Annals of Botany Volume 114 Number 6 2014

Phosphoinositides and cell walls

Supra-molecular assembly of
AGP31 in arabidopsis cell walls



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 at Centro Federal de Educação Tecnológica do Paraná 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.

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

A long-standing question in cellular biology is how well the
transcriptome is coupled to the proteome. Rajasundaram et al.
(pp. 1109–1123) 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.


1093/aob/mcu041 doi:10. 1161–1175) identify a group-2 PME (PME17) that is strongly co-expressed. 2014 The physiological function of fasciclin-like arabinogalactan proteins (FLAs) is largely unknown.5 in the processing of PME17 in planta. making chemical fixation for electron microscopy difficult. one interacting with them. ( pp. can be modified by pectin methylesterases (PMEs) and this plays a central role in both plant development and responses to stress. and highlights a need for identifying specific PME–SBT pairs. the degree of methylesterification of homogalacturonans. and shows that the tapetum and middle layer of the anther remain intact into the tricellular pollen and late-uninucleate microspore stages. biochemistry and proteomic approaches. The middle domain is masked by mixed-linkage glucan. Preserving the ultrastructure of the developing pollen cell wall presents challenges because the key cell types are embedded deep within the anther. 1135–1145) 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. Quilichini et al. suggesting that sialyltransferase-like proteins are required for proper pollen tube growth by maintaining the integrity of RG-II. Space for glucan is provided along the middle domain. Seifert et al. the proportion of which increases during cell elongation.5). Using a combination of functional genomics.1093/aob/mcu035 Mixed-linkage glucan and glucuronoarabinoxylan in roots In Arabidopsis thaliana. 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. 1147–1159) identify VTI13 as a vesicular soluble NSF attachment receptor (SNARE) protein required for root hair growth in Arabidopsis thaliana. . 1189–1201) 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. Analyses of the two heterozygous lines reveals a strong reduction in pollen germination and pollen tube growth in vitro and in vivo. Kozlova et al.1093/aob/mcu093 Plant cell enlargement is unambiguously coupled to changes in cell wall architecture. ( pp. possibly involved in the transfer on the homogalacturonan backbone of Kdo and/or Dha.oxfordjournals. Rhamnogalacturonan-II in pollen tube growth (Research in Context) doi:10. Se´ne´chal et al. with a specific subtilisin-type serine protease (SBT3.1093/aob/mcu042 Endosomal trafficking is required for polarized growth. with consequent effects on growth. They find that fla4 shows a reduced level of ABA responsive transcripts. two specific sugars of RG-II. both spatially and temporally. PME activity is modified in roots of knock-out mutants for both proteins. They demonstrate that VTI13 localizes to the vacuole membrane and the trans-Golgi network (TGN) or an early endosomal compartment. Synergism between ABA and FLA4 is further supported by suppression of the fla4 phenotype by chemical inhibition of ABA catabolism. and a middle domain between the two. Larson et al. 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. ( pp.Synergism between FLA4 and ABA signalling in roots PME and SBT expression in arabidopsis doi:10.1093/aob/mcu125 doi:10. SNARE protein VTI13 and root hair growth in arabidopsis Tapetum and pollen wall ultrastructure in arabidopsis doi:10. The technique reveals the ultrastructure of tapetosomes and at Centro Federal de Educação Tecnológica do Paraná on December 1. ( pp. respectively. and is likely to play a role in the transport of cargo to the large vacuole in root hairs. Downloaded from http://aob. In addition. but many of the proteins that control this process have yet to be identified. This suggests a role for SBT3. is important for TGN integrity. (pp. ii Rhamnogalacturonan type-II (RG-II) is the most complex pectic polysaccharide of the plant cell wall and is necessary for strengthening. The cell membrane-localized FLA4 might indirectly influence the sensitivity of cytosolic ABA signalling and thereby link cell wall biosynthesis with the stress response. ( pp. particularly in the roots. Dumont et al. 1177–1188) study pollen tubes of Arabidopsis thaliana and show the presence of specific sugars of RG-II. They also show that VTI13 function is essential for the maintenance of cell wall organization in root hair and root epidermal cells.1093/aob/mcu010 doi:10. they analyse two T-DNA insertion lines in At3g48820 encoding a putative sialyltransferase-like protein. the main pectic constituent of the cell wall.

The study demonstrates that a cortical cytoskeletal aggregate like the PPB appears in early divergent green plants such as charophytes. particularly in the model plant Arabidopsis thaliana. ( pp. doi:10. They report compositional differences between stem and leaf samples to be predominantly associated with structural carbohydrates. Moreover. ‘Cabernet Sauvignon’. 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. xyloglucan. doi:10. implying that some features of land plant cell walls evolved prior to the transition to land. iii Downloaded from http://aob. ( pp. North et al. A key trait for biomass conversion to biofuels and biomaterials is cell wall quality. and the cell wall was possibly a defining structure that enabled the green algal ancestor to colonize land.Cell-wall structure and evolution in brown algae Seed coat polysaccharide production (Review) doi:10. with respect to changes at three ripening stages: green berry touch stage.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. while lignin content does not correlate with ethanol production from leaf biomass. . ( pp. The potential for intra. ( pp. Costa et al. Like a pre-prophase band.1093/aob/mcu011 During seed coat differentiation the epidermal cells of certain species accumulate polysaccharides either to reinforce cell walls. They identify pectic-b-(1. ve´raison and full-ripe berries. Penium margaritaceum. respectively. elevated levels of pectin epitopes are detected in the ‘Crimson Seedless’ samples. Cell wall profiling of wine and table grapes Cortical cytoskeleton and wall dynamics in a green alga doi:10.1093/aob/mcu013 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).1093/aob/mcu171 Charophyte green algae (CGA) are the closest living relatives to land plants. Deniaud-Boue¨t et al. Moore et al. or for release during seed imbibition to form sticky mucilage.1093/aob/mcu053 Table grapes are bred for crunchy texture and physical appearance. mannan. this marks the future site of cell division and also serves as the focal point of cell wall deposition during ‘bi-directional’ cell expansion. Mikkelsen et al. They suggest that these two networks sustain distinct roles in the regulation of wall rigidity and osmotic stress responses.1093/aob/mcu054 Miscanthus represents one of the most promising dedicated lignocellulosic bioenergy crops. whereas wine grapes have been selected for small size and concentrated flavour and at Centro Federal de Educação Tecnológica do Paraná on December 1. 1265–1277) present data on cell wall compositional changes as a function of development and tissue type across 25 selected Miscanthus genotypes. 1203–1216) use a systematic approach to study polymer interlinks in five species of the order Fucales. The emergence of this cell wall structure may have been instrumental in the acquisition of multicellularity in brown algae. Overall. Whilst the general pattern of changes is highly conserved in both cultivars. Ochs et al. ( pp. ( pp. Variation of cell wall properties in Miscanthus Cell wall biosynthesis in charophyte green algae (Research in Context) doi:10. they provide the first evidence that all cellulose synthase-like genes present in early-divergent land plants were already present in CGA. 1251–1263) review the recent exploitation of these cells as a model system for the study of polysaccharide metabolism and properties. including cellulose. ‘Crimson Seedless’. versus a wine grape. 2014 Brown algae (Phaeophyceae) are marine organisms that are evolutionary distant from land plants and with a distinctive cell wall.1093/aob/mcu096 doi:10. They conclude that genetic developmental programming has a strong influence on the cell wall changes occurring in ripening Vitis vinifera grape berries. 1237 – 1249) study cell expansion in the unicellular charophyte green alga. the detailed organization of which is unclear. 1217 – 1236) provide genetic evidence that many of the most important core cell wall polysaccharides have their evolutionary origins in CGA. rather than having evolved as a result of selection pressures inherent in this transition. leaf tissue contributes significantly to total above-ground biomass at all developmental stages. xylan and pectin. extensins and arabinogalactan-proteins as polymers that differentiate the different phenological stages.4)-galactans. and find that alginates are mostly associated with phenolic compounds while sulfated fucans are tightly associated with proteins and cellulose. 1279–1294) apply cell wall profiling tools to characterize a table grape cultivar. as well as arabinonogalactan proteins (AGPs).oxfordjournals. and find a cortical cytoskeletal band of microtubules and actin filaments at the cell isthmus.

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. ( pp. 2014 Fern gametophytes and sporophytes are free-living and show many morphological and physiological differences. Recent evidence implicates receptor-like kinases (RLKs) in the regulatory networks associated with plant cell wall-related stress. to a lesser extent. such as dearabinosylation and deacetylation. lemon. reflect natural at Centro Federal de Educação Tecnológica do Paraná on December 1. 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. iv Deposition of callose. while xylans and pectic galactans are only detected in the sporophyte. mannan and xyloglucan present in gametophytic and sporophytic tissues. Phenotypic analysis of transgenic lines with altered expression of a rice XTH suggests that XTHs play redundant roles in rice growth. 1349– 1358) summarize data from 10 years of research. the xyloglucan endotransglucosylase/hydrolase (XTH) gene family in rice (Oryza sativa) is comparable in size to that of the eudicotyledon Arabidopsis thaliana. and. to the length of their homogalacturonan domains. a waste product from the juicing industry. 1327– 1337) study RGII from wine (Vitis vinifera. However. Ellinger and Voigt ( pp. focussing on callose deposition in response to pathogen attack in the model plant Arabidopsis thaliana. Some of these. are shown to be the consequence of acid treatment. 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. and hence macromolecular.1093/aob/mcu120 Pectin is used as a gelling. Hara et al. is involved in several fundamental biological process. Kaya et al. Receptor-like kinases and maintenance of cell wall integrity (Review) doi:10. ( pp. ‘Merlot’) and determine several modifications to its structure.1093/aob/mcu097 Xyloglucan endotransglucosylase/ hydrolases in rice doi:10. ( pp.3)-b-glucan cell wall polymer. but despite its importance detailed knowledge about the regulation of its biosynthesis in plants is rather limited. Engelsdorf and Hamann ( pp. ( pp. and hence potential functions of RLKs in cell wall integrity maintenance are discussed. 1319–1326) examine how pectin components vary in relation to the plant source (orange. They find that structural. Rhizoids and root hairs show a similar arabinogalactan protein and xyloglucan epitope distribution.oxfordjournals. variations within the different citrus pectin samples are mainly related to their rhamnogalacturonan I contents and integrity. thickening and emulsifying agent in a wide range of applications. 1339–1347) review recent developments in our knowledge on plant cell wall integrity maintenance with a specific focus on possible signal elicitors and receptors. They find pectic homogalacturonan. lime.1093/aob/mcu039 doi:10. Buffetto et al. whilst others. a (1. A range of RGII structures exhibiting specific physico-chemical properties are thus shown to co-exist in wine. Eeckhout et al. they are less abundant in Poales than in eudicotyledons.1093/aob/mcu043 Although xyloglucans are ubiquitous in land plants. Influence of extraction conditions on citrus pectins Regulation of stress-induced callose biosynthesis (Viewpoint) doi:10. 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.Fern cell wall composition and functional specialization Structural variation of RGII in wine (Research in Context) doi:10. . and current industrial extraction processes are based on fruit peel. such as methyl-esterification. grapefruit) and consider the influence of extraction conditions on the chemical and macromolecular characteristics of pectin samples. The results indicate that cell wall composition largely reflects functional specialization rather than genetic origin. Downloaded from http://aob. 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’. methyl-etherification and oxidation. This timing seems to be achieved through co-ordinated transport and formation of the callose synthase complex.1093/aob/mcu150 doi:10.

Pectins covalently bound to the cell wall appear to be the most affected. Odontites vernus and Melampyrum pratense. as determined by atomic force microscopy (AFM).1093/aob/mcu172 Certain membrane-associated arabinogalactan-proteins (AGPs) with lysine-rich subdomains participate in plant growth. Their results show extensive paramural and intercellular modifications and the presence of large intercellular globules suggestive of storage functions. Zhou et al.oxfordjournals. v .1093/aob/mcu121 Nano-structural changes in pectins during fruit softening (Review) doi:10. and pectins. pectin disassembly during fruit ripening and review the nano-structural characterization of pectins and its relationship with texture and postharvest fruit shelf life. 1375– 1383) discuss the main features of A surface-spread AGP with a lysine-rich subdomain Downloaded from http://aob. It can form composite rings of larger size when the AGP-specific Yariv reagent is added. Most AFM studies show a reduction in the length of individual pectin chains and in the frequency of aggregates during ripening. All of these structures are extremely rich in arabinogalactan proteins (AGPs). which may participate in nutrient transfer and metabolism in the hyaline bodies. a central parenchymatous tissue found in the haustoria of some parasitic plants. are extensively modified during this process. a major component of fruit cell walls. They conclude that AFM studies can provide valuable insights into the relationships between pectins and fruit ripening. They find that in the selected cell type it can form aligned perimembrane bands.Arabinogalactan proteins in parasitic plant haustoria doi:10. 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. at Centro Federal de Educação Tecnológica do Paraná on December 1. is hypothesized to process nutrients abstracted from the host. and viewed at high resolution on the artificial substrates it can form arcs. development and resistance to stress. in developing ideas about the assembly of this regulatory glycoprotein at the surface of the cell membrane. rings.1093/aob/mcu149 Textural changes during fruit ripening are mainly due to the dissolution of the middle lamella. 2014 The hyaline body. The nano-behavior of LeAGP1-EGFP on artificial substrates should be useful. 1359–1373) carry out the first cell wall-focussed immunocytochemical study of hyaline bodies using three hemiparasites of semi-natural grasslands: Rhinanthus minor. Pielach et al. in combination with other biophysical data. Paniagua et al. ( pp. ( pp. clusters and lacy sheets. a class of hyperglycosylated proteins. ( pp.