Transforming photosynthesis: frontier biology for a changing world Colin Osborne (University of Sheffield) Global Food Security in 21st Century Last 50 yrs: • global popn + grain prodn doubled • 1 billion people are hungry Next 40 yrs: • global population to stabilize at 9 bn • 70-100% increase in crop production http://gli.environment.umn.edu We already farm 40% of Earth’s land area http://gli.environment.umn.edu Yield stagnation in major crops Rice http://gli.environment.umn.edu Yield stagnation in major crops Wheat http://gli.environment.umn.edu Photosynthesis sunlight water + carbon dioxide sugars + oxygen “makes life and oxygen out of water and thin air” Cyanobacteria “a small step for a bacterium, but a giant leap for biology” O2 4 H+ + 4 e2 H2O ATP NADPH Cyanobacteria enslaved within plants as chloroplasts www.microscopy-uk.org.uk/mag/artnov00/dwelodea.html Photosynthesis powers life on our planet NASA SeaWIFS http://archive.org/details/SVS-3451 Rubisco: the most abundant protein on Earth Rubisco CO2 sugar Calvin cycle NADPH ATP • Ribulose-1,5-bisphosphate carboxylase/oxygenase • ~5-10 kg for every person on the planet • “C3 photosynthesis” http://cdiac.ornl.gov/trends/co2/sio-mlo.html CO2 in geological time Cretaceous chalk 100 myr ago Carboniferous limestone 300 myr ago CO2 and climate in geological time http://droyer.web.wesleyan.edu/GSA_Today.pdf Rubisco works well at high CO2 … Rubisco CO2 Calvin cycle … but it’s less efficient at low CO2 Rubisco CO2 photosynthesis O2 photorespiration • uses ATP, NADH • releases CO2 • CO2 : O2 in the modern atmosphere = 1 : 500 • photosynthesis : photorespiration = 3 : 1 Evolution of carbon-concentrating mechanisms ATP CO2 CO2 Evolution of carbon-concentrating mechanisms cyanobacteria green algae Evolution of carbon-concentrating mechanisms C4 plants CAM plants The C4 carbon-concentrating mechanism CO2 malate C4 pathway requires anatomical adaptations How many times did C4 photosynthesis evolve? • DNA sequences used to reconstruct tree of life C3 C4 C4 evolved • DNA sequences used to reconstruct tree of life C3 C4 C4 evolved C4 photosynthesis evolved more than 70 times! How did C4 pathway evolve so many times? C4 cycle delivers malate to bundle sheath Enzymes in bundle sheath remove CO2 from malate All plants have metabolic components of C4 cycle Xylem transports malate from roots to photosynthetic cells in stem Enzymes in these cells remove CO2 from malate CO2 is fixed by photosynthesis http://eprints.whiterose.ac.uk/113/1/quickwp1.pdf All plants have metabolic components of C4 cycle Xylem transports malate from roots to photosynthetic cells in stem Enzymes in these cells remove CO2 from malate CO2 is fixed by photosynthesis Evolution builds new machinery from existing parts! Why don’t all plants use C4 photosynthesis? Why are some lineages more likely to evolve C4? Why are some lineages more likely to evolve C4? C4 photosynthesis evolved after CO2 dropped C4 evolved http://www.sciencedirect.com/science/article/pii/S0960982207023445 C4 photosynthesis evolved in hot, sunny places Why don’t all plants use C4 photosynthesis? 1. Only some lineages had the necessary anatomical arrangement for C4 photosynthesis. 2. CO2 drop 30 myr ago drove evolution of C4 photosynthesis in hot, sunny environments. 3. C4 evolution made use of metabolic “parts” already present in every plant. C4 grasses are the most productive plants on Earth Echinochloa polystachya 100 t / ha 8 m water rise Miscanthus giganteus 60 t / ha [wheat ~20 t / ha] Can we harness C4 photosynthesis in agriculture? C4 crops already important for food and fuel maize sugarcane miscanthus Can we harness C4 photosynthesis in agriculture? • Rice uses C3 photosynthesis in hot, sunny climates • It has reached a yield plateau • C4 pathway could boost photosynthesis by 50% C4 Rice Project • Introduce C4 into rice • Discover genes involved • C4 rice crop in 20 years? www.3to4.org www.c4rice.irri.org Combining algal and plant photosynthesis Algal carbon-concentrating mechanisms into plants? http://cambridgecapp.wordpress.com osbornelab.group.shef.ac.uk @sheffieldplants
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