photosynthetic stramenopiles

Boudire L., Bott C. Y., Saidani N., Lajoie M., Marion J., Brhlin L., et al. Chromalveolates include very important photosynthetic organisms, such as diatoms, brown algae, and significant disease agents in animals and plants. Kory N., Farese R. V., Walther T. C. (2016). Radakovits R., Jinkerson R. E., Fuerstenberg S. I., Tae H., Settlage R. E., Boore J. L., et al. Bacillariophyceans (Diatoms). Data from yeast as well as animals suggest that such domains are sites of both LD and peroxisome biogenesis (Joshi et al., 2018; Wang et al., 2018), thus linking the two processes. Boudire L., Michaud M., Petroutsos D., Rbeill F., Falconet D., Bastien O., et al. Genome, functional gene annotation, and nuclear transformation of the heterokont oleaginous alga. As mentioned above, TAG production and storage are induced under stress conditions, which results in cell cycle slowdown or arrest. In yeast, the seipin homolog Fld1 function depends on its interaction with Ldb16 (Wang C.-W. et al., 2014). Disclaimer, National Library of Medicine In addition, DAG may derive from membrane glycerolipids, like PC, via the action of phospholipases C (PLC) or phosphatidylcholine:diacylglycerol choline transferases (PDAT). The central luminal part is involved in seipins oligomerization, thus forming a channel-like structure that stabilizes the ERLD junction (Binns et al., 2010; Sui et al., 2018; Yan et al., 2018) and is essential to ensure unidirectionality of the TAG and/or sterolester flow from the ER to the LD (Salo et al., 2019). It was later on shown to be likely involved in LD degradation through binding to autophagy-related protein 8 (ATG8) (Zienkiewicz et al., 2020). The cells have two heterokont flagella, inserted parallel to one another at the anterior, whose ultrastructure is a distinguishing characteristic of the group. Full-length transcriptome analysis of the bloom-forming dinoflagellate, Genome evolution of a nonparasitic secondary heterotroph, the diatom. Recently, the human DGAT1 enzyme (HsDGAT1) was structurally characterized (Sui et al., 2020) by cryo-electron microscopy, revealing that the protein contains nine transmembrane helices with the amino-terminus (N-ter) spreading in the cytosol and the carboxyl-terminus (C-ter) in the ER lumen. (2014). This stramenopile cell has a single hairy flagellum and a secondary smooth flagellum. After lense growth, complex mechanisms lead to the budding of an LD from the ER membrane at specialized domains. {"type":"entrez-protein","attrs":{"text":"AFB75402","term_id":"377774508","term_text":"AFB75402"}}, Predicted structures of major lipid droplet proteins found in stramenopile models. Thus, in the early step of the Kennedy pathway, LPAT isoforms seem to control the fate of the produced phosphatidic acid, upstream membrane glycerolipid, and/or TAG pathways. Leyland B., Boussiba S., Khozin-Goldberg I. This interaction is conserved in mammals for which Promethin/TMEM159/LDAF1 (lipid droplet assembly factor) has been identified as the Ldo45 homolog (Castro et al., 2019; Chung et al., 2019). (2013). Algal genomes reveal evolutionary mosaicism and the fate of nucleomorphs. (2019) on mature LDs (7 days of nitrogen starvation), StLDP and PtLDP1 were both recorded in the LD proteome, although no major LD protein was pointed out. ATPase activity associated with the uncoating of clathrin baskets by Hsp70. Oku M., Maeda Y., Kagohashi Y., Kondo T., Yamada M., Fujimoto T., et al. Analyses of TAG species in the OE lines show an increase in C48 containing TAG, suggesting that PtDGAT2B may prefer C16 species and C16-containing DAG as substrates (Haslam et al., 2020). LDs as well (Peled et al., 2011; Tsai et al., 2015; Lupette et al., 2019). Stramenopiles, also known as heterokonts, are a very large and diverse phylum (Derelle et al., 2016). Arrows show hypothetical conversions of membrane lipid down-products in the production of TAG, as well as in the increase in very long-chained PUFA (e.g., 20:5) in LDs. SEIPIN proteins mediate lipid droplet biogenesis to promote pollen transmission and reduce seed dormancy. (B,E,H) Predictions of transmembrane helices obtained with TMHMM prediction server (http://www.cbs.dtu.dk/services/TMHMM/). (2020). The .gov means its official. Effects of organic carbon sources on cell growth and eicosapentaenoic acid content of. Human health benefits of very-long-chain polyunsaturated fatty acids from microalgae. In addition to lipid metabolism enzymes as mentioned above, seipin interacts with different proteins depending on the organism. Weng L.-C., Pasaribu B., -Ping Lin I., Tsai C.-H., Chen C.-S., Jiang P.-L. (2015). The architecture of the secondary plastids is unique, and the outermost membrane, the EpM, is in continuity with the ER (see above). Brown Algae (Phaeophycean) Diversity. 2022 Aug;153(1-2):59-70. doi: 10.1007/s11120-022-00915-w. Epub 2022 Apr 7. In P. tricornutum, fucoxanthin, and -carotene are the main carotenoids in LD-enriched fractions, while only traces of other pigments were detected (Lupette et al., 2019). This view is now considered as partly incomplete, since numerous chloroplastic proteins, including the majority of enzymes involved in membrane lipid biosynthesis, are of non-cyanobacterial origin (Petroutsos et al., 2014; Cenci et al., 2017; Sato and Awai, 2017; Marechal, 2018). DGDG, digalactosyldiacylglycerol; EPA, eicosapentaenoic acid; MGDG, monogalactosyldiacylglycerol; PC, phosphatidylcholine; PG, phosphatidylglycerol; SQDG, sulfoquinovosyldiacylglycerol; TAG, triacylglycerol. Consistent with the presence of ubiquitination proteins, four 26S proteasome proteins are also present in the LD proteome (Lupette et al., 2019). Erdene-Ochir E, Shin BK, Huda MN, Lee EH, Song DG, Jung C, Pan CH. (2019a). (2018). The engulfed organisms underwent genetic reductions, a loss of autonomy to such an extent that they fully integrated the host cell, giving rise to organelles, usually photosynthetic, collectively termed plastids. The initial acquisition of a cyanobacterium by an unknown ancestral heterotrophic eukaryote occurred around 1 to 1.5 billion years ago (Jensen and Leister, 2014) leading to the emergence of the primary plastidthe so-called chloroplast (Figure 2). LDSP, StLDP, and PtLDP1 all contain hydrophobic sequences harboring a proline-rich region (Figure 6), similar to the proline knot motif found in plant oleosins (Abell et al., 2004). In P. tricornutum, genes coding for a putative PDAT have been predicted (Dolch et al., 2017b), but to our knowledge, not formally characterized. In P. tricornutum under phosphate deprivation, phospholipids are supposed to also provide part of the EPA in TAG molecular species (Abida et al., 2015), and this was also hypothesized in M. gaditana under nitrogen deprivation (Janssen et al., 2019a). We compared a pennate (Phaeodactylum tricornutum) and a centric diatom (Thalassiosira pseudonana) with those of other stramenopiles (oomycetes, Pelagophyceae, and Phaeophyceae (Ectocarpus siliculosus)) as well as to that of Emiliania huxleyi, a haptophyte that is evolutionarily related to the stramenopiles. In N. oceanica under nitrogen shortage, a homolog of the Arabidopsis galactolipase PSD1 is upregulated, coincidently with degradation of MGDG (Li et al., 2014). EpM, epiplastidial membrane; PPM, periplastidial membrane; P-oEM and P-iEM, plastid outer and inner envelope membranes, {"type":"entrez-nucleotide","attrs":{"text":"AY875644","term_id":"1127827640","term_text":"AY875644"}}, {"type":"entrez-nucleotide","attrs":{"text":"MN061782","term_id":"1818800654","term_text":"MN061782"}}, {"type":"entrez-nucleotide","attrs":{"text":"JX469835","term_id":"401836789","term_text":"JX469835"}}, {"type":"entrez-nucleotide","attrs":{"text":"JQ837823","term_id":"390994298","term_text":"JQ837823"}}, {"type":"entrez-nucleotide","attrs":{"text":"JX469836","term_id":"401836791","term_text":"JX469836"}}, {"type":"entrez-nucleotide","attrs":{"text":"JX469837","term_id":"401836793","term_text":"JX469837"}}, {"type":"entrez-nucleotide","attrs":{"text":"XM_002184438","term_id":"219128552","term_text":"XM_002184438"}}, {"type":"entrez-protein","attrs":{"text":"XP_002184474","term_id":"219128553","term_text":"XP_002184474"}}, Different hypotheses regarding lipid droplet (LD) biogenesis in. Glycerolipid conversions occur to rescue phospholipid degradation. Are oomycetes Stramenopiles? and transmitted securely. Cenci U., Bhattacharya D., Weber A. P. M., Colleoni C., Subtil A., Ball S. G. (2017). Cytosolic triacylglycerol biosynthetic pathway in oilseeds. Epub 2013 May 8. TAG composition in SFA, MUFA, and VLC-PUFA reflects therefore the metabolic context that gave rise to the formation of the LD, either via an early partitioning of carbon following FA de novo synthesis and/or a recycling of FA from membrane lipids, e.g., plastid galactolipids or endomembrane phosphor- or betaine lipids. Rapoport I., Boll W., Yu A., Bcking T., Kirchhausen T. (2008). Disentangling the mechanism of each of the DGAT proteins may not only increase our knowledge of stramenopile metabolisms but also can be functional to biotechnological aims. Flori S., Jouneau P.-H., Finazzi G., Marchal E., Falconet D. (2016). Besides de novo synthesis via DGATs, TAGs may accumulate through glycerolipid recycling via the activity of a so-called phospholipid:diacylglycerol acyltransferase, or PDAT (Dahlqvist et al., 2000; Ma et al., 2016; Zulu et al., 2018; Falarz et al., 2020). Nomaguchi T., Maeda Y., Liang Y., Yoshino T., Asahi T., Tanaka T. (2018). However, it is not possible to conclude if the helices are in the same direction. Prokaryotic Pathway is not prokaryotic: noncyanobacterial origin of the chloroplast lipid biosynthetic pathway revealed by comprehensive phylogenomic analysis. The Tree of Eukaryotes. (2019a). Transcriptional coordination of physiological responses in. oomycetes. Plastids in Plant Cells Function & Types | What do Plastids Do? In N. oceanica, the ATG8 expression level is induced directly after the transition from nitrogen-depleted condition to nitrogen replete (Zienkiewicz et al., 2020). copyright 2003-2022 Study.com. Schematic representation of primary and secondary endosymbiosis, and organelle architecture. Test Prep. Adl S. N., Simpson A. G. B., Farmer M. A., Andersen R. A., Anderson R. O., Barta J. R., et al. (2007). Photosynthetic stramenopiles, such as diatoms, brown algae, and red tide algae (Raphidophyceae), are predominant species in aquatic ecosystems. These components could be linked with the presence in the unfiltered proteome of SNARE proteins, whose function is necessary for fusion of transport vesicle with target membrane (Tanaka et al., 2015), and which play a role in COP1 transport (Lupette et al., 2019). official website and that any information you provide is encrypted Based on gene ontology, LD-associated proteins can be divided in seven large functional and structural groups: metabolism, membrane organelle, chaperones/protein folding/post translational modification/quality control, cytoskeleton, genomic information processing, plastoglobules, and unknown (Lupette et al., 2019). In this review, we summarize the current knowledge on the relationship between cytosolic lipid droplets (LDs) and the complex membrane compartmentalization within stramenopile cells, the metabolic routes leading to TAG accumulation, and the physiological conditions in which LDs are produced, in response to various environmental factors. However, different suppositions have been made about the origin of the additional bounding membranes. 2022 Oct 17;13:993914. doi: 10.3389/fmicb.2022.993914. A recent analysis in N. oceanica has focused on the four copies of LPATs, addressing their subcellular location and function in the synthesis of eukaryotic precursors, based on single and double knockout (KO) studies (Nobusawa et al., 2017). Changes in the cell composition of the marine microalga. Nitric oxide mediates nitrite-sensing and acclimation and triggers a remodeling of lipids. The outermost membrane, called the epiplastidial membrane (EpM), could derive from the host phagocytic membrane. The abundance of C16:2 on the betaine lipid DGTA found in the lipid droplet monolayer (Lupette et al., 2019) supports a plastid emerging hypothesis as this fatty acid is major in the plastid but very minor in endomembranes (Abida et al., 2015). LDSP, StLDP, and PtLDP1 gene expression can be induced by stress conditions leading to lipid accumulation (Yoneda et al., 2016; Wang et al., 2017; Zienkiewicz et al., 2020), and their function is related to the modulation of LD size (Vieler et al., 2012a; Yoneda et al., 2016; Wang et al., 2017). Diverse phospholipases A, C, and D (PLA, PLC, and PLD, respectively) were also identified in both P. tricornutum and Nannochloropsis s.l. For DOAP1 and PtLDP1, a beta barrel structure bordered with two alpha helices is predicted. -. Glycerolipids in photosynthesis: composition, synthesis and trafficking. Ultrastructure and taxonomy of a marine photosynthetic stramenopile Phaeomonas parva gen. et sp. Architecture of lipid droplets in endoplasmic reticulum is determined by phospholipid intrinsic curvature. Generating an ePub file may take a long time, please be patient. The origin and establishment of the plastid in algae and plants. However, stramenopiles can get much larger as well. (2020). Schematic representation of primary and secondary endosymbiosis, and, Architecture of the membrane bound O -acyl transferase (MBOAT)-core of human diacylglycerol acyltransferase, Different hypotheses regarding lipid droplet, Different hypotheses regarding lipid droplet (LD) biogenesis in Phaeodactylum tricornutum . Although they are only made of a single cell, both diatoms and golden algae can form massive ribbons and blooms composed of thousands of individual cells together. In the latest eToL, based almost entirely on multigene molecular phylogenies (Figure 1; Burki et al., 2020), microalgae cluster in numerous supergroups. Properties and biotechnological applications of Acyl-CoA:diacylglycerol Acyltransferase and Phospholipid:diacylglycerol Acyltransferase from terrestrial plants and microalgae. (2018b). (2018). Two components of the clathrin vesicle coating system (CHC and PTAP1/2BETA) were also identified (Lupette et al., 2019). Architecture of the membrane bound O-acyl transferase (MBOAT)-core of human diacylglycerol acyltransferase (DGAT1) enzyme (HsDGAT1). Purification of LDs consisted in (i) treatment of cells to nutrient starvation (mostly nitrogen deprivation) to trigger LD formation, (ii) cell disruption, and (iii) LD isolation, generally through sucrose density centrifugation. The LD core is mainly made of TAGs, but pigments have also been detected, and the presence of other hydrophobic compounds cannot be excluded. In addition, membrane contact sites were detected between the inner membrane of the nuclear envelope and the PPM (Flori et al., 2016). An OpenStax CNX book. In the present genome database, AUREO orthologs are only conserved in photosynthetic stramenopiles, such as P. tricornutum possesses several DGATs, namely, one PtDAGT1, five PtDGAT2, and a PtDGATX (Cui et al., 2018) or PtDGAT3 (Zhang et al., 2020), with a double wax ester synthase (WS) and DGAT function (Cui et al., 2013, 2018). Adapted from Sui et al. NoLPAT1 proved to be mainly involved in the transfer of 16:1 to the sn-2 position of LPA used for the synthesis of membrane glycerolipids, particularly PC and DGTS. stramenopiles. De novo synthesis produces long-chain saturated or monounsaturated fatty acids (SFA, MUFA), whereas subsequent maturation processes lead to very long-chain polyunsaturated FA (VLC-PUFA). The other subgroup of chromalveolates, the stramenopiles, includes photosynthetic marine algae and heterotrophic protists. Conversely, no homologs have been found in Nannochloropsis s.l. In Arabidopsis, AtSDP1 is a TAG lipase that plays a key role in TAG hydrolysis during seed germination (Eastmond, 2006). contain 16 carbons, whereas longer chained FAs are produced by an elongation process occurring in the ER (Dolch et al., 2017b). (2014). To test the idea of a photosynthetic ancestry for ciliates, we used the 27,446 predicted proteins from the macronuclear genome of Tetrahymena thermophila to query prokaryotic and eukaryotic genomes. In N. oceanica under nitrogen deprivation, several genes encoding for putative patatin and one putative lysophospholipase were upregulated under nitrogen deprivation (Li et al., 2014; Jia et al., 2015). Interestingly, a vesicular network forming a blob-like structure has been detected between the PPM and the EpM in P. tricornutum, but its function is still elusive (Kilian and Kroth, 2004; Flori et al., 2016). Leyland B, Zarka A, Didi-Cohen S, Boussiba S, Khozin-Goldberg I. J Phycol. At their vegetative state, thraustochytrids measure 4 to 20 m in diameter and are globose or subglobose in shape. Photosynthetic Stramenopiles; pp. Schematic representation of primary and secondary endosymbiosis, and organelle architecture. Conte M., Lupette J., Seddiki K., Me C., Dolch L.-J., Gros V., et al. Briefly, TAG and sterol ester synthesis occurs between the two leaflets of the ER membrane leading to the formation of lenses once they reach a certain concentration (Choudhary et al., 2015; Thiam and Fort, 2016). SUMMARY The Pinguiophyceae class. The ER pathway is often referred to as the eukaryotic pathway, and leads to glycerolipids with various signatures, depending on the species. Its like a teacher waved a magic wand and did the work for me. In P. tricornutum, MGDG, PG, and PE decrease, while other lipid classes are stable (Abida et al., 2015; Figure 5). Murine DGAT2 shows two TM domains and both N- and C- termini in the cytosol (Stone et al., 2006). The https:// ensures that you are connecting to the Photosynthetic Stramenopiles; pp. (2016). The novel annotation pointed out the presence in the N-ter of the protein of a pleckstrin homology (PH) domain (Zhang et al., 2020), involved in signaling and membrane binding. | {{course.flashcardSetCount}} Would you like email updates of new search results? Since PDAT allows a connection between TAG production and membrane lipid turnover, a role of the reorganization of cellular membranes needs also to be investigated. In support to the latter hypothesis, electron microscopy inspections show association of the LD with endomembranes in P. tricornutum (Lupette et al., 2019). 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. Su W.-C., Lin Y.-H., Pagac M., Wang C.-W. (2019). Pasquet V., Ulmann L., Mimouni V., Guihneuf F., Jacquette B., Morant-Manceau A., et al. Janssen J. H., Spoelder J., Koehorst J. J., Schaap P. J., Wijffels R. H., Barbosa M. J. That can be the case with stramenopiles, a large group of organisms composed of cells with a distinct form of chlorophyll. Proteins are inserted into or associated to the LD bounding lipid monolayer. However, stramenopile cells contain a complex plastid deriving from a secondary endosymbiosis, limited by four membranes, the outermost one being connected to the endomembrane system. Diatoms die, their hard frustules sink to the bottom of the ocean or lake, and build up layers of silica-rich sediment. Recent cell imaging and proteomic studies suggest that at least some cytosolic LDs might be associated to the surface of the complex plastid, via still uncharacterized contact sites. The characteristics of TAG and EPA accumulation in. Funding. Seipin and the membrane-shaping protein Pex30 cooperate in organelle budding from the endoplasmic reticulum. Nannochloropsis s.l. Xin Y., Lu Y., Lee Y.-Y., Wei L., Jia J., Wang Q., et al. The last step of TAG formation can alternatively occur using FA transferred from an existing membrane glycerolipid used as a donor. Laterally, a gate was identified that opens up in between the two membrane sheets (Figure 3) and driving the DAG to the core of the dimer, where the active histidine is located (Sui et al., 2020). Its function, activity, and specificity were characterized both in vitro and by heterologous expression in yeast (Zhang et al., 2020). The carbon length and number of double bonds of the acyl groups contained in the TAG molecules depend on their origin. nov., a new class of photosynthetic stramenopiles (chromophytes), is described. | Find, read and cite all the research you need on . Buitrago-Flrez FJ, Restrepo S, Riao-Pachn DM. By contrast, NoLPAT4 transfers 16:0 to the sn-2 position of LPA purely dedicated to TAG. The chromalveolates can be subdivided into alveolates and stramenopiles. Epub 2016 Jan 18. Zienkiewicz A., Zienkiewicz K., Poliner E., Pulman J. Sui X., Wang K., Gluchowski N. L., Elliott S. D., Liao M., Walther T. C., et al. (2019). Nel 2012 Diaphoretickes ha ricevuto la seguente . Cytosolic LD biogenesis is usually described as an ER process. Understanding the physiological contexts leading to the formation of TAG with various FA composition is therefore essential. 8. Another major peculiarity of stramenopiles is the biogenesis site of LD. Alternatively, down-products can be recycled to form TAG. Castro I., Eisenberg-Bord M., Persiani E., Rochford J., Schuldiner M., Bohnert M. (2019). Stepwise biogenesis of subpopulations of lipid droplets in nitrogen starved. Hydrophobicity level of AAs is represented with a color code in (C,F,I,J), ranging from dark blue for very hydrophobic AAs, light blue for hydrophobic AAs, green for poorly hydrophobic or hydrophilic AAs, yellow for hydrophilic AAs, and red for very hydrophilic AAs. Elongation of FAs to 18 or more carbons involves the export of C16 from the plastid to the ER. Chimeric origins of ochrophytes and haptophytes revealed through an ancient plastid proteome. Temperature variations were also reported to alter the extent of TAG accumulation (Pasquet et al., 2014; Gill et al., 2018). Enzymes have been purified, functionally and molecularly characterized, and metabolic pathways reconstructed at the cellular level. (2019) used inhibitors of membrane trafficking-related proteins and lipophagy (Nonoyama et al., 2019). Isogamy to oogamy are known in the sexual reproduction of stramenopiles. Front Microbiol. Characterization of endogenous promoters of GapC1 and GS for recombinant protein expression in Phaeodactylum tricornutum. The Tree of Eukaryotes. (2000). Nojima D., Yoshino T., Maeda Y., Tanaka M., Nemoto M., Tanaka T. (2013). Choreography of transcriptomes and lipidomes of. The number, speed, and impact of plastid endosymbioses in eukaryotic evolution. Stramenopiles Photosynthetic Stramenopiles Ochrophytes Synurophyceans SilicaScaled Algae Tribophyceans Boutet E., El Mourabit H., Prot M., Nemani M., Khallouf E., Colard O., et al. On the other hand, proteins involved in post translational modification such as ubiquitination (UBI3 and PUB39) and phosphorylation (STK and EPK2) were found, as well as proteins involved in the endoplasmic reticulum-associated degradation (ERAD) system (CDC48 and BIP) (Lupette et al., 2019). Aspects of phosphorus physiology associated with phosphate-induced polar lipid remodelling in marine microalgae. Ben Mbarek K., Ajjaji D., Chorlay A., Vanni S., Fort L., Thiam A. R. (2017). Such a profile differs from the general assumption of LD monolayer exclusively composed of phospholipids. In this review, we address the relationship between cytosolic LDs and the complex membrane compartmentalization within stramenopile cells, the metabolic routes leading to TAG accumulation, and the physiological conditions that trigger LD production, in response to various environmental factors. (2020). The membrane-spanning sequences of the beta barrel do not seem as enriched in hydrophobic residues as the alpha-helices of the hairpin structures of StLDP and LDSP, as already observed in the literature for these structures (Tamm et al., 2004; Wimley, 2009). In the genome of Phytophthora (Oomycetes), some plastid derived genes were found, although Oomycetes do not have any plastids (Tyler etal. Let's get to know some of our stramenopiles. The energy excess produced by photosynthesis that still works during the first phases of stress is stocked and constitutes a pool of carbon and energy available upon stress release. PtDGAT2B is the only one that rescued the phenotype of the H1246 mutant (Gong et al., 2013; Zhang et al., 2020), had an in vitro activity (Zhang et al., 2020), and presented a considerable overexpression in nitrogen starvation (Gong et al., 2013; Zhang et al., 2020). The first analysis identified nine proteins, among which the most abundant is the Stramenopile-type lipid droplet protein (StLDP, accession number {"type":"entrez-protein","attrs":{"text":"XP_002183367","term_id":"219126235","term_text":"XP_002183367"}}XP_002183367), named based on the homology found in six other stramenopiles including M. gaditana (Yoneda et al., 2016). In both yeast and humans, the abovementioned interaction of seipin may determine the sites of LD formation (Bohnert, 2020). Despite being single-cell organisms, diatoms are a major part of food chains and are part of the collective groups of marine organisms we call plankton. Microalgal triacylglycerols as feedstocks for biofuel production: perspectives and advances. Hu Q., Sommerfeld M., Jarvis E., Ghirardi M., Posewitz M., Seibert M., et al. Homologs of AtSDP1 have been characterized in stramenopiles, named tgl1 in P. tricornutum (Barka et al., 2016), and NoTGL1 and NoTGL2 in N. oceanica (Nobusawa et al., 2019). In P. tricornutum genome, the locus Phatr3_J49708, encoding for a predicted protein ({"type":"entrez-nucleotide","attrs":{"text":"XM_002184438","term_id":"219128552","term_text":"XM_002184438"}}XM_002184438, protein ID: {"type":"entrez-protein","attrs":{"text":"XP_002184474","term_id":"219128553","term_text":"XP_002184474"}}XP_002184474), was first isolated from cDNA (Cui et al., 2013) and characterized by heterologous expression in the H1246 mutant and in vitro assay (Cui et al., 2013; Zhang et al., 2020).

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