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Summary The ability to accumulate nutrients is a hallmark for living creatures and plants evolved highly effective nutrient transport systems, especially for the uptake of potassium (K+). However, plants also developed mechanisms that enable the rapid extrusion of K+ in combination with anions. The combined release of K+ and anions is probably an ancient extrusion system, as it is found in the Characeae that are closely related to land plants. We postulate that the ion extrusion mechanisms have developed as an emergency valve, which enabled plant cells to rapidly reduce their turgor, and prevent them from bursting. Later in evolution, seed plants adapted this system for various responses, such as the closure of stomata, long‐distance stress waves, dropping of leaves by pulvini, and loading of xylem vessels. We discuss the molecular nature of the transport proteins that are involved in ion extrusion‐based functions of plants and describe the functions that they obtained during evolution. [ABSTRACT FROM AUTHOR]

Summary Characteristic of all conifer needles, the transfusion tissue mediates the radial transport of water and sugar between the endodermis and axial vasculature. Physical constraints imposed by the needle's linear geometry introduce two potential extravascular bottlenecks where the opposition of sugar and water flows may frustrate sugar export: one at the vascular access point and the other at the endodermis. We developed a network model of the transfusion tissue to explore how its structure and composition affect the delivery of sugars to the axial phloem. To describe extravascular transport with cellular resolution, we construct networks from images of Pinus pinea needles obtained through tomographic microscopy, as well as fluorescence and electron microscopy. The transfusion tissue provides physically distinct pathways for sugar and water, reducing resistance between the vasculature and endodermis and mitigating flow constriction at the vascular flank. Dissipation of flow velocities through the transfusion tissue's branched structure allows for bidirectional transport of an inbound diffusive sugar flux against an outbound advective water flux across the endodermis. Our results clarify the structure–function relationships of the transfusion tissue under conditions free of physiological stress. The presented model framework is also applicable to different transfusion tissue morphologies in other gymnosperms. [ABSTRACT FROM AUTHOR]

Enhancing carbohydrate export from source to sink tissues is considered to be a realistic approach for improving photosynthetic efficiency and crop yield. The rice sucrose transporters OsSUT1, OsSWEET11a and OsSWEET14 contribute to sucrose phloem loading and seed filling. Crucially, Xanthomonas oryzae pv. oryzae (Xoo) infection in rice enhances the expression of OsSWEET11a and OsSWEET14 genes, and causes leaf blight. Here we show that co‐overexpression of OsSUT1, OsSWEET11a and OsSWEET14 in rice reduced sucrose synthesis and transport leading to lower growth and yield but reduced susceptibility to Xoo relative to controls. The immunity‐related hypersensitive response (HR) was enhanced in the transformed lines as indicated by the increased expression of defence genes, higher salicylic acid content and presence of HR lesions on the leaves. The results suggest that the increased expression of OsSWEET11a and OsSWEET14 in rice is perceived as a pathogen (Xoo) attack that triggers HR and results in constitutive activation of plant defences that are related to the signalling pathways of pathogen starvation. These findings provide a mechanistic basis for the trade‐off between plant growth and immunity because decreased susceptibility against Xoo compromised plant growth and yield. Summary statement: The study reported in this manuscript concerns the emerging topic of the roles played by SWEET sugar transporters in plant immunity. The data reported in this manuscript show that overexpression of Clade III SWEET sucrose transporters OsSWEET11a and OsSWEET14 that are exploited by bacterial blight pathogen Xanthomonas oryzae pv. oryzae to acquire sugars for its propagation elicits immune responses and activates plant defence responses leading to enhanced resistance against bacterial as well as fungal pathogens. The findings obtained by the overexpression of OsSWEET11a and OsSWEET14 in rice show that sucrose transport occurs via an apoplastic route, and overexpression of these sugar transporters inhibits sucrose synthesis and transport, which explains the inverse relationships between plant growth and yield, and resistance to pathogens. Moreover, our results show that sucrose transport and pathogen responses are coupled processes, and provide evidence for the concept that pathogen‐induced starvation is linked to plant defence. Also, to the best of our knowledge this is the first study where we show that disease susceptibility genes OsSWEET11a and OsSWEET14 can lead to hypersensitive reaction leading to enhanced plant immunity. [ABSTRACT FROM AUTHOR]

SUMMARY: High‐affinity K+ transporters/K+ uptake permeases/K+ transporters (HAK/KUP/KT) are important pathways mediating K+ transport across cell membranes, which function in maintaining K+ homeostasis during plant growth and stress response. An increasing number of studies have shown that HAK/KUP/KT transporters play crucial roles in root K+ uptake and root‐to‐shoot translocation. However, whether HAK/KUP/KT transporters also function in phloem K+ translocation remain unclear. In this study, we revealed that a phloem‐localized rice HAK/KUP/KT transporter, OsHAK18, mediated cell K+ uptake when expressed in yeast, Escherichia coli and Arabidopsis. It was localized at the plasma membrane. Disruption of OsHAK18 rendered rice seedlings insensitive to low‐K+ (LK) stress. After LK stress, some WT leaves showed severe wilting and chlorosis, whereas the corresponding leaves of oshak18 mutant lines (a Tos17 insertion line and two CRISPR lines) remained green and unwilted. Compared with WT, the oshak18 mutants accumulated more K+ in shoots but less K+ in roots after LK stress, leading to a higher shoot/root ratio of K+ per plant. Disruption of OsHAK18 does not affect root K+ uptake and K+ level in xylem sap, but it significantly decreases phloem K+ concentration and inhibits root‐to‐shoot‐to‐root K+ (Rb+) translocation in split‐root assay. These results reveal that OsHAK18 mediates phloem K+ loading and redistribution, whose disruption is in favor of shoot K+ retention under LK stress. Our findings expand the understanding of HAK/KUP/KT transporters' functions and provide a promising strategy for improving rice tolerance to K+ deficiency. Significance Statement: HAK/KUP/KT transporters play important roles in K+ uptake and root‐to‐shoot translocation, but whether HAK/KUP/KT transporters function in phloem K+ translocation remain unclear. Here, we reported that a phloem‐localized rice HAK/KUP/KT transporter, OsHAK18, mediated phloem K+ loading and affected shoot‐to‐root K+ redistribution, which expands the understanding of HAK/KUP/KT transporters' functions and provides a potential strategy for improving rice tolerance to K+ deficiency. [ABSTRACT FROM AUTHOR]

Phloem loading and transport of sugar from leaves to sink tissues such as fruits are crucial for yield formation. Camellia oleifera is an evergreen horticultural crop with high value; however, its low production limits the development of the C. oleifera industry. In this study, using a combination of ultrastructural observation, fluorescence loss in photobleaching (FLIP) and inhibitor treatment, we revealed that C. oleifera leaves mainly adopt a symplastic loading route from mesophyll cells to the surrounding vascular bundle cells in minor veins. HPLC assays showed that sucrose is the main sugar transported and only a small amount of raffinose or stachyose was detected in petioles, supporting a passive symplastic loading route in C. oleifera leaves. Compared to leaves grown this year (LT), the carbohydrate synthesis capacity in leaves grown last year (LL) was decreased while LL retained more soluble sugar, suggesting a decrease in transport capacity with leaf ageing. TEM and tissue staining showed that a reduction in plasmodesmata density leads to a decline in the degree of cellular coupling and is responsible for the weakening transport capacity in older leaves. RNA-seq revealed several differentially expressed genes (DEGs) including CoPDCB1-1, CoSUT1 and CoSWEET12, which are likely involved in the regulation of phloem loading and sugar transport. An expression correlation network is constructed between PD-callose binding protein genes, sugar transporter genes and senescence-associated genes. Collectively, this study provides the evidence of the passive symplastic phloem loading pathway in C. oleifera leaves and constructs the correlation between sugar transport and leaf ageing. [ABSTRACT FROM AUTHOR]

Summary: The sugars will eventually be exported transporters (SWEET) family of transporters in plants is identified as a novel class of sugar carriers capable of transporting sugars, sugar alcohols and hormones. Functioning in intercellular sugar transport, SWEETs influence a wide range of physiologically important processes. SWEETs regulate the development of sink organs by providing nutritional support from source leaves, responses to abiotic stresses by maintaining intracellular sugar concentrations, and host–pathogen interactions through the modulation of apoplastic sugar levels. Many bacterial and fungal pathogens activate the expression of SWEET genes in species such as rice and Arabidopsis to gain access to the nutrients that support virulence. The genetic manipulation of SWEETs has led to the generation of bacterial blight (BB)‐resistant rice varieties. Similarly, while the overexpression of the SWEETs involved in sucrose export from leaves and pathogenesis led to growth retardation and yield penalties, plants overexpressing SWEETs show improved disease resistance. Such findings demonstrate the complex functions of SWEETs in growth and stress tolerance. Here, we review the importance of SWEETs in plant–pathogen and source–sink interactions and abiotic stress resistance. We highlight the possible applications of SWEETs in crop improvement programmes aimed at improving sink and source strengths important for enhancing the sustainability of yield. We discuss how the adverse effects of the overexpression of SWEETs on plant growth may be overcome. [ABSTRACT FROM AUTHOR]

Bioenergy sorghum hybrids are being developed with enhanced drought tolerance and high levels of stem sugars. Raffinose family oligosaccharides (RFOs) contribute to plant environmental stress tolerance, sugar storage, transport, and signaling. To better understand the role of RFOs in sorghum, genes involved in myo-inositol and RFO metabolism were identified and relative transcript abundance analyzed during development. Genes involved in RFO biosynthesis (SbMIPS1, SbInsPase, SbGolS1, SbRS) were more highly expressed in leaves compared to stems and roots, with peak expression early in the morning in leaves. SbGolS, SbRS, SbAGA1 and SbAGA2 were also expressed at high levels in the leaf collar and leaf sheath. In leaf blades, genes involved in myo-inositol biosynthesis (SbMIPS1, SbInsPase) were expressed in bundle sheath cells, whereas genes involved in galactinol and raffinose synthesis (SbGolS1, SbRS) were expressed in mesophyll cells. Furthermore, SbAGA1 and SbAGA2, genes that encode neutral-alkaline alpha-galactosidases that hydrolyze raffinose, were differentially expressed in minor vein bundle sheath cells and major vein and mid-rib vascular and xylem parenchyma. This suggests that raffinose synthesized from sucrose and galactinol in mesophyll cells diffuses into vascular bundles where hydrolysis releases sucrose for long distance phloem transport. Increased expression (>20-fold) of SbAGA1 and SbAGA2 in stem storage pith parenchyma of sweet sorghum between floral initiation and grain maturity, and higher expression in sweet sorghum compared to grain sorghum, indicates these genes may play a key role in non-structural carbohydrate accumulation in stems. [ABSTRACT FROM AUTHOR]

Bioenergy sorghum hybrids are being developed with enhanced drought tolerance and high levels of stem sugars. Raffinose family oligosaccharides (RFOs) contribute to plant environmental stress tolerance, sugar storage, transport, and signaling. To better understand the role of RFOs in sorghum, genes involved in myo-inositol and RFO metabolism were identified and relative transcript abundance analyzed during development. Genes involved in RFO biosynthesis (SbMIPS1, SbInsPase, SbGolS1, SbRS) were more highly expressed in leaves compared to stems and roots, with peak expression early in the morning in leaves. SbGolS, SbRS, SbAGA1 and SbAGA2 were also expressed at high levels in the leaf collar and leaf sheath. In leaf blades, genes involved in myo-inositol biosynthesis (SbMIPS1, SbInsPase) were expressed in bundle sheath cells, whereas genes involved in galactinol and raffinose synthesis (SbGolS1, SbRS) were expressed in mesophyll cells. Furthermore, SbAGA1 and SbAGA2, genes that encode neutral-alkaline alpha-galactosidases that hydrolyze raffinose, were differentially expressed in minor vein bundle sheath cells and major vein and mid-rib vascular and xylem parenchyma. This suggests that raffinose synthesized from sucrose and galactinol in mesophyll cells diffuses into vascular bundles where hydrolysis releases sucrose for long distance phloem transport. Increased expression (>20-fold) of SbAGA1 and SbAGA2 in stem storage pith parenchyma of sweet sorghum between floral initiation and grain maturity, and higher expression in sweet sorghum compared to grain sorghum, indicates these genes may play a key role in non-structural carbohydrate accumulation in stems.

In Arabidopsis, polarized deposition of wall ingrowths in phloem parenchyma (PP) transfer cells (TCs) occurs adjacent to cells of the sieve element/companion cell (SE/CC) complex. However, the spatial relationships between these different cell types in minor veins, where phloem loading occurs, are poorly understood. PP TC development and wall ingrowth localization were compared with those of other phloem cells in leaves of Col-0 and the transgenic lines AtSUC2::AtSTP9-GFP (green fluorescent protein) and AtSWEET11::AtSWEET11-GFP that identify CCs and PP cells, respectively. The development of PP TCs in minor veins, indicated by deposition of wall ingrowths, proceeded basipetally in leaves. However, not all PP cells develop wall ingrowths, and higher levels of deposition occur in abaxial- compared with adaxial-positioned PP TCs. Furthermore, the deposition of wall ingrowths was exclusively initiated on and preferentially covered the PP TC/SE interface, rather than the PP TC/CC interface, and only occurred in PP cells that were adjacent to SEs. Collectively, these results demonstrate a tight association between SEs and wall ingrowth deposition in PP TCs and suggest the existence of two subtypes of PP cells in leaf minor veins. Compared with PP cells, PP TCs showed more abundant accumulation of AtSWEET11–GFP, indicating functional differences in phloem loading between PP and PP TCs. [ABSTRACT FROM AUTHOR]

The role of phloem–xylem interaction via sucrose exchanges in recovery of dehydration impacts, specifically xylem embolism, has not been directly investigated thus far. Most previous studies were indirect approaches leading to suggestive conclusions. We hypothesized that a block in phloem loading and so no exchange of sucrose with xylem affect tolerance and recovery of tobacco plants (Nicotiana tabacum) during dehydration and after the rehydration phase. Transgenic N. tabacum (αNtSUT1-antisense) plants, which showed impaired phloem loading and high accumulation of soluble sugars in leaves, were compared to the wild-type (WT) plants. The water status, osmotic adjustments, leaf turgor, stomatal conductance, xylem cavitation, and stem xylem sucrose content were determined during dehydration and after the rehydration phases. Results showed that retention of sucrose outside phloem conduits highly improved water status, osmotic adjustment and turgidity of the source leaves in the transgenics during drought period. However, no impact occurred on stomata function and tolerance to xylem cavitation in αNtSUT1. After the rehydration period, WT plants with free phloem transport and phloem–-xylem exchange of sucrose recovered better their water status, leaf turgidity, stomatal conductance and xylem functioning than αNtSUT1 plants. The accumulation of sucrose in leaves of transformants ameliorated their tolerance to water deficit by reinforcing the osmotic adjustment mechanism at the leaf level. However, lack of sucrose in phloem sieve resulted in impairment of hydraulic recovery of xylem from drought of αNtSUT1 after rehydration. This suggests a crucial role of the phloem–-xylem exchange of sucrose in refilling of embolized xylem vessels. [ABSTRACT FROM AUTHOR]

Sugars are the major source of energy in living organisms and play important roles in osmotic regulation, cell signaling and energy storage. SWEETs (Sugars Will Eventually be Exported Transporters) are the most recent family of sugar transporters that function as uniporters, facilitating the diffusion of sugar molecules across cell membranes. In plants, SWEETs play roles in multiple physiological processes including phloem loading, senescence, pollen nutrition, grain filling, nectar secretion, abiotic (drought, heat, cold, and salinity) and biotic stress regulation. In this review, we summarized the role of SWEET transporters in plant development and abiotic stress. The gene expression dynamics of various SWEET transporters under various abiotic stresses in different plant species are also discussed. Finally, we discuss the utilization of genome editing tools (TALENs and CRISPR/Cas9) to engineer SWEET genes that can facilitate trait improvement. Overall, recent advancements on SWEETs are highlighted, which could be used for crop trait improvement and abiotic stress tolerance. [ABSTRACT FROM AUTHOR]

SUMMARY: Photosynthetic efficiency and sink demand are tightly correlated with rates of phloem loading, where maintaining low cytosolic sugar concentrations is paramount to prevent the downregulation of photosynthesis. Sugars Will Eventually be Exported Transporters (SWEETs) are thought to have a pivotal role in the apoplastic phloem loading of C4 grasses. SWEETs have not been well studied in C4 species, and their investigation is complicated by photosynthesis taking place across two cell types and, therefore, photoassimilate export can occur from either one. SWEET13 homologues in C4 grasses have been proposed to facilitate apoplastic phloem loading. Here, we provide evidence for this hypothesis using the C4 grass Setaria viridis. Expression analyses on the leaf gradient of C4 species Setaria and Sorghum bicolor show abundant transcript levels for SWEET13 homologues. Carbohydrate profiling along the Setaria leaf shows total sugar content to be significantly higher in the mature leaf tip compared with the younger tissue at the base. We present the first known immunolocalization results for SvSWEET13a and SvSWEET13b using novel isoform‐specific antisera. These results show localization to the bundle sheath and phloem parenchyma cells of both minor and major veins. We further present the first transport kinetics study of C4 monocot SWEETs by using a Xenopus laevis oocyte heterologous expression system. We demonstrate that SvSWEET13a and SvSWEET13b are high‐capacity transporters of glucose and sucrose, with a higher apparent Vmax for sucrose, compared with glucose, typical of clade III SWEETs. Collectively, these results provide evidence for an apoplastic phloem loading pathway in Setaria and possibly other C4 species. [ABSTRACT FROM AUTHOR]

Phloem loading and transport are fundamental processes for allocating carbon from source organs to sink tissues. Cotton (Gossypium spp.) has a high sink demand for the cellulosic fibers that grow on the seed coat and for the storage reserves in the developing embryo, along with the demands of new growth in the shoots and roots. Addressing how cotton mobilizes resources from source leaves to sink organs provides insight into processes contributing to fiber and seed yield. Plasmodesmata frequencies between companion cells and flanking parenchyma in minor veins are higher than expected for an apoplastic loader, and cotton's close relatedness to Tilia spp. hints at passive loading. Suc was the only canonical transport sugar in leaves and constituted 87% of 14C-labeled photoassimilate being actively transported. [14C]Suc uptake coupled with autoradiography indicated active [14C]Suc accumulation in minor veins, suggesting Suc loading from the apoplast; esculin, a fluorescent Suc analog, did not accumulate in minor veins. Of the nine sucrose transporter (SUT) genes identified per diploid genome, only G hSUT1-L2 showed appreciable expression in mature leaves, and silencing GhSUT1-L2 yielded phenotypes characteristic of blocked phloem transport. Furthermore, only G hSUT1-L2 cDNA stimulated esculin and [14C]Suc uptake into yeast, and only the G hSUT1-L2 promoter caused uidA (β-glucuronidase) reporter gene expression in minor vein phloem of Arabidopsis thaliana. Collectively, these results argue that apoplastic phloem loading mediated by GhSUT1-L2 is the dominant mode of phloem loading in cotton. [ABSTRACT FROM AUTHOR]

Plant acclimation and stress responses depend on the dynamic optimization of carbon balance between source and sink organs. This optimization also applies to the leaf export rate of photosynthetically produced sugars. So far, investigations into the molecular mechanisms of how the rate is controlled have focused on sugar transporters responsible for loading sucrose into the phloem sieve element–companion cell complex of leaf veins. Here, we take a broader view of the various proteins with potential direct influence on the leaf sugar export rate in the model plant Arabidopsis thaliana , helped by the cell type-specific transcriptome data that have recently become available. Furthermore, we integrate current information on the regulation of these potential target proteins. Our analysis identifies putative control points and units of transcriptionally and post-transcriptionally co-regulated genes. Most notable is the potential regulatory unit of sucrose transporters (SUC2, SWEET11, SWEET12, and SUC4) and proton pumps (AHA3 and AVP1). Our analysis can guide future research aimed at understanding the regulatory network controlling leaf sugar export by providing starting points for characterizing regulatory strategies and identifying regulatory factors that link sugar export rate to the major signaling pathways. [ABSTRACT FROM AUTHOR]

Summary: The green leaves of plants are optimised for carbon fixation and the production of sugars, which are used as central units of carbon and energy throughout the plant. However, there are physical limits to this optimisation that remain insufficiently understood.Here, quantitative anatomical analysis combined with mathematical modelling and sugar transport rate measurements were used to determine how effectively sugars are exported from the needle‐shaped leaves of conifers in relation to leaf length.Mathematical modelling indicated that phloem anatomy constrains sugar export in long needles. However, we identified two mechanisms by which this constraint is overcome, even in needles longer than 20 cm: (1) the grouping of transport conduits, and (2) a shift in the diurnal rhythm of sugar metabolism and export in needle tips.The efficiency of sugar transport in the phloem can have a significant influence on leaf function. The constraints on sugar export described here for conifer needles are likely to also be relevant in other groups of plants, such as grasses and angiosperm trees. [ABSTRACT FROM AUTHOR]

Phloem loading is the first step in sucrose transport from source leaves to sink organs. The phloem loading strategy in rice remains unclear. To determine the potential phloem loading mechanism in rice, yeast invertase (INV) was overexpressed by a 35S promoter specifically in the cell wall to block sugar transmembrane loading in rice. The transgenic lines exhibited obvious phloem loading suppression characteristics accompanied by the accumulation of sucrose and starch, restricted vegetative growth and decreased grain yields. The decreased sucrose exudation rate with p-chloromercuribenzenesulfonic acid (PCMBS) treatment also indicated that rice actively transported sucrose into the phloem. OsSUT1 (SUCROSE TRANSPORTER 1) showed the highest mRNA levels of the plasma membrane-localized OsSUT s in source leaves. Cross sections of the OsSUT::GUS transgenic plants showed that the expression of OsSUT1 and OsSUT5 occurred in the phloem companion cells. Rice ossut1 mutants showed reduced growth and grain yield, supporting the hypothesis of OsSUT1 acting in phloem loading. Based on these results, we conclude that apoplastic phloem loading plays a major role in the export of sugar from rice leaves. [ABSTRACT FROM AUTHOR]

Sugars are the major source of energy in living organisms and play important roles in osmotic regulation, cell signaling and energy storage. SWEETs (Sugars Will Eventually be Exported Transporters) are the most recent family of sugar transporters that function as uniporters, facilitating the diffusion of sugar molecules across cell membranes. In plants, SWEETs play roles in multiple physiological processes including phloem loading, senescence, pollen nutrition, grain filling, nectar secretion, abiotic (drought, heat, cold, and salinity) and biotic stress regulation. In this review, we summarized the role of SWEET transporters in plant development and abiotic stress. The gene expression dynamics of various SWEET transporters under various abiotic stresses in different plant species are also discussed. Finally, we discuss the utilization of genome editing tools (TALENs and CRISPR/Cas9) to engineer SWEET genes that can facilitate trait improvement. Overall, recent advancements on SWEETs are highlighted, which could be used for crop trait improvement and abiotic stress tolerance.

Abstract Plant sucrose transporters are required for phloem loading, and therefore are essential for plant growth and development. In common beans (Phaseolus vulgaris) there are only two sucrose transporters functionally characterized. Through a previous RNA‐seq study, we identified a putative sucrose transporter in common bean, which we hypothesize to function in import of sucrose into plant cells. In silico analysis revealed that PvSUT1.1 is a putative sucrose‐proton co‐transporter distinct from other characterized sucrose transporters in common bean indicating that this is a previously undescribed transporter protein in beans. Further analysis revealed that PvSUT1.1 shares high protein sequence homology to the phloem loader Arabidopsis SUC2; both have 12 transmembrane domains, a typical characteristic of plant sucrose transporters. Heterologous expression in yeast further showed PvSUT1.1 to be functional and it imported sucrose into yeast cells with a Km of 0.7 mM sucrose. Import of sucrose through PvSUT1.1 is also pH‐dependent with highest uptake at pH 4.0, and activity is lost in the presence of the uncoupler carbonyl cyanide 3‐chlorophenylhydrazone. Consistent with identification of PvSUT1.1 as a Type I transporter, PvSUT1.1 also transports esculin. Finally, PvSUT1.1 showed expression in multiple tissues and the protein was localized to the plasma membrane. The results show that PvSUT1.1 is a sucrose transporter that is probably involved in the uptake of sucrose into source and sink cells. The potential role of PvSUT1.1 in leaf phloem loading of sucrose in common beans and its importance in heat tolerance of reproductive tissues are further discussed.

The SWEET (sugars will eventually be exported transporter) family was identified as a new class of sugar transporters that function as bidirectional uniporters/facilitators and facilitate the diffusion of sugars across cell membranes along a concentration gradient. SWEETs are found widely in plants and play central roles in many biochemical processes, including the phloem loading of sugar for long-distance transport, pollen nutrition, nectar secretion, seed filling, fruit development, plant–pathogen interactions and responses to abiotic stress. This review focuses on advances of the plant SWEETs, including details about their discovery, characteristics of protein structure, evolution and physiological functions. In addition, we discuss the applications of SWEET in plant breeding. This review provides more in-depth and comprehensive information to help elucidate the molecular basis of the function of SWEETs in plants.

The transmission of mobile wound signals along the phloem pathway is essential to the activation of wound-induced systemic response/resistance, which requires an upsurge of jasmonic acid (JA) in the distal undamaged leaves. Among these mobile signals, the electrical signal mediated by the glutamate-dependent activation of several clade three GLUTAMATE RECEPTOR-LIKE (GLR3) proteins is involved in the stimulation of JA production in distal leaves. However, whether JA acts as a mobile wound signal and, if so, how it is transmitted and interacts with the electrical signal remain unclear. Here, we show that JA was translocated from the local to distal leaves in Arabidopsis , and this process was predominantly regulated by two phloem-expressed and plasma membrane-localized jasmonate transporters, AtJAT3 and AtJAT4. In addition to the cooperation between AtJAT3/4 and GLR3.3 in the regulation of long-distance JA translocation, our findings indicate that importer-mediated cell–cell JA transport is important for driving the loading and translocation of JA in the phloem pathway in a self-propagating manner. Whether jasmonate (JA) can function as a mobile wound signal remains unclear. This study shows that two membrs of the jasmonate transporter family, AtJAT3 and AtJAT4, can import JA across the plasma membrane and regulate the translocation of wound-induced JA and systemic response/immunity. The findings suggest that transporter-mediated cell–cell transport drives the long-distance transmission of JA in a self-propagating manner. [ABSTRACT FROM AUTHOR]

Cell wall ingrowths, phloem loading, sucrose, transfer cells. [Extracted from the article]

In Arabidopsis thaliana , phloem parenchyma transfer cells (PPTCs) occur in leaf minor veins and play a pivotal role in phloem loading. Wall ingrowth formation in PPTCs is induced by the phloem loading activity of these cells, which is regulated by sucrose (Suc). The effects of endogenous versus exogenous Suc on wall ingrowth deposition, however, differ. Elevating endogenous Suc levels by increased light enhanced wall ingrowth formation, whereas lowering endogenous Suc levels by dark treatment or genetically in ch-1 resulted in lower levels of deposition. In contrast, exogenously applied Suc, or Suc derived from other organs, repressed wall ingrowth deposition. Analysis of pAtSUC2::GFP plants, used as a marker for phloem loading status, suggested that wall ingrowth formation is correlated with phloem loading activity. Gene expression analysis revealed that exogenous Suc down-regulated expression of AtSWEET11 and 12 , whereas endogenous Suc up-regulated AtSWEET11 expression. Analysis of a TREHALOSE 6-PHOSPHATE (T6P) SYNTHASE overexpression line and the hexokinase (HXK)-null mutant, gin2-1 , suggested that Suc signalling of wall ingrowth formation is independent of T6P and HXK. Collectively, these results are consistent with the conclusion that Suc regulates wall ingrowth formation via affecting Suc exporting activity in PPTCs. [ABSTRACT FROM AUTHOR]

Since C4 photosynthesis was first discovered >50 years ago, researchers have sought to understand how this complex trait evolved from the ancestral C3 photosynthetic machinery on >60 occasions. Despite its repeated emergence across the plant kingdom, C4 photosynthesis is notably rare in trees, with true C4 trees only existing in Euphorbia. Here we consider aspects of the C4 trait that could limit but not preclude the evolution of a C4 tree, including reduced quantum yield, increased energetic demand, reduced adaptive plasticity, evolutionary constraints, and a new theory that the passive symplastic phloem loading mechanism observed in trees, combined with difficulties in maintaining sugar and water transport over a long pathlength, could make C4 photosynthesis largely incompatible with the tree lifeform. We conclude that the transition to a tree habit within C4 lineages as well as the emergence of C4 photosynthesis within pre-existing trees would both face a series of challenges that together explain the global rarity of C4 photosynthesis in trees. The C4 trees in Euphorbia are therefore exceptional in how they have circumvented every potential barrier to the rare C4 tree lifeform. [ABSTRACT FROM AUTHOR]

Plant sucrose transporters are required for phloem loading, and therefore are essential for plant growth and development. In common beans (Phaseolus vulgaris) there are only two sucrose transporters functionally characterized. Through a previous RNA‐seq study, we identified a putative sucrose transporter in common bean, which we hypothesize to function in import of sucrose into plant cells. In silico analysis revealed that PvSUT1.1 is a putative sucrose‐proton co‐transporter distinct from other characterized sucrose transporters in common bean indicating that this is a previously undescribed transporter protein in beans. Further analysis revealed that PvSUT1.1 shares high protein sequence homology to the phloem loader Arabidopsis SUC2; both have 12 transmembrane domains, a typical characteristic of plant sucrose transporters. Heterologous expression in yeast further showed PvSUT1.1 to be functional and it imported sucrose into yeast cells with a Km of 0.7 mM sucrose. Import of sucrose through PvSUT1.1 is also pH‐dependent with highest uptake at pH 4.0, and activity is lost in the presence of the uncoupler carbonyl cyanide 3‐chlorophenylhydrazone. Consistent with identification of PvSUT1.1 as a Type I transporter, PvSUT1.1 also transports esculin. Finally, PvSUT1.1 showed expression in multiple tissues and the protein was localized to the plasma membrane. The results show that PvSUT1.1 is a sucrose transporter that is probably involved in the uptake of sucrose into source and sink cells. The potential role of PvSUT1.1 in leaf phloem loading of sucrose in common beans and its importance in heat tolerance of reproductive tissues are further discussed. [ABSTRACT FROM AUTHOR]

Nitrogen (N)-fixing soybean plants use the ureides allantoin and allantoic acid as major long-distance transport forms of N, but in non-fixing, non-nodulated plants amino acids mainly serve in source-to-sink N allocation. However, some ureides are still synthesized in roots of non-fixing soybean, and our study addresses the role of ureide transport processes in those plants. In previous work, legume ureide permeases (UPSs) were identified that are involved in cellular import of allantoin and allantoic acid. Here, UPS1 from common bean was expressed in the soybean phloem, which resulted in enhanced source-to-sink transport of ureides in the transgenic plants. This was accompanied by increased ureide synthesis and elevated allantoin and allantoic acid root-to-sink transport. Interestingly, amino acid assimilation, xylem transport, and phloem partitioning to sinks were also strongly up-regulated. In addition, photosynthesis and sucrose phloem transport were improved in the transgenic plants. These combined changes in source physiology and assimilate partitioning resulted in increased vegetative growth and improved seed numbers. Overall, the results support that ureide transport processes in non-fixing plants affect source N and carbon acquisition and assimilation as well as source-to-sink translocation of N and carbon assimilates with consequences for plant growth and seed development. [ABSTRACT FROM AUTHOR]

Sucrose/H+ Symporters (SUTs) play an important role in plant growth and yield. They are involved in long? distance transport of sucrose from source leaves to filling grains of cereals through a process called phloem loading. However, the molecular mechanism of sucrose transport through SUTs is not yet known. Understanding the key residues involved in sucrose transport can be helpful in developing high yielding varieties through genetic engineering, gene editing or allele mining. Here, the molecular model of OsSUT1 developed based on refined target-template alignment using Modeller software provides structural insights on the sucrose transport mechanism. We propose 13 putative sucrose binding residues and 11 putative H+ binding residues involved in sucrose/H+ co-transport in OsSUT1. [ABSTRACT FROM AUTHOR]

Sucrose synthase (SuSy) is one of two enzyme families capable of catalyzing the first degradative step in sucrose utilization. Several earlier studies examining SuSy mutants in Arabidopsis failed to identify obvious phenotypic abnormalities compared with wild-type plants in normal growth environments, and as such a functional role for SuSy in the previously proposed cellulose biosynthetic process remains unclear. Our study systematically evaluated the precise subcellular localization of all six isoforms of Arabidopsis SuSy via live-cell imaging. We showed that yellow fluorescent protein (YFP)-labeled SuSy1 and SuSy4 were expressed exclusively in phloem companion cells, and the sus1/sus4 double mutant accumulated sucrose under hypoxic conditions. SuSy5 and SuSy6 were found to be parietally localized in sieve elements and restricted only to the cytoplasm. SuSy2 was present in the endosperm and embryo of developing seeds, and SuSy3 was localized to the embryo and leaf stomata. No single isoform of SuSy was detected in developing xylem tissue of elongating stem, the primary site of cellulose deposition in plants. SuSy1 and SuSy4 were also undetectable in the protoxylem tracheary elements, which were induced by the vascular-related transcription factor VND7 during secondary cell wall formation. These findings implicate SuSy in the biological events related to sucrose translocation in phloem. [ABSTRACT FROM AUTHOR]

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-dependentmanner. 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, WALLASSOCIATED KINASE LIKE 8 (WAKL8), showed decreased phloem loading and growth. An in vivo assay based on a fluorescent sucrose analog confirmed that SUC2 phosphorylation byWAKL8 can increase transport activity. Both proteins are required for the up-regulation of phloemloading in response to increased light intensity. The molecular mechanism of SUC2 regulation elucidated here provides promising targets for the biotechnological enhancement of source strength. [ABSTRACT FROM AUTHOR]

Melon production is often compromised by viral diseases, which cannot be treated with chemicals. Therefore, the use of genetic resistances is the main strategy for generating crops resistant to viruses. Resistance to Cucumber mosaic virus (CMV) in melon is scarcely described in few accessions. Until recently, the only known resistant accessions were Freeman’s Cucumber and PI 161375, cultivar Songwhan Charmi (SC). Resistance to CMV in melon is recessive and generally oligogenic and quantitative. However, in SC, the resistance to CMV strains of subgroup II is monogenic, depending only on one gene, cmv1, which is able to stop CMV movement by restricting the virus to the bundle sheath cells and preventing a systemic infection. This restriction depends on the viral movement protein (MP). Chimeric viruses carrying the MP of subgroup II strains, like the strain LS (CMV-LS), are restricted in the bundle sheath cells, whereas those carrying MP from subgroup I, like the strain FNY (CMV-FNY), are able to overcome this restriction. cmv1 encodes a vacuolar protein sorting 41 (CmVPS41), a protein involved in the transport of cargo proteins from the Golgi to the vacuole through late endosomes. We have analyzed the variability of the gene CmVPS41 in a set of 52 melon accessions belonging to 15 melon groups, both from the spp melo and the spp agrestis. We have identified 16 different haplotypes, encoding 12 different CmVPS41 protein variants. Challenging members of all haplotypes with CMV-LS, we have identified nine new resistant accessions. The resistance correlates with the presence of two mutations, either L348R, previously found in the accession SC and present in other three melon genotypes, or G85E, present in Freeman’s Cucumber and found also in four additional melon genotypes. Moreover, the new resistant accessions belong to three different melon horticultural groups, Conomon, Makuwa, and Dudaim. In the new resistant accessions, the virus was able to replicate and move cell to cell, but was not able to reach the phloem. Therefore, resistance to phloem entry seems to be a general strategy in melon controlled by CmVPS41. Finally, the newly reported resistant accessions broaden the possibilities for the use of genetic resistances in new melon breeding strategies.

Generally, tree species load photoassimilates passively into the phloem, while herbaceous species load actively. These phloem loading strategies have implications for phloem sugar concentration and growth potential. Whereas, in previous research, phloem loading identification was performed with 14C-autoradiography, we suggest 11C-autoradiography, because of its compatibility with plant-PET (positron emission tomography) scans. Because 11C-autoradiography has been hardly used in plant sciences so far, it was tested in contrasting plant species: one temperate tree species, Populus tremula L., three tropical tree species, Erythrophleum suaveolens (Guill. & Perr.) Brenan, E. ivorense A. Chev., and Maesopsis eminii Engl., and two herbaceous crop species Solanum lycopersicum L. and S. tuberosum L. Our results confirmed that P. tremula is a passive loader, and Solanum spp. are active loaders. Erythrophleum spp. and young leaves of M. eminii showed the expected passive loading strategy, but the mature leaves of M. eminii showed an uncommon pattern. Images corrected for leaf tissue thickness supported that mature leaves of M. eminii used active phloem loading, which is linked to continuous investment in growth and new leaves, supporting the lower carbon storage levels often observed in tropical tree species. With this study, we demonstrate that 11C-autoradiography is a powerful tool to acquire detailed tracer distribution in leaves to typify phloem loading strategies in plant species.

Storage roots are the main sink for photo-assimilate accumulation and reflect cassava yield and productivity. Regulation of sugar partitioning from leaves to storage roots has not been elucidated. Cell wall invertases are involved in the hydrolysis of sugar during phloem unloading of vascular plants to control plant development and sink strength but have rarely been studied in root crops like cassava. MeCWINV3 encodes a typical cell wall invertase in cassava and is mainly expressed in vascular bundles. The gene is highly expressed in leaves, especially mature leaves, in response to diurnal rhythm. When MeCWINV3 was overexpressed in cassava, sugar export from leaves to storage roots was largely inhibited and sucrose hydrolysis in leaves was accelerated, leading to increased transient starch accumulation by blocking starch degradation and reduced overall plant growth. The progress of leaf senescence was promoted in the MeCWINV3 over-expressed cassava plants with increased expression of senescence-related genes. Storage root development was also delayed because of dramatically reduced sugar allocation from leaves. As a result, the transcriptional expression of starch biosynthetic genes such as small subunit ADP-glucose pyrophosphorylase, granule-bound starch synthase I, and starch branching enzyme I was reduced in accordance with insufficient sugar supply in the storage roots of the transgenic plants. These results show that MeCWINV3 regulates sugar allocation from source to sink and maintains sugar balance in cassava, thus affecting yield of cassava storage roots.

Here, isotopically labeled 68 ZnO NPs (ZnO NPs) and 68 ZnO NPs with a thin 68 Zn 3 (PO 4 ) 2 shell (ZnO_Ph NPs) were foliarly applied (40 μg Zn) to pepper plants ( Capsicum annuum ) to determine the effect of surface chemistry of ZnO NPs on the Zn uptake and systemic translocation to plant organs over 6 weeks. Despite similar dissolution of both Zn-based NPs after 3 weeks, the Zn 3 (PO 4 ) 2 shell on ZnO_Ph NPs (48 ± 12 nm; -18.1 ± 0.6 mV) enabled a leaf uptake of 2.31 ± 0.34 μg of Zn, which is 2.7 times higher than the 0.86 ± 0.18 μg of Zn observed for ZnO NPs (26 ± 8 nm; 14.6 ± 0.4 mV). Further, ZnO_Ph NPs led to higher Zn mobility and phloem loading, while Zn from ZnO NPs was stored in the epidermal tissues, possibly through cell wall immobilization as a storage strategy. These differences led to higher translocation of Zn from the ZnO_Ph NPs within all plant compartments. ZnO_Ph NPs were also more persistent as NPs in the exposed leaf and in the plant stem over time. As a result, the treatment of ZnO_Ph NPs induced significantly higher Zn transport to the fruit than ZnO NPs. As determined by spICP-TOFMS, Zn in the fruit was not in the NP form. These results suggest that the Zn 3 (PO 4 ) 2 shell on ZnO NPs can help promote the transport of Zn to pepper fruits when foliarly applied. This work provides insight into the role of Zn 3 (PO 4 ) 2 on the surface of ZnO NPs in foliar uptake and in planta biodistribution for improving Zn delivery to edible plant parts and ultimately improving the Zn content in food for human consumption.

Melon production is often compromised by viral diseases, which cannot be treated with chemicals. Therefore, the use of genetic resistances is the main strategy for generating crops resistant to viruses. Resistance to Cucumber mosaic virus (CMV) in melon is scarcely described in few accessions. Until recently, the only known resistant accessions were Freeman's Cucumber and PI 161375, cultivar Songwhan Charmi (SC). Resistance to CMV in melon is recessive and generally oligogenic and quantitative. However, in SC, the resistance to CMV strains of subgroup II is monogenic, depending only on one gene, cmv1 , which is able to stop CMV movement by restricting the virus to the bundle sheath cells and preventing a systemic infection. This restriction depends on the viral movement protein (MP). Chimeric viruses carrying the MP of subgroup II strains, like the strain LS (CMV-LS), are restricted in the bundle sheath cells, whereas those carrying MP from subgroup I, like the strain FNY (CMV-FNY), are able to overcome this restriction. cmv1 encodes a vacuolar protein sorting 41 (CmVPS41), a protein involved in the transport of cargo proteins from the Golgi to the vacuole through late endosomes. We have analyzed the variability of the gene CmVPS41 in a set of 52 melon accessions belonging to 15 melon groups, both from the spp melo and the spp agrestis. We have identified 16 different haplotypes, encoding 12 different CmVPS41 protein variants. Challenging members of all haplotypes with CMV-LS, we have identified nine new resistant accessions. The resistance correlates with the presence of two mutations, either L348R, previously found in the accession SC and present in other three melon genotypes, or G85E, present in Freeman's Cucumber and found also in four additional melon genotypes. Moreover, the new resistant accessions belong to three different melon horticultural groups, Conomon, Makuwa, and Dudaim. In the new resistant accessions, the virus was able to replicate and move cell to cell, but was not able to reach the phloem. Therefore, resistance to phloem entry seems to be a general strategy in melon controlled by CmVPS41. Finally, the newly reported resistant accessions broaden the possibilities for the use of genetic resistances in new melon breeding strategies. [ABSTRACT FROM AUTHOR]

Key message: Present review describes the structure, evolution, transport mechanism and physiological functions of SWEETs. Their application using TALENs and CRISPR/CAS9 based genomic editing approach is discussed. Sugars Will Eventually be Exported Transporters (SWEET) proteins were first identified in plants as the novel family of sugar transporters which mediates the translocation of sugars across cell membranes. The SWEET family of sugar transporters is unique in terms of their structure which contains seven predicted transmembrane domains with two internal triple-helix bundles which possibly originate due to prokaryotic gene duplication. SWEETs perform diverse physiological functions such as pollen nutrition, nectar secretion, seed filling, phloem loading, and pathogen nutrition which we have discussed in the present review. We also discuss how transcriptional activator-like effector nucleases (TALENs) and CRISPR/CAS9 genome editing tools are used to engineer SWEET mutants which modulate pathogen resistance in plants and its applications in the field of agriculture. The expression of SWEETs promises to implement insights into many other cellular transport mechanisms. To conclude, the present review highlights the recent aspects which will further develop better understanding of molecular evolution, structure, and function of SWEET transporters in plants. [ABSTRACT FROM AUTHOR]

Most conifer species have needle-shaped leaves that are only a few centimeters long. In general, variation in leaf size has been associated with environmental factors, such as cold or drought stress. However, it has recently been proposed that sugar export efficiency is the limiting factor for conifer needle length, based on the results obtained using a mathematical model of phloem transport. Here, phloem transport rates in long conifer needles were experimentally determined to test if the mathematical model accurately represents phloem transport. The validity of the model's assumptions was tested by anatomical analyses and sugar quantification. Furthermore, various environmental and physiological factors were tested for their correlation with needle length. The results indicate that needle length is not limited by sugar transport efficiency, but, instead, by winter temperatures and light availability. The identification of factors that influence needle size is instrumental for using this trait as a variable in breeding programs. [ABSTRACT FROM AUTHOR]

Amino acids are not only a nitrogen source that can be directly absorbed by plants, but also the major transport form of organic nitrogen in plants. A large number of amino acid transporters have been identified in different plant species. Despite belonging to different families, these amino acid transporters usually exhibit some general features, such as broad expression pattern and substrate selectivity. This review mainly focuses on transporters involved in amino acid uptake, phloem loading and unloading, xylem-phloem transfer, import into seed and intracellular transport in plants. We summarize the other physiological roles mediated by amino acid transporters, including development regulation, abiotic stress tolerance and defense response. Finally, we discuss the potential applications of amino acid transporters for crop genetic improvement.

High-affinity K + transporters/K + uptake permeases/K + transporters (HAK/KUP/KT) are important pathways mediating K + transport across cell membranes, which function in maintaining K + homeostasis during plant growth and stress response. An increasing number of studies have shown that HAK/KUP/KT transporters play crucial roles in root K + uptake and root-to-shoot translocation. However, whether HAK/KUP/KT transporters also function in phloem K + translocation remain unclear. In this study, we revealed that a phloem-localized rice HAK/KUP/KT transporter, OsHAK18, mediated cell K + uptake when expressed in yeast, Escherichia coli and Arabidopsis. It was localized at the plasma membrane. Disruption of OsHAK18 rendered rice seedlings insensitive to low-K + (LK) stress. After LK stress, some WT leaves showed severe wilting and chlorosis, whereas the corresponding leaves of oshak18 mutant lines (a Tos17 insertion line and two CRISPR lines) remained green and unwilted. Compared with WT, the oshak18 mutants accumulated more K + in shoots but less K + in roots after LK stress, leading to a higher shoot/root ratio of K + per plant. Disruption of OsHAK18 does not affect root K + uptake and K + level in xylem sap, but it significantly decreases phloem K + concentration and inhibits root-to-shoot-to-root K + (Rb + ) translocation in split-root assay. These results reveal that OsHAK18 mediates phloem K + loading and redistribution, whose disruption is in favor of shoot K + retention under LK stress. Our findings expand the understanding of HAK/KUP/KT transporters' functions and provide a promising strategy for improving rice tolerance to K + deficiency., (© 2023 Society for Experimental Biology and John Wiley & Sons Ltd.)

Main conclusion: Sucrose concentration in phloem sap was several times higher than in the cytosol of mesophyll cells. The results suggest that phloem loading involves active steps in the analyzed tree species.Phloem loading in source leaves is a key step for carbon partitioning and passive symplastic loading has been proposed for several tree species. However, experimental evidence to prove the potential for sucrose diffusion from mesophyll to phloem is rare. Here, we analyzed three tree species (two angiosperms, Fagus sylvatica, Magnolia kobus, and one gymnosperm, Gnetum gnemon) to investigate the proposed phloem loading mechanism. For this purpose, the minor vein structure and the sugar concentrations in phloem sap as well as in the subcellular compartments of mesophyll cells were investigated. The analyzed tree species belong to the open type minor vein subcategory. The sucrose concentration in the cytosol of mesophyll cells ranged between 75 and 165 mM and was almost equal to the vacuolar concentration. Phloem sap could be collected from F. sylvatica and M. kobus and the concentration of sucrose in phloem sap was about five- and 11-fold higher, respectively, than in the cytosol of mesophyll cells. Sugar exudation of cut leaves was decreased by p-chloromercuribenzenesulfonic acid, an inhibitor of sucrose-proton transporter. The results suggest that phloem loading of sucrose in the analyzed tree species involves active steps, and apoplastic phloem loading seems more likely. [ABSTRACT FROM AUTHOR]

Sucrose is produced in mesophyll cells and transferred into phloem cells before it is delivered long-distance to sink tissues. However, little is known about how sucrose transport is regulated in plants. Here, we identified a T-DNA insertional mutant of Oryza sativa DNA BINDING WITH ONE FINGER 11 ( OsDOF11 ), which is expressed in the vascular cells of photosynthetic organs and in various sink tissues. The osdof11 mutant plants are semi-dwarf and have fewer tillers and smaller panicles as compared with wild-type (WT) plants. Although sucrose enhanced root elongation in young WT seedlings, this enhancement did not occur in osdof11 seedlings due to reduced sucrose uptake. Sugar transport rate analyses revealed that less sugar was transported in osdof11 plants than in the WT. Expression of four Sucrose Transporter (SUT) genes— OsSUT1 , OsSUT3 , OsSUT4 , and OsSUT5— as well as two Sugars Will Eventually be Exported Transporters (SWEET) genes, OsSWEET11 and OsSWEET14 , was altered in various organs of the mutant, including the leaves. Chromatin immunoprecipitation assays showed that OsDOF11 directly binds the promoter regions of SUT1 , OsSWEET11 , and OsSWEET14 , indicating that the expression of these transporters responsible for sucrose transport via apoplastic loading is coordinately controlled by OsDOF11. We also observed that osdof11 mutant plants were less susceptible to infection by Xanthomonas oryzae pathovar oryzae , suggesting that OsDOF11 participates in sugar distribution during pathogenic invasion. Collectively, these results suggest that OsDOF11 modulates sugar transport by regulating the expression of both SUT and SWEET genes in rice. [ABSTRACT FROM AUTHOR]

Summary: Crop yield depends on efficient allocation of sucrose from leaves to seeds. In Arabidopsis, phloem loading is mediated by a combination of SWEET sucrose effluxers and subsequent uptake by SUT1/SUC2 sucrose/H+ symporters. ZmSUT1 is essential for carbon allocation in maize, but the relative contribution to apoplasmic phloem loading and retrieval of sucrose leaking from the translocation path is not known. Here we analysed the contribution of SWEETs to phloem loading in maize. We identified three leaf‐expressed SWEET sucrose transporters as key components of apoplasmic phloem loading in Zea mays L. ZmSWEET13 paralogues (a, b, c) are among the most highly expressed genes in the leaf vasculature. Genome‐edited triple knock‐out mutants were severely stunted. Photosynthesis of mutants was impaired and leaves accumulated high levels of soluble sugars and starch. RNA‐seq revealed profound transcriptional deregulation of genes associated with photosynthesis and carbohydrate metabolism. Genome‐wide association study (GWAS) analyses may indicate that variability in ZmSWEET13s correlates with agronomical traits, especifically flowering time and leaf angle. This work provides support for cooperation of three ZmSWEET13s with ZmSUT1 in phloem loading in Z. mays. [ABSTRACT FROM AUTHOR]

In compatible interactions, biotrophic microbial phytopathogens rely on the supply of assimilates by the colonized host tissue. It has been found in rice that phloem localized SWEET sucrose transporters can be reprogrammed by bacterial effectors to establish compatibility. We observed that sweet11/sweet12 double mutants, but not single mutants, exhibited increased resistance toward the fungal hemibiotroph Colletotrichum higginsianum (Ch), both in the biotrophic and the necrotrophic colonization phase. We therefore investigated if the phloem localized transporters AtSWEET11 and AtSWEET12 represent additive susceptibility factors in the interaction of Arabidopsis with Ch. AtSWEET12-YFP fusion protein driven by the endogenous promoter strongly accumulated at Ch infection sites and in the vasculature upon challenge with Ch. However, susceptibility of sweet12 single mutants to Ch was comparable to wild type, indicating that the accumulation of AtSWEET12 at Ch infection sites does not play a major role for compatibility. AtSWEET12-YFP reporter protein was not detectable at the plant–pathogen interface, suggesting that AtSWEET12 is not targeted by Ch effectors. AtSWEET11-YFP accumulation in pAtSWEET11:AtSWEET11-YFP plants were similar in Ch infected and mock control leaves. A close inspection of major carbohydrate metabolism in non-infected control plants revealed that soluble sugar and starch content were substantially elevated in sweet11/sweet12 double mutants during the entire diurnal cycle, that diurnal soluble sugar turnover was increased more than twofold in sweet11/sweet12, and that accumulation of free hexoses and sucrose was strongly expedited in double mutant leaves compared to wild type and both single mutants during the course of Ch infection. After 2 days of treatment, free and conjugated SA levels were significantly increased in infected and mock control leaves of sweet11/sweet12 relative to all other genotypes, respectively. Induced genes in mock treated sweet11/sweet12 leaves were highly significantly enriched for several GO terms associated with SA signaling and response compared to mock treated wild-type leaves, indicating sugar-mediated priming of the SA pathway in the double mutant. Infection assays with salicylic acid deficient sweet11/sweet12/sid2 triple mutants demonstrated that reduced susceptibility observed in sweet11/sweet12 was entirely dependent on the SA pathway. We suggest a model how defects in phloem loading of sucrose can influence SA priming and hence, compatibility.