Inter-tracheid pitting and the hydraulic efficiency of conifer wood: the role of tracheid allometry and cavitation protection.

Plant xylem must balance efficient delivery of water to the canopy against protection from air entry into the conduits via air-seeding. We investigated the relationship between tracheid allometry, end wall pitting, safety from air-seeding, and the hydraulic efficiency of conifer wood in order to better understand the trade-offs between effective transport and protection against air entry. Root and stem wood were sampled from conifers belonging to the Pinaceae, Cupressaceae, Podocarpaceae, and Araucariaceae. Hydraulic resistivity of tracheids decreased with increasing tracheid diameter and width, with 64 ± 4% residing in the end wall pitting regardless of tracheid size or phylogenetic affinity. This end-wall percentage was consistent with a near-optimal scaling between tracheid diameter and length that minimized flow resistance for a given tracheid length. There was no evidence that tracheid size and hydraulic efficiency were constrained by the role of the pits in protecting against cavitation by air-seeding. An increase in pit area resistance with safety from cavitation was observed only for species of the northern hemisphere (Pinaceae and Cupressaceae), but this variable was independent of tracheid size, and the increase in pit resistance did not significantly influence tracheid resistance. In contrast to recent work on angiosperm vessels, protection against air-seeding in conifer tracheids appears to be uncoupled from conduit size and conducting efficiency.

[1]  J. Sperry,et al.  Scaling of angiosperm xylem structure with safety and efficiency. , 2006, Tree physiology.

[2]  T. Dawson,et al.  Hydraulic efficiency and safety of branch xylem increases with height in Sequoia sempervirens (D. Don) crowns. , 2006, Plant, cell & environment.

[3]  J. Sperry,et al.  Inter‐vessel pitting and cavitation in woody Rosaceae and other vesselled plants: a basis for a safety versus efficiency trade‐off in xylem transport , 2005 .

[4]  J. Sperry,et al.  Comparative analysis of end wall resistivity in xylem conduits , 2005 .

[5]  J. Sperry,et al.  Analysis of circular bordered pit function II. Gymnosperm tracheids with torus-margo pit membranes. , 2004, American journal of botany.

[6]  Nicanor Z. Saliendra,et al.  Influence of leaf water status on stomatal response to humidity, hydraulic conductance, and soil drought in Betula occidentalis , 1995, Planta.

[7]  W. Liese,et al.  The morphological variability of the bordered pit membranes in gymnosperms , 1972, Wood Science and Technology.

[8]  W. Liese,et al.  On the closure of bordered pits in conifers , 1967, Wood Science and Technology.

[9]  J. Sperry,et al.  Tracheid diameter is the key trait determining the extent of freezing-induced embolism in conifers. , 2003, Tree physiology.

[10]  B. Choat,et al.  Pit Membrane Porosity and Water Stress-Induced Cavitation in Four Co-Existing Dry Rainforest Tree Species , 2003, Plant Physiology.

[11]  J. Domec,et al.  How do water transport and water storage differ in coniferous earlywood and latewood? , 2002, Journal of experimental botany.

[12]  N. Holbrook,et al.  Hydraulic and photosynthetic co‐ordination in seasonally dry tropical forest trees , 2002 .

[13]  I. Oliveras,et al.  Xylem hydraulic properties of roots and stems of nine Mediterranean woody species , 2002, Oecologia.

[14]  A. R. Ennos,et al.  Modelling the hydrodynamic resistance of bordered pits. , 2002, Journal of Experimental Botany.

[15]  Stefan Mayr,et al.  Winter-drought induced embolism in Norway spruce (Picea abies) at the Alpine timberline. , 2002, Physiologia plantarum.

[16]  N. Holbrook,et al.  Hydraulic properties of individual xylem vessels of Fraxinus americana. , 2001, Journal of experimental botany.

[17]  J. Sperry,et al.  Xylem Cavitation and Freezing in Conifers , 2001 .

[18]  Michael G. Ryan,et al.  Stomatal conductance and photosynthesis vary linearly with plant hydraulic conductance in ponderosa pine , 2001 .

[19]  J. Palmer,et al.  Seed plant phylogeny inferred from all three plant genomes: monophyly of extant gymnosperms and origin of Gnetales from conifers. , 2000, Proceedings of the National Academy of Sciences of the United States of America.

[20]  J. Sperry,et al.  Vulnerability to xylem cavitation and the distribution of Sonoran Desert vegetation. , 1996, American journal of botany.

[21]  D. Ellsworth,et al.  Influence of soil porosity on water use in Pinus taeda , 2000, Oecologia.

[22]  J. Sperry,et al.  The relationship between xylem conduit diameter and cavitation caused by freezing. , 1999, American journal of botany.

[23]  R. Hill,et al.  The importance of xylem constraints in the distribution of conifer species , 1999 .

[24]  R. Oren,et al.  Sap-flux-scaled transpiration responses to light, vapor pressure deficit, and leaf area reduction in a flooded Taxodium distichum forest. , 1999, Tree physiology.

[25]  R. Hill,et al.  The photosynthetic drought physiology of a diverse group of southern hemisphere conifer species is correlated with minimum seasonal rainfall , 1998 .

[26]  R. Hill,et al.  Light response characteristics of a morphologically diverse group of southern hemisphere conifers as measured by chlorophyll fluorescence , 1997, Oecologia.

[27]  William T. Pockman,et al.  Use of centrifugal force in the study of xylem cavitation , 1997 .

[28]  William T. Pockman,et al.  Sustained and significant negative water pressure in xylem , 1995, Nature.

[29]  J. A. Jarbeau,et al.  The mechanism of water‐stress‐induced embolism in two species of chaparral shrubs , 1995 .

[30]  J. Garrec,et al.  The growth and gas exchange response of soil-planted Norway spruce [Picea abies (L.) Karst.] and red oak (Quercus rubra L.) exposed to elevated CO2 and to naturally occurring drought. , 1995, The New phytologist.

[31]  J. Mauseth,et al.  Resin‐casting: a method for investigating apoplastic spaces , 1994 .

[32]  J. Sperry Water Transport in Plants under Climatic Stress: Winter xylem embolism and spring recovery in Betula cordifolia, Fagus grandifolia, Abies balsamea and Picea rubens , 1993 .

[33]  A. M. Lewis MEASURING THE HYDRAULIC DIAMETER OF A PORE OR CONDUIT. , 1992, American journal of botany.

[34]  M. Tyree,et al.  Use of positive pressures to establish vulnerability curves : further support for the air-seeding hypothesis and implications for pressure-volume analysis. , 1992, Plant physiology.

[35]  XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF COSTA RICAN BLECHNUM FERNS , 1990 .

[36]  Melvin T. Tyree,et al.  Water‐stress‐induced xylem embolism in three species of conifers , 1990 .

[37]  A. Tyree,et al.  Vulnerability of Xylem to Cavitation and Embolism , 1989 .

[38]  M. Hipkins,et al.  Gas penetration of pit membranes in the xylem of Rhododendron as the cause of acoustically detectable sap cavitation , 1985 .

[39]  John Finn Siau,et al.  Transport Processes in Wood , 1984, Springer Series in Wood Science.

[40]  M. Zimmermann Xylem Structure and the Ascent of Sap , 1983, Springer Series in Wood Science.

[41]  T. Sharkey,et al.  Stomatal conductance and photosynthesis , 1982 .

[42]  J. Petty The aspiration of bordered pits in conifer wood , 1972, Proceedings of the Royal Society of London. Series B. Biological Sciences.