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.

Photosynthesis and reflectance indices for rainforest species in ecosystems undergoing progression and retrogression along a soil fertility chronosequence in New Zealand.

Measurements of photosynthesis at saturating irradiance and CO2 partial pressure, Amax, "adjusted" normalised difference vegetation index, RaNDVI, and photochemical reflectance index, RPRI, were made on trees sampled along a soil chronosequence to investigate the relationship between carbon uptake and ecosystem development in relation to nutrient availability. Measurements were made on the three most dominant species at six sites along the sequence in South Westland, New Zealand with soil age ranging from < 6 to 120,000 years resulting from the retreat of the Franz Josef glacier. The decrease in soil phosphorus availability with increasing soil age and high soil nitrogen availability at the two youngest sites, due to the presence of a nitrogen-fixing species, provided marked differences in nutrient availability. Mean Amax was high at the two youngest sites, then decreased markedly with increasing site age. Analysis of the data for individual species within sites revealed separation of groups of species in the response of Amax to Nm and Pm, suggesting complex interactions between the two nutrients. There were strong linear relationships for leaf-level RaNDVI and RPRI with Amax, at high irradiance, showing that measurements of reflectance indices can be used to estimate Amax for foliage with a range in morphology and nutrient concentrations. Notwithstanding the change in species composition from angiosperms to conifers with increasing site age, the presence of nitrogen-fixing species, the variability in foliage morphology from flat leaves to imbricate scales and a wide range in foliar nitrogen and phosphorus concentrations, there were strong positive linear relationships between site average Amax and foliage nitrogen, Nm, and phosphorus, Pm, concentrations on a foliage mass basis. The results provide insights to interpret the regulation of photosynthesis across natural ecosystems with marked gradients in nitrogen and phosphorus availability.

Stomatal and non-stomatal limitations to photosynthesis in four tree species in a temperate rainforest dominated by Dacrydium cupressinum in New Zealand.

We assessed the relative limitations to photosynthesis imposed by stomatal and non-stomatal processes in Dacrydium cupressinum Lamb. (Podocarpaceae), which is the dominant species in a native, mixed conifer-broad-leaved rainforest in New Zealand. For comparison, we included three co-occurring broad-leaved tree species (Meterosideros umbellata Cav. (Myrtaceae), Weinmannia racemosa L.f. (Cunoniaceae) and Quintinia acutifolia Kirk (Escalloniaceae)) that differ in phylogeny and in leaf morphology from D. cupressinum. We found that low foliage phosphorus content on an area basis (P(a)) limited light-saturated photosynthesis on an area basis (A(sat)) in Q. acutifolia. Depth in the canopy did not generally affect A(sat) or the relative limitations to A(sat) because of stomatal and non-stomatal constraints, despite reductions in the ratio of foliage mass to area, foliar nitrogen on an area basis (N(a)) and P(a) with depth in the canopy. In the canopy-dominant conifer D. cupressinum, A(sat) was low, consistent with low values of the maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(cmax)). In comparison, the A(sat) response of the three broad-leaved tree species was quite variable. Although A(sat) was high in the canopy-dominant M. umbellata, it was low in the sub-canopy trees W. racemosa and Q. acutifolia. Relative stomatal limitation to photosynthesis was more pronounced in W. racemosa (40%) than in the other three species (28-33%). Despite differences in degree, non-stomatal limitation to A(sat) predominated in all tree species.

Respiration characteristics in temperate rainforest tree species differ along a long-term soil-development chronosequence.

We measured the response of dark respiration (R(d)) to temperature and foliage characteristics in the upper canopies of tree species in temperate rainforest communities in New Zealand along a soil chronosequence (six sites from 6 years to 120,000 years). The chronosequence provided a vegetation gradient characterised by significant changes in soil nutrition. This enabled us to examine the extent to which changes in dark respiration can be applied across forest biomes and the utility of scaling rules in whole-canopy carbon modelling. The response of respiration to temperature in the dominant tree species differed significantly between sites along the sequence. This involved changes in both R(d) at a reference temperature (R(10)) and the extent to which R(d) increased with temperature (described by E(o), a parameter related to the energy of activation, or the change in R(d) over a 10 degrees C range, Q(10)). Site averaged E(o) ranged from 44.4 kJ mol(-1) K(-1) at the 60-year-old site to 26.0 kJ mol(-1) K(-1) at the oldest, most nutrient poor, site. Relationships between respiratory and foliage characteristics indicated that both the temperature response of respiration (E(o) or Q(10)) and the instantaneous rate of respiration increased with both foliar nitrogen and phosphorus content. The ratio of photosynthetic capacity (Whitehead et al. in Oecologia 2005) to respiration (A(max)/R(d)) attained values in excess of 15 for species in the 6- to 120-year-old sites, but thereafter decreased significantly to around five at the 120,000-year-old site. This indicates that shoot carbon acquisition is regulated by nutrient limitations in the retrogressing ecosystems on the oldest sites. Our findings indicate that respiration and its temperature response will vary according to soil age and, therefore, to soil nutrient availability and the stage of forest development. Thus, variability in respiratory characteristics for canopies should be considered when using models to integrate respiration at large spatial scales.