The novel chloroplast glucose transporter pGlcT2 affects adaptation to extended light periods.

Intracellular sugar compartmentation is critical in plant development and acclimation to challenging environmental conditions. Sugar transport proteins are present in plasma membranes and in membranes of organelles such as vacuoles, the Golgi apparatus, and plastids. However, there may exist other transport proteins with uncharacterized roles in sugar compartmentation. Here we report one such novel transporter of the Monosaccharide Transporter Family, the closest phylogenetic homolog of which is the chloroplast-localized glucose transporter pGlcT and that we therefore term plastidic glucose transporter 2 (pGlcT2). We show, using gene-complemented glucose uptake deficiency of an Escherichia coli ptsG/manXYZ mutant strain and biochemical characterization, that this protein specifically facilitates glucose transport, whereas other sugars do not serve as substrates. In addition, we demonstrate pGlcT2-GFP localized to the chloroplast envelope and that pGlcT2 is mainly produced in seedlings and in the rosette center of mature Arabidopsis plants. Therefore, in conjunction with molecular and metabolic data, we propose pGlcT2 acts as a glucose importer that can limit cytosolic glucose availability in developing pGlcT2-overexpressing seedlings. Finally, we show both overexpression and deletion of pGlcT2 resulted in impaired growth efficiency under long day and continuous light conditions, suggesting pGlcT2 contributes to a release of glucose derived from starch mobilization late in the light phase. Together, these data indicate the facilitator pGlcT2 changes the direction in which it transports glucose during plant development and suggest the activity of pGlcT2 must be controlled spatially and temporarily in order to prevent developmental defects during adaptation to periods of extended light.

Carbohydrate distribution via SWEET17 is critical for Arabidopsis inflorescence branching under drought.

Sugars Will Eventually be Exported Transporters (SWEETs) are the most recently discovered family of plant sugar transporters. By acting as uniporters, SWEETs facilitate the diffusion of sugars across cell membranes and play an important role in various physiological processes such as abiotic stress adaptation. AtSWEET17, a vacuolar fructose facilitator, was shown to be involved in the modulation of the root system during drought. In addition, previous studies have shown that overexpression of an apple homolog leads to increased drought tolerance in tomato plants. Therefore, SWEET17 might be a molecular element involved in the plant´s drought response. However, the role and function of SWEET17 in aboveground tissues of Arabidopsis under drought stress remains elusive. By combining gene expression analysis and stem architecture with the sugar profiles of different aboveground tissues, we uncovered a putative role of SWEET17 in carbohydrate supply and thus cauline branch elongation, especially during periods of carbon limitation, as occurs under drought stress. Thus, SWEET17 seems to be involved in maintaining efficient plant reproduction under drought-stress conditions.

Vacuolar fructose transporter SWEET17 is critical for root development and drought tolerance.

Root growth and architecture are markedly influenced by both developmental and environmental cues. Sugars integrate different stimuli and are essential building blocks and signaling molecules for modulating the root system. Members from the SUGAR WILL EVENTUALLY BE EXPORTED TRANSPORTER (SWEET) family facilitate the transport of different sugars over cellular membranes and steer both inter and intracellular distribution of sugars. SWEET17 represents a fructose-specific sugar porter localized to the vacuolar membrane, the tonoplast. Here, we analyzed how SWEET17-dependent fructose released from vacuoles affects root growth during drought stress in Arabidopsis (Arabidopsis thaliana). We found that the SWEET17 gene was predominantly expressed in the root vasculature and in meristematic cells of the root tip. SWEET17 expression appeared markedly induced during lateral root (LR) outgrowth and under drought. Moreover, fructose repressed primary root growth but induced density and length of first order LRs. Consistently, sweet17 knock-out mutants exhibited reduced LR growth and a diminished expression of LR-development-related transcription factors during drought stress, resulting in impaired drought tolerance of sweet17 mutants. We discuss how SWEET17 activity integrates drought-induced cellular responses into fructose signaling necessary for modulation of the root system and maximal drought tolerance.

Overexpression of the vacuolar sugar importer BvTST1 from sugar beet in Camelina improves seed properties and leads to altered root characteristics.

Overexpression of the vacuolar sugar transporter TST1 in Arabidopsis leads to higher seed lipid levels and higher total seed yield per plant. However, effects on fruit biomass have not been observed in crop plants like melon, strawberry, cotton, apple, or tomato with increased tonoplast sugar transporter (TST) activity. Thus, it was unclear whether overexpression of TST in selected crops might lead to increased fruit yield, as observed in Arabidopsis. Here, we report that constitutive overexpression of TST1 from sugar beet in the important crop species Camelina sativa (false flax) resembles the seed characteristics observed for Arabidopsis upon increased TST activity. These effects go along with a stimulation of sugar export from source leaves and not only provoke optimised seed properties like higher lipid levels and increased overall seed yield per plant, but also modify the root architecture of BvTST1 overexpressing Camelina lines. Such mutants grew longer primary roots and showed an increased number of lateral roots, especially when developed under conditions of limited water supply. These changes in root properties result in a stabilisation of total seed yield under drought conditions. In summary, we demonstrate that increased vacuolar TST activity may lead to optimised yield of an oil-seed crop species with high levels of healthy ω3 fatty acids in storage lipids. Moreover, since BvTST1 overexpressing Camelina mutants, in addition, exhibit optimised yield under limited water availability, we might devise a strategy to create crops with improved tolerance against drought, representing one of the most challenging environmental cues today and in future.