Initial observations of increased requirements for light-energy dissipation in ryegrass (Lolium perenne) when source / sink ratios become high at a naturally grazed free air CO2 enrichment (FACE) site.

Although photosynthetic response to long-term elevated CO2 has been extensively studied, little attention has yet been directed at coordinated adjustments between the use of absorbed light for CO2 fixation, and the dissipation of potentially harmful excess light. In this study, we have performed an initial analysis of photosynthetic light use and excess light dissipation in response to grazing-induced variation in the source / sink ratio in ryegrass (Lolium perenne L.) after 6 years' exposure to Free Air CO2 Enrichment (FACE). Before grazing, when the source / sink ratio was relatively large, significant down-regulation of photosynthetic capacity (Amax) was observed in the FACE leaves compared with control leaves at the same stage of maturity. The decrease in Amax partly offset the direct stimulation of elevated CO2 on light-saturated photosynthesis, and was accompanied by a reduction in photochemical electron flow that was accompanied by a large increase in susceptibility to photoinhibition. This was indicated by large increases in both non-photochemical quenching (NPQ) and the de-epoxidised state of xanthophyll cycle (DEPS), and also by changes in the photochemical reflectance index (PRI). However, no significant increase in the xanthophyll pool size in FACE leaves was observed, despite the apparent large increase in requirements for photodissipation in FACE leaves. After grazing, when the source / sink ratio was relatively small, the CO2 fixation rates in both the FACE and control leaves were, as expected, significantly higher compared with those before grazing, and there was no down-regulation of photosynthetic capacity in the leaves under FACE conditions. In addition, the extent of photodissipation in the FACE and control leaves was not significantly different. Overall, the profile of leaf physiological and biochemical responses supports the hypothesis that the effect of long-term elevated CO2 can be significantly influenced by short-term variation in the source / sink ratio. As the xanthophyll pool size does not change significantly, this poses the question of whether the increased photodissipative demand observed here under even moderately elevated CO2 concentrations may lead to increased plant susceptibility to photoinhibition, and thus to an increased risk of damage to plant function, under conditions of low sink demand. This question clearly deserves further study.

Characteristics of photosynthesis and stomatal conductance in the shrubland species manuka (Leptospermum scoparium) and kanuka (Kunzea ericoides) for the estimation of annual canopy carbon uptake.

Responses of photosynthesis to carbon dioxide (CO2) partial pressure and irradiance were measured on leaves of 39-year-old trees of manuka (Leptospermum scoparium J. R. Forst. & G. Forst.) and kanuka (Kunzea ericoides var. ericoides (A. Rich.) J. Thompson) at a field site, and on leaves of young trees grown at three nitrogen supply rates in a nursery, to determine values for parameters in a model to estimate annual net carbon uptake. These secondary successional species belong to the same family and commonly co-occur. Mean (+/- standard error) values of the maximum rate of carboxylation (hemi-surface area basis) (Vcmax) and the maximum rate of electron transport (Jmax) at the field site were 47.3 +/- 1.9 micromol m(-2) s(-1) and 94.2 +/- 3.7 micromol m(-2) s(-1), respectively, with no significant differences between species. Both Vcmax and Jmax were positively related to leaf nitrogen concentration on a unit leaf area basis, and the slopes of these relationships did not differ significantly between species or between the trees in the field and young trees grown in the nursery. Mean values of Jmax/Vcmax measured at 20 degrees C were significantly lower (P < 0.01) for trees in the field (2.00 +/- 0.05) than for young trees in the nursery with similar leaf nitrogen concentrations (2.32 +/- 0.08). Stomatal conductance decreased sharply with increasing air saturation deficit, but the sensitivity of the response did not differ between species. These data were used to derive parameters for a coupled photosynthesis-stomatal conductance model to scale estimates of photosynthesis from leaves to the canopy, incorporating leaf respiration at night, site energy and water balances, to estimate net canopy carbon uptake. Over the course of a year, 76% of incident irradiance (400-700 nm) was absorbed by the canopy, annual net photosynthesis per unit ground area was 164.5 mol m(-2) (equivalent to 1.97 kg C m(-2)) and respiration loss from leaves at night was 37.5 mol m(-2) (equivalent to 0.45 kg m(-2)), or 23% of net carbon uptake. When modeled annual net carbon uptake for the trees was combined with annual respiration from the soil surface, estimated net primary productivity for the ecosystem (0.30 kg C m(-2)) was reasonably close to the annual estimate obtained from independent mensurational and biomass measurements made at the site (0.22 +/- 0.03 kg C m(-2)). The mean annual value for light-use efficiency calculated from the ratio of net carbon uptake (net photosynthesis minus respiration of leaves at night) and absorbed irradiance was 13.0 mmol C mol(-1) (equivalent to 0.72 kg C GJ(-1)). This is low compared with values reported for other temperate forests, but is consistent with limitations to photosynthesis in the canopy attributable mainly to low nitrogen availability and associated low leaf area index.

Estimating photosynthetic light-use efficiency using the photochemical reflectance index: variations among species.

Recent studies have shown that the photochemical reflectance index (PRI), derived from narrow waveband reflectance at 531 and 570 nm, can be used as a remote measure of photosynthetic light-use efficiency (LUE). However, uncertainty remains as to the consistency of the relationship between PRI and LUE across species. In this study we examined the relationship between the PRI and various photosynthetic parameters for a group of species with varying photosynthetic capacity. At constant irradiance, for the species group as a whole, the PRI was well correlated with LUE (r2=0.58) and with several other photosynthetic parameters, but best correlated with the ratio of carotenoids to chlorophylls contents (Caro / Chl). Despite the interspecific trends observed, determination of light response functions for the PRI in relation to photosynthetic parameters revealed that species-specific relationships were clearly stronger. For example, r2>0.90 for species-level PRI / LUE relationships. Also, the species-specific light-response data show that the magnitude of the PRI can be related to the magnitude of the saturated irradiance and the rate of CO2 uptake. As demonstrated here, a light response function provides a simple yet precise approach for characterising the relationship between the PRI and photosynthetic parameters, which should assist with improved evaluation of the usefulness of the PRI as a generalised measure of LUE.

Analysis of the growth of rimu (Dacrydium cupressinum) in South Westland, New Zealand, using process-based simulation models.

Two process-based models were used to identify the environmental variables limiting productivity in a pristine, mature forest dominated by rimu (Dacrydium cupressinum Sol. ex Lamb.) trees in South Westland, New Zealand. A model of canopy net carbon uptake, incorporating routines for radiation interception, photosynthesis and water balance was used to determine a value for quantum efficiency when climate variables were not limiting. The annual net carbon uptake by the canopy was estimated to be 1.1 kg C m(-2) and the quantum efficiency 22.6 mmol mol quanta(-1). This value of quantum efficiency, combined with other parameters obtainable from the literature, was then used in a model of forest productivity (3-PG), to simulate changes in net productivity and the allocation of carbon to tree components. The model was adjusted to match a measured stem increment of 10.6 Mg ha(-1) over a period of 13 years. To achieve this while maintaining a low, but stable value for leaf area index, it was necessary to set the site fertility rating very low and select high values for the parameters describing the proportional allocation of total carbon to roots. This approach highlighted nutrient availability as the principal constraint on productivity for the ecosystem and identified critical measurements that will be necessary for using the model to predict the effects of climate change on carbon sequestration. The low rates of carbon uptake and productivity are consistent with the low nutrient supply available from the highly leached, acid soils, most likely attributable to frequent saturation and a very shallow aerobic zone.