title: Hydrocarbons Are Essential for Optimal Cell Size, Division, and Growth of Cyanobacteria.
abstract: Cyanobacteria are intricately organized, incorporating an array of internal thylakoid membranes, the site of photosynthesis, into cells no larger than other bacteria. They also synthesize C15-C19 alkanes and alkenes, which results in substantial production of hydrocarbons in the environment. All sequenced cyanobacteria encode hydrocarbon biosynthesis pathways, suggesting an important, undefined physiological role for these compounds. Here, we demonstrate that hydrocarbon-deficient mutants of Synechococcus sp. PCC 7002 and Synechocystis sp. PCC 6803 exhibit significant phenotypic differences from wild type, including enlarged cell size, reduced growth, and increased division defects. Photosynthetic rates were similar between strains, although a minor reduction in energy transfer between the soluble light harvesting phycobilisome complex and membrane-bound photosystems was observed. Hydrocarbons were shown to accumulate in thylakoid and cytoplasmic membranes. Modeling of membranes suggests these compounds aggregate in the center of the lipid bilayer, potentially promoting membrane flexibility and facilitating curvature. In vivo measurements confirmed that Synechococcus sp. PCC 7002 mutants lacking hydrocarbons exhibit reduced thylakoid membrane curvature compared to wild type. We propose that hydrocarbons may have a role in inducing the flexibility in membranes required for optimal cell division, size, and growth, and efficient association of soluble and membrane bound proteins. The recent identification of C15-C17 alkanes and alkenes in microalgal species suggests hydrocarbons may serve a similar function in a broad range of photosynthetic organisms.
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.); djl63@cam.ac.uk.|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.); djl63@cam.ac.uk.|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.); djl63@cam.ac.uk.|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.); djl63@cam.ac.uk.|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.); djl63@cam.ac.uk.|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.); djl63@cam.ac.uk.|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and djl63@cam.ac.uk.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.) djl63@cam.ac.uk.
authors.forename: David J
authors.initials: DJ
authors.lastname: Lea-Smith
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Maite L
authors.identifier: 0000-0002-2980-9749
authors.initials: ML
authors.lastname: Ortiz-Suarez
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Tchern
authors.identifier: 0000-0002-5827-4616
authors.initials: T
authors.lastname: Lenn
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Dennis J
authors.initials: DJ
authors.lastname: Nürnberg
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Laura L
authors.initials: LL
authors.lastname: Baers
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Matthew P
authors.initials: MP
authors.lastname: Davey
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Lucia
authors.initials: L
authors.lastname: Parolini
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Roland G
authors.initials: RG
authors.lastname: Huber
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Charles A R
authors.initials: CA
authors.lastname: Cotton
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Giulia
authors.initials: G
authors.lastname: Mastroianni
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Paolo
authors.initials: P
authors.lastname: Bombelli
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Petra
authors.initials: P
authors.lastname: Ungerer
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Tim J
authors.initials: TJ
authors.lastname: Stevens
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Alison G
authors.identifier: 0000-0001-6511-5704
authors.initials: AG
authors.lastname: Smith
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Peter J
authors.identifier: 0000-0003-2900-098X
authors.initials: PJ
authors.lastname: Bond
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Conrad W
authors.identifier: 0000-0001-7194-9916
authors.initials: CW
authors.lastname: Mullineaux
authors.affiliation: Department of Biochemistry, University of Cambridge, Cambridge CB2 1QW, United Kingdom (D.J.L.-S., L.L.B., C.A.R.C., P.B., C.J.H.).|Centre for Molecular Science Informatics, Department of Chemistry, University of Cambridge, Cambridge CB2 1EW, United Kingdom (M.L.O.-S., P.J.B.).|School of Biological and Chemical Sciences, Queen Mary University of London, London E1 4NS, United Kingdom (T.L., D.J.N., G.M., P.U., C.W.M.).|Department of Plant Sciences, University of Cambridge, Cambridge CB2 3EA, United Kingdom (M.P.D., A.G.S.).|Department of Physics, University of Cambridge, Cambridge CB3 0HE, United Kingdom (L.P.).|Bioinformatics Institute, A*STAR, Singapore 138671 (R.G.H., P.J.B.).|MRC Laboratory of Molecular Biology, Cambridge CB2 0QH, United Kingdom (T.J.S.); and.|National University of Singapore, Department of Biological Sciences, Singapore 117543 (P.J.B.).
authors.forename: Christopher J
authors.initials: CJ
authors.lastname: Howe
mesh_terms: D053898:Biosynthetic Pathways / Q000187:drug effects; D002455:Cell Division / Q000187:drug effects*; D049109:Cell Proliferation / Q000187:drug effects; D006838:Hydrocarbons / Q000494:pharmacology*; D007425:Intracellular Membranes / Q000187:drug effects / Q000378:metabolism; D008051:Lipid Bilayers / Q000378:metabolism; D009154:Mutation / Q000235:genetics; D010788:Photosynthesis / Q000187:drug effects; D046939:Synechocystis / Q000166:cytology* / Q000187:drug effects / Q000254:growth & development* / Q000378:metabolism; D020524:Thylakoids / Q000187:drug effects / Q000378:metabolism
chemical_list: D006838:Hydrocarbons; D008051:Lipid Bilayers