Drought (water deficits) and heat (high temperatures) stress are the prime abiotic constraints, under the current and climate change scenario in future. Any further increase in the occurrence, and extremity of these stresses, either individually or in combination, would severely reduce the crop productivity and food security, globally. Although, they obstruct productivity at all crop growth stages, the extent of damage at reproductive phase of crop growth, mainly the seed filling phase, is critical and causes considerable yield losses. Drought and heat stress substantially affect the seed yields by reducing seed size and number, eventually affecting the commercial trait ‘100 seed weight’ and seed quality. Seed filling is influenced by various metabolic processes occurring in the leaves, especially production and translocation of photoassimilates, importing precursors for biosynthesis of seed reserves, minerals and other functional constituents. These processes are highly sensitive to drought and heat, due to involvement of array of diverse enzymes and transporters, located in the leaves and seeds. We highlight here the findings in various food crops showing how their seed composition is drastically impacted at various cellular levels due to drought and heat stresses, applied separately, or in combination. The combined stresses are extremely detrimental for seed yield and its quality, and thus need more attention. Understanding the precise target sites regulating seed filling events in leaves and seeds, and how they are affected by abiotic stresses, is imperative to enhance the seed quality. It is vital to know the physiological, biochemical and genetic mechanisms, which govern the various seed filling events under stress environments, to devise strategies to improve stress tolerance. Converging modern advances in physiology, biochemistry and biotechnology, especially the “omics” technologies might provide a strong impetus to research on this aspect. Such application, along with effective agronomic management system would pave the way in developing crop genotypes/varieties with improved productivity under drought and/or heat stresses.

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Drought or/and heat-stress effects on seed filling in food crops: Impacts on functional biochemistry, seed yields, and nutritional quality

On-going climate change is increasing the risk of drought stress across large areas worldwide. Such drought events decrease ecosystem productivity and have been increasingly linked to tree mortality. Understanding how trees respond to water shortage is key to predicting the future of ecosystem functions. Phloem is at the core of the tree functions, moving resources such as non-structural carbohydrates, nutrients, and defence and information molecules across the whole plant. Phloem function and ability to transport resources is tightly controlled by the balance of carbon and water fluxes within the tree. As such, drought is expected to impact phloem function by decreasing the amount of available water and new photoassimilates. Yet, the effect of drought on the phloem has received surprisingly little attention in the last decades. Here we review existing knowledge on drought impacts on phloem transport from loading and unloading processes at cellular level to possible effects on long-distance transport and consequences to ecosystems via ecophysiological feedbacks. We also point to new research frontiers that need to be explored to improve our understanding of phloem function under drought. In particular, we show how phloem transport is affected differently by increasing drought intensity, from no response to a slowdown, and explore how severe drought might actually disrupt the phloem transport enough to threaten tree survival. Because transport of resources affects other organisms interacting with the tree, we also review the ecological consequences of phloem response to drought and especially predatory, mutualistic and competitive relations. Finally, as phloem is the main path for carbon from sources to sink, we show how drought can affect biogeochemical cycles through changes in phloem transport. Overall, existing knowledge is consistent with the hypotheses that phloem response to drought matters for understanding tree and ecosystem function. However, future research on a large range of species and ecosystems is urgently needed to gain a comprehensive understanding of the question.

/pdf/drought-impacts-on-tree-phloem-from-cell-level-responses-to-29d2wobog5.pdf

Drought impacts on tree phloem: from cell-level responses to ecological significance.

Abiotic stresses, including drought and salinity, negatively affect plant development and physiology at molecular and metabolic levels. Sucrose transport, mediating distribution of photosynthates in plant, is a key physiological process impacted by drought and salinity stresses, as sucrose is a prime energy and signaling molecule as well as an osmolyte. Therefore, understanding the effects of abiotic stresses on sucrose transport and transporters, and underlying genetic and molecular mechanisms, is imperative to maintain sugar homeostasis in plants under stress. Here, we investigated the effects of drought and salinity stresses on sucrose transport and distribution, and on expression levels of genes encoding Sugars Will Eventually be Exported Transporters (SWEETs), along with a potential transcription factor regulating SWEET expression in rice. We observed that drought and salinity stresses increased the sucrose content in leaf and root tissues and in phloem sap of rice indica varieties. Expression analyses of SWEET genes and histochemical analysis of β-glucuronidase-reporter transgenic plants suggested that OsSWEET13 and OsSWEET15 are major SWEET transporters regulating the sucrose transport and levels in response to the abiotic stresses. Transactivation analyses showed that an abscisic acid (ABA)-responsive transcription factor OsbZIP72 directly binds to the promoters of OsSWEET13 and OsSWEET15 and activates their expression. Taken together, the results showed that the higher expressions of OsSWEET13 and OsSWEET15 genes, induced by binding of an ABA-responsive transcription factor OsbZIP72 to the promoters, potentially modulate sucrose transport and distribution in response to the abiotic stresses. The mechanism could possibly be targeted for maintaining sugar homeostasis in rice under drought and salinity stresses.

Sucrose transport in response to drought and salt stress involves ABA-mediated induction of OsSWEET13 and OsSWEET15 in rice.

All multicellular organisms keep a balance between sink and source activities by controlling nutrient transport at strategic positions. In most plants, photosynthetically produced sucrose is the predominant carbon and energy source, whose transport from leaves to carbon sink organs depends on sucrose transporters. In the model plant Arabidopsis thaliana, transport of sucrose into the phloem vascular tissue by SUCROSE TRANSPORTER 2 (SUC2) sets the rate of carbon export from source leaves, just like the SUC2 homologs of most crop plants. Despite their importance, little is known about the proteins that regulate these sucrose transporters. Here, identification and characterization of SUC2-interaction partners revealed that SUC2 activity is regulated via its protein turnover rate and phosphorylation state. UBIQUITIN-CONJUGATING ENZYME 34 (UBC34) was found to trigger turnover of SUC2 in a light-dependent manner. The E2 enzyme UBC34 could ubiquitinate SUC2 in vitro, a function generally associated with E3 ubiquitin ligases. ubc34 mutants showed increased phloem loading, as well as increased biomass and yield. In contrast, mutants of another SUC2-interaction partner, WALL-ASSOCIATED KINASE LIKE 8 (WAKL8), showed decreased phloem loading and growth. An in vivo assay based on a fluorescent sucrose analog confirmed that SUC2 phosphorylation by WAKL8 can increase transport activity. Both proteins are required for the up-regulation of phloem loading in response to increased light intensity. The molecular mechanism of SUC2 regulation elucidated here provides promising targets for the biotechnological enhancement of source strength.

https://www.pnas.org/content/pnas/117/11/6223.full.pdf

Carbon export from leaves is controlled via ubiquitination and phosphorylation of sucrose transporter SUC2.

Carbon partitioning mechanisms in POTATO under drought stress

Suc transporters (SUTs) play a key role in the allocation and partitioning of photosynthetically fixed carbon in plants. While a function could be assigned to many members of the SUT family, almost no information is available on their regulation. Here, the transcriptional regulation of SUTs in response to various environmental stimuli in the leaves of five dicots (Arabidopsis [Arabidopsis thaliana], soybean [Glycine max], potato [Solanum tuberosum], tomato [Solanumlycopersicum], and poplar [Populus spp.]) and four monocots (maize [Zeamays], rice [Oryza sativa], wheat [Triticum aestivum], and barley [Hordeum vulgare]) was investigated. Extensive data on expression of SUTs in relation to changes of environmental conditions were obtained through a global analysis of 168 transcriptomics data sets. Results were validated by quantitative PCR measurements and extended by the measurement of photosynthesis rate and phloem sugar content to draw insight on the correlation of SUT expression and sugar export from leaves. For the apoplasmic phloem loaders, a clear difference in transcriptional regulation in response to different environmental stimuli was observed. The consistent patterns of SUT expression under abiotic stress indicates which types of SUTs are involved in the regulation of leaf sugar status and in stress signaling. Furthermore, it is shown that down-regulation of phloem loading is likely to be caused by transcriptional regulation of SUTs, while up-regulation depends on post-transcriptional regulation. In poplar, expression of PtaSUT4 was found to consistently respond to environmental stimuli, suggesting a significant role in the regulation of sugar export from leaves in this passive symplasmic phloem loader.

Regulation of Sucrose Transporters and Phloem Loading in Response to Environmental Cues

Sub-Cellular Distribution of Zinc in Different Vegetative Organs and Their Contribution to Grains Zinc Accumulation in Rice Under Different Nitrogen and Zinc Supply

Siyuan Chen

Papers

Regulation of Sucrose Transporters and Phloem Loading in Response to Environmental Cues

Sub-Cellular Distribution of Zinc in Different Vegetative Organs and Their Contribution to Grains Zinc Accumulation in Rice Under Different Nitrogen and Zinc Supply

Alleviating effects of zinc and 24-epibrassionlide on cadmium accumulation in rice plants under nitrogen application.