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14 Pàg. Centro de Investigación en Sanidad Animal (CISA), Metabolic changes underpinning drought-induced variations in stem respiration (Rs ) are unknown. We measured Rs rates and metabolite and gene expression profiles in Ulmus minor Mill. and Quercus ilex L. seedlings subjected to increasing levels of drought stress to better understand how carbon, nitrogen and energy metabolism interact during drought. In both species, only plants showing extreme stress symptoms - i.e. negligible rates of leaf stomatal conductance and photosynthesis, and high stem dehydration (30-50% of maximum water storage) and contraction (50-150 μm week-1 ) - exhibited lower Rs rates than well-watered plants. Abundance of low-molecular weight sugars (e.g. glucose and fructose) and sugar alcohols (e.g. mannitol) increased with drought, at more moderate stress and to a higher extent in Q. ilex than U. minor. Abundance of amino acids increased at more severe stress, more abruptly, and to a higher extent in U. minor, coinciding with leaf senescence, which did not occur in Q. ilex. Organic acids changed less in response to drought: threonate and glycerate increased, and citrate decreased although slightly in both species. Transcripts of genes coding for enzymes of the Krebs cycle decreased in Q. ilex and increased in U. minor in conditions of extreme drought stress. The maintenance of Rs under severe growth and photosynthetic restrictions reveals the importance of stem mitochondrial activity in drought acclimation. The eventual decline in Rs diverts carbon substrates from entering the Krebs cycle that may help to cope with osmotic and oxidative stress during severe drought and to recover hydraulic functionality afterwards., Funding was provided by the project AGL2015‐66925‐R MINECO/FEDER, UE. CA acknowledges support by national funds from Fundação para a Ciência e Tecnologia (FCT) through the FCT Investigator Programme (contract IF/00376/2012/CP0165/CT0003) and the ITQB NOVA R&D unit GREEN‐IT ‘Bioresources for sustainability’ (UID/Multi/04551/2013). AMR acknowledges FCT for the PhD fellowship (PD/BD/114417/2016) and the ITQB NOVA International PhD Programme ‘Plants for Life’ (PD/00035/2013).

Resprouting woody species adapted to perturbations store large amounts of carbon reserves to sustain plant metabolism during and resprout following perturbations. However, it is not known how carbon sources and sinks re-adjust in response to perturbations. Here, we studied when and where different reserve pools were mobilized after an aboveground perturbation, and how were new assimilates used after resprouting. Leaf gas exchange, plant organ carbon isotopic composition (δ13C) and nonstructural carbohydrate concentrations [NSC] were measured over one year in Pinus canariensis saplings subjected to complete defoliation (D) or complete defoliation plus stem damage (DD). Growth stopped completely in D and DD trees by the second month after the treatment application and during subsequent crown development. Growth resumed in most trees (60%) by the end of the growing season. Relative to control (C) trees, stem xylem [starch] decreased in D and DD during resprouting of new buds. Stem and root [starch] were also lower during growth resumption in these trees. One year after aboveground damages, leaf and root δ13C were significantly lower in D and DD, whereas [starch] recovered to values of C trees, except in roots of DD trees. Mobilization of carbon reserves from woody organs allows for resprouting and contributes to crown regeneration, stem growth resumption and wound healing in DD trees. New 13C-depleted photoassimilates contribute to these processes, and to refilling carbohydrates reserves. High [starch], plasticity in growth during and recovery of [starch] after severe aboveground disturbance, highlights high resilience of P. canariensis to perturbation.

Leaf growth, predawn leaf water potential (Ψpd), evapotranspiration, stem maximum permeability, and its percentage loss of hydraulic conductivity (PLC) were measured in rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered and drought conditions. Drought significantly reduced evapotranspiration, leaf growth and maximum permeability. E. globulus clones lost up to 70% of conductivity at values of Ψpd less negative than –1 MPa. PLC values higher than 85% could not be measured without causing leaf shedding. The coefficient related to the slope of the vulnerability curves ranged from 1.52–2.23. The lowest value was measured in the most drought-resistant clone, as estimated from field trials. Plants from this clone displayed higher drought-induced reductions in maximum permeability than plants from other clones, had significantly smaller leaves and maintained higher values of predawn leaf water potential as soil water content (SWC) declined.

Dutch elm disease (DED) is a vascular disease that has killed over 1 billion elm trees. The pathogen spreads throughout the xylem network triggering vessel blockage, which results in water stress, tissue dehydration and extensive leaf wilting in susceptible genotypes. We investigated the differences between four Ulmus minor Mill. clones of contrasting susceptibility to Ophiostoma novo-ulmi Brasier regarding morphological, anatomical and physiological traits affecting water transport, in order to gain a better understanding of the mechanisms underlying DED susceptibility. We analyzed the differential response to water shortage and increased air vapor pressure deficit (VPD) to investigate whether resistance to water stress might be related to DED tolerance. Sixteen plants per clone, aged 2 years, were grown inside a greenhouse under differential watering. Stomatal conductance was measured under ambient and increased VPD. Growth, bark water content and stem hydraulic and anatomical parameters were measured 22 days after starting differential watering. Vessel lumen area, lumen fraction and hydraulic conductance were highest in susceptible clones. Stomatal conductance was lowest under low VPD and decreased faster under increased VPD in resistant clones. We found a negative relationship between the decrease in stomatal conductance at increased VPD and specific hydraulic conductance, revealing a narrower hydraulic margin for sustaining transpiration in resistant clones. The effect of water shortage was greater on radial stem growth than on leaf area, which could be explained through an extensive use of capacitance water to buffer xylem water potential. Water shortage reduced stomatal conductance and vessel lumen area. Bark water content under conditions of water shortage only decreased in susceptible clones. Higher hydraulic constraints to sap flow in resistant clones may determine higher stomatal sensitivity to VPD and so contribute to DED resistance by limiting pathogen expansion and reducing water loss and metabolic impairment in cells involved in fighting against infection.

Combining hydraulic- and carbon-related measurements helps to understand drought-induced plant mortality. Here, we investigated the role that plant respiration (R) plays in determining carbon budgets under drought. We measured the hydraulic conductivity of stems and roots, and gas exchange and nonstructural carbohydrate (NSC) concentrations of leaves, stems and roots of seedlings of two resprouting species exposed to drought or well-watered conditions Ulmus minor (riparian tree) and Quercus ilex (dryland tree). With increasing water stress (occurring more rapidly in larger U. minor), declines in leaf, stem and root R were less pronounced than that in leaf net photosynthetic CO2 uptake (Pn). Daytime whole-plant carbon gain was negative below −4 and −6 MPa midday xylem water potential in U. minor and Q. ilex, respectively. Relative to controls, seedlings exhibiting shoot dieback suffered c. 80% loss of hydraulic conductivity in both species, and reductions in NSC concentrations in U. minor. Higher drought-induced depletion of NSC reserves in U. minor was related to higher plant R, faster stomatal closure, and premature leaf-shedding. Differences in drought resistance relied on the ability to maintain hydraulic conductivity during drought, rather than tolerating conductivity loss. Root hydraulic failure elicited shoot dieback and precluded resprouting without root NSC reserves being apparently limiting for R. © 2016 The Authors. New Phytologist © 2016 New Phytologist Trust

Key message Flooding reduces leaf photosynthesis but initially enhances plant respiration. Differences in flood-induced restrictions to plant net carbon gain could underlie distinct susceptibility to flooding between species.

Agriculture and forestry traditionally focus on improving plant growth traits based on an anthropocentric point of view. This paradigm has led to global problems associated to soil overexploitation such as soil losses, reductions of the C stock in soils, and the generalized use of fertilizers, which particularly increases the costs of production and pollution treatment. This view may also have limited our understanding of mutualistic symbioses of plants and microorganisms assuming that the main role of non-photosynthetic symbionts is to mobilize the nutrients that are necessary for plant growth and development, and being plants the dominant agents of the symbiotic relationship. In response to these issues, this chapter offers an alternative approach taking advantage of the “rhizo-centric” point of view, where non-photosynthetic partners are the main protagonists in play; and secondly, it builds a multidisciplinary body of knowledge that could be called “rhizoculture”, which includes techniques focussing on the intensification of the development and activity of roots, mycorrhizae, and other symbiotic and free living rhizosphere organisms. In short, rhizoculture may lead to decrease plant production dependence on fertilization and provides other benefits to agriculture, forestry, and the environment. Within this conceptual framework, the first objective of this book chapter is to explore whether there is a “paradox of calcium salts” (i.e., Ca2+ and its salts are simultaneously nutrients, promoters, and stressors for the host plants) that would explain a dominance of mycorrhizal fungi over plants based on inducing a Ca(pH)–mediated chlorosis to the host plants. If this paradigm shifting hypothesis were finally fully verified, it would provide conceptual bases to reconsider our current technologies in agriculture and forestry by introducing the “rhizocultural” approach, based on the management of roots (introducing alternative cultural practices), Ca2+ salts (using liming and other techniques), rock-eating mycorrhizae, organic matter, and the soil microbiome (increasing the presence of symbiotic microorganisms against saprophytes), N and P contents (by aquaculture and smart recycling of organic waste), and the physical properties of the soil (by the activity of soil symbiotic microorganisms and soil fauna, such as ants, termites and earthworms). The development of such new technological approaches in rhizoculture would significantly decrease the high cost and associated pollution of the application of fertilizers and phytochemicals; as well as it would increase soil C stocks, improve the resilience of agricultural and forest systems to environmental disturbances, such as climate change, and enhance food production and security.

Climate change is expected to lead to increases in drought frequency and severity, with deleterious effects on many ecosystems. Stomatal responses to changing environmental conditions form the backbone of all ecosystem models, but are based on empirical relationships and are not well-tested during drought conditions. Here, we use a dataset of 34 woody plant species spanning global forest biomes to examine the effect of leaf water potential on stomatal conductance and test the predictive accuracy of three major stomatal models and a recently proposed model. We find that current leaf-level empirical models have consistent biases of over-prediction of stomatal conductance during dry conditions, particularly at low soil water potentials. Furthermore, the recently proposed stomatal conductance model yields increases in predictive capability compared to current models, and with particular improvement during drought conditions. Our results reveal that including stomatal sensitivity to declining water potential and consequent impairment of plant water transport will improve predictions during drought conditions and show that many biomes contain a diversity of plant stomatal strategies that range from risky to conservative stomatal regulation during water stress. Such improvements in stomatal simulation are greatly needed to help unravel and predict the response of ecosystems to future climate extremes. Funding for this research was provided by NSF DEB EF-1340270 and the Climate Mitigation Initiative at the Princeton Environmental Institute, Princeton University. SL acknowledges financial support from the China Scholarship Council (CSC). VRD acknowledges funding from Ramón y Cajal fellowship (RYC-2012-10970). BTW was supported by the Next Generation Ecosystem Experiments-Tropics, funded by the U.S. Department of Energy, Office of Science, Office of Biological and Environmental Research. DJC acknowledges funding from the National Science Centre, Poland (NN309 713340). WRLA was supported in part by NSF DEB 1714972.

The ability of trees to grow and survive during periods of water scarcity is determined by the water use strategy of each individual. In this study we evaluated how water use strategies can influence the establishment phase of three coexisting species in forests where harvesting intensity and silvicultural interventions have been reduced over recent decades. We conducted a greenhouse experiment between February and July, applying two watering regimes (moist and dry) to current year seedlings in 1.5 m high pots to avoid root growth constraint. At the same time, the physiological performance of seedlings of the same species was monitored under natural conditions in the field over a period of decreasing soil moisture. Our results indicated that resprouter species allocated more than 40% of total biomass to roots whereas the non-resprouter P. pinea allocated only 26%. The plasticity index for the root traits was highest in Q. ilex, in which the highest stomatal conductance was measured in early summer. The slowest growth and lowest root plasticity, root hydraulic conductance and leaf water potential were measured in J. oxycedrus. The three species exhibited a positive carbon balance under low water availability. Coexisting species displayed a continuum in the water use strategies adopted. P. pinea would be the least competitive species under the forecasted increasing drought conditions in contrast to J. oxycedrus, which presents traits more suited to coping with droughts. © 2016 Elsevier B.V.

Background and Aims Stomatal conductance has long been considered of key interest in the study of plant adaptation to water stress. The expected increase in extreme meteorological events under a climate change scenario may compromise survival in Eucalyptus globulus plantations established in south-western Spain. We investigated to what extent changes in stomatal conductance in response to high vapour pressure deficits and water shortage are mediated by hydraulic and chemical signals in greenhouse-grown E. globulus clones. Methods Rooted cuttings were grown in pots and submitted to two watering regimes. Stomatal conductance, shoot water potential, sap pH and hydraulic conductance were measured consecutively in each plant over 4 weeks under vapour pressure deficits ranging 0 42 to 2 25 kPa. Evapotranspiration, growth in leaf area and shoot biomass were also determined. Key Results There was a significant effect of both clone and watering regime in stomatal conductance and leaf-specific hydraulic conductance, but not in sap pH. Sap pH decreased as water potential and stomatal conductance decreased under increasing vapour pressure deficit. There was no significant relationship between stomatal conductance and leaf-specific hydraulic conductance. Stomata closure precluded shoot water potential from falling below -1 8 MPa. The percentage loss of hydraulic conductance ranged from 40 to 85 %. The highest and lowest leaf-specific hydraulic conductances were measured in clones from the same half-sib families. Water shortage reduced growth and evapotranspiration, decreases in evapotranspiration ranging from 14 to 32 % in the five clones tested. Conclusions Changes in sap pH seemed to be a response to changes in atmospheric conditions rather than soil water in the species. Stomata closed after a considerable amount of hydraulic conductance was lost, although intraspecific differences in leaf-specific hydraulic conductance suggest the possibility of selection for improved productivity under water-limiting conditions combined with high temperatures in the early stages of growth. © The Author 2016. Published by Oxford University Press on behalf of the Annals of Botany Company.

The photosynthesis source-sink relationship in young Pinus canariensis seedlings was modified by stem girdling to investigate sprouting and cambial activity, feedback inhibition of photosynthesis, and stem and root hydraulic capacity. Removal of bark tissue showed a trade-off between sprouting and diameter growth. Above the girdle, growth was accelerated but the number of sprouts was almost negligible, whereas below the girdle the response was reversed. Girdling resulted in a sharp decrease in whole plant transpiration and root hydraulic conductance. The reduction of leaf area after girdling was strengthened by the high levels of abscisic acid found in buds which pointed to stronger bud dormancy, preventing a new needle flush. Accumulation of sugars in leaves led to a coordinated reduction in net photosynthesis (AN) and stomatal conductance (gS) in the short term, but later (gS below 0.07 mol m(-2) s(-1)) AN decreased faster. The decrease in maximal efficiency of photosystem II (FV/FM) and the operating quantum efficiency of photosystem II (?PSII) in girdled plants could suggest photoprotection of leaves, as shown by the vigorous recovery of AN and ?PSII after reconnection of the phloem. Stem girdling did not affect xylem embolism but increased stem hydraulic conductance above the girdle. This study shows that stem girdling affects not only the carbon balance, but also the water status of the plant.

Background Wounds caused by fire, herbivorism, rock impacts, etc. cause the direct loss of photosynthetic, storage and/or vascular tissue. In addition, they may entail other damages, such as desiccation of the exposed internal parts, or become a gateway to infection by fungi and other pathogens. To successfully overcome such injuries, plants must reorganize their meristems or even differentiate new ones, producing new traumatic tissues to cover the wound and restore the vascular connection. Results In this work we analyse the anatomical growth response in conifers after debarking and injuring the vascular cambium, using Pinus canariensis as model species, due to its high wound recovery ability. Conversely to angiosperm woody species, this process is initiated and largely driven by the damaged vascular cambium and not by proliferation in the wound surface. We have detected alterations and switches in the divisions of cambial cells, associated to their position relative to the surface and edges of the wound, resulting in disordered traumatic xylem. We also describe the formation of column-like structures, after girdling, which are in part formed by the proliferation of xylem parenchymatous cells, associated to axial resin ducts. Conclusions Abundant resinosis on the wound surface, typical of conifers, is an efficient barrier against opportunistic fungi, insects, etc. but it also hinders the healing process directly from the surface. Thus, wound closure must be largely carried out from the wound margins, being a much slower process, which very often remains unconcluded for long years. This work also describes for the first time the proliferation of inner parenchymatous cells to form column-like structures, which accelerates wound closure in girdled P. canariensis. Irregularities in the surface of the healing edge or column-like structures result in the production of disordered vascular tissues, compromising their future functionality, and which must be overcome through the fast restoration of the proper polarity in vascular cambium.

Sustainable forest management requires knowledge of forest structure and dynamics as well as an estimation of growing stock. The forest inventory provides the data for estimating stand variables. The measurement device MU2005-01738, patented by the Center for Forest Research (INIA-CIFOR), provides stereoscopic hemispherical images which can allow the 3D restoration of the stand around the sampling point. The aim of this study is to develop a methodology for forest stand mapping as well as tree diameter and height measurement along with volume estimation from the stereoscopic hemispherical images provided by the MU2005-01738. Using the MU2005-01738, Eucalyptus globulus Labill. plantations were sampled. Distance, diameter, height and volume were derived from the stereoscopic hemispherical images of 30 trees located at distances ranging from 0 to 15 m from the device. These variables were then compared with field measurements and the estimation errors analyzed. The (root-mean-squared error) RMSE was 0.23 m (8.95 %) for tree position and 1.51 cm (10.43 %) for diameter at breast height measurement at distances of less than 8 m. In the case of stem height and individual tree volume estimation, the RMSE was 2.59 m (23.05 %) and 0.025 m3 (17.94 %), respectively. The analysis of measurement errors indicated that the measurement precision decreases beyond 8 m from the device as well as for directions close to the baseline (the line between the optical centers of the two images), whereas the precision was highest for directions near to the line which is perpendicular to the base line. Future research should focus on improving measurement accuracy and possible applications in the field of forestry of the techniques presented in this study. © 2014 Springer-Verlag Berlin Heidelberg.

Changes in leaf size, specific leaf area (SLA), transpiration and tissue water relations were studied in leaves of rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered or drought conditions in a greenhouse. Significant differences between clones were found in leaf expansion and transpiration. There was a significant clone × treatment interaction on SLA. Water stress significantly reduced osmotic potential at the turgor loss point (Π 0 ) and at full turgor (Π 100 ), and significantly increased relative water content at the turgor loss point and maximum bulk elastic modulus. Differences in tissue water relations between clones were significant only in the mild drought treatment. Among clones in the drought treatments, the highest leaf expansion and the highest increase in transpiration during the experiment were measured in those clones that showed an early and large decrease in Π 0 and Π 100 .

In this study a stand level approach is proposed for estimating leaf area index (LAI) in even-aged Scots pine stands using gap fraction data derived from hemispherical photographs. The approach includes both the effect of the spatial distribution of the foliage elements as well as the slope of the terrain. Two chronosequences consisting of 11 plots were established in two Scots pine (Pinus sylvestris L.) forests in the Central Mountain Range of Spain. 12 hemispherical photographs were taken in each plot and variables related to the stage of development, density, relative spacing and site index were calculated. The gap frequency Poisson model was used to estimate the leaf area index. A new function is proposed to assess the effect of the foliage inclination on light transmittance for Scots pine, based on the assumption that the foliage elements show circular horizontal cross-sections. A new model was also developed to restrict the clumping effect on the vertical component in the Poisson model. The effect of the slope on LAI estimates was determined and included in the model. The results indicate that both the clumping effect correction and the slope correction improve the fitting of the Poisson model to the gap frequency experimental curves, and that the function proposed for the inclination angle correction may provide an acceptable approach for some coniferous species. The relationship between the parameters of the gap fraction model (the foliage inclination angle and the clumping coefficient) and stand level variables, that is, the stage of development, density, spacing of the stand and the site index, is analysed. © 2007 Elsevier B.V. All rights reserved.