{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T04:19:19Z","timestamp":1787026759890,"version":"build-2736575974"},"reference-count":80,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T00:00:00Z","timestamp":1779148800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:sec>\n                    <jats:title>State of the art<\/jats:title>\n                    <jats:p>Photosynthetic oxygenic microorganisms are recognized to be the functional core of Bioregenerative life support systems (BLSS) for sustaining long-term human presence in space. Among these organisms, microalgae are particularly promising due to their elevated growth rates, highly efficient light use, and ability to synthesize a plethora of valuable compounds. Their cultivation could lead to the sustainable production of food enriched in essential nutraceuticals while ensuring oxygen regeneration in confined environments. Understanding their responses to different light spectra is crucial for optimizing their cultivation in close systems and reducing the energetic costs. Considering the growing interest in the potential contribution of Far-Red (FR) light to the photosynthetic process and biomass production, we investigated the acclimations of different microalgal species under light spectra including this low-energy waveband in combination with visible (VIS) light.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Research methodologies<\/jats:title>\n                    <jats:p>\n                      Four phylogenetically diverse microalgal species, either capable (\n                      <jats:italic>Chromera velia<\/jats:italic>\n                      and\n                      <jats:italic>Nannochloropsis gaditana<\/jats:italic>\n                      ) or incapable (\n                      <jats:italic>Dixoniella grisea<\/jats:italic>\n                      and\n                      <jats:italic>Chlorella vulgaris<\/jats:italic>\n                      ) of utilizing solely FR light beside VIS radiation, were cultivated for 10\u202fdays under three light regimes: (i) simulated solar light (SOL); (ii) low-VIS, FR-enriched light (FR-e); (iii) FR light peaking at 730\u202fnm (FR). All treatments were characterized by the same total photon flux, but differed in spectral distribution and total available energy. The culture growth during acclimation to different lights was assessed. At the beginning and at the end of experiments, samples were investigated through optical microscopy, spectrophotometry and HPLC techniques. Changes in cell morphology and photosynthetic apparatus\n                      <jats:italic>in vivo<\/jats:italic>\n                      absorption of cultures were analyzed. Photosynthetic pigments were characterized and quantified.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      The results showed that microalgal growth under FR-e conditions was consistently higher than expected, being not proportional to the reduced fraction of available VIS photons (37.0% in FR-e vs. 84.3% in SOL). Specifically, growth under FR-e with respect to SOL, reached 61% in\n                      <jats:italic>C. vulgaris<\/jats:italic>\n                      , 82% in\n                      <jats:italic>D. grisea<\/jats:italic>\n                      , 91% in\n                      <jats:italic>N. gaditana<\/jats:italic>\n                      , and 93% in\n                      <jats:italic>C. velia<\/jats:italic>\n                      . Considering that the total energy of the FR-e spectrum is 83.5% of SOL (expressed in W m\n                      <jats:sup>\u22122<\/jats:sup>\n                      ) this light regime results particularly advantageous for the cultivation of FR-users microalgal species. The assessment of cell features and pigment composition showed similar characteristics of the produced biomass in SOL and FR-e for all tested species. Finally, the experiments also revealed a FR dose dependent photoacclimation in\n                      <jats:italic>C. velia<\/jats:italic>\n                      , whose FR light absorption capacity was enhanced at increasing FR doses.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key findings<\/jats:title>\n                    <jats:p>\n                      The results demonstrate that the growth of diverse microalgal species is differently influenced by a low-VIS FR-enriched spectrum. For FR-user species under this light regime the energy costs can be reduced for the efficient production of biomass, while maintaining a comparable quality. The most promising species turned out to be\n                      <jats:italic>N. gaditana<\/jats:italic>\n                      , due to its high growth rate and the capability to use FR light beside the VIS one. These findings are crucial for a sustainable microalgal cultivation in the frame of BLSS.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fmicb.2026.1833797","type":"journal-article","created":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T05:44:10Z","timestamp":1779169450000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Growth of phylogenetically diverse microalgae under Far-Red light enriched spectra: implication for space missions\u2019 sustainability"],"prefix":"10.3389","volume":"17","author":[{"given":"Beatrice","family":"Boccia","sequence":"first","affiliation":[{"name":"Department of Biology, University of Padua","place":["Padua, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mariano","family":"Battistuzzi","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Padua","place":["Padua, Italy"]},{"name":"National Institute for Astrophysics, Astrophysical Observatory of Arcetri (INAF-OAA)","place":["Florence, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elisabetta","family":"Liistro","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Padua","place":["Padua, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Riccardo","family":"Claudi","sequence":"additional","affiliation":[{"name":"National Institute for Astrophysics, Astronomical Observatory of Padua (INAF-OAPd)","place":["Padua, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lorenzo","family":"Cocola","sequence":"additional","affiliation":[{"name":"National Research Council \u2013 Institute for Photonics and Nanotechnologies (CNR\u2013IFN)","place":["Padua, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luca","family":"Poletto","sequence":"additional","affiliation":[{"name":"National Research Council \u2013 Institute for Photonics and Nanotechnologies (CNR\u2013IFN)","place":["Padua, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicoletta","family":"La Rocca","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Padua","place":["Padua, Italy"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,5,19]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"3539","DOI":"10.1007\/s10811-025-03569-y","article-title":"Physiological, biochemical, and morphological responses to nitrogen starvation and biomass-specific photon supply rates of Nannochloropsis oceanica and Microchloropsis gaditana","volume":"37","author":"Abdelkarim","year":"2025","journal-title":"J. Appl. Phycol."},{"key":"ref2","doi-asserted-by":"publisher","first-page":"2468","DOI":"10.1104\/pp.16.00599","article-title":"Light remodels lipid biosynthesis in Nannochloropsis gaditana by modulating carbon partitioning between organelles","volume":"171","author":"Alboresi","year":"2016","journal-title":"Plant Physiol."},{"key":"ref3","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/0022-5193(61)90026-1","article-title":"Methods for measuring and correcting the absorption spectrum of scattering suspensions","volume":"1","author":"Amesz","year":"1961","journal-title":"J. Theor. Biol."},{"key":"ref4","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1007\/s42452-025-06916-3","article-title":"Harnessing microalgae as a bioresource for nutraceuticals: advancing bioactive compound exploration and shaping the future of health and functional food innovation","volume":"7","author":"Ayub","year":"2025","journal-title":"Discov. Appl. Sci."},{"key":"ref5","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1007\/s12010-015-1751-7","article-title":"Effects of temperature and other operational parameters on Chlorella vulgaris mass cultivation in a simple and low-cost column photobioreactor","volume":"177","author":"Bamba","year":"2015","journal-title":"Appl. Biochem. Biotechnol."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"306","DOI":"10.1016\/j.bbabio.2013.11.019","article-title":"Characterization of the photosynthetic apparatus of the Eustigmatophycean Nannochloropsis gaditana: evidence of convergent evolution in the supramolecular organization of photosystem I","volume":"1837","author":"Basso","year":"2014","journal-title":"Biochim. Biophys. Acta"},{"key":"ref7","doi-asserted-by":"publisher","first-page":"1070359","DOI":"10.3389\/fpls.2023.1070359","article-title":"Oxygenic photosynthetic responses of cyanobacteria exposed under an M-dwarf starlight simulator: implications for exoplanet\u2019s habitability","volume":"14","author":"Battistuzzi","year":"","journal-title":"Front. Plant Sci."},{"key":"ref8","doi-asserted-by":"publisher","first-page":"1641","DOI":"10.3390\/life13081641","article-title":"Growth and photosynthetic efficiency of microalgae and plants with different levels of complexity exposed to a simulated M-dwarf starlight","volume":"13","author":"Battistuzzi","year":"","journal-title":"Life"},{"key":"ref9","doi-asserted-by":"publisher","first-page":"182","DOI":"10.3389\/fpls.2020.00182","article-title":"A new remote sensing-based system for the monitoring and analysis of growth and gas exchange rates of photosynthetic microorganisms under simulated non-terrestrial conditions","volume":"11","author":"Battistuzzi","year":"2020","journal-title":"Front. Plant Sci."},{"key":"ref10","doi-asserted-by":"publisher","first-page":"1322052","DOI":"10.3389\/fpls.2023.1322052","article-title":"Transcriptomic and photosynthetic analyses of Synechocystis sp. PCC6803 and Chlorogloeopsis fritschii sp. PCC6912 exposed to an M-dwarf spectrum under an anoxic atmosphere","volume":"14","author":"Battistuzzi","year":"2024","journal-title":"Front. Plant Sci."},{"key":"ref11","doi-asserted-by":"publisher","first-page":"952","DOI":"10.1111\/1462-2920.14774","article-title":"Life in the dark: far-red absorbing cyanobacteria extend photic zones deep into terrestrial caves","volume":"22","author":"Behrendt","year":"2020","journal-title":"Environ. Microbiol."},{"key":"ref12","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1016\/j.actaastro.2014.04.023","article-title":"Physicochemical and biological technologies for future exploration missions","volume":"101","author":"Belz","year":"2014","journal-title":"Acta Astronaut."},{"key":"ref9016","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1083\/jcb.58.2.419","article-title":"Complementary chromatic adaptation in a filamentous blue-green alga","volume":"58","author":"Bennett","year":"1973","journal-title":"J. Cell Biol."},{"key":"ref13","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1016\/j.lssr.2024.06.003","article-title":"Cultivation and nutritional characteristics of Chlorella vulgaris cultivated using Martian regolith and synthetic urine","volume":"42","author":"Casula","year":"2024","journal-title":"Life Sci. Space Res."},{"key":"ref14","doi-asserted-by":"publisher","first-page":"17348","DOI":"10.1016\/S0021-9258(19)85341-3","article-title":"Molecular and biochemical characterization of herbicide-resistant mutants of cyanobacteria reveals that phytoene desaturation is a rate-limiting step in carotenoid biosynthesis","volume":"268","author":"Chamovitz","year":"1993","journal-title":"J. Biol. Chem."},{"key":"ref15","doi-asserted-by":"publisher","first-page":"113486","DOI":"10.1016\/j.isci.2025.113486","article-title":"Previously unrecognized diversity of far-red light-using terrestrial green algae and cyanobacteria","volume":"28","author":"Chen","year":"2025","journal-title":"iScience"},{"key":"ref16","doi-asserted-by":"publisher","first-page":"5481","DOI":"10.1007\/s11033-023-08456-8","article-title":"The current situations and limitations of genetic engineering in cyanobacteria: a mini review","volume":"50","author":"Cheng","year":"2023","journal-title":"Mol. Biol. Rep."},{"key":"ref17","doi-asserted-by":"publisher","first-page":"10","DOI":"10.3390\/life11010010","article-title":"Super-earths, M dwarfs, and photosynthetic organisms: habitability in the lab","volume":"11","author":"Claudi","year":"2020","journal-title":"Life"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"733244","DOI":"10.3389\/fmicb.2021.733244","article-title":"Investigating the growth of algae under low atmospheric pressures for potential food and oxygen production on Mars","volume":"12","author":"Cycil","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref19","doi-asserted-by":"publisher","first-page":"700579","DOI":"10.3389\/fspas.2021.700579","article-title":"Chlorella Vulgaris photobioreactor for oxygen and food production on a moon base\u2014potential and challenges","volume":"8","author":"Detrell","year":"2021","journal-title":"Front. Astron. Space Sci."},{"key":"ref20","doi-asserted-by":"publisher","first-page":"22","DOI":"10.2216\/06-12.1","article-title":"Ecophysiological performance of the primitive red alga Dixoniella grisea (Rhodellophyceae) to irradiance, temperature and salinity stress: growth responses and the osmotic role of mannitol","volume":"46","author":"Eggert","year":"2007","journal-title":"Phycologia"},{"key":"ref9017","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1073\/pnas.43.1.133","article-title":"Some factors influencing the long-wave limit of photosynthesis","volume":"43","author":"Emerson","year":"1957","journal-title":"Procee National Acade Sci."},{"key":"ref21","doi-asserted-by":"publisher","first-page":"699525","DOI":"10.3389\/fmicb.2021.699525","article-title":"Use of photobioreactors in regenerative life support systems for human space exploration","volume":"12","author":"Fahrion","year":"2021","journal-title":"Front. Microbiol."},{"key":"ref22","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1038\/s41526-025-00550-4","article-title":"Superfood potential of Chlorella vulgaris: productivity and antioxidant boost under simulated moon and microgravity conditions","volume":"12","author":"Fais","year":"2025","journal-title":"NPJ Microgravity"},{"key":"ref23","doi-asserted-by":"publisher","first-page":"454","DOI":"10.1016\/j.actaastro.2022.09.058","article-title":"A novel process to grow edible microalgae on Mars by exploiting in situ-available resources: experimental investigation","volume":"201","author":"Fais","year":"2022","journal-title":"Acta Astronaut."},{"key":"ref24","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/S0005-2728(97)00093-5","article-title":"The xanthophyll cycle of higher plants: influence of antenna size and membrane organization","volume":"1363","author":"F\u00e4rber","year":"1998","journal-title":"Biochim. Biophys. Acta"},{"key":"ref25","doi-asserted-by":"publisher","first-page":"392","DOI":"10.1093\/pcp\/pch043","article-title":"On the 710 nm fluorescence emitted by the diatom Phaeodactylum tricornutum at room temperature","volume":"45","author":"Fujita","year":"2004","journal-title":"Plant Cell Physiol."},{"key":"ref26","doi-asserted-by":"publisher","first-page":"3450","DOI":"10.1111\/1462-2920.12992","article-title":"Adaptive and acclimative responses of cyanobacteria to far-red light: responses of cyanobacteria to far-red light","volume":"17","author":"Gan","year":"2015","journal-title":"Environ. Microbiol."},{"key":"ref27","doi-asserted-by":"publisher","first-page":"1312","DOI":"10.1126\/science.1256963","article-title":"Extensive remodeling of a cyanobacterial photosynthetic apparatus in far-red light","volume":"345","author":"Gan","year":"2014","journal-title":"Science"},{"key":"ref28","doi-asserted-by":"publisher","first-page":"1836","DOI":"10.3390\/plants10091836","article-title":"Potential of the red alga Dixoniella grisea for the production of additives for lubricants","volume":"10","author":"Gaval\u00e1s-Olea","year":"2021","journal-title":"Plants"},{"key":"ref29","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1089\/ast.2021.0184","article-title":"Absorption and scattering of UV and visible light through simulated Martian Regoliths and rock samples","volume":"23","author":"Godin","year":"2023","journal-title":"Astrobiology"},{"key":"ref30","doi-asserted-by":"publisher","first-page":"229","DOI":"10.1139\/m62-029","article-title":"Studies of marine planktonic diatoms. I. Cyclotella nana Hustedt, and Detonula confervacea (cleve) gran","volume":"8","author":"Guillard","year":"1962","journal-title":"Can. J. Microbiol."},{"key":"ref31","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1016\/j.lssr.2019.08.001","article-title":"High density long-term cultivation of Chlorella vulgaris SAG 211-12 in a novel microgravity-capable membrane raceway photobioreactor for future bioregenerative life support in SPACE","volume":"24","author":"Helisch","year":"2020","journal-title":"Life Sci. Space Res."},{"key":"ref32","doi-asserted-by":"publisher","first-page":"465","DOI":"10.3389\/fmicb.2019.00465","article-title":"Global transcriptional profiling of the cyanobacterium Chlorogloeopsis fritschii PCC 9212 in far-red light: insights into the regulation of chlorophyll d synthesis","volume":"10","author":"Ho","year":"2019","journal-title":"Front. Microbiol."},{"key":"ref33","doi-asserted-by":"publisher","first-page":"2936","DOI":"10.1111\/pce.14909","article-title":"Far-red light enrichment affects gene expression and architecture as well as growth and photosynthesis in rice","volume":"47","author":"Huber","year":"2024","journal-title":"Plant Cell Environ."},{"key":"ref34","article-title":"Phytoplankton pigments in oceanography: Guidelines to modern methods","volume-title":"UNESCO Monographs on Oceanographic Methodology","author":"Jeffrey","year":"1997"},{"key":"ref35","doi-asserted-by":"publisher","first-page":"604","DOI":"10.1016\/j.asr.2025.10.061","article-title":"Exploiting biocementation for Martian construction: pathways, challenges, and research gaps","volume":"77","author":"Khoshtinat","year":"2026","journal-title":"Adv. Space Res."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"734","DOI":"10.1016\/j.bbabio.2014.01.012","article-title":"Novel type of red-shifted chlorophyll a antenna complex from Chromera velia. I. Physiological relevance and functional connection to photosystems","volume":"1837","author":"Kotabov\u00e1","year":"2014","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref37","doi-asserted-by":"publisher","first-page":"651","DOI":"10.1002\/elsc.201400057","article-title":"Far-red light (720 or 740 nm) improves growth and changes the chemical composition of Chlorella vulgaris","volume":"14","author":"Kula","year":"2014","journal-title":"Eng. Life Sci."},{"key":"ref38","doi-asserted-by":"publisher","first-page":"102976","DOI":"10.1016\/j.algal.2023.102976","article-title":"Chlorella vulgaris acclimated cultivation under flashing light: an in-depth investigation under iso-actinic conditions","volume":"70","author":"Levasseur","year":"2023","journal-title":"Algal Res."},{"key":"ref39","doi-asserted-by":"publisher","first-page":"7405","DOI":"10.3390\/molecules26237405","article-title":"Supplementary far-red and blue lights influence the biomass and phytochemical profiles of two lettuce cultivars in plant factory","volume":"26","author":"Li","year":"2021","journal-title":"Molecules"},{"key":"ref40","doi-asserted-by":"publisher","first-page":"108793","DOI":"10.1016\/j.plaphy.2024.108793","article-title":"Synechococcus sp. PCC7335 responses to far-red enriched spectra and anoxic\/microoxic atmospheres: potential for astrobiotechnological applications","volume":"213","author":"Liistro","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref41","doi-asserted-by":"publisher","first-page":"1785","DOI":"10.1111\/nph.70786","article-title":"Thylakoids reorganization enables driving photosynthesis under far-red light in the microalga Nannochloropsis gaditana","volume":"249","author":"Liistro","year":"2026","journal-title":"New Phytol."},{"key":"ref42","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1007\/s11120-016-0234-1","article-title":"Architecture of the light-harvesting apparatus of the eustigmatophyte alga Nannochloropsis oceanica","volume":"130","author":"Litv\u00edn","year":"2016","journal-title":"Photosynth. Res."},{"key":"ref43","doi-asserted-by":"publisher","first-page":"1791","DOI":"10.1007\/s10811-017-1114-6","article-title":"The effect of environmental factors on fatty acid composition of Chromera velia (Chromeridae)","volume":"29","author":"Luke\u0161","year":"2017","journal-title":"J. Appl. Phycol."},{"key":"ref44","doi-asserted-by":"publisher","first-page":"690191","DOI":"10.3389\/fenvs.2021.690191","article-title":"Interaction effects of temperature, light, nutrients, and pH on growth and competition of Chlorella vulgaris and Anabaena sp. Strain PCC","volume":"9","author":"Meng","year":"2021","journal-title":"Front. Environ. Sci."},{"key":"ref45","doi-asserted-by":"publisher","first-page":"959","DOI":"10.1038\/nature06635","article-title":"A photosynthetic alveolate closely related to apicomplexan parasites","volume":"451","author":"Moore","year":"2008","journal-title":"Nature"},{"key":"ref46","doi-asserted-by":"publisher","first-page":"8","DOI":"10.1016\/j.lssr.2017.10.002","article-title":"Spacecraft cabin environment effects on the growth and behavior of Chlorella vulgaris for life support applications","volume":"16","author":"Niederwieser","year":"2018","journal-title":"Life Sci. Space Res."},{"key":"ref47","doi-asserted-by":"publisher","first-page":"2368","DOI":"10.1038\/ismej.2017.98","article-title":"A niche for cyanobacteria producing chlorophyll f within a microbial mat","volume":"11","author":"Ohkubo","year":"2017","journal-title":"ISME J."},{"key":"ref48","first-page":"53","article-title":"Light Harvesting Complexes and chromatic Adaptation of Eustigmatophyte alga Trachydiscus minutus","volume-title":"Mgr. Thesis","author":"Pazdern\u00edk","year":"2015"},{"key":"ref49","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1088\/0067-0049\/208\/1\/9","article-title":"Intrinsic colors, temperatures, and bolometric corrections of pre-main-sequence stars","volume":"208","author":"Pecaut","year":"2013","journal-title":"Astrophys. J. Suppl. Ser."},{"key":"ref50","doi-asserted-by":"publisher","first-page":"384","DOI":"10.1016\/S0005-2728(89)80347-0","article-title":"Determination of accurate extinction coefficients and simultaneous equations for assaying chlorophylls a and b extracted with four different solvents: verification of the concentration of chlorophyll standards by atomic absorption spectroscopy","volume":"975","author":"Porra","year":"1989","journal-title":"Biochim. Biophys. Acta"},{"key":"ref51","doi-asserted-by":"publisher","first-page":"e47036","DOI":"10.1371\/journal.pone.0047036","article-title":"Photosynthesis in Chromera velia represents a simple system with high efficiency","volume":"7","author":"Quigg","year":"2012","journal-title":"PLoS One"},{"key":"ref52","doi-asserted-by":"publisher","first-page":"103332","DOI":"10.1016\/j.algal.2023.103332","article-title":"Microalgae in bioregenerative life support systems for space applications","volume":"77","author":"Revellame","year":"2024","journal-title":"Algal Res."},{"key":"ref53","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1099\/00221287-111-1-1","article-title":"Generic assignments, strain histories and properties of pure cultures of cyanobacteria","volume":"111","author":"Rippka","year":"1979","journal-title":"Microbiology"},{"key":"ref54","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1016\/S1389-0344(03)00061-3","article-title":"Growth aspects of the marine microalga Nannochloropsis gaditana","volume":"20","author":"Rocha","year":"2003","journal-title":"Biomol. Eng."},{"key":"ref55","doi-asserted-by":"publisher","first-page":"977429","DOI":"10.3389\/fbioe.2022.977429","article-title":"Microalgae growth in ultra-thin steady-state continuous photobioreactors: assessing self-shading effects","volume":"10","author":"Saccardo","year":"2022","journal-title":"Front. Bioeng. Biotechnol."},{"key":"ref56","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1016\/j.rser.2014.04.007","article-title":"Morphology, composition, production, processing and applications of Chlorella vulgaris: a review","volume":"35","author":"Safi","year":"2014","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref57","doi-asserted-by":"publisher","first-page":"419","DOI":"10.1034\/j.1399-3054.1996.980226.x","article-title":"Far-red light inhibits growth but promotes carotenoid accumulation in the green microalga Dunaliella bardawil","volume":"98","author":"S\u00e1nchez-Saavedra","year":"1996","journal-title":"Physiol. Plant."},{"key":"ref58","doi-asserted-by":"publisher","first-page":"e0171872","DOI":"10.1371\/journal.pone.0171872","article-title":"Community analysis of pigment patterns from 37 microalgae strains reveals new carotenoids and porphyrins characteristic of distinct strains and taxonomic groups","volume":"12","author":"Serive","year":"2017","journal-title":"PLoS One"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1002\/9780470110232.ch3","volume-title":"Methods of Biochemical Analysis","author":"Shibata","year":"1959"},{"key":"ref60","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.physrep.2016.10.003","article-title":"The habitability of planets orbiting M-dwarf stars","volume":"663","author":"Shields","year":"2016","journal-title":"Phys. Rep."},{"key":"ref61","doi-asserted-by":"publisher","first-page":"170","DOI":"10.3390\/metabo9080170","article-title":"Far-red light acclimation for improved mass cultivation of cyanobacteria","volume":"9","author":"Silkina","year":"2019","journal-title":"Meta"},{"key":"ref62","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1016\/0273-1177(84)90569-6","article-title":"BLSS: a contribution to future life support","volume":"4","author":"Skoog","year":"1984","journal-title":"Adv. Space Res."},{"key":"ref63","volume-title":"NMR Structure of Chromeraxanthin\u2014a Unique Allenic Carotenoid of the Apicomplexan Alga","author":"Takacs","year":"2025"},{"key":"ref64","doi-asserted-by":"publisher","first-page":"1067782","DOI":"10.3389\/fmicb.2022.1067782","article-title":"Microalgae production in human urine: fundamentals, opportunities, and perspectives","volume":"13","author":"Tao","year":"2022","journal-title":"Front. Microbiol."},{"key":"ref65","doi-asserted-by":"publisher","first-page":"440","DOI":"10.3390\/md21080440","article-title":"Microalgae: a promising source of bioactive phycobiliproteins","volume":"21","author":"Tounsi","year":"2023","journal-title":"Mar. Drugs"},{"key":"ref66","doi-asserted-by":"publisher","first-page":"58","DOI":"10.1080\/09670262.2017.1376255","article-title":"Life at elevated CO2 modifies the cell composition of Chromera velia (Chromerida)","volume":"53","author":"Venuleo","year":"2018","journal-title":"Eur. J. Phycol."},{"key":"ref67","doi-asserted-by":"publisher","first-page":"977364","DOI":"10.3389\/fspas.2022.977364","article-title":"Editorial: bioregenerative life-support systems for crewed missions to the moon and Mars","volume":"9","author":"Verseux","year":"2022","journal-title":"Front. Astron. Space Sci."},{"key":"ref68","doi-asserted-by":"publisher","first-page":"114658","DOI":"10.1016\/j.icarus.2021.114658","article-title":"The ultraviolet radiation environment and shielding in pit craters and cave skylights on Mars","volume":"370","author":"Vi\u00fadez-Moreiras","year":"2021","journal-title":"Icarus"},{"key":"ref69","doi-asserted-by":"publisher","first-page":"8231","DOI":"10.1111\/1750-3841.17529","article-title":"Chlorella vulgaris as a food substitute: applications and benefits in the food industry","volume":"89","author":"Wang","year":"2024","journal-title":"J. Food Sci."},{"key":"ref70","doi-asserted-by":"publisher","first-page":"316","DOI":"10.3389\/fmicb.2012.00316","article-title":"Light gradients and optical microniches in coral tissues","volume":"3","author":"Wangpraseurt","year":"2012","journal-title":"Front. Microbiol."},{"key":"ref71","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1016\/S0176-1617(11)81192-2","article-title":"The spectral determination of chlorophylls a and b, as well as total carotenoids, using various solvents with spectrophotometers of different resolution","volume":"144","author":"Wellburn","year":"1994","journal-title":"J. Plant Physiol."},{"key":"ref72","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1007\/s11120-005-9002-3","article-title":"Uphill energy transfer from long-wavelength absorbing chlorophylls to PS II in Ostreobium sp. is functional in carbon assimilation","volume":"87","author":"Wilhelm","year":"2006","journal-title":"Photosynth. Res."},{"key":"ref73","doi-asserted-by":"publisher","first-page":"015005","DOI":"10.1088\/1758-5090\/ae1e32","article-title":"Regolith-supported biofabrication: 3D bioprinting of living, photosynthetically active structures using lunar and Martian regolith simulants","volume":"18","author":"Windisch","year":"2026","journal-title":"Biofabrication"},{"key":"ref74","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1007\/s11120-019-00653-6","article-title":"Far-red light acclimation in diverse oxygenic photosynthetic organisms","volume":"142","author":"Wolf","year":"2019","journal-title":"Photosynth. Res."},{"key":"ref75","doi-asserted-by":"publisher","first-page":"183","DOI":"10.3354\/meps077183","article-title":"Improved HPLC method for the analysis of chlorophylls and carotenoids from marine phytoplankton","volume":"77","author":"Wright","year":"1991","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"ref76","doi-asserted-by":"publisher","first-page":"531","DOI":"10.3390\/stresses2040036","article-title":"Mechanisms of stress tolerance in cyanobacteria under extreme conditions","volume":"2","author":"Yadav","year":"2022","journal-title":"Stress"},{"key":"ref77","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1007\/978-90-481-3795-4_2","volume-title":"Red Algae in the Genomic Age","author":"Yoon","year":"2010"},{"key":"ref9018","doi-asserted-by":"crossref","first-page":"538","DOI":"10.1111\/nph.18375","article-title":"Photosynthesis in sun and shade: the surprising importance of far\u2010red photons","volume":"236","author":"Zhen","year":"2022","journal-title":"New Phytologist."}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1833797\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,19]],"date-time":"2026-05-19T05:44:22Z","timestamp":1779169462000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1833797\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,19]]},"references-count":80,"alternative-id":["10.3389\/fmicb.2026.1833797"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2026.1833797","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,19]]},"article-number":"1833797"}}