Local adaptation shapes functional traits and resource allocation in black spruce.

Climate change is rapidly altering weather patterns, resulting in shifts in climatic zones. The survival of trees in specific locations depends on their functional traits. Local populations exhibit trait adaptations that ensure their survival and accomplishment of growth and reproduction processes during the growing season. Studying these traits offers valuable insights into species responses to present and future environmental conditions, aiding the implementation of measures to ensure forest resilience and productivity. This study investigates the variability in functional traits among five black spruce (Picea mariana (Mill.) B.S.P.) provenances originating from a latitudinal gradient along the boreal forest, and planted in a common garden in Quebec, Canada. We examined differences in bud phenology, growth performance, lifetime first reproduction, and the impact of a late-frost event on tree growth and phenological adjustments. The findings revealed that trees from northern sites exhibit earlier budbreak, lower growth increments, and reach reproductive maturity earlier than those from southern sites. Late-frost damage affected growth performance, but no phenological adjustment was observed in the successive year. Local adaptation in the functional traits may lead to maladaptation of black spruce under future climate conditions or serve as a potent evolutionary force promoting rapid adaptation under changing environmental conditions.

Small understorey trees have greater capacity than canopy trees to adjust hydraulic traits following prolonged experimental drought in a tropical forest.

Future climate change predictions for tropical forests highlight increased frequency and intensity of extreme drought events. However, it remains unclear whether large and small trees have differential strategies to tolerate drought due to the different niches they occupy. The future of tropical forests is ultimately dependent on the capacity of small trees (<10 cm in diameter) to adjust their hydraulic system to tolerate drought. To address this question, we evaluated whether the drought tolerance of neotropical small trees can adjust to experimental water stress and was different from tall trees. We measured multiple drought resistance-related hydraulic traits across nine common neotropical genera at the world's longest-running tropical forest throughfall-exclusion experiment and compared their responses with surviving large canopy trees. Small understorey trees in both the control and the throughfall-exclusion treatment had lower minimum stomatal conductance and maximum hydraulic leaf-specific conductivity relative to large trees of the same genera, as well as a greater hydraulic safety margin (HSM), percentage loss of conductivity and embolism resistance, demonstrating that they occupy a distinct hydraulic niche. Surprisingly, in response to the drought treatment, small trees increased specific hydraulic conductivity by 56.3% and leaf:sapwood area ratio by 45.6%. The greater HSM of small understorey trees relative to large canopy trees likely enabled them to adjust other aspects of their hydraulic systems to increase hydraulic conductivity and take advantage of increases in light availability in the understorey resulting from the drought-induced mortality of canopy trees. Our results demonstrate that differences in hydraulic strategies between small understorey and large canopy trees drive hydraulic niche segregation. Small understorey trees can adjust their hydraulic systems in response to changes in water and light availability, indicating that natural regeneration of tropical forests following long-term drought may be possible.

Changes in tree resistance, recovery and resilience across three successive extreme droughts in the northeast Iberian Peninsula.

Understanding which variables affect forest resilience to extreme drought is key to predict future dynamics under ongoing climate change. In this study, we analyzed how tree resistance, recovery and resilience to drought have changed along three consecutive droughts and how they were affected by species, tree size, plot basal area (as a proxy for competition) and climate. We focused on the three most abundant pine species in the northeast Iberian Peninsula: Pinus halepensis, P. nigra and P. sylvestris during the three most extreme droughts recorded in the period 1951-2010 (occurred in 1986, 1994, and 2005-2006). We cored trees from permanent sample plots and used dendrochronological techniques to estimate resistance (ability to maintain growth level during drought), recovery (growth increase after drought) and resilience (capacity to recover pre-drought growth levels) in terms of tree stem basal area increment. Mixed-effects models were used to determine which tree- and plot-level variables were the main determinants of resistance, recovery and resilience, and to test for differences among the studied droughts. Larger trees were significantly less resistant and resilient. Plot basal area effects were only observed for resilience, with a negative impact only during the last drought. Resistance, recovery and resilience differed across the studied drought events, so that the studied populations became less resistant, less resilient and recovered worse during the last two droughts. This pattern suggests an increased vulnerability to drought after successive drought episodes.

Death from drought in tropical forests is triggered by hydraulics not carbon starvation.

Drought threatens tropical rainforests over seasonal to decadal timescales, but the drivers of tree mortality following drought remain poorly understood. It has been suggested that reduced availability of non-structural carbohydrates (NSC) critically increases mortality risk through insufficient carbon supply to metabolism ('carbon starvation'). However, little is known about how NSC stores are affected by drought, especially over the long term, and whether they are more important than hydraulic processes in determining drought-induced mortality. Using data from the world's longest-running experimental drought study in tropical rainforest (in the Brazilian Amazon), we test whether carbon starvation or deterioration of the water-conducting pathways from soil to leaf trigger tree mortality. Biomass loss from mortality in the experimentally droughted forest increased substantially after >10 years of reduced soil moisture availability. The mortality signal was dominated by the death of large trees, which were at a much greater risk of hydraulic deterioration than smaller trees. However, we find no evidence that the droughted trees suffered carbon starvation, as their NSC concentrations were similar to those of non-droughted trees, and growth rates did not decline in either living or dying trees. Our results indicate that hydraulics, rather than carbon starvation, triggers tree death from drought in tropical rainforest.

Green spruce aphid infestations cause larger growth reductions to Sitka spruce under shade.

Light availability and infestation by the green spruce aphid (Elatobium abietinum) are key factors affecting the growth of Sitka spruce (Picea sitchensis) seedlings under a mature tree canopy, but their combined effect on seedling growth has not previously been quantified. A controlled outdoor experiment in which light levels (high light (HL): 100%, intermediate light (IL): 24%) and aphid infestation (absence/presence) were manipulated was conducted over 2 years to look at the effects on seedling growth and biomass distribution patterns. Aphid population assessments showed a significantly increased population density under IL, with three to four times higher cumulative aphid densities than that under HL. Defoliation rates of infested seedlings were directly related to aphid density. Total seedling biomass was strongly reduced in IL, and aphid infestation caused additional reductions in the biomass of particular components of the seedlings. Dry weight (DW) of older (≥1-year-old) needles in infested trees was significantly decreased in both years. Total root DW at the end of the second year was significantly affected by aphid infestation, and the reduction (14-18%) was similar in IL and HL treatments despite large differences in aphid density. Biomass distribution patterns in infested trees were similar to that of uninfested trees within each light treatment, indicating that the relative decreases in root biomass were accompanied by similar reductions in distribution to the above-ground parts of the seedlings. Leader extension growth of infested seedlings was reduced by 15-17% compared with uninfested seedlings under IL, whereas only a 2-3% reduction in leader extension of infested seedlings under HL was observed. The results showed that the response of seedlings to E. abietinum were primarily dependent on the light environment. The significant reduction caused by aphids on the total DW of older needles and roots, and on leader extension growth, does suggest the potential for effects to accumulate over time.

Hydraulic adjustment of Scots pine across Europe.

* The variability of branch-level hydraulic properties was assessed across 12 Scots pine populations covering a wide range of environmental conditions, including some of the southernmost populations of the species. The aims were to relate this variability to differences in climate, and to study the potential tradeoffs between traits. * Traits measured included wood density, radial growth, xylem anatomy, sapwood- and leaf-specific hydraulic conductivity (K(S) and K(L)), vulnerability to embolism, leaf-to-sapwood area ratio (A(L) : A(S)), needle carbon isotope discrimination (Delta13C) and nitrogen content, and specific leaf area. * Between-population variability was high for most of the hydraulic traits studied, but it was directly associated with climate dryness (defined as a combination of atmospheric moisture demand and availability) only for A(L) : A(S), K(L) and Delta13C. Shoot radial growth and A(L) : A(S) declined with stand development, which is consistent with a strategy to avoid exceedingly low water potentials as tree size increases. In addition, we did not find evidence at the intraspecific level of some associations between hydraulic traits that have been commonly reported across species. * The adjustment of Scots pine's hydraulic system to local climatic conditions occurred primarily through modifications of A(L) : A(S) and direct stomatal control, whereas intraspecific variation in vulnerability to embolism and leaf physiology appears to be limited.

Linking phloem function to structure: analysis with a coupled xylem-phloem transport model.

We carried out a theoretical analysis of phloem transport based on Münch hypothesis by developing a coupled xylem-phloem transport model. Results showed that the maximum sugar transport rate of the phloem was limited by solution viscosity and that transport requirements were strongly affected by prevailing xylem water potential. The minimum number of xylem and phloem conduits required to sustain transpiration and assimilation, respectively, were calculated. At its maximum sugar transport rate, the phloem functioned with a high turgor pressure difference between the sugar sources and sinks but the turgor pressure difference was reduced if additional parallel conduits were added or solute relays were introduced. Solute relays were shown to decrease the number of parallel sieve tubes needed for phloem transport, leading to a more uniform turgor pressure and allowing faster information transmission within the phloem. Because xylem water potential affected both xylem and phloem transport, the conductance of the two systems was found to be coupled such that large structural investments in the xylem reduced the need for investment in the phloem and vice versa.

Plasticity in hydraulic architecture of Scots pine across Eurasia.

Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G(s)) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G(s) to D (m/G (s,ref), where m is the slope and G(s,ref) is reference G(s) at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G (s,ref), and its absolute sensitivity to D(m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites.

Plant size, not age, regulates growth and gas exchange in grafted Scots pine trees.

We studied the effect of scion donor-tree age on the physiology and growth of 6- to 7-year-old grafted Scots pine (Pinus sylvestris L.) trees (4 and 5 years after grafting). Physiological measurements included photosynthethetic rate, stomatal conductance, transpiration, whole plant hydraulic conductance, needle nitrogen concentration and carbon isotope composition. Growth measurements included total and component biomasses, relative growth rates and growth efficiency. Scion donor trees ranged in age from 36 to 269 years at the time of grafting. Hydraulic conductance was measured gravimetrically, applying the Ohm's law analogy, and directly, with a high-pressure flow meter. We found no effect of scion donor-tree age on any of the variables measured. There was, however, great variation within scion donor-tree age groups, which was related to the size of the grafted trees. Differences in size may have been caused by variable initial grafting success, but there was no indication that grafting success and age were related. At the stem level, hydraulic conductance scaled with total leaf area so that total conductance per unit leaf area did not vary with crown size. However, leaf specific hydraulic conductance (gravimetric), transpiration, photosynthesis and stomatal conductance declined with increasing total tree leaf area and needle width. We hypothesize that needle width is inversely related to mesophyll conductance. We conclude that canopy and needle size and not scion donor-tree age determined gas exchange in our grafted trees.

Size-mediated ageing reduces vigour in trees.

There is increasing interest in understanding the costs and benefits of increased size and prolonged lifespan for plants. Some species of trees can grow more than 100 m in height and can live for several millennia, however whether these achievements are obtained at the cost of some other physiological functions is currently unclear. As increases in size are usually associated with ageing, it is also unclear whether observed reductions in growth rates and increased mortality rates are a function of size or of age per se. One theory proposes that reduced growth after the start of the reproductive phase is caused by cellular senescence. A second set of theories has focussed instead on plant size and the increased respiratory burdens or excessive height. We report on experimental manipulations to separate the effects of extrinsic factors such as size from those of intrinsic factors such as age for four tree species of contrasting phylogeny and life history. For each species, we measured growth, gas exchange and leaf biochemical properties for trees of different ages and sizes in the field and on propagated material obtained from the same genetic individuals but now all of small similar size in our common gardens. For all species, evidence indicated that size, not cellular senescence, accounted for the observed age-related declines in relative growth rates and net assimilation rates. Two species exhibited evidence of genetic control on leaf characters such as specific leaf area, although size also exerted an independent, and stronger, effect. We found partial support for the theory of hydraulic limitations to tree growth. The lack of a marked separation of soma and germline, an unlimited proliferation potential of meristem cells and the exponential increase in reproductive effort with size all help explain the lack of a senescence-induced decline in trees. It is possible that trees much older than the ones we sampled exhibit senescence symptoms.

Field measurements of ultrasonic acoustic emissions and stem diameter variations. New insight into the relationship between xylem tensions and embolism.

We examined interrelated xylem water tensions and embolism dynamics under field conditions by simultaneously monitoring ultra-acoustic emissions and changes in stem xylem diameter. Variation in stem xylem diameter was measured with linear displacement transducers to estimate variation in sap tension. Measured ultrasonic acoustic emissions coincided well with changes in xylem diameter, indicating that individual peaks in embolism occurred simultaneously with peaks in water tension. The good time resolution between measurements makes this method especially suitable for observing embolism dynamics on a short timescale. Longer lasting measurements can also be made to monitor inter-daily patterns in water tension and embolism because the techniques are non-destructive. Ultra-acoustic emissions occurred mainly during periods of decreasing stem xylem diameter, i.e., increasing water tension, when the water tension was high enough. Embolism also occurred during periods of increasing xylem diameter, i.e., decreasing water tension, but the number of embolizing conduits under these conditions was small.

Biomechanical and hydraulic determinants of tree structure in Scots pine: anatomical characteristics.

The development of anatomical, hydraulic and biomechanical properties in Scots pine (Pinus sylvestris L.) stems aged 7 to 59 years was followed. The hydraulic diameter and length of tracheids increased with age to a maximum at 15 and 35 years, respectively. Number of tracheids per unit of sapwood area decreased with age to a minimum of 500-600 tracheids mm(-2). Variations in specific hydraulic conductivity and Young's modulus of stems were associated with variation in anatomical properties. Over the time sequence considered, hydraulic and mechanical properties were positively related to each other and followed a similar developmental pattern, with no suggestion of a trade-off between the two. For most of the tree's life-cycle, heartwood made only a small contribution to whole-section mechanical stiffness because of its location close to the flexural neutral axis, and because of the presence of juvenile wood.

Hydraulic conductance, light interception and needle nutrient concentration in Scots pine stands and their relations with net primary productivity.

Aboveground xylem hydraulic conductance was determined in Scots pine (Pinus sylvestris L.) trees and stands from 7 to about 60 years of age. At the stand scale, leaf area index and net primary productivity (NPP, above- plus belowground) increased and reached a plateau at about 25-30 and 15-20 years, respectively; both parameters declined in mature stands. Stand hydraulic conductance followed a similar trend to NPP, with a maximum at about 15-20 years and a pronounced reduction in old stands. At the tree scale, annual biomass growth per unit of leaf area (growth efficiency) declined with tree age, whereas aboveground sapwood volume per unit leaf area, which is linearly related to maintenance respiration costs, steadily increased. Radiation interception per unit leaf area increased significantly with reduced leaf area index of mature stands, despite increased foliage clumping in the canopies of mature trees. Needle nutrient concentration did not change in the chronosequence. Tree hydraulic conductance per unit leaf area was strongly and positively correlated with growth efficiency. We discuss our findings in the context of growth reductions in mature and old trees, and suggest that increased hydraulic resistance and maintenance respiration costs may be the main causes of reduced carbon gain in mature and old trees.

Climate influences the leaf area/sapwood area ratio in Scots pine.

We tested the hypothesis that the leaf area/sapwood area ratio in Scots pine (Pinus sylvestris L.) is influenced by site differences in water vapor pressure deficit of the air (D). Two stands of the same provenance were selected, one in western Scotland and one in eastern England, so that effects resulting from age, genetic variability, density and fertility were minimized. Compared with the Scots pine trees at the cooler and wetter site in Scotland, the trees at the warmer and drier site in England produced less leaf area per unit of conducting sapwood area both at a stem height of 1.3 m and at the base of the live crown, whereas stem permeability was similar at both sites. Also, trees at the drier site had less leaf area per unit branch cross-sectional area at the branch base than trees at the wetter site. For each site, the average values for leaf area, sapwood area and permeability were used, together with values of transpiration rates at different D, to calculate average stem water potential gradients. Changes in the leaf area/sapwood area ratio acted to maintain a similar water potential gradient in the stems of trees at both sites despite climatic differences between the sites.

Evaporative demand determines branchiness of Scots pine.

Analysis of the branch area/stem area ratio of Scots pine growing in different climatic conditions in Europe and Siberia indicates that the branch area supported by a stem increases in warmer and drier conditions. The ratio was significantly correlated with several climatic variables, especially with potential evapotranspiration (E p). The ratio was negatively correlated with stand density (d s). A regression model combining E p and d s accounted for 85% of the total variation. These trends are believed to reflect hydraulic segmentation of trees and may represent a strategy to avoid cavitation in the tree, especially in the branches.

Increase of rooting ability in the woody species kiwi (Actinidia deliciosa A. Chev.) by transformation with Agrobacterium rhizogenes rol genes.

The woody species kiwi (Actinidia deliciosa A. Chev.), a male and late flowering clone of the cv Hayward, has been transformed by a T-DNA fragment encompassing rol A, B, C genes of A. rhizogenes. Transgenic plants, regenerated from leaf disc callus, showed the typical "hairy root" phenotype as described in herbaceous species. Explants from these plants (both leaf discs or 3 to 4 node leafy microcuttings) showed an increased ability to produce roots. Since root formation is one of the limiting factors in the vegetative propagation of woody species, the results have been discussed in relation to the use of A. rhizogenes rol genes in improving root morphogenesis in trees.