The temporal response to drought in a Mediterranean evergreen tree: comparing a regional precipitation gradient and a throughfall exclusion experiment (original) (raw)
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Tree Physiology, 2016
Adjustment in leaf water status parameters and modification in xylem structure and functioning can be important elements of a tree's response to continued water limitation. In a growth trial with saplings of five co-occurring temperate broad-leaved tree species (genera Fraxinus, Acer, Carpinus, Tilia and Fagus) conducted in moist or dry soil, we compared the drought acclimation in several leaf water status and stem hydraulic parameters. Considering the extremes in the species responses, Fraxinus excelsior L. improved its leaf tissue hydration in the dry treatment through osmotic, elastic and apoplastic adjustment while Fagus sylvatica L. solely modified its xylem anatomy, which resulted in increased embolism resistance at the cost of hydraulic efficiency. Our results demonstrate the contrasting response strategies of coexisting tree species and how variable trait plasticity among species can be. The comparison of plants grown either in monoculture or in five-species mixture showed that the neighbouring species diversity can significantly influence a tree's hydraulic architecture and leaf water status regulation. Droughted Carpinus betulus L. (and to a lesser extent, Acer pseudoplatanus L.) plants developed a more efficient stem hydraulic system in heterospecific neighbourhoods, while that of F. sylvatica was generally more efficient in conspecific than heterospecific neighbourhoods. We conclude that co-occurring tree species may develop a high diversity of drought-response strategies, and exploring the full diversity of trait characteristics requires synchronous study of acclimation at the leaf and stem (and possibly also the root) levels, and consideration of physiological as well as morphological and anatomical modifications.
Annals of Forest Science, 2006
The extreme drought event that occurred in Western Europe during 2003 highlighted the need to understand the key processes that may allow trees and stands to overcome such severe water shortages. We therefore reviewed the current knowledge available about such processes. First, impact of drought on exchanges at soil-root and canopy-atmosphere interfaces are presented and illustrated with examples from water and CO 2 flux measurements. The decline in transpiration and water uptake and in net carbon assimilation due to stomatal closure has been quantified and modelled. The resulting models were used to compute water balance at stand level basing on the 2003 climate in nine European forest sites from the CARBOEUROPE network. Estimates of soil water deficit were produced and provided a quantitative index of soil water shortage and therefore of the intensity of drought stress experienced by trees during summer 2003. In a second section, we review the irreversible damage that could be imposed on water transfer within trees and particularly within xylem. A special attention was paid to the inter-specific variability of these properties among a wide range of tree species. The inter-specific diversity of hydraulic and stomatal responses to soil water deficit is also discussed as it might reflect a large diversity in traits potentially related to drought tolerance. Finally, tree decline and mortality due to recurrent or extreme drought events are discussed on the basis of a literature review and recent decline studies. The potential involvement of hydraulic dysfunctions or of deficits in carbon storage as causes for the observed long term (several years) decline of tree growth and development and for the onset of tree dieback is discussed. As an example, the starch content in stem tissues recorded at the end of the 2003's summer was used to predict crown conditions of oak trees during the following spring: low starch contents were correlated with large twig and branch decline in the crown of trees.
Biologia plantarum, 2005
Plant water potential (ψ), its components, and gas exchange data of two Mediterranean co-occurring woody species (Quercus ilex L. and Phillyrea latifolia L.) were measured in response to seasonal changes in water availability over two consecutive years. The relative contribution of physiological and morphological adjustments to drought resistance was assessed through Principal Component Analyses. There were large adjustments in stomatal conductance (~36 % of accounted variance). Net photosynthetic rate and water use efficiency were closely tuned to water availability and accounted for ~17 % of variance. The slope of the water potential vs. relative water content (dψ/dRWC 0) below zero pressure potential increased as a result of seasonal and ontogenic increases in apoplastic water fraction and accounted for ~20 % variance. This tolerance mechanism was accompanied by an increased range of positive pressure potential, suggesting a functional role of sclerophylly in these Mediterranean evergreens. Similarly, changes in the slope of dψ/dRWC in the range of positive pressure potential (~13 % of accounted variance) were associated to variations in cell wall elasticity and resulted in lower RWC at zero pressure potential. When considering the species studied separately, the results indicated the primary role of stomatal regulation in the drought resistance of Q. ilex, while increased apoplastic water fraction had a major contribution in the drought resistance of P. latifolia.
Tree Physiology, 2020
Nowadays, evergreen sclerophyllous and winter-deciduous malacophyllous oaks with different paleogeographical origins coexist under Mediterranean-type climates, such as the mixed forests of the evergreen Quercus ilex subsp. rotundifolia Lam. and the winter-deciduous Quercus faginea Lam. Both Mediterranean oaks constitute two examples of contrasting leaf habit, so it could be expected that they would have different functional strategies to cope with summer drought. In this study, we analysed photosynthetic, photochemical and hydraulic traits of different organs for Q. faginea and Q. ilex subsp. rotundifolia under well-watered conditions and subjected to very severe drought. The coordinated response between photosynthetic and hydraulic traits explained the higher photosynthetic capacity of Q. faginea under well-watered conditions, which compensated its shorter leaf life span at the expense of higher water consumption. The progressive imposition of water stress evidenced that both types...
Oecologia, 2000
We investigated scaling of physiological parameters between age classes of Quercus rubra by combining in situ field measurements with an experimental approach. In the in situ field study, we investigated changes in drought response with age in seedlings, juveniles, and mature trees of Q. rubra. Throughout the particularly dry summer of 1995 and the unusually wet summer of 1996 in New England, we measured water potential of leaves (Ψ Leaf) and gas exchange of plants at three sites at the Harvard Forest in Petersham, Massachusetts. In order to determine what fraction of the measured differences in gas exchange between seedlings and mature trees was due to environment versus ontogeny, an experiment was conducted in which seedlings were grown under light and soil moisture regimes simulating the environment of mature trees. The photosynthetic capacity of mature trees was threefold greater than that of seedlings during the wet year, and six-fold greater during the drought year. The seedling experiment demonstrated that the difference in photosynthetic capacity between seedlings and mature trees is comprised equally of an environmental component (50%) and an ontogenetic component (50%) in the absence of water limitation. Photosynthesis was depressed more severely in seedlings than in mature trees in the drought year relative to the wet year, while juveniles showed an intermediate response. Throughout the drought, the predawn leaf water potential (Ψ PD) of seedlings became increasingly negative (-0.4 to-1.6 MPa), while that of mature trees became only slightly more negative (-0.2 to-0.5 MPa). Again, juveniles showed an intermediate response (-0.25 to-0.8 MPa). During the wet summer of 1996, however, there was no difference in Ψ PD between seedlings, juveniles and mature trees. During the dry summer of 1995, seedlings were more responsive to a major rain event than mature trees in terms of Ψ Leaf , suggesting that the two age classes depend on different water sources. In all age classes, instantaneous measurements of intrinsic water use efficiency (WUE i), defined as C assimilation rate divided by stomatal conductance, increased as the drought progressed, and all age classes had higher WUE i during the drought year than in the wet year. Mature trees, however, showed a greater ability to increase their WUE i in response to drought. Integrated measurements of WUE from C isotope discrimination (∆) of leaves indicated higher WUE in mature trees than juveniles and seedlings. Differences between years, however, could not be distinguished, probably due to the strong bias in C isotope fractionation at the time of leaf production, which occurred prior to the onset of drought conditions in 1995. From this study, we arrive at two main conclusions: 1. Different age classes of Q. rubra use different strategies for responding to drought. Seedlings resist drought by closing stomata early in the day at the expense of C uptake; mature trees avoid drought conditions by accessing deeper water reserves and adjusting WUE. 2. Only through studies which separate environmental differences from ontogenetic differences can parameters measured on seedlings be scaled to mature trees.
Journal of Experimental Botany, 2011
Plants may exhibit some degree of acclimation after experiencing drought, but physiological adjustments to consecutive cycles of drought and re-watering (recovery) have scarcely been studied. The Mediterranean evergreen holm oak (Q. ilex) and the semi-deciduous rockrose (C. albidus) showed some degree of acclimation after the first of three drought cycles (S1, S2, and S3). For instance, during S2 and S3 both species retained higher relative leaf water contents than during S1, despite reaching similar leaf water potentials. However, both species showed remarkable differences in their photosynthetic acclimation to repeated drought cycles. Both species decreased photosynthesis to a similar extent during the three cycles (20-40% of control values). However, after S1 and S2, photosynthesis recovered only to 80% of control values in holm oak, due to persistently low stomatal (g s) and mesophyll (g m) conductances to CO 2. Moreover, leaf intrinsic water use efficiency (WUE) was kept almost constant in this species during the entire experiment. By contrast, photosynthesis of rockrose recovered almost completely after each drought cycle (90-100% of control values), while the WUE was largely and permanently increased (by 50-150%, depending on the day) after S1. This was due to a regulation which consisted in keeping g s low (recovering to 50-60% of control values after re-watering) while maintaining a high g m (even exceeding control values during rewatering). While the mechanisms to achieve such particular regulation of water and CO 2 diffusion in leaves are unknown, it clearly represents a unique acclimation feature of this species after a drought cycle, which allows it a much better performance during successive drought events. Thus, differences in the photosynthetic acclimation to repeated drought cycles can have important consequences on the relative fitness of different Mediterranean species or growth forms within the frame of climate change scenarios.
Mediterranean trees coping with severe drought: Avoidance might not be safe
Environmental and Experimental Botany, 2018
Plants coexisting in Mediterranean regions differ widely in their sensitivity to drought and in their ability to maintain carbon assimilation and hydraulic function ranging from avoidance to tolerance of stressful periods. We studied the response of three coexisting Mediterranean tree species (Quercus ilex L. spp. Ballota (Desf.) Samp., Quercus faginea Lam. and Pinus nigra ssp. Salzmannii J.F. Arnold) under natural conditions for three years to test their functional strategies in dealing with extremely dry years. Despite P. nigra following a droughtavoiding strategy, it was unable to effectively avoid the hampering effects of a severe drought in the driest year, and experienced significant drops in the efficiency of the hydraulic system, in gas exchange at the leaf level and in tree water use estimated from daily sap flow. In contrast, the two Quercus species showed a more droughttolerant strategy by maintaining a more profligate water use at the leaf and tree level, with Q. ilex having a larger tolerance threshold to severe drought. The main decrease in carbon assimilation was due to stomatal closure, with a minor effect of photochemistry damage or down regulation in the three species. In the case of the two oaks, seasonal osmoregulation was inferred from changes in the osmotic potential at full turgor (π o), bringing about the lowest leaf osmotic potentials at turgor lost (π tlp) at late summer in both species, and maintenance of leaf turgor under the lowest midday leaf water potential (Ψ mid). Seasonal changes in osmotic potentials were observed in the three years, more during the driest year. Plasticity in modulating the tolerance to leaf dehydration as the season advances, especially in response to the intensity of inter-annual summer droughts was higher for the evergreen Q. ilex than in the semi-deciduous Q. faginea. In spite of the intensity of the drought and the associated stress experienced by the three forest tree species, especially by P. nigra, mortality or severe defoliation were not observed after the extreme dry year. This reveals a large operational range of coping with water stress by the three species regardless of the hydric strategy, which is crucial for survival under the increasing drought expected in the most likely climate change scenarios for the region.
Functional Plant Biology, 2012
Drought frequency and intensity are expected to increase in the Mediterranean as a consequence of global climate change. To understand how photosynthetic capacity responds to long-term water stress, we measured seasonal patterns of stomatal (S L ), mesophyll (MC L ) and biochemical limitations (B L ) to net photosynthesis (A max ) in three Quercus ilex (L.) populations from sites differing in annual rainfall. In the absence of water stress, stomatal conductance (g s ), maximum carboxylation capacity (V cmax ), photosynthetic electron transport rate (J max ) and A max were similar among populations. However, as leaf predawn water potential (Y l,pd ) declined, the population from the wettest site showed steeper declines in g s , V cmax , J max and A max than those from the drier sites. Consequently, S L , MC L and B L increased most steeply in response to decreasing Y l,pd in the population from the wettest site. The higher sensitivity of A max to drought was primarily the result of stronger stomatal regulation of water loss. Among-population differences were not observed when g s was used instead of Y l,pd as a drought stress indicator. Given that higher growth rates, stature and leaf area index were observed at the wettest site, we speculate that hydraulic architecture may explain the greater drought sensitivity of this population. Collectively, these results highlight the importance of considering among-population differences in photosynthetic responses to seasonal drought in large scale process-based models of forest ecosystem function.
Annals of botany, 2014
Vulnerability of the leaf hydraulic pathway to water-stress-induced dysfunction is a key component of drought tolerance in plants and may be important in defining species' climatic range. However, the generality of the association between leaf hydraulic vulnerability and climate across species and sites remains to be tested. Leaf hydraulic vulnerability to drought (P50leaf, the water potential inducing 50 % loss in hydraulic function) was measured in a diverse group of 92 woody, mostly evergreen angiosperms from sites across a wide range of habitats. These new data together with some previously published were tested against key climate indices related to water availability. Differences in within-site variability in P50leaf between sites were also examined. Values of hydraulic vulnerability to drought in leaves decreased strongly (i.e. became more negative) with decreasing annual rainfall and increasing aridity across sites. The standard deviation in P50leaf values recorded withi...
Journal of Experimental Botany, 2010
Understanding the responses of cork oak (Quercus suber L.) to actual and predicted summer conditions is essential to determine the future sustainability of cork oak woodlands in Iberia. Thermal imaging may provide a rapid method for monitoring the extent of stress. The ecophysiology of cork trees was studied over three years. Three treatments were applied by means of rainfall capture and irrigation, with plots receiving 120%, 100%, or 80% of natural precipitation. Despite stomatal closure, detected using both thermal imaging and porometry, leaf water potential fell during the summer, most drastically during the third year of accumulative stress. The quantum efficiency (ΦPSII) and the maximum efficiency Formula of photosystem II also fell more intensely over the third summer, while non-photochemical quenching (NPQ) increased. The reduced precipitation treatment sporadically further reduced leaf water potential, stomatal conductance (gs), IG (an index of gs derived from thermal imaging), ΦPSII, and Formula, and increased leaf temperature and NPQ. It is concluded that these are very resilient trees since they were only severely affected in the third year of severe drought (the third year registering 45% less rainfall than average), and removing 20% of rainfall had a limited impact.