Beverley Green

Professor Emeritus

Academic History

  • B.Sc.(Hons. Chemistry), Univ. of B.C. 1960; 
  • Ph.D. (Biochemistry) Univ. of Washington, 1965.

Contact Information


Genomics, molecular evolution and chloroplast protein import in algae with chlorophyll c (diatoms, dinoflagellates and other chromists). Evolution of genes for light-harvesting antenna proteins and their stress-induced relatives. Photoacclimation in diatoms. Replication and transcription of minicircular chloroplast genes in dinoflagellates.

Research Interests

All my research is concerned with chloroplasts, the photosynthetic organelles of eukaryotic cells. It brings together the structure of photosynthetic membrane proteins with the evolution of the gene families that encode them, as well as the evolutionary origins of the chloroplasts themselves. We are currently investigating the amazing minicircular genes found in dinoflagellate chloroplasts‹each gene on its own individual minicircle of DNA, “one gene-one circle”! These were the first chloroplast gene sequences ever reported for dinoflagellates, a large group of unicellular marine algae.

Another project concerns how nuclear-encoded proteins are imported into the chloroplasts of chromophyte (Chl a/c-containing) algae, which are bounded by four envelope membranes rather than two. Both these groups of algae are believed to have originated by secondary endosymbiosis, where a non-photosynthetic eukaryote engulfed a photosynthetic eukaryote rather than a prokaryote. We are looking for molecular clues about how this process might have occurred.

Most of my earlier research concerned the extended family of chlorophyll (Chl) a/b and Chl a/c light-harvesting proteins in plants and algae. Now we are looking more closely at the Chl a/c family of the chromophyte alga Heterosigma, to try to determine if there is preferential association of any particular members of this family with Photosystem I or Photosystem II. From phylogenetic trees, it is clear that the Chl a/b and the Chl a/c proteins diverged from each other early in the evolution of the photosynthetic eukaryotic cell, so the organization of the chlorophyll a/c protein complexes in the thylakoid membrane may be quite different.

Selected Publications

Books and Chapters: 

B.R. Green (2007) Evolution of light-harvesting antennas in an oxygen world. In:“Evolution of Aquatic Photoautotrophs”. P.G. Falkowski and A.H. Knoll (Editors). Academic Press, in press.

B.K Chaal and B.R. Green (2007) Protein targeting in “Secondary” or “Complex” Chloroplasts. In: “Protein Targeting Protocols: 2nd Edition”. M. van der Giezen (Editor). Humana Press, in press.

B. R. Green and E. Gantt (2005) Distal Antennas of Photosystem II. In “Photosystem II: The Water/Plastoquinone Oxido-Reductase In Photosynthesis”. T. Wydrzynski and K. Satoh (Editors). Kluwer Academic Press, Dordrecht.

“Light-harvesting Antennas in Photosynthesis” (2003) Eds. B. R. Green and W. W. Parson. Kluwer Academic Press, Dordrecht. Authored two chapters of that book:

B. R. Green, J.A. Anderson, W.W. Parson (2003) Photosynthetic membranes and their light-harvesting antennas. Ibid, pp 1-28

B. R. Green (2003) The evolution of light-harvesting antennas. Ibid, pp129-168

Martha J. Nelson, Yunkun Dang, Elena Filek, Zhaoduo Zhang, Vionnie Yu, Ken-ichiro Ishida and Beverley R. Green. (2007) Identification and transcription of transfer RNA genes in dinoflagellate plastid minicircles. Gene 392: 291-298.

Marie-Pierre Oudot-Le Secq, Jane Grimwood, Harris Shapiro, E.Virginia Armbrust, Chris Bowler and Beverley R. Green. (2007) Chloroplast genomes of the diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana: Comparison with other plastid genomes of the red lineage. Mol. Gen. Gen. 277:427–439.

M. Oborník and B.R. Green. (2005) “Mosaic Origin of the Heme Biosynthesis Pathway in Photosynthetic Eukaryotes”. Mol. Biol. Evol. 22: 2343-2353.

M. J. Nelson and B.R.Green. (2005) “Double hairpin elements and tandem repeats in the non-coding region of Adenoides eludens chloroplast gene minicircles.” Gene 358: 102-110.

P. Juneau, B. R. Green and P. J. Harrison. (2005) Simulation of pulse-amplitude-modulated (PAM) fluorescence: Limitations of some PAM-parameters in studying environmental stresss effects. Photosynthetica 43: 75-83.

B.K. Chaal and B.R. Green. (2005) “Protein import pathways in ‘complex’ chloroplasts derived from secondary endosymbiosis involving a red algal ancestor.” Plant Molecular Biology 57: 333-342.

F. Bentley, J. Harnett, K. Ishida and B.R.Green (2005) “Investigation of PSI-associated light-harvesting proteins in a chromophyte alga.” In: Photosynthesis: Fundamental Aspects to Global Perspectives (Eds. A. van der Est and D. Bruce), pp.161-163.

E.V.Armbrust, J.A.Berges, C. Bowler, B.R. Green, D. Martinez and 38 others. (2004) “The genome of the diatom Thalassiosira pseudonana: ecology, evolution, and metabolism”. Science 306: 79-86

B.R. Green (2004) The chloroplast genome of dinoflagellates – A reduced instruction set? Protist 155: 23-31

E. L. Camm and B.R. Green (2004) How the chlorophyll-proteins got their names. Photosyn Res 80: 189-196

B. K. Chaal, K. Ishida and B.R.Green (2003) A thylakoidal processing peptidase from the heterokont alga Heterosigma akashiwo. Plant Mol Biol 52: 463-472

K. Ishida and B.R. Green (2002) Second- and third-hand chloroplasts in dinoflagellates: Phylogeney of oxygen-evolving enhancer 1 (PsbO) protein reveals replacement of a nuclear-encoded plastid gene by that of a haptophyte tertiary endosymbiont. Proc Natl Acad Sci USA 99: 9294–9299

Z. Zhang, T. Cavalier-Smith and B.R.Green (2002)“Evolution of dinoflagellate unigenic minicircles and the partially concerted divergence of their putative replicon origins.” Mol Biol Evol 19: 489-500

Z. Zhang, T. Cavalier-Smith and B.R. Green (2001) “A family of selfish minicircular chromosomes with jumbled chloroplast gene fragments from a dinoflagellate.” Mol Biol Evol 18:1558-65

B.R. Green (2001) “Was ‘molecular opportunism’ a factor in the evolution of different photosynthetic light-harvesting pigment systems?” Proc. Natl. Acad. Sci. USA 98: 2119-2121

Z. Zhang, B.R. Green and T. Cavalier-Smith (2000) “Phylogeny of Ultra-rapidly Evolving Dinoflagellate Chloroplast Genes: A Possible Common Origin for Spoorozoan and Dinflagellate Plastids.” J. Mol. Evol. 51: 26-40

K. Ishida, T. Cavalier-Smith and B.R.Green (2000) “Endomembrane Structure and the Chloroplast protein Targeting Pathway in Heterosigma akashiwo (Raphidophyceae, Chromista). J. Phycol. 36; 1135-1144

B.R. Green and E. Gantt (2000) “Is Photosynthesis Really Derived from Purple Bacteria?”. J. Phycol.36: 983-985

Z. Zhang, B.R.Green and T. Cavalier-Smith (1999) “Single Gene Circles in Dinoflagellate Chloroplast Genomes” Nature 400: 155-159

K. Ishida, B.R.Green and T. Cavalier-Smith (1999) “Diversification of a Chimaeric Algal Group, the Chlorarachniophytes: Phylogeny of Nuclear and Nucleomorph Small Subunit RRNA Genes.” Mol. Biol. Evol. 16: 321-331.

D.G. Durnford, J.A. Deane, S. Tan, G.I. McFadden, E. Gantt, B.R.Green, (1999) “A Phylogenetic Assessment of Eukaryotic Light-harvesting Antenna Proteins, with implications for Plastid Evolution”, J. Mol. Evol. 48: 59-68.

B.R. Green, S. Singh, I Babic, C. Bladen, A. Johnson-Flanagan, (1998)“Relationship of chlorophyll, seed moisture and ABA levels in the maturing Brassica seed and effect of a mild freezing stress”. Physiol. Plant. 104: 125-133.

B.R. Green (1998) “Solutions to the Light-harvesting Problem: Mix, Match and Duplicate”. (Invited Symposium talk), published in Proceedings of XIth International Congress on Photosynthesis, Budapest, Aug. 1998.

G.W.M. van der Staay, N. Yurkova, B.R.Green (1998) “The 38 kDa Chlorophyll a/b Protein of the Prokaryote Prochlorothrix hollandica is Encoded by a Divergent pcb Gene”, Plant Mol. Biol. 36: 775-783.

R.D.Dinkins, H. Bandaranayake, L. Baeza, A.J.F. Griffiths, B.R. Green (1997) “hcf5, a Nuclear Photosynthetic Electron Transport Mutant of Arabidopsis thaliana with a Pleiotropic Effect on Chloroplast Gene Expression”. Plant Physiol. 113: 1023-1031 (and on the cover!).

B.R. Green and D. Durnford (1996) “Chlorophyll-carotenoid Proteins in Oxygenic Photosynthesis”. Annu. Rev. Plant Physiology, Plant Mol. Biol. 47:685-714.

J. La Roche, G.W.M. van der Staay, F. Partensky, A. Ducret, R. Aebersold, R. Li, S.S.Golden, R.G.Hiller, P.M. Wrench, A.W.D.Larkum, B.R.Green (1996) “Independent Evolution of the Prochlorophyte and Green Plant Chlorophyll a/b Light-harvesting Proteins” Proc. Natl. Acad. Sci. USA 93: 15244-15248.

D.G. Durnford, R. Aebersold, B.R.Green (1996) “The Fucoxanthin-chlorophyll Proteins from a Chromophyte Alga Are Part of a Large Multigene Family: Structural and Evolutionary Relationships to Other Light Harvesting Antennae” Mol. Gen. Genet. 253: 377-386.