Bio-communication of Plants

Plants communicate with a great variety of symbiotic partners, above and below ground. Constant monitoring of signals of biotic origin as well as abiotic environmental influences allows plants to generate appropriate response behavior. These communication processes are primarily sign-mediated interactions and not simply an exchange of information. They involve active coordination and active organization of a great variety of different behavioural patterns – mediated by signs

Callose homeostasis at plasmodesmata: molecular regulators and developmental relevance

Plasmodesmata are membrane-lined channels that are located in the plant cell wall and that physically interconnect the cytoplasm and the endoplasmic reticulum (ER) of adjacent cells. Operating as controllable gates, plasmodesmata regulate the symplastic trafficking of micro- and macromolecules, such as endogenous proteins [transcription factors (TFs)] and RNA-based signals (mRNA, siRNA, etc.), hence mediating direct cell-to-cell communication and long distance signaling. Besides this physiological role, plasmodesmata also form gateways through which viral genomes can pass, largely facilitating the pernicious spread of viral infections. Plasmodesmatal trafficking is either passive (e.g., diffusion) or active and responses both to developmental and environmental stimuli. In general, plasmodesmatal conductivity is regulated by the controlled build-up of callose at the plasmodesmatal neck, largely mediated by the antagonistic action of callose synthases (CalSs) and beta-1,3-glucanases. Here, in this theory and hypothesis paper, we outline the importance of callose metabolism in PD SEL control, and highlight the main molecular factors involved. In addition, we also review other proteins that regulate symplastic PD transport, both in a developmental and stress-responsive framework, and discuss on their putative role in the modulation of PD callose turn-over. Finally, we hypothesize on the role of structural sterols in the regulation of (PD) callose deposition and outline putative mechanisms by which this regulation may occur

Symplastic communication in organ formation and tissue patterning.

Communication between cells is a crucial step to coordinate organ formation and tissue patterning. In plants, the intercellular transport of metabolites and signalling molecules occur symplastically through membranous structures (named plasmodesmata) that traverse the cell wall to connect the cytoplasm and endoplasmic reticulum of neighbouring cells. This review aims to highlight the importance of symplastic communication in plant development. We revisit current literature reporting the effects of changing plasmodesmata in cell morphogenesis, organ initiation and meristem maintenance and comment on recent work involving the identification of novel plasmodesmata regulators and of mobile developmental proteins and RNA molecules. New opportunities for unravelling the dynamic regulation and function of plasmodesmata are also discussed.EPSRC (Grant ID: EP/MO27740/1)This is the author accepted manuscript. The final version is available from Elsevier via http://dx.doi.org/10.1016/j.pbi.2015.10.00

Comparative ultrastructure of plasmodesmata of Chara and selected bryophytes: towards an elucidation of the evolutionary origin of plant plasmodesmata

We have used transmission electron microscopy to examine plasmodesmata of the charophycean green alga Chara zeylanica, and of the putatively early divergent bryophytes Monoclea gottschei (liverwort), Notothylas orbicularis (hornwort), and Sphagnum fimbriatum (moss), in an attempt to learn when seed plant plasmodesmata may have originated. The three bryophytes examined have desmotubules. In addition, Monoclea was found to have branched plasmodesmata, and plasmodesmata of Sphagnum displayed densely staining regions around the neck region, as well as ring-like wall specializations. In Chara, longitudinal sections revealed endoplasmic reticulum (ER) that sometimes appeared to be associated with plasmodesmata, but this was rare, despite abundant ER at the cell periphery. Across all three fixation methods, cross-sectional views showed an internal central structure, which in some cases appeared to be connected to the plasma membrane via spoke-like structures. Plasmodesmata were present even in the incompletely formed reticulum of forming cell plates, from which we conclude that primary plasmodesmata are formed at cytokinesis in Chara zeylanica. Based on these results it appears that plasmodesmata of Chara may be less specialized than those of seed plants, and that complex plasmodesmata probably evolved in the ancestor of land plants before extant lineages of bryophytes diverged

Towards reconciliation of structure with function in plasmodesmata—who is the gatekeeper?

Whilst the structure of higher plant plasmodesmata was first described by Robards (1963. Desmotubule—a plasmodesmatal substructure. Nature 218, 784), and despite many subsequent intensive investigations, there is still much that remains unclear relating to their ultrastructure and functioning in higher plants. We have examined chemically fixed plant material, and suggest that the conformational changes seen in plasmodesmatal substructure, particularly the deposition of electron-dense extra-plasmodesmal material, is linked to either manipulation of the hormonal balance (as in Avocado fruit), or of osmotic potential in leaf blade material. These changes result in the deposition of β 1,3-glucan (callose) at the neck region of these plasmodesmata. This electron-dense material is deposited at the neck region of plasmodesmata, and forms a collar-like structure. The formation of a collar is shown to be coupled with loss of lucence within the cytoplasmic sleeve. The formation of a collar at the plasmodesmatal orifice thus results in encapsulation and closure of the plasmodesmatal orifice. Closure of the orifice coincides with a loss of electron-lucence and a lack of resolution of the desmotubule. These ultrastructural changes are potentially significant and could contribute to, result in, or assist in the down-regulation of cell to cell trafficking via plasmodesmata

Phloem loading in the sucrose-export-defective (SXD-1) mutant maize is limited by callose deposition at plasmodesmata in bundle sheath-vascular parenchyma interface

Using Lucifer Yellow we have demonstrated that the phloem-loading pathway from the mesophyll to the bundle sheath-vascular parenchyma interface in Zea mays source leaves follows a symplasmic route in small and intermediate vascular bundles in control as well as in the green sections of mutant sucrose-export-defective (SXD-1) plants. In the anthocyanin-rich mutant leaf sections, Lucifer Yellow transport was prohibited along the same path, at the bundle sheath-vascular parenchyma interface in particular. Plasmodesmata at the latter interface in SXD-1 anthocyanin-rich leaf sections appear to be structurally altered through callose deposition at the plasmodesmal orifices. We suggest that a transport bottleneck at the bundle sheath-vascular parenchyma interface is thus orchestrated and regulated through callose formation, preventing symplasmic transport across this important loading interface

A Thioredoxin Domain-Containing Protein Interacts with Pepino mosaic virus Triple Gene Block Protein 1

Pepino mosaic virus (PepMV) is a mechanically-transmitted tomato pathogen of importance worldwide. Interactions between the PepMV coat protein and triple gene block protein (TGBp1) with the host heat shock cognate protein 70 and catalase 1 (CAT1), respectively, have been previously reported by our lab. In this study, a novel tomato interactor (SlTXND9) was shown to bind the PepMV TGBp1 in yeast-two-hybrid screening, in vitro pull-down and bimolecular fluorescent complementation (BiFC) assays. SlTXND9 possesses part of the conserved thioredoxin (TRX) active site sequence (W__PC vs. WCXPC), and TXND9 orthologues cluster within the TRX phylogenetic superfamilyclosesttophosducin-likeprotein-3. InPepMV-infectedandhealthyNicotianabenthamiana plants,NbTXND9mRNAlevelswerecomparable,andexpressionlevelsremainedstableinbothlocal and systemic leaves for 10 days post inoculation (dpi), as was also the case for catalase 1 (CAT1). To localize the TXND9 in plant cells, a polyclonal antiserum was produced. Purified α-SlTXND9 immunoglobulin (IgG) consistently detected a set of three protein bands in the range of 27–35 kDa, in the 1000 and 30,000 g pellets, and the soluble fraction of extracts of healthy and PepMV-infected N. benthamiana leaves, but not in the cell wall. These bands likely consist of the homologous protein NbTXND9 and its post-translationally modified derivatives. On electron microscopy, immuno-gold labellingofultrathinsectionsofPepMV-infectedN.benthamianaleavesusingα-SlTXND9IgGrevealed particle accumulation close to plasmodesmata, suggesting a role in virus movement. Taken together, this study highlights a novel tomato-PepMV protein interaction and provides data on its localization in planta. Currently, studies focusing on the biological function of this interaction during PepMV infection are in progress

Traffic into silence: endomembranes and post-transcriptional RNA silencing.

microRNAs (miRNAs) and small interfering RNAs (siRNAs) are small RNAs that repress gene expression at the post-transcriptional level in plants and animals. Small RNAs guide Argonaute-containing RNA-induced silencing complexes to target RNAs in a sequence-specific manner, resulting in mRNA deadenylation followed by exonucleolytic decay, mRNA endonucleolytic cleavage, or translational inhibition. Although our knowledge of small RNA biogenesis, turnover, and mechanisms of action has dramatically expanded in the past decade, the subcellular location of small RNA-mediated RNA silencing still needs to be defined. In contrast to the prevalent presumption that RNA silencing occurs in the cytosol, emerging evidence reveals connections between the endomembrane system and small RNA activities in plants and animals. Here, we summarize the work that uncovered this link between small RNAs and endomembrane compartments and present an overview of the involvement of the endomembrane system in various aspects of RNA silencing. We propose that the endomembrane system is an integral component of RNA silencing that has been long overlooked and predict that a marriage between cell biology and RNA biology holds the key to a full understanding of post-transcriptional gene regulation by small RNAs