Growth and maintenance respiration in leaves of northern red oak seedlings and mature trees after 3 years of ozone exposure

A two-component model of growth and maintenance respiration is used to study the response of northern red oak (Quercus rubra L.) seedlings and 32-year-old trees to sub-ambient (10 μmol h; cumulative dose based on 7 h daily mean), ambient (43 μmol h), and twice-ambient (85 μmolh) ozone. The relative growth rates (RGR) of leaves sampled from seedlings and trees were similar across treatments, as were specific leaf respiration rates (SRR). Growth coefficients estimated from the SRR versus RGR relationship averaged 25-3 mol CO2 kg^?1 leaf dry mass produced for seedlings and 21-5 mol kg^?1 for trees. Maintenance coefficients ranged from 0-89 to 1-07 mol CO₂ kg^?1 leaf dry mass d^?1 for seedlings and from 0-64 to 0-84 mol kg-1 d^?1 for trees. Neither coefficient was affected by ozone. Leaves sampled throughout the growing season also showed little response of respiration to ozone. This occurred despite a 30% reduction in net photosynthesis for trees grown at twice-ambient ozone. These results suggest that growth and maintenance respiration in young northern red oak leaves are not affected by ozone and that in older leaves injury can occur without a parallel increase in so-called ‘maintenance’ respiration.     

Energetic Costs of Tissue Construction in Yellow-poplar and White Oak Trees Exposed to Long-term CO2Enrichment

Two methods were used to estimate construction costs for leaves,stems, branches and woody roots of yellow-poplar (LiriodendrontulipiferaL.) trees grown at ambient (35 Pa) and elevated (65Pa) CO₂for 2.7 years and trees of white oak (Quercus albaL.)grown at these same CO₂partial pressures for 4 years. Samplecombustion in a bomb calorimeter combined with measurementsof ash and nitrogen content provided the primary method of estimatingtissue construction costs (W_G; g glucose g^-1dry mass). Thesevalues were compared with a second, simpler method in whichcost estimates were derived from tissue ash, carbon and nitrogencontent (V_G). Estimates of W_Gwere lower for leaves, branchesand roots of yellow-poplar and for leaves of white oak grownat elevated compared with ambient CO₂partial pressures. TheseCO₂-induced differences in W_Granged from 3.7% in yellow-poplarroots to 2.1% in white oak leaves. Only in the case of yellow-poplarleaves, however, were differences in V_Gobserved between CO₂treatments.Leaf V_Gwas 1.46 g glucose g^-1dry mass in ambient-grown treescompared with 1.41 g glucose g^-1dry mass for CO₂-enriched trees.Although paired-estimates of W_Gand V_Gclustered about a 1:1 linefor leaves and branches, estimates of V_Gwere consistently lowerthan W_Gfor stems and roots. Construction costs per unit leafarea were 95 g glucose m^-2for yellow-poplar trees grown at ambientCO₂and 106 g glucose m^-2for trees grown at elevated CO₂partialpressures. No differences in area-based construction costs wereobserved for white oak. Whole-plant energy content was 1220g glucose per tree in ambient-grown white oak compared with2840 g glucose per tree for those grown at elevated CO₂partialpressures. These differences were driven largely by CO₂-inducedchanges in total biomass. We conclude that while constructioncosts were lower at elevated CO₂partial pressures, the magnitudeof this response argues against an increased efficiency of carbonuse in the growth processes of trees exposed to CO₂enrichment. Bomb calorimeter; construction costs; elevated CO₂; energy allocation; global change; growth respiration; heat of combustion; respiration; Liriodendron tulipifera; Quercus alba     

Environmental controls on water use efficiency during severe drought in an Ozark Forest in Missouri,USA

To accurately predict ecosystem responses induced by climate warming at local‐to‐global scales, models are in need of more precise knowledge of response during periods of environmental stress such as drought. In this paper, we studied environmental control of canopy‐level water use efficiency (WUE) during drought at an eddy flux site in an oak‐hickory forest in central Missouri, USA. Two consecutive severe droughts in the summers of 2006 and 2007 afforded coverage of a broad range of environmental conditions. We stratified data to obtain subranges that minimized cross‐correlations among putative WUE‐controlling factors. Our results showed that WUE was subject to control by atmospheric saturation deficit (ASD), soil water potential (SWP) and the ratio of diffuse to total photosynthetically active radiation (I_f/I_t). Generally, WUE was found to scale with 1/(ASD)^0.5, consistent with predictions from stomatal optimization theory. In contrast, SWP and I_f/I_t were related to WUE in a linear fashion. ASD was better correlated with WUE than either of the other two factors. It was also observed that the relationship between WUE and any single controlling factor was subject to influence of the other two. One such example was an opposite response of WUE to SWP between low and high ASD values, suggesting a breakdown of stomatal optimality under severe environmental stresses and a shift from optimal stomatal regulation to nonstomatal regulation at leaf scale. We have demonstrated that different data handling (stratified vs. nonstratified) or selection (hourly vs. daily) could lead to different conclusions on the relationship between WUE and its controls. For this reason, we recommend modelers to be cautious when applying WUE‐response formulas at environmental conditions or at time scales different from those at which they are derived.     

Reliable estimation of biochemical parameters from C3 leaf photosynthesis–intercellular carbon dioxide response curves

The Farquhar–von Caemmerer–Berry (FvCB) model of photosynthesis is a change‐point model and structurally overparameterized for interpreting the response of leaf net assimilation (A) to intercellular CO₂ concentration (Ci). The use of conventional fitting methods may lead not only to incorrect parameters but also several previously unrecognized consequences. For example, the relationships between key parameters may be fixed computationally and certain fits may be produced in which the estimated parameters result in contradictory identification of the limitation states of the data. Here we describe a new approach that is better suited to the FvCB model characteristics. It consists of four main steps: (1) enumeration of all possible distributions of limitation states; (2) fitting the FvCB model to each limitation state distribution by minimizing a distribution‐wise cost function that has desirable properties for parameter estimation; (3) identification and correction of inadmissible fits; and (4) selection of the best fit from all possible limitation state distributions. The new approach implemented theoretical parameter resolvability with numerical procedures that maximally use the information content of the data. It was tested with model simulations, sampled A/Ci curves, and chlorophyll fluorescence measurements of different tree species. The new approach is accessible through the automated website leafweb.ornl.gov.     

Temperature-controlled open-top chambers for global change research

To enable experiments on the interactive effects of elevated atmospheric CO₂ and increased air temperature on physiological processes in trees to be carried out, we altered the standard design of open-top chambers by replacing blowers with evaporative coolers and in-line heaters, with a feedback control system to maintain ambient or elevated air temperatures within the chambers. Ambient and elevated (+ 4 °C) temperature regimes were attained consistently and reliably throughout the growing season, with high reproducibility between chambers. From May through December the average of nearly 300,000 temperature measurements was 18.5 °C in ambient air, 18.9 ± 0.6 °C in six ambient chambers, and 22.4 ± 0.9 °C in six elevated temperature chambers. The difference in soil temperature between ambient and elevated chambers was 1.2 °C. Absolute humidity (vapour pressure) in the chambers was higher than that of ambient air, but it was generally similar between temperature treatments. Vapour pressure deficit therefore was higher in elevated temperature chambers than in ambient chambers, and this difference is considered an inseparable part of the temperature treatment. The addition of a temperature control system to open-top chambers removes what has been an important flaw in this important tool for global change research.     

Sensitivity of canopy transpiration to altered precipitation in an upland oak forest: evidence from a long-term field manipulation study

Climate‐induced changes in regional precipitation could have important implications for the carbon, water, and nutrient cycles of forest ecosystems. However, few studies have examined the response of deciduous forests to increases or decreases in precipitation. Therefore, the throughfall displacement experiment (TDE) was established in 1993 near Oak Ridge, Tennessee to examine the sensitivity of an upland oak (Quercus spp.) forest to ambient, wet (+33%), and dry (?33%) precipitation regimes. Sap flux measurements on co‐occurring tree species were scaled using species‐specific estimates of stand sapwood area to derive daily and seasonal rates of canopy transpiration (E_C) from 2000 to 2003. With the exception of 2003, which was an extremely wet year, daily E_C in the dry plot, and occasionally during extended droughts in the ambient and wet plots, declined as water potential in the upper 0.35 m soil profile approached ‐3.0 MPa. Seasonal patterns of soil water potential and treatment‐specific differences in E_C were dependent on precipitation frequency and intensity. Supplemental precipitation added to the wet plot increased seasonal E_C on average by 9% (range ?1% to 19%), whereas extended periods of drought on the dry plot in 2000, 2001, and 2002 were sufficient to reduce seasonal E_C by 26–30% compared with the ambient plot. There was a strong correlation between seasonal E_C and the water stress integral, a cumulative index of drought severity and duration. A polynomial fitted to these data indicated that reductions in seasonal E_C on the order of 40% were possible given TDE‐imposed reductions in soil water potential. Application of this equation to all years of the TDE (1994–2003) revealed considerable interannual and treatment‐specific variation in canopy transpiration. In general, a 33% removal of throughfall on the dry plot during 1995, 1998, and 2002 resulted in a 23–32% reduction in seasonal E_C compared with the ambient plot. While droughts in deciduous forests are often limited in duration and tend to occur late in the growing season, soil water deficits of the magnitude observed in this study have the potential to impact local and regional forest water budgets.     

The citrus leaf cuticle in relation to measurement of leaf water potential using thermocouple psychrometers*

Abstract. Cuticular resistance to water vapour diffusion is an important aspect of thermocouple psychrometry and may introduce significant error in the measurement of leaf water potential (Ψ). The effect of the citrus (Citrus mitis Blanco) leaf cuticle on water vapour movement was studied using the times required for vapour pressure equilibration during thermocouple psychrometric measurement of Ψ. Cuticular abrasion with various carborundum powders was used to reduce the diffusive resistance of both the adaxial and abaxial leaf surfaces, and the extent of the disruption to the leaf was investigated with light and electron microscopy. Cuticular abrasion resulted in reduced equilibration times due to decreased cuticular resistance and greater water vapour movement between the leaf and the psychrometer chamber. Equilibration times were reduced from over 5 h in the unabraded control leaves to 1 h with cuticle abrasion. This was associated with the decrease in diffusive resistance with cuticular abrasion from over 55 s cm^?1 to less than 8 s cm^?1 for both the adaxial and abaxial leaf surfaces. Scanning electron micrographs of the abraded leaf tissue revealed considerable disruption of the stomatal ledge and of the guard cells, surface smoothing and displacement of waxes into the stomatal aperture, and damage to veins. Observations with the transmission electron microscope revealed frequent disruption of epidermal cell walls, and damage to both the cytoplasmic and vacuolar membranes.     

Electron Microscope Study of Cuticular Abrasion on Cotton Leaves in Relation to Water Potential Measurements

Wullschleger, S. D. and Oosterhuis, D. M. 1987. Electron microscopestudy of cuticular abrasion on cotton leaves in relation towater potential measurements.—J. exp. Bot 38: 660–667. Accurate determination of plant water potential using thermocouplepsychrometers requires vapour equilibrium between the tissuesample and the sensing psychrometer junction. Failure to achievethis equilibrium due to cuticular resistances to vapour movementmay introduce significant errors into psychrometrically measured. The effect of cuticular abrasion on W equilibration timesfor cotton (Gossypium hirsutum L.) was studied with three typesof thermocouple psychrometers, and the extent of surface andcuticular damage was determined with electron microscopy. Water vapour equilibration between the leaf and psychrometerchamber was achieved in approximately 4 h for unabraded controls,whereas abrasion of the leaf with carborundum powder consistentlyreduced equilibration times to below 2?5 h for all three typesof psychrometers. Microscopic observation of abraded leaf tissueindicated that substantial damage to surface structures occurredduring the cuticular abrasion process. Scanning electron micrographsrevealed localized cellular damage to anastomosing leaf veinsand physical disruption of both the stomatal complex and glandulartrichomes. Transverse sections viewed with a transmission electronmicroscope indicated substantial direct damage to the cuticlewith large sections of cell wall devoid of a cuticular layer.Although the exposed cell walls were intact, the lateral cellwalls were physically compressed and distorted during abrasion.In addition, the cytoplasmic and vacuolar membranes of the epidermalcells were also frequently ruptured. Evaluation of the damagefollowing abrasion indicated that the release of turgor by theaffected cells may contribute to increased sample variabilityand possibly to errors in measurements. Key words: Leaf water potential, cuticular resistance, thermocouple psychrometer