOURCE TISSUE
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
SOURCE
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stems and flowers contain two- to threefold more thermospermine compared to whole seedlings and mature leaves
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strong ACL5 expression is present in the megagametophyte cells destined to die via morphologically necrotic cell death
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in developing seeds, the expression of both ACL5 and diamine oxidase increases as embryogenesis proceeds
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strong ACL5 expression is present in the procambial cells of the embryo and in the megagametophyte cells destined to die via morphologically necrotic cell death
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strong ACL5 expression is present in the procambial cells of the embryo
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-
stems and flowers contain two- to threefold more thermospermine compared to whole seedlings and mature leaves
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stems and flowers contain two- to threefold more thermospermine compared to whole seedlings and mature leaves
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-
stems and flowers contain two- to threefold more thermospermine compared to whole seedlings and mature leaves
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in the hypocotyl as well as in the inflorescence stem, ACL5 is expressed not just broadly with respect to vasculature, but specifically in the xylem vessel elements at a strictly defined developmental stage, suggesting direct involvement of ACL5 in xylem vessel differentiation. The acl5 mutant displays severe overall inhibition of the secondary growth of the vascular tissues, dramatic alteration in the morphology of the vessel elements and complete lack of xylem fibers
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the enzyme is expressed specifically in early developing xylem vessel elements
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GENERAL INFORMATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
malfunction
physiological function
malfunction
plants containing the mutation of AtACL5 have defective elongation of their stem internodes because cell expansion is reduced
malfunction
the increase in height in GhACL5-overexpressing plants cannot be attributed to early germination rates. GhACL5-silenced plants show a severe dwarf phenotype
malfunction
-
the increase in height in GhACL5-overexpressing plants cannot be attributed to early germination rates. GhACL5-silenced plants show a severe dwarf phenotype
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malfunction
-
plants containing the mutation of AtACL5 have defective elongation of their stem internodes because cell expansion is reduced
-
physiological function
thermospermine-deficient mutant acl5 shows severe dwarfism and excessive xylem differentiation. Some auxin analogs remarkably enhance xylem vessel differentiation in the acl5 mutant but not in the wild type. The xylem-inducing effect of auxin analogs is clearly suppressed by thermospermine. Auxin analogs promote protoxylem differentiation in roots and cotyledons in the acl5 mutant. The opposite action between thermospermine and auxin in xylem differentiation may be common in different organs and thermospermine might be required for the suppression of protoxylem differentiation
physiological function
the enzyme is involved in synthesis of thermospermidine, which has a toxic effect on pthogen Verticillium dahliae development in a dose-dependent manner
physiological function
thermospermine levels are controlled by an auxin-dependent feedback loop mechanism in Populus xylem. Negative feedback regulation, increased POPACAULIS5 expression functions to reduce indole-3-acetic acid levels, which in turn prevents further expression of POPACAULIS5, a regulatory mechanism, mediated through auxin and PttHB8, for maintenance of thermospermine homeostasis in secondary xylem tissues of the stem
physiological function
involvement of ACL5 in morphologically necrotic cell death. Thermospermine synthase (ACL5) and diamine oxidase (DAO) expression is needed for zygotic embryogenesis and vascular development in scots pine
physiological function
-
the enzyme is involved in synthesis of thermospermidine, which has a toxic effect on pthogen Verticillium dahliae development in a dose-dependent manner
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UNIPROT
ENTRY NAME
ORGANISM
NO. OF AA
NO. OF TRANSM. HELICES
MOLECULAR WEIGHT[Da]
SOURCE
SEQUENCE
LOCALIZATION PREDICTION
The reliability class of the localization prediction ranges from 1 (very reliable) to 5 (not reliable).
The reliability class of the localization prediction ranges from 1 (very reliable) to 5 (not reliable). ACL5_ARATH
339
0
38547
Swiss-Prot
other Location (Reliability: 2)
SPEE1_HYPBU
Hyperthermus butylicus (strain DSM 5456 / JCM 9403 / PLM1-5)
292
0
32856
Swiss-Prot
-
SPEE_SACS2
Saccharolobus solfataricus (strain ATCC 35092 / DSM 1617 / JCM 11322 / P2)
301
0
34854
Swiss-Prot
-
SPEE_THET8
Thermus thermophilus (strain ATCC 27634 / DSM 579 / HB8)
314
0
36006
Swiss-Prot
-
SPEE_PYRAE
Pyrobaculum aerophilum (strain ATCC 51768 / DSM 7523 / JCM 9630 / CIP 104966 / NBRC 100827 / IM2)
291
0
32806
Swiss-Prot
-
A0A5B7BTQ1_DAVIN
365
0
41424
TrEMBL
Secretory Pathway (Reliability: 1)
A0A2G9HI57_9LAMI
329
0
37264
TrEMBL
other Location (Reliability: 1)
A0A2G9G805_9LAMI
342
0
39023
TrEMBL
other Location (Reliability: 2)
A0A0C4JZU4_GOSHI
339
0
38421
TrEMBL
Secretory Pathway (Reliability: 4)
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MOLECULAR WEIGHT
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
48000
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SUBUNIT
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
tetramer
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CRYSTALLIZATION (Commentary)
ORGANISM
UNIPROT
LITERATURE
crystal structures, one without ligands and five in complexes with either reaction substrate, reaction product, or cofactor analogs (5'-methylthioadenosine, S-adenosylthiopropylamine, and adenosine). TSPS is a symmetric homotetramer with an interdomain eight-stranded beta-barrel
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PROTEIN VARIANTS
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
E156D
-
besides main product thermospermine, mutant is able to synthesize some spermine
additional information
additional information
contrary to wild-type, seedlings of acl5-1 loss-of-function mutant do not contain thermospermine. Daily application of thermospermine onto the shoot apex partially rescues the dwarf phenotype of the mutant, while application of spermine has no effect. The acl5-1 transcript level in acl5-1 seedlings, which is much higher than the ACL5 transcript level in wild-type seedlings, is reduced by exogenous thermospermine
additional information
-
contrary to wild-type, seedlings of acl5-1 loss-of-function mutant do not contain thermospermine. Daily application of thermospermine onto the shoot apex partially rescues the dwarf phenotype of the mutant, while application of spermine has no effect. The acl5-1 transcript level in acl5-1 seedlings, which is much higher than the ACL5 transcript level in wild-type seedlings, is reduced by exogenous thermospermine
additional information
constitutive expression of the cotton ACL5 gene (GhACL5) in transgenic Arabidopsis thaliana ecotype Columbia-0, via Agrobacterium tumefaciens strain GV3101 transfection using the CaMV 35S promoter, significantly increases plant height and elevates the level of thermospermidine, quantitative real-time RT-PCR expression analysis of transgenic GhACL5 and endogenous AtACL5 transcripts, transcript levels of endogenous AtACL5 are reduced in plants that ectopically express gene GhACL5, seedling phenotypes, overview. Germination rate for seeds from GhACL5-overexpressing plants is lower when compared with control after 3 days
additional information
-
constitutive expression of the cotton ACL5 gene (GhACL5) in transgenic Arabidopsis thaliana ecotype Columbia-0, via Agrobacterium tumefaciens strain GV3101 transfection using the CaMV 35S promoter, significantly increases plant height and elevates the level of thermospermidine, quantitative real-time RT-PCR expression analysis of transgenic GhACL5 and endogenous AtACL5 transcripts, transcript levels of endogenous AtACL5 are reduced in plants that ectopically express gene GhACL5, seedling phenotypes, overview. Germination rate for seeds from GhACL5-overexpressing plants is lower when compared with control after 3 days
additional information
-
constitutive expression of the cotton ACL5 gene (GhACL5) in transgenic Arabidopsis thaliana ecotype Columbia-0, via Agrobacterium tumefaciens strain GV3101 transfection using the CaMV 35S promoter, significantly increases plant height and elevates the level of thermospermidine, quantitative real-time RT-PCR expression analysis of transgenic GhACL5 and endogenous AtACL5 transcripts, transcript levels of endogenous AtACL5 are reduced in plants that ectopically express gene GhACL5, seedling phenotypes, overview. Germination rate for seeds from GhACL5-overexpressing plants is lower when compared with control after 3 days
-
additional information
generation of GhACL5-sielenced plants endogenous levels of putrescine, spermidine, and spermine are similar between GhACL5-silenced and control plants. GhACL5-silenced plants show a severe dwarf phenotype
additional information
-
generation of GhACL5-sielenced plants endogenous levels of putrescine, spermidine, and spermine are similar between GhACL5-silenced and control plants. GhACL5-silenced plants show a severe dwarf phenotype
additional information
-
generation of GhACL5-sielenced plants endogenous levels of putrescine, spermidine, and spermine are similar between GhACL5-silenced and control plants. GhACL5-silenced plants show a severe dwarf phenotype
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additional information
enzme overexpression under the control of the 35S CaMV constitutive promoter in Populus tremula x Populus tremuloides plants leads to reduced overall growth and slight defects in xylem development. Ectopic expression of POPACAULIS5 results in the dramatic changes in shoot and root development, and the transgenic lines shows (severe) dwarf phenotypes, overview. Some dwarf transgenic plants are able to develop a rooting system and show partial restoration of the wild-type phenotype once transferred to a plant growth regulator-free medium. Thermospermine accumulation is suppressed in the 35S::POPACAULIS5 Populus woody stem, but not in leaves. Indole-3-acetic acid levels are strongly reduced in the leaves and in the secondary xylem tissues of the transgenic trees
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CLONED (Commentary)
ORGANISM
UNIPROT
LITERATURE
expression in Escherichia coli
gene GhACL5, DNA and amino acid sequence determination and analysis, phylogenetic analysis, sequence comparisons, constitutive expression of the cotton ACL5 gene (GhACL5) in Arabidopsis thaliana ecotype Columbia-0, via Agrobacterium tumefaciens strain GV3101 transfection using the CaMV 35S promoter, significantly increases plant height and elevates the level of thermospermidine, quantitative real-time RT-PCR expression analysis of transgenic GhACL5 and endogenous AtACL5 transcripts, transcript levels of endogenous AtACL5 are reduced in plants that ectopically express gene GhACL5, seedling phenotypes, overview
gene POPACAULIS5 or PttACL5, sequence comparisons and phylogenetic analysis, quantitative real-time RT-PCR expression analysis, functional expression in Saccharomyces cerevisiae cells
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EXPRESSION
ORGANISM
UNIPROT
LITERATURE
ACL5 expression is associated with morphologically necrotic cell death
exogenous supply of auxin results in a strong increase in POPACAULIS5 transcript levels after 4 h in transgenic overexpressing lines but not in the wild-type. Class III HD-Zip PttHB8 over-expression stimulates POPACAULIS5 expression, mechanism, overview. Expression of PttHB8 is rapidly induced by exogenous auxin and it is therefore possible that the suppression of PttHB8 expression by POPACAULIS5 overexpression is mediated through Indole-3-acetic acid
jasmonic acid methyl ester and ethephon affect the enzyme expression
salicylic acid, jasmonic acid methyl ester, and ethephon induce the enzyme expression
jasmonic acid methyl ester and ethephon affect the enzyme expression
jasmonic acid methyl ester and ethephon affect the enzyme expression
-
-
salicylic acid, jasmonic acid methyl ester, and ethephon induce the enzyme expression
salicylic acid, jasmonic acid methyl ester, and ethephon induce the enzyme expression
-
-
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APPLICATION
ORGANISM
UNIPROT
COMMENTARY
LITERATURE
analysis
-
simple method to separate spermine and thermospermine in extracts from plant tissues of less than 500 mg. Polyamines extracted from plant tissues are benzoylated, and the derivatives are completely resolved by high-performance liquid chromatography on a C18 reverse-phase column. With this method, the thermospermine contents in various tissues of Arabidopsis thaliana have been monitored. Stems and flowers contain two- to three-fold more thermospermine compared to whole seedlings and mature leaves
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REF.
AUTHORS
TITLE
JOURNAL
VOL.
PAGES
YEAR
ORGANISM (UNIPROT)
PUBMED ID
SOURCE
Knott, J.M.; Roemer, P.; Sumper, M.
Putative spermine synthases from Thalassiosira pseudonana and Arabidopsis thaliana synthesize thermospermine rather than spermine
FEBS Lett.
581
3081-3086
2007
Thalassiosira pseudonana, Arabidopsis thaliana (Q9S7X6)
brenda
Romer, P.; Faltermeier, A.; Mertins, V.; Gedrange, T.; Mai, R.; Proff, P.
Investigations about N-aminopropyl transferases probably involved in biomineralization
J. Physiol. Pharmacol.
59 Suppl 5
27-37
2008
Thalassiosira pseudonana
brenda
Kakehi, J.; Kuwashiro, Y.; Niitsu, M.; Takahashi, T.
Thermospermine is required for stem elongation in Arabidopsis thaliana
Plant Cell Physiol.
49
1342-1349
2008
Arabidopsis thaliana (Q9S7X6), Arabidopsis thaliana
brenda
Takahashi, T.; Kakehi, J.
Polyamines: ubiquitous polycations with unique roles in growth and stress responses
Ann. Bot.
105
1-6
2010
Arabidopsis thaliana
brenda
Muniz, L.; Minguet, E.G.; Singh, S.K.; Pesquet, E.; Vera-Sirera, F.; Moreau-Courtois, C.L.; Carbonell, J.; Blazquez, M.A.; Tuominen, H.
ACAULIS5 controls Arabidopsis xylem specification through the prevention of premature cell death
Development
135
2573-2582
2008
Arabidopsis thaliana
brenda
Yoshimoto, K.; Noutoshi, Y.; Hayashi, K.; Shirasu, K.; Takahashi, T.; Motose, H.
A chemical biology approach reveals an opposite action between thermospermine and auxin in xylem development in Arabidopsis thaliana
Plant Cell Physiol.
53
635-645
2012
Arabidopsis thaliana (Q9S7X6), Arabidopsis thaliana
brenda
Naka, Y.; Watanabe, K.; Sagor, G.H.; Niitsu, M.; Pillai, M.A.; Kusano, T.; Takahashi, Y.
Quantitative analysis of plant polyamines including thermospermine during growth and salinity stress
Plant Physiol. Biochem.
48
527-533
2010
Arabidopsis thaliana
brenda
Mo, H.; Wang, X.; Zhang, Y.; Yang, J.; Ma, Z.
Cotton ACAULIS5 is involved in stem elongation and the plant defense response to Verticillium dahliae through thermospermine alteration
Plant Cell Rep.
34
1975-1985
2015
Gossypium hirsutum (A0A0C4JZU4), Gossypium hirsutum, Arabidopsis thaliana (Q9S7X6), Arabidopsis thaliana, Gossypium hirsutum Jimian 20 (A0A0C4JZU4), Arabidopsis thaliana Columbia-0 (Q9S7X6)
brenda
Milhinhos, A.; Prestele, J.; Bollhoener, B.; Matos, A.; Vera-Sirera, F.; Rambla, J.L.; Ljung, K.; Carbonell, J.; Blazquez, M.A.; Tuominen, H.; Miguel, C.M.
Thermospermine levels are controlled by an auxin-dependent feedback loop mechanism in Populus xylem
Plant J.
75
685-698
2013
Populus tremula x Populus tremuloides (T1PPE6)
brenda
Sekula, B.; Dauter, Z.
Crystal structure of thermospermine synthase from Medicago truncatula and substrate discriminatory features of plant aminopropyltransferases
Biochem. J.
475
787-802
2018
Medicago truncatula (G7K2D1)
brenda
Vuosku, J.; Muilu-Maekelae, R.; Avia, K.; Suokas, M.; Kestilae, J.; Laeaerae, E.; Haeggman, H.; Savolainen, O.; Sarjala, T.
Thermospermine synthase (ACL5) and diamine oxidase (DAO) expression is needed for zygotic embryogenesis and vascular development in Scots pine
Front. Plant Sci.
10
1600
2019
Pinus sylvestris (G8C849), Pinus sylvestris
brenda
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