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A tribute to Robert John Porra (august 7, 1931–may 16, 2019)


Robert John Porra (7.8.1931–16.5.2019) is probably best known for his substantial practical contributions to plant physiology and photosynthesis by addressing the problems of both the accurate spectroscopic estimation and the extractability of chlorophylls in many organisms. Physiological data and global productivity estimates, in particular of marine primary productivity, are often quoted on a chlorophyll basis. He also made his impact by work on all stages of tetrapyrrole biosynthesis: he proved the C5 pathway to chlorophylls, detected an alternative route to protoporphyrin in anaerobes and the different origin of the oxygen atoms in anaerobes and aerobes. A brief review of his work is supplemented by personal memories of the authors.

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  • Akutsu H, Park JS, Sano S (1993) l-Methionine methyl is specifically incorporated into the C-2 and C-7 positions of the porphyrin of cytochrome-c(3) in a strictly anaerobic bacterium, Desulfovibrio vulgaris. J Am Chem Soc 115:12185–12186

    CAS  Article  Google Scholar 

  • Arnon PI (1949) Copper enzymes in isolated chloroplasts. Polyphenoloxidase in Beta vulgaris. Plant Physiol 24:1–15

    CAS  Article  Google Scholar 

  • Beale SI, Castelfranco PA, Granick S (1975) The biosynthesis of d-aminolevulinic acid from the intact carbon skeleton of glutamate in greening barley. Proc Natl Acad Sci USA 72:2719–2723

    CAS  Article  Google Scholar 

  • Chow WS, Horton P, Barrett M, Osmond CB (2016) Remembering Joan (Jan) Mary Anderson (1932–2015). Photosynth Res 129:129–146.

    CAS  Article  PubMed  Google Scholar 

  • Esteban R, García-Plazaola JI, Hernández A, Fernández-Marín B (2018) On the recalcitrant use of Arnon’s method for chlorophyll determination. New Phytol 217:474–476.

    Article  PubMed  Google Scholar 

  • Grimm B, Porra R, Rüdiger W, Scheer H (2006) Chlorophylls and bacteriochlorophylls: biochemistry, biophysics, functions and applications. In: Advances in photosynthesis, vol 25. Springer, Dordrecht

  • Jeffrey SW, Humphrey GF (1975) New spectroscopic equations for determining Chlorophylls a, b, c1, c2 in higher plants, algae and natural phytoblankton. Biochem Physiol Pflanzen 167:191–194

    CAS  Article  Google Scholar 

  • Kühner M, Haufschildt K, Neumann A, Storbeck S, Streif J, Layer G (2014) The alternative route to heme in the methanogenic archaeon Methanosarcina barkeri. Archaea 2014:327637–327637.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  • Larkum AWD (1991) Evolution of chlorophylls. In: Scheer H (ed) Chlorophylls. CRC Press, Boca Raton, FL, pp 367–384

    Google Scholar 

  • Larkum AWD (1992) The evolution of chlorophylls, light-harvesting systems and photoreaction centers. In: Murata N (ed) Research in photosynthesis. Kluwer, Dordrecht, pp 475–482

    Chapter  Google Scholar 

  • Porra RJ (1965) Porphyrinogen and corrin biosynthesis in Clostridium tetanomorphum. Biochim Biophys Acta 107:176–179

    CAS  Article  Google Scholar 

  • Porra RJ (1990a) The extraction and assay of refractory Chlorophylls and a simple method to correct data from Arnon’s equations. Curr Res Photosynth.

    Article  Google Scholar 

  • Porra RJ (1990b) The assay of Chlorophylls a and b converted to their respective magnesium-rhodochlorin derivatives by extraction from recalcitrant algal cells with aqueous alkaline methanol: prevention of allomerization with reductants. Biochim Biophys Acta 1015:493–502

    CAS  Article  Google Scholar 

  • Porra RJ (1991) Recent advances and re-assessments in Chlorophyll extraction and assay procedures for terrestrial, aquatic, and marine organisms, including recalcitrant algae. In: Scheer H (ed) Chlorophylls. CRC-Press, Boca Raton, pp 31–58

    Google Scholar 

  • Porra RJ (2002) The chequered history of the development and use of simultaneous equations for the accurate determination of chlorophylls a and b. Photosynth Res 73:149

    CAS  Article  Google Scholar 

  • Porra RJ, Falk JE (1964) The enzymic conversion of coproporphyrinogen III into protoporphyrin IX. Biochem J 90:69–75.

    CAS  Article  PubMed  PubMed Central  Google Scholar 

  • Porra RJ, Grimme LH (1978) Tetrapyrrole biosynthesis in algae and higher plants: a discussion of the importance of the 5-Aminolaevulinate synthase and the Dioxovalerate transaminase pathways in the biosynthesis of Chlorophyll. Int J Biochem 9:883–886

    CAS  Article  Google Scholar 

  • Porra RJ, Jones OTG (1963) Studies on ferrochelatase 1. Assay and properties of ferrochelatase from a pig-liver mitochondrial extract. Biochem J 87:181–185

    CAS  Article  Google Scholar 

  • Porra RJ, Scheer H (2000) 18O and mass spectrometry in chlorophyll research: derivation and loss of oxygen atoms at the periphery of the chlorophyll macrocycle during biosynthesis, degradation and adaptation. Photosynth Res 66:159–175

    CAS  Article  Google Scholar 

  • Porra RJ, Scheer H (2019) Towards a more accurate future for chlorophyll a and b determinations: the inaccuracies of Daniel Arnon’s assay. Photosynth Res 140:215–219.

    CAS  Article  PubMed  Google Scholar 

  • Porra RJ, Klein O, Wright PE (1983) The proof by 13C-nuclear magnetic resonance spectroscopy of the predominance of the C5-pathway over the Shemin pathway in Chlorophyll biosynthesis and the formation of the methyl ester group of Chlorophyll from Glycine. Eur J Biochem 130:509–516

    CAS  Article  Google Scholar 

  • Porra RJ, Thompson WA, Kriedemann PE (1989) 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. Biochim Biophys Acta 975:384–394

    CAS  Article  Google Scholar 

  • Ritchie RJ (2006) Consistent sets of spectrophotometric chlorophyll equations for acetone, methanol and ethanol solvents. Photosynth Res 89:27–41

    CAS  Article  Google Scholar 

  • Ritchie RJ (2008) Universal chlorophyll equations for estimating chlorophylls a, b, c and d and total chlorophylls in natural assemblages of photosynthetic organisms using acetone, methanol or ethanol solvents. Photosynthetica 46:115–126

    CAS  Article  Google Scholar 

  • Ritchie RJ, Sma-Air S (2020a) Using integrating sphere spectrophotometry in unicellular algal research. J Appl Phycol 32:2947–2958.

    CAS  Article  Google Scholar 

  • Ritchie RJ, Sma-Air S (2020b) Solvent-free chlorophyll spectrometry in unicellular algal research. J Appl Phycol 32:2711–2723.

    CAS  Article  Google Scholar 

  • Strid Å, Porra RJ (1992) Alterations in pigment content in leaves of Pisum sativum after exposure to supplementary UV-B. Plant Cell Physiol 33:1015–1023

    CAS  Google Scholar 

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Correspondence to Hugo Scheer.

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Chow, W.S., Larkum, A.W.D., Pfündel, E. et al. A tribute to Robert John Porra (august 7, 1931–may 16, 2019). Photosynth Res 147, 125–130 (2021).

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  • Chlorophyll analytics
  • Oxygen-dependent pathways
  • Stable isotopes
  • Tetrapyrrole biosynthesis