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1,1,1-kestopentaose + H2O
?
Substrates: 18% activity compared to levan
Products: -
?
1,1,1-kestose + H2O
?
-
Substrates: 45% of the activity with levan
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?
1,1-kestose + H2O
?
-
Substrates: 39% of the activity with levan
Products: -
?
1,1-kestotetraose + H2O
?
1,6G-kestotetraose + H2O
1-kestotriose + fructose
-
Substrates: -
Products: -
?
6,6,6-kestopentaose + H2O
?
-
Substrates: -
Products: -
?
6,6-kestotetraose + H2O
?
6-kestose + H2O
sucrose + D-fructose
Substrates: as active as levan with native enzyme, 85% of the activity with levan with the heterologous enzyme
Products: -
?
6G,1,6-kestopentaose + H2O
fructose + 6G-kestotriose + ?
-
Substrates: -
Products: -
?
6G,6-kestotetraoase + H2O
fructose + 6G-kestotriose
-
Substrates: -
Products: -
?
6G-kestotriose + H2O
?
Substrates: -
Products: -
?
6G-kestotriose + H2O
sucrose + fructose
-
Substrates: -
Products: -
?
bacterial levan + H2O
?
Substrates: -
Products: -
?
fructan + H2O
?
Substrates: -
Products: -
?
fructan + H2O
D-fructose + ?
-
Substrates: Agave tequilana fructans
Products: -
?
garlic fructan + H2O
?
-
Substrates: 60% of the activity with levan
Products: -
?
graminan + H2O
?
-
Substrates: -
Products: -
?
inulin + H2O
D-fructose + ?
-
Substrates: -
Products: -
?
inulin + H2O
fructose + ?
levanbiose + H2O
beta-D-fructose
levansucrase + H2O
?
Substrates: Bacillus subtilis
Products: -
?
neokestose + H2O
?
-
Substrates: native enzyme shows 93% of the activity with levan
Products: -
?
sucrose + H2O
D-fructose + D-glucose
additional information
?
-
1,1-kestotetraose + H2O
?
Substrates: low activity
Products: -
?
1,1-kestotetraose + H2O
?
-
Substrates: 5fold lower activity than with 6,6-kestotetraose
Products: -
?
1,1-kestotetraose + H2O
?
Substrates: 18% activity compared to levan
Products: -
?
1,1-kestotetraose + H2O
?
-
Substrates: minor hydrolase activity (16% activity compared to 6-kestotriose)
Products: -
?
1-kestose + H2O
?
-
Substrates: native enzyme shows 12% of the activity with levan
Products: -
?
1-kestose + H2O
?
-
Substrates: 12% of the activity with levan
Products: -
?
1-kestotriose + H2O
?
Substrates: low activity
Products: -
?
1-kestotriose + H2O
?
Substrates: 7% activity compared to levan
Products: -
?
1-kestotriose + H2O
?
-
Substrates: minor hydrolase activity (8% activity compared to 6-kestotriose)
Products: -
?
6,6-kestotetraose + H2O
?
-
Substrates: -
Products: -
?
6,6-kestotetraose + H2O
?
Substrates: -
Products: -
?
6-kestose + H2O
?
Substrates: -
Products: -
?
6-kestose + H2O
?
-
Substrates: native enzyme shows 77% of the activity with levan
Products: -
?
6-kestotriose + H2O
?
Substrates: best substrate
Products: -
?
6-kestotriose + H2O
?
Substrates: 25% activity compared to levan
Products: -
?
6-kestotriose + H2O
?
-
Substrates: preferred substrate (100% activity)
Products: -
?
inulin + H2O
?
-
Substrates: 11.9% of the activity with 6G,6-kestotetraoase
Products: -
?
inulin + H2O
?
-
Substrates: 9% of the activity with levan
Products: -
?
inulin + H2O
?
Substrates: 6% activity compared to levan
Products: -
?
inulin + H2O
fructose + ?
-
Substrates: -
Products: -
?
inulin + H2O
fructose + ?
-
Substrates: 13% of the activity with levan
Products: -
?
inulin + H2O
fructose + ?
-
Substrates: 34% of the activity with levan
Products: -
?
levan + H2O
?
-
Substrates: -
Products: -
?
levan + H2O
?
-
Substrates: 8.5% of the activity with 6G,6-kestotetraoase
Products: -
?
levan + H2O
?
-
Substrates: -
Products: -
?
levan + H2O
?
Substrates: -
Products: -
?
levan + H2O
?
Substrates: from Erwinia herbicola
Products: -
?
levan + H2O
?
Substrates: 100% activity, the enzyme shows 6-FEH activity against levan (mean degree of polymerization is 20) that is 4fold higher than against 6-kestotriose(mean degree of polymerization is 3)
Products: -
?
levan + H2O
fructose + ?
-
Substrates: best substrate
Products: -
?
levan + H2O
fructose + ?
-
Substrates: -
Products: -
?
levan + H2O
fructose + ?
-
Substrates: exo-type mechanism
Products: -
?
levanbiose + H2O
beta-D-fructose
-
Substrates: native enzyme shows 63% of the activity with levan
Products: -
?
levanbiose + H2O
beta-D-fructose
Substrates: i.e. O-beta-D-fructofuranosyl-(2-6)-beta-D-fructofuranoside. 63% of the activity with levan with the native enzyme, 57% of the activity with levan with the heterologous enzyme
Products: -
?
neokestin + H2O
?
-
Substrates: -
Products: -
?
neokestin + H2O
?
-
Substrates: native enzyme shows 25% of the activity with levan
Products: -
?
phlein + H2O
?
-
Substrates: native enzyme shows 84% of the activity with levan
Products: -
?
phlein + H2O
?
Substrates: 95% of the activity levan with native enzyme, 120% of the activity with levan with the heterologous enzyme
Products: -
?
raffinose + H2O
?
-
Substrates: 12% of the activity with levan
Products: -
?
raffinose + H2O
?
-
Substrates: 5% of the activity with levan
Products: -
?
sucrose + H2O
D-fructose + D-glucose
Substrates: -
Products: -
?
sucrose + H2O
D-fructose + D-glucose
-
Substrates: 3% of the activity with levan
Products: -
?
sucrose + H2O
D-fructose + D-glucose
Substrates: in vitro incubations with sucrose as a single substrate show a prominent formation of 6-kestotriose compared to 1-kestotriose and 6G-kestotriose (neokestose) and the formation of 1- and 6-kestotetraose from sucrose and 1-kestotriose
Products: -
?
sucrose + H2O
D-fructose + D-glucose
Substrates: 15% activity with 100 mM sucrose and less than 1% activity with 1 mM sucrose compared to levan
Products: -
?
sucrose + H2O
D-fructose + D-glucose
-
Substrates: 21% of the activity with levan
Products: -
?
sucrose + H2O
D-fructose + D-glucose
-
Substrates: 9% of the activity with levan
Products: -
?
sucrose + H2O
D-fructose + D-glucose
-
Substrates: 2% activity compared to 6-kestotriose
Products: -
?
additional information
?
-
-
Substrates: no cleavage of inulin, dextran, sucrsoe, raffinose and melezitose
Products: -
?
additional information
?
-
-
Substrates: Agave tequilana fructans are a polydisperse mixture of long degree of polymerization (DP) polymers and fructooligosaccharides (FOSs) with both b(2-1) and b(2-6) linkages. This mixture also contains highly branched agavins and graminans, and agavins are the most abundant of all. Agave tequilana fructans exhibit fluctuation of their total reducing sugar content and DP according to the plant age
Products: -
?
additional information
?
-
-
Substrates: hydrolysis of terminal, non-reducing 2,6-linked beta-D-fructofuranose residues in fructans
Products: -
?
additional information
?
-
-
Substrates: no activity with sucrose
Products: -
?
additional information
?
-
-
Substrates: no activity with sucrose, 1,1-nystose and inulin
Products: -
?
additional information
?
-
Substrates: hydrolytic cleavage of terminal fructosyl residues off inulin
Products: -
?
additional information
?
-
-
Substrates: hydrolytic cleavage of terminal fructosyl residues off inulin
Products: -
?
additional information
?
-
-
Substrates: low ability to hydrolyze sucrose
Products: -
?
additional information
?
-
Substrates: purification of and assay with fructans from Lolium perenne, overview
Products: -
?
additional information
?
-
-
Substrates: purification of and assay with fructans from Lolium perenne, overview
Products: -
?
additional information
?
-
Substrates: the enzyme has significant activity against beta(2,1)-linked fructans, but considerably less than against beta(2,6)-linked fructans and shows weak invertase activity
Products: -
?
additional information
?
-
-
Substrates: less than 1% activity towards inulin and levan compared to 6-kestotriose (poor invertase activity)
Products: -
?
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evolution
the enzyme's the catalytic triad is conserved among glycoside hydrolase family 32, GH32, members
evolution
plant FEHs are classified into two types: 1-FEH (EC 3.2.1.153) and 6-FEH (EC 3.2.1.154), which hydrolyze a terminal of beta(2->1)-fructosyl linkage and beta(2->6)-fructosyl linkage, respectively
metabolism
6-FEHs are key enzymes associated with perennity of forage species
metabolism
model for fructan and primary carbohydrate metabolism in sink cells of perennial ryegrass, overview
metabolism
-
proposed model for the biosynthesis of fructooligosaccharides (FOSs) in Agave tequilana Weber Blue variety, overview
metabolism
regulation of the expression of FEH genes is a crucial factor for overwintering ability of fructan-accumulating cereals andgrasses. The regulation of the expression of FEH genes is a crucial factor for overwintering ability of fructan-accumulating cereals andgrasses. The coordinated expression of FEH genes in wheat is related to the regulation of water-soluble carbohydrate accumulation from autumn to early winter and fructan consumption under snow cover as well as energy supply. Wheat FEHs also play an important role in the varietal difference in freezing tolerance and snow mold resistance. Cooperative expression of 6-FEH and 1-FEH genes might be related to the seasonal changes and varietal difference in mono- and disaccharide contents
metabolism
-
6-FEHs are key enzymes associated with perennity of forage species
-
physiological function
-
the enzyme is associated with degradation of fructans in wheat leaf tissues during inoculation and incubation under snow cover
physiological function
the enzyme plays a role in the degradation of fructans and the mobilization of carbon sources for regrowth after defoliation in timothy
physiological function
Lolium perenne is a major forage grass species that accumulates fructans, mainly composed of beta(2,6)-linked fructose units. Fructans are mobilized through strongly increased activities of fructan exohydrolases (FEHs), sustaining regrowth following defoliation. Differences in the regulation of FEH activity among forage grasses influence their tolerance to defoliation
physiological function
absence of strong 6-FEH activity increases (on a fresh weight basis), especially in the lower parts, during the most critical period of the onset of fructan degradation and fructose accumulation under drought. The combined 1-FEH and 6-FEH activities are particularly important during the later stages in drought treated DH 338. FEH dynamics under drought may play a more essential role in var. DH 307 than in var. DH 338
physiological function
fructan exohydrolase, FEH, gene plays a key role in fructan metabolism associated with wintering ability, especially for snow mold resistance
physiological function
-
fructan metabolism in Agave tequilana exhibits changes in fructan content, type, degree of polymerization (DP), and molecular structure, overview. Analysis of the specific activities of involved vacuolar fructan active enzymes (FAZY) in Agave tequilana plants of different age and the biosynthesis of fructooligosaccharides (FOSs). Fructan hydrolysis is carried out by FEH enzymes, that remove terminal fructosyl units from fructan chains, to fulfill a diverse set of functions in the plant, such as energy supply during plant growth, maintenance of the osmotic pressure in the vacuoles, and modulation of the oligofructans amounts under oxidative stress and freezing tolerance
physiological function
fructans are polymers of fructose and one of the main constituents of water-soluble carbohydrates in forage grasses and cereal crops of temperate climates. Fructans are involved in cold and drought resistance, regrowth following defoliation and early spring growth, seed filling, they have beneficial effects on human health and are used for industrial processes. Fructan metabolism is under the control of both synthesis by fructosyltransferases (FTs) and breakdown through fructan exohydrolases (FEHs). The accumulation of fructans can be triggered by high sucrose levels and abiotic stress conditions such as drought and cold stress. The activities of enzymes involved in fructan synthesis and breakdown, the expression levels for the corresponding genes and levels for water-soluble carbohydrates are determined following pulse treatments with abscisic acid (ABA), auxin (AUX), ethylene (ET), gibberellic acid (GA), or kinetin (KIN)
physiological function
-
Lolium perenne is a major forage grass species that accumulates fructans, mainly composed of beta(2,6)-linked fructose units. Fructans are mobilized through strongly increased activities of fructan exohydrolases (FEHs), sustaining regrowth following defoliation. Differences in the regulation of FEH activity among forage grasses influence their tolerance to defoliation
-
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Igarashi, T.; Takahashi, M.; Yamamoto, A.; Etoh, Y.; Takamori, K.
Purification and characterization of levanase from Actinomyces viscosus ATCC19246
Infect. Immun.
55
3001-3005
1987
Actinomyces viscosus
brenda
Burne, R.A.; Schilling, K.; Bowen, W.H.; Yasbin, R.E.
Expression, purification, and characterization of an exo-beta-D-fructosidase of Streptococcus mutans
J. Bacteriol.
169
4507-4517
1987
Streptococcus mutans, Streptococcus mutans GS-5
brenda
Igarashi, T.; Yakamoto, A.; Goto, N.
Characterization of an exo-beta-D-fructosidase from Streptococcus mutants ingbritt
Microbiol. Immunol.
36
643-647
1992
Streptococcus mutans
brenda
Henson, C.A.; Livingston III, D.P.
Purification and characterization of an oat fructan exohydrolase that preferentially hydrolyzes beta-2,6-fructans
Plant Physiol.
110
639-644
1996
Avena sativa
brenda
Van den Ende, W.; De Coninck, B.; Clerens, S.; Vergauwen, R.; Van Laere, A.
Unexpected presence of fructan 6-exohydrolases (6-FEHs) in non-fructan plants: characterization, cloning, mass mapping and functional analysis of a novel 'cell-wall invertase-like' specific 6-FEH from sugar beet (Beta vulgaris L.)
Plant J.
36
697-710
2003
Beta vulgaris
brenda
Paludan-Muller, C.; Gram, L.; Rattray, F.P.
Purification and characterisation of an extracellular fructan beta-fructosidase from a Lactobacillus pentosus strain isolated from fermented fish
Syst. Appl. Microbiol.
25
13-20
2002
Lactiplantibacillus pentosus, Lactiplantibacillus pentosus B235
brenda
Bonnett, G.D.; Simpson, R.J.
Fructan exohydrolase activities from Lolium rigidum that hydrolyze ??2, 1? and ??2, 6?glycosidic linkages at different rates
New Phytol.
131
199-209
1995
Lolium rigidum
brenda
Marx, S.P.; Nosberger, J.; Frehner, M.
Hydrolysis of fructan in grasses: a beta-(2-6)-linkage specific fructan-beta-fructosidase from stubble of Lolium perenne
New Phytol.
135
279-290
1997
Lolium perenne
brenda
Mller, M.; Seyfarth, W.
Purification and substrate specificity of an extracellular fructanhydrolase from Lactobacillus paracasei ssp. paracasei P 4134
New Phytol.
136
89-96
1997
Lacticaseibacillus paracasei
brenda
Van Riet, L.; Nagaraj, V.; Van den Ende, W.; Clerens, S.; Wiemken, A.; van Laere, A.
Purification, cloning and functional characterization of a fructan 6-exohydrolase from wheat (Triticum aestivum L.)
J. Exp. Bot.
57
213-223
2006
Triticum aestivum (Q2UXF7)
brenda
De Coninck, B.; Le Roy, K.; Francis, I.; Clerens, S.; Vergauwen, R.; Halliday, A.M.; Smith, S.M.; Van Laere, A.; van den Ende, W.
Arabidopsis AtcwINV3 and 6 are not invertases but are fructan exohydrolases (FEHs) with different substrate specificities
Plant Cell Environ.
28
432-443
2005
Arabidopsis thaliana
-
brenda
Le Roy, K.; Lammens, W.; Van Laere, A.; Van den Ende, W.
Influencing the binding configuration of sucrose in the active sites of chicory fructan 1-exohydrolase and sugar beet fructan 6-exohydrolase
New Phytol.
178
572-580
2008
Beta vulgaris (Q70XE6), Beta vulgaris
brenda
Tamura, K.I.; Sanada, Y.; Tase, K.; Komatsu, T.; Yoshida, M.
Pp6-FEH1 encodes an enzyme for degradation of highly polymerized levan and is transcriptionally induced by defoliation in timothy (Phleum pratense L.)
J. Exp. Bot.
62
3421-3431
2011
Phleum pratense (E9RGV6)
brenda
Kawakami, A.; Yoshida, M.
Graminan breakdown by fructan exohydrolase induced in winter wheat inoculated with snow mold
J. Plant Physiol.
169
294-302
2012
Triticum aestivum
brenda
Van den Ende, W.; Coopman, M.; Clerens, S.; Vergauwen, R.; Le Roy, K.; Lammens, W.; Van Laere, A.
Unexpected presence of graminan- and levan-type fructans in the evergreen frost-hardy eudicot Pachysandra terminalis (Buxaceae): purification, cloning, and functional analysis of a 6-SST/6-SFT enzyme
Plant Physiol.
155
603-614
2011
Pachysandra terminalis (E3PQS3)
brenda
Lothier, J.; Van Laere, A.; Prudhomme, M.P.; Van den Ende, W.; Morvan-Bertrand, A.
Cloning and characterization of a novel fructan 6-exohydrolase strongly inhibited by sucrose in Lolium perenne
Planta
240
629-643
2014
Lolium perenne (D0PQE8), Lolium perenne
brenda
Gasperl, A.; Morvan-Bertrand, A.; Prudhomme, M.P.; van der Graaff, E.; Roitsch, T.
Exogenous classic phytohormones have limited regulatory effects on fructan and primary carbohydrate metabolism in perennial ryegrass (Lolium perenne L.)
Front. Plant Sci.
6
1251
2015
Lolium perenne (D0PQE8)
brenda
Zhang, J.; Chen, W.; Dell, B.; Vergauwen, R.; Zhang, X.; Mayer, J.E.; Van den Ende, W.
Wheat genotypic variation in dynamic fluxes of WSC components in different stem segments under drought during grain filling
Front. Plant Sci.
6
624
2015
Triticum aestivum (Q2UXF7)
brenda
Meguro-Maoka, A.; Yoshida, M.
Analysis of seasonal expression levels of wheat fructan exohydrolase (FEH) genes regulating fructan metabolism involved in wintering ability
J. Plant Physiol.
191
54-62
2016
Triticum aestivum (Q2UXF7)
brenda
Mellado-Mojica, E.; Gonzalez de la Vara, L.E.; Lopez, M.G.
Fructan active enzymes (FAZY) activities and biosynthesis of fructooligosaccharides in the vacuoles of Agave tequilana c variety plants of different age
Planta
245
265-281
2017
Agave tequilana
brenda