Mesophyll conductance: a third player in the photosynthesis game

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1 Mesophyll conductance: a third player in the photosynthesis game Jaume Flexas The China - EU Workshop on Phenotypic Profiling and Technology Transfer on Crop Breeding, Barcelona

2 Photosynthesis traditionally viewed as a sugar factory with only TWO limiting steps: CO2 supply (STOMATA) Demand function Supply function Photosynthetic machinery functioning SUGAR

3 Photosynthesis as a diffusion process A = gs (Ca Ci) = gm (Ci Cc) Traditionally considered LARGE and CONSTANT But: IT IS NOT SO LARGE...

4 But CO2 is used in photosynthesis INSIDE chloroplasts There is long way accross!!!! CO2 enters leaves through stomata

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10 SUB-STOMATAL CAVITY: Where CO2 initially ARRIVES CHLOROPLASTS: Where photosynthesis OCCURS How to REACH photosynthetic sites?

11 The CO2 path A Ca gb B Cs GAS PHASE gs gias gw LIPIDLIQUID PHASE gliq Cc Ci Ci s gm cw Cc st pm ce pg 50 mm 0.2 mm Flexas et al Plant Cell Environ. 31,

12 Evans et al. (2009) J. Exp. Bot. 60,

13 0,5 0,4 a ab 0,3 A A abc A A bc 0,2 A c c A 0,1 d Se H er m i-d bs ec id uo us D ec id uo us Ev er gr ee Li n ve C rw on or ife ts rs /H or nw or ts se s 0,0 G ra s gs or gm (mol CO2 m-2 s-1) Stomatal conductance Mesophyll conductance Group Flexas et al Plant Science 196, 70-84

14 CO2 assimilation Restricted gm = photosynthetsis inefficiency [CO2] (proxy Ci)

15 STRUCTURAL CANDIDATES FOR SETTING gm RESISTANCES ALONG THE LIQUID PHASE Evans et al. (2009) J. Exp. Bot. 60,

16 Tomàs et al. (submitted)

17 However gm is only partly dependend of leaf anatomy gm (mol CO2 m s bar ) ? LMA (g m ) Flexas et al Plant Cell Environ. 31,

18 ... AND IT IS NOT SO CONSTANT! - Salinity (Bongi & Loreto 1989 Plant Physiol. 90, ) - Leaf ontogeny (Loreto et al Photosynth. Res. 41, ) - Altitude (Kogami et al Plant Cell Environ. 24, ) - Shade (Hanba et al Plant Cell Environ. 25, ) - Drought (Flexas et al Funct. Plant Biol. 29, ) - Temperature (Bernacchi et al Plant Physiol. 130, ) - CO2 (Singsaas et al Plant Cell Environ. 27, 41-50) - O3 (Eichelmann et al Plant Cell Environ. 27, )

19 High CO2 applied Ambient CO2 restored gm (mol CO2 m s ) -2-1 gs (mol CO2 m s ) B 6000 Time (seconds) Flexas et al. (2007) Plant Cell Environ. 30,

20 METABOLIC CANDIDATES FOR RAPID CHANGES IN gm - Carbonic anhydrase (Gillon and Yakir 2000 Plant Physiol. 123, ) - Aquaporins (Flexas et al Plant J. 48, ) - Others (Flexas et al Plant Cell Environ. In press)

21 AQUAPORINS NtAQP1 transformed plants Anti-sense (AS) Control Antisense (CAS) Over-express (O) Control Overexpress (CO)

22 NtAQP1 affects mesophyll conductance to CO2 GENOTYPE gm (mol CO2 m-2 s-1 bar-1) AS a CAS b CO c O d Flexas et al. (2006) Plant J. 48,

23 Aquaporing activity (GATING) under water stress Miyazawa et al Funct. Plant Biol. 35,

24 OVERALL: gm limits photosynthesis as much as stomata and more than biochemistry Species Factor Total photosynthesis limitation (%) SL (%) MCL (%) BL (%) Reference Arabidopsis thaliana Senescent leaves Flexas et al Beta maritima Severe water stress Galmés et al Lavatera maritima Severe water stress Galmés et al Phlomis italica Severe water stress Galmés et al Vitis berlandieri x rupestris Moderate water stress Flexas et al Vitis berlandieri x rupestris Severe water stress Flexas et al Nicotiana sylvestris Rewatering High Light Gallé et al Nicotiana sylvestris Rewatering Mid Light Gallé et al Nicotiana sylvestris Rewatering Low Light Gallé et al Olea europaea Water stress High VPD Pérez-Martín et al Olea europaea Water stress Low VPD Pérez-Martín et al Rhamnus ludovici-salvatoris Shade leaves Mattos et al. unpub. Beta vulgaris Excess Zinc (300 µm) Sagardoy et al. submit. AVERAGE STRESS

25 gm to improve photosynthesis and yield

26 gm to improve photosynthesis and yield

27 gm to improve photosynthesis and yield Barbour et al Plant Cell Environ 33,

28 gm to improve photosynthesis and yield Low gs & High gm High gs & gm Low gs & gm Barbour et al Plant Cell Environ 33,

29 GLOBAL ANALYSIS SHOWS HOW TO IMPROVE WATER USE EFFICIENCY AN / gs ( mol CO2 mol-1) gm / gs (mol CO2 mol ) Flexas et al., unpublished 5

30 Take-home messages Mesophyll conductance (gm) is as FINITE and RAPIDLY VARIABLE as stomatal conductance gm is a really important LIMITATION to photosynthesis having a GENETIC BASIS (structural + metabolic) Hence improving gm is a TARGET to genetically improve photosynthesis In addition, it is a perfect target to improve WUE as well, specially under drought-prone conditions

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