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Thermodynamic Limits Around M-dwarf Stars: Acknowledgements

by Photosynthesis Technology: It's not just for plants! August 1st, 2024

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In this study, researchers explore the feasibility and potential characteristics of photosynthetic light-harvesting on exo-planets.

featured image - Thermodynamic Limits Around M-dwarf Stars: Acknowledgements

(1) Samir Chitnavis, School of Biological and Behavioural Sciences, Queen Mary University of London, Mile End, London E1 4NS, UK & Digital Environment Research Institute, Queen Mary University of London, Empire House, Whitechapel E1 1HH, UK;

(2) Thomas J. Haworth, Astronomy Unit, Queen Mary University of London, Mile End Road, London E1 4NS, UK;

(3) Edward Gillen, Astronomy Unit, Queen Mary University of London, Mile End Road, London E1 4NS, UK;

(4) Conrad W. Mullineaux, School of Biological and Behavioural Sciences, Queen Mary University of London, Mile End, London E1 4NS, UK;

(5) Christopher D. P. Duffy, School of Biological and Behavioural Sciences, Queen Mary University of London, Mile End, London E1 4NS, UK & Digital Environment Research Institute, Queen Mary University of London, Empire House, Whitechapel E1 1HH, UK (Email: [email protected]).

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5 Acknowledgements

CDPD and SC would like to acknowledge support from the BBSRC London Interdisciplinary Bioscience PhD Consortium (LIDo). Additionally, CDPD would like to acknowledge support from BBSRC for funding work (grant BB/T000023/1) that developed the key ideas in this work. TJH is funded by a Royal Society Dorothy Hodgkin Fellowship. EG gratefully acknowledges support from the UK Science and Technology Facilities Council (STFC; project reference ST/W001047/1).

6 Author Contribution Statement

CDPD, TJH and in discussion with EG and CWM, devised the project. TJH and EG provided stellar spectral data. Light-harvesting model development and simulations were carried out by CDPD and SC. TJH, EG and CWM provided additional interpretation of results. All authors were involved in Manuscript drafting.

7 Authors disclosure statement

There are no conflicts of interest to disclose.

References

E. R. Adams, S. Seager, and L. Elkins-Tanton. Ocean Planet or Thick Atmosphere: On the Mass-Radius Relationship for Solid Exoplanets with Massive Atmospheres. Astrophysical Journal, 673(2):1160–1164, February 2008. doi: 10.1086/524925.

Guillem Anglada-Escud´e, Pedro J. Amado, John Barnes, Zaira M. Berdi˜nas, R. Paul Butler, Gavin A. L. Coleman, Ignacio de La Cueva, Stefan Dreizler, Michael Endl, Benjamin Giesers, Sandra V. Jeffers, James S. Jenkins, Hugh R. A. Jones, Marcin Kiraga, Martin K¨urster, Mar´ıa J. L´opez-Gonz´alez, Christopher J. Marvin, Nicol´as Morales, Julien Morin, Richard P. Nelson, Jos´e L. Ortiz, Aviv Ofir, Sijme-Jan Paardekooper, Ansgar Reiners, Eloy Rodr´ıguez, Cristina Rodr´ıguez-L´opez, Luis F. Sarmiento, John P. Strachan, Yiannis Tsapras, Mikko Tuomi, and Mathias Zechmeister. A terrestrial planet candidate in a temperate orbit around Proxima Centauri. Nature, 536(7617): 437–440, August 2016. doi: 10.1038/nature19106.

Arnold, L., Gillet, S., Lardi`ere, O., Riaud, P., and Schneider, J. A test for the search for life on extrasolar planets* - looking for the terrestrial vegetation signature in the earthshine spectrum. A&A, 392(1):231–237, 2002. doi: 10.1051/0004-6361:20020933. URL https://doi.org/10.1051/0004-6361: 20020933.

Eva-Mari Aro, Ivar Virgin, and Bertil Andersson. Photoinhibition of photosystem II. inactivation, protein damage and turnover. Biochimica et Biophysica Acta (BBA) - Bioenergetics, 1143(2):113–134, 1993. doi: 10.1016/ 0005-2728(93)90134-2.

Trevor B. Arp, Jed Kistner-Morris, Vivek Aji, Richard J. Cogdell, Rienk van Grondelle, and Nathaniel M. Gabor. Quieting a noisy antenna reproduces photosynthetic light-harvesting spectra. Science, 368(6498):1490–1495, 2020. doi: 10.1126/science.aba6630. URL https://www.science.org/doi/abs/ 10.1126/science.aba6630.

S. Bailey, R. G. Walters, S. Jansson, and P. Horton. Acclimation of Arabidopsis thaliana to the light environment: the existence of separate low light and high light responses. . Planta, 213(5):794–801, 2001. doi: 10.1007/s004250100556.

Neil R. Baker and Eva Rosenqvist. Applications of chlorophyll fluorescence can improve crop production strategies: an examination of future possibilities. Journal of Experimental Botany, 55(403):1607–1621, 07 2004. doi: 10.1093/ jxb/erh196.

V. Barzda, V. Gulbinas, R. Kananavicius, V. Cervinskas, H. van Amerongen, R. van Grondelle, and L. Valkunas. Singlet-singlet annihilation kinetics in aggregates and trimers of lhcii. Biophys. J., 80:2409–2421, 2001.

Mariano Battistuzzi, Lorenzo Cocola, Riccardo Claudi, Anna Caterina Pozzer, Anna Segalla, Diana Simionato, Tomas Morosinotto, Luca Poletto, and Nicoletta La Rocca. Oxygenic photosynthetic responses of cyanobacteria exposed under an m-dwarf starlight simulator: Implications for exoplanet’s habitability. Frontiers in Plant Science, 14, 2023.

J. Thomas Beatty, J¨org Overmann, Michael T. Lince, Ann K. Manske, Andrew S. Lang, Robert E. Blankenship, Cindy L. Van Dover, Tracey A. Martinson, and F. Gerald Plumley. An obligately photosynthetic bacterial anaerobe from a deep-sea hydrothermal vent. Proceedings of the National Academy of Sciences, 102(26):9306–9310, 2005.

Bjoern Benneke and Sara Seager. Atmospheric Retrieval for Super-Earths: Uniquely Constraining the Atmospheric Composition with Transmission Spectroscopy. Astrophysical Journal, 753(2):100, July 2012. doi: 10.1088/ 0004-637X/753/2/100.

Bj¨orn. Why are plants green? relationships between pigment absorption and photosynthetic efficiency. Photosynthetica, 10:121–129, 1976.

G. Ozan Bozdag, Eric Libby, Rozenn Pineau, Christopher T. Reinhard, and William C. Ratcliff. Oxygen suppression of macroscopic multicellularity. Nature Communications, 12:2041–1723, 2021.

Koen Broess, Gediminas Trinkunas, Chantal D. van der Weij-de Wit, Jan P. Dekker, Arie van Hoek, and Herbert van Amerongen. Excitation energy transfer and charge separation in photosystem ii membranes. Biophys. J., 91: 3776–3786, 2006.

D. A. Bryant and N. U. Frigaard. Prokaryotic photosynthesis and phototrophy illuminated. Trends Microbiol., 14:488–496, 2006.

NJ Butterfield. Oxygen, animals and oceanic ventilation: an alternative view, 2009.

T. Cardona, A. Sedoud, and A. W. Cox, N.and Rutherford. Charge separation in photosystem ii: a comparative and evolutionary overview. Biochim Biophys Acta., 1817:26–43, 2012. doi: doi:10.1016/j.bbabio.2011.07.012.

Gilles Chabrier. Galactic Stellar and Substellar Initial Mass Function. Publications of the Astronomical Society of the Pacific, 115(809):763–795, 2003. doi: 10.1086/376392.

Charles S. Cockell, Lisa Kaltenegger, and John A. Raven. Cryptic Photosynthesis-Extrasolar Planetary Oxygen Without a Surface Biological Signature. Astrobiology, 9(7):623–636, September 2009. doi: 10.1089/ast. 2008.0273.

Devon B Cole, Daniel B Mills, Douglas H Erwin, Erik A Sperling, Susannah M Porter, Christopher T Reinhard, and Noah J Planavsky. On the co-evolution of surface oxygen levels and animals. Geobiology, 18(3):260–281, 2020.

Giovanni Covone, Riccardo M Ienco, Luca Cacciapuoti, and Laura Inno. Efficiency of the oxygenic photosynthesis on Earth-like planets in the habitable zone. Monthly Notices of the Royal Astronomical Society, 505(3):3329–3335, 2021.

Jan P. Dekker and Egbert J. Boekema. Supramolecular organization of thylakoid membrane proteins in green plants. Biochimica et Biophysica Acta (BBA) - Bioenergetics, 1706(1):12–39, 2005. ISSN 0005-2728. doi: https://doi.org/10. 1016/j.bbabio.2004.09.009.

Jason A. Dittmann, Jonathan M. Irwin, David Charbonneau, Xavier Bonfils, Nicola Astudillo-Defru, Rapha¨elle D. Haywood, Zachory K. Berta-Thompson, Elisabeth R. Newton, Joseph E. Rodriguez, Jennifer G. Winters, ThiamGuan Tan, Jose-Manuel Almenara, Fran¸cois Bouchy, Xavier Delfosse, Thierry Forveille, Christophe Lovis, Felipe Murgas, Francesco Pepe, Nuno C. Santos, Stephane Udry, Ana¨el W¨unsche, Gilbert A. Esquerdo, David W. Latham, and Courtney D. Dressing. A temperate rocky super-Earth transiting a nearby cool star. Nature, 544(7650):333–336, April 2017. doi: 10.1038/nature22055.

Christopher D. P. Duffy, Gregoire Canchon, Thomas J. Haworth, Edward Gillen, Samir Chitnavis, and Conrad W. Mullineaux. Photosynthesis under a red sun: Predicting the absorption characteristics of an extraterrestrial lightharvesting antenna, 2023.

J. P. Faria, A. Su´arez Mascare˜no, P. Figueira, A. M. Silva, M. Damasso, O. Demangeon, F. Pepe, N. C. Santos, R. Rebolo, S. Cristiani, V. Adibekyan, Y. Alibert, R. Allart, S. C. C. Barros, A. Cabral, V. D’Odorico, P. Di Marcantonio, X. Dumusque, D. Ehrenreich, J. I. Gonz´alez Hern´andez, N. Hara, J. Lillo-Box, G. Lo Curto, C. Lovis, C. J. A. P. Martins, D. M´egevand, A. Mehner, G. Micela, P. Molaro, N. J. Nunes, E. Pall´e, E. Poretti, S. G. Sousa, A. Sozzetti, H. Tabernero, S. Udry, and M. R. Zapatero Osorio. A candidate short-period sub-Earth orbiting Proxima Centauri. Astronomy and Astrophysics, 658:A115, February 2022. doi: 10.1051/0004-6361/202142337.

Graham R. Fleming, Gabriela S. Schlau-Cohen, Kapil Amarnath, and Julia Zaks. Design principles of photosynthetic light-harvesting. Faraday Discuss., 155:27–41, 2012.

Micha¨el Gillon, Amaury H. M. J. Triaud, Brice-Olivier Demory, Emmanu¨el Jehin, Eric Agol, Katherine M. Deck, Susan M. Lederer, Julien de Wit, Artem Burdanov, James G. Ingalls, Emeline Bolmont, Jeremy Leconte, Sean N. Raymond, Franck Selsis, Martin Turbet, Khalid Barkaoui, Adam Burgasser, Matthew R. Burleigh, Sean J. Carey, Aleksander Chaushev, Chris M. Copperwheat, Laetitia Delrez, Catarina S. Fernandes, Daniel L. Holdsworth, Enrico J. Kotze, Val´erie Van Grootel, Yaseen Almleaky, Zouhair Benkhaldoun, Pierre Magain, and Didier Queloz. Seven temperate terrestrial planets around the nearby ultracool dwarf star TRAPPIST-1. Nature, 542(7642):456–460, February 2017. doi: 10.1038/nature21360.

Lea Gregersen, Donald Bryant, and Niels-Ulrik Frigaard. Mechanisms and evolution of oxidative sulfur metabolism in green sulfur bacteria. Frontiers in Microbiology, 2, 2011. ISSN 1664-302X. doi: 10.3389/fmicb.2011.00116. URL https://www.frontiersin.org/articles/10.3389/fmicb.2011.00116.

Simon L. Grimm, Brice-Olivier Demory, Micha¨el Gillon, Caroline Dorn, Eric Agol, Artem Burdanov, Laetitia Delrez, Marko Sestovic, Amaury H. M. J. Triaud, Martin Turbet, Emeline Bolmont, Anthony Caldas, Julien de Wit, ´ Emmanu¨el Jehin, J´er´emy Leconte, Sean N. Raymond, Val´erie Van Grootel, Adam J. Burgasser, Sean Carey, Daniel Fabrycky, Kevin Heng, David M. Hernandez, James G. Ingalls, Susan Lederer, Franck Selsis, and Didier Queloz. The nature of the TRAPPIST-1 exoplanets. Astronomy and Astrophysics, 613:A68, May 2018. doi: 10.1051/0004-6361/201732233.

C. Hall, P. C. Stancil, J. P. Terry, and C. K. Ellison. A new definition of exoplanet habitability: Introducing the photosynthetic habitable zone, 2023. URL https://arxiv.org/abs/2301.13836.

Sasha Hinkley, Aarynn L. Carter, Shrishmoy Ray, Andrew Skemer, Beth Biller, Elodie Choquet, Maxwell A. Millar-Blanchaer, Stephanie Sallum, Brittany Miles, Niall Whiteford, Polychronis Patapis, Marshall Perrin, Laurent Pueyo, Glenn Schneider, Karl Stapelfeldt, Jason Wang, Kimberly Ward-Duong, Brendan P. Bowler, Anthony Boccaletti, Julien H. Girard, Dean Hines, Paul Kalas, Jens Kammerer, Pierre Kervella, Jarron Leisenring, Eric Pantin, Yifan Zhou, Michael Meyer, Michael C. Liu, Mickael Bonnefoy, Thayne Currie, Michael McElwain, Stanimir Metchev, Mark Wyatt, Olivier Absil, Jea Adams, Travis Barman, Isabelle Baraffe, Mariangela Bonavita, Mark Booth, Marta Bryan, Gael Chauvin, Christine Chen, Camilla Danielski, Matthew De Furio, Samuel M. Factor, Michael P. Fitzgerald, Jonathan J. Fortney, Carol Grady, Alexandra Greenbaum, Thomas Henning, Kielan K. W. Hoch, Markus Janson, Grant Kennedy, Matthew Kenworthy, Adam Kraus, Masayuki Kuzuhara, Pierre-Olivier Lagage, Anne-Marie Lagrange, Ralf Launhardt, Cecilia Lazzoni, James Lloyd, Sebastian Marino, Mark Marley, Raquel Martinez, Christian Marois, Brenda Matthews, Elisabeth C. Matthews, Dimitri Mawet, Johan Mazoyer, Mark Phillips, Simon Petrus, Sascha P. Quanz, Andreas Quirrenbach, Julien Rameau, Isabel Rebollido, Emily Rickman, Matthias Samland, B. Sargent, Joshua E. Schlieder, Anand Sivaramakrishnan, Jordan M. Stone, Motohide Tamura, Pascal Tremblin, Taichi Uyama, Malavika Vasist, Arthur Vigan, Kevin Wagner, and Marie Ygouf. The JWST Early Release Science Program for the Direct Imaging and Spectroscopy of Exoplanetary Systems. Publications of the Astronomical Society of the Pacific, 134(1039):095003, September 2022. doi: 10.1088/1538-3873/ac77bd.

Danley C. Hsu, Eric B. Ford, Darin Ragozzine, and Keir Ashby. Occurrence Rates of Planets Orbiting FGK Stars: Combining Kepler DR25, Gaia DR2, and Bayesian Inference. Astronomical Journal, 158(3):109, September 2019. doi: 10.3847/1538-3881/ab31ab.

Danley C. Hsu, Eric B. Ford, and Ryan Terrien. Occurrence rates of planets orbiting M Stars: applying ABC to Kepler DR25, Gaia DR2, and 2MASS data. Monthly Noticesofthe Royal Astronomical Society, 498(2):2249–2262, October 2020. doi: 10.1093/mnras/staa2391.

Xiche Hu, Thorsten Ritz, Ana Damjanovi´c, Felix Autenrieth, and Klaus Schulten. Photosynthetic apparatus of purple bacteria. Quarterly Reviews of Biophysics, 35(1):1–62, 2002. doi: 10.1017/S0033583501003754.

Husser, T.-O., Wende-von Berg, S., Dreizler, S., Homeier, D., Reiners, A., Barman, T., and Hauschildt, P. H. A new extensive library of phoenix stellar atmospheres and synthetic spectra. A&A, 553:A6, 2013. doi: 10. 1051/0004-6361/201219058. URL https://doi.org/10.1051/0004-6361/ 201219058.

Stefan Jansson. The light-harvesting chlorophyll ab-binding proteins. Biochimica et Biophysica Acta (BBA)-Bioenergetics, 1184(1):1–19, 1994.

N. Y. Kiang, A. Segura, G. Tinetti, Govindjee, R. E. Blankenship, M. Cohen, J. Siefert, D. Crisp, and V. S. Meadows. Spectral signatures of photosynthesis ii: Coevolution with other stars and the atmosphere on extrasolar worlds. Astrobiology, 7:252–274, 2007a. doi: 10.1089/ast.2006.0108.

Nancy Y. Kiang. Looking for life elsewhere: Photosynthesis and astrobiology. The Biochemist, 36(6):24–30, 12 2014. ISSN 0954-982X. doi: 10.1042/BIO03606024. URL https://doi.org/10.1042/BIO03606024.

Nancy Y. Kiang, Janet Siefert, Govindjee, and Robert E. Blankenship. Spectral signatures of photosynthesis. i. review of earth organisms. Astrobiology, 7(1): 222–251, 2007b.

R. S. Knox and B. Q Spring. Dipole strengths in the chlorophylls. Photochem. Photobiol., 77, 2003.

Yuval Kolodny, Yoav Avrahami, Hagit Zer, Miguel J. Frada, Yossi Paltiel, and Nir Keren. Phycobilisome light-harvesting efficiency in natural populations of the marine cyanobacteria synechococcus increases with depth. Communications Biology, 5:727, 2022.

Masaharu Kondo, Haruka Matsuda, Tomoyasu Noji, Mamoru Nango, and Takehisa Dewa. Photocatalytic activity of the light-harvesting complex of photosystem ii (lhcii) monomer. Journal of Photochemistry and Photobiology A: Chemistry, 406:112926, 2021.

Roman Kouˇril, Jan P. Dekker, and Egbert J. Boekema. Supramolecular organization of photosystem ii in green plants. Biochimica et Biophysica Acta (BBA) - Bioenergetics, 1817(1):2–12, 2012. doi: https://doi.org/10.1016/j.bbabio. 2011.05.024. URL https://www.sciencedirect.com/science/article/ pii/S0005272811001514.

Roman Kouˇril, Emilie Wientjes, Jelle B. Bultema, Roberta Croce, and Egbert J. Boekema. High-light vs. low-light: Effect of light acclimation on photosystem ii composition and organization in arabidopsis thaliana. Biochimica et Biophysica Acta (BBA) - Bioenergetics, 1827(3):411–419, 2013. doi: https://doi.org/10.1016/j.bbabio.2012.12.003.

Pavel Kroupa. On the variation of the initial mass function. Monthly Notices of the Royal Astronomical Society, 322(2):231–246, April 2001. doi: 10.1046/ j.1365-8711.2001.04022.x.

Owen R. Lehmer, David C. Catling, Mary N. Parenteau, Nancy Y. Kiang, and Tori M. Hoehler. The peak absorbance wavelength of photosynthetic pigments around other stars from spectral optimization. Frontiers in Astronomy and Space Sciences, 8, 2021.

Xu Li, Tong Liang, and Hongtao Liu. How plants coordinate their development in response to light and temperature signals. The Plant Cell, 34(3):955–966, 12 2021. doi: 10.1093/plcell/koab302.

Manasvi Lingam, Amedeo Balbi, and Swadesh M. Mahajan. Excitation properties of photopigments and their possible dependence on the host star. Astrophysical Journal Letters, 921(2):L41, 2021.

Zhenfeng Liu, Hanchi Yan, Kebin Wang, Tingyun Kuang, Jiping Zhang, Lulu Gui, Xiaomin An, and Wenrui Chang. Crystal structure of spinach major light-harvesting complex at 2.72 ˚a resolution. Nature, 428(6980):287–292, 2004.

Heiko Lokstein, Gernot Renger, and Jan P. G¨otze. Photosynthetic lightharvesting (antenna) complexes—structures and functions. Molecules, 26 (11), 2021. doi: 10.3390/molecules26113378. URL https://www.mdpi.com/ 1420-3049/26/11/3378.

Timothy W Lyons, Christopher T Reinhard, and Noah J Planavsky. The rise of oxygen in earth’s early ocean and atmosphere. Nature, 506(7488):307–315, 2014.

M. A. Marosv¨olgyi and H. J. van Gorkom. Cost and color of photosynthesis. Photosynthesis Research, 103:105–109, 2010.

Vincenzo Mascoli, Luca Bersanini, and Roberta Croce. Far-red absorption and light-use efficiency trade-offs in chlorophyll f photosynthesis. Nature Plants, 6, 08 2020. doi: 10.1038/s41477-020-0718-z.

Vincenzo Mascoli, Ahmad Farhan Bhatti, Luca Bersanini, Herbert van Amerongen, and Roberta Croce. The antenna of far-red absorbing cyanobacteria increases both absorption and quantum efficiency of photosystem ii. Nature Communications, 13:3562, 2022.

K.J. McCree. A solarimeter for measuring photosynthetically active radiation. Agricultural Meteorology, 3(5):353–366, 1966. doi: https://doi.org/10.1016/ 0002-1571(66)90017-3.

K.J. McCree. The action spectrum, absorptance and quantum yield of photosynthesis in crop plants. Agricultural Meteorology, 9:191–216, 1971. doi: https://doi.org/10.1016/0002-1571(71)90022-7.

Christopher P. McKay. Requirements and limits for life in the context of exoplanets. Proceedings of the National Academy of Sciences, 111(35):12628– 12633, 2014. doi: 10.1073/pnas.1304212111.

Victoria S. Meadows and Rory K. Barnes. Factors Affecting Exoplanet Habitability, pages 1–24. Springer International Publishing, 2018. doi: 10.1007/ 978-3-319-30648-3 57-1.

Michael Mendillo, Paul Withers, and Paul A. Dalba. Atomic oxygen ions as ionospheric biomarkers on exoplanets. Nature Astronomy, 2:287––291, 2018. doi: 10.1038/s41550-017-0375-y.

Daniel B Mills and Donald E Canfield. Oxygen and animal evolution: Did a rise of atmospheric oxygen “trigger” the origin of animals? BioEssays, 36 (12):1145–1155, 2014.

Gijs D. Mulders, Ilaria Pascucci, and D´aniel Apai. A Stellar-mass-dependent Drop in Planet Occurrence Rates. Astrophysical Journal, 798(2):112, January 2015. doi: 10.1088/0004-637X/798/2/112.

NASA Exoplanet Archive. https://exoplanetarchive.ipac.caltech.edu/.

Nikolay Nikolov, David K. Sing, Neale P. Gibson, Jonathan J. Fortney, Thomas M. Evans, Joanna K. Barstow, Tiffany Kataria, and Paul A. Wilson. VLT FORS2 Comparative Transmission Spectroscopy: Detection of Na in the Atmosphere of WASP-39b from the Ground. Astrophysical Journal, 832(2): 191, December 2016. doi: 10.3847/0004-637X/832/2/191.

Vladimir Novoderezhkin, Alessandro Marin, and Rienk van Grondelle. Intraand inter-monomeric transfers in the light harvesting LHCII complex: the redfield–f¨orster picture. Physical Chemistry Chemical Physics, 13(38):17093– 17103, 2011. doi: 10.1039/C1CP21079C.

D. Noy, C. C. Moser, and P. L. Dutton. Design and engineering of photosynthetic light-harvesting and electron transfer using length, time, and energy scales. Biochim. Biophys. Acta., 1757:90–105, 2006. doi: 10.1016/j.bbabio.2005.11. 010.

Dennis J N¨urnberg, Jennifer Morton, Stefano Santabarbara, Alison Telfer, Pierre Joliot, Laura A Antonaru, Alexander V Ruban, Tanai Cardona, Elmars Krausz, Alain Boussac, et al. Photochemistry beyond the red limit in chlorophyll f–containing photosystems. Science, 360(6394):1210–1213, 2018.

Vello Oja and Agu Laisk. Oxygen yield from single turnover flashes in leaves: non-photochemical excitation quenching and the number of active psii. Biochimica et Biophysica Acta (BBA)-Bioenergetics, 1460(2-3):291–301, 2000.

SL Olson, EW Schwieterman, CT Reinhard, and TW Lyons. Handbook of exoplanets ed h. Deeg and J. Belmont (Cham: Springer), 189, 2018a.

Stephanie L. Olson, Edward W. Schwieterman, Christopher T. Reinhard, Andy Ridgwell, Stephen R. Kane, Victoria S. Meadows, and Timothy W. Lyons. Atmospheric seasonality as an exoplanet biosignature. Astrophysical Journal Letters, 858:2, 2018b.

Jack T. O’Malley-James and Lisa Kaltenegger. The vegetation red edge biosignature through time on earth and exoplanets. Astrobiology, 18(9):1123–1136, 2018. doi: 10.1089/ast.2017.1798. URL https://doi.org/10.1089/ast. 2017.1798. PMID: 30204495.

Gert T Oostergetel, Herbert Van Amerongen, and Egbert J Boekema. The chlorosome: a prototype for efficient light harvesting in photosynthesis. Photosynthesis research, 104:245–255, 2010.

J¨org Overmann and Ferran Garcia-Pichel. The Phototrophic Way of Life, pages 203–257. Springer Berlin Heidelberg, Berlin, Heidelberg, 2013. ISBN 978-3- 642-30123-0. doi: 10.1007/978-3-642-30123-0 51. URL https://doi.org/ 10.1007/978-3-642-30123-0_51.

Kazumi Ozaki, Katharine J Thompson, Rachel L Simister, Sean A Crowe, and Christopher T Reinhard. Anoxygenic photosynthesis and the delayed oxygenation of earth’s atmosphere. Nature communications, 10(1):3026, 2019.

Ankit Rohatgi. Webplotdigitizer: Version 4.6, 2022. URL https://automeris. io/WebPlotDigitizer.

Alexander V. Ruban. Nonphotochemical chlorophyll fluorescence quenching: Mechanism and effectiveness in protecting plants from photodamage. Plant Physiology, 170(4):1903–1916, 2016. doi: 10.1104/pp.15.01935. URL https: //doi.org/10.1104/pp.15.01935.

Alexander V Ruban and Sam Wilson. The mechanism of non-photochemical quenching in plants: localization and driving forces. Plant and Cell Physiology, 62(7):1063–1072, 2021.

Rafael G Saer and Robert E Blankenship. Light harvesting in phototrophic bacteria: structure and function. Biochemical Journal, 474(13):2107–2131, 2017.

C. Sagan, W. R. Thompson, R. Carlson, D. Gurnett, and C. Hord. A search for life on earth from the galileo spacecraft. Nature, 365:715–721, 1993.

P. S´anchez-Baracaldo and T. Cardona. On the origin of oxygenic photosynthesis and cyanobacteria. New Phytol, 225:1440–1446, 2019.

Edward W. Schwieterman, Nancy Y. Kiang, Mary N. Parenteau, Chester E. Harman, Shiladitya DasSarma, Theresa M. Fisher, Giada N. Arney, Hilairy E. Hartnett, Christopher T. Reinhard, Stephanie L. Olson, Victoria S. Meadows, Charles S. Cockell, Sara I. Walker, John Lee Grenfell, Siddharth Hegde, Sarah Rugheimer, Renyu Hu, and Timothy W. Lyons. Exoplanet biosignatures: A review of remotely detectable signs of life. Astrobiology, 18(6):663–708, 2018. doi: 10.1089/ast.2017.1729.

S. Seager, E. L. Turner, J. Schafer, and E. B. Ford. Vegetation’s Red Edge: A Possible Spectroscopic Biosignature of Extraterrestrial Plants. Astrobiology, 5(3):372–390, 2005. doi: 10.1089/ast.2005.5.372.

S. Seager, M. Kuchner, C. A. Hier-Majumder, and B. Militzer. Mass-Radius Relationships for Solid Exoplanets. Astrophysical Journal, 669(2):1279–1297, November 2007. doi: 10.1086/521346.

J. Semer, M. Navr´atil, V. Spunda, and M. ˇ Stroch. Chlorophyll fluorescence ˇ parameters to assess utilization of excitation energy in photosystem ii independently of changes in leaf absorption. J. Photochem. Photobiol. B., 197 (111535), 2019. doi: doi:10.1016/j.jphotobiol.2019.111535.

Sourav Sil, Ryan W Tilluck, Nila Mohan TM, Chase H Leslie, Justin B Rose, Maria Agustina Dom´ınguez-Mart´ın, Wenjing Lou, Cheryl A Kerfeld, and Warren F Beck. Excitation energy transfer and vibronic coherence in intact phycobilisomes. Nature Chemistry, 14(11):1286–1294, 2022.

Rochelle M. Soo, James Hemp, Donovan H. Parks, Woodward W. Fischer, and Philip Hugenholtz. On the origins of oxygenic photosynthesis and aerobic respiration in cyanobacteria. Science, 355(6332):1436–1440, 2017. doi: 10. 1126/science.aal3794.

Martijn Tros, Vincenzo Mascoli, Gaozhong Shen, Ming-Yang Ho, Luca Bersanini, Christopher J. Gisriel, Donald A. Bryant, and Roberta Croce. Breaking the red limit: Efficient trapping of long-wavelength excitations in chlorophyll-f-containing photosystem i. Chem, 7:155–173, 2021. ISSN 2451- 9294.

Leonas Valkunas, Gediminas Trinkunas, Jevgenij Chmeliov, and Alexander V. Ruban. Modeling of exciton quenching in photosystem ii. Phys. Chem. Chem. Phys., 11:7576–7584, 2009. doi: 10.1039/B901848D.

Silvere Vialet-Chabrand, Jack SA Matthews, Andrew J Simkin, Christine A Raines, and Tracy Lawson. Importance of fluctuations in light on plant photosynthetic acclimation. Plant physiology, 173(4):2163–2179, 2017.

Stefania Viola, William Roseby, Stefano Santabarbara, Dennis N¨urnberg, Ricardo Assun¸c˜ao, Holger Dau, Julien Sell´es, Alain Boussac, Andrea Fantuzzi, and A William Rutherford. Impact of energy limitations on function and resilience in long-wavelength photosystem ii. eLife, 11:e79890, jul 2022. doi: 10.7554/eLife.79890.

X. Wei, X. Su, P. Cao, X. Liu, W. Chang, M. Li, X. Zhang, and Z. Liu. Structure of spinach Photosystem II-LHCII supercomplex at 3.2 ˚A resolution. Nature, 534:69–74, 2016.

Lauren M. Weiss and Geoffrey W. Marcy. The Mass-Radius Relation for 65 Exoplanets Smaller than 4 Earth Radii. Astrophysical Journal Letters, 783 (1):L6, March 2014. doi: 10.1088/2041-8205/783/1/L6.

G. R. Wolfe, F. X. Cunningham, D. G. Durnford, Green B. R., and E. Gantt. Evidence for a common origin of chloroplasts with light-harvesting complexes of different pigmentation. Nature, 367:566–568, 1994.

R.D. Wolstencroft and J.A. Raven. Photosynthesis: Likelihood of occurrence and possibility of detection on earth-like planets. Icarus, 157:535–548, 2002

Rachel Wood, Philip CJ Donoghue, Timothy M Lenton, Alexander G Liu, and Simon W Poulton. The origin and rise of complex life: progress requires interdisciplinary integration and hypothesis testing, 2020.

Shi-Wei Wu, Arjan Bik, Joachim M. Bestenlehner, Thomas Henning, Anna Pasquali, Wolfgang Brandner, and Andrea Stolte. The massive stellar population of W49: A spectroscopic survey. Astronomy and Astrophysics, 589: A16, May 2016. doi: 10.1051/0004-6361/201527823.

Xiaolong Yang, Lihua Liu, Zhikai Yin, Xingyu Wang, Shoubing Wang, and Zipiao Ye. Quantifying photosynthetic performance of phytoplankton based on photosynthesis–irradiance response models. Environmental Sciences Europe, 32:24, 2020.