C O2. These sugars have also been shown to rapidly affect the expression of many genes, even at concentrations as low as 1 mM (Kunz et al., 2014). Plant Cell Environ. Genet. Overexpression of cotton SUS in tobacco also led to an increased growth rate and taller plants (Coleman et al., 2006). Breed. SuSy is also the main enzyme metabolizing Suc in the phloem of Ricinus communis (Geigenberger et al., 1993). A. S. Basra (New York, NY: Food Products Press), 85–112. Plant Sci., 08 February 2019 Transgenic tomato plants with decreased sucrose synthase are unaltered in starch and sugar accumulation in the fruit. (2016)]. These features of SuSy may help to control the amount of Suc consumed in different organs, for example, in stems and petioles. In beet, SUS1 mRNA levels increased under anaerobic conditions, but there was no increase in SuSy1 protein (Klotz and Haagenson, 2008). 33, 1870–1874. Oxygen is passed into the atmosphere and the hydrogen is used to assimilate carbon dioxide in a dark (non photosynthetic) reaction that forms starch, sucrose, and another disaccharide called maltose. SuSy activity is feedback-inhibited by its product, Fru, and its activity is also reversible. Evidence for a role of SuSy in callose deposition was found in an Arabidopsis double mutant of phloem-specific SUS (sus5 sus6). (2017). G1P is then converted into ADP-G, in a reaction catalyzed by ADP-G pyrophosphorylase (AGPase). (1999). The occurrence of strong end-product inhibition appears to be correlated with high acid-invertase activity in fully expanded leaves. FEBS Lett. Molecules 23, 1–16. Plant Biol. To summarize, plant SuSy activity has been shown to play important roles in plant sugar metabolism, primarily in sink tissues. Plant Cell Physiol. The signal metabolite trehalose-6-phosphate inhibits the sucrolytic activity of sucrose synthase from developing castor beans. In addition, five maize starch-deficient endosperm mutants were screened for metabolic enzyme activity and all showed reduced SuSy activity (Doehlert and Kuo, 1990). 2), 503–509. Plant J. Sucrose synthase (SuSy) is a glycosyl transferase enzyme that plays a key role in sugar metabolism, primarily in sink tissues. The source-sink ratio of l-year-old, potted `Montmorency' sour cherry (Prunus cerasus) trees was manipulated by partial defoliation (D) or continuous lighting (CL) to investigate the phenomenon of end-product inhibition of photosynthesis. doi: 10.1146/annurev.pp.39.060188.002035, Huang, D. Y., and Wang, A. Y. Biophys. Mol. Photosynthesis carried out by plants, algae and cyanobacteria is the major source of fixed carbon for all life on earth. Sucrose synthase, a cytosolic enzyme in protoplasts of Jerusalem artichoke tubers (Helianthus tuberosus L.). 1.6. doi: 10.1016/0378-1119(90)90028-P, Martin, T., Frommer, W. B., Salanoubat, M., and Willmitzer, L. (1993). doi: 10.1007/s00425-011-1356-5, Goren, S., Lugassi, N., Stein, O., Yeselson, Y., Schaffer, A. Sugar metabolism in developing tubers of Solanum tuberosum. Lysine-containing proteins in maize endosperm: a major contribution from cytoskeleton-associated carbohydrate-metabolizing enzymes. 46, 1366–1376. Thus the carbohydrates appeared in the order of their complexity. ): structure, expression, and evolution. (2012). Plant Mol. doi: 10.1080/17429140902898429, Yu, W. P., Wang, A. Y., Juang, R. H., Sung, H. Y., and Su, J. C. (1992). Because Suc cleavage by SuSy yields UDP-G, which is a direct substrate for both cellulose (β1-4) and callose (β1-3) glucans, it is widely assumed that SuSy plays a role in the synthesis of both of these polysaccharides. The Formula. Biotechnol. A recent study found that the Suc-cleavage activity of a castor bean SuSy, RcSUS1, is inhibited by trehalose 6-phosphate, suggesting another mechanism by which Suc flux can be controlled in heterotrophic tissues (Fedosejevs et al., 2018). Plant Mol. doi: 10.1007/BF00047725, Wang, F., Smith, A. G., and Brenner, M. L. (1994). Expression of an Arabidopsis sucrose synthase gene indicates a role in metabolization of sucrose both during phloem loading and in sink organs. OS and DG jointly wrote the manuscript, and read and approved the final manuscript. Biochem. Tissue-specific expression of two genes for sucrose synthase in carrot (Daucus carota L.). That double mutant had less callose in its phloem plasmodesmata and in response to leaf wounding, as compared with WT or quadruple mutant (sus1, sus2, sus3, and sus4) plants (Barratt et al., 2009). FEBS Lett. doi: 10.1046/j.1365-313x.2001.01002.x, Regmi, K. C., Zhang, S., and Gaxiola, R. A. The hexoses can be phosphorylated to hexose phosphates (hex-P), directed to starch synthesis in the plastid or to glycolysis and then respiration in the mitochondria or directed to other metabolic pathways. 104, 535–540. doi: 10.1104/pp.95.3.669, Ricard, B., Toai, T. V., Chourey, P., and Saglio, P. (1998). SuSy proteins have also been detected in citrus (Citrus paradisi) and maize phloem companion cells using immunohistological analysis (Nolte and Koch, 1993). However, SuSy and INV also co-localize to the cytosol. There are following products of photosynthesis: I. Hexoses: According Weevers the first Carbohydrates were the. is the raw material for photosynthesis and glucose and oxygen are the products. 4, 87–101. Oxygen deficiency (hypoxia) and a complete absence of oxygen (anoxia) are forms of serious abiotic stress that often cause reduced plant growth and productivity. Abdullah, M., Cao, Y., Cheng, X., Meng, D., Chen, Y., Shakoor, A., et al. Sucrose synthase may play a number of different roles in the phloem involving the regulation of sink strength and phloem unloading (see section “ Role of SuSy in Sink Strength”), supplying hexoses for companion cell respiration and supplying precursors for complex carbohydrates, such as callose and cellulose (see section “Roles of SuSy in Cellulose and Callose Metabolism”). (2014). Suc can also pass directly from the phloem to sink cells through plasmodesmata (Figure 1). doi: 10.1016/S0031-9422(00)81212-1, Munoz, F. J., Baroja-Fernandez, E., Moran-Zorzano, M. T., Viale, A. M., Etxeberria, E., Alonso-Casajus, N., et al. Only a few studies have used mutant and transgenic plants to elucidate the roles of SuSy in phloem. In the parasitic plant Phelipanche ramosa, PrSUS1 transcript was found in the xylem of developing roots (Peron et al., 2012). Phloem loading is thought to be highly important for defining sink strength and the breakdown of Suc in sink organs may also contribute to sink strength. There is much more evidence linking SuSy to starch synthesis in non-photosynthetic tissues or sink tissues. Sink- and vascular-associated sucrose synthase functions are encoded by different gene classes in potato. 4, 113–117. Plant Physiol. Purification of sucrose synthase from thermotolerant wheat grains and its characterization. Plant Biol. Yang, C. L., and Su, J. C. (1980). The rapid generation of mutation data matrices from protein sequences. Bot. (1999). Planta 217, 628–638. 174, 534–543. (2006). (2011). Inside the cell, Suc can be stored in the vacuole or hydrolyzed by vINV. A., Luan, S., Wi, S. G., Bae, H., Lee, D. S., and Bae, H. J. Biol. The transgenic plants overexpressing AtSUS1 showed increased chlorophyll levels, as well as increased photosynthesis, TSS (total soluble sugars), starch, Suc and Fru, as well as increased enzymatic activity of SPS and SPP in leaves, indicating increased sugar production in the transgenic plants. 34, 837–846. 65, 33–67. doi: 10.1105/tpc.010108, Salnikov, V. V., Grimson, M. J., Delmer, D. P., and Haigler, C. H. (2001). 17, 847–856. 44, 500–509. Sci. Companion-cell specific localization of sucrose synthase in zones of phloem loading and unloading. The development of cotton fibers starts with the initiation and elongation of the epidermal cells, followed by secondary growth and maturation marked by massive cellulose production. These unique phylogenetic trees raise fundamental questions about the evolution of SuSy in plants. This review focuses on all aspects of the SuSy found in plants, including their evolution, gene families, protein structure and tissue and subcellular localization, as well as their roles in plant development and sugar signaling. In red beet, it was estimated that 7% of SuSy protein may be tonoplast-bound and it was suggested that SuSy may play a role in the mobilization of Suc from the vacuole (Etxeberria and Gonzalez, 2003). Plant Microbe Interact. Phytochemistry 57, 823–833. Plant Mol. These plants exhibited abnormal leaf development and irregular auxin patterning, suggesting that altered sugar signaling in the SAM or primordia, rather than lower sugar metabolism, is likely to be the cause of these developmental changes. Subjective And Short Questions For Photosynthesis, DARK REACTION OR CALVIN – BENSON CYCLE OR C3 CYCLE, DEFINITIONS AND KEY POINTS FOR Photosynthesis, Answer of Question of Reproduction & Development, DEFINITIONS AND KEY POINTS FOR OBJECTIVES. Antisense inhibition of tomato fruit sucrose synthase decreases fruit setting and the sucrose unloading capacity of young fruit. A study of potato tubers of transgenic plants overexpressing INV or Suc pyrophosphorylase, which allows a way to bypass the degradation of Suc by SuSy, revealed a steeper reduction in oxygen levels inside the tubers, reduced starch synthesis and a lower ATP to ADP ratio, underscoring the importance of SuSy under low-oxygen conditions (Bologa et al., 2003). doi: 10.1104/pp.110.171371, Carlson, S. J., and Chourey, P. S. (1996). Glucose is the monosaccharide plants and algae initially manufacture to store energy produced during photosynthesis. The SlSUS4 transcript was shown to be present asymmetrically and localized to the primordia from very early stages of development using in situ hybridization with an SlSUS4 antisense probe (Pien et al., 2001). Work with isoform-specific antibodies has revealed that specific SuSy isoforms are more abundant in the phloem. However, at least five of the Chinese pear SUS genes cannot be functional, as the predicted proteins are too short to contain both the SuSy domain and the glycosyl-transferase domain. There is a lot of evidence that SUS are highly expressed in vascular tissues. (2014). Plant Physiol. Mutations at the rug4 locus alter the carbon and nitrogen metabolism of pea plants through an effect on sucrose synthase. doi: 10.1093/pcp/pcs180, Lunn, J. E., Feil, R., Hendriks, J. H., Gibon, Y., Morcuende, R., Osuna, D., et al. This situation encouraged us to create a more comprehensive phylogenetic tree. Pronounced phenotypic changes in transgenic tobacco plants overexpressing sucrose synthase may reveal a novel sugar signaling pathway. 114, 55–62. Enzymic properties of amyloplasts form suspension cultures of soybean. 132, 2058–2072. doi: 10.1071/FP06234, Ruan, Y. L. (2014). doi: 10.1104/pp.111.178574, Buckeridge, M. S., Vergara, C. E., and Carpita, N. C. (1999). Plant Physiol. Adv. doi: 10.1104/pp.108.2.735, Haigler, C. H., Ivanova-Datcheva, M., Hogan, P. S., Salnikov, V. V., Hwang, S., Martin, K., et al. Chem. Plant Growth Regul. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. Front. Similarly, overexpression of aspen (Popolus tremuloides) SUS in Arabidopsis resulted in an increased growth rate and increased plant biomass, and also induced early flowering (Xu and Joshi, 2010). Additional amino acid sequences were retrieved using the Plaza 3.0 tool for gene-family analysis (Van Bel et al., 2017) using “sucrose synthase” in a keyword search. *Correspondence: David Granot, granot@agri.gov.il, Front. Identification of actively filling sucrose sinks. DEFINITIONS AND KEY POINTS FOR OBJECTIVES OF PLANT DIVERSITY. Sucrose is the end product of photosynthesis and the primary sugar transported in the phloem of most plants. Sucrose synthase belongs to the glycosyltransferase-4 subfamily of glycosyltransferases, a large family that includes a wide variety of glycosyltransferases, including SPS, trehalose synthase, and trehalose phosphorylase. Normal growth of Arabidopsis requires cytosolic invertase but not sucrose synthase. (1986). In the cytosol, two triose-P molecules produce one fructose 1,6-bisphosphate (F1,6BP) molecule in a reaction catalyzed by aldolase. Simplified schematic presentation of sugar metabolism in sink tissue cells toward cellulose, callose and starch synthesis. For example, a reduction in SuSy activity reduced starch content in potato tubers, carrot taproots and maize endosperm (Chourey and Nelson, 1976; Zrenner et al., 1995; Tang and Sturm, 1999). (1994). Plant Physiol. 89, 1117–1121. BMC Plant Biol. SuSy proteins are typically considered homotetramers (Schmolzer et al., 2016), although some SuSy isoforms have been reported to act as heterotetramers in barley (Hordeum vulgare) (Guerin and Carbonero, 1997), maize (Zea mays) (Duncan et al., 2006), rice (Oryza sativa) (Huang and Wang, 1998) and bird cherry (Prunus padus) (Sytykiewicz et al., 2008). (1999). Transcript levels of some SUS genes have been found to increase in response to low levels of oxygen in potato (Biemelt et al., 1999), maize (McCarty et al., 1986; Bailey-Serres et al., 1988; Zeng et al., 1998), rice (Ricard et al., 1991; Hirose et al., 2008), carrot (Sturm et al., 1999), Arabidopsis (Martin et al., 1993; Dejardin et al., 1999; Baud et al., 2004), wheat (Marana et al., 1990), beet (Hesse and Willmitzer, 1996; Klotz and Haagenson, 2008), pigeon pea (Cajanus cajan) (Kumutha et al., 2008) and pondweed (Potamogeton distinctus ) (Harada et al., 2004, 2005). U.S.A. 106, 13124–13129. 39, 355–378. Sucrose is made by binding glucose with fructose. Harada, T., Satoh, S., Yoshioka, T., and Ishizawa, K. (2004). doi: 10.1007/BF00485135, Ciereszko, I., and Kleczkowski, L. A. 46, D1190–D1196. PLoS One 12:e0182334. doi: 10.1007/BF00224782, Gerber, L., Zhang, B., Roach, M., Rende, U., Gorzsas, A., Kumar, M., et al. Chem. Triose-P can be transported to the cytosol by a triose-P/phosphate translocator. Analysis of the sucrose synthase gene family in tobacco: structure, phylogeny, and expression patterns. Plant SuSy have also been found to play a role in mutualism with symbiotic organisms like Rhizobium bacteria. Fu, H., and Park, W. D. (1995). (2016), it was shown that both sucrose and light affect WUS expression in the SAM, although Suc is unable to release SAM dormancy in the dark. Sucrose is the end product of photosynthesis and the primary sugar transported in the phloem of most plants. doi: 10.1104/pp.121.2.599, Zhang, C. H., Yu, M. L., Ma, R. J., Shen, Z. J., Zhang, B. Two plasma-membrane SuSy proteins were also detected in maize endosperm by activity assays and monoclonal antibodies; those proteins were also found in the cytosol (Carlson and Chourey, 1996). D-fructose, sucrose and starch are commonly are formed in the green cells during photosynthesis. Spinach, a sucrose storer, showed the least inhibition in both girdled and excised leaf systems, which indicates that sucrose is probably not directly responsible for the end-product inhibition of photosynthesis. doi: 10.1007/s10725-016-0189-4, Zheng, Y., Anderson, S., Zhang, Y., and Garavito, R. M. (2011). Physiol. Acad. doi: 10.1007/s004250050652, Bieniawska, Z., Paul Barratt, D. H., Garlick, A. P., Thole, V., Kruger, N. J., Martin, C., et al. Structure and expression profile of the sucrose synthase gene family in the rubber tree: indicative of roles in stress response and sucrose utilization in the laticifers. Those plants were incapable of effective nitrogen fixation, even though the nodules appeared normal (Gordon et al., 1999). Biol. Suc can be converted into starch via different pathways, which also differ between chloroplasts and heterotrophic tissues (For a comprehensive review of starch synthesis, see Bahaji et al., 2014b). Biochem. 39, 459–466. doi: 10.1007/BF00018467, Zeng, Y., Wu, Y., Avigne, W. T., and Koch, K. E. (1998). The sucrose synthase gene family in Populus: structure, expression, and evolution. That SuSy exhibited optimum activity at 37°C and was stable at temperatures up to 50°C (Verma et al., 2018), unlike other SuSy proteins, whose stability decreases at temperatures above 30°C (Schmolzer et al., 2016). 11, 67–75. Sucrose and starch are products of two physically separated gluconeogenic pathways, sucrose in the cytosol and starch in the chloroplast. (1998). The enzymatic deficiency conditioned by the shrunken-1 mutations of maize. 35, 588–603. Sucrose is metabolised by sucrose synthase and glycolysis within the phloem complex of Ricinus communis L. seedlings. doi: 10.1093/pcp/pcj068, Etxeberria, E., and Gonzalez, P. (2003). Biochim. doi: 10.4236/abb.2010.15056, Xu, S. M., Brill, E., Llewellyn, D. J., Furbank, R. T., and Ruan, Y. L. (2012). (2011). Plant Physiol. Alternatively, Suc can be brought into the sink cell by a Suc transporter or enter through plasmodesmata. Sucrose synthase levels do not limit or regulate carbon transfer in the arbuscular mycorrhizal symbiosis. HPLC-MS/MS analyses show that the near-starchless aps1 and pgm leaves accumulate wild-type levels of ADPglucose: further evidence for the occurrence of important ADPglucose biosynthetic pathway(s) alternative to the pPGI-pPGM-AGP pathway. Evidence for a tonoplast-associated form of sucrose synthase and its potential involvement in sucrose mobilization from the vacuole. doi: 10.1002/1873-3468.13197, Fedosejevs, E. T., Ying, S., Park, J., Anderson, E. M., Mullen, R. T., She, Y. M., et al. Sucrose synthase activity does not restrict glycolysis in roots of transgenic potato plants under hypoxic conditions. Sink strength mainly relies on two parameters, sink organ size as a physical constraint and activity as a physiological constraint (Ho, 1988). Turquoise arcs indicate eudicot species and red arcs indicate monocot species. Evidence for multiple sites of glucosyl transfer in the synthase complex. The role of SuSy in the synthesis of cellulose and callose has been thoroughly investigated in cotton, with cotton fibers serving as a model for these processes. doi: 10.1016/S2095-3119(17)61755-6, Van Bel, M., Diels, T., Vancaester, E., Kreft, L., Botzki, A., Van De Peer, Y., et al. doi: 10.1073/pnas.92.20.9353, An, X., Chen, Z., Wang, J., Ye, M., Ji, L., Wang, J., et al. Similarly, overexpression of poplar xylem SUS in tobacco plants also resulted in increased cellulose content and xylem cell-wall thickness (Wei et al., 2015). The strong association of SuSy to the plasma membrane and a transmembrane prediction analysis led the researchers to hypothesize that part of the SuSy may be transmembrane (Carlson and Chourey, 1996). 45, S151–S151. doi: 10.1093/jxb/45.5.623, Sturm, A., Lienhard, S., Schatt, S., and Hardegger, M. (1999). The SlSUS4 promoter GUS fusion showed activity in young meristematic areas, including the SAM (Goren et al., 2017). (2001). SuSy activity was also suggested as an indicator for high rice grain yield in rice breeding programs (Counce and Gravois, 2006). 64, 31–37. Plant J. Plant J. “Cloning and sequencing of two differentially expressed sucrose synthase genes from potato,” in Proceedings of the Third International Congress of Plant Molecular Biology, Tucson, AZ. Herba Pol. Invertase/sucrose synthase balance, sugar signaling potential, and seedling survival. Role of the sucrose synthase encoding PrSus1 gene in the development of the parasitic plant Phelipanche ramosa L. (Pomel). It is a major end product of photosynthesis and functions as a primary transport sugar and in some cases as a direct or indirect regulator of gene expression. 90, 127–135. Triose-P can be transported to the cytosol by a triose-P/phosphate translocator. doi: 10.1093/pcp/pcr067, Bailey-Serres, J., Kloeckener-Gruissem, B., and Freeling, M. (1988). doi: 10.1023/A:1006327606696, Tong, X. L., Wang, Z. Y., Ma, B. Q., Zhang, C. X., Zhu, L. C., Ma, F. W., et al. (2014b). J. Chin. But still it is not clear which the first product of photosynthesis. In Arabidopsis, AtSUS5 and AtSUS6 are reported to be phloem specific (Barratt et al., 2009). 134, 1146–1152. In the phloem, SuSy may play a role in the maintenance of equilibrium between Suc and its breakdown products, supplying hexoses for energy production in companion cells and substrates for complex carbohydrates, like callose (Nolte and Koch, 1993). 117, 85–90. Haplotype analysis of sucrose synthase gene family in three Saccharum species. Phytochemistry 25, 1579–1585. Photosynthesis uses light energy and carbon dioxide to make triose phospates (G3P). Sucrose synthase of soybean nodules. doi: 10.1016/j.gene.2014.01.062, Zhang, J., Arro, J., Chen, Y., and Ming, R. (2013). Plant J. accumulationandturnover in sucrose storers andother plants. Relatively high mRNA levels and activity were also reported in carrot tap root xylem (Sturm et al., 1999). The second site is also a serine, at around position 170, and is thought to regulate protein degradation (Hardin et al., 2003). Plant Cell Physiol. Sucrose synthase is an integral component of the cellulose synthesis machinery. J. Environ. Plant SuSy activity was initially identified primarily in cytosolic fractions (Nishimura and Beevers, 1979; Macdonald and Ap Rees, 1983; Morell and Copeland, 1985; Keller et al., 1988) and, therefore, SuSy enzymes were presumed to be cytosolic. Biol. (1999). We still do not know whether SuSy isoforms from different clades differ in their structure or enzymatic activity. The direction of SuSy activity may also be regulated by pH; its optimal Suc-synthesis activity is observed between pH 7.5 and 9.5 and optimal Suc degradation occurs at pH values between 5.5 and 7.5 (Schmolzer et al., 2016). doi: 10.1023/A:1006136524725, Cho, L. H., Pasriga, R., Yoon, J., Jeon, J. S., and An, G. (2018). Sucrose synthase activity as a potential indicator of high rice grain yield. (2014). The spatial distribution of sucrose synthase isozymes in barley. doi: 10.1046/j.1365-313X.2003.01765.x, Gordon, A. J., Minchin, F. R., James, C. L., and Komina, O. Structure of the sucrose synthase gene on chromosome 9 of Zea mays L. EMBO J. doi: 10.1111/j.1399-3054.1997.tb01066.x, Schmolzer, K., Gutmann, A., Diricks, M., Desmet, T., and Nidetzky, B. On the other hand, Amor et al. 51, 294–301. 203, 1220–1230. Biochemical and molecular characterization of RcSUS1, a cytosolic sucrose synthase phosphorylated in vivo at serine 11 in developing castor oil seeds. None of those mutants exhibit any significant phenotype that suggesting redundancy between the different clades (Bieniawska et al., 2007). Sucrose synthesis is predominant in leaves, but the ability to synthesize sucrose is fairly widespread among plant cells. doi: 10.1104/pp.88.2.239, Kleines, M., Elster, R. C., Rodrigo, M. J., Blervacq, A. S., Salamini, F., and Bartels, D. (1999).

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