Co-ordination of growth, gas exchange and hydraulics define the carbon safety margin in tree species with contrasting drought strategies.

Gas exchange, growth, water transport and carbon (C) metabolism diminish during drought according to their respective sensitivities to declining water status. The timing of this sequence of declining physiological functions may determine how water and C relations compromise plant survival. In this paper, we test the hypothesis that the degree of asynchrony between declining C supply (photosynthesis) and C demand (growth and respiration) determines the rate and magnitude of changes in whole-plant non-structural carbohydrates (NSC) during drought. Two complementary experiments using two tree species (Eucalyptus globulus Labill. and Pinus radiata D. Don) with contrasting drought response strategies were performed to (i) assess changes in radial stem growth, transpiration, leaf water potential and gas exchange in response to chronic drought, and (ii) evaluate the concomitant impacts of these drought responses on the temporal patterns of NSC during terminal drought. The three distinct phases of water stress were delineated by thresholds of growth cessation and stomatal closure that defined the 'carbon safety margin' (i.e., the difference between leaf water potential when growth is zero and leaf water potential when net photosynthesis is zero). A wider C safety margin in E. globulus was defined by an earlier cessation of growth relative to photosynthesis that reduced the demand for NSC while maintaining C acquisition. By contrast, the narrower C safety margin in P. radiata was characterized by a synchronous decline in growth and photosynthesis, whereby growth continued under a declining supply of NSC from photosynthesis. The narrower C safety margin in P. radiata was associated with declines in starch concentrations after ∼ 90 days of chronic drought and significant depletion of starch in all organs at mortality. The observed divergence in the sensitivity of drought responses is indicative of a potential trade-off between maintaining hydraulic safety and adequate C availability.

Down-regulation of photosynthesis following girdling, but contrasting effects on fruit set and retention, in two sweet cherry cultivars.

Sweet cherry (Prunus avium) trees were manipulated to analyse the contribution of soluble sugars to sink feedback down-regulation of leaf net CO2 assimilation rate (Anet) and fruit set and quality attributes. Total soluble sugar concentration and Anet were measured in the morning on fully expanded leaves of girdled branches in two sweet cherry cultivars, 'Kordia' and 'Sylvia' characterised typically by low and high crop load, respectively. Leaves on girdled trees had higher soluble sugar concentrations and reduced Anet than leaves on non-girdled trees. Moreover, RuBP carboxylation capacity of Rubisco (Vcmax) and triose-phosphate utilisation (TPU) were repressed in the girdled treatments, despite Jmax remaining unchanged; suggesting an impairment of photosynthetic capacity in response to the girdling treatment. Leaf Anet was negatively correlated to soluble sugars, suggesting a sink feedback regulatory control of photosynthesis. Although there were significantly less fruit set and retained in 'Kordia' than 'Sylvia'; girdling had contrasting effects in each cultivar. Girdling significantly increased fruit set and fruitlet retention in 'Sylvia' cultivar, but had no effect in 'Kordia' cultivar. We propose that low inherent sink demand for photoassimilates of 'Kordia' fruit could have contributed to the low fruit retention rate, since both non-girdled and girdled trees exhibited similar retention rate and that increases in foliar carbohydrates was observed above the girdle. In 'Sylvia' cultivar, the carbohydrate status may be a limiting factor for 'Sylvia' fruit, since girdling improved both fruit set and retention, and leaf soluble solids accumulation.

A latitudinal cline in disease resistance of a host tree.

The possible drivers and implications of an observed latitudinal cline in disease resistance of a host tree were examined. Mycosphaerella leaf disease (MLD) damage, caused by Teratosphaeria species, was assessed in five Eucalyptus globulus (Tasmanian blue gum) common garden trials containing open-pollinated progeny from 13 native-forest populations. Significant population and family within population variation in MLD resistance was detected, which was relatively stable across different combinations of trial sites, ages, seasons and epidemics. A distinct genetic-based latitudinal cline in MLD damage among host populations was evident. Two lines of evidence argue that the observed genetic-based latitudinal trend was the result of direct pathogen-imposed selection for MLD resistance. First, MLD damage was positively associated with temperature and negatively associated with a prediction of disease risk in the native environment of these populations; and, second, the quantitative inbreeding coefficient (QST) significantly exceeded neutral marker FST at the trial that exhibited the greatest MLD damage, suggesting that diversifying selection contributed to differentiation in MLD resistance among populations. This study highlights the potential for spatial variation in pathogen risk to drive adaptive differentiation across the geographic range of a foundation host tree species.

Interactive effects of water supply and defoliation on photosynthesis, plant water status and growth of Eucalyptus globulus Labill.

Increased climatic variability, including extended periods of drought stress, may compromise on the health of forest ecosystems. The effects of defoliating pests on plantations may also impact on forest productivity. Interactions between climate signals and pest activity are poorly understood. In this study, we examined the combined effects of reduced water availability and defoliation on maximum photosynthetic rate (A(sat)), stomatal conductance (g(s)), plant water status and growth of Eucalyptus globulus Labill. Field-grown plants were subjected to two water-availability regimes, rain-fed (W-) and irrigated (W+). In the summer of the second year of growth, leaves from 75% of crown length removed from trees in both watering treatments and physiological responses within the canopies were examined. We hypothesized that defoliation would result in improved plant water status providing a mechanistic insight into leaf- and canopy-scale gas-exchange responses. Defoliated trees in the W+ treatment exhibited higher A(sat) and g(s) compared with non-defoliated trees, but these responses were not observed in the W- treatment. In contrast, at the whole-plant scale, maximum rates of transpiration (E(max)) and canopy conductance (G(Cmax)) and soil-to-leaf hydraulic conductance (K(P)) increased in both treatments following defoliation. As a result, plant water status was unaffected by defoliation and trees in the defoliated treatments exhibited homeostasis in this respect. Whole-plant soil-to-leaf hydraulic conductance was strongly correlated with leaf scale g(s) and A(sat) following the defoliation, providing a mechanistic insight into compensatory up-regulation of photosynthesis. Above-ground height and diameter growth were unaffected by defoliation in both water availability treatments, suggesting that plants use a range of responses to compensate for the impacts of defoliation.

Estimating forest net primary production under changing climate: adding pests into the equation.

The current approach to modelling pest impacts on forest net primary production (NPP) is to apply a constant modifier. This does not capture the large spatial and temporal variability in pest abundance and activity that can occur, meaning that overestimates or underestimates of pest impacts on forest NPP are likely. Taking a more mechanistic approach that incorporates an understanding of how physiology is influenced by pest attack, enables us to better capture system feedbacks and dynamics, thereby improving the capacity to predict into novel situations such as changing climate, and to account for both changes in pest activity and host responses to the growing environment now and into the future. We reviewed the effects of pests on forest NPP and found a range of responses and physiological mechanisms underlying those responses. Pest outbreaks can clearly be a major perturbation to forest NPP, and it seems likely that the frequency and intensity of pest outbreaks, and the ways in which host species respond to pest damage, will change in the future. We summarized these impacts in the form of a conceptual model at leaf, tree and stand scales, and compared the physiological processes embedded within that framework with the capacity of a representative range of NPP models to capture those processes. We found that some models can encapsulate some of the processes, but no model can comprehensively account for the range of physiological responses to pest attack experienced by trees. This is not surprising, given the paucity of empirical data for most of the world's forests, and that the models were developed primarily for other purposes. We conclude with a list of the key physiological processes and pathways that need to be included in forest growth models in order to adequately capture pest impacts on forest NPP under current and future climate scenarios, the equations that might enable this and the empirical data required to support them.

Are gas exchange responses to resource limitation and defoliation linked to source:sink relationships?

Productivity of trees can be affected by limitations in resources such as water and nutrients, and herbivory. However, there is little understanding of their interactive effects on carbon uptake and growth. We hypothesized that: (1) in the absence of defoliation, photosynthetic rate and leaf respiration would be governed by limiting resource(s) and their impact on sink limitation; (2) photosynthetic responses to defoliation would be a consequence of changing source:sink relationships and increased availability of limiting resources; and (3) photosynthesis and leaf respiration would be adjusted in response to limiting resources and defoliation so that growth could be maintained. We tested these hypotheses by examining how leaf photosynthetic processes, respiration, carbohydrate concentrations and growth rates of Eucalyptus globulus were influenced by high or low water and nitrogen (N) availability, and/or defoliation. Photosynthesis of saplings grown with low water was primarily sink limited, whereas photosynthetic responses of saplings grown with low N were suggestive of source limitation. Defoliation resulted in source limitation. Net photosynthetic responses to defoliation were linked to the degree of resource availability, with the largest responses measured in treatments where saplings were ultimately source rather than sink limited. There was good evidence of acclimation to stress, enabling higher rates of C uptake than might otherwise have occurred.

Photosynthesis of Eucalyptus globulus with Mycosphaerella leaf disease.

Mycosphaerella leaf disease (MLD) is a major cause of foliage damage in Eucalyptus globulus plantations. Our study is the first to describe the physiological effects of MLD on E. globulus leaves. It involved measurements on both field and potted plants. Changes in photosynthetic parameters in response to MLD were quantified in a study using gas exchange techniques. There was a negative linear relationship between light-saturated photosynthesis (A(max)) and leaf-level damage from MLD. Reductions in A(max) were proportionally greater than might be expected from the reduction in green leaf area as a result of the disease, indicating that asymptomatic tissue also was affected by MLD. The reductions in A(max) were not related to increases in stomatal resistance, but were a result of reduced activity of ribulose bisphosphate carboxylase (Rubisco) and changes in the capacity for ribulose bisphosphate (RuBP) regeneration. Changes in mesophyll resistance to CO2 were also implicated. The effect of MLD was similar at different sites and irrespective of tree-level infection, suggesting a general leaf-level response of E. globulus to MLD.

Precision and accuracy of pest and pathogen damage assessment in young eucalypt plantations.

Fungal pathogens, browsing mammals, birds, insects, nutrient deficiencies, drought, frost and waterlogging are all damaging agents to plantation species. The subsequent loss in leaf tissue or reduced photosynthetic potential can reduce growth and potentially lead to tree death. The Crown Damage Index (CDI) was developed in Australia to quantify damage in young eucalypt plantations. The accuracy and precision of assessing damage at a tree level were determined to ensure the reliability, objectivity and repeatability of the CDI method. Nine assessors, with varying levels of experience, estimated damage on three plots of fifty trees each, to obtain an understanding of the subjectivity of assessing damage caused by insects (e.g. Chrysophtharta spp.) and fungal pathogens (e.g. Mycosphaerella spp.) on Eucalyptus globulus. Damage levels were measured by destructive sampling to enable direct comparisons between estimates and damage levels to be made. The most experienced assessors provided the most repeatable estimates and were generally the most accurate. The incidence of foliar necrosis was the least subjective measure while defoliation was the most subjective and the least accurate of the indices measured. All assessors, regardless of experience, were able to predict the Crown Damage Index (a combined index of all damage classes) to within 12% of measured damage levels.

Modeling the effect of physiological responses to green pruning on net biomass production of Eucalyptus nitens.

Green pruning of Eucalyptus nitens (Deane and Maiden) Maiden increases instantaneous rates of light-saturated CO(2) assimilation (A), and changes patterns of total leaf area and foliage distribution. We investigated the importance of such changes on the rate of recovery of growth following pruning. A simple process-based model was developed to estimate daily net biomass production (G(d)) of three-year-old plantation-grown trees over a 20-month period. The trees had been pruned by removal of 0, 50 or 70% of the length of green crown, equivalent to removal of 0, 55 or 88% of leaf area, respectively, when the plantation verged on canopy closure. Total G(d) was reduced by only 20% immediately following the 50%-pruning treatment, as a result of both the high leaf dark respiration and low A in the portion of the crown removed compared to the top of the crown. Pruning at the time of canopy closure preempted a natural and rapid decline in G(d) of the lower crown. Although leaf area index (L) was approximately 6.0 at the time of pruning, high light interception (95%) occurred with an L of 4.0. The 50%-pruning treatment reduced L to 3.5, but the physiological responses to pruning were sufficient to compensate fully for the reduction in intercepted radiation within 110 days of pruning. The 70%-pruning treatment reduced L to 1.9, and reduced G(d) by 77%, reflecting the removal of branches with high A in the mid and upper crown. Physiological responses to the 70%-pruning treatment were insufficient to increase G(d) to the value of unpruned trees during the study. Model sensitivity analysis showed that increases in A following pruning increased G(d) by 20 and 25% in the 50- and 70%-pruned trees, respectively, 20 months after pruning. Changes in leaf area/foliage distribution had a greater effect on G(d) of 50%-pruned trees (47% increase) than did changes in A. However, the reduction in photosynthetic potential associated with the 70%-pruning treatment resulted in only small changes in leaf area/foliage distribution, which consequently had little effect on G(d). The effects of physiological processes occurring within the crown and in response to green pruning on G(d) are discussed with respect to pruning of plantations.