Ann Bot 2014 Content I V

ContentSnapshots
Annals of Botany Volume 114 Number 6 2014
Phosphoinositides and cell walls

(Review)
Supra-molecular assembly of
AGP31 in arabidopsis cell walls
doi:10.1093/aob/mcu055
Trafficking of the cellulose synthase

complex (Review)
AGP31 (arabinogalactan protein 31) is a remarkable cell wall

protein that displays a multi-domain organization unique in
Arabidopsis thaliana. Hijazi et al. ( pp. 1087 1097) demonstrate
that its C-terminal PAC (PRP-AGP containing Cys) domain
interacts in vitro with galactans and with its own central
O-glycosylated (Hyp-O-Gal/Ara-rich motifs) domain. The
interaction of AGP31 with galactans (which are branches of
rhamnogalacturonans I) and with itself suggests that it forms
non-covalent networks in cell walls. Thus a model is proposed of of
interactions of AGP31 with different cell wall components, where
AGP31 participates in complex supra-molecular scaffolds. Such
scaffolds could contribute to the strengthening of cell walls of
quickly growing organs such as etiolated hypocotyls.
Downloaded from http://aob.oxfordjournals.org/ at Centro Federal de Educao Tecnolgica do Paran on December 1, 2014
Recent studies have suggested the involvement of

phosphoinositides (PIs), a class of membrane lipids, as
key regulatory molecules during secretion and assembly of
cell wall polymers, and even recycling processes in plants.
Krishnamoorthy et al. ( pp. 1049 1057) review the current
state of knowledge of how PIs regulate vesicle trafficking,
and their potential influence on plant cell wall architecture.
They consider first how PIs are formed in plants and then
examine their role in the control of vesicle trafficking.
Interactions between PIs and the actin cytoskeleton and
small GTPases are also discussed. Future challenges for
research are suggested.
MYB46/MYB83-mediated
regulation of secondary wall
biosynthesis (Review)
Cellulose synthase complexes (CSCs) are assembled in the Golgi
apparatus but are thought to only synthesize cellulose when
localized at the plasma membrane. Hence the delivery and
endocytosis of CSCs to and from the membrane are important
aspects for the regulation of cellulose biosynthesis. Bashline et al.
( pp. 1059 1067) review recent findings related to CSC
localization and behaviour, as well as recent advances in
understanding associated trafficking pathways and mechanisms.
Topics such as the implications of the Golgi and trans-Golgi
network in CSC assembly and trafficking, and the possible
mechanisms and pathways of CSC secretion, endocytosis and
recycling are also considered.
Formation of secondary cell walls requires co-ordinated

transcriptional regulation of the genes involved in the biosynthesis of
cellulose, hemicellulose and lignin, and the transcription factor
MYB46 (At5g12870) and its paralog MYB83 (At3g08500) have
been shown to function as a master switch for the secondary wall
biosynthetic program in Arabidopsis thaliana. Ko et al. (pp. 1099
1107) review our current understanding of the MYB46-mediated
transcriptional regulatory network, including upstream regulators,
downstream targets and negative regulators of MYB46. They
conclude that because of its ability to directly regulate the
biosynthesis genes for the major components, MYB46 may be useful
in pathway-specific manipulation of secondary wall biosynthesis.
Dynamic calcium recycling by an

AGP-Ca21 oscillator (Review)
Cell-specific transcription and

translation in root cells
Glycomodules of cell surface arabinogalactan proteins (AGPs)

bind Ca2+ stoichiometrically at pH 5, and low pH dissociates the
AGP-Ca2+ capacitor and hence is the primary source of cytosolic
Ca2+ waves. Lamport et al. ( pp. 1069 1085) thus consider that
dynamic recycling of cytosolic Ca2+ by an AGP-Ca2+ oscillator
determines the Ca2+ flux and may be a crucial component in
the regulation of plant growth. The link between AGPs and
Ca2+-signalling includes a wide range of auxin-dependent plant
processes. This solves the problem of classical AGP function at
the molecular level and accounts for the wide involvement of
AGPs in plant morphogenesis, including tropic and nastic
movements.
A long-standing question in cellular biology is how well the

transcriptome is coupled to the proteome. Rajasundaram et al.
(pp. 11091123) assess the degree of co-ordination and divergence
between these two levels of cellular organization by using cell-type
specific datasets of the root transcriptome and translatome in
Arabidopsis thaliana, with particular reference to cell wall biology. In
agreement with previous studies in animal cells, they find that the
majority of genes exhibit uncorrelated transcription and translation
levels. However, components and processes are also identified that are
under co-ordinated transcriptional and translational control in plant
root cells, such as the development of secondary cell wall biosynthesis.
Synergism between FLA4 and ABA

signalling in roots
PME and SBT expression in

arabidopsis
Mixed-linkage glucan and

glucuronoarabinoxylan in roots
In Arabidopsis thaliana, the degree of methylesterification of

homogalacturonans, the main pectic constituent of the cell wall,
can be modified by pectin methylesterases (PMEs) and this plays a
central role in both plant development and responses to stress.
Using a combination of functional genomics, biochemistry and
proteomic approaches, Senechal et al. ( pp. 11611175) identify a
group-2 PME (PME17) that is strongly co-expressed, both spatially
and temporally, with a specific subtilisin-type serine protease
(SBT3.5), particularly in the roots. PME activity is modified in
roots of knock-out mutants for both proteins, with consequent
effects on growth. This suggests a role for SBT3.5 in the processing
of PME17 in planta, and highlights a need for identifying specific
PMESBT pairs.
Rhamnogalacturonan-II in pollen
tube growth (Research in Context)
Plant cell enlargement is unambiguously coupled to changes in cell
wall architecture. Kozlova et al. ( pp. 11351145) study the
correspondence between the fine structure of cell walls and the
course of the elongation process in roots of maize (Zea mays) and
determine the presence of three domains of glucuronoarabinoxylan
molecules: one separating cellulose microfibrils, one interacting
with them, and a middle domain between the two. The middle
domain is masked by mixed-linkage glucan. A model is proposed in
which the mixed-linkage glucan serves as a gel-like filler of the space
between the separating domain of the glucuronoarabinoxylan and
the cellulose microfibrils. Space for glucan is provided along the
middle domain, the proportion of which increases during cell
elongation.
SNARE protein VTI13 and root hair

growth in arabidopsis
Tapetum and pollen wall

ultrastructure in arabidopsis
Endosomal trafficking is required for polarized growth, but many

of the proteins that control this process have yet to be identified.
Larson et al. ( pp. 11471159) identify VTI13 as a vesicular
soluble NSF attachment receptor (SNARE) protein required for
root hair growth in Arabidopsis thaliana. They demonstrate that
VTI13 localizes to the vacuole membrane and the trans-Golgi
network (TGN) or an early endosomal compartment, is important
for TGN integrity, and is likely to play a role in the transport of
cargo to the large vacuole in root hairs. They also show that VTI13
function is essential for the maintenance of cell wall organization
in root hair and root epidermal cells.
ii
Rhamnogalacturonan type-II (RG-II) is the most complex pectic

polysaccharide of the plant cell wall and is necessary for
strengthening. Dumont et al. ( pp. 11771188) study pollen tubes
of Arabidopsis thaliana and show the presence of specific sugars of
RG-II. In addition, they analyse two T-DNA insertion lines in
At3g48820 encoding a putative sialyltransferase-like protein,
possibly involved in the transfer on the homogalacturonan
backbone of Kdo and/or Dha, two specific sugars of RG-II.
Analyses of the two heterozygous lines reveals a strong reduction in
pollen germination and pollen tube growth in vitro and in vivo,
suggesting that sialyltransferase-like proteins are required for
proper pollen tube growth by maintaining the integrity of RG-II.
Preserving the ultrastructure of the developing pollen cell wall

presents challenges because the key cell types are embedded deep
within the anther, making chemical fixation for electron microscopy
difficult. Quilichini et al. (pp. 11891201) use a high-pressure
freezing/freeze-substitution technique in order to preserve the
delicate tapetum tissue and developing pollen grains during
production of the complex pollen cell wall in the anther of wildtype
Arabidopsis thaliana. The technique reveals the ultrastructure of
tapetosomes and elaioplasts, and shows that the tapetum and middle
layer of the anther remain intact into the tricellular pollen and
late-uninucleate microspore stages, respectively.
The physiological function of fasciclin-like arabinogalactan

proteins (FLAs) is largely unknown. Seifert et al. ( pp. 1125
1133) study the salt-oversensitive mutant fla4 and show that
externally applied abscisic acid (ABA) as well as loss of function of
negative ABA regulators suppresses the fla4 phenotype. They find
that fla4 shows a reduced level of ABA responsive transcripts.
Synergism between ABA and FLA4 is further supported by
suppression of the fla4 phenotype by chemical inhibition of ABA
catabolism. The cell membrane-localized FLA4 might indirectly
influence the sensitivity of cytosolic ABA signalling and thereby
link cell wall biosynthesis with the stress response.
Cell-wall structure and evolution in

brown algae
Seed coat polysaccharide production

(Review)
During seed coat differentiation the epidermal cells of certain
species accumulate polysaccharides either to reinforce cell walls,
or for release during seed imbibition to form sticky mucilage.
North et al. ( pp. 12511263) review the recent exploitation of
these cells as a model system for the study of polysaccharide
metabolism and properties, particularly in the model plant
Arabidopsis thaliana. The potential for intra- and interspecies
natural variation in these polysaccharides as a resource to extend
the use of this model and to improve our knowledge of seed
mucilage ecophysiological function is examined.
Variation of cell wall properties

in Miscanthus
Cell wall biosynthesis in charophyte
green algae (Research in Context)
Charophyte green algae (CGA) are the closest living relatives to
land plants, and the cell wall was possibly a defining structure that
enabled the green algal ancestor to colonize land. Mikkelsen
et al. ( pp. 1217 1236) provide genetic evidence that many of
the most important core cell wall polysaccharides have their
evolutionary origins in CGA, including cellulose, mannan,
xyloglucan, xylan and pectin, as well as arabinonogalactan
proteins (AGPs). Moreover, they provide the first evidence that all
cellulose synthase-like genes present in early-divergent land
plants were already present in CGA, implying that some features
of land plant cell walls evolved prior to the transition to land,
rather than having evolved as a result of selection pressures
inherent in this transition.
Miscanthus represents one of the most promising dedicated
lignocellulosic bioenergy crops. A key trait for biomass conversion
to biofuels and biomaterials is cell wall quality. Costa et al.
( pp. 12651277) present data on cell wall compositional changes
as a function of development and tissue type across 25 selected
Miscanthus genotypes. They report compositional differences
between stem and leaf samples to be predominantly associated with
structural carbohydrates, while lignin content does not correlate
with ethanol production from leaf biomass. Overall, leaf tissue
contributes significantly to total above-ground biomass at all
developmental stages. These factors highlight the importance of
examining leaf and stem biomass composition separately in order
to infer gene trait associations relating to cell wall quality of
lignocellulosic biomass.
Cell wall profiling of wine and table

grapes
Cortical cytoskeleton and wall
dynamics in a green alga
The site of cell division in plant cells is often marked or
predicted by a cortical band of microtubules called the
pre-prophase band (PPB). Ochs et al. ( pp. 1237 1249) study cell
expansion in the unicellular charophyte green alga, Penium
margaritaceum, and find a cortical cytoskeletal band of
microtubules and actin filaments at the cell isthmus. Like a
pre-prophase band, this marks the future site of cell division and
also serves as the focal point of cell wall deposition during
bi-directional cell expansion. The study demonstrates that a
cortical cytoskeletal aggregate like the PPB appears in early
divergent green plants such as charophytes.
Table grapes are bred for crunchy texture and physical appearance,
whereas wine grapes have been selected for small size and
concentrated flavour and aroma. Moore et al. ( pp. 12791294)
apply cell wall profiling tools to characterize a table grape cultivar,
Crimson Seedless, versus a wine grape, Cabernet Sauvignon,
with respect to changes at three ripening stages: green berry
touch stage, veraison and full-ripe berries. They identify
pectic-b-(1,4)-galactans, extensins and arabinogalactan-proteins
as polymers that differentiate the different phenological stages.
Whilst the general pattern of changes is highly conserved in both
cultivars, elevated levels of pectin epitopes are detected in the
Crimson Seedless samples. They conclude that genetic
developmental programming has a strong influence on the cell wall
changes occurring in ripening Vitis vinifera grape berries.
iii
Brown algae (Phaeophyceae) are marine organisms that are

evolutionary distant from land plants and with a distinctive cell
wall, the detailed organization of which is unclear. Deniaud-Bouet
et al. ( pp. 12031216) use a systematic approach to study polymer
interlinks in five species of the order Fucales, and find that alginates
are mostly associated with phenolic compounds while sulfated
fucans are tightly associated with proteins and cellulose. They
suggest that these two networks sustain distinct roles in the
regulation of wall rigidity and osmotic stress responses,
respectively. The emergence of this cell wall structure may have
been instrumental in the acquisition of multicellularity in brown
algae.
Fern cell wall composition and

functional specialization
Structural variation of RGII in wine

(Research in Context)
Xyloglucan endotransglucosylase/
hydrolases in rice
doi:10.1093/aob/mct292
Rhamnogalacturonan II (RGII) is a structurally complex pectic

sub-domain composed of more than 12 different sugars and
20 different linkages distributed in five side chains along a
homogalacturonan backbone. Buffetto et al. ( pp. 1327 1337)
study RGII from wine (Vitis vinifera, Merlot) and determine
several modifications to its structure. Some of these, such as
dearabinosylation and deacetylation, are shown to be the
consequence of acid treatment, whilst others, such as
methyl-esterification, methyl-etherification and oxidation, reflect
natural diversity. A range of RGII structures exhibiting specific
physico-chemical properties are thus shown to co-exist in wine.
Receptor-like kinases and

maintenance of cell wall integrity
(Review)
Although xyloglucans are ubiquitous in land plants, they are less

abundant in Poales than in eudicotyledons. However,
the xyloglucan endotransglucosylase/hydrolase (XTH) gene
family in rice (Oryza sativa) is comparable in size to that of the
eudicotyledon Arabidopsis thaliana. Hara et al. ( pp. 1309 1318)
study the function of three representative rice XTHs and find all
have xyloglucan endohydrolase (XEH) activity and one has both
XEH and xyloglucan endotransglycosylase activities. Phenotypic
analysis of transgenic lines with altered expression of a rice XTH
suggests that XTHs play redundant roles in rice growth.
Plant cell walls are exposed to a wide range of stress stimuli that
have to be detected by a suitable receptor in order to induce specific
reactions appropriate to the organ affected and the developmental
state of the plant. Engelsdorf and Hamann ( pp. 13391347)
review recent developments in our knowledge on plant cell wall
integrity maintenance with a specific focus on possible signal
elicitors and receptors. Recent evidence implicates receptor-like
kinases (RLKs) in the regulatory networks associated with plant
cell wall-related stress, and hence potential functions of RLKs
in cell wall integrity maintenance are discussed.
Influence of extraction conditions on

citrus pectins
Regulation of stress-induced callose

biosynthesis (Viewpoint)
Pectin is used as a gelling, thickening and emulsifying agent in

a wide range of applications, and current industrial extraction
processes are based on fruit peel, a waste product from the juicing
industry. Kaya et al. ( pp. 13191326) examine how pectin
components vary in relation to the plant source (orange, lemon,
lime, grapefruit) and consider the influence of extraction
conditions on the chemical and macromolecular characteristics
of pectin samples. They find that structural, and hence
macromolecular, variations within the different citrus pectin
samples are mainly related to their rhamnogalacturonan I contents
and integrity, and, to a lesser extent, to the length of their
homogalacturonan domains.
iv
Deposition of callose, a (1,3)-b-glucan cell wall polymer, is

involved in several fundamental biological process, but despite
its importance detailed knowledge about the regulation of its
biosynthesis in plants is rather limited. Ellinger and Voigt
( pp. 1349 1358) summarize data from 10 years of research,
focussing on callose deposition in response to pathogen attack in
the model plant Arabidopsis thaliana. They consider that growing
evidence has been found that the timing of callose deposition in the
multilayered system of plant defence responses could be the key
parameter for optimal effectiveness. This timing seems to be
achieved through co-ordinated transport and formation of the
callose synthase complex.
Fern gametophytes and sporophytes are free-living and show many

morphological and physiological differences. Eeckhout et al.
( pp. 1295 1307) combine glycan microarray analysis with in situ
immunolabelling in order to compare the presence and distribution
of glycan epitopes between both generations in the fern model
system Ceratopteris richardii C-Fern. They find pectic
homogalacturonan, mannan and xyloglucan present in
gametophytic and sporophytic tissues, while xylans and pectic
galactans are only detected in the sporophyte. Rhizoids and root
hairs show a similar arabinogalactan protein and xyloglucan
epitope distribution. The results indicate that cell wall composition
largely reflects functional specialization rather than genetic origin.
Arabinogalactan proteins in
parasitic plant haustoria
Nano-structural changes in pectins

during fruit softening (Review)
Textural changes during fruit ripening are mainly due to the
dissolution of the middle lamella, and pectins, a major component
of fruit cell walls, are extensively modified during this process.
Paniagua et al. ( pp. 1375 1383) discuss the main features of
A surface-spread AGP with a

lysine-rich subdomain
The hyaline body, a central parenchymatous tissue found in the

haustoria of some parasitic plants, is hypothesized to process
nutrients abstracted from the host. Pielach et al. ( pp. 13591373)
carry out the first cell wall-focussed immunocytochemical study of
hyaline bodies using three hemiparasites of semi-natural
grasslands: Rhinanthus minor, Odontites vernus and Melampyrum
pratense. Their results show extensive paramural and intercellular
modifications and the presence of large intercellular globules
suggestive of storage functions. All of these structures are
extremely rich in arabinogalactan proteins (AGPs), a class of
hyperglycosylated proteins, which may participate in nutrient
transfer and metabolism in the hyaline bodies.
pectin disassembly during fruit ripening and review the

nano-structural characterization of pectins and its relationship with
texture and postharvest fruit shelf life, as determined by atomic
force microscopy (AFM). Most AFM studies show a reduction in
the length of individual pectin chains and in the frequency of
aggregates during ripening. Pectins covalently bound to the cell
wall appear to be the most affected. They conclude that AFM
studies can provide valuable insights into the relationships between
pectins and fruit ripening.
Certain membrane-associated arabinogalactan-proteins (AGPs)
with lysine-rich subdomains participate in plant growth,
development and resistance to stress. Zhou et al. ( pp. 1385 1397)
observe one such AGP labelled with EGFP (LeAGP1-EGFP) in a
living leaf cell of Arabidopsis thaliana using confocal microscopy
and on inert chips using atomic force microscopy. They find that in
the selected cell type it can form aligned perimembrane bands, and
viewed at high resolution on the artificial substrates it can form arcs,
rings, clusters and lacy sheets. It can form composite rings of larger
size when the AGP-specific Yariv reagent is added. The
nano-behavior of LeAGP1-EGFP on artificial substrates should be
useful, in combination with other biophysical data, in developing
ideas about the assembly of this regulatory glycoprotein at the
surface of the cell membrane.

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ContentSnapshots

Annals of Botany Volume 114 Number 6 2014

Phosphoinositides and cell walls

Trafficking of the cellulose synthase

AGP31 (arabinogalactan protein 31) is a remarkable cell wall

Recent studies have suggested the involvement of

Formation of secondary cell walls requires co-ordinated

Dynamic calcium recycling by an

Cell-specific transcription and

Glycomodules of cell surface arabinogalactan proteins (AGPs)

A long-standing question in cellular biology is how well the

Synergism between FLA4 and ABA

PME and SBT expression in

Mixed-linkage glucan and

In Arabidopsis thaliana, the degree of methylesterification of

SNARE protein VTI13 and root hair

Tapetum and pollen wall

Endosomal trafficking is required for polarized growth, but many

Rhamnogalacturonan type-II (RG-II) is the most complex pectic

Preserving the ultrastructure of the developing pollen cell wall

The physiological function of fasciclin-like arabinogalactan

Cell-wall structure and evolution in

Seed coat polysaccharide production

Variation of cell wall properties

Cell wall profiling of wine and table

Brown algae (Phaeophyceae) are marine organisms that are

Fern cell wall composition and

Structural variation of RGII in wine

Rhamnogalacturonan II (RGII) is a structurally complex pectic

Receptor-like kinases and

Although xyloglucans are ubiquitous in land plants, they are less

Influence of extraction conditions on

Regulation of stress-induced callose

Pectin is used as a gelling, thickening and emulsifying agent in

Deposition of callose, a (1,3)-b-glucan cell wall polymer, is

Fern gametophytes and sporophytes are free-living and show many

Nano-structural changes in pectins

A surface-spread AGP with a

The hyaline body, a central parenchymatous tissue found in the

pectin disassembly during fruit ripening and review the

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