Joana Zaragoza-Castells

Is Shade Beneficial for Mediterranean Shrubs Experiencing Periods of Extreme Drought and Late-winter Frosts?

BACKGROUND AND AIMS Plants are naturally exposed to multiple, frequently interactive stress factors, most of which are becoming more severe due to global change. Established plants have been reported to facilitate the establishment of juvenile plants, but net effects of plant-plant interactions are difficult to assess due to complex interactions among environmental factors. An investigation was carried out in order to determine how two dominant evergreen shrubs (Quercus ilex and Arctostaphylos uva-ursi) co-occurring in continental, Mediterranean habitats respond to multiple abiotic stresses and whether the shaded understorey conditions ameliorate the negative effects of drought and winter frosts on the physiology of leaves. METHODS Microclimate and ecophysiology of sun and shade plants were studied at a continental plateau in central Spain during 2004-2005, with 2005 being one of the driest and hottest years on record; several late-winter frosts also occurred in 2005. KEY RESULTS Daytime air temperature and vapour pressure deficit were lower in the shade than in the sun, but soil moisture was also lower in the shade during the spring and summer of 2005, and night-time temperatures were higher in the shade. Water potential, photochemical efficiency, light-saturated photosynthesis, stomatal conductance and leaf 13C composition differed between sun and shade individuals throughout the seasons, but differences were species specific. Shade was beneficial for leaf-level physiology in Q. ilex during winter, detrimental during spring for both species, and of little consequence in summer. CONCLUSIONS The results suggest that beneficial effects of shade can be eclipsed by reduced soil moisture during dry years, which are expected to be more frequent in the most likely climate change scenarios for the Mediterranean region.

The productivity, metabolism and carbon cycle of two lowland tropical forest plots in south-western Amazonia, Peru

Background: The forests of western Amazonia are known to be more dynamic that the better-studied forests of eastern Amazonia, but there has been no comprehensive description of the carbon cycle of a western Amazonian forest. Aims: We present the carbon budget of two forest plots in Tambopata in south-eastern Peru, western Amazonia. In particular, we present, for the first time, the seasonal variation in the detailed carbon budget of a tropical forest. Methods: We measured the major components of net primary production (NPP) and total autotrophic respiration over 3–6 years. Results: The NPP for the two plots was 15.1 ± 0.8 and 14.2 ± 1.0 Mg C ha−1 year−1, the gross primary productivity (GPP) was 35.5 ± 3.6 and 34.5 ± 3.5 Mg C ha−1 year−1, and the carbon use efficiency (CUE) was 0.42 ± 0.05 and 0.41 ± 0.05. NPP and CUE showed a large degree of seasonality. Conclusions: The two plots were similar in carbon cycling characteristics despite the different soils, the most notable difference being high allocation of NPP to canopy and low allocation to fine roots in the Holocene floodplain plot. The timing of the minima in the wet–dry transition suggests they are driven by phenological rhythms rather than being driven directly by water stress. When compared with results from forests on infertile forests in humid lowland eastern Amazonia, the plots have slightly higher GPP, but similar patterns of CUE and carbon allocation.

Respiration in Photosynthetic Cells: Gas Exchange Components, Interactions with Photorespiration and the Operation of Mitochondria in the Light

According to gas exchange measurements, mitochondrial oxygen consumption in the light is always fast, while respiratory CO2 evolution is markedly decreased (compared with rates in darkness). We analyze the metabolic events that lead to such contrasting responses. In the light, the generation of NADH in mitochondria, both in the glycine decarboxylase reaction and in the tricarboxylic acid cycle, leads to increased NAD(P)H levels, which may increase the activity of the rotenone-insensitive NAD(P)H dehydrogenases. The resulting increase of the reduction level of ubiquinone activates the alternative oxidase. Stabilization of (photo)respiratory flux during the transition from darkness to light takes place at higher NADH/NAD+ and ATP/ADP ratios. Maintenance of fast rates of mitochondrial electron transport in the light is facilitated by the import of oxaloacetate (OAA) from the cytosol to remove NADH, and by the export of citrate to the cytosol. This reduces the flow of metabolites in the tricarboxylic acid cycle, decreasing decarboxylation rates, while the rate of oxygen consumption reactions remain fast.

Light inhibition of leaf respiration in field-grown Eucalyptus saligna in whole-tree chambers under elevated atmospheric CO2 and summer drought

We investigated whether the degree of light inhibition of leaf respiration (R) differs among large Eucalyptus saligna grown in whole-tree chambers and exposed to present and future atmospheric [CO(2) ] and summer drought. Associated with month-to-month changes in temperature were concomitant changes in R in the light (R(light) ) and darkness (R(dark) ), with both processes being more temperature dependent in well-watered trees than under drought. Overall rates of R(light) and R(dark) were not significantly affected by [CO(2) ]. By contrast, overall rates of R(dark) (averaged across both [CO(2) ]) were ca. 25% lower under drought than in well-watered trees. During summer, the degree of light inhibition of leaf R was greater in droughted (ca. 80% inhibition) than well-watered trees (ca. 50% inhibition). Notwithstanding these treatment differences, an overall positive relationship was observed between R(light) and R(dark) when data from all months/treatments were combined (R(2)  = 0.8). Variations in R(light) were also positively correlated with rates of Rubisco activity and nitrogen concentration. Light inhibition resulted in a marked decrease in the proportion of light-saturated photosynthesis respired (i.e. reduced R/A(sat) ). Collectively, these results highlight the need to account for light inhibition when assessing impacts of global change drivers on the carbon economy of tree canopies.

Thermal acclimation of leaf dark respiration of beech seedlings experiencing summer drought in high and low light environments

Little is known about how environmental factors shape the short- and long-term responses of leaf respiration to temperature under field conditions despite the importance of respiration for plant and stand carbon balances. Impacts of water availability and canopy cover on leaf dark respiration (R) and temperature sensitivity were assessed in beech (Fagus sylvatica L.) seedlings in a sub-Mediterranean population. We studied seedlings established within canopy gaps (39% global site factor; GSF) that were subject to either no watering (unwatered plants; UW) or regular watering (2-10% higher volumetric topsoil water content as summer progressed; W plants) and seedlings established beneath the adjacent understorey (12% GSF). Leaf R rose exponentially with diurnal increases in temperature; the same temperature sensitivity (Q(10): 2.2) was found for understorey and gap plants, irrespective of watering treatment. Respiration estimated at 25 degrees C (R(25)) was lower in the understorey than the gaps and was significantly lower in the unwatered than in the watered gap plants by the end of summer (0.65 versus 0.80 micromol m(-2) s(-1)). R(25) declined with increasing summer temperature in all plants; however, respiration estimated at the prevailing ambient temperature did not change through the summer. There were parallel declines in R(25) and concentrations of starch and soluble sugars with increasing summer temperature for gap plants. We conclude that seasonal shifts in temperature-response curves of beech leaf R occur in both low- and high-light environments; since leaf R decreased with increasing plant water deficit, such shifts are likely to be greater whenever plants experience summer drought compared to scenarios where plants experience high rainfall in summer.

Evidence for strong seasonality in the carbon storage and carbon use efficiency of an Amazonian forest

The relative contribution of gross primary production and ecosystem respiration to seasonal changes in the net carbon flux of tropical forests remains poorly quantified by both modelling and field studies. We use data assimilation to combine nine ecological time series from an eastern Amazonian forest, with mass balance constraints from an ecosystem carbon cycle model. The resulting analysis quantifies, with uncertainty estimates, the seasonal changes in the net carbon flux of a tropical rainforest which experiences a pronounced dry season. We show that the carbon accumulation in this forest was four times greater in the dry season than in the wet season and that this was accompanied by a 5% increase in the carbon use efficiency. This seasonal response was caused by a dry season increase in gross primary productivity, in response to radiation and a similar magnitude decrease in heterotrophic respiration, in response to drying soils. The analysis also predicts increased carbon allocation to leaves and wood in the wet season, and greater allocation to fine roots in the dry season. This study demonstrates implementation of seasonal variations in parameters better enables models to simulate observed patterns in data. In particular, we highlight the necessity to simulate the seasonal patterns of heterotrophic respiration to accurately simulate the net carbon flux seasonal tropical forest.

Transient shade and drought have divergent impacts on the temperature sensitivity of dark respiration in leaves of Geum urbanum

The respiratory response of plants to temperature is a critical biotic feedback in the study of global climate change. Few studies, however, have investigated the effects of environmental stresses on the short-term temperature response of dark respiration (Rdark) at the leaf level. We investigated the effect of shade and transient drought on the temperature sensitivity (Q10; the proportional increase in respiration per 10°C increase in temperature) of Rdark of Geum urbanum L. in controlled experiments. Shade effects were most pronounced following sustained, near-darkness, when rates of leaf Rdark at a set measuring temperature (25°C) and the Q10 of Rdark were both reduced. By contrast, rates of leaf Rdark and the Q10 of Rdark both increased in response to the onset of severe water stress. Water stress was associated with a rapid (but reversible) decline in rates of light-saturated photosynthesis (Psat), stomatal closure (gs) and progressive wilting. Re-watering resulted in a rapid recovery of Psat, gs and a decline in the Q10 of Rdark (due to larger proportional reductions in the rate of Rdark measured at 25°C compared with those measured at 14°C). The concentration of soluble sugars in leaves did not decline during drought (5-7 day cycles) or shading, but during drought the starch concentration dropped, suggesting starch to sugar conversion helped to maintain homeostatic concentrations of soluble sugars. Thus, the drought and shade induced changes in Rdark were unlikely to be due to stress-induced changes in substrate supply. Collectively, the data highlight the dynamic responses of respiratory Q10 values to changes in water supply and sustained reductions in growth irradiance. If widespread, such changes in the Q10 of leaf respiration could have important implications for predicted rates of ecosystem carbon exchange in the future, particularly in areas that experience more frequent droughts.

Leaf respiration in darkness and in the light under pre-industrial, current and elevated atmospheric CO2 concentrations

Our study sought to understand how past, low atmospheric CO2 concentrations ([CO2]) impact respiration (R) of soybean (Glycine max), when compared to plants grown under current and future [CO2]s. Experiments were conducted using plants grown under 290, 400 and 700 ppm [CO2]. Leaf R was measured in both darkness (RD) and in the light (RL; using the Kok method), with short-term changes in measurement [CO2] and [O2] being used to explore the relationship between light inhibition of leaf R and photorespiration. Root R, photosynthesis (A), leaf [N] and biomass allocation traits were also quantified. In contrast to the inhibitory effect of low growth [CO2] on A, growth [CO2] had no significant effect on leaf RD or root R. Irrespective of growth [CO2], RL was always lower than RD, with light inhibiting leaf R by 17-47%. Importantly, the degree of light inhibition of leaf R was lowest in plants grown under low [CO2], with variations in RL being positively correlated with RD and photorespiration. Irrespective of whether leaf R was measured in the light or dark, a greater proportion of the carbon fixed by leaf photosynthesis was released by leaf R in plants grown under low [CO2] than under current/future [CO2]s. Collectively, our results highlight the differential responses of A and R to growth of plants under low to elevated atmospheric [CO2].

Scaling leaf respiration with nitrogen and phosphorus in tropical forests across two continents

Summary Leaf dark respiration (R dark) represents an important component controlling the carbon balance in tropical forests. Here, we test how nitrogen (N) and phosphorus (P) affect R dark and its relationship with photosynthesis using three widely separated tropical forests which differ in soil fertility. R dark was measured on 431 rainforest canopy trees, from 182 species, in French Guiana, Peru and Australia. The variation in R dark was examined in relation to leaf N and P content, leaf structure and maximum photosynthetic rates at ambient and saturating atmospheric CO 2 concentration. We found that the site with the lowest fertility (French Guiana) exhibited greater rates of R dark per unit leaf N, P and photosynthesis. The data from Australia, for which there were no phylogenetic overlaps with the samples from the South American sites, yielded the most distinct relationships of R dark with the measured leaf traits. Our data indicate that no single universal scaling relationship accounts for variation in R dark across this large biogeographical space. Variability between sites in the absolute rates of R dark and the R dark : photosynthesis ratio were driven by variations in N‐ and P‐use efficiency, which were related to both taxonomic and environmental variability.

Light inhibition of foliar respiration in response to soil water availability and seasonal changes in temperature in Mediterranean holm oak (Quercus ilex) forest

In the present study we investigated variations in leaf respiration in darkness (RD) and light (RL), and associated traits in response to season, and along a gradient of soil moisture, in Mediterranean woodland dominated by holm oak (Quercus ilex L.) in central and north-eastern Spain respectively. On seven occasions during the year in the central Spain site, and along the soil moisture gradient in north-eastern Spain, we measured rates of leaf RD, RL (using the Kok method), light-saturated photosynthesis (A) and related light response characteristics, leaf mass per unit area (MA) and leaf nitrogen (N) content. At the central Spain site, significant seasonal changes in soil water content and ambient temperature (T) were associated with changes in MA, foliar N, A and stomatal conductance. RD measured at the prevailing daily T and in instantaneous R-T responses, displayed signs of partial acclimation and was not significantly affected by time of year. RL was always less than, and strongly related to, RD, and RL/RD did not vary significantly or systematically with seasonal changes in T or soil water content. Averaged over the year, RL/RD was 0.66±0.05s.e. (n=14) at the central Spain site. At the north-eastern Spain site, the soil moisture gradient was characterised by increasing MA and RD, and reduced foliar N, A, and stomatal conductance as soil water availability decreased. Light inhibition of R occurred across all sites (mean RL/RD=0.69±0.01s.e. (n=18)), resulting in ratios of RL/A being lower than for RD/A. Importantly, the degree of light inhibition was largely insensitive to changes in soil water content. Our findings provide evidence for a relatively constrained degree of light inhibition of R (RL/RD ~ 0.7, or inhibition of ~30%) across gradients of water availability, although the combined impacts of seasonal changes in both T and soil water content increase the range of values expressed. The findings thus have implications in terms of the assumptions made by predictive models that seek to account for light inhibition of R, and for our understanding of how environmental gradients impact on leaf trait relationships in Mediterranean plant communities.