{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T13:02:46Z","timestamp":1790254966925,"version":"4.1.0"},"reference-count":38,"publisher":"American Chemical Society (ACS)","issue":"38","license":[{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP23H02423"],"award-info":[{"award-number":["JP23H02423"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP24H02075"],"award-info":[{"award-number":["JP24H02075"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP24H02086"],"award-info":[{"award-number":["JP24H02086"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP26H01085"],"award-info":[{"award-number":["JP26H01085"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP26H01094"],"award-info":[{"award-number":["JP26H01094"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,24]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Dark-heterotrophic growth allows photosynthetic pigment\u2013protein complexes to be examined when photosynthetic energy conversion is not required for growth. A dark-adapted variant, dg211, of Leptolyngbya boryana has been reported to exhibit altered carotenoid composition, but its effect on excitation-energy transfer remains unclear. Here, we analyzed light-grown wild-type (WT-light) and dg211 (dg211-light), and dark-grown dg211 (dg211-dark) cells using immunoblotting, pigment analysis, absorption spectroscopy, 77 K steady-state and time-resolved fluorescence spectroscopy, and fluorescence decay-associated spectra. \u03b2-Carotene was detected in the WT-light and dg211-light cells but not in the dg211-dark cells, although chlorophyll a and phycobilisome-related spectral features were retained. In the dg211-dark cells, D1 accumulation was markedly reduced, whereas PsaA\/B remained detectable. PSII-delayed fluorescence disappeared, while PSI-associated 729\u2013730 nm fluorescence persisted, indicating a severe depletion of functional PSII reaction-center units with retention of PSI-associated features. Pronounced 687 nm nanosecond fluorescence was attributed mainly to prolonged excitation-energy retention in phycobilisome-related emitting states. These findings reveal substantial reorganization of photosynthetic excitation-energy transfer in dark-grown dg211.<\/jats:p>","DOI":"10.1021\/acs.jpcb.6c03891","type":"journal-article","created":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T09:02:14Z","timestamp":1788944534000},"page":"9843-9850","source":"Crossref","is-referenced-by-count":0,"title":["Altered Excitation-Energy\nTransfer in the Dark-Adapted\n                    <i>Leptolyngbya boryana<\/i>\n                    Variant dg211"],"prefix":"10.1021","volume":"130","author":[{"given":"Kano N.","family":"Toyoizumi","sequence":"first","affiliation":[{"name":"Shizuoka University , , ,","place":["Shizuoka, Japan, 422-8529"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuichi","family":"Fujita","sequence":"additional","affiliation":[{"name":"Nagoya University , , ,","place":["Aichi, Japan, 464-8601"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8951-8978","authenticated-orcid":true,"given":"Seiji","family":"Akimoto","sequence":"additional","affiliation":[{"name":"Kobe University , , ,","place":["Hyogo, Japan, 657-8501"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8212-3001","authenticated-orcid":true,"given":"Ryo","family":"Nagao","sequence":"additional","affiliation":[{"name":"Shizuoka University , , ,","place":["Shizuoka, Japan, 422-8529"]},{"name":"Shizuoka University , , ,","place":["Shizuoka, Japan, 422-8529"]},{"name":"Shizuoka University , , ,","place":["Shizuoka, Japan, 422-8529"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,9,9]]},"reference":[{"key":"2026092408094438100_cit1","volume-title":"Molecular\nMechanisms of Photosynthesis","author":"Blankenship","year":"2021","edition":"3rd"},{"key":"2026092408094438100_cit2","doi-asserted-by":"publisher","first-page":"33098","DOI":"10.1021\/acsomega.3c02205","article-title":"Heterotrophy among Cyanobacteria","volume":"8","author":"Stebegg","year":"2023","journal-title":"ACS Omega"},{"key":"2026092408094438100_cit3","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1099\/00221287-111-1-1","article-title":"Generic\nassignments, strain histories and properties of pure cultures of cyanobacteria","volume":"111","author":"Rippka","year":"1979","journal-title":"J. Gen. Microbiol."},{"key":"2026092408094438100_cit4","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1093\/oxfordjournals.pcp.a078224","article-title":"The nifH-like (frxC) gene is involved in the\nbiosynthesis of chlorophyll in the filamentous cyanobacterium Plectonema boryanum","volume":"33","author":"Fujita","year":"1992","journal-title":"Plant Cell Physiol."},{"key":"2026092408094438100_cit5","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1111\/j.1574-6968.1998.tb12959.x","article-title":"Obligate phototrophy\nin cyanobacteria:\nmore than a lack of sugar transport","volume":"161","author":"Zhang","year":"1998","journal-title":"FEMS Microbiol.\nLett."},{"key":"2026092408094438100_cit6","doi-asserted-by":"publisher","first-page":"334","DOI":"10.1093\/pcp\/pcu165","article-title":"Loss of cytochrome  cM stimulates\ncyanobacterial heterotrophic growth in the dark","volume":"56","author":"Hiraide","year":"2015","journal-title":"Plant Cell Physiol."},{"key":"2026092408094438100_cit7","doi-asserted-by":"publisher","DOI":"10.1101\/2024.04.08.588626","article-title":"Microevolution toward loss of photosynthesis:\nmutations promoting dark-heterotrophic growth and suppressing photosynthetic\ngrowth in cyanobacteria.","author":"Hida","journal-title":"bioRxiv"},{"key":"2026092408094438100_cit8","doi-asserted-by":"publisher","first-page":"1102","DOI":"10.1093\/pcp\/pcaf043","article-title":"Genome analysis of\ndark-adapted variants identifies the phosphatase gene phsP involved in the regulation of photosynthetic and dark-heterotrophic\ngrowth in the cyanobacterium Leptolyngbya boryana","volume":"66","author":"Hida","year":"2025","journal-title":"Plant Cell Physiol."},{"key":"2026092408094438100_cit9","doi-asserted-by":"publisher","first-page":"1558","DOI":"10.1104\/pp.125.4.1558","article-title":"Non-photochemical\nquenching. A response\nto excess light energy","volume":"125","author":"Mu\u0308ller","year":"2001","journal-title":"Plant Physiol."},{"key":"2026092408094438100_cit10","doi-asserted-by":"publisher","first-page":"2021","DOI":"10.1021\/cr020674n","article-title":"Ultrafast\ndynamics of carotenoid excited states\u2013From\nsolution to natural and artificial systems","volume":"104","author":"Poli\u0301vka","year":"2004","journal-title":"Chem. Rev."},{"key":"2026092408094438100_cit11","doi-asserted-by":"publisher","first-page":"1101","DOI":"10.3390\/md9061101","article-title":"Carotenoids\nin algae: distributions, biosyntheses and functions","volume":"9","author":"Takaichi","year":"2011","journal-title":"Mar. Drugs"},{"key":"2026092408094438100_cit12","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1016\/S0005-2728(01)00195-5","article-title":"Structure of photosystem I","volume":"1507","author":"Fromme","year":"2001","journal-title":"Biochim. Biophys. 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PCC 6803","volume":"51","author":"Sozer","year":"2010","journal-title":"Plant Cell Physiol."},{"key":"2026092408094438100_cit15","doi-asserted-by":"publisher","first-page":"1153","DOI":"10.1016\/j.bbabio.2015.05.020","article-title":"Carotenoids\nare essential for the assembly of cyanobacterial photosynthetic complexes","volume":"1847","author":"To\u0301th","year":"2015","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit16","doi-asserted-by":"publisher","first-page":"1093","DOI":"10.1093\/oxfordjournals.pcp.a078173","article-title":"Cloning, nucleotide\nsequences and differential expression of the nifH and nifH-like (frxC) genes from\nthe filamentous nitrogen-fixing cyanobacterium Plectonema boryanum","volume":"32","author":"Fujita","year":"1991","journal-title":"Plant Cell Physiol."},{"key":"2026092408094438100_cit17","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1128\/br.35.2.171-205.1971","article-title":"Purification\nand properties of unicellular blue-green algae (order Chroococcales)","volume":"35","author":"Stanier","year":"1971","journal-title":"Bacteriol. Rev."},{"key":"2026092408094438100_cit18","doi-asserted-by":"publisher","first-page":"7474","DOI":"10.1074\/jbc.M117.781062","article-title":"Genetically introduced hydrogen bond interactions reveal\nan asymmetric charge distribution on the radical cation of the special-pair\nchlorophyll P680","volume":"292","author":"Nagao","year":"2017","journal-title":"J. Biol. Chem."},{"key":"2026092408094438100_cit19","doi-asserted-by":"publisher","first-page":"148509","DOI":"10.1016\/j.bbabio.2021.148509","article-title":"Excitation-energy\ntransfer in heterocysts\nisolated from the cyanobacterium Anabaena sp. PCC\n7120 as studied by time-resolved fluorescence spectroscopy","volume":"1863","author":"Nagao","year":"2022","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit20","doi-asserted-by":"publisher","first-page":"55","DOI":"10.1007\/s11120-023-01025-x","article-title":"Isolation\nand characterization of\ntrimeric and monomeric PSI cores from Acaryochloris marina MBIC11017","volume":"157","author":"Nagao","year":"2023","journal-title":"Photosynth. Res."},{"key":"2026092408094438100_cit21","doi-asserted-by":"publisher","first-page":"99","DOI":"10.1016\/0014-5793(88)81039-1","article-title":"A new photosystem II\nreaction center component (4.8 kDa protein)\nencoded by chloroplast genome","volume":"241","author":"Ikeuchi","year":"1988","journal-title":"FEBS Lett."},{"key":"2026092408094438100_cit22","doi-asserted-by":"publisher","first-page":"148327","DOI":"10.1016\/j.bbabio.2020.148327","article-title":"Molecular organizations\nand function of iron-stress-induced-A protein family in Anabaena sp. PCC 7120","volume":"1862","author":"Nagao","year":"2021","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit23","doi-asserted-by":"publisher","first-page":"1353","DOI":"10.1016\/j.bbabio.2007.10.007","article-title":"Isolation and characterization\nof\noxygen-evolving thylakoid membranes and Photosystem II particles from\na marine diatom Chaetoceros gracilis","volume":"1767","author":"Nagao","year":"2007","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit24","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1016\/j.jim.2006.07.027","article-title":"A cost effective non-commercial\nECL-solution for Western\nblot detections yielding strong signals and low background","volume":"318","author":"Haan","year":"2007","journal-title":"J. Immunol. Methods"},{"key":"2026092408094438100_cit25","doi-asserted-by":"publisher","first-page":"6888","DOI":"10.1021\/jp403923q","article-title":"High excitation\nenergy\nquenching in fucoxanthin chlorophyll a\/c-binding protein complexes from the diatom Chaetoceros gracilis","volume":"117","author":"Nagao","year":"2013","journal-title":"J. Phys. Chem. B"},{"key":"2026092408094438100_cit26","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1021\/acs.jpcb.8b09253","article-title":"Low-energy\nchlorophylls in fucoxanthin chlorophyll a\/c-binding protein conduct excitation energy transfer to\nphotosystem I in diatoms","volume":"123","author":"Nagao","year":"2019","journal-title":"J. Phys. Chem. B"},{"key":"2026092408094438100_cit27","doi-asserted-by":"publisher","first-page":"29","DOI":"10.3354\/meps195029","article-title":"Separation of chlorophylls and carotenoids\nfrom marine phytoplankton: a new HPLC method using a reversed phase\nC8 column and pyridine-containing mobile phases","volume":"195","author":"Zapata","year":"2000","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2026092408094438100_cit28","doi-asserted-by":"publisher","first-page":"60","DOI":"10.1007\/s11120-025-01182-1","article-title":"Changes in\nlight-harvesting and energy-transfer processes of the chlorophyll d-dominated cyanobacterium Acaryochloris marina in response to different light qualities","volume":"163","author":"Wang","year":"2025","journal-title":"Photosynth. Res."},{"key":"2026092408094438100_cit29","doi-asserted-by":"publisher","first-page":"5179","DOI":"10.1021\/acs.jpcb.5c08754","article-title":"Analysis of optical spectra of intramolecular\ncharge transfer system for Gibbs energy evaluation; solvent-independent\ncharge transfer band","volume":"130","author":"Takamoto","year":"2026","journal-title":"J. Phys. Chem. B"},{"key":"2026092408094438100_cit30","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1016\/j.bbabio.2004.08.009","article-title":"P700+-and 3P700-induced quenching\nof the fluorescence at 760 nm in trimeric Photosystem I complexes\nfrom the cyanobacterium Arthrospira platensis","volume":"1706","author":"Schlodder","year":"2005","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit31","doi-asserted-by":"publisher","first-page":"732","DOI":"10.1016\/j.bbabio.2007.01.013","article-title":"Steady-state\nand transient polarized absorption spectroscopy of photosystem I complexes\nfrom the cyanobacteria Arthrospira platensis and Thermosynechococcus elongatus","volume":"1767","author":"Schlodder","year":"2007","journal-title":"Biochim.\nBiophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit32","doi-asserted-by":"publisher","first-page":"5279","DOI":"10.1038\/s41467-020-18884-w","article-title":"The structure of a red-shifted photosystem\nI reveals a red site in the core antenna","volume":"11","author":"Toporik","year":"2020","journal-title":"Nat.\nCommun."},{"key":"2026092408094438100_cit33","doi-asserted-by":"publisher","first-page":"e73990","DOI":"10.7554\/eLife.73990","article-title":"Structural basis for the absence of low-energy chlorophylls\nin a photosystem I trimer from Gloeobacter violaceus","volume":"11","author":"Kato","year":"2022","journal-title":"elife"},{"key":"2026092408094438100_cit34","doi-asserted-by":"publisher","first-page":"2112","DOI":"10.1104\/pp.104.046110","article-title":"Phycobilisome\ndiffusion is required for light-state\ntransitions in cyanobacteria","volume":"135","author":"Joshua","year":"2004","journal-title":"Plant Physiol."},{"key":"2026092408094438100_cit35","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1007\/s11120-013-9830-5","article-title":"Modification of energy-transfer\nprocesses in the cyanobacterium, Arthrospira platensis, to adapt to light conditions, probed by time-resolved fluorescence\nspectroscopy","volume":"117","author":"Akimoto","year":"2013","journal-title":"Photosynth. Res."},{"key":"2026092408094438100_cit36","doi-asserted-by":"publisher","first-page":"327","DOI":"10.1016\/j.bbabio.2007.02.012","article-title":"Delayed fluorescence\nobserved in the nanosecond time\nregion at 77 K originates directly from the photosystem II reaction\ncenter","volume":"1767","author":"Mimuro","year":"2007","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit37","doi-asserted-by":"publisher","first-page":"847","DOI":"10.1016\/j.bbabio.2011.03.014","article-title":"Excitation\nenergy transfer between\nphotosystem II and photosystem I in red algae: larger amounts of phycobilisome\nenhance spillover","volume":"1807","author":"Yokono","year":"2011","journal-title":"Biochim. Biophys. Acta, Bioenerg."},{"key":"2026092408094438100_cit38","doi-asserted-by":"publisher","first-page":"6675","DOI":"10.1038\/ncomms7675","article-title":"A megacomplex composed\nof both photosystem reaction centres in higher plants","volume":"6","author":"Yokono","year":"2015","journal-title":"Nat. Commun."}],"container-title":["The Journal of Physical\nChemistry B"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acs.jpcb.6c03891","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/jpcbfk\/article-pdf\/130\/38\/9843\/68455217\/acs.jpcb.6c03891.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/jpcbfk\/article-pdf\/130\/38\/9843\/68455217\/acs.jpcb.6c03891.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T12:23:19Z","timestamp":1790252599000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/jpcbfk\/article\/130\/38\/9843\/5426756\/Altered-Excitation-Energy-Transfer-in-the-Dark"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,9]]},"references-count":38,"journal-issue":{"issue":"38","published-online":{"date-parts":[[2026,9,9]]},"published-print":{"date-parts":[[2026,9,24]]}},"URL":"https:\/\/doi.org\/10.1021\/acs.jpcb.6c03891","relation":{},"ISSN":["1520-6106","1520-5207"],"issn-type":[{"value":"1520-6106","type":"print"},{"value":"1520-5207","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,9]]}}}