BLBS102-c12 BLBS102-Simpson March 21, 2012 13:11 Trim: 276mm X 219mm Printer Name: Yet to Come
12 Pectic Enzymes in Tomatoes 233
Key:
Cellulose microfibril
Xyloglucans
Extensin
Arabinosides
RG I
with arabinogalactans
Junction zone
Pectins
Arabinose-rich runs
that open pores
Glucuronoarabinoxylan
Region of
H-bonding
β-Glucans
Cellotriosyl- and
cellotetraosyl-rich
Longer
cellodextrins
H-bonding
Phenolic
network
Ca^2 ++-linked
RG I
with arabinans
Figure 12.1.Current structural model of cell walls of flowering plants. (Reprinted from Buchanan et al. 2000).
be identified (Fig. 12.1), the cellulose–matrix glycan network,
consisting of cellulose microfibrils held together by matrix gly-
cans (formerly referred to as hemicelluloses), and the pectin
network (Carpita and Gibeaut 1993). The way by which these
polymers interact with each other is still unclear. Several cell
wall models are proposed in an attempt to explain measured
physical properties of the wall (Cosgrove 2000, Vincken et al.
2003, Thompson 2005). The cell wall provides shape and struc-
tural integrity to the cell, functions as a protective barrier to
pathogen invasion and environmental stress, and provides sig-
nal molecules important in cell-to-cell signaling, plant–symbiot,
and plant–pathogen interactions.
Pectin is an elaborate network of highly hydrated polysac-
charides rich in galacturonic acid (GalA). Pectins can be
classified in four major groups: homogalacturonan (HGA),
rhamnogalacturonan I (RGI), rhamnogalacturonan II (RGII),
and xylogalacturonan (XGA; Mohnen 2008). HGA is a ho-
mopolymer of 1,4-α-d-GalA (Fig. 12.2A) usually forming long
chains of approximately 100 GalA residues. Galacturonosyl
residues are esterified to various extents at the carboxyl group
with methanol and may be acetylated atO-2 orO-3. Non-
esterified regions of HGA chains can be associated with each
other by ionic interactions via calcium ion bridges, forming
egg-box-like structures (Fig. 12.2A). At least ten contiguous
unesterified galacturonyl residues are required in order to form
a stable cross link between HGA chains (Liners et al. 1992).
RGI is composed of repeating disaccharide units of 1,2-α-d-
rhamnose–1,4-α-d-GalA. Rhamnosyl residues on the backbone
of this polymer can be further substituted withβ-d-galactose-
andα-l-arabinose-rich side chains (Fig. 12.2B). RGII is a minor