# John Raven, FRS, FRSE: a truly great innovator in plant physiology, photosynthesis and much more

## TL;DR

This paper honors John Raven, a pioneering plant biologist who made significant contributions to understanding photosynthesis, algae evolution, and environmental impacts on plant physiology.

## Contribution

John Raven's novel work includes advancing knowledge on phytoplankton evolution, CO2 concentrating mechanisms in algae, and the effects of ocean acidification.

## Key findings

- Raven's research elucidated the evolution and physiology of phytoplankton and land plants.
- He contributed significantly to understanding carbon utilization and the impact of global change on ocean acidification.
- His work on ion transport and nutrient acquisition strategies in plants has been highly influential.

## 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 1Ten best cited articles (from google scholar)CitationsDateAquatic Photosynthesis, 3rd EditionP.G. Falkowski & J.A. RavenPrinceton University Press, 201338542013The evolution of modern eukaryotic phytoplanktonP.G. Falkowski, M.E. Katz, A.H. Knoll, A. Quigg, J.A. Raven, et alScience 305, 354–36017902004CO2 concentrating mechanisms in algae:mechanisms, environmental modulation, and evolutionM. Giordano, J. Beardall & J.A. RavenAnnu. Rev. Plant Biol. 56 (1), 99–13116482005Algae 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, 949–98215272017Plant Nutrient acquisition strategies change with soil ageH. Lambers, J.A. Raven, G.R. Shaver & S.E. SmithTrends in ecology & evolution 23, 95–10314882008Ocean acidification due to increasing atmospheric carbon dioxideJ. Raven, K. Caldeira, H. Elderfield, O. Hoegh-Guldberg, P. Liss, et alThe Royal Society, Policy Document, June 200514702005Phytoplankton 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, 119–13711982010Opportunities for improving phosphorus efficiency in crop plantsE.J. Veneklaas, H. Lambers, J. Bragg, P.M. Finnegan, C.E. Lovelock, et alNew phytologist 195, 306–3209512012Adaptation of unicellular algae to irradiance: an analysis of strategiesK. Richardson, J. Beardall & J.A. RavenNew Phytologist 93, 157–1919141983Nitrogen assimilation and transport in vascular land plants in relation to Intracellular pH regulationJ.A. Raven & F.A. SmithNew Phytologist 76, 415–4318931976Temperature and algal growthJ.A. Raven & R.J. GeiderNew phytologist 110, 441–4618671988The role of trace metals in photosynthetic electron transport in O2 -evolving organismsJ.A. Raven, M.C.W. Evans & R.E. KorbPhotosynthesis Research 60, 111–1508401999

Ten best cited articles (from google scholar)

Aquatic Photosynthesis, 3rd Edition

P.G. Falkowski & J.A. Raven

Princeton University Press, 2013

The evolution of modern eukaryotic phytoplankton

P.G. Falkowski, M.E. Katz, A.H. Knoll, A. Quigg, J.A. Raven, et al

Science 305, 354–360

CO2 concentrating mechanisms in algae:

mechanisms, environmental modulation, and evolution

M. Giordano, J. Beardall & J.A. Raven

Annu. Rev. Plant Biol. 56 (1), 99–131

Algae as nutritional food sources: revisiting our understanding

M.L. Wells, P. Potin, J.S. Craigie, J.A. Raven, S.S. Merchant, et al

Journal of applied phycology 29, 949–982

Plant Nutrient acquisition strategies change with soil age

H. Lambers, J.A. Raven, G.R. Shaver & S.E. Smith

Trends in ecology & evolution 23, 95–103

Ocean acidification due to increasing atmospheric carbon dioxide

J. Raven, K. Caldeira, H. Elderfield, O. Hoegh-Guldberg, P. Liss, et al

The Royal Society, Policy Document, June 2005

Phytoplankton in a changing world: cell size and elemental stoichiometry

Z.V. Finkel, J. Beardall, K.J. Flynn, A. Quigg, T.A.V. Rees & J.A. Raven

Journal of plankton research 32, 119–137

Opportunities for improving phosphorus efficiency in crop plants

E.J. Veneklaas, H. Lambers, J. Bragg, P.M. Finnegan, C.E. Lovelock, et al

New phytologist 195, 306–320

Adaptation of unicellular algae to irradiance: an analysis of strategies

K. Richardson, J. Beardall & J.A. Raven

New Phytologist 93, 157–191

Nitrogen assimilation and transport in vascular land plants in relation to Intracellular pH regulation

J.A. Raven & F.A. Smith

New Phytologist 76, 415–431

Temperature and algal growth

J.A. Raven & R.J. Geider

New phytologist 110, 441–461

The role of trace metals in photosynthetic electron transport in O2 -evolving organisms

J.A. Raven, M.C.W. Evans & R.E. Korb

Photosynthesis Research 60, 111–150