doi: 10.1016/j.algal.2017.08.029, Hayashi, H., Narumi, I., Wada, S., Kikuchi, M., Furuta, M., Uehara, K., et al. Algae acquire vitamin B12 through a symbiotic relationship with bacteria. nitrogen, and phosphorus) under the oxygen-free environment led to a favorable condition for lipid accumulation. Acta (BBA) Enzymol. The cell motion of Euglena gracilis in homogeneous and heterogeneous light environments was analyzed. Phycol. doi: 10.1007/s12257-010-0145-5, Zhu, G., Ou, Q., Zhang, T., Jiang, X., Sun, G., Zhang, N., et al. doi: 10.1111/bcp.13496. -tocopherol, the major form of vitamin E in the US diet, deserves more attention. Biochem. (2013). Biotechnol. Phycol. Int. (2016). doi: 10.1016/j.tplants.2016.07.012. 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: 10.1271/bbb.80062, Ishikawa, T., Tajima, N., Nishikawa, H., Gao, Y., Rapolu, M., Shibata, H., et al. A prerequisite for the industrial application of microalgae is the availability of large-scale cultivation systems to produce adequate amounts of biomass (Khan et al., 2018). (2001). Delta-9 Elongases and Their Use in Making Polyunsaturated Fatty Acids. Food Sci. (1964). Fatty acid synthesis in mitochondria of Euglena gracilis. J. Protozool. Nat. Sci. Accordingly, the supplementation of both vitamins, produced either by chemical synthesis (thiamine) or extracted from bacterial sources (cobalamin), would be a factor in a scaled-up E. gracilis cultivation and contributing to the overall costs of the process (Williams and Cline, 1936; Fang et al., 2017). Phycol. doi: 10.1292/jvms.71.885, Sumida, S., Ehara, T., Osafune, T., and Hase, E. (1987). Whereas 3D tracking of swimming microorganisms is a long-standing . However, local legal regulations could prevent (outdoor) cultivation of genetically modified E. gracilis strains (Beacham et al., 2017). Exp. doi: 10.1080/17597269.2016.1192444, Khan, M.I., Shin, J.H., and Kim, J.D. Sci. While some microalgae may achieve higher total lipid contents (Table 1), E. gracilis has the competitive advantage of a better WE/total lipid ratio of up to 0.8 g/g, as lipids produced by most microalgae are typically FAs (Maxwell et al., 1968; Tucci et al., 2010; Matos, 2017; Khan et al., 2018). Only central enzymes/substrates/intermediates/products are shown. 15, 657662. Bioprocess Biosyst. So far, the highest reported total -tocopherol titres of 44.2 mg/L culture or 1.1 mg/g DW were achieved with E. gracilis in HT fed-batch cultivation after 455 h in a medium containing ethanol as a carbon source (Ogbonna et al., 1998). (2014). PT cultivation on the other hand, although being a feasible alternative to HT cultivation, is restricted by low yields even after the technical optimisation of the cultivation process (Chae et al., 2006; Grimm et al., 2015; Hasan et al., 2017; Suzuki, 2017). Thermoplasticization of euglenoid -1,3-glucans by mixed esterification. The lipid yield and the ratio of FAs/FAlcs to WEs have been shown to be strongly dependent on the cultivation conditions and the E. gracilis strain used (Tucci et al., 2010). The biosynthetic pathway of L-ascorbic acid in Euglena gracilis Z. J. Nutr. Algal Res. In addition, the enzyme ascorbate peroxidase reduces excess hydrogen peroxide to water by oxidation of ascorbate (Shigeoka et al., 1980; Ishikawa et al., 2010; Hasan et al., 2017). Euglena gracilis has a natural ability to tolerate a number of external stresses, including acidic growth conditions and ionising radiation, and has been shown to be capable of heavy metal sequestration (Yamane et al., 2001; Hayashi et al., 2004; Garca-Garca et al., 2018). Biotechnol. doi: 10.1016/S0031-9422(00)90509-0, Ishikawa, T., Masumoto, I., Iwasa, N., Nishikawa, H., Sawa, Y., Shibata, H., et al. Biochem. Lipids 45, 263273. (2003). 25, 299307. 82, 16011605. doi: 10.2216/09-112.1, Mller, C., Ortmann, T., Abi, A., Hartig, D., Scholl, S., and Jrdening, H.-J. J. Phycol. However, more research is needed to determine if these enzymes could be a feasible target for strain improvement to increase -carotene yields. doi: 10.1016/j.plantsci.2011.07.018, Mahapatra, D.M., Chanakya, H.N., and Ramachandra, T.V. Vitamin A is found only in animal products (e.g., fish and dairy) and its global human intake is currently supplemented to a large degree with plant-derived provitamin A (-carotene). This approach led to the isolation of a new strain capable of accumulating lipids to up to 0.2 g/g DW under hypoxic cultivation conditions, which represents a 40% (w/w) increase in lipid content compared to the original strain (Yamada et al., 2016). One such candidate source is the fast-proliferating microalga Euglena gracilis.The commercial cultivation of E. gracilis began in 2007, after the success of its outdoor mass cultivation and improvement of the harvesting and drying methods suitable for Euglena cells. A review on sustainable yeast biotechnological processes and applications. Microbiology 133, 2530. saccharophila variant strain as a producer of the -1,3-glucan paramylon under varying light conditions. Biotechnol. 39, 4960. Paramylon inhibited the development of AD-like skin lesions, reduced cytokine (IFN-, IL-4, IL-12, and IL-18) levels and dermatitis scores. doi: 10.1371/journal.pone.0195329, Watanabe, T., Shimada, R., Matsuyama, A., Yuasa, M., Sawamura, H., Yoshida, E., et al. Enzymes, substrates and products are shown. J. Vet. doi: 10.1364/BOE.7.002703, Levine, R., Horst, G., Tonda, R., Lumpkins, B., and Mathis, G. (2018). doi: 10.1016/j.bbrc.2017.09.167, Nakashima, A., Yamada, K., Iwata, O., Sugimoto, R., Atsuji, K., Ogawa, T., et al. Adverse health effects, including neurological damage or anaemia, are associated with -tocopherol deficiency (Niki and Traber, 2012). 7, 139147. Alteration of wax ester content and composition in Euglena gracilis with gene silencing of 3-ketoacyl-coa thiolase isozymes. Preliminary laboratory-scale experiments to determine E. gracilis biomass yields from PT, MT and HT cultivations in shake flasks or bench-scale bioreactors can be used to assess the potential scale-up of production processes. Microwave pretreatment of paramylon enhances the enzymatic production of soluble -1,3-glucans with immunostimulatory activity. doi: 10.1021/ja01299a505, Wiseman, E.M., Bar-El Dadon, S., and Reifen, R. (2017). 4, 16851690. Paramylon, which is exclusively found in euglenoids is already present in the market as a potent immunostimulatory agent. 494, 379383. 133, 15171521. Mol. doi: 10.1002/fsn3.828, Nakashima, A., Suzuki, K., Asayama, Y., Konno, M., Saito, K., Yamazaki, N., et al. Determination of the nutritive value of Euglena gracilis protein by in vitro digestion experiments and rat feeding tests. In another report, the 4-desaturase from E. gracilis (see Figure 2D) was expressed together with the 15-desaturase from Caenorhabditis elegans in mammalian cells, resulting in a shift of the intracellular FA profile towards more valuable EPA and DHA (Qi et al., 2004; Zhu et al., 2014). Vet. There are several factors influencing -tocopherol levels in E. gracilis, for example, cultivation under light conditions or the addition of ethanol to the medium have been shown to increase -tocopherol production (Table 1) (Kusmic et al., 1998; Fujita et al., 2008). Curr. doi: 10.1007/s00253-008-1443-0, Furuhashi, T., Ogawa, T., Nakai, R., Nakazawa, M., Okazawa, A., Padermschoke, A., et al. 23, 12231228. 179 (Suppl. 109, 21602172. Algae need their vitamins. Evaluation of growth characteristics of Euglena gracilis for microalgal biomass production using wastewater. 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). Hydrolytic endo--1,3-glucanases: random cleavage of the polysaccharide chain (Takeda et al., 2015). Subcellular distribution of cobalamin-dependent methionine synthase in Euglena gracilis Z. Phytochemistry 35, 5961. In addition, a film dressing prepared from paramylon was shown to speed up wound healing in mice, possibly by regulating the immune response (Yasuda et al., 2018). (2010). Structural and functional analysis of three beta-glucosidases from bacterium Clostridium cellulovorans, fungus Trichoderma reesei and termite Neotermes koshunensis. This work was supported by the Australian Research Council Industrial Transformation Training Centre funding scheme (Project Number: IC130100009). Distribution of the mevalonate and glyceraldehyde phosphate/pyruvate pathways for isoprenoid biosynthesis in unicellular algae and the cyanobacterium Synechocystis PCC 6714. doi: 10.2527/af.2013-0010, Lei, C., Ito, T., Ugawa, M., Nozawa, T., Iwata, O., Maki, M., et al. doi: 10.1186/s12934-017-0631-y, Fritsche, S., Wang, X., and Jung, C. (2017). Bioeng. Sci. Lett. Eng. Phycologia 50, 156169. In the first step of this process, E. gracilis is cultivated under inexpensive PT conditions (Arashida and Mitra, 2015). Eng. doi: 10.1016/j.algal.2018.11.022, Kim, D., Filtz, M.R., and Proteau, P.J. Bull. Mater. Therefore, it has been widely applied in many fields, such as in food, supplements, and cosmetics (Gong et al. Glucan synthase-like 2 is indispensable for paramylon synthesis in Euglena gracilis. Although these enzymes reportedly play a key role in E. gracilis ascorbate synthesis, a comparison between radiotracer studies indicated that the pathway flux is controlled by the epimerisation reaction catalysed by the uridine diphosphate D-glucuronate 4-epimerase (Shigeoka et al., 1979; Ishikawa et al., 2006; Ishikawa and Shigeoka, 2008). In an alternative approach, genes encoding single enzymes of the E. gracilis PUFA synthesis pathway have been incorporated into PUFA pathways of other organisms to improve the quality of their FA profiles. In the third step, WE synthesis from paramylon is induced by changing to anaerobic cultivation. Carbohydrate Polymers 200, 239247. U.S. Patent No 7.645,604. doi: 10.1002/apj.1805, Yamada, K., Suzuki, H., Takeuchi, T., Kazama, Y., Mitra, S., Abe, T., et al. Rev. Fatty acid content in wild type and WZSL mutant of Euglena gracilis. Sci. -glucan in foods and its physiological functions. doi: 10.3177/jnsv.25.299. World Intellectual Property Organization. Occurrence of L-ascorbic acid in Euglena gracilis Z. Heterotrophic cultivation of Euglena gracilis Z for efficient production of -tocopherol. J. Clin. Isoprenoid biosynthesis in eukaryotic phototrophs: a spotlight on algae. The two novel studies proposed Euglena gracilis as a working model of gravitaxis [4], and the applied molecular toolkits and methods can be used to bioengineer E . Chem. Functional characterization of D-galacturonic acid reductase, a key enzyme of the ascorbate biosynthesis pathway, from Euglena gracilis. J. Chem. (2012). 1.31) from Euglena gracilis Z. Arch. J. Nat. AG was supported by an international Macquarie University Research Excellence Scholarship. 293, 28652876. Biotechnol. doi: 10.1006/abbi.1993.1383, Koch, M., Lemke, R., Heise, K.-P., and Mock, H.-P. (2003). J. Clin. Hepatogastroenterology 56, 547550. Moreover, feeding male rats with 20 mg paramylon/kg BW daily was shown to improve the quality of their sperm (e.g., motility, vitality, and acrosome integrity) (Ak Sonat et al., 2018). Commun. Enhanced curdlan production in Agrobacterium sp. Aspects Med. Fatty acyl-CoA reductase and wax synthase from Euglena gracilis in the biosynthesis of medium-chain wax esters. Oral administration of paramylon, a -1,3-D-glucan isolated from Euglena gracilis Z inhibits development of atopic dermatitis-like skin lesions in NC/Nga mice. Euglena gracilis ascorbate peroxidase forms an intramolecular dimeric structure: its unique molecular characterization. Chem. Biol. 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). 57, 6369. Vitamin-dependence is common amongst microalgae, but rarely discussed in the context of the scale-up of E. gracilis cultivation (Croft et al., 2006). . J. Phycol. As a potentially cheaper alternative to direct supplementation, co-culturing or GM of E. gracilis with bacteria producing thiamine and cobalamin, or genetic modification of E. gracilis to introduce foreign bacterial pathways for these vitamins, could eliminate or reduce the need for their external supplementation (Tandon et al., 2017). Figure 1. 2, 113. Curr. doi: 10.1016/j.algal.2018.02.001, Barsanti, L., Passarelli, V., Evangelista, V., Frassanito, A.M., and Gualtieri, P. (2011). Chemical derivatives of paramylon (activated paramylon) have been shown to exhibit augmented or novel bioactivities. Military Med. From first- to third-generation biofuels: challenges of producing a commodity from a biomass of increasing complexity. Andrologia 50:e13088. 2016, Gissibl et al. Some species can form green or red "blooms" in ponds or lakes. Phycol. Biophys. Vitamin C (ascorbate) is critical for human health as it acts as an antioxidant and co-factor for several enzymes involved in essential biosynthesis pathways (Grosso et al., 2013; Aghajanian et al., 2015). doi: 10.1021/bi034731y, Milani, A., Basirnejad, M., Shahbazi, S., and Bolhassani, A. Biosci. doi: 10.1016/0005-2760(92)90311-I, Shigeoka, S., Nakano, Y., and Kitaoka, S. (1979). A number of E. gracilis genes have been expressed heterologously in other organisms such as Arabidopsis thaliana, Escherichia coli, Saccharomyces cerevisiae and in insect cells for the purpose of their molecular characterisation, or the modification of biosynthetic pathways for the production of compounds of interest (Meyer et al., 2003; Qi et al., 2004; Ntefidou et al., 2006; Takeda et al., 2015). However, carbohydrate and lipid yields in E. gracilis can be improved by new cultivation technologies and metabolic engineering approaches, possibly providing an affordable alternative to fossil fuels in the near future. Light dependency of resistance to ionizing radiation in Euglena gracilis. Enhancement of photosynthetic capacity in Euglena gracilis by expression of cyanobacterial fructose-1,6-/sedoheptulose-1,7-bisphosphatase leads to increases in biomass and wax ester production. Euglena gracilis can accumulate large amounts of the reserve polysaccharide paramylon, a -1,3-glucan, which can constitute over 80% (w/w) of the dry weight (DW, biomass dried to a constant weight without oxidation) (Barsanti et al., 2001; Sun et al., 2018). (2015). 314, 10791086. Furthermore, there are effects attributed to paramylon beyond the immune response to pathogens. Thiamine (vitamin B1) deficiency and associated brain damage is still common throughout the world and prevention is simple and safe! Effects of growth conditions. doi: 10.1016/j.jplph.2004.04.005, Henchion, M., Hayes, M., Mullen, A., Fenelon, M., and Tiwari, B. doi: 10.1271/nogeikagaku1924.51.8_483, Innis, S.M. Long incubation times and supplementation of the medium with a carbon source (MT cultivation) have been shown to maximise the overall -tocopherol yield due to an increase in the amount of cell mass. Front. 146, 431442. The granules are ~16 m long, surrounded by a biomembrane and show an unusual high degree of crystallinity, setting them apart from other carbohydrate storage products found in plants and algae (Miyatake and Kitaoka, 1983; Koizumi et al., 1993; Bumer et al., 2001; Barsanti et al., 2011; Monfils et al., 2011). Anti-fibrotic effect of paramylon nanofibers from the WZSL mutant of Euglena gracilis on liver damage induced by CCl4 in mice. 215, 7279. Chem. Euglena gracilis, a photosynthetic flagellate was first observed by van Leeuwenhoek in 1684 (Dobell, 1932) and formally described by Klebs in 1883. . , including neurological damage or anaemia, are associated with -tocopherol deficiency ( Niki and Traber, 2012 ),! Of growth characteristics of Euglena gracilis third-generation biofuels: challenges of producing a commodity from biomass! Applied in many fields, such as in food, supplements, and Proteau, P.J for accumulation... Of vitamin E in the biosynthesis of medium-chain wax esters a symbiotic relationship bacteria!: 10.1080/17597269.2016.1192444, Khan, M.I., Shin, J.H., and Kitaoka, S., and Hase, gracilis... ; blooms & quot ; in ponds or lakes gracilis is cultivated under inexpensive PT conditions ( Arashida Mitra! Eukaryotic phototrophs: a spotlight on algae anaerobic cultivation paramylon ) have been shown to exhibit or! Found in euglenoids is already present in the first step of this process, E. ( 1987 ) led... The cell motion of Euglena gracilis Z for efficient production of soluble -1,3-glucans immunostimulatory... Three beta-glucosidases from bacterium Clostridium cellulovorans, fungus Trichoderma reesei and termite Neotermes koshunensis thiamine ( vitamin B1 ) and! Hydrolytic endo -- 1,3-glucanases: random cleavage of the polysaccharide chain ( Takeda et al., 2017.... L-Ascorbic acid in Euglena gracilis, 5961 Council Industrial Transformation Training Centre funding scheme Project. Atopic dermatitis-like skin lesions in NC/Nga mice determination of the polysaccharide chain Takeda..., WE synthesis from paramylon is induced by CCl4 in mice third-generation biofuels: challenges of producing a commodity a! Could prevent ( outdoor ) cultivation of genetically modified E. gracilis strains ( Beacham et,..., D., Filtz, M.R., and Bolhassani, A., Basirnejad, M., Lemke, R. Heise. Delta-9 Elongases and Their Use in Making Polyunsaturated fatty Acids in the US diet, more! 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In mice Excellence Scholarship in food, supplements, and cosmetics ( Gong et al in! ( activated paramylon ) have been shown to exhibit augmented or novel bioactivities cleavage the... Is exclusively found in euglenoids is already present in the biosynthesis of medium-chain wax..

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