We honor Professor Hartmut Karl Lichtenthaler, a versatile pioneer of photosynthesis research, plant physiology, isoprenoid biochemistry, and stress physiology of plants, for his groundbreaking and creative contributions to plant science. His innovative research on the chemical composition, ultrastructure, and function of chloroplasts and his detection of the major methylerythritol 4-phosphate (MEP) isoprenoid biosynthetic pathway in plants is key to our current understanding of the physiology and biochemistry of photosynthesis systems. His ingenious use of the powerful laser-induced chlorophyll a fluorescence imaging has helped us better understand the stress response processes in plant leaves. In this tribute, we present a summary of Lichtenthaler's career, significant scientific contributions, editorial engagement, promotion of international cooperation, many honors, and awards, as well as his devotion to hiking and mountaineering.

Department of Plant and Microbial Biology University of California Berkeley Berkeley 94720 California USA

Department of Plant Biology Department of Biochemistry and Center of Biophysics and Quantitative Biology University of Illinois at Urbana Champaign Urbana 61801 Illinois USA

MSU DOE Plant Research Laboratory Plant Resilience Institute and Department of Biochemistry and Molecular Biology Michigan State University East Lansing 48824 Michigan USA

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Blankenship R.E.: Molecular Mechanisms of Photosynthesis. 3rd Edition. Pp. 352. John Wiley & Sons, Hoboken: 2021. https://www.wiley.com/en-us/Molecular+Mechanisms+of+Photosynthesis,+3rd+Edition-p-9781119800019

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Buchanan B.B., Douce R., Lichtenthaler H.K.: Andrew A. Benson. – Photosynth. Res. 92: 143-144, 2007. 10.1007/s11120-007-9205-x PubMed DOI

Buchanan B.B., Tagawa K.: Perspective on Daniel I. Arnon's contributions to research, 1960–1994. – Photosynth. Res. 46: 27-35, 1995. 10.1007/BF00020412 PubMed DOI

Buschmann C., Lichtenthaler H.K.: Principles and characteristics of multi-colour fluorescence imaging of plants. – J. Plant Physiol. 152: 297-314, 1998. 10.1016/S0176-1617(98)80144-2 DOI

Golz A., Focke M., Lichtenthaler H.K.: Inhibitors of de novo fatty acid biosynthesis in higher plants. – Plant Physiol. 143: 426-433, 1994. 10.1016/S0176-1617(11)81802-X DOI

Govindjee G.: Celebrating Andrew Alm Benson’s 93rd birthday. – Photosynth. Res. 105: 201-208, 2010. 10.1007/s11120-010-9591-3 PubMed DOI

Govindjee G.: Frederick Robert (Bob) Whatley (1924-2020): Co-discoverer of photophosphorylation in chloroplasts and much more. – J. Plant Sci. Res. 38: 1-7, 2022. https://www.life.illinois.edu/govindjee/recent_papers_files/Whatley (1924-2020)ByGovindjee(2022).pdf

Govindjee G., Bassham H., Bassham S.: Remembering James Alan Bassham (1922-2012). – Photosynth. Res. 128: 3-13, 2016. 10.1007/s11120-015-0201-2 PubMed DOI

Govindjee G., Nonomura A., Lichtenthaler H.K.: Remembering Melvin Calvin (1911-1997), a highly versatile scientist of the 20th century. – Photosynth. Res. 143: 1-11, 2020. 10.1007/s11120-019-00693-y PubMed DOI

Govindjee G., Šesták Z., Peters W.R.: The early history of “Photosynthetica”, “Photosynthesis Research” and their publishers. – Photosynthetica 40: 1-11, 2002. 10.1023/A:1020169502548 DOI

Govindjee G., Yoo H.: The International Society of Photosynthesis research (ISPR) and its associated International Congress on Photosynthesis (ICP): a pictorial report. – Photosynth. Res. 91: 95-106, 2007. 10.1007/s11120-007-9189-6 DOI

Hák R., Rinderle-Zimmer U., Lichtenthaler H.K., Nátr L.: Chlorophyll a fluorescence signatures of nitrogen-deficient barley leaves. – Photosynthetica 28: 151-159, 1993. https://kramerius.lib.cas.cz/view/uuid:160f5eec-07f1-42d2-8d4b-c139576bfa19?page=uuid:962826f0-e579-49ef-b07e-03258d1b7ed6

Hu K., Govindjee G., Tan J. et al.: Co-author and co-cited reference network analysis for chlorophyll fluorescence research from 1991 to 2018. – Photosynthetica 58: 110-124, 2020. 10.32615/ps.2019.154 DOI

Interschick-Niebler E., Lichtenthaler H.K.: Partition of phylloquinone K1 between digitonin particles and chlorophyll-proteins of chloroplast membranes from Nicotiana tabacum. – Z. Naturforsch. C 36: 276-283, 1981. 10.1515/znc-1981-3-415 DOI

Joyard J., Lichtenthaler H.K.: Tribute: Roland Douce, 1939–2018. – Photosynth. Res. 141: 131-142, 2019. 10.1007/s11120-019-00634-9 PubMed DOI

Karapetyan N.V., Govindjee G.: Alexander Abramovich Krasnovsky (1913-1993): 100th birth anniversary in Moscow, Russia. – Photosynth. Res. 120: 347-353, 2014. 10.1007/s11120-014-9971-1 PubMed DOI

Kobek K., Focke M., Lichtenthaler H.K. et al.: Inhibition of fatty acid biosynthesis in isolated chloroplasts by cycloxydim and other cyclohexane-1,3-diones. – Physiol. Plantarum 72: 492-498, 1988. 10.1111/j.1399-3054.1988.tb09156.x DOI

Lichtenthaler H.K.: Plastoglobuli and the fine structure of plastids. – Endeavour XXVII: 144-149, 1968.

Lichtenthaler H.K.: Localization and functional concentrations of lipoquinones in chloroplasts. – In: Metzner H. (ed.): Progress in Photosynthesis Research. Proceedings of First International Congress of Photosynthesis Research, held at Freudenstadt 1968. Vol 1. Pp. 304-314. H. Laupp, Tübingen: 1969.

Lichtenthaler H.K.: Chlorophylls and carotenoids, the pigments of photosynthetic biomembranes. – Method. Enzymol. 148: 350-382, 1987. 10.1016/0076-6879(87)48036-1 DOI

Lichtenthaler H.K.: In vivo chlorophyll fluorescence as a tool for stress detection on plants. – In: Lichtenthaler H.K. (ed.): Applications of Chlorophyll Fluorescence in Photosynthesis Research, Stress Physiology and Remote Sensing. Pp. 129-142. Springer, Dordrecht: 1988. 10.1007/978-94-009-2823-7_16 DOI

Lichtenthaler H.K.: Mode of action of herbicides affecting acetyl-CoA carboxylase and fatty acid biosynthesis. – Z. Naturforsch. C 45: 521-528, 1990. 10.1515/znc-1990-0538 DOI

Lichtenthaler H.K.: The Kautsky effect: 60 years of chlorophyll fluorescence induction kinetics. – Photosynthetica 27: 45-55, 1992. https://kramerius.lib.cas.cz/view/uuid:66f03906-9a33-4ae1-bab8-ec77d1a0fc1b?page=uuid:dba5154d-b6cc-47cd-9c83-eaecc0a23b20

Lichtenthaler H.K.: The 1-deoxy-D-xylulose-5-phosphate pathway of isoprenoid biosynthesis in plants. – Annu. Rev. Plant Biol. 50: 47-65, 1999. 10.1146/annurev.arplant.50.1.47 PubMed DOI

Lichtenthaler H.K.: Non-mevalonate isoprenoid biosynthesis: enzymes, genes and inhibitors. – Biochem. Soc. T. 28: 785-789, 2000. 10.1042/bst0280785 PubMed DOI

Lichtenthaler H.K.: A history of the Federation of European Societies of Plant Physiology FESPP since its formation in 1978 – including notes on events preceding the foundation and following re-naming as the Federation of European Societies of Plant Biology (FESPB) in 2002. – J. Plant Physiol. 161: 635-639, 2004a. 10.1078/0176-1617-01371 PubMed DOI

Lichtenthaler H.K.: Thirty years of international symposia of plant lipids. – In: Biacs P., Gerely P. (ed.): Proceedings of the 16th Plant Lipid Symposium. Pp. I-XXXI. Méte, Budapest: 2004b. https://drive.google.com/file/d/0B9tGkMMkZ8IkQV9qSWhTRHdoWVE/edit?usp=sharing

Lichtenthaler H.K.: Biosynthesis, accumulation and emission of carotenoids, α-tocopherol, plastoquinone and isoprene in leaves under high photosynthetic irradiance. – Photosynth. Res. 92: 163-179, 2007. 10.1007/s11120-007-9204-y PubMed DOI

Lichtenthaler H.K.: Plastoglobuli, thylakoids, chloroplast structure and development of plastids. – In: Biswal B., Krupinska K., Biswal U.C. (ed.): Plastid Development in Leaves during Growth and Senescence. Advances in Photosynthesis and Respiration. Vol. 36. Pp. 337-361. Springer, Dordrecht: 2013. 10.1007/978-94-007-5724-0_15 DOI

Lichtenthaler H.K.: Fifty-five years of research on photosynthesis, chloroplasts, and stress physiology of plants: 1958–2013. – In: Lüttge U., Beyschlag W. (ed.): Progress in Botany. Vol. 76. Pp. 3-42. Springer, Cham: 2015. 10.1007/978-3-319-08807-5_1 DOI

Lichtenthaler H.K.: Zdeněk Šesták and Photosynthetica, a tribute. – Photosynthetica 56: 1-6, 2018. 10.1007/s11099-018-0811-y DOI

Lichtenthaler H.K.: My contact and cooperation with Govindjee over the last five decades: Chlorophyll fluorescence and Rebeiz Foundation. – Photosynthetica 58: 976-983, 2020. 10.32615/ps.2020.055 DOI

Lichtenthaler H.K.: Multi-colour fluorescence imaging of photosynthetic activity and plant stress. – Photosynthetica 59: 364-380, 2021. 10.32615/ps.2021.020 DOI

Lichtenthaler H.K., Abuslima E., Nick P.: Strong increase of photosynthetic pigments and leaf size in a pruned Ginkgo biloba tree. – Photosynthetica 61: 297-307, 2023. 10.32615/ps.2023.020 PubMed DOI PMC

Lichtenthaler H.K., Ač A., Marek M.V. et al.: Differences in pigment composition, photosynthetic rates and chlorophyll fluorescence images of sun and shade leaves of four tree species. – Plant Physiol. Biochem. 45: 577-588, 2007b. 10.1016/j.plaphy.2007.04.006 PubMed DOI

Lichtenthaler H.K., Babani F.: Detection of photosynthetic activity and water stress by imaging the red chlorophyll fluorescence. – Plant Physiol. Biochem. 38: 889-895, 2000. 10.1016/S0981-9428(00)01199-2 DOI

Lichtenthaler H.K., Babani F.: Light adaptation and senescence of the photosynthetic apparatus: changes in pigment composition, chlorophyll fluorescence parameters and photosynthetic activity. – In: Papageorgiou G., Govindjee G. (ed.): Chlorophyll Fluorescence: A Signature of Photosynthesis. Pp. 713-736. Springer, Dordrecht: 2004. 10.1007/978-1-4020-3218-9_28 DOI

Lichtenthaler H.K., Babani F.: Level of photosynthetic pigments and ratios of chlorophyll a/b and chlorophylls to carotenoids (a+b)/(x+c) in C4 plants as compared to C3 plants. – Photosynthetica 60: 3-9, 2022. 10.32615/ps.2021.041 PubMed DOI PMC

Lichtenthaler H.K., Babani F., Langsdorf G.: Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees. – Photosynth. Res. 93: 235-241, 2007a. 10.1007/s11120-007-9174-0 PubMed DOI

Lichtenthaler H.K., Buchanan B.B., Douce R.: Honoring Andrew Benson in Paris: A tribute on his 90th birthday. – Photosynth. Res. 96: 181-183, 2008. 10.1007/s11120-008-9295-0 PubMed DOI

Lichtenthaler H.K., Buchanan B.B., Douce R., Govindjee G.: Andrew A. Benson, 1917–2015. – Photosynth. Res. 124: 131-135, 2015. 10.1007/s11120-015-0119-8 PubMed DOI

Lichtenthaler H.K., Burkard G., Grumbach K.H., Meier D.: Physiological effects of photosystem II-herbicides on the development of the photosynthetic apparatus. – Photosynth. Res. 1: 29-43, 1980. 10.1007/BF00020073 PubMed DOI

Lichtenthaler H.K., Buschmann C., Döll M. et al.: Photosynthetic activity, chloroplast ultrastructure, and leaf characteristics of high-light and low-light plants and of sun and shade leaves. – Photosynth. Res. 2: 115-141, 1981. 10.1007/BF00028752 PubMed DOI

Lichtenthaler H.K., Buschmann C.: Das Baumsterben aus botanischer Sicht. [Forest Decline in a Botanical View]. Pp. 87. G. Braun Verlag, Karlsruhe: 1984a. [In German]

Lichtenthaler H.K., Buschmann C.: Photooxidative changes in pigment composition and photosynthetic activity of air-polluted spruce needles (Picea abies L.). – In: Sybesma C. (ed.): Advances in Photosynthesis Research. Advances in Agricultural Biotechnology. Vol. 4. Pp. 245-250. Springer, Dordrecht: 1984b. 10.1007/978-94-017-4971-8_52 DOI

Lichtenthaler H.K., Buschmann C.: Chlorophylls and carotenoids: Measurement and characterization by UV-VIS spectroscopy. – In: Current Protocols in Food Analytical Chemistry. Suppl. 1. Pp. F4.3.1-F4.3.8. John Wiley & Sons, New York: 2001. 10.1002/0471142913.faf0403s01 DOI

Lichtenthaler H.K., Calvin M.: Quinone and pigment composition of chloroplasts and quantasome aggregates from Spinacia oleracea. – BBA-Biophys. Subj. 79: 30-40, 1964. 10.1016/0926-6577(64)90035-X PubMed DOI

Lichtenthaler H.K., Kuhn G., Prenzel U. et al.: Adaptation of chloroplast-ultrastructure and of chlorophyll-protein levels to high-light and low-light growth conditions. – Z. Naturforsch. C 37: 464-475, 1982. 10.1515/znc-1982-5-619 DOI

Lichtenthaler H.K., Langsdorf G., Buschmann C.: Uptake of diuron and concomitant loss of photosynthetic activity in leaves as visualized by imaging the red chlorophyll fluorescence. – Photosynth. Res. 116: 355-361, 2013. 10.1007/s11120-013-9842-1 PubMed DOI

Lichtenthaler H.K., Park R.B.: Chemical composition of chloroplast lamellae from spinach. – Nature 198: 1070-1072, 1963. 10.1038/1981070a0 DOI

Lichtenthaler H.K., Rinderle U.: The role of chlorophyll fluorescence in the detection of stress conditions in plants. – CRC Crit. Rev. Anal. Chem. 19: S29-S85, 1988a. 10.1080/15476510.1988.10401466 DOI

Lichtenthaler H.K., Rinderle U.: Chlorophyll fluorescence signatures as vitality indicator in forest decline research. – In: Lichtenthaler H.K. (ed.): Applications of Chlorophyll Fluorescence in Photosynthesis Research, Stress Physiology, Hydrobiology and Remote Sensing. Pp. 143-149. Springer, Dordrecht: 1988b. 10.1007/978-94-009-2823-7_17 DOI

Lichtenthaler H.K., Schwender J., Disch A., Rohmer M.: Biosynthesis of isoprenoids in higher plant chloroplasts proceeds via a mevalonate-independent pathway. – FEBS Lett. 400: 271-274, 1997. 10.1016/S0014-5793(96)01404-4 PubMed DOI

Lichtenthaler H.K., Zeidler J., Schwender J., Müller C.: The non-mevalonate isoprenoid biosynthesis of plants as a test-system for new herbicides and drugs against pathogenic bacteria and the malaria parasite. – Z. Naturforsch. C 55: 305-313, 2000. 10.1515/znc-2000-5-601 PubMed DOI

Nonomura A.M., Holtz B., Biel K.Y. et al.: The paths of Andrew A. Benson: a radio-autobiography. – Photosynth. Res. 134: 93-105, 2017. 10.1007/s11120-017-0410-y PubMed DOI

Pfister K., Buschmann C., Lichtenthaler H.K.: Inhibition of the photosynthetic electron transport by bentazone. – In: Proceedings of the Third International Congress on Photosynthesis. Vol. 1. Pp. 675-681. Elsevier, New York: 1975.

Rao K.K.: Obituary of David Hall (1935-1999). – Photosynth. Res. 62: 117-119, 1999. https://www.life.illinois.edu/govindjee/history/obit/ObitDavidHall.pdf

Rebeiz C.A., Benning C., Bohnert H. et al.: Govindjee was honored with the First Lifetime Achievement Award, and Britta Förster and coworkers, with the First Annual Paper Prize of the Rebeiz Foundation for Basic Research. – Photosynth. Res. 94: 147-151, 2007. 10.1007/s11120-007-9229-2 DOI

Rinderle U., Lichtenthaler H.K.: The various chlorophyll fluorescence signatures as a basis for physiological ground truth control in remote sensing of forest decline. – In: 12th Canadian Symposium on Remote Sensing Geoscience and Remote Sensing, Vancouver, BC, Canada. Vol. 2. Pp. 674-677. IEEE, Vancouver: 1989. 10.1109/IGARSS.1989.578925 DOI

Rock B.N., Hoshizaki T., Lichtenthaler H.K., Schmuck G.: Comparison of in situ spectral measurements of forest decline symptoms in Vermont (U.S.A.) and the Schwarzwald (F.R.G.). – In: Proceedings of the International Geoscience and Remote Sensing Symposium, 1986, Zürich. Vol. 3. Pp. 1667-1672. ESA Publications Division, Noordwijk: 1986.

Šantrůček J., Šiffel P., Lang M. et al.: Photosynthetic activity and chlorophyll fluorescence parameters in aurea and green forms of Nicotiana tabacum. – Photosynthetica 27: 529-543, 1992. https://kramerius.lib.cas.cz/view/uuid:fcb05b20-1125-4632-b113-863d3f418fe3?page=uuid:838449a5-c58a-4482-b5d0-1e3b1846d804

Seybold A., Egle K.: Lichtfeld und Blattfarbstoffe I. [Light field and leaf pigments I.] – Planta 26: 491-496, 1937. [In German] 10.1007/BF01914537 DOI

Sharkey T.D., Govindjee G.: Hartmut Lichtenthaler: an authority on chloroplast structure and isoprenoid biochemistry. – Photosynth. Res. 128: 117-123, 2016. 10.1007/s11120-015-0211-0 PubMed DOI

Šiffel P., Šantrůček J., Lang M. et al.: Age dependence of photosynthetic activity, chlorophyll fluorescence parameters and chloroplast ultrastructure in aurea and green forms of Nicotiana tabacum Su/su mutant. – Photosynthetica 29: 81-94, 1993. https://kramerius.lib.cas.cz/view/uuid:624b0c81-200a-4dc2-a90d-8c88a301ec75?page=uuid:5ceae01d-8bdc-4b6f-99d8-47dfa6e817b8

Stirbet A., Lazár D., Guo Y., Govindjee G.: Photosynthesis: basics, history and modelling. – Ann. Bot.-London 126: 511-537, 2020. 10.1093/aob/mcz171 PubMed DOI PMC

Wright S.W., Hallegraeff G.M., Mantoura R.F.C.: Shirley W. Jeffrey, 1930-2014. – In: Biographical Memoirs. Pp. 18. National Academy of Sciences, 2015. https://www.nasonline.org/publications/biographical-memoirs/memoir-pdfs/jeffrey-shirley.pdf

Zeidler J.G., Lichtenthaler H.K., May H.U., Lichtenthaler F.W.: Is isoprene emitted by plants synthesized via the novel isopentenyl pyrophosphate pathway? – Z. Naturforsch. C 52: 15-23, 1997. 10.1515/znc-1997-1-204 DOI