Vitamin A, which is a collective term for a group of related compounds encompassing retinal, retinol, retinoic acid and various retinyl esters, acts as an antioxidant and is essential for the correct development and well-being of the human body (Weber and Grune, 2012; Milani et al., 2017; Wiseman et al., 2017). Arch. 78, 110. Wax ester and lipophilic compound profiling of Euglena gracilis by gas chromatography-mass spectrometry: toward understanding of wax ester fermentation under hypoxia. Rep. 28, 457466. Washington, DC: United States Patent and Trademark Office. doi: 10.1007/s12257-010-0145-5, Zhu, G., Ou, Q., Zhang, T., Jiang, X., Sun, G., Zhang, N., et al. Until now, the research on the health benefits of paramylon has focussed mainly on applications to treat human conditions, but it has been suggested that it also could be beneficial to livestock and fish health. 5, 205214. Lipids 50, 483492. Biomass production in mixotrophic culture of Euglena gracilis under acidic condition and its growth energetics. Aquacult. Biotechnol. Military Med. Nature 438, 90. doi: 10.1038/nature04056, Croft, M.T., Warren, M.J., and Smith, A.G. (2006). doi: 10.3389/fenrg.2016.00036, Andreani, L., and Rocha, J.D. Creation of straight-chain cationic polysaccharide-based bile salt sequestrants made from euglenoid -1,3-glucan as potential antidiabetic agents. Fuel 80, 747756. Photosynthesis of Euglena gracilis under cobalamin-sufficient and -limited growing conditions. Paramylon is the water-insoluble storage polysaccharide of E. gracilis, which consists of -1,3-linked glucose subunits, and has an estimated molecular weight between 100 and 500 kDa (Figure 3A) (Miyatake and Kitaoka, 1983; Koizumi et al., 1993; Barsanti et al., 2011). doi: 10.1016/j.algal.2018.02.001, Barsanti, L., Passarelli, V., Evangelista, V., Frassanito, A.M., and Gualtieri, P. (2011). However, the current chemical synthesis of oligosaccharides is still laborious and product yields are very low despite some advances in the field, whereas enzymatic synthesis has been considered as a technically feasible option (Plante et al., 2001; Ogawa et al., 2014). Foods 6, 121. 58, 532538. Evaluation of growth characteristics of Euglena gracilis for microalgal biomass production using wastewater. J. Appl. Fatty acid content in wild type and WZSL mutant of Euglena gracilis. Two alternative metabolic routes to -carotene have been identified biochemically or via transcriptomics in E. gracilis: the methylerythritol phosphate and the mevalonate pathway (Disch et al., 1998; Kim et al., 2004; O'Neill et al., 2015b). 22, 590594. J. Appl. doi: 10.3390/antiox6040099, Fujita, T., Aoyagi, H., Ogbonna, J.C., and Tanaka, H. (2008). 68, 11441148. (2018). At present, only two enzymes of this pathway, the D-galacturonate reductase and aldonolactonase, have been biochemically characterised in vitro. Euglena gracilis cell mass has been shown to be a suitable source for the production of biogas by anaerobic digestion, producing around 650 or 800 mL biogas/g DW under PT or HT cultivation conditions, respectively (Grimm et al., 2015). Paramylon (-1,3-glucan) content in wild type and WZSL mutant of Euglena gracilis. In recent years, the versatile phototrophic protist Euglena gracilis has emerged as an interesting candidate for application-driven research and commercialisation, as it is an excellent source of dietary protein, pro(vitamins), lipids, and the -1,3-glucan paramylon only found in euglenoids. doi: 10.1111/j.1365-2672.2010.04848.x, Rosenberg, A. Kataoka, H., Shimura, T., Mizoshita, T., Kubota, E., Mori, Y., Mizushima, T., et al. While a draft genome assembly and initial features of the genome have been made available, a complete annotated genome sequence is not on hand as yet (O'Neill et al., 2015a; Ebenezer et al., 2017). Euglena gracilis is used as model organism for various microbiological, molecular biological and biotechnological studies. doi: 10.3945/jn.111.148080, Disch, A., Schwender, J., Mller, C., Lichtenthaler, H.K., and Rohmer, M. (1998). Euglena gracilis cell confined in a glass capillary and imaged using bright-field reflected light microscopy. Antioxidants 6, 118. J. Clin. doi: 10.1016/j.pisc.2015.07.002, O'Neill, E.C., Trick, M., Hill, L., Rejzek, M., Dusi, R.G., Hamilton, C.J., et al. doi: 10.1002/fsn3.383. doi: 10.1111/j.1550-7408.2009.00452.x. Carbohydrate Polymers 200, 239247. Bioresource Technol. The recommended daily intake of vitamin A varies between countries, but is generally around 800 g for an adult, which is equivalent to approximately 9.6 mg of the provitamin -carotene or around 3 g of dehydrated E. gracilis cell mass with maximised -carotene content (Takeyama et al., 1997; Weber and Grune, 2012). From first- to third-generation biofuels: challenges of producing a commodity from a biomass of increasing complexity. doi: 10.1039/C5MB00319A, Ooi, V.E., and Liu, F. (2000). J. Pharmacol. 13, 195205. Ammonia- and light-induced degradation of paramylum in Euglena gracilis. In the absence of complete genetic information, transcriptomic and proteomic studies have provided valuable insights into the complex metabolic pathways of E. gracilis and their regulation under different growth conditions (O'Neill et al., 2015b; Yoshida et al., 2016; Hasan et al., 2017). For example, silencing of the genes coding for the E. gracilis 3-ketoacyl-CoA thiolase (EgKAT) isozymes EgKAT 1, EgKAT 2 and EgKAT 3 via RNAi lead to an increased production of short-chain WEs, and an artificial biodiesel inferred from the lipid composition before and after gene silencing indicated an improvement of cold flow properties (Nakazawa et al., 2015). Int. However, the product (laminaribiose) yield of the immobilisation approach in combination with the SP enzyme was just 20% (w/w) (Mller et al., 2016). 25, 90100. doi: 10.1016/j.fsi.2018.04.003, Yamane, Y.-,I., Utsunomiya, T., Watanabe, M., and Sasaki, K. (2001). doi: 10.1002/(SICI)1097-0290(19970120)53:2<185::AID-BIT8>3.0.CO;2-K, Tanaka, Y., Ogawa, T., Maruta, T., Yoshida, Y., Arakawa, K., and Ishikawa, T. (2017). Furthermore, substantial research has been performed to improve the WE yields specifically. (1989). (1964). saccharophila. (2009). Euglena gracilis growth and cell composition under different temperature, light and trophic conditions. (2012). Barras, D.R., and Stone, B.A. Anim. (2012). However, local legal regulations could prevent (outdoor) cultivation of genetically modified E. gracilis strains (Beacham et al., 2017). Chem. J. 10, 165170. Molecular structure of paramylon (A). Characterization of paramylon morphological diversity in photosynthetic euglenoids (Euglenales, Euglenophyta). J. Nat. 25, 855865. Euglena gracilis as source of the antioxidant vitamin E. Effects of culture conditions in the wild strain and in the natural mutant WZSL. Biological, and Environmental Sciences, Faculty of Agriculture, Tottori . Outdoor PT cultivation is considered the most economical option for the bulk production of microalgal biomass (Apel and Weuster-Botz, 2015). 25, 299307. doi: 10.1093/oxfordjournals.pcp.a077454, Sun, A., Hasan, M.T., Hobba, G., Nevalainen, H., and Te'o, J. Appl. doi: 10.1002/1873-3468.12659, Tandon, P., Jin, Q., and Huang, L. (2017). (2018). Subcellular distribution of cobalamin-dependent methionine synthase in Euglena gracilis Z. Phytochemistry 35, 5961. Production of wax esters via microbial oil synthesis from food industry waste and by-product streams. As another target for a metabolic engineering approach, trans-2-enoyl-CoA reductase has been identified as a potentially rate-limiting enzyme in the FA synthesis pathway of E. gracilis (Inui et al., 2017). doi: 10.1364/BOE.7.002703, Levine, R., Horst, G., Tonda, R., Lumpkins, B., and Mathis, G. (2018). doi: 10.3177/jnsv.64.8, Nakazawa, M., Andoh, H., Koyama, K., Watanabe, Y., Nakai, T., Ueda, M., et al. Purification and properties of the -1,3-glucan exo-hydrolase. J. Biochem. Exp. Eur. Sci. Influence of fatty acid composition of raw materials on biodiesel properties. However, DW titres of E. gracilis under PT conditions are significantly lower than those obtained with MT/HT growth (Table 1) (Takeyama et al., 1997; Hasan et al., 2017). This work was supported by the Australian Research Council Industrial Transformation Training Centre funding scheme (Project Number: IC130100009). Euglena. 270, 8492. Metabolomics 11, 175183. Biochem. Implications for the recognition of fungi by innate immunity. The alga Euglena gracilis (E. gracilis) has recently gained attention as a health food, but its effects on human gut microbiota remain unknown. For example, intracellular lipids of a mutant strain generated by Fe-ion irradiation of the WT parental strain were stained with a boron-dipyrromethene dye (BODIPY505/515) to isolate high-producers using fluorescence-activated cell sorting. U.S. Patent No 9,045,784. As some microalgal species, including Chlorella sp. (Table 1) (de Oliveira et al., 1999; Tokuoglu and nal, 2003; Ramazanov and Ramazanov, 2006;Rodrguez-Zavala et al., 2010). These eukaryotic algae undergo cell division asexually in the dark . Isolation and characterization of a starchless mutant of Chlorella pyrenoidosa STL-PI with a high growth rate, and high protein and polyunsaturated fatty acid content. Acta (BBA) Enzymol. doi: 10.1371/journal.pone.0191462, Takeda, T., Nakano, Y., Takahashi, M., Konno, N., Sakamoto, Y., Arashida, R., et al. Alteration of wax ester content and composition in Euglena gracilis with gene silencing of 3-ketoacyl-coa thiolase isozymes. Fish Shellfish Immunol. 58, 15041505. -glucan in foods and its physiological functions. Microbial production of vitamin B12: a review and future perspectives. Metallomics 6, 604616. Both species were acid tolerant, growing optimally at pH 2.5-7. At pH values typical of AMD (pH 2.5-4) in the absence of elevated metals, E. gracilis outcompeted E. mutabilis (growth rates of 1.0 and 0.8 div d 1, respectively). doi: 10.1371/journal.pone.0195329, Watanabe, T., Shimada, R., Matsuyama, A., Yuasa, M., Sawamura, H., Yoshida, E., et al. 15, 657662. (2018). Impact of -1,3-glucan isolated from Euglena gracilis on corneal epithelial cell migration and on wound healing in a rat alkali burn model. We would like to thank the Macquarie University Microscopy Unit for its contribution to Figure 3. . doi: 10.1002/apj.1805, Yamada, K., Suzuki, H., Takeuchi, T., Kazama, Y., Mitra, S., Abe, T., et al. Comparison of the methods for determining degree of polymerization of water-insoluble -1,3-glucans. doi: 10.1016/j.algal.2017.04.028, Behera, S., Singh, R., Arora, R., Sharma, N.K., Shukla, M., and Kumar, S. (2014). Sci. J. Nutr. Biol. doi: 10.1016/S0016-2361(00)00142-3, Maxwell, J.R., Douglas, A.G., Eglinton, G., and McCormick, A. doi: 10.1021/bi034731y, Milani, A., Basirnejad, M., Shahbazi, S., and Bolhassani, A. There is an emerging market for E. gracilis-derived products, which is reflected in the recent founding of new companies specialised in the cultivation and commercialisation of E. gracilis. These factors need to be taken into account when determining the most favourable time of harvest. Cell Factories 16, 114. doi: 10.1111/bph.13625. J. Shibakami, M., Shibata, K., Akashi, A., Onaka, N., Takezaki, J., Tsubouchi, G., et al. Nevertheless, this study offered that lipid . For example, knock-down of the starch degradation kinases 1 and 2 using RNAi was shown to lead to a 1.2 and 1.4-fold increase in WE accumulation, respectively, under anaerobic growth conditions (Kimura and Ishikawa, 2018). Adaptive laboratory evolution - harnessing the power of biology for metabolic engineering. doi: 10.1074/jbc.RA117.000936, Kusmic, C., Barsacchi, R., Barsanti, L., Gualtieri, P., and Passarelli, V. (1998). However, at present, most studies only extract the paramylon of E. gracilis, which is a waste of resources.In this study, a method of sequential extraction of lipid and paramylon from E. gracilis was studied, and the extraction conditions were optimized. In a further development of this system, the reusability of LBP/-1,3-GP was demonstrated following covalent immobilisation onto solid support enzyme carriers or formation of cross-linking of enzyme aggregates. However, the metabolic conversion efficiency of -carotene to vitamin A, generally considered to be 12:1, is dependent on several factors (e.g., bioavailability) and always needs to be taken into account in a dietary context (Weber and Grune, 2012). doi: 10.1186/s12870-015-0698-8, Keshavarz, B., and Khalesi, M. (2016). doi: 10.1002/elsc.201100025, Santini, A., Cammarata, S.M., Capone, G., Ianaro, A., Tenore, G.C., Pani, L., et al. doi: 10.1016/j.micres.2017.11.013, Niki, E., and Traber, M.G. J. Phycol. To circumvent this issue, a fed-batch two-step cultivation method with a transition from MT to PT growth conditions has been developed and shown to increase the -carotene titre per L culture more than 6-fold (Table 1) (Takeyama et al., 1997). doi: 10.1016/0005-2744(69)90253-8, Barsanti, L., Bastianini, A., Passarelli, V., Tredici, M.R., and Gualtieri, P. (2000). Phytochemistry 116, 2127. Sci. Am. Wax ester fermentation and its application for biofuel production, in Euglena: Biochemistry, Cell and Molecular Biology, eds S. D. Schwartzbach and S. Shigeoka (Cham: Springer International Publishing), 269283. Fish Dis. There are a few isolated reports on genetic manipulation of E. gracilis. 4, 115. 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