Genotypic, Developmental and Drought-Induced Differences in Root Hydraulic Conductance of Contrasting Sugarcane Cultivars

Hydraulic properties of entire root systems and isolated rootsof three contrasting sugarcane clones were evaluated using transpiration-induceddifferences in hydrostatic pressure across intact root systems,root pressure-generated xylem sap exudation, and pressure-fluxrelationships. Regardless of the measurement technique employed,the clones were ranked in the same order on the basis of theirleaf area–specific total root system hydraulic conductance(C_root). All methods employed detected large developmental changesin G_rootroot with maximum values occurring in plants with approximately02 m₂ total leaf area. Genotypic ranking according to G_root,was reflected as a similar ranking according to root length-specifichydraulic conductance (L) of individual excised roots. Genotypicdifferences in G_root and L were consistent with anatomical characteristicsobserved in individual roots. Patterns of G_root, during soildrying and following re-irrigation suggested that the declinein G_root, observed during soil drying occurred within the rootsrather than at the soil–root interface and may have beencaused in part by xylem cavitation in the roots. Key words: Root hydraulic conductance, Saccharum spp, transpiration, root pressure, pressure-flux     

New evidence for large negative xylem pressures and their measurement by the pressure chamber method

Pressure probe measurements have been interpreted as showing that xylem pressures below c. –0.4 MPa do not exist and that pressure chamber measurements of lower negative pressures are invalid. We present new evidence supporting the pressure chamber technique and the existence of xylem pressures well below –0.4 MPa. We deduced xylem pressures in water-stressed stem xylem from the following experiment: (1) loss of hydraulic conductivity in hydrated stem xylem (xylem pressure = atmospheric pressure) was induced by forcing compressed air into intact xylem conduits; (2) loss of hydraulic conductivity from cavitation and embolism in dehydrating stems was measured, and (3) the xylem pressure in dehydrated stems was deduced as being equal and opposite to the air pressure causing the same loss of hydraulic conductivity in hydrated stems. Pressures determined in this way are only valid if cavitation was caused by air entering the xylem conduits (air-seeding). Deduced xylem pressure showed a one-to-one correspondence with pressure chamber measurements for 12 species (woody angiosperms and gymnosperms); data extended to c. –10 MPa. The same correspondence was obtained under field conditions in Betula occidentalis Hook., where pressure differences between air- and water-filled conduits were induced by a combination of in situ xylem water pressure and applied positive air pressure. It is difficult to explain these results if xylem pressures were above –0.4 MPa, if the pressure chamber was inaccurate, and if cavitation occurred by some mechanism other than air-seeding. A probable reason why the pressure probe does not register large negative pressures is that, just as cavitation within the probe limits its calibration to pressures above c. –0.5 MPa, cavitation limits its measurement range in situ.     

Symplast Volume, Turgor, Stomatal Conductance and Growth in Relation to Osmotic and Elastic Adjustment in Droughted Sugarcane1

Leaf water relations, stomatal conductance (g) and shoot growthrate (SGR) were monitored during a soil drying cycle in threesugarcane cultivars growing in pots in a greenhouse. The pressure-volumetechnique was used to evaluate diurnal and droughtinduced variationin leaf water relations characteristics. Leaf solute contentand bulk elasticity varied diurnally in both irrigated and droughtedplants and were highest at midday. Solute accumulation and increasedelasticity were also observed as leaf water deficits developedmore slowly during soil drying. This osmotic and elastic adjustmentmaintained symplast volume essentially constant both diurnallyand during soil drying, whereas turgor was only partially maintained.The extent of osmotic adjustment associated with drought wasnot reflected in the leaf osmotic potential at full turgor becausethe concurrent increase in tissue elasticity resulted in a largersymplast volume at full turgor. Cultivar responses over therange of leaf water deficits imposed did not provide conclusiveevidence for genotypic variation in osmotic and elastic adjustment.It appeared that behavioural differences in rates of water usemay have determined the magnitude of osmotic and elastic adjustmentin response to drought. In the early stages of soil drying,reductions in SGR and g were not accompanied by significantreductions in bulk leaf water status. This suggested that otherfactors, presumably signals originating from the roots, mayhave regulated SGR and g.     

Physiological and environmental regulation of interannual variability in CO2 exchange on rangelands in the western United States

For most ecosystems, net ecosystem exchange of CO₂ (NEE) varies within and among years in response to environmental change. We analyzed measurements of CO₂ exchange from eight native rangeland ecosystems in the western United States (58 site‐years of data) in order to determine the contributions of photosynthetic and respiratory (physiological) components of CO₂ exchange to environmentally caused variation in NEE. Rangelands included Great Plains grasslands, desert shrubland, desert grasslands, and sagebrush steppe. We predicted that (1) week‐to‐week change in NEE and among‐year variation in the response of NEE to temperature, net radiation, and other environmental drivers would be better explained by change in maximum rates of ecosystem photosynthesis (A_max) than by change in apparent light‐use efficiency (α) or ecosystem respiration at 10 °C (R₁₀) and (2) among‐year variation in the responses of NEE, A_max, and α to environmental drivers would be explained by changes in leaf area index (LAI). As predicted, NEE was better correlated with A_max than α or R₁₀ for six of the eight rangelands. Week‐to‐week variation in NEE and physiological parameters correlated mainly with time‐lagged indices of precipitation and water‐related environmental variables, like potential evapotranspiration, for desert sites and with net radiation and temperature for Great Plains grasslands. For most rangelands, the response of NEE to a given change in temperature, net radiation, or evaporative demand differed among years because the response of photosynthetic parameters (A_max, α) to environmental drivers differed among years. Differences in photosynthetic responses were not explained by variation in LAI alone. A better understanding of controls on canopy photosynthesis will be required to predict variation in NEE of rangeland ecosystems.     

Intra- and inter-plant variation in xylem cavitation in Betula occidentalis

A modified version of a method that uses positive air pressures to determine the complete cavitation response of a single axis is presented. Application of the method to Betula occidentalis Hook, gave a cavitation response indistinguishable from that obtained by dehydration, thus verifying the technique and providing additional evidence that cavitation under tension occurs by air entry through interconduit pits. Incidentally, this also verified pressure-bomb estimates of xylem tension and confirmed the existence of large (i.e. >0·4 MPa) tensions in xylem, which have been questioned in recent pressure-probe studies. The air injection method was used to investigate variation within and amongst individuals of B. occidentalis. Within an individual, the average cavitation tension increased from 0·66±0·27 MPa in roots (3·9 to 10·7 mm diameter), to 1·17±0·10 MPa in trunks (12 to 16 mm diameter), to 1·36±0·04 MPa in twigs (3·9 to 5 mm diameter). Cavitation tension was negatively correlated with the hydraulically weighted mean of the vessel diameter, and was negatively correlated with the conductance of the xylem per xylem area. Native cavitation was within the range predicted from the measured cavitation response and in situ maximum xylem tensions: roots were significantly cavitated compared with minimal cavitation in trunks and twigs. Leaf turgor pressure declined to zero at the xylem tensions predicted to initiate cavitation in petiole xylem (1·5 MPa). Amongst individuals within B. occidentalis, average cavitation tension in the main axis varied from 0·90 to 1·90 MPa and showed no correlation with vessel diameter. The main axes of juveniles (2–3 years old) had significantly narrower vessel diameters than those of adults, but there was no difference in the average cavitation tension. However, juvenile xylem retained hydraulic conductance to a much higher xylem tension (3·25 MPa) than did adult xylem (2·25 MPa), which could facilitate drought survival during establishment.