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Xylem structure-function relations and trade-offs in conifers
Hervé Cochard The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012
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Structure Function Environment
The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012
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Xylem structure Trunk structure Ring structure Cells structure
Sapwood/heartwood Ring structure Ring width, density Earlywood/Latewood Compression wood Cells structure Tracheid dimensions (length/width/volume) Ray cells (volume, size, grouping) Resin duck ? Cell wall (ultra) structure Secondary wall structure (thickness, microfibril angle) and composition (chemical, isotopic) Pit number/structure/chemical composition The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 3 3
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Tree environment Abiotic factors Biotic factors
Climate temperature, precipitation, wind, sun, vpd, fire… Soil Depth, structure, texture, fertility, water reserve.. Biotic factors Pathogens Competition Growth period, Water stresses (drought, flooding), Mechanical stresses (wind, snow), Biotic stresses The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 4 4
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Major xylem functions water Transport Storage Mechanical Support
nutriments hormones CO2 carbohydrates Storage C and N reserves heat ? Mechanical Support The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012
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Trade-offs efficiency safety ‘energy’ Transport Storage Mechanics
The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 6 6
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How to proceed with xylem hydraulic functions?
Measure them directly (cf growth, density) Derive them from correlations with xylem structure The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 7 7
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Xylem water transport efficiency
Ks: Specific axial hydraulic conductivity Mayr & Cochard PCE 2003 Can be measured directly Problems with native embolism and heartwood The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 8 8
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efficiency Ks can be derived from tracheids dimensions Interspecific
Intraspecific Domec et al 2006 Pittermann et al 2006 The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 9 9
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efficiency What is the functional significance of Ks? Flux
Pleaf<< 0 Psol < 0 Flux Kroot-soil Kxylem Kleaf Impact of Kradial on Kxylem? Kxylem>>Ktree The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 10
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Xylem water transport safety
Cavitation/Embolism Collapse Utsumi et al 2003 Cochard et al 2004 The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 11 11
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safety P50: Xylem vulnerability to embolism
Can be measured directly (eg, with a Cavitron) The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 12 12
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safety Problems with: native embolism ageing heartwood HC, unpublished
The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 13 13
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safety Can cavitation resistance be derived from xylem structure? Interspecific r²=0.25 Tracheid Lumen area, mm² Pittermann et al 2006 Delzon et al 2010 The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 14 14
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safety Intraspecific (Scot Pine) Sterck et al 2012
The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 15 15
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safety Intra-ring (Douglas) Nano CT Native state -2 MPa -2.5 MPa
Badel et al, unpublished The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 16 16
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safety Looking closer Delzon et al 2010
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safety Looking even closer Jansen, Lamy et al 2012
The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 18 18
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safety Frost resistance Pittermann & Sperry 2003
The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 19 19
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Efficiency versus safety
interspecific The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 20 20
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Efficiency versus safety
Intraspecific (maritime pine) Within a tree (douglas fir) Corcuera et al 2011 The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 21 21
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Safety versus energy Interspecific Delzon et al (in prep)
Hacke et al 2001 Delzon et al (in prep) The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 22 22
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Safety versus energy Intraspecific maritime pine douglas
Dalla Salda et al 2011 Lamy et al (in prep) The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 23 23
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Xylem structure-function relations and trade-offs in conifers
Concluding remarks Xylem structure-function relations and trade-offs in conifers The situation is not (yet) totally clear Lot of noise from techniques and plant material (species, populations, individuals) To assess phenotypic plasticity = same genotypes Data on other xylem functions are largely missing (storage) Data are still largely missing at the ring levels The significance of xylem hydraulic plasticity for reconstructing past environments: Kippel, May 15th – 17th 2012 24 24
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