John Raven, FRS, FRSE: a truly great innovator in plant physiology, photosynthesis and much more
Authors: Larkum A. W. D. , Falkowski P. G. , Edwards Dianne , Osmond C. B. , Lambers H. , Sanchez-Baracaldo P. , Ritchie R. J. , Runcie J. W. , Ralph P. J. , Westoby M. , Maberly S. , Griffiths H. , Smith F. A. , Beardall J. 
Published: February 2025

Abstract
This is a tribute to a truly inspirational plant biologist, Prof. John A. Raven, FRS, FRSE (25th June 1941– 23rd May 2024), who died at the age of 82. He was a leader in the field of evolution and physiology of algae and land plants. His research touched on many areas including photosynthesis, ion transport, carbon utilisation, mineral use, such as silicon, iron and molybdenum, the evolution of phytoplankton, the evolution of root systems, the impact of global change, especially on the acidification of the oceans, carbon gain and water use in early land plants, and ways of detecting extraterrestrial photosynthesis. Beginning his research career in the Botany School, University of Cambridge, John studied ion uptake in a giant algal cell. This was at the time of great strides brought about by Peter Mitchell (1920–1992) in elucidating the role of energy generation in mitochondria and chloroplasts and the coupling of ion transport systems to energy generation. With Enid MacRobbie and Andrew Smith, John pioneered early work on the involvement of ion transport in the growth and metabolism of plant cells.On leaving Cambridge John took up a lectureship at the University of Dundee in 1971, where he was still attached upon his death. His primary focus over the years, with one of us (Paul Falkowski), was on phytoplankton, the photosynthetic microalgae of the oceans. Still, his publication list of 5 books and over 600 scientific papers spans a very broad range. The many highly cited papers (see Table 1) attest to an outstanding innovator, who influenced a multitude of students and coworkers and a very wide readership worldwide. At the personal level, John Raven was a wonderful human being; he had an extraordinary memory, dredging up facts and little-known scientific papers, like a scientific magician, but at the same time making humorous jokes and involving his colleagues in fun and sympathetic appreciation.
Table 1

Ten best cited articles (from google scholar)

Citations
Date

Aquatic Photosynthesis, 3rd EditionP.G. Falkowski & J.A. RavenPrinceton University Press, 2013

3854

2013

The evolution of modern eukaryotic phytoplanktonP.G. Falkowski, M.E. Katz, A.H. Knoll, A. Quigg, J.A. Raven, et alScience 305, 354–360

1790

2004

CO2 concentrating mechanisms in algae:mechanisms, environmental modulation, and evolutionM. Giordano, J. Beardall & J.A. RavenAnnu. Rev. Plant Biol. 56 (1), 99–131

1648

2005

Algae as nutritional food sources: revisiting our understandingM.L. Wells, P. Potin, J.S. Craigie, J.A. Raven, S.S. Merchant, et alJournal of applied phycology 29, 949–982

1527

2017

Plant Nutrient acquisition strategies change with soil ageH. Lambers, J.A. Raven, G.R. Shaver & S.E. SmithTrends in ecology & evolution 23, 95–103

1488

2008

Ocean acidification due to increasing atmospheric carbon dioxideJ. Raven, K. Caldeira, H. Elderfield, O. Hoegh-Guldberg, P. Liss, et alThe Royal Society, Policy Document, June 2005

1470

2005

Phytoplankton in a changing world: cell size and elemental stoichiometryZ.V. Finkel, J. Beardall, K.J. Flynn, A. Quigg, T.A.V. Rees & J.A. RavenJournal of plankton research 32, 119–137

1198

2010

Opportunities for improving phosphorus efficiency in crop plantsE.J. Veneklaas, H. Lambers, J. Bragg, P.M. Finnegan, C.E. Lovelock, et alNew phytologist 195, 306–320

951

2012

Adaptation of unicellular algae to irradiance: an analysis of strategiesK. Richardson, J. Beardall & J.A. RavenNew Phytologist 93, 157–191

914

1983

Nitrogen assimilation and transport in vascular land plants in relation to Intracellular pH regulationJ.A. Raven & F.A. SmithNew Phytologist 76, 415–431

893

1976

Temperature and algal growthJ.A. Raven & R.J. GeiderNew phytologist 110, 441–461

867

1988

The role of trace metals in photosynthetic electron transport in O2 -evolving organismsJ.A. Raven, M.C.W. Evans & R.E. KorbPhotosynthesis Research 60, 111–150

840

1999