Does leaf gas exchange correlate with petiole xylem structural traits in Ulmus laevis seedlings under well-watered and drought stress conditions?

Several studies have shown that petiole xylem structure could be an important predictor of leaf gas exchange capacity, but the question of how petiole xylem structure relates to leaf gas exchange under different environment conditions remains unresolved. Moreover, knowledge of the amount of leaf gas exchange and structural variation that exists within a single species is also limited. In this study, we investigated the intraspecies coordination of leaf gas exchange and petiole xylem traits in 2-year-old seedlings of Ulmus laevis Pall. under well-watered and drought conditions. It was found that all studied petiole xylem traits of the elm seedlings were positively correlated with each other. This shows that the development of petiole xylem structure is internally well-coordinated. Nevertheless, the lower correlation coefficients between some petiole xylem traits indicate that the coordination is also individually driven. Drought stress reduced all studied leaf gas exchange traits and significantly increased intraspecies variation. In addition, drought stress also shifted the relationships between physiological traits and exhibited more structure-function relationships. This indicates the importance of petiole xylem structure in dictating water loss during drought stress and could partly explain the inconsistencies between leaf structure-function relationships studied under optimal conditions. Although several structure-function traits were related, the wide ranges of correlation coefficients indicate that the internal coordination of these traits substantially differs between individual elm seedlings. These findings are very important in the context of expected climatic change, as some degree of intraspecies variation in structure-function relationships could ensure the survival of some individuals under different environmental conditions.

Biotic threats for 23 major non-native tree species in Europe.

For non-native tree species with an origin outside of Europe a detailed compilation of enemy species including the severity of their attack is lacking up to now. We collected information on native and non-native species attacking non-native trees, i.e. type, extent and time of first observation of damage for 23 important non-native trees in 27 European countries. Our database includes about 2300 synthesised attack records (synthesised per biotic threat, tree and country) from over 800 species. Insects (49%) and fungi (45%) are the main observed biotic threats, but also arachnids, bacteria including phytoplasmas, mammals, nematodes, plants and viruses have been recorded. This information will be valuable to identify patterns and drivers of attacks, and trees with a lower current health risk to be considered for planting. In addition, our database will provide a baseline to which future impacts on non-native tree species could be compared with and thus will allow to analyse temporal trends of impacts.

Low resistance but high resilience to drought of flushing Norway spruce seedlings.

Spring drought episodes are becoming more frequent and intensive in European temperate forests. To study tree resilience to spring drought, Norway spruce seedlings were exposed to three levels of drought stress (well-watered (W), moderately stressed (M) and severely stressed (S)) for 42 days and then fully irrigated for 14 days. Drought strongly reduced gas exchange parameters for both M and S seedlings. After 42 days, stomatal conductance was lower by 83 and 97% in M and S, respectively, than in W seedlings. Respiration prevailed over photosynthesis in S seedlings at the end of the drought period. Drought mostly reduced longitudinal growth, especially in shoots and needles. Xylem growth reduction was caused mainly by a lower number of newly produced tracheids, not by changes in their size. Norway spruce seedlings showed good resilience to spring drought, as the observed physiological parameters started to recover after rewatering and seedlings started to sprout and form new tracheids. In M seedlings, all physiological traits recovered to the level of W seedlings during the 14-day irrigation period but the recovery took longer in S seedlings. Shoots and needles did not regrow in length but leaf mass per area increased during the recovery phase. To conclude, Norway spruce seedlings showed good resilience to spring single-drought event, but time necessary to full recovery from stress could make seedlings more vulnerable to recurrent drought events.

Machine Learning Guidance for Connection Tableaux.

Connection calculi allow for very compact implementations of goal-directed proof search. We give an overview of our work related to connection tableaux calculi: first, we show optimised functional implementations of connection tableaux proof search, including a consistent Skolemisation procedure for machine learning. Then, we show two guidance methods based on machine learning, namely reordering of proof steps with Naive Bayesian probabilities, and expansion of a proof search tree with Monte Carlo Tree Search.

The resistance and resilience of European beech seedlings to drought stress during the period of leaf development.

Spring drought is becoming a frequently occurring stress factor in temperate forests. However, the understanding of tree resistance and resilience to the spring drought remains insufficient. In this study, European beech (Fagus sylvatica L.) seedlings at the early stage of leaf development were moderately and severely drought stressed for 1 month and then subjected to a 2-week recovery period after rewatering. The study aimed to disentangle the complex relationships between leaf gas exchange, vascular anatomy, tree morphology and patterns of biomass allocation. Stomatal conductance decreased by 80 and 85% upon moderate and severe drought stress, respectively, which brought about a decline in net photosynthesis. However, drought did not affect the indices of slow chlorophyll fluorescence, indicating no permanent damage to the light part of the photosynthetic apparatus. Stem hydraulic conductivity decreased by more than 92% at both drought levels. Consequently, the cambial activity of stressed seedlings declined, which led to lower stem biomass, reduced tree ring width and a lower number of vessels in the current tree ring, these latter also with smaller dimensions. In contrast, the petiole structure was not affected, but at the cost of reduced leaf biomass. Root biomass was reduced only by severe drought. After rewatering, the recovery of gas exchange and regrowth of the current tree ring were observed, all delayed by several days and by lower magnitudes in severely stressed seedlings. The reduced stem hydraulic conductivity inhibited the recovery of gas exchange, but xylem function started to recover by regrowth and refilling of embolized vessels. Despite the damage to conductive xylem, no mortality occurred. These results suggest the low resistance but high resilience of European beech to spring drought. Nevertheless, beech resilience could be weakened if the period between drought events is short, as the recovery of severely stressed seedlings took longer than 14 days.

SoilTemp: A global database of near-surface temperature.

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long-term average thermal conditions at coarse spatial resolutions only. Hence, many climate-forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold-air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free-air temperatures, microclimatic ground and near-surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near-surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes.

Global effects of non-native tree species on multiple ecosystem services.

Non-native tree (NNT) species have been transported worldwide to create or enhance services that are fundamental for human well-being, such as timber provision, erosion control or ornamental value; yet NNTs can also produce undesired effects, such as fire proneness or pollen allergenicity. Despite the variety of effects that NNTs have on multiple ecosystem services, a global quantitative assessment of their costs and benefits is still lacking. Such information is critical for decision-making, management and sustainable exploitation of NNTs. We present here a global assessment of NNT effects on the three main categories of ecosystem services, including regulating (RES), provisioning (PES) and cultural services (CES), and on an ecosystem disservice (EDS), i.e. pollen allergenicity. By searching the scientific literature, country forestry reports, and social media, we compiled a global data set of 1683 case studies from over 125 NNT species, covering 44 countries, all continents but Antarctica, and seven biomes. Using different meta-analysis techniques, we found that, while NNTs increase most RES (e.g. climate regulation, soil erosion control, fertility and formation), they decrease PES (e.g. NNTs contribute less than native trees to global timber provision). Also, they have different effects on CES (e.g. increase aesthetic values but decrease scientific interest), and no effect on the EDS considered. NNT effects on each ecosystem (dis)service showed a strong context dependency, varying across NNT types, biomes and socio-economic conditions. For instance, some RES are increased more by NNTs able to fix atmospheric nitrogen, and when the ecosystem is located in low-latitude biomes; some CES are increased more by NNTs in less-wealthy countries or in countries with higher gross domestic products. The effects of NNTs on several ecosystem (dis)services exhibited some synergies (e.g. among soil fertility, soil formation and climate regulation or between aesthetic values and pollen allergenicity), but also trade-offs (e.g. between fire regulation and soil erosion control). Our analyses provide a quantitative understanding of the complex synergies, trade-offs and context dependencies involved for the effects of NNTs that is essential for attaining a sustained provision of ecosystem services.

Ecohydrological consequences of tree removal in an urban park evaluated using open data, free software and a minimalist measuring campaign.

With ongoing global climate change and an increasingly urbanized population, the importance of city parks and other forms of urban vegetation increases. Trees in urban parks can play an important role in mitigating runoff and delivering other ecosystem services. Park managers, E-NGOs, citizen scientists and others are increasingly called upon to evaluate the possible consequences of changes in park management such as, e.g., tree removal. Here, we present an unorthodox approach to hydrological modelling and its potential use in local policy making regarding urban greenery. The approach consists of a minimalist field campaign to characterize vegetation and soil moisture status combined with a novel model calibration using freely available data and software. During modelling, we were able to obtain coefficients of determination (R2) of 0.66 and 0.73 for probe-measured and simulated soil moisture under tree stand and park lawn land covers respectively. The results demonstrated that tree cover had a significant positive effect on the hydrological regime of the locality through interception, transpiration and effects on soil moisture. Simulations suggested that tree cover was twice as effective at mitigating runoff than park lawn and almost seven times better than impervious surfaces. In the case of a potential replacement of tree vegetation in favour of park lawn or impervious surfaces an increase in runoff of 14% and 81% respectively could be expected. The main conclusion drawn from our study was that such an approach can be a very useful tool for supporting local decision-making processes as it offers a freely available, cheap and relatively easy-to-use way to describe the hydrological consequences of landcover change (e.g., tree removal) with sufficient accuracy.

Tree differences in primary and secondary growth drive convergent scaling in leaf area to sapwood area across Europe.

Trees scale leaf (AL ) and xylem (AX ) areas to couple leaf transpiration and carbon gain with xylem water transport. Some species are known to acclimate in AL  : AX balance in response to climate conditions, but whether trees of different species acclimate in AL  : AX in similar ways over their entire (continental) distributions is unknown. We analyzed the species and climate effects on the scaling of AL vs AX in branches of conifers (Pinus sylvestris, Picea abies) and broadleaved (Betula pendula, Populus tremula) sampled across a continental wide transect in Europe. Along the branch axis, AL and AX change in equal proportion (isometric scaling: b Ëœ 1) as for trees. Branches of similar length converged in the scaling of AL vs AX with an exponent of b = 0.58 across European climates irrespective of species. Branches of slow-growing trees from Northern and Southern regions preferentially allocated into new leaf rather than xylem area, with older xylem rings contributing to maintaining total xylem conductivity. In conclusion, trees in contrasting climates adjust their functional balance between water transport and leaf transpiration by maintaining biomass allocation to leaves, and adjusting their growth rate and xylem production to maintain xylem conductance.

Stomatal conductance increases with rising temperature.

Stomatal conductance directly modifies plant water relations and photosynthesis. Many environmental factors affecting the stomatal conductance have been intensively studied but temperature has been largely neglected, even though it is one of the fastest changing environmental variables and it is rising due to climate change. In this study, we describe how stomata open when the temperature increases. Stomatal conductance increased by ca 40% in a broadleaf and a coniferous species, poplar (Populus deltoides x nigra) and loblolly pine (Pinus taeda) when temperature was increased by 10 °C, from 30 °C to 40 °C at a constant vapor pressure deficit of 1 kPa. The mechanism of regulating stomatal conductance by temperature was, at least partly, independent of other known mechanisms linked to water status and carbon metabolism. Stomatal conductance increased with rising temperature despite the decrease in leaf water potential, increase in transpiration, increase in intercellular CO2 concentration and was decoupled from photosynthesis. Increase in xylem and mesophyll hydraulic conductance coming from lower water viscosity may to some degree explain temperature dependent opening of stomata. The direct stomatal response to temperature allows plants to benefit from increased evaporative cooling during the heat waves and from lower stomatal limitations to photosynthesis but they may be jeopardized by faster depletion of soil water.

Increase in leaf temperature opens stomata and decouples net photosynthesis from stomatal conductance in Pinus taeda and Populus deltoides x nigra.

The effect of temperature on stomatal conductance (gs) and corresponding gas exchange parameters was studied in two tree species with contrasting leaf anatomy and ecophysiology-a broadleaf angiosperm, Populus deltoides x nigra (poplar), and a needle-leaf gymnosperm, Pinus taeda (loblolly pine). Experiments were conducted in growth chambers across a leaf temperature range of 19-48°C. Manipulations of temperature were done in well-watered and drought soil conditions and under ambient (400 ppm) and elevated (800 ppm) air CO2 concentrations. Increases in leaf temperature caused stomatal opening at both ambient and elevated [CO2]. The gs increased by 42% in poplar and by 40% in loblolly pine when leaf temperature increased from 30°C to 40°C at a vapour pressure difference of 1 kPa. Stomatal limitation to photosynthesis decreased in elevated temperature in loblolly pine but not in poplar. The ratio of net photosynthesis to gs depended on leaf temperature, especially at high temperatures. Evaporative cooling of transpiring leaves resulted in reductions in leaf temperature up to 9°C in well-watered poplar but only 1°C in drought-stressed poplar and in loblolly pine. As global mean temperatures rise and temperature extremes become more frequent and severe, understanding the effect of temperature on gs, and modelling that relationship, will become increasingly important.

Osmolality and Non-Structural Carbohydrate Composition in the Secondary Phloem of Trees across a Latitudinal Gradient in Europe.

Phloem osmolality and its components are involved in basic cell metabolism, cell growth, and in various physiological processes including the ability of living cells to withstand drought and frost. Osmolality and sugar composition responses to environmental stresses have been extensively studied for leaves, but less for the secondary phloem of plant stems and branches. Leaf osmotic concentration and the share of pinitol and raffinose among soluble sugars increase with increasing drought or cold stress, and osmotic concentration is adjusted with osmoregulation. We hypothesize that similar responses occur in the secondary phloem of branches. We collected living bark samples from branches of adult Pinus sylvestris, Picea abies, Betula pendula and Populus tremula trees across Europe, from boreal Northern Finland to Mediterranean Portugal. In all studied species, the observed variation in phloem osmolality was mainly driven by variation in phloem water content, while tissue solute content was rather constant across regions. Osmoregulation, in which osmolality is controlled by variable tissue solute content, was stronger for Betula and Populus in comparison to the evergreen conifers. Osmolality was lowest in mid-latitude region, and from there increased by 37% toward northern Europe and 38% toward southern Europe due to low phloem water content in these regions. The ratio of raffinose to all soluble sugars was negligible at mid-latitudes and increased toward north and south, reflecting its role in cold and drought tolerance. For pinitol, another sugar known for contributing to stress tolerance, no such latitudinal pattern was observed. The proportion of sucrose was remarkably low and that of hexoses (i.e., glucose and fructose) high at mid-latitudes. The ratio of starch to all non-structural carbohydrates increased toward the northern latitudes in agreement with the build-up of osmotically inactive C reservoir that can be converted into soluble sugars during winter acclimation in these cold regions. Present results for the secondary phloem of trees suggest that adjustment with tissue water content plays an important role in osmolality dynamics. Furthermore, trees acclimated to dry and cold climate showed high phloem osmolality and raffinose proportion.

Long-term impact of Ophiostoma novo-ulmi on leaf traits and transpiration of branches in the Dutch elm hybrid 'Dodoens'.

To better understand the long-term impact of Ophiostoma novo-ulmi Brasier on leaf physiology in 'Dodoens', a Dutch elm disease-tolerant hybrid, measurements of leaf area, leaf dry mass, petiole anatomy, petiole hydraulic conductivity, leaf and branch water potential, and branch sap flow were performed 3 years following an initial artificial inoculation. Although fungal hyphae were detected in fully expanded leaves, neither anatomical nor morphological traits were affected, indicating that there was no impact from the fungal hyphae on the leaves during leaf expansion. In contrast, however, infected trees showed both a lower transpiration rate of branches and a lower sap flow density. The long-term persistence of fungal hyphae inside vessels decreased the xylem hydraulic conductivity, but stomatal regulation of transpiration appeared to be unaffected as the leaf water potential in both infected and non-infected trees was similarly driven by the transpirational demands. Regardless of the fungal infection, leaves with a higher leaf mass per area ratio tended to have a higher leaf area-specific conductivity. Smaller leaves had an increased number of conduits with smaller diameters and thicker cell walls. Such a pattern could increase tolerance towards hydraulic dysfunction. Measurements of water potential and theoretical xylem conductivity revealed that petiole anatomy could predict the maximal transpiration rate. Three years following fungal inoculation, phenotypic expressions for the majority of the examined traits revealed a constitutive nature for their possible role in Dutch elm disease tolerance of 'Dodoens' trees.

Effects of prolonged drought on the anatomy of sun and shade needles in young Norway spruce trees.

Predicted increases in the frequency and duration of drought are expected to negatively affect tree vitality, but we know little about how water shortage will influence needle anatomy and thereby the trees' photosynthetic and hydraulic capacity. In this study, we evaluated anatomical changes in sun and shade needles of 20-year-old Norway spruce trees exposed to artificial drought stress. Canopy position was found to be important for needle structure, as sun needles had significantly higher values than shade needles for all anatomical traits (i.e., cross-sectional needle area, number of tracheids in needle, needle hydraulic conductivity, and tracheid lumen area), except proportion of xylem area per cross-sectional needle area. In sun needles, drought reduced all trait values by 10-40%, whereas in shade needles, only tracheid maximum diameter was reduced by drought. Due to the relatively weaker response of shade needles than sun needles in drought-stressed trees, the difference between the two needle types was reduced by 25% in the drought-stressed trees compared to the control trees. The observed changes in needle anatomy provide new understanding of how Norway spruce adapts to drought stress and may improve predictions of how forests will respond to global climate change.

Learning-assisted theorem proving with millions of lemmas.

Large formal mathematical libraries consist of millions of atomic inference steps that give rise to a corresponding number of proved statements (lemmas). Analogously to the informal mathematical practice, only a tiny fraction of such statements is named and re-used in later proofs by formal mathematicians. In this work, we suggest and implement criteria defining the estimated usefulness of the HOL Light lemmas for proving further theorems. We use these criteria to mine the large inference graph of the lemmas in the HOL Light and Flyspeck libraries, adding up to millions of the best lemmas to the pool of statements that can be re-used in later proofs. We show that in combination with learning-based relevance filtering, such methods significantly strengthen automated theorem proving of new conjectures over large formal mathematical libraries such as Flyspeck.

Leaf trait dissimilarities between Dutch elm hybrids with a contrasting tolerance to Dutch elm disease.

BACKGROUND AND AIMS: Previous studies have shown that Ophiostoma novo-ulmi, the causative agent of Dutch elm disease (DED), is able to colonize remote areas in infected plants of Ulmus such as the leaf midrib and secondary veins. The objective of this study was to compare the performances in leaf traits between two Dutch elm hybrids 'Groeneveld' and 'Dodoens' which possess a contrasting tolerance to DED. Trait linkages were also tested with leaf mass per area (LMA) and with the reduced Young's modulus of elasticity (MOE) as a result of structural, developmental or functional linkages. METHODS: Measurements and comparisons were made of leaf growth traits, primary xylem density components, gas exchange variables and chlorophyll a fluorescence yields between mature plants of 'Groeneveld' and 'Dodoens' grown under field conditions. A recently developed atomic force microscopy technique, PeakForce quantitative nanomechanical mapping, was used to reveal nanomechanical properties of the cell walls of tracheary elements such as MOE, adhesion and dissipation. KEY RESULTS: 'Dodoens' had significantly higher values for LMA, leaf tissue thickness variables, tracheary element lumen area (A), relative hydraulic conductivity (RC), gas exchange variables and chlorophyll a fluorescence yields. 'Groeneveld' had stiffer cell walls of tracheary elements, and higher values for water-use efficiency and leaf water potential. Leaves with a large carbon and nutrient investment in LMA tended to have a greater leaf thickness and a higher net photosynthetic rate, but LMA was independent of RC. Significant linkages were also found between the MOE and some vascular traits such as RC, A and the number of tracheary elements per unit area. CONCLUSIONS: Strong dissimilarities in leaf trait performances were observed between the examined Dutch elm hybrids. Both hybrids were clearly separated from each other in the multivariate leaf trait space. Leaf growth, vascular and gas exchange traits in the infected plants of 'Dodoens' were unaffected by the DED fungus. 'Dodoens' proved to be a valuable elm germplasm for further breeding strategies.

Effect of thinning on anatomical adaptations of Norway spruce needles.

Conifers and other trees are constantly adapting to changes in light conditions, water/nutrient supply and temperatures by physiological and morphological modifications of their foliage. However, the relationship between physiological processes and anatomical characteristics of foliage has been little explored in trees. In this study we evaluated needle structure and function in Norway spruce families exposed to different light conditions and transpiration regimes. We compared needle characteristics of sun-exposed and shaded current-year needles in a control plot and a thinned plot with 50% reduction in stand density. Whole-tree transpiration rates remained similar across plots, but increased transpiration of lower branches after thinning implies that sun-exposed needles in the thinned plot were subjected to higher water stress than sun-exposed needles in the control plot. In general, morphological and anatomical needle parameters increased with increasing tree height and light intensity. Needle width, needle cross-section area, needle stele area and needle flatness (the ratio of needle thickness to needle width) differed most between the upper and lower canopy. The parameters that were most sensitive to the altered needle water status of the upper canopy after thinning were needle thickness, needle flatness and percentage of stele area in needle area. These results show that studies comparing needle structure or function between tree species should consider not only tree height and light gradients, but also needle water status. Unaccounted for differences in needle water status may have contributed to the variable relationship between needle structure and irradiance that has been observed among conifers.

Assessing the applicability of the earth impedance method for in situ studies of tree root systems.

Several electrical methods have been introduced as non-invasive techniques to overcome the limited accessibility to root systems. Among them, the earth impedance method (EIM) represents the most recent development. Applying an electrical field between a cormus and the rooted soil, the EIM measures the absorptive root surface area (ARSA) from grounding resistance patterns. Allometric relationships suggested that this method was a valuable tool. Crucial assumptions for the applicability of the EIM, however, have not been tested experimentally. Focusing on tree root systems, the present study assesses the applicability of the EIM. Six hypotheses, deduced from the EIM approach, were tested in several experiments and the results were compared with conventional methods. None of the hypotheses could be verified and the results allow two major conclusions. First, in terms of an analogue electrical circuit, a tree-root-soil continuum appears as a serial circuit with xylem and soil resistance being the dominant components. Allometric variation in contact resistance, with the latter being the proxy for root surface area, are thus overruled by the spatial and seasonal variation of soil and xylem resistances. Second, in a tree-root-soil continuum, distal roots conduct only a negligible portion of the electric charge. Most of charge carriers leave the root system in the proximal parts of the root-soil interface.