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The formation and development of storage roots is the most important physiological process in sweetpotato production. Sucrose transporters (SUTs) regulate sucrose transport from source to sink organs and play important roles in growth and development of plants. However, whether SUTs involved in sweetpotato storage roots formation is so far unknown. In this study, we show that IbSUT1, a SUT, is localized to the plasma membrane. Overexpression of IbSUT1 in sweetpotato promotes the sucrose efflux rate from leaves, leading to increased sucrose levels in roots, thus induces lignin deposition in the stele, which inhibits the storage roots formation and compromises the yield. Heterologous expression of IbSUT1 in Arabidopsis significantly increases the sucrose accumulation and promotes lignification in the inflorescence stems. RNA-seq and biochemical analysis further demonstrated that IbMYB1 negatively regulates the expression of IbSUT1. Overexpression of IbMYB1 in Arabidopsis reduces the sucrose accumulation and lignification degree in the inflorescence stems. Moreover, co-overexpression of IbMYB1 and IbSUT1 restores the phenotype of lignin over-deposition in Arabidopsis. Collectively, our results reveal that IbSUT1 regulates source-sink sucrose transport and participates in the formation of sweetpotato storage roots and highlight the potential application of IbSUT1 in improving sweetpotato yield in the future., (© 2024 Society for Experimental Biology and John Wiley & Sons Ltd.)

The transport, compartmentation and allocation of sugar are critical for plant growth and development, as well as for stress resistance, but sugar transporter genes have not been comprehensively characterized in soybean. Here, we performed a genome-wide identification and expression analyses of sugar transporter genes in soybean in order to reveal their putative functions. A total of 122 genes encoding sucrose transporters (SUTs) and monosaccharide transporters (MSTs) were identified in soybean. They were classified into 8 subfamilies according to their phylogenetic relationships and their conserved motifs. Comparative genomics analysis indicated that whole genome duplication/segmental duplication and tandem duplication contributed to the expansion of sugar transporter genes in soybean. Expression analysis by retrieving transcriptome datasets suggested that most of these sugar transporter genes were expressed in various tissues, and a number of genes exhibited tissue-specific expression patterns. Several genes including GmSTP21, GmSFP8, and GmPLT5/6/7/8/9 were predominantly expressed in nodules, and GmPLT8 was significantly induced by rhizobia inoculation in root hairs. Transcript profiling and qRT-PCR analyses suggested that half of these sugar transporter genes were significantly induced or repressed under stresses like salt, drought, and cold. In addition, GmSTP22 was found to be localized in the plasma membrane, and its overexpression promoted plant growth and salt tolerance in transgenic Arabidopsis under the supplement with glucose or sucrose. This study provides insights into the evolutionary expansion, expression pattern and functional divergence of sugar transporter gene family, and will enable further understanding of their biological functions in the regulation of growth, yield formation and stress resistance of soybean., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Houqing Zeng reports financial support was provided by Hangzhou Normal University. If there are other authors, they declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Masson SAS. All rights reserved.)

Photosynthetic products in most fleshy fruits are unloaded via the apoplasmic pathway. Sugar transporters play an important role in the apoplasmic unloading pathway and are involved in sugar transport for fruit development. The MdSWEET23 , cloned from ''Hanfu'' apple ( Malus × domestica Borkh.) fruits, belongs to Clade III of the SWEET family. Subcellular localization revealed that MdSWEET23 is localized on the plasma membrane. β-glucuronidase activity assays showed that MdSWEET23 was primarily expressed in the sepal and carpel vascular bundle of apple fruits. Heterologous expression assays in yeast showed that MdSWEET23 functions in sucrose transport. The overexpression of MdSWEET23 in the ''Orin" calli increased the soluble sugar content. The silencing of MdSWEET23 significantly reduced the contents of sucrose and sorbitol in apple fruits. Ectopic overexpression of MdSWEET23 in tomato altered sugar metabolism and distribution in leaves and fruits, causing a reduction in photosynthetic rates and plant height, enhanced cold stress tolerance, and increased the content of sucrose, fructose, and glucose in breaking color fruits, but did not increase sugar sink potency of tomato fruits., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The reviewer X-FW declared a past co-authorship with the author XX to the handling editor., (Copyright © 2023 Nie, Wang, Li, Lyu, Qin and Xue.)

Sweet potato is an important food crop that can also be used as an industrial raw material. Sucrose is the main form of long-distance carbohydrate transport in plants, and sucrose transporter (SUT) regulates the transmembrane transport and distribution of sucrose during plant growth and metabolism. Moreover, SUT plays a key role in phloem mediated source-to-sink sucrose transport and physiological activities, supplying sucrose for the sink tissues. In this study, the full-length cDNA sequences of IbSUT 62788 and IbSUT 81616 were obtained by rapid amplification of cDNA ends (RACE) cloning according to the transcripts of the two SUT coding genes which were differentially expressed in sweet potato storage roots with different starch properties. Phylogenetic analysis was performed to clarify the classification of IbSUT 62788 and IbSUT 81616 . The subcellular localization of IbSUT 62788 and IbSUT 81616 was determined by transient expression in Nicotiana benthamiana . The function of IbSUT 62788 and IbSUT 81616 in sucrose and hexose absorption and transport was identified using yeast functional complementarity system. The expression pattern of IbSUT 62788 and IbSUT 81616 in sweet potato organs were analyzed by real-time fluorescence quantitative PCR (RT-qPCR). Arabidopsis plants heterologous expressing IbSUT 62788 and IbSUT 81616 genes were obtained using floral dip method. The differences in starch and sugar contents between transgenic and wild-type Arabidopsis were compared. The results showed IbSUT 62788 and IbSUT 81616 encoded SUT proteins with a length of 505 and 521 amino acids, respectively, and both proteins belonged to the SUT1 subfamily. IbSUT 62788 and IbSUT 81616 were located in the cell membrane and were able to transport sucrose, glucose and fructose in the yeast system. In addition, IbSUT 62788 was also able to transport mannose. The expression of IbSUT 62788 was higher in leaves, lateral branches and main stems, and the expression of IbSUT 81616 was higher in lateral branches, stems and storage roots. After IbSUT 62788 and IbSUT 81616 were heterologously expressed in Arabidopsis , the plants grew normally, but the biomass increased. The heterologous expression of IbSUT 62788 increased the soluble sugar content, leaf size and 1 000-seed weight of Arabidopsis plants. Heterologous expression of IbSUT 81616 increased starch accumulation in leaves and root tips and 1 000-seed weight of seeds, but decreased soluble sugar content. The results obtained in this study showed that IbSUT 62788 and IbSUT 81616 might be important genes regulating sucrose and sugar content traits in sweet potato. They might carry out physiological functions on cell membrane, such as transmembrane transport of sucrose, sucrose into and out of sink tissue, as well as transport and unloading of sucrose into phloem. The changes in traits result from their heterologous expression in Arabidopsis indicates their potential in improving the yield of other plants or crops. The results obtained in this study provide important information for revealing the functions of IbSUT 62788 and IbSUT 81616 in starch and glucose metabolism and formation mechanism of important quality traits in sweet potato.

In the developing seeds of all higher plants, filial cells are symplastically isolated from the maternal tissue supplying photosynthate to the reproductive structure. Photoassimilates must be transported apoplastically, crossing several membrane barriers, a process facilitated by sugar transporters. Sugars Will Eventually be Exported Transporters (SWEETs) have been proposed to play a crucial role in apoplastic sugar transport during phloem unloading and the post-phloem pathway in sink tissues. Evidence for this is presented here for developing seeds of the C4 model grass Setaria viridis. Using immunolocalization, SvSWEET4 was detected in various maternal and filial tissues within the seed along the sugar transport pathway, in the vascular parenchyma of the pedicel, and in the xylem parenchyma of the stem. Expression of SvSWEET4a in Xenopus laevis oocytes indicated that it functions as a high-capacity glucose and sucrose transporter. Carbohydrate and transcriptional profiling of Setaria seed heads showed that there were some developmental shifts in hexose and sucrose content and consistent expression of SvSWEET4 homologues. Collectively, these results provide evidence for the involvement of SWEETs in the apoplastic transport pathway of sink tissues and allow a pathway for post-phloem sugar transport into the seed to be proposed., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Phloem loading and sugar distribution are key steps for carbon partitioning in herbaceous and woody species. Although the phloem loading mechanisms in herbs are well studied, less is known for trees. It was shown for saplings of Fagus sylvatica L. and Quercus robur L. that the sucrose concentration in the phloem sap was higher than in the mesophyll cells, which suggests that phloem loading of sucrose involves active steps. However, the question remains whether this also applies for tall trees. To approach this question, tissue-specific sugar and starch contents of small and tall trees of F. sylvatica and Q. robur as well as the sugar concentration in the subcellular compartments of mesophyll cells were examined. Moreover, sucrose uptake transporters (SUTs) were analyzed by heterology expression in yeast and the tissue-specific expressions of SUTs were investigated. Sugar content in leaves of the canopy (11 and 26 m height) was up to 25% higher compared with that of leaves of small trees of F. sylvatica and Q. robur (2 m height). The sucrose concentration in the cytosol of mesophyll cells from tall trees was between 120 and 240 mM and about 4- to 8-fold lower than the sucrose concentration in the phloem sap of saplings. The analyzed SUT sequences of both tree species cluster into three types, similar to SUTs from other plant species. Heterologous expression in yeast confirmed that all analyzed SUTs are functional sucrose transporters. Moreover, all SUTs were expressed in leaves, bark and wood of the canopy and the expression levels in small and tall trees were similar. The results show that the phloem loading in leaves of tall trees of F. sylvatica and Q. robur probably involves active steps, because there is an uphill concentration gradient for sucrose. SUTs may be involved in phloem loading., (© The Author(s) 2022. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Starch is an important reserve of sugar, and starch-sugar conversion in plants plays an important role in the response of plants to various abiotic stresses. Nicosulfuron is a post-emergence herbicide commonly applied to maize fields. However, it is unclear how sucrose and starch in sweet corn are converted to accommodate nicosulfuron stress. Field and pot experiments were conducted to study the effects of nicosulfuron on the sugar metabolism enzymes, starch metabolism enzymes, non-enzyme substances, and expression of key enzyme genes in leaves and roots of sweet maize seedlings. Accordingly, this research compared the responses of the sister lines HK301 and HK320, which are nicosulfuron tolerant and sensitive, respectively. Under nicosulfuron stress, compared with HK301 seedlings, the accumulation of stem and root dry matter of HK320 seedlings was significantly reduced, resulting in a lower root-to-shoot ratio. Compared with HK320 seedlings, nicosulfuron stress significantly increased the sucrose, soluble sugar, and starch contents in HK301 leaves and roots. This may be related to the enhanced carbohydrate metabolism under nicosulfuron stress, including significant changes in sugar metabolism enzyme activity and the levels of SPS and SuSys expression. Further, under nicosulfuron stress, sucrose transporter genes (SUC 1, SUC 2, SWEET 13a, and SWEET 13b) in the leaves and roots of HK301 seedlings were significantly upregulated. Our results emphasize that changes in sugar distribution, metabolism, and transport can improve the adaptability of sweet maize to nicosulfuron stress., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The pollen grains of Phalaenopsis orchids are clumped tightly together, packed in pollen dispersal units called pollinia. In this study, the morphology, cytology, biochemistry, and sucrose transporters in pollinia of Phalaenopsis orchids were investigated. Histochemical detection was used to characterize the distribution of sugars and callose at the different development stages of pollinia. Ultra-performance liquid chromatography-high resolution-tandem mass spectrometry data indicated that P. aphrodite accumulated abundant saccharides such as sucrose, galactinol, myo-inositol, and glucose, and trace amounts of raffinose and trehalose in mature pollinia. We found that galactinol synthase (PAXXG304680) and trehalose-6-phosphate phosphatase (PAXXG016120) genes were preferentially expressed in mature pollinia. The P. aphrodite genome was identified as having 11 sucrose transporters (SUTs). Our qRT-PCR confirmed that two SUTs (PAXXG030250 and PAXXG195390) were preferentially expressed in the pollinia. Pollinia germinated in pollen germination media (PGM) supplemented with 10% sucrose showed increased callose production and enhanced pollinia germination, but there was no callose or germination in PGM without sucrose. We show that P. aphrodite accumulates high levels of sugars in mature pollinia, providing nutrients and enhanced SUT gene expression for pollinia germination and tube growth., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Background: Sucrose transporters (SUTs) mediate sucrose phloem loading in source tissue and sucrose unloading into sink tissue in potatoes and higher plants, thus playing a crucial role in plant growth and development. In potatoes, the physiological function of the sucrose transporters StSUT1 and StSUT4 has been clarified, whereas the physiological role of StSUT2 is not yet fully understood., Methods and Results: This study analyzed the relative expression of StSUT2 compared to that of StSUT1 and StSUT4 in different tissues from potatoes and its impact on different physiological characteristics by using StSUT2-RNA interference lines. Here, we report a negative effect of StSUT2-RNA interference on plant height, fresh weight, internodes number, leaf area, flowering time, and tuber yield. However, our data indicate that StSUT2 is not involved in carbohydrate accumulation in potato leaves and tubers. In addition, the data of the RNA-seq between the StSUT2-RNA interference line and WT showed that 152 genes were differentially expressed, of which 128 genes were upregulated and 24 genes were downregulated, and the GO and KEGG analyses revealed that differentially expressed genes were mainly related to cell wall composition metabolism., Conclusions: Thus, StSUT2 functions in potato plant growth, flowering time, and tuber yield without affecting carbohydrate accumulation in the leaves and tubers but may be involved in cell wall composition metabolism., Competing Interests: The authors declare that they have no conflicts of interest.

Interdependent metabolic and transport processes of carbon (C) and nitrogen (N) regulate plant growth and development, while the regulatory pathways remain poorly defined. We previously reported that rice circadian clock N-mediated heading date-1 (Nhd1) regulates growth duration-dependent N use efficiency. Here, we report that knockout of Nhd1 in rice reduced the rate of photosynthesis and the sucrose ratio of sheaths to blades, but increased the total C to N ratio and free amino acids. Leaf RNA-seq analysis indicated that mutation of Nhd1 dramatically altered expression of the genes linked to starch and sucrose metabolism, circadian rhythm, and amino acid metabolic pathways. We identified that Nhd1 can directly activate the transcriptional expression of sucrose transporter-1 (OsSUT1). Knockout of Nhd1 suppressed OsSUT1 expression, and both nhd1 and ossut1 mutants showed similar shorter height, and lower shoot biomass and sucrose concentration in comparison with the wild type, while overexpression of OsSUT1 can restore the defective sucrose transport and partially ameliorate the reduced growth of nhd1 mutants. The Nhd1-binding site of the OsSUT1 promoter is conserved in all known rice genomes. The positively related variation of Nhd1 and OsSUT1 expression among randomly selected indica and japonica varieties suggests a common regulatory module of Nhd1-OsSUT1-mediated C and N balance in rice., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

At the end of the summer season, grapevine buds (Vitis vinifera L) grown in temperate climates enter a state of winter recess or endodormancy (ED), which is induced by the shortening of the photoperiod, and during this period, the buds accumulate sucrose. In this study, we investigated whether the shortening of the photoperiod regulates the accumulation of sucrose in the buds in the same way as it regulates its entry into the ED. Because sucrose accumulation is regulated by genes that control its transport and degradation, the effect of the SD photoperiod and the transition of buds from paradormancy (PD) to ED on the expression of sucrose transporter (VvSUTs) and invertase genes (VvINVs) was studied. To analyze the possible role of sucrose during ED development, its effect on bud swelling and sprouting was studied on dormant and nondormant buds under forced growth conditions. The results showed that the SD photoperiod upregulates the expression of the VvSUT genes and downregulates that of the VvINV genes in grapevine buds. Additionally, during the transition of buds from PD to ED, the sucrose content increased, the expression of the VvINV genes decreased, and the expression of the VvSUT genes did not change significantly. Sucrose delayed bud swelling and sprouting when applied to dormant buds but had no effect when applied to nondormant buds. Therefore, we concluded that ED development and sucrose accumulation were synchronized events induced by the SD photoperiod and that a sucrose peak marks the end of ED development in grapevine buds., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Elsevier Masson SAS. All rights reserved.)

Plants secrete sugars from their roots into the soil, presumably to support beneficial plant-microbe interactions. Accordingly, manipulation of sugar secretion might be a viable strategy to enhance plant health and productivity. To evaluate the effect of increased root sugar secretion on plant performance and the soil microbiome, we overexpressed glucose and sucrose-specific membrane transporters in root epidermal cells of the model plant Arabidopsis thaliana. These plants showed strongly increased rates of sugar secretion in a hydroponic culture system. When grown on soil, the transporter-overexpressor plants displayed a higher photosynthesis rate, but reduced shoot growth compared to the wild-type control. Amplicon sequencing and qPCR analysis of rhizosphere soil samples indicated a limited effect on the total abundance of bacteria and fungi, but a strong effect on community structure in soil samples associated with the overexpressors. Notable changes included the increased abundance of bacteria belonging to the genus Rhodanobacter and the fungi belonging to the genus Cutaneotrichosporon, while Candida species abundance was reduced. The potential influences of the altered soil microbiome on plant health and productivity are discussed. The results indicate that the engineering of sugar secretion can be a viable pathway to enhancing the carbon sequestration rate and optimizing the soil microbiome., (© 2022 Wiley-VCH GmbH.)

Phloem unloading plays an important role in photoassimilate partitioning and grain yield improvements in cereal crops. The phloem unloading strategy and its effects on photoassimilate translocation and yield formation remain unclear in rice. In this study, plasmodesmata were observed at the interface between the sieve elements (SEs) and companion cells (CCs), and between the SE-CC complex and surrounding parenchyma cells (PCs) in phloem of the dorsal vascular bundle in developing caryopses. Carboxyfluorescein (CF) signal was detected in the phloem of caryopses, which showed that CF was unloaded into caryopses. These results indicated that the SE-CC complex was symplasmically connected with adjacent PCs by plasmodesmata. Gene expression for sucrose transporter (SUT) and cell wall invertase (CWI), and OsSUT1 and OsCIN1 proteins were detected in developing caryopses, indicating that rice plants might actively unload sucrose into caryopses by the apoplasmic pathway. Among three rice recombinant inbred lines, R201 exhibited lower plasmodesmal densities at the boundaries between cell types (SE-CC, SE-PC and CC-PC) in developing caryopses than R91 and R156. R201 also had lower expression of SUT and CWI genes and lower protein levels of OsSUT1 and OsCIN1, as well as CWI activity, than R91 and R156. These data agreed with stem non-structural carbohydrate (NSC) translocation and grain yields for the three lines. The nitrogen application rate had no significant effect on plasmodesmal densities at the interfaces between different cells types, and did not affect CF unloading in the phloem of developing caryopses. Low nitrogen treatment enhanced expression levels of OsSUT and OsCIN genes in the three lines. These results suggested that nitrogen application had no substantial effect on symplasmic unloading but affected apoplasmic unloading. Therefore, we concluded that poor symplasmic and apoplasmic unloading in developing caryopses might result in low stem NSC translocation and poor grain yield formation of R201., (© The Author(s) 2022. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Sucrose transporter (SUT) is a type of transmembrane protein that exists widely in plants and plays a significant role in the transportation of sucrose and the specific signal sensing process of sucrose. Therefore, identifying sucrose transporter is significant to the study of seed development and plant flowering and growth. In this study, a random forest-based model named ISTRF was proposed to identify sucrose transporter. First, a database containing 382 SUT proteins and 911 non-SUT proteins was constructed based on the UniProt and PFAM databases. Second, k-separated-bigrams-PSSM was exploited to represent protein sequence. Third, to overcome the influence of imbalance of samples on identification performance, the Borderline-SMOTE algorithm was used to overcome the shortcoming of imbalance training data. Finally, the random forest algorithm was used to train the identification model. It was proved by 10-fold cross-validation results that k-separated-bigrams-PSSM was the most distinguishable feature for identifying sucrose transporters. The Borderline-SMOTE algorithm can improve the performance of the identification model. Furthermore, random forest was superior to other classifiers on almost all indicators. Compared with other identification models, ISTRF has the best general performance and makes great improvements in identifying sucrose transporter proteins., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Chen, Li and Chen.)

The ever-increasing human population alongside environmental deterioration has presented a pressing demand for increased food production per unit area. As a consequence, considerable research effort is currently being expended in assessing approaches to enhance crop yields. One such approach is to harness the allelic variation lost in domestication. This is of particular importance since crop wild relatives often exhibit better tolerance to abiotic stresses. Here, we wanted to address the question as to why wild rice species have decreased grain production despite being characterized by enhanced rates of photosynthesis. In order to do so, we selected ten rice species on the basis of the presence of genome information, life span, the prominence of distribution, and habitat type and evaluated the expression of genes in photosynthesis, photorespiration, sucrose and starch synthesis, sucrose transport, and primary and secondary cell walls. We additionally measured the levels of a range of primary metabolites via gas chromatography-mass spectrometry. The results revealed that the wild rice species exhibited not only higher photosynthesis but also superior CO 2 recovery by photorespiration; showed greater production of photosynthates such as soluble sugars and starch and quick transportation to the sink organs with a possibility of transporting forms such as RFOs, revealing the preferential consumption of soluble sugars to develop both primary and secondary cell walls; and, finally, displayed high glutamine/glutamic acid ratios, indicating that they likely exhibited high N-use efficiency. The findings from the current study thus identify directions for future rice improvement through breeding.

The grain filling of rice depends on photoassimilates from leaves and stems. Phloem loading is the first crucial step for the transportation of sucrose to grains. However, phloem loading mechanisms in rice leaves and stems and their response to nitrogen (N) remain unclear. Here, using a combination of electron microscopy, transportation of a phloem tracer and 13C labeling, phloem loading was studied in rice leaves and stems. The results showed that the sieve element-companion cell complex lacked a symplastic connection with surrounding parenchyma cells in leaves and stems. The genes expression and protein levels of sucrose transporter (SUTs) and sugars will eventually be exported transporters (SWEETs) were detected in the vascular bundle of leaves and stems. A decrease in the 13C isotope remobilization from leaves to stems and panicles following treatment with p-chloromercuribenzenesulfonic acid indicated that rice leaves and stems actively transport sucrose into the phloem. Under low-N (LN) treatment, the activities of α-amylase, β-amylase and sucrose phosphate synthase (SPS) in stems and activity of SPS in leaves increased; genes expression and protein levels of SUTs and SWEETs in leaves and stems increased; the 13C isotope reallocation in panicles increased. These indicated that LN enhanced apoplastic phloem loading in stems and leaves. This improved the translocation of photoassimilates and consequently increased grain filling percentage, grain weight and harvest index. This study provides evidence that rice leaves and stems utilize an apoplastic loading strategy and respond to N stimuli by regulating the genes expression and protein levels of SUTs and SWEETs., (© The Author(s) 2022. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Introduction: Photosynthesis provides the energy basis for the life activities of plants by producing organic compounds, mainly sugar. As the main energy form of photosynthesis, sugar affects the growth and development of plants. During long-distance transportation, sucrose is the main form of transportation. The rate of sugar transport and the allocation of carbohydrates affect the biomass of crops and are closely related to the reproductive growth of crops., Main Text: The transportation of sugar is divided into active transportation and passive transportation. So how does the sucrose transporters (SUT) genes, which are the main carriers of sucrose in active transportation, affect the performance of rice agronomic traits is still to be explored. In this article, we describe the structure of inflorescence and review the transport forms and metabolic processes of sucrose in rice, such as how CO 2 is fixed, carbohydrate assimilation, and transport of organic matter. Sucrose transporters exhibited remarkable effects on the development of reproductive organs in rice., Conclusions: Here, the effects of different factors, such as the effects of anthers morphology on starch enrichment of pollen, effects of biotic and abiotic factors on sucrose transporters, effects of changes in trace elements on sucrose transporters, were discussed. Moreover, the regulation of transcription or translation level provides ideas for future research on sucrose transporters., (© 2022. The Author(s), under exclusive licence to Springer Nature B.V.)

Sucrose, an essential carbohydrate, is transported from source to sink organs in the phloem and is involved in a variety of physiological and metabolic processes in plants. Sucrose transporter proteins (SUTs) may play significant parts in the phloem loading and unloading of sucrose. In our study, the SUT gene family was identified in four Solanaceae species ( Capsicum annuum, Solanum lycopersicum, S. melongena , and S. tuberosum ) and other 14 plant species ranged from lower and high plants. The comprehensive analysis was performed by integration of chromosomal distribution, gene structure, conserved motifs, evolutionary relationship and expression profiles during pepper growth under stresses. Chromosome mapping revealed that SUT genes in Solanaceae were distributed on chromosomes 4, 10 and 11. Gene structure analysis showed that the subgroup 1 members have the same number of introns and exons. All the SUTs had 12 transmembrane structural domains exception from CaSUT2 and SmSUT2 , indicating that a structure variation might occurred among the Solanaceae SUT proteins. We also found a total of 20 conserved motifs, with over half of them shared by all SUT proteins, and the SUT proteins from the same subgroup shared common motifs. Phylogenetic analysis divided a total of 72 SUT genes in the plant species tested into three groups, and subgroup 1 might have diverged from a single common ancestor prior to the mono-dicot split. Finally, expression levels of CaSUTs were induced significantly under heat, cold, and salt treatments, indicating diverse functions of the CaSUTs to adapt to adverse environments.

SWEETs play important roles in intercellular sugar transport. Induction of SWEET sugar transporters by Transcription Activator-Like effectors (TALe) of Xanthomonas ssp. is key for virulence in rice, cassava and cotton. We identified OsSWEET11b with roles in male fertility and potential bacterial blight (BB) susceptibility in rice. While single ossweet11a or 11b mutants were fertile, double mutants were sterile. As clade III SWEETs can transport gibberellin (GA), a key hormone for spikelet fertility, sterility and BB susceptibility might be explained by GA transport deficiencies. However, in contrast with the Arabidopsis homologues, OsSWEET11b did not mediate detectable GA transport. Fertility and susceptibility therefore are likely to depend on sucrose transport activity. Ectopic induction of OsSWEET11b by designer TALe enabled TALe-free Xanthomonas oryzae pv. oryzae (Xoo) to cause disease, identifying OsSWEET11b as a potential BB susceptibility gene and demonstrating that the induction of host sucrose uniporter activity is key to virulence of Xoo. Notably, only three of six clade III SWEETs are targeted by known Xoo strains from Asia and Africa. The identification of OsSWEET11b is relevant for fertility and for protecting rice against emerging Xoo strains that target OsSWEET11b., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

The sugar distribution mechanism in fruits has been the focus of research worldwide; however, it remains unclear. In order to elucidate the relevant mechanisms in grape berries, the expression, localization, function, and regulation of three sucrose transporters were studied in three representative Vitis varieties. Both SUC11 and SUC12 expression levels were positively correlated with sugar accumulation in grape berries, whereas SUC27 showed a negative relationship. The alignment analysis and sucrose transport ability of isolated SUCs were determined to reflect coding region variations among V. vinifera , V. amurensis Ruper, and V. riparia , indicating that functional variation existed in one SUT from different varieties. Furthermore, potentially oligomerized abilities of VvSUCs colocalized in the sieve elements of the phloem as plasma membrane proteins were verified. The effects of oligomerization on transport properties were characterized in yeast. VvSUC11 and VvSUC12 are high-affinity/low-capacity types of SUTs that stimulate each other by upregulating V max and K m , inhibiting sucrose transport, and downregulating the K m of VvSUC27. Thus, changes in the distribution of different SUTs in the same cell govern functional regulation. The activation and inhibition of sucrose transport could be achieved in different stages and tissues of grape development to achieve an effective distribution of sugar.

Plant photosynthesis processes play vital roles in crop plant development. Understanding carbohydrate partitioning via sugar transport is one of the potential ways to modify crop biomass, which is tightly linked to plant architecture, such as plant height and panicle size. Based on the literature, we highlight recent findings to summarize phloem loading by sucrose transport in rice. In rice, sucrose transporters, OsSUTs (sucrose transporters) and OsSWEETs (sugars are eventually exported transporters) import sucrose and export cells between phloem parenchyma cells and companion cells. Before sucrose transporters perform their functions, several transcription factors can induce sucrose transporter gene transcription levels, such as Oryza sativa DNA binding with one finger 11 ( OsDOF11 ) and Oryza sativa Nuclear Factor Y B1 ( OsNF-YB1 ). In addition to native regulator genes, environmental factors, such as CO 2 concentration, drought stress and increased temperature, also affect sucrose transporter gene transcription levels. However, more research work is needed on formation regulation webs. Elucidation of the phloem loading mechanism could improve our understanding of rice development under multiple conditions and facilitate its manipulation to increase crop productivity.

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., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Natural rubber is an important industrial raw material and is commercially produced by rubber trees (Hevea brasiliensis). The sucrose transporter HbSUT3 plays an essential role in rubber production. Its expression in latex (cytoplasm of rubber-producing laticifers) is induced by bark treatment with Ethrel, an ethylene releaser, and the inducing effect correlates well with Ethrel-stimulated rubber yield increase. However, the mechanisms of ethylene induction on HbSUT3 expression are not known. Here, five Ethylene Response Factor (ERF) genes were identified from the cDNA library of Hevea latex by yeast one-hybrid screening with the promoter of HbSUT3 gene as bait. As revealed in a tobacco (Nicotiana tabacum) protoplast transient expression system, these HbERFs were mainly localized in the nucleus and four of them exhibited apparent transactivation activity. Of the five HbERF genes, HbERF-IXc4 was the most frequently screened in yeast one-hybrid, accounting for 65% of the ERF clones obtained. Moreover, among the five HbERFs, HbERF-IXc4 showed the strongest transactivation capacity when expressed in tobacco protoplast, the highest transcript abundance in latex and a close expressional correlation with its target gene, HbSUT3, in response to the Ethrel treatment. Taken together, our results indicate that ERFs, especially HbERF-IXc4, are critically involved in the activation of HbSUT3 expression in latex after Ethrel treatment on Hevea bark, and thus the stimulated latex yield., (Published by Oxford University Press 2021.)

Citrus fruit has a unique structure with soft leathery peel and pulp containing vascular bundles and several segments with many juice sacs. The function and morphology of each fruit tissue are different. Therefore, analysis at the organ-wide or mixed-tissue level inevitably obscures many tissue-specific phenomena. High-throughput RNA sequencing was used to profile Citrus sinensis fruit development based on four fruit tissue types and six development stages from young fruits to ripe fruits. Using a coexpression network analysis, modules of coexpressed genes and hub genes of tissue-specific networks were identified. Of particular, importance is the discovery of the regulatory network of phytohormones during citrus fruit development and ripening. A model was proposed to illustrate how ABF2 mediates the ABA signalling involved in sucrose transport, chlorophyll degradation, auxin homoeostasis, carotenoid and ABA biosynthesis, and cell wall metabolism during citrus fruit development. Moreover, we depicted the detailed spatiotemporal expression patterns of the genes involved in sucrose and citric acid metabolism in citrus fruit and identified several key genes that may play crucial roles in sucrose and citric acid accumulation in the juice sac, such as SWEET15 and CsPH8. The high spatial and temporal resolution of our data provides important insights into the molecular networks underlying citrus fruit development and ripening., (© 2021 The Authors. Plant Biotechnology Journal published by Society for Experimental Biology and The Association of Applied Biologists and John Wiley & Sons Ltd.)

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 OsSUTs 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., (© The Author(s) 2021. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

In potato plants, the phloem-mobile miR172 is involved in the sugar-dependent transmission of flower and tuber inducing signal transduction pathways and a clear link between solute transport and the induction of flowering and tuberization was demonstrated. The sucrose transporter StSUT4 seems to play an important role in the photoperiod-dependent triggering of both developmental processes, flowering and tuberization, and the phenotype of StSUT4 -inhibited potato plants is reminiscent to miR172 overexpressing plants. The first aim of this study was the determination of the level of miR172 in sink and source leaves of StSUT4 -silenced as well as StSUT4 -overexpressing plants in comparison to Solanum tuberosum ssp. Andigena wild type plants. The second aim was to investigate the effect of sugars on the level of miRNA172 in whole cut leaves, as well as in whole in vitro plantlets that were supplemented with exogenous sugars. Experiments clearly show a sucrose-dependent induction of the level of mature miR172 in short time as well as long time experiments. A sucrose-dependent accumulation of miR172 was also measured in mature leaves of StSUT4 -silenced plants where sucrose export is delayed and sucrose accumulates at the end of the light period.

Mucoromycotina is one of the earliest fungi to establish a mutualistic relationship with plants in the ancient land. However, the detailed information on their carbon supply from the host plants is largely unknown. In this research, a free-living Mucoromycotina called Gongronella sp. w5 (w5) was employed to explore its effect on Medicago truncatula growth and carbon source utilization from its host plant during the interaction process. W5 promoted M. truncatula growth and caused the sucrose accumulation in M. truncatula root tissue at 16 days post-inoculation (dpi). The transportation of photosynthetic product sucrose to the rhizosphere by M. truncatula root cells seemed accelerated by upregulating the SWEET gene. A predicted cytoplasmic invertase ( GspInv ) gene and a sucrose transporter ( GspSUT1 ) homology gene in the w5 genome upregulated significantly at the transcriptional level during w5- M. truncatula interaction at 16 dpi, indicating the possibility of utilizing plant sucrose directly by w5 as the carbon source. Further investigation showed that the purified GspInv displayed an optimal pH of 5.0 and a specific activity of 3380 ± 26 U/mg toward sucrose. The heterologous expression of GspInv and GspSUT1 in Saccharomyces cerevisiae confirmed the function of GspInv as invertase and GspSUT1 as sugar transporter with high affinity to sucrose in vivo . Phylogenetic tree analysis showed that the ability of Mucoromycotina to utilize sucrose from its host plant underwent a process of "loss and gain." These results demonstrated the capacity of Mucoromycotina to interact with extant land higher plants and may employ a novel strategy of directly up-taking and assimilating sucrose from the host plant during the interaction., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Wang, Fang, Liu, Liu, Fang, Fang and Xiao.)

Sucrose controls various developmental and metabolic processes in plants. In this review, we evaluate whether sucrose could be a preferred signaling molecule that controls processes like carbohydrate metabolism, accumulation of storage proteins, sucrose transport, anthocyanin accumulation, and floral induction. We summarize putative sucrose-dependent signaling pathways. Sucrose, but not other sugars, stimulates the genes that encode ADP-glucose pyrophosphorylase (AGPase), granule-bound starch synthase I, and UDP-glucose pyrophosphorylase in several species. The class-1 patatin promoter is induced under high sucrose conditions in potato (Solanum tuberosum). Exogenous sucrose reduces the loading of sucrose to the phloem by inhibiting the expression of the sucrose transporter and its protein activity in sugar beet (Beta vulgaris). Sucrose also influences a wide range of growth processes, including cell division, ribosome synthesis, cotyledon development, far-red light signaling, and tuber development. Floral induction is promoted by sucrose in several species. The molecular mechanisms by which sucrose functions as a signal are largely unknown. Sucrose enhances the expression of transcription factors such as AtWRKY20 and MYB75, which function upstream of the sucrose-responsive genes. Sucrose controls the expression of AtbZIP11 at the post-transcriptional level by the peptide encoded by uORF2. Sucrose levels affect translation of a group of mRNAs in Arabidopsis. Sucrose increases the activity of AGPase by posttranslational redox-modification. Sucrose interrupts the interaction between sucrose transporter SUT4 and cytochrome b5. In addition, the SNF-related protein kinase-1 appears to be involved in sucrose-dependent pathways by controlling sucrose synthase (SUS4) expression., (Copyright © 2020 Elsevier B.V. All rights reserved.)

In 'Crimson Seedless' grapes, the appearance of senescence caused by abnormal dark red color, the loss of crisp taste caused by the decrease in firmness, and the fading of sweetness caused by the decrease in total soluble sugar (TSS) are the main problems affecting its edible qualities after storage. In the mesocarp, burdock fructooligosaccharide (BFO) restricted sucrose export; therefore, more carbohydrates were retained directly leading to higher TSS and sweetness, and cell osmotic pressure and firmness were retained indirectly. In the exocarp, BFO restricted sucrose import; therefore, the signal molecule sucrose was reduced and the senescence-related processes were inhibited. The downregulation of SUC12 and SUC27 by BFO may play an important role in restricting sucrose transportation. The opposing effects exhibited by exogenous sucrose treatments compared to those of BFO further verified these mechanisms. Based on the above mechanisms, sucrose transportation mediates the fresh-keeping effects of BFO in 'Crimson Seedless' grapes., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Early fruit development is critical for determining crop yield. Cell wall invertase (CWIN) and sugar transporters both play important roles in carbon allocation and plant development. However, there is little information about the relationship between CWIN and those functionally related sugar transporters during fruit development. By using transgenic tomato with an elevated CWIN activity, we investigated how an increase in CWIN activity may regulate the expression of sugar transporter genes during fruit development. Our analyses indicate that CWIN activity may be under tight regulation by multiple regulators, including two invertase inhibitors (INVINHs) and one defective CWIN (deCWIN) in tomato ovaries prior to anthesis. Among the sugar transporters, expression of SlSWEET12c for sucrose efflux and SlHT2 for hexose uptake was enhanced by the elevated CWIN activity at 10 and 15 days after anthesis of tomato fruit development, respectively. The findings show that some specific sugars will eventually be exported transporters (SWEETs) and hexose transporters (HTs) respond to elevate CWIN activity probably to promote rapid fruit expansion when sucrose efflux from phloem and hexose uptake by parenchyma cell are in high demand. The analyses provide new leads for improving crop yield by manipulating CWIN-responsive sugar transporters, together with CWIN itself, to enhance fruit development and sugar accumulation., (Copyright © 2020 Ru, He, Zhu, Patrick and Ruan.)

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 K m 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., Competing Interests: The authors declare no conflict of interest., (© 2020 The Authors. Plant Direct published by American Society of Plant Biologists and the Society for Experimental Biology and John Wiley & Sons Ltd.)

The expression of AtSUC1 is controlled by the promoter and intragenic sequences. AtSUC1 is expressed in roots, pollen and trichomes. However, AtSUC1 promoter-GUS transgenics only show expression in trichomes and pollen. Here, we show that the root expression of AtSUC1 is controlled by an interaction between the AtSUC1 promoter and two short introns. The deletion of either intron from whole-gene-GUS constructs results in no root expression, showing that both introns are required. The two introns in tandem, fused to GUS, produce high constitutive expression throughout the vegetative parts of the plant. When combined with the promoter, the expression driven by the introns is reduced and localized to the roots. In Arabidopsis seedlings, exogenously applied sucrose induces the expression of AtSUC1 in roots and causes anthocyanin accumulation. atsuc1 loss-of-function mutants are defective in sucrose-induced anthocyanin accumulation. We show that an AtSUC1 whole-gene-GUS construct expressing a nonfunctional AtSUC1 (D152N) mutant, that is transport inactive, is defective in sucrose-induced AtSUC1 expression when expressed in an atsuc1-null background. We also show that the transport-defective allele does not complement the loss of sucrose-induced anthocyanin accumulation in null atsuc1 mutants. The results indicate that sucrose uptake via AtSUC1 is required for sucrose-induced AtSUC1 expression and sucrose-induced anthocyanin accumulation and that the site for sucrose detection is intracellular., (© The Author(s) 2020. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Short and concentrated natural fluorescence hinders tree peony (Paeonia suffruticosa) annual production, and defoliation and gibberellin (GA) application is used to induce its reflowering in autumn. Here, the individual roles of defoliation and GA treatment were determined by monitoring morphological and soluble sugar changes in buds and leaves, and by investigating carbon allocation- and metabolism-related gene expression. Both defoliation and GA treatment induced early bud development, but induction was faster using the GA treatment. Only defoliation, not GA treatment, induced the final reflowering, although their combination accelerated it. Furthermore, defoliation decreased the sucrose content in buds much faster than the GA treatment. This sucrose reduction may play a key role in tree peony reflowering, and the higher carbon metabolism activity in young leaves after defoliation may further help the reflowering process. Defoliation enhanced the expression of sucrose transporters PsSUT4 and PsSWEET12 in buds, and their expression in young leaves was greater than after GA treatment. This indicated that PsSUT4 and PsSWEET12 may help transport carbon into buds after defoliation. In addition, the invertases, PsCIN2 and PsCWIN1 in young leaves were more highly expressed after defoliation, indicating that they may contribute to reflowering after defoliation by accelerating sucrose hydrolysis in young leaves. In addition, the expression levels of PsVIN1 and PsVIN2 in leaves, and PsVIN2 in buds were more highly induced by GA treatment than by defoliation, indicating that PsVINs may mainly respond to GA treatment. These results may help improve the tree peony forcing culture technology and related industrial production., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

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., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Post-translational regulation of sucrose transporters represents one possibility to adapt transporter activity in a very short time frame. This can occur either via phosphorylation/dephosphorylation, oligomerization, protein-protein interactions, endocytosis/exocytosis, or degradation. It is also known that StSUT1 can change its compartmentalization at the plasma membrane and concentrate in membrane microdomains in response to changing redox conditions. A systematic screen for protein-protein-interactions of plant sucrose transporters revealed that the interactome of all three known sucrose transporters from the Solanaceous species Solanum tuberosum and Solanum represents a specific subset of interaction partners, suggesting different functions for the three different sucrose transporters. Here, we focus on factors that affect the subcellular distribution of the transporters. It was already known that sucrose transporters are able to form homo- as well as heterodimers. Here, we reveal the consequences of homo- and heterodimer formation and the fact that the responses of individual sucrose transporters will respond differently. Sucrose transporter SlSUT2 is mainly found in intracellular vesicles and several of its interaction partners are involved in vesicle traffic and subcellular targeting. The impact of interaction partners such as SNARE/VAMP proteins on the localization of SlSUT2 protein will be investigated, as well as the impact of inhibitors, excess of substrate, or divalent cations which are known to inhibit SUT1-mediated sucrose transport in yeast cells. Thereby we are able to identify factors regulating sucrose transporter activity via a change of their subcellular distribution.lycopersicum represents a specific subset of interaction partners, suggesting different functions for the three different sucrose transporters. Here, we focus on factors that affect the subcellular distribution of the transporters. It was already known that sucrose transporters are able to form homo- as well as heterodimers. Here, we reveal the consequences of homo- and heterodimer formation and the fact that the responses of individual sucrose transporters will respond differently. Sucrose transporter SlSUT2 is mainly found in intracellular vesicles and several of its interaction partners are involved in vesicle traffic and subcellular targeting. The impact of interaction partners such as SNARE/VAMP proteins on the localization of SlSUT2 protein will be investigated, as well as the impact of inhibitors, excess of substrate, or divalent cations which are known to inhibit SUT1-mediated sucrose transport in yeast cells. Thereby we are able to identify factors regulating sucrose transporter activity via a change of their subcellular distribution., Competing Interests: The authors declare no conflict of interest.

Sucrose is the main photosynthesis product of plants and the fundamental carbon skeleton monomer and energy supply for seed formation and development. Drought stress induces decreased photosynthetic carbon assimilation capacity, and seriously affects seed weight in soybean. However, little is known about the relationship between decreases in soybean seed yield and disruption of sucrose metabolism and transport balance in leaves and seeds during the reproductive stages of crop growth. Three soybean cultivars with similar growth periods, "Shennong17", "Shennong8", and "Shennong12", were subjected to drought stress during reproductive growth for 45 days. Drought stress significantly reduced leaf photosynthetic rate, shoot biomass, and seed weight by 63.93, 33.53, and 41.65%, respectively. Drought stress increased soluble sugar contents, the activities of sucrose phosphate synthase, sucrose synthase, and acid invertase enzymes, and up-regulated the expression levels of GmSPS1 , GmSuSy2 , and GmA-INV , but decreased starch content by 15.13% in leaves. Drought stress decreased the contents of starch, fructose, and glucose in seeds during the late seed filling stages, while it induced sucrose accumulation, which resulted in a decreased hexose-to-sucrose ratio. In developing seeds, the activities of sucrose synthesis and degradation enzymes, the expression levels of genes related to metabolism, and the expression levels of sucrose transporter genes were enhanced during early seed development under drought stress; however, under prolonged drought stress, all of them decreased. These results demonstrated that drought stress enhances the capacity for unloading sucrose into seeds and activated sucrose metabolism during early seed development. At the middle and late seed filling stages, sucrose flow from leaves to seeds was diminished, and the balance of sucrose metabolism was impaired in seeds, resulting in seed mass reduction. The different regulation strategies in sucrose allocation, metabolism, and transport during different seed development stages may be one of the physiological mechanisms for soybean plants to resist drought stress.

Sucrose is the main photosynthetic product in plants, and acts as a major energy substrate and signaling regulator of plant growth. Furthermore, sucrose is involved in the responses to various abiotic stresses. However, the role of sucrose in soybean (Glycine max L.) growth and development under drought stress remains largely unknown. In this study, the two soybean cultivars, Shennong8 (CV.SN8) and Shennong12 (CV.SN12), were grown in pot culture and subjected to three water treatments for 15 days: soil moisture contents of 75 ± 5% (CK), 45 ± 5% (MD), and 30 ± 5% (SD) of field capacity. Under drought stress, the reduction in shoot biomass was more pronounced than the reduction of biomass in the root of both soybean cultivars, resulting in higher root/shoot (R/S) ratio. Drought stress increased the contents of soluble sugar and sucrose in the leaves, but decreased starch content; in the roots, all of these parameters were increased. This may be related to the enhanced carbohydrate metabolism activity under drought stress, including notable changes in the activities of sugar metabolism enzymes and expression levels of GmSPS, GmSuSy, GmC-INV, GmA-INV, GmAMY3, and GmBAM1. Furthermore, the expression levels of sucrose transporter genes (GmSUC2, GmSWEET6, and GmSWEET15) in leaves and roots of soybean seedlings were up-regulated under drought stress. In conclusion, our results highlight that the increase in R/S ratio caused by the changes of sugar allocation, metabolism, and transport under drought stress contributes towards drought resistance of soybean., (Copyright © 2019 Elsevier Masson SAS. All rights reserved.)

Background: Sucrose (Suc), as the precursor molecule for rubber biosynthesis in Hevea brasiliensis, is transported via phloem-mediated long-distance transport from leaves to laticifers in trunk bark, where latex (cytoplasm of laticifers) is tapped for rubber. In our previous report, six Suc transporter (SUT) genes have been cloned in Hevea tree, among which HbSUT3 is verified to play an active role in Suc loading to the laticifers. In this study, another latex-abundant SUT isoform, HbSUT5, with expressions only inferior to HbSUT3 was characterized especially for its roles in latex production., Results: Both phylogenetic analysis and subcellular localization identify HbSUT5 as a tonoplast-localized SUT protein under the SUT4-clade (=type III). Suc uptake assay in baker's yeast reveals HbSUT5 to be a typical Suc-H + symporter, but its high affinity for Suc (Km = 2.03 mM at pH 5.5) and the similar efficiency in transporting both Suc and maltose making it a peculiar SUT under the SUT4-clade. At the transcript level, HbSUT5 is abundantly and preferentially expressed in Hevea barks. The transcripts of HbSUT5 are conspicuously decreased both in Hevea latex and bark by two yield-stimulating treatments of tapping and ethephon, the patterns of which are contrary to HbSUT3. Under the ethephon treatment, the Suc level in latex cytosol decreases significantly, but that in latex lutoids (polydispersed vacuoles) changes little, suggesting a role of the decreased HbSUT5 expression in Suc compartmentalization in the lutoids and thus enhancing the Suc sink strength in laticifers., Conclusions: Our findings provide insights into the roles of a vacuolar sucrose transporter, HbSUT5, in Suc exchange between lutoids and cytosol in rubber-producing laticifers.

A convenient and effective sucrose transport assay for Clostridium strains is needed. Traditional methods, such as 14 C-sucrose isotope labelling, use radioactive materials and are not convenient for many laboratories. Here, a sucrose transporter from potato was introduced into Clostridium , and a fluorescence assay based on esculin was used for the analysis of sucrose transport in Clostridium strains. This showed that the heterologously expressed potato sucrose transporter is functional in Clostridium . Recombinant engineering of high-level sucrose transport would aid sucrose fermentation in Clostridium strains. The assay described herein provides an important technological platform for studying sucrose transporter function following heterologous expression in Clostridium .

Phloem loading, as the first step of transporting photoassimilates from mesophyll cells to sieve element-companion cell complex, creates a driving force for long-distance nutrient transport. Three loading strategies have been proposed: passive symplastic loading, apoplastic loading and symplastic transfer followed by polymer-trapping of stachyose and raffinose. Although individual species are generally referred to as using a single phloem loading mechanism, it has been suggested that some plants may use more than one, i.e. 'mixed loading'. Here, by using a combination of electron microscopy, reverse genetics and 14 C labeling, loading strategies were studied in cucumber, a polymer-trapping loading species. The results indicate that intermediary cells (ICs), which mediate polymer-trapping, and ordinary companion cells, which mediate apoplastic loading, were mainly found in the fifth and third order veins, respectively. Accordingly, a cucumber galactinol synthase gene (CsGolS1) and a sucrose transporter gene (CsSUT2) were expressed mainly in the fifth/third and the third order veins, respectively. Immunolocalization analysis indicated that CsGolS1 was localized in companion cells (CCs) while CsSUT2 was in CCs and sieve elements (SEs). Suppressing CsGolS1 significantly decreased the stachyose level and increased sucrose content, while suppressing CsSUT2 decreased the sucrose level and increased the stachyose content in leaves. After 14 CO 2 labeling, [ 14 C]sucrose export increased and [ 14 C]stachyose export reduced from petioles in CsGolS1i plants, but [ 14 C]sucrose export decreased and [ 14 C]stachyose export increased into petioles in CsSUT2i plants. Similar results were also observed after pre-treating the CsGolS1i leaves with PCMBS (transporter inhibitor). These results demonstrate that cucumber phloem loading depends on both polymer-trapping and apoplastic loading strategies., (© 2019 The Authors The Plant Journal © 2019 John Wiley & Sons Ltd.)

The sucrose carrier AtSUC2 of Arabidopsis thaliana is localized in the phloem, where it catalyzes the uptake of sucrose from the apoplast into companion cells. Imported sucrose moves passively via plasmodesmata from the companion cells into the neighboring sieve elements that distribute this disaccharide to the different sink organs. Phloem loading of sucrose by the AtSUC2 protein is an essential process, and mutants lacking this protein stay tiny, develop no or only few flowers, and have a strongly reduced root system. The promoter of the AtSUC2 gene is active exclusively in companion cells of the phloem. Moreover, it drives very strong expression not only in Arabidopsis, but also in all plant species tested so far, including monocot species. Due to these features, the AtSUC2 promoter has become an important tool in diverse areas of plant research during the last two decades. It was used to study phloem development and function including phloem loading and unloading. Furthermore, it was helpful in analyzing the pathways of posttranscriptional silencing by RNA interference, the regulation of flowering, mechanisms of nutrient withdrawal by phloem-feeding pathogens, and other physiological functions that are related to long distance transport. The present paper gives an overview of different approaches in plant research that utilized the strong and companion cell-specific expression of own or foreign genes driven by the AtSUC2 promoter.

Immunolocalization of proteins in differentiated phloem cells is a challenging task given their special anatomy, organellar infrastructure, and the phloem tissue's heterogeneity. Incorporation of specific wall components in the thickened cell walls of phloem cells is often the source of unspecific labeling, leading to erroneous localization. Therefore, special care is required regarding generation and purification of specific antibodies. In addition, tissue preservation of phloem cells, which contain a high osmotic pressure in their functional state, is a very challenging task prone to various pitfalls. This chapter provides practical advice for cautious tissue preparation and antibody purification. Furthermore, methods that can be used to verify immunohistochemical localization data, such as promoter-reporter studies or activity tests, are discussed. Such confirmation experiments are essential for unambiguous determination of protein location in cells of the phloem.

Key Message: The homologous genes to OsSUT1-5 in wheat were identified and detailed analysed. TaSUT1 was the predominant sucrose transporter group and it illustrated the genotypic variations towards drought during grain filling. Sucrose transporters (SUT) play crucial roles in wheat stem water soluble carbohydrate (WSC) remobilization to grain. To determine the major functional SUT gene groups in shoot parts of wheat during grain development, drought tolerant varieties, Westonia and Kauz, were investigated in field drought experiments. Fourteen homologous genes to OsSUT1-5 were identified on five homeologous groups, namely TaSUT1_4A, TaSUT1_4B, TaSUT1_4D; TaSUT2_5A, TaSUT2_5B, TaSUT2_5D; TaSUT3_1A, TaSUT3_1D; TaSUT4_6A, TaSUT4_6B, TaSUT4_6D; TaSUT5_2A, TaSUT5_2B, and TaSUT5_2D, and their gene structures were analysed. Wheat plants above the ground were harvested from pre-anthesis to grain maturity and the stem, leaf sheath, rachis, lemma and developing grain were used for analysing TaSUT gene expression. Grain weight, thousand grain weight, kernel number per spike, biomass and stem WSC were characterized. The study showed that among the five TaSUT groups, TaSUT1 was the predominant sucrose transporting group in all organs sampled, and the expression was particularly high in the developing grain. In contrast to TaSUT1, the gene expression levels of TaSUT2, TaSUT3 and TaSUT4 were lower, except for TaSUT3 which showed preferential expression in the lemma before anthesis. The TaSUT5 gene group was very weakly expressed in all tissues. The upregulated gene expression of TaSUT1 Westonia type in stem and grain reveal a crucial role in stem WSC remobilization to grain under drought. The high TaSUT1 gene expression and the significant correlations with thousand grain weight (TGW) and kernel number per spike demonstrated the contribution in Kauz's high grain yield in an irrigated environment and high TGW in Westonia under drought stress. Further molecular level identification is required for gene marker development.

Sucrose transporters (SUTs) play key roles in allocating the translocation of assimilates from source to sink tissues. Although the characteristics and biological roles of SUTs have been intensively investigated in higher plants, this gene family has not been functionally characterized in cotton. In this study, we performed a comprehensive analysis of SUT genes in the tetraploid cotton Gossypium hirsutum . A total of 18 G. hirsutum SUT genes were identified and classified into three groups based on their evolutionary relationships. Up to eight SUT genes in G. hirsutum were placed in the dicot-specific SUT1 group, while four and six SUT genes were, respectively, clustered into SUT4 and SUT2 groups together with members from both dicot and monocot species. The G. hirsutum SUT genes within the same group displayed similar exon/intron characteristics, and homologous genes in G. hirsutum At and Dt subgenomes, G. arboreum , and G. raimondii exhibited one-to-one relationships. Additionally, the duplicated genes in the diploid and polyploid cotton species have evolved through purifying selection, suggesting the strong conservation of SUT loci in these species. Expression analysis in different tissues indicated that SUT genes might play significant roles in cotton fiber elongation. Moreover, analyses of cis -acting regulatory elements in promoter regions and expression profiling under different abiotic stress and exogenous phytohormone treatments implied that SUT genes, especially GhSUT6A/D , might participate in plant responses to diverse abiotic stresses and phytohormones. Our findings provide valuable information for future studies on the evolution and function of SUT genes in cotton.

Despite the relevance of seed-filling mechanisms for crop yield, we still have only a rudimentary understanding of the transport processes that supply the caryopsis with sugars. We hypothesized that SWEET sucrose transporters may play important roles in nutrient import pathways in the rice caryopsis. We used a combination of mRNA quantification, histochemical analyses, translational promoter-reporter fusions and analysis of knockout mutants created by genomic editing to evaluate the contribution of SWEET transporters to seed filling. In rice caryopses, SWEET11 and 15 had the highest mRNA levels and proteins localized to four key sites: all regions of the nucellus at early stages; the nucellar projection close to the dorsal vein; the nucellar epidermis that surrounds the endosperm; and the aleurone. ossweet11;15 double knockout lines accumulated starch in the pericarp, whereas caryopses did not contain a functional endosperm. Jointly, SWEET11 and 15 show all the hallmarks of being necessary for seed filling with sucrose efflux functions at the nucellar projection and a role in transfer across the nucellar epidermis/aleurone interface, delineating two major steps for apoplasmic seed filling, observations that are discussed in relation to observations made in rice and barley regarding the relative prevalence of these two potential import routes., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

A putative sucrose transporter PdSUT1 included in the same clade that Sut1p from Schizosaccharomyces pombe was identified in Penicillium digitatum, the major citrus postharvest pathogen. PdSUT1 gene was characterized using target gene disruption and gene overexpression. The ΔPdSUT1 mutants generated by gene elimination showed reduction in fungal virulence during citrus fruit infection assayed in mature fruit at 20°C. However, the overexpression mutants did not increased disease severity neither in the mutants coming from a high virulent nor from a low virulent P. digitatum progenitor strains. Moreover, fungicide sensitivity was affected in the deletant mutants but not in the overexpression transformants. The expression analysis of several genes involved in fungicide resistance showed an intensification of MFS transporters and a decrease of sterol demethylases transcriptional abundance in the ΔPdSUT1 mutants compare to the parental wild type strain. PdSUT1 appear not to be directly involved in fungicide resistance although can affect the gene expression of fungicide related genes. These results indicate that PdSUT1 contribute to P. digitatum fungal virulence and influence fungicide sensitivity through carbohydrate uptake and MFS transporters gene activation., (Copyright © 2017 Elsevier GmbH. All rights reserved.)

Sucrose transport and partitioning are crucial for seed filling. While many plasma-membrane-localised sucrose transporters (SUT1 family members) have been analysed in seeds, the functions of vacuolar SUT2 members are still obscure. In barley grains, expression of HvSUT1 and HvSUT2 overlap temporally and spatially, suggesting concerted functions to regulate sucrose homeostasis. Using HvSUT2-RNAi plants, we found that grains were also deficient in HvSUT1 expression and seemingly sucrose-limited during mid-to-late grain filling. Transgenic endosperms accumulated less starch and dry weight, although overall sucrose and hexose contents were higher. Comprehensive transcript and metabolite profiling revealed that genes related to glycolysis, the tricarboxylic acid cycle, starch and amino acid synthesis, grain maturation, and abscisic acid signalling were down-regulated together with most glycolytic intermediates and amino acids. Sucrose was increased along the sucrose delivery route in the nucellar projection, the endosperm transfer cells, and the starchy endosperm, indicating that suppressed transporter activity diminished sucrose efflux from vacuoles, which generated sugar deficiency in the cytoplasm. Thus, endosperm vacuoles may buffer sucrose concentrations to regulate homeostasis at grain filling. Transcriptional changes revealed that limited endosperm sucrose initiated sugar starvation responses, such as sugar recycling from starch, hemicelluloses and celluloses together with vacuolar protein degradation, thereby supporting formation of nucleotide sugars. Barley endosperm cells can thus suppress certain pathways to retrieve resources to maintain essential cell functions., (© The Author 2017. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

In higher plants, sugars (mainly sucrose) are produced by photosynthetically assimilated carbon in mesophyll cells of leaves and translocated to heterotrophic organs to ensure plant growth and development. Sucrose transporters, or sucrose carriers (SUCs), play an important role in the long-distance transportation of sucrose from source organs to sink organs, thereby affecting crop yield and quality. The identification, characterization, and molecular function analysis of sucrose transporter genes have been reported for monocot and dicot plants. However, no relevant study has been reported on sucrose transporter genes in Brassica rapa var. rapa , a cruciferous root crop used mainly as vegetables and fodder. We identified and cloned 12 sucrose transporter genes from turnips, named BrrSUC1.1 to BrrSUC6.2 according to the SUC gene sequences of B. rapa pekinensis . We constructed a phylogenetic tree and analyzed conserved motifs for all 12 sucrose transporter genes identified. Real-time quantitative polymerase chain reaction was conducted to understand the expression levels of SUC genes in different tissues and developmental phases of the turnip. These findings add to our understanding of the genetics and physiology of sugar transport during taproot formation in turnips.

An essential step for the distribution of carbon throughout the whole plant is the loading of sugars into the phloem in source organs. In many plants, accumulation of sugars in the sieve element-companion cell (SE-CC) complex is mediated and regulated by active processes. However, for poplar and many other tree species, a passive symplasmic mechanism of phloem loading has been proposed, characterized by symplasmic continuity along the pre-phloem pathway and the absence of active sugar accumulation in the SE-CC complex. A high overall leaf sugar concentration is thought to enable diffusion of sucrose into the phloem. In this review, we critically evaluate current evidence regarding the mechanism of passive symplasmic phloem loading, with a focus on the potential influence of active sugar transport and plasmodesmal regulation. The limited experimental data, combined with theoretical considerations, suggest that a concomitant operation of passive symplasmic and active phloem loading in the same minor vein is unlikely. However, active sugar transport could well play an important role in how passively loading plants might modulate the rate of sugar export from leaves. Insights into the operation of this mechanism has direct implications for our understanding of how these plants utilize assimilated carbon., (© 2017 Institute of Botany, Chinese Academy of Sciences.)

Background and Aims: Pisum sativum L. (pea) seed is a source of carbohydrate and protein for the developing plant. By studying pea seeds inoculated by the cytokinin-producing bacterium, Rhodococcus fascians , we sought to determine the impact of both an epiphytic (avirulent) strain and a pathogenic strain on source-sink activity within the cotyledons during and following germination., Methods: Bacterial spread was monitored microscopically, and real-time reverse transcription-quantitative PCR was used to determine the expression of cytokinin biosynthesis, degradation and response regulator gene family members, along with expression of family members of SWEET , SUT , CWINV and AAP genes - gene families identified initially in pea by transcriptomic analysis. The endogenous cytokinin content was also determined., Key Results: The cotyledons infected by the virulent strain remained intact and turned green, while multiple shoots were formed and root growth was reduced. The epiphytic strain had no such marked impact. Isopentenyl adenine was elevated in the cotyledons infected by the virulent strain. Strong expression of RfIPT , RfLOG and RfCKX was detected in the cotyledons infected by the virulent strain throughout the experiment, with elevated expression also observed for PsSWEET , PsSUT and PsINV gene family members. The epiphytic strain had some impact on the expression of these genes, especially at the later stages of reserve mobilization from the cotyledons., Conclusions: The pathogenic strain retained the cotyledons as a sink tissue for the pathogen rather than the cotyledon converting completely to a source tissue for the germinating plant. We suggest that the interaction of cytokinins, CWINVs and SWEETs may lead to the loss of apical dominance and the appearance of multiple shoots., (© The Author 2016. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com)