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Xylans: Xiao Liu 10/22/2010

Xylans are a family of polysaccharides that make up hemicellulose in plant cell walls. This document summarizes research on xylan biosynthesis in Arabidopsis. It identifies two Golgi-localized glycosyltransferases, GUX1 and GUX2, that add glucuronic acid and methylglucuronic acid branches to the xylan backbone. Knocking out both GUX1 and GUX2 results in xylan with reduced substitution but unchanged backbone quantity. This simpler xylan structure is more easily extracted and broken down, suggesting it could improve biomass processing for biofuels. Altering crop xylan structure may thus facilitate sugar release from lignocellulose.

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Surya Negara
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127 views

Xylans: Xiao Liu 10/22/2010

Xylans are a family of polysaccharides that make up hemicellulose in plant cell walls. This document summarizes research on xylan biosynthesis in Arabidopsis. It identifies two Golgi-localized glycosyltransferases, GUX1 and GUX2, that add glucuronic acid and methylglucuronic acid branches to the xylan backbone. Knocking out both GUX1 and GUX2 results in xylan with reduced substitution but unchanged backbone quantity. This simpler xylan structure is more easily extracted and broken down, suggesting it could improve biomass processing for biofuels. Altering crop xylan structure may thus facilitate sugar release from lignocellulose.

Uploaded by

Surya Negara
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
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Download as PPTX, PDF, TXT or read online on Scribd
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Xylans

Xiao Liu
10/22/2010

Big picture
Other
Proteins proteins

Plan
t
cell
wall

GRPs

Structural
proteins

PRPs
HRGP Extens
s
in
AGPs
Cellulose

Polysaccharid
es

Pectin
Hemicellulo
se

Xylogluc
an
Xylans
Mannans,
Glucomann
ans

How do xylans affect our well-being?


Xylans can help reduce some diseases in hum
ans
Xylans are important functional ingredients in
baked products.
Xylans impact brewing properties of grains.
Xylans can be converted to xylitol, a natural f
ood sweetener.
Xylans are major constituents in the nonnutri
tional constituent of feed in monogastric ani
mals.

Introduction of Xylans
Xylans are a family of structurally divers
e plant polysaccharides with a backbone
composed of 1,4-linked -D-xylosyl resid
ues.

http://www.scientificpsychic.com/fitness/carbohydr
ates2.html

Major types of xylans


In almost all cases, the backbone is substi
tuted with monosaccharide or disaccharid
e side chains, to varying degrees.
glucuronic acid and 4-O-methyl glucuroni
c acid (glucuronoxylan, GX),
arabinose (arabinoxylan, AX),
a combination of acidic and neutral sugar
s (glucuronoarabinoxylan, GAX).

Arabinoxylan
Arabinoxylan is the predominant hemicellu
lose of grasses.
L-arabinofuranose attached randomly by 1
2 and/or 13 linkages to the xylose u
nits throughout the chain.
Side chains containing arabinosyl, galactos
yl, glucosyluronic acid, and 4-O-methyl gluc
osyluronic acid residues have been identifi
ed.

Arabinoxylan

http://www.btinternet.com/~martin.chaplin/hyara.h
tml

Glucuronoxylan
Major components of the secondary cell walls of
dicots (15%-30%):
(1,2)-linked D-glucuronyl (GlcA)
4-O-methyl-GlcA (MeGlcA) residues attached to
C-2 every 10 Xyl residues
70% contain one O-acetyl group at C-2 or C-3
.
Contain a distinct glycosyl sequence at the re
ducing end.
Devoid of Ara units

Glucuronoxylan
Two-domain architecture: a typical
polymer domain and a distinct
glycosyl sequence domain:

William S York and Malcolm A ONeill, Biochemical control of xylan biosynthesis


which end is up?

Glucuronoarabinoxylan (GAX)

Ara units are added to the O-3 position of the xylosyl units of the
backbone
Feruloyl groups are esterified to the O-5 position of the Ara units
in about every 50 units. (55 biphenyl bond forms cross-link
between two ferulic acid residues)

GAX in type II walls


The major cross-linking glycans of
the primary cell walls of
commelinoid monocots.
Type II walls :
cellulose microfibrils + crosslinking GAXs
Characteristics:
Pectin-poor matrix
Little structural protein
GlcA units contribute to charge
density (interconnecting)
GAXs:
1. Branched GAXs: cross-linking is
blocked (during elongation)
2. Unbranched GAXs: hydrogenbond to cellulose or to each
other

Xylan biosynthesis
Synthesized in Golgi Apparatus.
Backbone synthesis:
IRX9, IRX10, IRX10L, IRX14
Synthesis of side chain primer or terminat
or oligosaccharide:
IRX7, IRX8, PARVUS

GX biosynthesis
Serveral glycosyltransferases may be involv
ed in the initiation and elongation of the po
lymer backbone; other enzymes for the add
ition and /or modification of the side chain.
Five genes (FRA8, IRX8, IRX9, PARVUS, IRX1
4) in Arabidopsis have been identified to be
involved in GX synthesis. They encode putat
ive GTs that may have a role in forming red
ucing end.

Two models of GX biosynthesis mechani


sm

Model (a): GX is synthesized by transfer of xylosyl residues to the


reducing end of the chain.
Model (b): the glycosyl sequence acts as a primer; xylosyl residues are

GAX biosynthesis

Newly synthesizsed GAX polymers have re


gular structures
After treatment of
endoxylanase III, only three
kinds of oligosaccharides are
released
Possible explanation of the
formation of GAX fragments:
two unbranched Xyl residues at
the reducing end and one or
two unbranced Xyl residue at
the nonreducing end.
This type of regular structure is
usually the result of a
cooperative mechanism
between enzymes:
XylT, AraT, GlcAT.

AX biosynthesis
Xylosyltransferase and arabinosyl transfer
ase activities have been detected in micro
somal fractions isolated from wheat and
barley.

Why are we desired to know how Xylans


are synthesized ?
Contribution to the recalcitrance in biofuel
production.
In paper manufacture, decrease in brightne
ss of final product

MeGlcA
Hexenuronic Acid
In animal feed, loss of nutrition

Absence of branches from xyla


n in Arabidopsis gux mutants re
veals potential for simplificatio
n of lignocellulosic biomass
Jennifer C. Mortimer, Godfrey P. Miles, David M. Brown,
Zhinong Zhang, Marcelo P. Segura, Thilo Weimar, Xiaola
n Yu, Keith A. Seffeen, Elaine Stephen, Simon R. Turner,
and Paul Dupree

Identification of candidate secondary cel


l wall GTs

GT8 family

Subcellular localization of GUX1 and GUX


2

Knockout: gux1, gux2, gux1 gux


2

Morphology and Phenotype

Double mutant is slightly weaker than W


T

TEM:
secondary cell wall of
xylem fibers

Four-point bending test:


: WT stems
: gux1 gux2 stems

Xylan structure and quantity in stem of W


T and gux mutant plants.
AIR was characterized
with PACE using GH:
Decrease in intensity of
[Me]GlcA(Xyl)4

Quantification of Xyl residues substituted a


nd Xyl backbone

Monosaccharide analysis of WT and gu


x1 and gux2 stem

Xyl backbone unaltered; GlcA


reduced.

MALDI-TOF MS analysis of xylan structur


e

GuxT activity in WT and gux1 gux2

GuxT activity is strongly reduced


in the double mutant

The activity of XylT was


unaffected in the double mutant

GuxT activity in WT and gux1 gu


x2

Properties of WT and gux1 gux2 xylan

Double mutant shows improved

Properties of WT and gux1 gux2 xyl


an

GlcA/[Me]GlcA contribute to
solubility

gux1 gux2 could be hydrolyzed to


monosaccharide in the presence of

Conclusions
Identification of two Golgi-localized putative
glycosyltransferases, GUX1 and GUX2, that a
re required for the addition of both glucuro
nic acid and 4-O-methylglucuronic acid bran
ches to xylan in Arabidopsis stem cell wall.
gux1 gux2 double mutants show loss of xyla
n glucuronyltansferase activity and lack alm
ost all detectable xylan substitution, but no
change in xylan backbone quantity.

The xylan in gux1 gux2 shows improved e


xtractibility
Xylan chain extension and substitution ar
e not obligatorily coupled during synthesi
s.
Fermentable sugar release from lignocell
ulose can be increased by reducing xylan
branching.
Alterations in crop xylan structure could b
e a feasible goal for the bioprocessing ind
ustry.

Reference
Plant Cell Walls, Peter Albersheim, etc,
Biochemistry & Molecular Biology of Plants, B. Buchana
n, W. Gruissem, R. Jones,Eds.
Biochemical control of xylan biosynthesis- which end is
up? William S York, etc.
Xylan biosynthesis: News from the grass. Ahmed Faik.

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