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Plant Cell Walls

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Plant Cell Walls
Vaibhav Srivastava, Division of Glycoscience, School of Biotechnology, KTH
Royal Institute of Technology, Stockholm, Sweden
Lauren S McKee, Division of Glycoscience, School of Biotechnology, KTH Royal
Institute of Technology, Stockholm, Sweden
Vincent Bulone, Division of Glycoscience, School of Biotechnology, KTH Royal
Institute of Technology, Stockholm, Sweden
Based in part on the previous versions of this eLS article ‘Plant Cell
Walls’ (2012).
Common to all plant species, the cell wall is the
tough outer coat that protects the plant cell. The
cell wall is mostly carbohydrate-based, compris-
ing three major classes of polysaccharides: cel-
lulose, hemicellulose and pectin. There are also
important structural proteins as well as pheno-
lic and aliphatic polymers. The cell wall provides
mechanical strength to the plant body, allowing
upright growth and structure formation, and also
plays important roles in cellular processes such
as cell expansion, tissue differentiation, intercellu-
lar communication, water movement and defence
responses against pests or pathogens. Cell walls
may even be involved in signal sensing during pat-
tern formation in plant development.
Introduction
The plant cell wall is a strong external layer of the plant cell,
located outside the plasma membrane. The cell wall is thick,
complex and dynamic and acts as a semipermeable barrier to
physically constrain and protect the cell. The wall comprises
polysaccharides, highly glycosylated proteins and lignin. The
precise composition of the wall is highly variable, depending on
plant species, tissue and developmental stage. Typically, plants
comprise around 35 cell types, each of which has a characteris-
tic size, shape, position and wall composition (Cosgrove, 2005).
Compared to cell membranes, which are <0.01 μm thick, cell
walls range from ∼0.1 μm to over 10 μm in thickness. The cell
eLS subject area: Plant Science
How to cite:
Srivastava, Vaibhav; McKee, Lauren S; and Bulone, Vincent (July
2017) Plant Cell Walls. In: eLS. John Wiley & Sons, Ltd:
Chichester.
DOI: 10.1002/9780470015902.a0001682.pub3
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
• Introduction
• General Morphology: Cell Wall Proper, Middle
Lamella
• Cell Wall Architecture: Primary and Secondary
Walls
• Major Components and Their Properties
• Macromolecular Organisation of the Wall Matrix
• Cell Wall Expansion
• Regulation of Cell Wall Expansion
• Functions of Cell Walls
Online posting date: 17th July 2017
walls of living plants play diverse and subtle roles in plant growth,
development and defence. And even after the death of the proto-
plast, the wall will continue to serve its particular biological role
for many years. This can be seen clearly in the way that cork
(the outer layer of bark) covers and protects the tree trunk for the
lifespan of the tree. See also: Plant Cell: Overview
This article considers the cell walls of seed plants only. The
walls of ferns, liverworts and charophycean green algae do resem-
ble those of seed plants but have been investigated in less detail.
General Morphology: Cell Wall
Proper, Middle Lamella
The cell wall is composed of a network of cellulose microfibrils
embedded in a matrix of other carbohydrates, proteins and other
polymers. The long cellulose microfibrils (see below), which lie
in the plane of the cell surface, form the rigid scaffolding of
the cell wall. These microfibrils are typically 4–10 nm thick and
spaced roughly 30 nm apart. Many plant cells are roughly cylin-
drical, with the surrounding microfibrils deposited in parallel to
each other so as to form cyclical ‘hoops’ around the body of the
cell (Figure 1). This microfibrillar arrangement dictates the direc-
tion in which the cell can expand, as will be discussed. See also:
Cellulose: Structure and Distribution
New cell walls are usually formed soon after cellular mitosis
at a location that divides the mother cell into two daughter
cells. The location of the new wall is directed by a cluster of
microtubules, known as the phragmoplast. Indeed, the deposition
of cellulose microfibrils in general is thought to be directed by
the microtubules (see the following discussion). See also: Plant
Mitosis, Cytokinesis and Cell Plate Formation
The thickness of a cell wall can be increased by the depo-
sition of new microfibrils on the inner face. Walls can also
expand laterally, allowing cellular expansion for plant growth
and development. This can be anisotropic (unidirectional) to
produce cylindrical cells, isotropic (roughly equal expansion
in all directions to produce spherical cells, e.g. in potato tuber
parenchyma) or irregular (e.g. in spongy mesophyll cells). In
the actively growing cell wall, the structure of the noncellulosic
1
 Plant Cell Walls
Figure 1 Model showing the arrangement of cellulose microfibrils in the walls of a plant cell. Microfibrils are inextensible, so this spiral-like arrangement
dictates that the cylindrical cell will expand predominantly in one dimension, as shown. Many of the cells in an elongating stem or root conform to this
model.
polysaccharides is modified by a cocktail of enzymes, which
affect polysaccharide interactions with each other and with
cellulose. The resulting ‘loosening’ of the cell wall matrix allows
for an increase in cellular volume (see the following discussion).
The cessation of cell expansion is linked with a reduced expres-
sion of wall-loosening genes and changes in polysaccharides,
including a tightening of the cellulose–matrix network. In some
anisotropically growing cells, wall expansion occurs uniformly
over the area of the side walls (e.g. in the parenchyma and
epidermis of a stem or coleoptile); in others (e.g. fibre cells,
pollen tubes and root hairs), it is confined to the tip(s). See also:
Plant Cell Growth and Elongation
The inner face of the cell wall is usually in contact with the
plasma membrane, while the outer face contacts a neighbouring
wall. Many proteins that are important for cell wall biosynthe-
sis are located at the plasma membrane, allowing newly formed
polysaccharide chains to be easily delivered to the wall. How-
ever, most of the cell wall area is not directly attached to the
underlying plasma membrane. During plasmolysis induced by
osmotic stress, the membrane of the shrinking protoplast read-
ily withdraws from the wall. By comparison, adhesion between
neighbouring cell walls is much stronger. This tissue cohesion is
imparted by the middle lamella, a glue-like layer between cells
which is laid down before the primary cell wall. The middle
lamella is highly enriched in acidic gel-forming pectins, con-
tributing to its adhesive properties. However, the application of
pectin-degrading enzymes is often not sufficient to separate plant
cells, indicating that there are other factors involved in adhesion.
See also: Plant Plasma Membrane; Pectic Substances
In some tissues (e.g. mesophyll, cortex and especially
aerenchyma), there is a large air-filled space between neigh-
2 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
bouring cells, whereas others (e.g. meristems, the stele and
collenchyma) tend not to have any obvious air spaces. The
formation of air spaces between a pair of sister cells involves
the partial disintegration of middle lamella and primary wall
material. Similarly, many tissue types display plasmodesmata.
These are channels through the cell walls of adjacent cells, which
allow signals and metabolites to be sent between cells. These
are lined with specific polysaccharides which give additional
strength to the local wall area and may contain different proteins
than the main wall. See also: Plasmodesmata
Cell Wall Architecture: Primary
and Secondary Walls
Primary cell walls
The wall that surrounds cells that are actively growing or capable
of growth is called the primary cell wall. The cellulose microfib-
rils of the primary cell wall are deposited during this time of
potential growth. After deposition, the primary wall may lose its
ability to grow owing to an increase in rigidity or wall thick-
ness after the addition of other layers (see below). Thus, all
growing walls are primary, but not all primary walls can grow.
Primary walls are often approximately 0.1 μm thick, but col-
lenchyma primary walls can be up to 10 μm thick. Additional
components such as lignin, cutin, suberin and extensin may in
some cases be inserted between the microfibrils of an existing
primary wall, but no additional microfibrils will be added. See
also: Collenchyma

Secondary cell walls
Once the cell stops growing, additional wall material is deposited
within the primary wall and is therefore in contact with the inner
face of the primary wall and the outer face of the plasma mem-
brane. This new layer constitutes the secondary cell wall. In
some cases, the secondary wall will comprise up to three lay-
ers, termed S1 , S2 and S3 , in which the cellulose microfibrils
are deposited with different orientations. Secondary walls are
much thicker than primary walls, providing significant additional
strength but reduced flexibility. Only those cells that require addi-
tional mechanical strength and structural reinforcement will typi-
cally form secondary walls. Secondary walls of fibre cells can be
10–20 μm thick and can take up most of the volume of the cell.
Secondary walls are critical for the vertical transport of water and
nutrients and to allow upright growth. As such, xylem vessels and
tracheids are heavily reinforced with secondary cell wall thicken-
ings and account for a significant proportion of the mass of mature
wood. Secondary walls have a different carbohydrate composi-
tion than primary walls, and many are further strengthened by
the incorporation of lignin, which also acts as a waterproofing
agent (Barros et al., 2015). Lignin may be deposited in rings
around the cell wall, in helical spirals, or as pitted layers over the
whole wall to prevent vessel collapse during transpiration. See
also: Secondary Cell Walls
Architecture
Cell walls from different plant species, tissues and developmental
stages differ in thickness, composition and microfibrillar orienta-
tion. Cells of the structural tissues collenchyma, sclerenchyma,
tracheary elements (tracheids and vessels), fibres and cork cells
have especially thick walls, as they require additional physical
strength. The outer facing cell wall of stem and leaf tissue is
thicker than the inner facing wall, owing to the contact with
the external atmosphere. The outer walls are also optimised to
minimise water loss and exclude pathogens (see the following
discussion). The guard cells which surround stomata have asym-
metrical thickness of the secondary wall, so that modulations
in turgor pressure will allow the pores to open and close for
gas exchange. Seeds also tend to have very thick walls, which
may contain fine polysaccharide structures not found elsewhere
in a plant. These are used as energy reserve carbohydrates and
are enzymatically hydrolysed after germination to provide sugar
to the young plant. See also: Cork; Meristems; Parenchyma;
Sclerenchyma; Seeds; Xylem Structure and Function; Plant
Storage Products (Carbohydrates, Oils and Proteins)
The orientation of cellulose microfibrils in the primary cell wall
is controlled by cortical microtubules connected to the cellulose
synthase complex (CSC) and is used to control cell expansion.
Cellular expansion generally proceeds in a direction perpendicu-
lar to the microfibrillar orientation (Panteris et al., 2015). Treat-
ment of dwarf pea plants with the plant hormone gibberellic acid
causes new microfibrils to be deposited mainly perpendicular to
the long axis of the stem; the cells then grow in length, leading to
a tall, thin stem. Treatment with ethylene causes new microfibrils
to be deposited more randomly; the cells then balloon, and the
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Plant Cell Walls
stem swells in diameter. See also: Plant Growth Factors and
Receptors
Major Components and Their
Properties
The dry matter of the primary cell wall comprises around 90%
polysaccharides and 10% glycoproteins (see the following dis-
cussion). The carbohydrate components are broadly classified as
cellulose, hemicellulose and pectin. Their structures and com-
ponents are summarised in Table 1. Secondary cell walls also
contain significant amounts of lignin (see the following discus-
sion).
Polysaccharides
Cellulose
The major load-bearing carbohydrate component of cell walls is
cellulose, the most abundant organic compound on the planet.
Cellulose is a linear polymer of D-glucose [I] monosaccharides
found in all primary plant cell walls, comprising around 50%
of the total mass of wood and around 90% of cotton. Cellulose
(glucan) chains aggregate after synthesis and are connected by
hydrogen bonds to form strong bundles known as microfibrils.
Cellulose is insoluble in water and most organic solvents and can
be collected as the residue after alkali extraction of hemicellu-
loses. The major industrial uses of cellulose are in traditional
areas such as pulp and paper production and textile manufac-
turing. Cellulose nanofibres (CNF) and cellulose nanocrystals
(CNC) are also used in the production of more modern materials
such as functional nanopapers. Cellulose can also be hydrolysed
by chemical or enzymatic means to glucose, which can be fer-
mented to ethanol for fuel. Cellulose microfibrils can exceed
100 μm in length, far longer than any individual glucan chain
(typically 1–5 μm), so there tend to be multiple chain termini
along the length of a microfibril. The glucan chains within a
microfibril lie parallel to each other, with all reducing ends point-
ing towards the same fibrillary end. See Guerriero et al. (2010) for
a review of cellulose structure and synthesis. See also: Cellulose:
Biogenesis and Biodegradation; Cellulose: Structure and Dis-
tribution
CH2OH
O OH
OH
HO
OH
[I]
β-D-Glucose
Cellulose in plants is synthesised by the CSC in the plasma
membrane. The CSC appears to comprise 36 individual enzymes,
each of which may be actively synthesising a glucan chain. A
microfibril formed by this complex might therefore be expected
to comprise 36 glucan chains, but the measured dimensions
3
 Plant Cell Walls

Table 1 Dominant polysaccharides and structural proteins of plant cell walls

Polymer Amount (%)a Chargeb Major components Notes
Polysaccharides
Microfibrillar
Cellulose Matrix 30 0 β-Glc Linear (1→4) glucan chain

Pectins
Homogalacturonan (HG) 16 – α-GalA Linear (1→4) GalA chain. Cleaved by
EPG. Some modifications with
methylester or O-acetyl groups, which
hinder enzymatic hydrolysis of the
backbone
Rhamnogalacturonan I 10 – α-GalA, α-Rha, β-Gal, α-Araf, Backbone of
(RGI) β-Arap (−GalA-(1→4)-Rha-(1→2)–). Half of
the Rha residues are decorated with
extended chains of arabinan, galactan
or arabinogalactan, which can be
cross-linked. Backbone GalA residues
are O-acetylated. Not cleaved by EPGc
Rhamnogalacturonan II 4 – α & β-GalA, α & β-Rha, α-Gal, HG backbone with highly complex side
(RGII) α-Fuc, α-Arap, β-Araf, β-Apif, chains. See Figure 2. Some Fuc and
β-GlcA, KDO, β-Acef A, all Xyl residues are 2-O-methyl ethers.
α-Xyl and β-DHA Some Apif residues are esterified with
borate. Not cleaved by EPGc
Apiogalacturonan (AG) ± – α-GalA, Apif HG backbone with (1→2) and/or (1→3)
linked Apif
Hemicelluloses
Xyloglucans 20 0 β-Glc, α-Xyl, β-Gal, α-Fuc, Mostly found in primary cell walls or as
α-Araf and β-Xyl seed energy storage polysaccharide.
Backbone of (1→4) Glc, regularly
decorated with (1→6) Xyl, which can
be further decorated with Gal, Fuc
and/or Ara, depending on the tissue
and species (Figure 3)
Xylans 8 – β-Xyl, α-Araf and α-GlcA Dominant secondary wall hemicellulose
in hardwood trees, less important in
primary walls. Backbone of (1→4)
Xyl, with (1→2) and/or (1→3) linked
Araf and/or (1→2) linked GlcA. Often
heavily Xyl O-acetylated (Figure 3)
Mannans ± 0 β-Man, β-Glc and α-Gal Dominant secondary wall hemicellulose
in softwood trees, not common in
primary walls. Storage polysaccharide
in some seeds. Backbone of (1→4)
linked Man or alternating Man and
Glc, with (1→6) linked Gal
substitutions. Often heavily Man
O-acetylated
Mixed-linkage glucans ±∼ 0 β-Glc Only found in graminaceous monocots.
(MLGs) Linear polymer of (1→4) and (1→3)
linkages in different ratios. Energy
storage polysaccharide in some grains
Callose ± 0 β-Glc Linear polymer of exclusively (1→3)
linkages
(continued overleaf )

4 eLS © 2017, John Wiley & Sons, Ltd. www.els.net

 Plant Cell Walls

Table 1 (continued)
Polymer Amount (%)a Chargeb Major components Notes
Proteins
Extensins ± + 40–50% protein: Hyp, Ser, Lys, Basic polypeptide backbone.
Tyr, Val and His Tetrasaccharide (αAraf-(1→3)-βAraf-
50–60% sugar: β and α-Araf, (1→2)-βAraf-(−1→2)-βAraf–)
α-Gal O-linked to most Hyp residues. Single
Gal attached to some Ser residues
Arabinogalactan ± – 2–10% protein: Hyp, Ser, Asp, Short acidic polypeptide backbone.
proteins (AGPs) Thr and Gly Polysaccharide groups (rich in (1→3)
90–98% sugar: β-Gal, and (1→6)-linked Gal) O-linked to
α-Araf, GlcA Hyp. Some AGPs are covalently
attached to lipids
Proline-rich proteins ± + Pro, Hyp, Val, Tyr and Lys Hyp:Pro ratio approximately 1:1. May be
(PRPs) covalently cross-linked, possibly via
Tyr residues. Little or no sugar.
Sometimes associated with lignin
polymer
Glycine-rich proteins ± – Gly; some Ser, Ala, etc. 60–70% Gly. Not glycosylated.
(GRPs) Sometimes associated with lignin
polymer
a Rough guide to amount of polymer present, as percentage of dry weight of a typical dicot primary cell wall from a rapidly growing cell culture. ±,

not always present; ∼, amount varies greatly.

b Charge on polymer molecule (at physiological pH): −, negative; +, positive and 0, uncharged.
c These pectin domains are not cleaved by EPG but can be released from cell walls by EPG.

Ala, L-alanine; Api, D-apiose; Ara, L-arabinose; Asp, L-aspartic acid; D- and L-, optical isomers; DHA, 3-deoxy-2-D-heptulosaric acid; EPG,
endo-polygalacturonase enzyme; f, furanose ring form; Fuc, L-fucose; Gal, galactose (D unless otherwise stated); GalA, D-galacturonic acid; Glc,
D-glucose; GlcA, D-glucuronic acid; Gly, glycine; His, L-histidine; Hyp, L-hydroxyproline; KDO, D-ketodeoxyoctulosonic acid; Lys, L-lysine; Man,
D-mannose; O-, via oxygen atom; p, pyranose ring form; Pro, L-proline; Rha, L-rhamnose; Ser, L-serine; Thr, L-threonine; Tyr, L-tyrosine; Val, L-valine
and Xyl, D-xylose. All sugar residues are in the pyranose ring form unless indicated.

of cellulose microfibrils suggest that typically 18–24 chains region of pectin, comprising a glycosidically linked linear chain
are produced by a CSC. See Carpita (2011) and Nixon et al. of D-galacturonic acid residues [II]. Rhamnogalacturonan-I
(2016) for a summary of plant cell wall biosynthesis. See also: (RGI) comprises repeating units of [II] and rhamnose [III],
Glycosyltransferases in Plant Cell Wall Synthesis decorated with long chains of arabinan and arabinogalactan.
Finally, rhamnogalacturonan II (RGII) comprises an HG back-
bone with four long, elaborate and highly conserved side chains
Pectins named A–D (Pabst et al., 2013) (Figure 2). See Mohnen (2008)
Pectic polysaccharides are found in primary cell walls and are for a review of pectin structure and function, and see Harholt
highly enriched in the middle lamella. Owing to the very high et al. (2010) for an update on pectin biosynthesis. See also:
degree of branching (Figure 2), extracted pectins are highly Pectic Substances; Polysaccharides; Polysaccharides: Plant
soluble in water. In the cell wall, pectin forms an extensively Noncellulosic; Plant Gums
cross-linked network in which other polymers are embedded. COOH
The pectin network has adhesive properties and is strong and
HO O HO O OH
flexible. Pectin has long found use in food manufacturing, in
jam production especially, as it complexes with ions of metals OH CH3
such as calcium to form a strong gel. In the cell wall, borate ions OH
serve this purpose. Pectins can therefore be partially extracted
from cell wall material simply using cold water and a chelating OH OH OH
agent such as ethylenediaminetetraacetic acid (EDTA). More [II] [III]
efficient extraction requires the use of hot water or an enzyme α-D-Galacturonic acid α-L-Rhamnose
(polygalacturonase, EPG (endo-polygalacturonase enzyme)),
but these will cleave the galacturonan backbone and reduce
molecular weight. Broadly, three classes of pectins have been Hemicelluloses
defined, which may form separate domains of a continuous poly- The term ‘hemicellulose’ refers to the noncellulosic, nonpectic
meric chain (Table 1). Homogalacturonan (HG) is the ‘smooth’ polysaccharides found in primary and secondary cell walls. See

eLS © 2017, John Wiley & Sons, Ltd. www.els.net 5

 Plant Cell Walls

(a) -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)-

-α(1 5)-
-α
(1
4)
-
-α(1 2)- -β Ar

-α(1 5)-
(1
ab
4) in
- og
Arabinan

-α(1 3)- -β al

-
(1 ac

3)
ta
-α(1 3)-

-α(1 5)-
4) n

(1
-

-α
-β

-
(1

5)
4)

(1
-

-α
-α(1 5)-

-β

-
(1

3)
4)

(1
-

-α
-α -β
(1 (1
(b) 4) 4)
- -
-α -α -α -α
(1 (1 (1
Backbone

(1

-
-
-
-

2)
2)
2)
2)

4) 4) 4) 4)

(1
(1
-
(1
(1

- - -

-α
-α
-α
-α

-α -α -α -α
(1 (1 (1 (1

-
-
-
-

4)
4)
4)
4)

2) 2) 2) 2)

(1
(1
(1
(1

- - - -

-α
-α
-α
-α

(c) -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)- -α(1 4)-
-β(1 2)-

-α(2 3)-

-β(2 3)-
-β(1 2)-
-β(1 3)-

-β(1 3)-

-α(1 5)-

-α(1 5)-
-β(2 1)-
-α(2 1)-
Side chains

-α(3
-α(1 3)-
-α(1 4)-

1)-
C D
-β(2 1)-
-α(1 4)-

(Gal replaces Fuc

in some species) -α(3
-β(1 4)-

1)-
-β(2 1)-
-α(2 1)-
-α(3
-α(2 1)- 1)-

A B

D-Galactopyranose L-Fucopyranose D-Glucuronic acid

D-Galacturonic acid Possible acetylation Possible methylation
Possible acetylation Possible methylation
Possible methylation

D-Xylopyranose L-Aceric acid L-Arabinofuranose

Possible methylation Acetylation

L-Arabinopyranose D-Apiofuranose L-Rhamnopyranose

3-Deoxy-D-manno- 3-Deoxy-D-lyxo2-
octulopyranosylonic acid (KDO) heptulopyranosaric acid (DHA)

Figure 2 Schematic models for pectic polymers. (a) Homogalacturonan can be extensively methyl-esterified and acetylated. (b) Rhamnogalacturonan I has
long arabinan and arabinogalactan side branches, which can be interlinked. (c) Rhamnogalacturonan II has highly complex side chains which show some
limited variability between plant species.

6 eLS © 2017, John Wiley & Sons, Ltd. www.els.net

 Plant Cell Walls

Scheller and Ulvskov (2010) for an in-depth review of hemi- although there is evidence for some repeated patterning in xylan
cellulose structure, function and synthesis. In general, hemicel- decoration (Bromley et al., 2013) (Figure 3).
luloses have β-(1→4)-linked backbones of glucose [I], xylose The backbone structure and molecular weight of a hemicel-
[IV] and mannose [V], extensively decorated with arabinose [VI], lulose, as well as the degree of substitution, affect the structural
galactose [VII], fucose (6-deoxygalactose) and/or glucuronic properties of the polymer. Hemicelluloses can be extracted from
acid [VIII]. All hemicelluloses may also be decorated with non- cell walls by aqueous alkali treatment, such as soaking in 6 M
sugar components, including methyl esters [IX] and ethers [X], NaOH, but this will cause the loss of ester-linked side groups.
acetyl esters [XI] and feruloyl esters [XII]. Extraction in hot water is preferable where native substitutions
should remain intact, such as in the production of materials
with specific properties. For example, the strength and flexibility
O OH
of films produced from xylan are dependent on the degrees of
O OH CH2OH arabinosylation and acetylation. See also: Polysaccharides;
OH Plant Storage Products (Carbohydrates, Oils and Proteins);
OH O OH
H 2C Plant Gums
HO OH HO The fine structure of hemicelluloses also influences how cell
HO OH
OH wall components interact with each other in the wall. Xyloglu-
HO
[VI] can in particular is known to interact very strongly with cellulose,
[IV] [V] α-L-Arabinose forming hydrogen bonds between their respective β-glucan back-
β-D-Xylose β-D-Mannose [furanose form] bone chains. The molecular weight of the polymer and the nature
of the xyloglucan side groups have a significant influence on the
CH2OH COOH strength of the xyloglucan–cellulose interaction (Lopez et al.,
OH O
2010). Xylan also interacts with cellulose, and this is affected by
HO O
the degree and spacing of substitutions on the xylan backbone
OH OH (Simmons et al., 2016).
HO OH Callose is another linear glucan polymer, comprising
β-(1→3)-linked D-glucose [I]. Callose is not a permanent
OH OH
cell wall component but is produced at a forming cell plate
[VII] [VIII] during cytokinesis; other polymers may then be laid down upon
β-D-Galactose α-D-Glucuronic acid this scaffold. Callose is also deposited in response to wounding,
infection or stress, to temporarily reinforce the wall. Callose
COOH COOH synthase activity is induced rapidly around the site of fungal
O O CH3
C O HO O penetration to resist fungal attack. The cell wall surrounding
HO O OH plasmodesmata is also enriched in callose. See also: Callose and
OH Related Glucans
OH OH
O OH
OH H 3C C O Cell wall proteins
CH3 OH
OH O
[IX] [X] [XI] Cell wall proteins are important structural and functional
Methyl α-D- 4-O-Methyl α-D- 2-O-Acetyl-α-D- constituents of plant cell walls. They play important roles in
galacturonate glucuronic acid galacturonic acid modifying cell wall structure and its components and in sig-
nalling events such as response to biotic and abiotic stress. Most
O OH cell wall proteins are highly glycosylated and interact extensively
with other cell wall components. Proteomic experiments to iden-
OH tify all proteins present at the cell wall are highly challenging;
H2C proteins can be destroyed during cell wall preparation, or they
O O CH3 may be difficult to extract owing to strong covalent linkages with
OH
C other cell wall components. Nonetheless, more than 2000 dif-
O OH ferent cell wall proteins have been identified in different plants,
including dicot and monocot species. The classes of proteins
typically identified in cell walls are hydrolases, proteases, oxi-
[XII]
5-O-Feruloyl- doreductases, signalling proteins and structural proteins (Jamet
α-L-arabinose et al., 2008; Albenne et al., 2014).
One of the best studied groups of cell wall structural pro-
teins is the extensins, which are rod-shaped, strongly basic,
Xyloglucan, which has a linear β-(1→4)-linked glucan back- tightly cross-linked glycoproteins. They are rich in hydroxypro-
bone, is heavily decorated, with regular repeating patterns of side line residues, which are the site for the extensive O-glycosylation.
groups (Schultink et al., 2014) (Figure 3). Substitutions on xylan Extensins are involved in cell wall assembly, the control and
and mannan polysaccharides are thought to be more random, maintenance of cell shape and cell growth. Some knockout

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8
Plant Cell Walls

(a)

-α(1 2)-
-β(1 2)- - β(1 2)-

-α(1 6)-

-α(1 6)-
-α(1 6)-

-α(1 6)-

-α(1 6)-
-α(1 6)-
-β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)-

-α(1 6)-

-α(1 6)-

-α(1 6)-
-β(1 2)-

XXXG XXLG XLFG

D-Glucopyranose D-Xylopyranose D-Galactopyranose L-Fucopyranose

Possible O-acetylation

(b)
-α(1 3)-

-α(1 3)-
-α(1 2)-
-α(1 2)-

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-β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)- -β(1 4)-

D-Glucuronic acid D-Xylopyranose L-Arabinofuranose

Figure 3 Schematic structural models for xyloglucan (a) and xylan (b).

mutants of extension proteins are embryolethal because of an
inability to form cross walls between cells during cell divi-
sion (Hall and Cannon, 2002). Stretches of amino acids with
alternating hydrophilic and hydrophobic properties encourage
interactions between extensin molecules and facilitate extensin
self-assembly into branched networks that are further stabilised
by intermolecular covalent bonds between tyrosine residues. This
may act as a scaffold for initial cell wall assembly during cell
division: ionic interactions with the negatively charged groups
found in pectin might attract the polymer to the site of pri-
mary wall deposition. Correct O-glycosylation on extensins is
essential for cell wall self-assembly and root hair elongation in
Arabidopsis (Velasquez et al., 2011). Extensins have also been
proposed to play important roles in cell wall structural integrity,
wound healing and plant defence. See Lamport et al. (2011)
for a full review of extensin structure and function. See also:
Glycoproteins; Plant Golgi Apparatus; Protein Glycosylation,
an Overview
Another important group of cell wall structural proteins is the
glycine-rich proteins (GRPs), polypeptides which comprise up
to 70% glycine residues arranged in short repeating units. It is
believed that in GRPs, the repetitive nature of these glycine-rich
domains allows the formation of pleated β-sheets, which in turn
provide elasticity and tensile strength during vascular develop-
ment. Using immunocytochemistry, many studies have shown
that GRPs are associated with the protoxylem, putatively due to a
role in the plant repair system utilised during the stretching phase
(Ringli et al., 2001).
Another prominent class of cell wall glycoproteins is the ara-
binogalactan proteins (AGPs). These consist of a proline-rich
polypeptide backbone which is post-translationally modified
to hydroxyproline [XIII], permitting extensive O-glycosylation.
The addition of long chains of arabinogalactan occurs in the
Golgi apparatus before AGP secretion to the cell wall. The gly-
can chains of AGPs are very long, highly branched and highly
variable. The arabinogalactans attached to AGPs show signifi-
cant structural differences to the pectic arabinogalactans shown
in Figure 2, with different backbone linkages and side-chain
structures. Many AGPs additionally have glycosylphosphatidyli-
nositol (GPI) anchors, which tether the proteins to the plasma
membrane. GPI-anchored AGPs may be involved in signalling
interactions between the cytoplasm and the cell wall compartment
through interactions with receptor-like kinases and/or micro-
tubules, but AGPs have traditionally been considered to have a
primarily structural role. In Arabidopsis, some AGPs are linked
covalently with pectins and xylans to form complexes called
Arabinoxylan Pectin Arabinogalactan Protein1 (APAP1). An
increased extractability of pectin and xylan is found in apap1
mutant plants, indicating structural roles of AGPs by acting
as cross-linker in cell walls (Tan et al., 2012, 2013). Some
recent functional studies suggest roles for AGPs in diverse sig-
nalling processes, including programmed cell death, pattern for-
mation, cell differentiation, pollen growth and guidance and inter-
actions with microbes. See Seifert and Roberts (2007) for a
review of AGP function, and see Canut et al. (2016) for a com-
plete review of posttranslational modifications in plant cell wall
proteins.
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Plant Cell Walls
HO
COOH
N
H
[XIII]
L-Hydroxyproline
Other functional classes of cell wall proteins involved in cell
wall modification and in the regulation of wall expansion are
discussed in further subsections. These proteins are vital for rein-
forcing the wall, but certain specialised walls are also reinforced
by components that are neither polysaccharide nor protein in
nature. These include lignin, cutin, suberin and silica, also dis-
cussed in later sections. See also: Cutin and Suberin
Chemical differences between different
cell walls
The cell wall structures described above are ‘averages’, but there
is a lot of taxonomic variation in primary wall composition. The
cell walls of grasses (including cereals) are identifiable by their
mixed-linkage glucans (MLGs). Grass cell walls also contain
more xylans and less pectins and xyloglucans than do the walls
of dicots and nongraminaceous monocots. Grass-derived xylans
contain much more ferulic acid, giving antioxidant properties.
Grass xyloglucans contain relatively little fucose, while those
of the Solanaceae (potato, tomato, tobacco, etc.) are enriched in
arabinose. Duckweed floating plants are unusual in having sig-
nificant amounts of apiogalacturonan. Secondary walls of xylem
vessel cells contain large amounts of cellulose, xylans and man-
nans, but little or no pectin or xyloglucan. Xylem secondary cell
walls also contain large amounts of reinforcing lignin arranged
in annular rings, helical spirals and pitted plates. See also: Plant
Tracheary Elements; Secondary Cell Walls; Xylem: Differen-
tiation, Water Transport and Ecology
Macromolecular Organisation
of the Wall Matrix
While the cellulose microfibrils are carefully ordered (see ear-
lier), the cell wall matrix as a whole, containing the majority
of the cell’s water, is more amorphous. However, the pectins
and hemicelluloses of the matrix do favour preferred orientations
and interactions. Most polysaccharides and glycoproteins of the
matrix are water soluble after purification but are firmly bound
in the wall and are therefore not water extractable. This implies
extensive cross-linking of matrix polymers in the wall. These
cross-links, which can be covalent or noncovalent (see the fol-
lowing discussion), determine the physical properties of the wall.
Noncovalent cross-links
Hydrogen bonds
Hydrogen bonds form between suitably oriented functional
groups that undergo fractional ionisation, specifically a hydroxyl
9
 Plant Cell Walls

group and an ether-bonded oxygen. other polysaccharides and proteins. Certain sugar residues,
especially apiose, form relatively stable borate di-esters: pairs
− −
R–O𝛿 –H𝛿+ … 𝛿 O < of rhamnogalacturonan-II molecules are thereby cross-linked
through apiose–borate–apiose diester bridges. This connection
In vitro, most hemicelluloses (including xyloglucans, MLGs between borate and pectin was first acknowledged in the 1930s
and low-arabinose xylans) readily hydrogen bond with cellulose. and was later confirmed to have a structural and mechanical
While an individual hydrogen bond is weak and transient, the effect on the cell wall (Hu et al., 1996).
extensive hydrogen bonding which occurs between two polysac-
charide molecules can add up to a firm and stable connection. Glycosidic bonds
Xyloglucan in vivo appears to be hydrogen bonded to the surfaces
A glycosidic linkage is a covalent bond connecting a sugar to
of cellulose microfibrils. As xyloglucan chains are long (typi-
another molecule, in the case of polysaccharides another sugar.
cally 300 nm) compared with the spacing between microfibrils
In cell wall carbohydrates, these are O-glycosidic bonds. These
(typically 30 nm), a xyloglucan chain may tether neighbouring
bonds are synthesised in newly forming carbohydrate chains
microfibrils. This would give xyloglucan a key position in wall
by glycosyltransferases. A glycosidic bond can also be formed
architecture. See Park and Cosgrove (2015) for a full review of the
between an existing polysaccharide through its reducing terminus
interactions between xyloglucan and other cell wall components.
to a nonreducing sugar residue of another polymer. In this way,
formerly separate polymer molecules can theoretically join to
Full ionic bonds form a linear or branched structure.
Full ionic bonds between amino and carboxyl groups also exist in
the wall matrix. The lysine residues of extensins are attracted to
the galacturonate residues of pectins. In addition, Ca2+ ions form Cell Wall Expansion
cross-links between pairs of negatively charged HGs.
During cell growth, wall extensibility can be defined as the abil-
R–NH3 + –− OOC –R′ ity of the cell wall to increase in surface area, in an irreversible
process. The plant cell grows in surface area by rearrangement
of cellulose microfibrils and other associated matrix components
Covalent cross-links in the plane of the wall. This does not necessarily require the
Phenolic coupling products addition of new polymers. Wall extensibility should not be con-
fused with wall elasticity, which is a property of both nongrowing
Phenolic groups in the cell wall include tyrosine residues of pro- and growing cells that allows the cell wall to accommodate tran-
teins and feruloyl esters [XII] of some polysaccharides. These sient changes in cell volume (Cosgrove, 2015a). The pliant walls
groups may be oxidatively coupled (dimerised) in vivo: tyro- of growing cells are under significant tensile stress, resulting
sine can form isodityrosine [XIV], which can itself dimerise to from the stretching of wall polymers as they resist cell turgor
di-isodityrosine. Ferulate groups form several diferulates, such as pressure (see the following discussion). The irreversible expan-
the example shown in compound [XV]. Such coupling may lead sion of the primary wall requires neighbouring microfibrils to
to covalent cross-linking of wall polymers. move apart from one another. To ease this stress on the wall,
O load-bearing polymers are rearranged, which loosens the cell wall
C O matrix and reduces cell stress and turgor pressure. In response
O Ara to this reduced intracellular pressure, water flows into the cell,
Ara O
OH C causing a volumetric expansion of the cell wall chamber and
restoration of turgor and wall stress. Proposed wall-loosening
O
agents that could enable this movement include hydrolases, trans-
O glycosylases, expansins and hydroxyl radicals (see the following
O CH3
CH2 discussion). This process of cell wall expansion may also be
coupled with the intercalation of new cell wall polymers (Cos-
H2C CHNH2
O CH3 grove, 2015a, 2015b). Cell expansion is fundamentally essential
CH COOH COOH
OH
for plant growth and morphogenesis, so the existence of multi-
NH2 ple mechanisms to achieve wall expansion offers multiple control
[XIV] [XV] points for the plant. See also: Plant Exocytosis, Endocytosis and
L,L-Isodityrosine A dimer of Membrane Recycling in Turgid Cells
feruloyl arabinose
Acid growth theory
Ester and amide cross-links The localised acidification of the cell wall is one mechanism
Other potential covalent cross-links in the cell wall include proposed to mediate auxin-induced wall loosening. The so-called
O-galacturonoyl-sugar esters and N𝜀 -galacturonoyl-lysine acid growth occurs when plant cells enlarge rapidly due to an
amides, by which pectins may, respectively, be bonded to increased extensibility of the cell wall caused by a pH change that

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activates the expansin proteins (see the following discussion).
The pH of the cell wall of growing cells is usually between 4.5 and
6, which is sufficient to activate expansin activity. Treatment of
excised stem and coleoptile segments with the growth-promoting
hormone auxin results in the rapid acidification of the wall. Inter-
estingly, even in the absence of auxin, treatment of tissue
with mildly acidic buffers rapidly promotes wall loosen-
ing. Conversely, the treatment of tissue with a neutral buffer
tends to suppress wall loosening in the presence of auxin. See
Sanchez-Rodríguez et al. (2010) for a review of hormones and the
cell wall. See also: Auxin; Plant Cell Growth and Elongation
Wall tightening
After cell maturation, cell walls lose the ability to expand, result-
ing in a cessation of growth, which is generally irreversible and
is typically accompanied by a tightening of the cell wall. Sev-
eral modifications to cell wall structure related to tightening
during maturation have been proposed, with changes in the hemi-
cellulose network most often observed. Tightening is thought
to involve the formation of more cross-links, probably includ-
ing phenolic coupling products such as iso-dityrosine, diferulate
and lignin, by the action of peroxidase enzymes. Indeed, there
are many examples of negative correlation between growth rate
and peroxidase activity. Another possible mechanism of wall
tightening is the de-esterification of methyl-esterified HG by
pectin methylesterase (PME), which consequently strengthens
pectin-calcium networks (see the following discussion).
Regulation of Cell Wall Expansion
Glycoside hydrolases
Glycoside hydrolases, also known as glycosyl hydrolases or
glycosidases, are extremely common in almost all forms of life.
These are the enzymes that cleave glycosidic bonds. During cell
wall biosynthesis and expansion, glycosyltransferases and gly-
coside hydrolases are, respectively, responsible for the synthesis
and breakage of glycosidic bonds. These opposing activities can
even work together to produce specific carbohydrate structures:
a membrane-bound glycosidase that cleaves β-glucans (called
KORRIGAN) is an integral part of the primary cell wall CSC
and influences the organisation of cellulose in the wall (Vain
et al., 2014).
(a) o XTH
(b) XEH + H2O
(c) XET + o
Figure 4 Mechanism of wall loosening caused by the activity of xyloglucan endo-transglycosylase (XET).
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Plant Cell Walls
A glycoside hydrolase may hydrolyse a bond within the back-
bone of a polysaccharide (endo-type activity) or it may remove
a terminal sugar residue (exo-type activity) from a side chain
or oligosaccharide. These endo and exo type enzymes are often
referred to as glycanases and glycosidases, respectively (e.g. a
xylanase and a xylosidase) (Franková and Fry, 2013). See also:
Glycosidases: Functions, Families and Folds
In the cell wall, where cellulose microfibrils are tethered by
hemicellulose chains, enzymes that can deconstruct hemicellu-
loses can permit cell wall expansion. Regulating the expression
of genes encoding these enzymes is one important way for the
plant to regulate cell wall expansion. Endo-type glycanases can
loosen the cell wall by reducing polysaccharide molecular weight
to increase solubility. Cellulases (endo-β-(1→4)-D-glucanases)
cleave the backbones of β-glucans such as xyloglucan and MLGs,
which may loosen the wall and facilitate cell expansion. Primary
cell walls also contain numerous glycosidases. As these enzymes
remove single monosaccharides from nonreducing chain termini,
their effects on large polysaccharide chains in the cell wall are
small, but they may be involved in uncoupling wall components
from each other. However, the inhibition of β-D-glucosidase and
β-D-galactosidase activities does not block the growth-promoting
action of auxin, so their function at the wall remains unclear.
Some glycosidases at the wall may be involved in the final ‘trim-
ming’ of structures such as xyloglucan before their initial depo-
sition at the wall.
Xyloglucan endo-transglycosylase/
hydrolase (XTH)
The xyloglucan endo-transglycosylase/hydrolase (XTH) enzyme
class includes enzymes with xyloglucan endo-transglycosylase
(XET) and xyloglucan endo-hydrolase (XEH) activities
(Figure 4). XET enzymes perform nonhydrolytic cleavage
and religation of xyloglucan chains, resulting in the extension
of xyloglucan polymers. By contrast, XEH enzymes cleave
the xyloglucan backbone, leading to a reduced chain length.
Higher plants have large numbers of XTH genes, underlining the
importance of these enzymes in cellular processes. An in-depth
comparison of the subtle differences between XET and XEH
enzymes can be found in Eklöf and Brumer (2010).
A role for XTH enzymes in cell wall expansion has been
suggested since the initial discovery of the enzymes in the
early 1990s. Models of cell wall structure which propose a
o
o
11
 Plant Cell Walls
‘tethered network’ say that the modification of xyloglucan by
XET enzymes leads to a loosening of the wall, allowing for
cellular expansion. By the action of a XET enzyme, a xyloglu-
can tether is cleaved, allowing incremental wall expansion and
molecular realignment of the attached cellulose microfibrils.
This is followed by reformation of the tether using a second
xyloglucan chain. The theory is supported by evidence that XET
activity often correlates with the rate of cell expansion, and
XET-mediated rearrangement of xyloglucan structure in the cell
wall has been demonstrated in developing wood (Nishikubo et al.,
2011). Research on the role of XTH enzymes in cell wall exten-
sibility is summarised in Miedes et al. (2011).
However, some recent insights may challenge this theory. The
direct application of an XTH enzyme leads to very limited
loosening of the cell wall (Saladie et al., 2006). In addition,
xyloglucan-deficient mutants of Arabidopsis thaliana show very
mild growth phenotypes. A recent alternative model proposes
that xyloglucan and cellulose interact closely in ‘biomechanical
hotspots’, areas of tight adherence which provide great mechani-
cal strength and may be the target of expansins (discussed in the
following section).
In recent years, it has been proposed that there are enzymes
similar to the XET which act to remodel other hemicelluloses.
A mannan endo-transglycosylase has been characterised and
detected in fruits and flowers from a range of species (Schröder
et al., 2004). Similarly, an endo-transglycosylase acting on xylan
has been purified from ripe papaya fruit and detected in the pri-
mary cell walls of some other fruit, vegetable and cereal tissues
(Johnston et al., 2013). These may play a similar role to the XET
enzymes, but their contribution to cell wall extensibility remains
to be verified. For an in-depth review into the importance of
enzymes in plant cell wall extensibility, see Cosgrove (2015a).
Expansins
It is more certain that the expansin proteins act as wall-loosening
agents. As their name suggests, they regulate cell wall expan-
sion in growing cells. The addition of exogenous expansin to
growing cells stimulates their growth, while the suppression of
expansin production by gene silencing decreases plant growth.
Expansins act optimally at low pH and may therefore mediate
‘acid growth’ (see earlier). Expansins act rapidly to promote the
relaxation of wall stress during cellular expansion, most likely by
the cleavage of hydrogen bonds between cell wall polysaccha-
ride complexes, allowing microfibrils to move without the lysis
of wall polymers.
Plant expansins are classified into two major families. The
α-expansins (EXPAs) are hydrophobic and firmly wall bound,
while the β-expansins (EXPBs) are hydrophilic glycopro-
teins. EXPAs are involved in acid-induced wall loosening. See
also Cosgrove (2015b) for diversity and interactions of plant
expansins. See also: Plant Peptide Signals
Pectin methylesterases
As described earlier, pectin in the cell wall forms a soft and
viscous gel-like matrix in which structural proteins and the
cellulose–hemicellulose network are embedded. HG is exten-
sively methyl esterified before being delivered to the cell surface
12 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
(Table 1, Figure 2). In the cell wall, PME enzymes hydrolyse
the methyl-ester bonds on the HG chain, converting neutral
methyl-esterified regions of HG into polyanionic domains with
exposed charged carboxylic acid groups that can be strongly
cross-linked by Ca2+ bridges. This de-esterification process may
proceed along the chain in a linear manner or may occur ran-
domly along the length of the chain. When PMEs act randomly
on HG, protons are released that promote the action of EPG and
contribute to cell wall loosening. Conversely, when PMEs act
linearly on methylated pectin, they expose many free carboxyl
groups that can interact with Ca2+ or borate ions, creating a
pectin gel that contributes to cell wall stiffening. In this way,
PMEs may both promote growth through cell wall loosening
and inhibit growth through wall stiffness, and so the regulation
of PME activity is vital in controlling cell wall expansion. This
regulation is achieved by spatially and temporally differential
expression of PME genes. Many studies have shown a strong cor-
relation between PME gene expression and enzyme activity with
physiological processes such as fruit maturation, seed germina-
tion, pollen tube growth, cambial cell differentiation, hypocotyl
elongation, cellular adhesion and response to pathogen attack. In
addition, PMEs have many industrial applications. As pectin is
mainly responsible for the cloudiness of fruit juices, PMEs are
used in combination with pectinases to clarify juice. Furthermore,
pectins with a reduced methyl-ester content are used for the pro-
duction of jellies and jams with a low sugar content. See Pelloux
et al. (2007) for a comprehensive review of PME function. See
also: Pectic Substances; Plant Cell Growth and Elongation
Nonenzymatic scission of wall
polysaccharides
In addition to protein-mediated wall loosening, cell wall polysac-
charides may be subjected in vivo to nonenzymatic scission
mediated by hydroxyl radicals (⋅ OH). ⋅ OH is a highly active form
of reactive oxygen species (ROS) and plays important roles in
signalling and cell death. It has been demonstrated in vitro that
cell wall polysaccharides can be broken down by ⋅ OH. This rad-
ical can be produced from hydrogen peroxide (H2 O2 ) in the cell
wall by a Fenton reaction using reduced copper ions (Cu+ ).
Cu+ + H2 O2 → Cu2+ + OH− + OH
•
⋅
OH radicals can also be produced by the combined action of
NAD(P)H-oxidases, superoxide dismutases and peroxidases. The
plasma membrane-localised NAD(P)H-oxidase reduces molecu-
lar oxygen to superoxide radicals (O2 − ), which are then converted
to H2 O2 by an extracellular superoxide dismutase. Superoxide
and H2 O2 can be converted by apoplastic peroxidases into highly
reactive ⋅ OH. See also: Enzymatic Free Radical Reactions;
Ozone and Reactive Oxygen Species
The production of ⋅ OH can be controlled by the presence of
Cu+ in the close vicinity of cell wall polysaccharides. Once
produced, the highly reactive ⋅ OH will diffuse scarcely 1 nm
before reacting with polysaccharides. This shows that if ⋅ OH is
generated at the right time and place within the cell wall, it can
be specifically targeted to the scission of a particular cell wall

area or structure. In vitro, ⋅ OH cleaves xyloglucan chains and
can solubilise pectins. The occurrence of ⋅ OH in cell walls in
vivo has been demonstrated spectroscopically, and the artificial
generation of ⋅ OH in cell walls stimulates the expansion of
isolated cell walls, as well as living cells. Recent studies have
shown that ⋅ OH can be involved in cell wall loosening during
fruit ripening, cell enlargement, extension growth of roots and
seed germination. See del Rio and Pusso (2009) for additional
insight into the roles of ROS in plant signalling.
Lignification
After a cell has ceased growth and expansion is no longer
required, lignin may be deposited in some tissues. Lignin is a
polyphenolic polymer deposited directly within the cell walls of
specialised cell types such as tracheary elements (vessels and
tracheids), fibres and other sclerenchymatous cells to provide
strength and rigidity. Small amounts of lignin have also been
found in some growing tissues (Figure 5). Lignin is a signifi-
cant component of woody biomass and may be the second most
abundant terrestrial biopolymer after cellulose. The major func-
tions of lignin are to strengthen and waterproof the cell wall of
specialised cell types including conducting vessels. Lignin con-
tributes to weight-bearing in trees by enabling stress transfer
between neighbouring microfibrils and confers physical resis-
tance to the buckling of walls under compressive stress. Partly
due to its randomised structure, lignin is highly resistant to enzy-
matic digestion, allowing it to also serve as a physical barrier
against initial pathogen colonisation. See Bonawitz and Chapple
(2010) for a review of lignin function and synthesis. See also:
Lignins; Plant Cell Wall Biosynthesis; Plant Tracheary Ele-
ments; Secondary Cell Walls
S1
S2
Figure 5 Immunogold localisation of lignin in the secondary cell walls of
poplar wood fibre cells. The image shows a cross section of a fibre cell wall
which has been treated with a preparation of gold particles coupled with
antibodies that recognise lignin. The gold particles are visible as black dots
in the S1 and S2 layers of the secondary cell wall, indicating the presence
of lignin there, but are absent from the primary wall and middle lamella
between two neighbouring cells. Bar is 0.5 μm. Reproduced from Joseleau
et al. (2004) © Springer.
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Plant Cell Walls
Lignification is a part of the normal differentiation process
in cell types requiring additional mechanical strength, but it
can also be triggered as a response to various biotic and abi-
otic stresses. Lignin is produced by the oxidative polymerisation
of three monolignols, p-coumaryl [XVI], coniferyl [XVII] and
sinapyl [XVIII] alcohols. These monomers are quite randomly
associated within the lignin polymer, owing to the radical nature
of the synthesis reactions involved, although there do appear to be
some more ordered domains. The monolignols are synthesised in
the cytosol and then transported into cell walls, where the poly-
merisation reaction takes place, catalysed by oxidases including
laccases and peroxidases (Bonawitz and Chapple, 2010). The
lignin polymer is classified into three types: guaiacyl (G) lignin,
a polymer of coniferyl alcohol; syringyl (S) lignin, a polymer of
sinapyl alcohol and p-hydroxyphenyl (H) lignin, a polymer of
p-coumaryl alcohol. The ratio of G, S and H units within a lignin
polymer can differ significantly among different plant species and
even among different cell types in the same species. For example,
in dicots, lignin is usually composed of moieties of both G and
S lignins, while gymnosperms contain G lignin but lack S lignin.
H lignin is particularly common in the secondary walls of grass
species (Zhong and Ye, 2015).
CH2OH
CH2OH
HO
HO
H3C O
[XVI] [XVII]
p-Coumaryl alcohol Coniferyl alcohol
H3C O
CH2OH
HO
H3 C O
[XVIII]
Sinapyl alcohol
Functions of Cell Walls
Cell and tissue integrity
One major function of the cell wall is to provide the plant with a
physical form by maintaining cell and tissue integrity. The com-
position and structure of plant cell walls gives them extraordinary
physical strength, allowing for cell turgor, upright growth and
the vertical transport of water and solutes. As a result of their
mechanical strength, cell walls can resist deformation under ten-
sile and compressive stress, conferring stability to plant cells,
tissues and organs.
Turgor pressure
Plant cells have the ability to accumulate solutes at higher concen-
trations within the vacuole than exists outside of the cell, resulting
in the movement of water across semipermeable membranes by
osmosis. In the absence of a cell wall, this water uptake would
cause the protoplast to swell and eventually burst. In an intact
13
 Plant Cell Walls
cell, however, the cell wall contains the swelling protoplast and
exerts a back pressure, allowing a positive hydrostatic pressure to
build up in the cell. This pressure is termed turgor pressure and
is responsible for growth, transport, movement and responses to
environmental stress. The opening and closing of stomatal pores
is an important turgor-driven process, which regulates water loss
from, and carbon dioxide entry into, leaves. See also: Turgor
Pressure
Cell growth and morphogenesis
Plant growth and morphogenesis are dependent on the physical
characteristics of the cell wall. Isolated protoplasts (where the
cell wall has been degraded by enzymes) are always spherical,
showing that the original characteristic shape (morphology) of the
cell was dictated by the wall. A new primary cell wall is generated
in the cell during cell division and quickly increases in surface
area during cell expansion. As discussed earlier, the expansion
behaviour of the cell wall not only controls the rate and direction
of cell growth but also regulates cell volume. In a growing plant
cell, the cellulose microfibrils are distributed directionally. For
this reason, the stiffness of the wall will vary depending on the
direction of the stress. In the presence of turgor pressure driving
cell expansion from within the cell, this generates differential
wall expansion and contributes to the generation of nonspherical
cell shapes during morphogenesis. See also: Biomechanics of
Plant Cell Growth; Plant Biomechanics; Plant Cell Growth
and Elongation
Reversible (elastic) cell expansion:
maintenance of cell volume
Plant cells expand by deforming reversibly (elastically) during
growth or whenever turgor pressure changes (e.g. during times of
drought). The new size is reached quickly, as the rubber band-like
wall has elastic properties. In the living plant, swelling cells press
against one another to maintain structural integrity over a whole
tissue. Elastic cell expansion is not permanent and reverses when
the deforming force is removed. Thus, the cell wall determines
and maintains cell volume under turgor pressure. See also: Cell
Size Control; Turgor Pressure
Irreversible (plastic) cell expansion: cell
volume growth
Plant cell growth is an irreversible expansion of the stiff cell wall
under tension. Plant cells control their expansion by controlling
the structure and material properties of their wall. When wall
material from an immature tissue is stretched, both elastic and
plastic expansion occur, the latter being the increase in wall area
that is not reversed when the stretching force is removed. A wall
that is capable of plastic expansion is described as loose; the
opposite is described as tight. Plastic expansion is the biophysical
basis of cell and plant growth (defined as an irreversible increase
in cell volume). It is slow, occurring over a period of hours,
and is dependent on the synthesis of new cell wall material
and the activity of wall-modifying proteins. Cellulose synthesis
appears to be directly related to the amount of cell expansion,
14 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
and impairments in cellulose production usually lead to stunted
growth. See also: Biomechanics of Plant Cell Growth; Plant
Cell Growth and Elongation
Growing walls expand irreversibly under tensile stress through
a process termed ‘polymer creep’ which involves the reg-
ulated breaking and reforming of bonds between cell wall
polymers. This is thought to involve enzymatic cleavage of
cell wall polysaccharide backbones, XET activity to break and
reform xyloglucan chains, the transient enzymatic disruption
of hydrogen-bonding networks within the cell wall matrix,
structural modification and calcium loss from pectins and
ROS-mediated scission of cell wall polymers (see earlier). See
also: Plant Biomechanics; Plant Cell Growth and Elongation
The expansion of a plant organ may be largely dictated by
the cell wall properties of one specific tissue – in shoots, this
is the epidermis. Thus, removal of the epidermis from a stem
or leaf enables a rapid burst of (mainly elastic) expansion in the
underlying tissues. See Cosgrove (2005) for a general review on
cell growth and cell wall expansion. See also: Epidermis: Outer
Cell Layer of the Plant; Shoots and Buds in Arabidopsis
Permeability
Plant cell walls are semipermeable and allow small molecules
and proteins to pass through in size-dependent channels. This
allows the free passage of most small, water-soluble molecules
and ions such as oxygen, carbon dioxide, nitrate, phosphate, sug-
ars and amino acids into and out of the cell, while excluding
larger molecules. The upper size limit for transport through the
wall is thought to be in the range 5000–50 000 Da. Localised pH
is another critical factor governing the transport of molecules
through walls. The permeability of plant cell walls to macro-
molecules may restrict the ability of the plant’s own enzymes to
change the physical and biochemical properties of the wall. See
also: Plasmodesmata
Cation binding
The plant cell wall acts as an ion exchanger where the charge of
cell wall components dictates their interactions with exchange-
able ions in the surrounding environment. Walls have a high
affinity for certain inorganic cations and may be a major loca-
tion of some of these, such as Ca2+ , Cu2+ and Al3+ . The action of
several wall-modifying enzymes can directly influence the affin-
ity of the cell wall for these cations by affecting the abundance
of negatively charged groups in the cell wall. Electrical potential
in cell walls is also involved in swelling and extension capabili-
ties of the cell wall matrix and is therefore thought to play a role
in pectin bonding, texture and intercellular adhesion. Cell wall
affinity for cations is partly governed by the activity of PMEs, a
major factor in the genotypic differences in aluminium sensitiv-
ity between crop cultivars. See also: Heavy Metal Adaptation;
Pectic Substances
Defence
The cell wall represents a first line of defence for plant cells
against mechanical injury and pathogen attack. In addition to

acting as a physical barrier to ingress, active local modifica-
tions of the cell wall can rapidly occur in response to pathogen
attack or wounding, notably the deposition of callose and lignin
at sites of penetration by fungal appressoria and the forma-
tion of water-proof cork at sites of wounding. The virulence of
many plant pathogens depends on their ability to degrade plant
cell walls using secreted hydrolytic enzymes, while the resis-
tance of plants against many pathogens depends on dynamic
wall modifications (e.g. callose deposition) that make the wall
more resistant to degradation. Plants may also produce cell
wall-targeted hydrolytic enzymes aimed at degrading the cell
walls of invading pathogens. Chitinase enzyme production is
often induced in the early stages of pathogenesis to attack the
chitinous cell walls of fungi and the exoskeletons of insects. Of
course, pathogens have developed counterstrategies to overcome
these defences. See also: Pectic Substances; Plant Disease and
Defence; Coevolution: Plant-Insect
The first line of defence in plants is the recognition of con-
served molecules characteristic of many microbes. These elic-
itors are also known as microbe-associated molecular patterns
(MAMPs) or pathogen-associated molecular patterns (PAMPs).
MAMPs are recognised by specific receptors on the surface
of plant cells. This first phase of defence induction is called
MAMP-triggered immunity (MTI). Several specific oligosac-
charides possess hormone-like or signalling activity, regulating
growth, development and metabolism. Such oligosaccharides are
termed oligosaccharins for ‘oligosaccharide elicitins of plant
defence’. Pectins are primary targets of attack by invading
microbes and their breakdown products (oligogalacturonides)
function as potent elicitors of plant defence responses. The abun-
dance of methyl and acetyl ester groups on pectic polymers
affects their susceptibility to hydrolytic enzymes, and highly
de-esterified forms of pectin are associated with wall tighten-
ing in response to pathogen attack. Oligosaccharide products of
chitin degradation have also been shown to induce plant defence
responses. Infection can induce the deposition of nonconstitu-
tive wall components. For example, lignin is deposited in wheat
leaves challenged by the pathogenic fungus Puccinia graminis,
and extensin production is induced in melons upon attempted
infection by Colletotrichum lindemuthianum. In addition, struc-
tural proteins of the wall can be rapidly cross-linked through
the formation of di-isodityrosine bridges when cells are exposed
to fungal elicitors. See also Hematy et al. (2009) for a review
of pathogens and the cell wall. See also: Fungal Pathogens of
Plants; Plant Defences against Fungal Attack: Biochemistry
Seed nutrition
Some atypical cell wall polysaccharides are found in the
endosperm cell walls of some seeds in disproportionately large
amounts and function as storage reserves to be rapidly broken
down during seed germination. These polymers may partly or
completely replace the traditional energy storage molecules
such as starch, seed oil and storage proteins. Seed cell wall
storage polysaccharides can be roughly categorised based on
their monosaccharide composition and structural complexity:
glucans (xyloglucan and MLG), linear glucomannans, branched
galactomannans and galactans. In malting barley endosperm,
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Plant Cell Walls
energy reserve MLGs are broken down before starch or any
other cell wall polysaccharides. Seed mannans can also provide
additional hardness as in palm (Phoenix dactylifera) and coffee
(Coffea arabica) seeds. In legumes, energy storage galactoman-
nans can represent up to 30% of the seed dry weight in the
endosperm cell wall of many species. Some plants may have
evolved to store these storage carbohydrates instead of starch as
many herbivores are incapable of degrading these more complex
carbohydrate polymers (for more detailed discussion of this
topic, see Buckeridge, 2010). See also: Plant Storage Products
(Carbohydrates, Oils and Proteins); Seed Germination and
Reserve Mobilization; Seeds
Other mechanical roles
Some specialised cell walls have particular mechanical roles such
as lubrication of the passage of roots through soil by mucilagi-
nous root-cap cell walls, the control of texture in fruit during
ripening and tissue disintegration to enable leaf and fruit abscis-
sion. Plants that have moving tissues, such as pine cones which
respond to changes in humidity, function by elegant deposition
of cell wall polymers in careful layers of different degrees of
hydrophobicity. See also: Plant Biomechanics; Roots: Contri-
bution to the Rhizosphere
Injury
Plants undergo many different types of mechanical injury caused
by both biotic and abiotic factors, such as parasite invasion, insect
attack, wind and fire. As the cell wall is the primary site of
mechanical injury, modifications to polysaccharide components
and matrices are among the main adaptive responses in cells, typi-
cally leading to the strengthening of the cell wall. The wall can be
strengthened by increased deposition of structural proteins (e.g.
hydroxyproline-rich and proline-rich proteins) and other poly-
mers (e.g. suberin, lignin and callose). Suberin is often induced
in newly exposed cells as a wound response, for example when
a potato tuber is cut open, resulting in the transformation of cells
adjacent to the wounding site into cork cells. The production
of cork at wounding sites seals the wound and prevents water
loss from adjacent tissues, as well as impeding the penetration
of pathogenic microbes into the wound site. Another extremely
rapid response to wounding is the induction of callose deposi-
tion. This is particularly effective at blocking severed phloem
vessels, preventing the loss of nutritious phloem sap. In a lab-
oratory setting, touch stimulation of plants, perhaps mimicking
the action of wind or passing animals, can induce XET expres-
sion and the resulting molecular rearrangement of xyloglucans
may confer mechanical strength. See also: Callose and Related
Glucans; Cork; Plant Responses to Wounding
Developmental patterning
It is currently thought that the cell wall may be involved in devel-
opmental patterning of plant tissues. Atomic force microscopy
has been used to show that the cell walls in the tissues of the
shoot apical meristem are subject to tensile and compressive
stresses in a well-defined pattern. Microtubules respond to this
15
 Plant Cell Walls
stress and orient themselves in a pattern that runs parallel to the
stress direction, influencing the deposition of cellulose microfib-
rils. The resulting wall structure can resist the applied stress.
The patterned distribution of stresses in the tissue is thought to
provide positional information for the initiation of leaves api-
cal to the meristem. Experimental manipulation of stress gradi-
ents at the meristem by applying compressive forces can induce
the initiation of leaf formation. Moreover, localised cell wall
loosening by the direct application of an expansin or by the
induction of expansin expression has been shown to initiate the
entire developmental programme of leaf formation and alter phyl-
lotaxy. See Mirabet et al. (2011) for a review of cell pattern-
ing and the cell wall. See also: Apical Meristems; Meristems;
Phyllotaxy; Plant Organ Primordia; Positional Information
in Plant Development; Regulatory Mechanisms of Phyllotaxy
References
Albenne C, Canut H, Hoffmann L and Jamet E (2014) Plant cell wall
proteins: a large body of data, but what about runaways? Proteomes
2: 224–242.
Barros J, Serk H, Granlund I and Pesquet E (2015) The cell biology
of lignification in higher plants. Annals of Botany 115: 1053–1074.
Bonawitz ND and Chapple C (2010) The genetics of lignin biosynthe-
sis: connecting genotype to phenotype. Annual Review of Genetics
44: 337–363.
Bromley JR, Busse-Wicher M, Tryfona T, et al. (2013) GUX1 and
GUX2 glucuronyltransferases decorate distinct domains of glu-
curonoxylan with different substitution patterns. The Plant Journal
74: 423–434.
Buckeridge MS (2010) Seed cell wall storage polysaccharides: mod-
els to understand cell wall biosynthesis and degradation. Plant
Physiology 154: 1017–1023.
Canut H, Albenne C and Jamet E (2016) Post-translational modifi-
cations of plant cell wall proteins and peptides: a survey from a
proteomics point of view. Biochimica et Biophysica Acta 1864:
983–990.
Carpita NC (2011) Update on mechanisms of plant cell wall biosyn-
thesis: how plants make cellulose and other (1→4)-β-D-glycans.
Plant Physiology 155: 171–184.
Cosgrove DJ (2005) Growth of the plant cell wall. Nature Reviews
Molecular Cell Biology 6: 850–861.
Cosgrove DJ (2015a) Plant cell wall extensibility: connecting plant
cell growth with cell wall structure, mechanics, and the action
of wall modifying enzymes. Journal of Experimental Botany 67:
463–476.
Cosgrove DJ (2015b) Plant expansins: diversity and interactions with
plant cell walls. Current Opinion in Plant Biology 25: 162–172.
Eklöf JM and Brumer H (2010) The XTH gene family: an update on
enzyme structure, function, and phylogeny in xyloglucan remod-
elling. Plant Physiology 153: 456–466.
Franková L and Fry SC (2013) Biochemistry and physiological roles
of enzymes that ‘cut and paste’ plant cell-wall polysaccharides.
Journal of Experimental Botany 64: 3519–3550.
Guerriero G, Fugelstad J and Bulone V (2010) What do we really
know about cellulose biosynthesis in higher plants? Journal of
Integrative Plant Biology 52: 161–175.
Harholt J, Suttangkakul A and Scheller HV (2010) Biosynthesis of
pectin. Plant Physiology 153: 384–395.
16 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Hall Q and Cannon MC (2002) The cell wall hydroxyproline-rich
glycoprotein RSH is essential for normal embryo development in
Arabidopsis. The Plant Cell 14: 1–12.
Hematy K, Cherk C and Somerville S (2009) Host-pathogen war-
fare at the plant cell wall. Current Opinion in Plant Biology 12:
406–413.
Hu H, Brown PH and Labavitch JM (1996) Species variability in
boron requirement is correlated with cell wall pectin. Journal of
Experimental Botany 47: 227–232.
Jamet E, Albenne C, Boudart G, et al. (2008) Recent advances in
plant cell wall proteomics. Proteomics 8: 893–908.
Johnston SL, Prakash R, Chen NJ, et al. (2013) An enzyme activity
capable of endotransglycosylation of heteroxylan polysaccharides
is present in plant primary cell walls. Planta 237: 173–187.
Joseleau JP, Imai T, Kuroda K and Ruel K (2004) Detection in situ
and characterization of lignin in the G-layer of tension wood fibres
of Populus deltoides. Planta 219: 338–345.
Lamport DT, Kieliszewski MJ, Chen Y and Cannon MC (2011) Role
of the extensin superfamily in primary cell wall architecture. Plant
Physiology 156: 11–19.
Lopez M, Bizot H, Chambat G, et al. (2010) Enthalpic studies
of xyloglucan – cellulose interactions. Biomacromolecules 11:
1417–1428.
Miedes E, Zarra I, Hoson T, et al. (2011) Xyloglucan endotransglu-
cosylase and cell wall extensibility. Journal of Plant Physiology
168: 196–203.
Mirabet V, Das P, Boudaoud A and Hamant O (2011) The role of
mechanical forces in plant morphogenesis. Annual Review of Plant
Biology 62: 365–385.
Mohnen D (2008) Pectin structure and biosynthesis. Current Opinion
in Plant Biology 11: 266–277.
Nishikubo N, Takahashi J, Roos AA, et al. (2011) Xyloglu-
can endo-transglycosylase-mediated xyloglucan rearrangements in
developingwood of hybrid aspen. Plant Physiology 155: 399–413.
Nixon BT, Mansouri K, Singh A, et al. (2016) Comparative structural
and computational analysis supports eighteen cellulose synthases
in the plant cellulose synthesis complex. Scientific Reports 6:
28696.
Pabst M, Fischl RM, Brecker L, et al. (2013) Rhamnogalacturonan
II structure shows variation in the side chains monosaccharide
composition and methylation status within and across different
plant species. The Plant Journal 76: 61–72.
Panteris E, Adamakis ID, Daras G and Rigas S (2015) Cortical micro-
tubule patterning in roots of Arabidopsis thaliana primary cell
wall mutants reveals the bidirectional interplay with cell expan-
sion. Plant Signaling & Behavior 10: e1028701.
Park YB and Cosgrove DJ (2015) Xyloglucan and its interactions
with other components of the growing cell wall. Plant and Cell
Physiology 56: 180–194.
Pelloux J, Rusterucci C and Mellerowicz EJ (2007) New insights
into pectin methylesterase structure and function. Trends in Plant
Science 12: 267–277.
Ringli C, Keller B and Ryser U (2001) Glycine-rich proteins as struc-
tural components of plant cell walls Cell. Cellular and Molecular
Life Sciences 58: 1430–1441.
del Rio LA and Pusso A (2009) Reactive Oxygen Species in Plant
Signaling. Heidelberg: Springer Verlag.
Saladie M, Rose JK, Cosgrove DJ and Catala C (2006) Characteriza-
tion of a new xyloglucan endotransglucosylase/hydrolase (XTH)
from ripening tomato fruit and implications for the diverse modes
of enzymic action. The Plant Journal 47: 282–295.

Sanchez-Rodríguez C, Rubio-Somoza I, Sibout R and Persson S
(2010) Phytohormones and the cell wall in Arabidopsis during
seedling growth. Trends in Plant Science 15: 291–301.
Scheller HV and Ulvskov P (2010) Hemicelluloses. Annual Review
of Plant Biology 61: 263–289.
Schröder R, Wegrzyn TF, Bolitho KM and Redgwell RJ (2004)
Mannan transglycosylase: a novel enzyme activity in cell walls of
higher plants. Planta 219: 590–600.
Schultink A, Liu L, Zhu L and Pauly M (2014) Structural diver-
sity and function of xyloglucan sidechain substituents. Plants 3:
526–542.
Seifert GJ and Roberts K (2007) The biology of arabinogalactan
proteins. Annual Review of Plant Biology 58: 137–161.
Simmons TJ, Mortimer JC, Bernardinelli OD, et al. (2016) Folding of
xylan onto cellulose fibrils in plant cell walls revealed by solid-state
NMR. Nature Communications 21: article 13902.
Tan L, Showalter A, Egelund J, et al. (2012)
Arabinogalactan-proteins and the research challenges for these
enigmatic plant cell surface proteoglycans. Frontiers in Plant
Science 3: e140.
Tan L, Eberhard S, Pattathil S, et al. (2013) An Arabidopsis cell wall
proteoglycan consists of pectin and arabinoxylan covalently linked
to an arabinogalactan protein. The Plant Cell 25: 270–287.
Vain T, Crowell EF, Timpano H, et al. (2014) The cellulase KORRI-
GAN is part of the cellulose synthase complex. Plant Physiology
165: 1521–1532.
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
View publication stats
Plant Cell Walls
Velasquez S, Ricardi MM, Dorosz JG, et al. (2011) O-Glycosylated
cell wall proteins are essential in root hair growth. Science 332:
1401–1403.
Zhong R and Ye ZH (2015) Secondary cell walls: biosynthesis,
patterned deposition and transcriptional regulation. Plant and Cell
Physiology 56: 195–214.
Further Reading
Morgan JL, Strumillo J and Zimmer J (2013) Crystallographic snap-
shot of cellulose synthesis and membrane translocation. Nature
493: 181–186.
Purushotham P, Cho SH, Díaz-Moreno SM, et al. (2016) A single
heterologously expressed plant cellulose synthase isoform is suf-
ficient for cellulose microfibril formation in vitro. Proceedings of
the National Academy of Sciences of the United States of America
113: 11360–11365.
Schuetz M, Benske A, Smith R, et al. (2014) Laccases direct lignifi-
cation in the discrete secondary cell wall domains of protoxylem.
Plant Physiology 166: 798–807.
Wolf S, Hematy K and Hofte H (2012) Growth control and cell wall
signaling in plants. Annual Review of Plant Biology 63: 381–407.
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