Summary• Vulnerability to cavitation and conductive efficiency depend on xylem anatomy. We tested a large range of structure-function hypotheses, some for the first time, within a single genus to minimize phylogenetic 'noise' and maximize detection of functionally relevant variation.• This integrative study combined in-depth anatomical observations using light, scanning and transmission electron microscopy of seven Acer taxa, and compared these observations with empirical measures of xylem hydraulics.• Our results reveal a 2 MPa range in species' mean cavitation pressure (MCP). MCP was strongly correlated with intervessel pit structure (membrane thickness and porosity, chamber depth), weakly correlated with pit number per vessel, and not related to pit area per vessel. At the tissue level, there was a strong correlation between MCP and mechanical strength parameters, and some of the first evidence is provided for the functional significance of vessel grouping and thickenings on inner vessel walls. In addition, a strong trade-off was observed between xylemspecific conductivity and MCP. Vessel length and intervessel wall characteristics were implicated in this safety-efficiency trade-off.• Cavitation resistance and hydraulic conductivity in Acer appear to be controlled by a very complex interaction between tissue, vessel network and pit characteristics.
Summary• Eudicot angiosperms with greater vulnerability to xylem cavitation tend to have vessels with greater total area of inter-vessel pits, which inspired the 'rare pit' hypothesis: the more pits per vessel, by chance the leakier will be the vessel's single air-seeding pit and the lower the air-seeding threshold for cavitation to spread between vessels.• Here, we demonstrate the feasibility of the hypothesis, using probability theory to model the axial propagation of air through air-injected stems. In the presence of rare, leaky pits, air-seeding pressures through short stems with few vessel ends in series should be low; pressures should increase in longer stems as more end-walls must be breached.• Measurements on three Acer species conformed closely to model predictions, confirming the rare presence of leaky pits. The model indicated that pits air-seeding at or below the mean cavitation pressure (MCP) occurred at similarly low frequencies in all species. Average end-wall air-seeding pressures predicted by the model closely matched species' MCPs.• Differences in species' vulnerability were primarily attributed to differences in frequency of the leakiest pits rather than pit number or area per vessel. Adjustments in membrane properties and extent of pitting per vessel apparently combine to influence cavitation resistance across species.
Summary The ferns comprise one of the most ancient tracheophytic plant lineages, and occupy habitats ranging from tundra to deserts and the equatorial tropics. Like their nearest relatives the conifers, modern ferns possess tracheid‐based xylem but the structure–function relationships of fern xylem are poorly understood. Here, we sampled the fronds (megaphylls) of 16 species across the fern phylogeny, and examined the relationships among hydraulic transport, drought‐induced cavitation resistance, the xylem anatomy of the stipe, and the gas‐exchange response of the pinnae. For comparison, the results are presented alongside a similar suite of conifer data. Fern xylem is as resistant to cavitation as conifer xylem, but exhibits none of the hydraulic or structural trade‐offs associated with resistance to cavitation. On a conduit diameter basis, fern xylem can exhibit greater hydraulic efficiency than conifer and angiosperm xylem. In ferns, wide and long tracheids compensate in part for the lack of secondary xylem and allow ferns to exhibit transport rates on a par with those of conifers. We suspect that it is the arrangement of the primary xylem, in addition to the intrinsic traits of the conduits themselves, that may help explain the broad range of cavitation resistance in ferns.
Vulnerability curves using the 'Cavitron' centrifuge rotor yield anomalous results when vessels extend from the end of the stem segment to the centre ('open-to-centre' vessels). Curves showing a decline in conductivity at modest xylem pressures ('r' shaped) have been attributed to this artefact. We determined whether the original centrifugal method with its different rotor is influenced by open-to-centre vessels. Increasing the proportion of open-to-centre vessels by shortening stems had no substantial effect in four species. Nor was there more embolism at the segment end versus centre as seen in the Cavitron. The dehydration method yielded an 'r' shaped curve in Quercus gambelii that was similar to centrifuged stems with 86% opento-centre vessels. Both 'r' and 's' (sigmoidal) curves from Cercocarpus intricatus were consistent with each other, differing only in whether native embolism had been removed. An 'r' shaped centrifuge curve in Olea europaea was indistinguishable from the loss of conductivity caused by forcing air directly across vessel end-walls. We conclude that centrifuge curves on long-vesselled material are not always prone to the open vessel artefact when the original rotor design is used, and 'r' shaped curves are not necessarily artefacts. Nevertheless, confirming curves with native embolism and dehydration data is recommended.
Summary• The rare pit hypothesis predicts that the extensive inter-vessel pitting in large early-wood vessels of ring-porous trees should render many of these vessels extremely vulnerable to cavitation by air-seeding. This prediction was tested in Quercus gambelii.• Cavitation was assessed from native hydraulic conductivity at field sap tension and in dehydrated branches. Single-vessel air injections gave air-seeding pressures through vessel files; these data were used to estimate air-seeding pressures for inter-vessel walls and pits.• Extensive cavitation occurred at xylem sap tensions below 1 MPa. Refilling occurred below 0.5 MPa and was inhibited by phloem girdling. Remaining vessels cavitated over a wide range to above 4 MPa. Similarly, 40% of injected vessel files air-seeded below 1.0 MPa, whereas the remainder seeded over a wide range exceeding 5 MPa. Inter-vessel walls averaged 1.02 MPa air-seeding pressure, similar and opposite to the mean cavitation tension of 1.22 MPa. Consistent with the rare pit hypothesis, only 7% of inter-vessel pits were estimated to air-seed by 1.22 MPa.• The results confirm the rare pit prediction that a significant fraction of large vessels in Q. gambelii experience high probability of failure by air-seeding.
During vessel evolution in angiosperms, scalariform perforation plates with many slit-like openings transformed into simple plates with a single circular opening. The transition is hypothesized to have resulted from selection for decreased hydraulic resistance. Previously, additional resistivity of scalariform plates was estimated to be smallgenerally 10% or less above lumen resistivity -based on numerical and physical models. Here, using the singlevessel technique, we directly measured the hydraulic resistance of individual xylem vessels. The resistivity of simple-plated lumens was not significantly different from the Hagen-Poiseuille (HP) prediction (+6 Ϯ 3.3% mean deviation). In the 13 scalariform-plated species measured, plate resistivity averaged 99 Ϯ 13.7% higher than HP lumen resistivity. Scalariform species also showed higher resistivity than simple species at the whole vessel (+340%) and sapwood (+580%) levels. The strongest predictor of scalariform plate resistance was vessel diameter (r 2 = 0.84), followed by plate angle (r 2 = 0.60). An equation based on laminar flow through periodic slits predicted single-vessel measurements reasonably well (r 2 = 0.79) and indicated that Baileyan trends in scalariform plate evolution maintain an approximate balance between lumen and plate resistances. In summary, we found scalariform plates of diverse morphology essentially double lumen flow resistance, impeding xylem flow much more than previously estimated.Key-words: ecological wood anatomy; hydraulic conductivity; plant water transport; vessel evolution; xylem flow resistance; xylem transport.Abbreviations: A, cross-sectional lumen area; a, major axis of ellipse; b, minor axis of ellipse; D, vessel diameter; h, width of slit plus bar thickness; k, width of slit/h; L, length of scalariform slit opening; Le, spacing between perforation plates; Lv, length of the vessel (=stem segment measured); p, perimeter of vessel lumen; P, pressure head; rL, resistance of the lumen; RL, resistivity of the lumen; rp, resistance of the plate; Rp, resistivity of the plate; rtotal, resistance of vessel and capillary tube; rcapillary, resistance of capillary tube; rvessel, resistance of vessel; Rv, resistivity of the vessel; t, time; V, volume; m, viscosity of water.
Night-time leaf conductance (gnight) and transpiration may have several adaptive benefits related to plant water, nutrient and carbon relations. Little is known, however, about genetic variation in gnight and whether this variation correlates with other gas exchange traits related to water use and/or native habitat climate. We investigated gnight in 12 natural accessions and three near isogenic lines (NILs) of Arabidopsis thaliana. Genetic variation in gnight was found for the natural accessions, and gnight was negatively correlated with native habitat atmospheric vapour pressure deficit (VPDair), suggesting lower gnight may be favoured by natural selection in drier habitats. However, there were also significant genetic correlations of gnight with daytime gas exchange traits expected to affect plant fitness [i.e. daytime leaf conductance, photosynthesis and intrinsic water-use efficiency (WUEi)], indicating that selection on daytime gas exchange traits may result in indirect selection on gnight. The comparison of three NILs to their parental genotypes identified one quantitative trait locus (QTL) contributing to variation in gnight. Further characterization of genetic variation in gnight within and among populations and species, and of associations with other traits and native habitats will be needed to understand gnight as a putatively adaptive trait.
Increasing clarity of Delta waters, the emergence of harmful algal blooms, the proliferation of aquatic water weeds, and the altered food web of the Delta have brought nutrient dynamics to the forefront. This paper focuses on the sources of nutrients, the transformation and uptake of nutrients, and the links of nutrients to primary producers. The largest loads of nutrients to the Delta come from the Sacramento River with the San Joaquin River seasonally important, especially in the summer. Nutrient concentrations reflect riverine inputs in winter and internal biological processes during periods of lower flow with internal nitrogen losses within the Delta estimated at approximately 30% annually. Light regime, grazing pressure, and nutrient availability influence rates of primary production at different times and locations within the Delta. The roles of the chemical form of dissolved inorganic nitrogen in growth rates of primary producers in the Delta and the structure of the open-water algal community are currently topics of much interest and considerable debate. Harmful algal blooms have been noted since the late 1990s, and the extent of invasive aquatic macrophytes (both submerged and free-floating forms) has increased especially during years of drought. Elevated nutrient loads must be considered in terms of their ability to support this excess biomass. Modern sensor technology and networks are now deployed that make high-frequency measurements of nitrate, ammonium, and phosphate. Data from such instruments allow a much more detailed assessment of the spatial and temporal dynamics of nutrients. Four fruitful directions for future research include utilizing continuous sensor data to estimate rates of primary production and ecosystem respiration, linking hydrodynamic models of the Delta with the transport and fate of dissolved nutrients, studying nutrient dynamics in various habitat types, and exploring the use of stable isotopes to trace the movement and fate of effluent-derived nutrients.
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