The influence of nitrogen and phosphorus supply and genotype on mesophyll conductance limitations to photosynthesis in Pinus radiata.

Mesophyll conductance, g(m), may pose significant limitations to photosynthesis and may be differentially affected by nutrition and genotype in Pinus radiata D. Don. Simultaneous measurements of gas exchange and chlorophyll fluorescence were made to determine g(m), using the constant J method (Harley, P.C., F. Loreto, G. Di Marco and T.D. Sharkey. 1992. Theoretical considerations when estimating the mesophyll conductance to CO(2) flux by analysis of the response of photosynthesis to CO(2). Plant Physiol. 98:1429-1436), in a fast- and a slow-growing clone of P. radiata grown in a greenhouse with a factorial combination of nitrogen (N) and phosphorus (P) supply. Values of g(m) increased linearly with the rate of photosynthesis at saturating irradiance and ambient CO(2) concentration, A(sat) (g(m) = 0.020A(sat), r(2) = 0.25, P < 0.001) and with stomatal conductance to CO(2) transfer, g(s) (g(m) = 1.16g(s), r(2) = 0.14, P < 0.001). Values of g(m) were greater than those of stomatal conductance, g(s), and the ratio (g(m)/g(s)) was not influenced by single or combined N and P additions or clone with a mean (+/-SE) value of 1.22 +/- 0.06. Relative limitations to mesophyll conductance, L(m) (16%) to photosynthesis, were generally greater than those imposed by stomata, L(s) (13%). The mean (+/-SE) CO(2) concentration in the intercellular air spaces (C(i)) was 53 +/- 3 mumol mol(-1) lower than that in the atmosphere (C(a)). Mean (+/-SE) CO(2) concentration in the chloroplasts (C(c)) was 48 +/- 2 mumol mol(-1) lower than C(i). Values of L(s), L(m) and CO(2) diffusion gradients posed by g(s) (C(a) - C(i)) and g(m) (C(i) - C(c)) did not significantly differ with nutrient supply or clone. Mean values of V(cmax) and J(max) calculated on a C(c) basis were 15.4% and 3.1% greater than those calculated on a C(i) basis, which translated into different slopes of the J(max)/V(cmax) relationship (C(c) basis: J(max) = 2.11V(cmax), r(2) = 0.88, P < 0.001; C(i) basis: J(max) = 2.43V(cmax), r(2) = 0.86, P < 0.001). These results will be useful for correcting estimates of V(cmax) and J(max) used to characterize the biochemical properties of photosynthesis for P. radiata.

The influence of N and P supply and genotype on carbon flux and partitioning in potted Pinus radiata plants.

Carbon (C) flux and partitioning responses of Pinus radiata (D. Don) clones to a factorial combination of nitrogen (N) and phosphorus (P) supply were estimated in small trees growing in a greenhouse over 44 weeks. Our objective was to use a C budget approach at the plant level to examine how a factorial combination of N and P additions and genotype modify gross primary production (GPP), net primary production (NPP), absolute C fluxes apportioned to aboveground net primary production (ANPP), aboveground plant respiration (APR), total belowground carbon flux (TBCF) and the partitioning of GPP to ANPP, APR and TBCF. Single N or P additions increased plant NPP and GPP similarly, but their combined effects exceeded those of their individual contributions. Nitrogen and to a lesser extent P additions enhanced carbon-use efficiency (CUE, NPP:GPP) and C partitioning to ANPP at the expense of TBCF. The fraction of GPP partitioned to APR was invariant to N or P additions. The ratio of soil respiration (FS) to TBCF was significantly greater in the low-N low-P addition treatment (61%) than in those treatments with single or combined N and P additions (49%). The slowest growing clone partitioned a significantly smaller fraction of GPP to ANPP (29%) than one of the faster-growing genotypes (33%). This research provides insight into how N and P regulate the C fluxes and partitioning in individual plants. Our results contribute to explaining clonal variation in aboveground growth rates and suggest that greater gains in CUE and partitioning to ANPP occur with addition of N rather than P supply.

Partititioning concurrent influences of nitrogen and phosphorus supply on photosynthetic model parameters of Pinus radiata.

Responses of photosynthesis (A) to intercellular CO(2) concentration (C(i)) were measured in a fast- and a slow-growing clone of Pinus radiata D. Don cultivated in a greenhouse with a factorial combination of nitrogen and phosphorus supply. Stomatal limitations scaled with nitrogen and phosphorus supply as a fixed proportion of the light-saturated photosynthetic rate (18.5%) independent of clone. Photosynthetic rates at ambient CO(2) concentration were mainly in the V(cmax)-limited portion of the CO(2) response curve at low-nitrogen supply and at the transition between V(cmax) and J(max) at high-nitrogen supply. Nutrient limitations to photosynthesis were partitioned based on the ratio of foliage nitrogen to phosphorus expressed on a leaf area basis (N(a)/P(a)), by minimizing the mean square error of segmented linear models relating photosynthetic parameters (V(cmax), J(max), T(p)) to foliar nitrogen and phosphorus concentrations. A value of N(a)/P(a) equal to 23 (mole basis) was identified as the threshold separating nitrogen (N(a)/P(a) < or = 23) from phosphorus (N(a)/P(a) > 23) limitations independent of clones. On an area basis, there were significant positive linear relationships between the parameters, V(cmax), J(max), T(p) and N(a) and P(a), but only the relationships between T(p) and N(a) and P(a) differed significantly between clones. These findings suggest that, in genotypes with contrasting growth, the responses of V(cmax) and J(max) to nutrient limitation are equivalent. The relationships between the parameters V(cmax), J(max), T(p) and foliage nutrient concentration on a mass basis were unaffected by clone, because the slow-growing clone had a significantly greater leaf area to mass ratio than the fast-growing clone. These results may be useful in discriminating nitrogen-limited photosynthesis from phosphorus-limited photosynthesis.

Modelling environmental variation in Young's modulus for Pinus radiata and implications for determination of critical buckling height.

BACKGROUND AND AIMS: Although density-specific stiffness, E/rho, (where E is Young's modulus and rho is wood density) is often assumed constant by the elastic similarity model, and in determination of critical buckling height (H(crit)), few studies have tested this assumption within species. Here this assumption is tested for Pinus radiata growing across an environmental gradient, and theory is combined with data to develop a model of Young's modulus. METHODS: Analyses use an extensive series of environmental plots covering the range of climatic and edaphic conditions over which P. radiata is grown in New Zealand. Reduced major axis regression was used to determine scaling exponents between log-log plots of H(crit) vs. groundline diameter (D), and E/rho vs. D. Path analysis was used to identify significant direct and indirect (through stem slenderness) edaphic and climatic influences on E. KEY RESULTS: Density-specific stiffness exhibited 3-fold variation. As E/rho scaled positively with D, the exponent of 0.95 between H(crit) and D exceeded the assumed value of 0.67 under constant E/rho. The final path analysis model included mean air temperature in early autumn (T(aut)) and slenderness as significant (P < 0.05) positive direct influences on E. Tree leaf area index and T(aut) were indirectly associated with E through their significant (P < 0.05) positive direct relationship with stem slenderness. Young's modulus was most sensitive to T(aut), followed by stem slenderness then leaf area index, and the final model explained 76 % of the variance in E. CONCLUSIONS: The findings suggest that within species E/rho variation may influence H(crit) and the scaling exponent between D and H(crit) so important in assumptions regarding allometric relationships. The model presented may provide a useful means of determining variation in E, E/rho and H(crit) across environmental gradients.