{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,3]],"date-time":"2026-06-03T11:32:45Z","timestamp":1780486365553,"version":"3.54.1"},"reference-count":44,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2021,4,13]],"date-time":"2021-04-13T00:00:00Z","timestamp":1618272000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001501","name":"University Grants Commission","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001501","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>Microalgae are used as a source of lipids for the production of biofuels. Most algae produce neutral lipids under stress conditions. Here, lipid accumulation by the unicellular alga <jats:italic>Chlamydomonas reinhardtii<\/jats:italic> was examined during cultivation under iron-limiting conditions. Severe iron stress caused the cells to accumulate a significant amount of lipid, specifically triacylglycerols (TAGs), by compromising the growth. Semi-quantitative measurements by Fourier transform infrared (FTIR) spectroscopy showed an increase in both carbohydrate and lipid content in iron-stressed <jats:italic>C. reinhardtii<\/jats:italic> cells compared to control. Analysis by flow cytometry and thin layer chromatography confirmed that severe iron deficiency-induced TAG accumulation to fourfold higher than in iron-replete control cells. This accumulation of TAGs was mostly degraded from chloroplast lipids accompanied by overexpression of diacylglycerol acyltransferase (DGAT2A) protein. Furthermore, liquid chromatography-mass spectrometry (LC-MS) analysis demonstrated significantly enhanced levels of C16:0, C18:2, and C18:3 fatty acids (FAs). These results indicate that iron stress triggers the rapid accumulation of TAGs in <jats:italic>C. reinhardtii<\/jats:italic> cells. The enhanced production of these lipids caused by the iron deficiency may contribute to the efficient production of algal biofuels if we escalate to the photobioreactor\u2019s growth conditions.<\/jats:p>","DOI":"10.3389\/fpls.2021.615577","type":"journal-article","created":{"date-parts":[[2021,4,13]],"date-time":"2021-04-13T06:36:15Z","timestamp":1618295775000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":31,"title":["Enhanced Lipid Production in Chlamydomonas reinhardtii Caused by Severe Iron Deficiency"],"prefix":"10.3389","volume":"12","author":[{"given":"Elsinraju","family":"Devadasu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rajagopal","family":"Subramanyam","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2021,4,13]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"316","DOI":"10.1016\/j.biortech.2018.09.140","article-title":"Sustainable production of biomass and biodiesel by acclimation of non-acidophilic microalgae to acidic conditions.","volume":"271","author":"Abinandan","year":"2019","journal-title":"Bioresour. Technol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"911","DOI":"10.1139\/y59-099","article-title":"A rapid method of total lipid extraction and purification.","volume":"37","author":"Bligh","year":"1959","journal-title":"Can. J. Biochem. Physiol."},{"key":"B3","first-page":"19","author":"Bozzola","year":"1998","journal-title":"Electron Microscopy: Principles and Techniques for Biologists"},{"key":"B4","doi-asserted-by":"publisher","first-page":"1947","DOI":"10.1002\/bit.24474","article-title":"Differential effects of nitrogen and sulfur deprivation on growth and biodiesel feedstock production of Chlamydomonas reinhardtii.","volume":"109","author":"Cakmak","year":"2012","journal-title":"Biotechnol. Bioeng."},{"key":"B5","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1016\/j.biortech.2013.12.093","article-title":"Induction of triacylglycerol production in Chlamydomonas reinhardtii: comparative analysis of different element regimes.","volume":"155","author":"\u00c7akmak","year":"2014","journal-title":"Bioresour. Technol."},{"key":"B6","doi-asserted-by":"publisher","first-page":"1649","DOI":"10.1016\/j.biortech.2010.09.062","article-title":"Effect of nutrients on growth and lipid accumulation in the green algae Dunaliella tertiolecta.","volume":"102","author":"Chen","year":"2011","journal-title":"Bioresour. Technol."},{"key":"B7","doi-asserted-by":"publisher","first-page":"1355","DOI":"10.1093\/jxb\/erx372","article-title":"Autophagic flux is required for the synthesis of triacylglycerols and ribosomal protein turnover in Chlamydomonas.","volume":"69","author":"Couso","year":"2018","journal-title":"J. Exp. Bot."},{"key":"B8","doi-asserted-by":"publisher","first-page":"4499","DOI":"10.1016\/j.biortech.2010.01.065","article-title":"Using FTIR spectroscopy for rapid determination of lipid accumulation in response to nitrogen limitation in freshwater microalgae.","volume":"101","author":"Dean","year":"2010","journal-title":"Bioresour. Technol."},{"key":"B9","first-page":"260","article-title":"The effects of nutritional restriction on neutral lipid accumulation in Chlamydomonas and Chlorella.","volume":"5","author":"Deng","year":"2011","journal-title":"Afr. J. Microbiol. Res."},{"key":"B10","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1007\/s11120-018-0580-2","article-title":"Changes in the photosynthetic apparatus and lipid droplet formation in Chlamydomonas reinhardtii under iron deficiency.","volume":"139","author":"Devadasu","year":"2019","journal-title":"Photosynth. Res."},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbabio.2020.148331","article-title":"Restoration of photosynthetic activity and supercomplexes from severe iron starvation in Chlamydomonas reinhardtii.","volume":"1862","author":"Devadasu","year":"2021","journal-title":"Biochim. Biophys. Acta"},{"key":"B12","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1007\/s11120-016-0284-4","article-title":"Iron deficiency cause changes in photochemistry, thylakoid organization, and accumulation of photosystem II proteins in Chlamydomonas reinhardtii.","volume":"130","author":"Devadasu","year":"2016","journal-title":"Photosynth. Res."},{"key":"B13","doi-asserted-by":"publisher","first-page":"302","DOI":"10.1016\/j.jbiosc.2017.04.009","article-title":"Acclimation to NaCl and light stress of heterotrophic Chlamydomonas reinhardtii for lipid accumulation.","volume":"124","author":"Fan","year":"2017","journal-title":"J. Biosci. Bioeng."},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2013.00337","article-title":"Iron economy in Chlamydomonas reinhardtii.","volume":"4","author":"Glaesener","year":"2013","journal-title":"Front. Plant Sci."},{"key":"B15","doi-asserted-by":"publisher","first-page":"1887","DOI":"10.1080\/09593330.2013.812668","article-title":"Effects of calcium, magnesium and sodium chloride in enhancing lipid accumulation in two green microalgae.","volume":"34","author":"Gorain","year":"2013","journal-title":"Environ. Technol."},{"key":"B16","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.ecoenv.2017.05.048","article-title":"Sensitivity of two green microalgae to copper stress: growth, oxidative and antioxidants analyses.","volume":"144","author":"Hamed","year":"2017","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"B17","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1146\/annurev-arplant-050312-120053","article-title":"Endoplasmic reticulum stress responses in plants.","volume":"64","author":"Howell","year":"2013","journal-title":"Annu. Rev. Plant Biol."},{"key":"B18","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1007\/s10811-012-9846-9","article-title":"Effects of nitrogen supplementation of the culture medium on the growth, total lipid content and fatty acid profiles of three microalgae (Tetraselmis subcordiformis, Nannochloropsis oculata and Pavlova viridis).","volume":"25","author":"Huang","year":"2013","journal-title":"J. Appl. Phycol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"3343","DOI":"10.1016\/j.biortech.2010.11.051","article-title":"Fatty acid profiling of Chlamydomonas reinhardtii under nitrogen deprivation.","volume":"102","author":"James","year":"2011","journal-title":"Bioresour. Technol."},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0182423","article-title":"LC-MS\/MS versus TLC plus GC methods: consistency of glycerolipid and fatty acid profiles in microalgae and higher plant cells and effect of a nitrogen starvation.","volume":"12","author":"Jouhet","year":"2017","journal-title":"PLoS One"},{"key":"B21","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1046\/j.1365-2427.1997.00231.x","article-title":"Effects of nutrient limitation on biochemical constituents of Ankistrodesmus falcatus.","volume":"38","author":"Kilham","year":"1997","journal-title":"Freshw. Biol."},{"key":"B22","doi-asserted-by":"publisher","first-page":"770","DOI":"10.1111\/j.1365-313x.2011.04537.x","article-title":"A revised mineral nutrient supplement increases biomass and growth rate in Chlamydomonas reinhardtii.","volume":"66","author":"Kropat","year":"2011","journal-title":"Plant J."},{"key":"B23","doi-asserted-by":"publisher","first-page":"4670","DOI":"10.1105\/tpc.112.105106","article-title":"A galactoglycerolipid lipase is required for triacylglycerol accumulation and survival following nitrogen deprivation in Chlamydomonas reinhardtii.","volume":"24","author":"Li","year":"2012","journal-title":"Plant Cell"},{"key":"B24","doi-asserted-by":"publisher","first-page":"1213","DOI":"10.1007\/s00449-018-1950-z","article-title":"Effects of fundamental nutrient stresses on the lipid accumulation profiles in two diatom species Thalassiosira weissflogii and Chaetoceros muelleri.","volume":"41","author":"Lin","year":"2018","journal-title":"Bioprocess Biosyst. Eng."},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0103389","article-title":"Lipid profile remodeling in response to nitrogen deprivation in the microalgae Chlorella sp. (Trebouxiophyceae) and Nannochloropsis sp. (Eustigmatophyceae).","volume":"9","author":"Martin","year":"2014","journal-title":"PLoS One"},{"key":"B26","doi-asserted-by":"publisher","first-page":"2854","DOI":"10.1093\/jxb\/eraa082","article-title":"Reserve lipids and plant autophagy.","volume":"71","author":"Masclaux-Daubresse","year":"2020","journal-title":"J. Exp. Bot."},{"key":"B27","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1126\/science.1143609","article-title":"The Chlamydomonas genome reveals the evolution of key animal and plant functions.","volume":"318","author":"Merchant","year":"2007","journal-title":"Science"},{"key":"B28","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1128\/ec.00203-09","article-title":"RNA interference silencing of a major lipid droplet protein affects lipid droplet size in Chlamydomonas reinhardtii.","volume":"9","author":"Moellering","year":"2010","journal-title":"Eukaryot Cell"},{"key":"B29","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1016\/j.algal.2016.03.001","article-title":"Effect of temperature and nitrogen concentration on lipid productivity and fatty acid composition in three Chlorella strains.","volume":"16","author":"\u00d6rd\u00f6g","year":"2016","journal-title":"Algal Res."},{"key":"B30","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1111\/tpj.12747","article-title":"The response of Chlamydomonas reinhardtii to nitrogen deprivation: a systems biology analysis.","volume":"81","author":"Park","year":"2015","journal-title":"Plant J."},{"key":"B31","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1007\/s00248-003-0144-6","article-title":"Fatty acid patterns in Chlamydomonas sp. as a marker for nutritional regimes and temperature under extremely acidic conditions.","volume":"48","author":"Poerschmann","year":"2004","journal-title":"Microb. Ecol."},{"key":"B32","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1007\/s12010-014-1283-6","article-title":"Nitrogen starvation for lipid accumulation in the microalga species Desmodesmus sp.","volume":"175","author":"Rios","year":"2015","journal-title":"Appl. Biochem. Biotechnol."},{"key":"B33","doi-asserted-by":"publisher","first-page":"1532","DOI":"10.3390\/en5051532","article-title":"High lipid induction in microalgae for biodiesel production.","volume":"5","author":"Sharma","year":"2012","journal-title":"Energies"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1186\/1472-6750-11-7","article-title":"Oil accumulation in the model green alga Chlamydomonas reinhardtii: characterization, variability between common laboratory strains and relationship with starch reserves.","volume":"11","author":"Siaut","year":"2011","journal-title":"BMC Biotechnol."},{"key":"B35","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1016\/j.pbi.2005.01.012","article-title":"Metabolic engineering of new fatty acids in plants.","volume":"8","author":"Singh","year":"2005","journal-title":"Curr. Opin. Plant Biol."},{"key":"B36","doi-asserted-by":"crossref","DOI":"10.1186\/s13068-016-0671-2","article-title":"The dilemma for lipid productivity in green microalgae: importance of substrate provision in improving oil yield without sacrificing growth.","volume":"9","author":"Tan","year":"2016","journal-title":"Biotechnol. Biofuels"},{"key":"B37","doi-asserted-by":"publisher","first-page":"4350","DOI":"10.1073\/pnas.76.9.4350","article-title":"Electrophoretic transfer of proteins from polyacrylamide gels to nitrocellulose sheets: procedure and some applications.","volume":"76","author":"Towbin","year":"1979","journal-title":"Proc. Natl. Acad. Sci. U.S.A."},{"key":"B38","doi-asserted-by":"publisher","first-page":"30246","DOI":"10.1074\/jbc.m113.490425","article-title":"Remodeling of membrane lipids in iron-starved Chlamydomonas.","volume":"288","author":"Urzica","year":"2013","journal-title":"J. Biol. Chem."},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1186\/preaccept-1279724404120676","article-title":"Systematically programmed adaptive evolution reveals potential role of carbon and nitrogen pathways during lipid accumulation in Chlamydomonas reinhardtii.","volume":"7","author":"Velmurugan","year":"2014","journal-title":"Biotechnol. Biofuels"},{"key":"B40","doi-asserted-by":"publisher","first-page":"544","DOI":"10.1128\/jb.175.2.544-547.1993","article-title":"In vitro ferredoxin- dependent desaturation of fatty acids in cyanobacterial thylakoid membranes.","volume":"175","author":"Wada","year":"1993","journal-title":"J. Bacteriol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.algal.2015.06.002","article-title":"Phenotypicy screening identifies Brefeldin A\/Ascotoxin as an inducer of lipid storage in the algae Chlamydomonas reinhardtii. Algal Res.","author":"Wase","year":"2015"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0035084","article-title":"Alteration of proteins and pigments influence the function of photosystem I under iron deficiency from Chlamydomonas reinhardtii.","volume":"7","author":"Yadavalli","year":"2012","journal-title":"PLoS One"},{"key":"B43","doi-asserted-by":"publisher","first-page":"1478","DOI":"10.1016\/j.bbalip.2018.09.009","article-title":"Triacylglycerol accumulates exclusively outside the chloroplast in short-term nitrogen-deprived Chlamydomonas reinhardtii.","volume":"1863","author":"Yang","year":"2018","journal-title":"Biochim. Biophys. Acta Mol. Cell Biol. Lipids"},{"key":"B44","doi-asserted-by":"publisher","first-page":"3885","DOI":"10.1105\/tpc.109.071795","article-title":"Acyltransferases have overlapping functions in Arabidopsis triacylglycerol biosynthesis and are essential for normal pollen and seed development.","volume":"21","author":"Zhang","year":"2009","journal-title":"Plant Cell"}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2021.615577\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,13]],"date-time":"2021-04-13T06:36:20Z","timestamp":1618295780000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2021.615577\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,4,13]]},"references-count":44,"alternative-id":["10.3389\/fpls.2021.615577"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2021.615577","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,4,13]]},"article-number":"615577"}}