In plant post-embryonic epidermis mitogen-activated protein kinase (MAPK) signaling promotes differentiation of pavement cells and inhibits initiation of stomata. Stomata are cells specialized to modulate gas exchange and water loss. Arabidopsis MAPKs MPK3 and MPK6 are at the core of the signaling cascade; however, it is not well understood how the activity of these pleiotropic MAPKs is constrained spatially so that pavement cell differentiation is promoted only outside the stomata lineage. Here we identified a PP2C-type phosphatase termed AP2C3 (Arabidopsis protein phosphatase 2C) that is expressed distinctively during stomata development as well as interacts and inactivates MPK3, MPK4 and MPK6. AP2C3 co-localizes with MAPKs within the nucleus and this localization depends on its N-terminal extension. We show that other closely related phosphatases AP2C2 and AP2C4 are also MAPK phosphatases acting on MPK6, but have a distinct expression pattern from AP2C3. In accordance with this, only AP2C3 ectopic expression is able to stimulate cell proliferation leading to excess stomata development. This function of AP2C3 relies on the domains required for MAPK docking and intracellular localization. Concomitantly, the constitutive and inducible AP2C3 expression deregulates E2F-RB pathway, promotes the abundance and activity of CDKA, as well as changes of CDKB1;1 forms. We suggest that AP2C3 downregulates the MAPK signaling activity to help maintain the balance between differentiation of stomata and pavement cells.
Main conclusionPlant RSH proteins are able to synthetize and/or hydrolyze unusual nucleotides called (p)ppGpp or alarmones. These molecules regulate nuclear and chloroplast transcription, chloroplast translation and plant development and stress response.Homologs of bacterial RelA/SpoT proteins, designated RSH, and products of their activity, (p)ppGpp—guanosine tetra—and pentaphosphates, have been found in algae and higher plants. (p)ppGpp were first identified in bacteria as the effectors of the stringent response, a mechanism that orchestrates pleiotropic adaptations to nutritional deprivation and various stress conditions. (p)ppGpp accumulation in bacteria decreases transcription—with exception to genes that help to withstand or overcome current stressful situations, which are upregulated—and translation as well as DNA replication and eventually reduces metabolism and growth but promotes adaptive responses. In plants, RSH are nuclei-encoded and function in chloroplasts, where alarmones are produced and decrease transcription, translation, hormone, lipid and metabolites accumulation and affect photosynthetic efficiency and eventually plant growth and development. During senescence, alarmones coordinate nutrient remobilization and relocation from vegetative tissues into seeds. Despite the high conservancy of RSH protein domains among bacteria and plants as well as the bacterial origin of plant chloroplasts, in plants, unlike in bacteria, (p)ppGpp promote chloroplast DNA replication and division. Next, (p)ppGpp may also perform their functions in cytoplasm, where they would promote plant growth inhibition. Furthermore, (p)ppGpp accumulation also affects nuclear gene expression, i.a., decreases the level of Arabidopsis defense gene transcripts, and promotes plants susceptibility towards Turnip mosaic virus. In this review, we summarize recent findings that show the importance of RSH and (p)ppGpp in plant growth and development, and open an area of research aiming to understand the function of plant RSH in response to stress.
Germination is a crucial process in crop plant life cycle that determines future yield. Hydrogen peroxide plays a significant role in both dormancy release and seed ageing. The potential of the rapeseed metallothioneins (BnMT1‐4) as reactive oxygen species scavengers was investigated. In the presence of H2O2, bacterial cells expressing BnMT1‐4 had higher growth rate than control cells. However, such effect was not observed in an oxidative stress oversensitive yap‐1Δ yeast mutant strain expressing BnMT1‐4. Next, it was shown that H2O2, up to 100 and 10 mM, promotes rapeseed germination rate and seedling growth, respectively. Moreover, it was demonstrated that in seeds germinating in the presence of H2O2, the expression of seed‐specific BnMT4 decreases to a high extent in a dose‐ and time‐dependent manner. It was shown that the decline in BnMT4 mRNA level, accelerated by up to 10 mM H2O2 treatment, correlates positively with increased rapeseed germination and early growth, underscoring the plausible role of BnMT4 as a H2O2 scavenging protein that level has to be downregulated in conditions where the compound is needed. Finally, it was demonstrated that H2O2 could be used as a priming agent of rapeseeds.
Knowledge about past climates, especially at a seasonal time scale, is important as it allows informed decisions to be made to mitigate future climate change. However, globally, and especially in semi-arid Tropics, instrumental climatic data are scarce. A dendroclimatic approach may fill this gap, but tropical dendrochronological data are rare and do not yet provide fine resolution intra-annual information about past climates. Unlike in the Tropics, in the Mediterranean, temperate, alpine, and arctic regions, dendroanatomy and quantitative wood anatomy (QWA) are progressing fast attaining an intra-annual resolution, which allows a better understanding of seasonal climate dynamics and climate–growth relationships. The existing dendroanatomical and QWA methods aren’t suitable for tropical trees because they do not consider the high variation in tree ring width and the frequent occurrence of micro-rings containing only a few tracheids per radial file. The available tracheid analysis programs generally fail to provide multiple sectors for micro-rings and they are unable to compute most of the useful dendroanatomical parameters at fine temporal resolutions. Here, we present a program (SabaTracheid) that addresses the three main standard tasks that are necessary for QWA and dendroanatomy before running a climate analysis: (1) tracheidogram standardization, (2) sectoring, and (3) computing QWA and dendroanatomical variables. SabaTracheid is demonstrated on African Juniper (Juniperus procera Hochst. ex Endl), but it is potentially able to provide fine-resolution QWA and dendroanatomic data that could be used for dendroanatomical studies in all regions of the world. SabaTracheid is a freeware that quickly and accurately standardizes tracheidograms, divides tree rings into multiple regular sectors, computes useful dendroanatomic and QWA variables for the whole tree rings, early- and latewood portions, and each sector separately. This program is particularly adapted to deal with high inter-annual growth variations observed in tropical trees so that it assures the provision of complete sectoral QWA and dendroanatomical data for micro-rings as well. We demonstrate SabaTracheid using a dataset of 30 Juniperus procera tree rings from the Blue Nile basin, in Ethiopia. SabaTracheid’s ability to provide fine resolution QWA and dendroanatomic data will help the discipline develop in tropical as well as in the Mediterranean and temperate regions.
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