Coordination between water relations strategy and carbon investment in leaf and stem in six fruit tree species.

The water relation strategy is a key issue in climate change. Given the difficulty of determining water relations strategy, there is a need for simple traits with a solid theoretical basis to estimate it. Traits associated with resource allocation patterns along a 'fast-slow' plant economics spectrum are particularly compelling, reflecting trade-offs between growth rate and carbon allocation. Avocado (Persea americana ), fig tree (Ficus carica ), mandarin (Citrus reticulata ), olive (Olea europaea ), pomegranate (Punica granatum ), and grapevine (Vitis vinifera ) were characterised in terms of iso-anisohydric strategy through stomatal behaviour, water potential at the turgor loss point (TLP), and hydroscape area. Additionally, the association of these metrics with leaf mass per area (LMA) and wood density (WDen) was explored. We observed high coordination between LMA and WDen, and both traits were related to metrics of water relation strategy. More anisohydric species tended to invest more carbon per unit leaf area or unit stem volume, which has implications for hydraulic efficiency and water stress tolerance. WDen and TLP were the most powerful traits in estimating the water relation strategy for six fruit species. These traits are easy to measure, time-cost efficient, and appear central to coordinating multiple traits and behaviours along the water relations strategies.

Linkages between stem vulnerability curves and tree demography and their implications for plant physiological modeling.

Vulnerability curves (VCs) have been measured extensively to describe the differences in plant vulnerability to cavitation. Although the roles of hydraulic conductivity (Ks,max) and hydraulic safety (P50, embolism resistance), both of which are parameters of VCs ('sigmoidal' type), in tree demography have been evaluated across different forests, the direct linkages between VCs and tree demography are rarely explored. In this study, we combined measured VCs and plot data of 16 tree species in Panamanian seasonal tropical forests to investigate the connections between VCs and tree mortality, recruitment and growth. We found that the mortality and recruitment rates of evergreen species were most significantly positively correlated with P50. However, the mortality and recruitment rates of deciduous species only exhibited significant positive correlations with parameter a, which describes the steepness of VCs and indicates the sensitivity of conductivity loss with water potential decline, but is often neglected. These differences among evergreen and deciduous species may contribute to the poor performance of existing quantitative relationships (such as the fitting relationships for all 16 species) in capturing tree mortality and recruitment dynamics. Additionally, evergreen species presented a significant positive relationship between relative growth rate (RGR) and Ks,max, while deciduous species did not display such relationship. The RGR of both evergreen and deciduous species also displayed no significant correlations with P50 and a. Further analysis demonstrated that species with steeper VCs tended to have high mortality and recruitment rates, while species with flatter VCs were usually those with low mortality and recruitment rates. Our results highlight the important role of parameter a in tree demography, especially for deciduous species. Given that VC is a key component of plant hydraulic models, integrating measured VC rather than optimizing its parameters will help improve the ability to simulate and predict forest response to water availability.

Linking root and stem hydraulic traits to leaf physiological parameters in Coffea canephora clones with contrasting drought tolerance.

Knowing the key hydraulic traits of different genotypes at early seedling stages can potentially provide crucial information and save time for breeding programs. In the current study we investigated: (1) how root, stem and whole plant conductivities are linked to xylem traits, and (2) how the integrated hydraulic system impacts leaf water potential, gas exchange, chlorophyll a fluorescence and the growth of three coffee cultivars (clones of Coffea canephora Pierre ex Froehner cv. Conilon) with known differences in drought tolerance. The Conilon clones CL 14, CL 5 V and CL 109A, classified as tolerant, moderately tolerant, and sensitive to drought respectively, were grown under non-limiting soil-water supply but high atmospheric demand (i.e., high VPDair). CL 14 and CL 5 V displayed higher root and stem hydraulic conductance and conductivity, and higher whole plant conductivity than CL 109A, and these differences were associated with higher root growth traits. In addition, CL 109A exhibited a non-significant trend towards wider vessels. Collectively, these responses likely contributed to reduce leaf water potential in CL 109A, and in turn, reduced leaf gas exchange, especially during elevated VPDair. Even when grown under well-watered conditions, the elevated VPDair observed during this study resulted in key differences in the hydraulic traits between the cultivars corresponding to differences in plant water status, gas exchange, and photochemical activity. Together these results suggest that coffee hydraulic traits, even when grown under non-water stress conditions, can be considered in breeding programs targeting more productive and efficient genotypes under drought and high atmospheric demand.

The Pneumatron: An automated pneumatic apparatus for estimating xylem vulnerability to embolism at high temporal resolution.

Xylem vulnerability to embolism represents an important trait to determine species distribution patterns and drought resistance. However, estimating embolism resistance frequently requires time-consuming and ambiguous hydraulic lab measurements. Based on a recently developed pneumatic method, we present and test the "Pneumatron", a device that generates high time-resolution and fully automated vulnerability curves. Embolism resistance is estimated by applying a partial vacuum to extract air from an excised xylem sample, while monitoring the pressure change over time. Although the amount of gas extracted is strongly correlated with the percentage loss of xylem conductivity, validation of the Pneumatron was performed by comparison with the optical method for Eucalyptus camaldulensis leaves. The Pneumatron improved the precision of the pneumatic method considerably, facilitating the detection of small differences in the (percentage of air discharged [PAD] < 0.47%). Hence, the Pneumatron can directly measure the 50% PAD without any fitting of vulnerability curves. PAD and embolism frequency based on the optical method were strongly correlated (r2 = 0.93) for E. camaldulensis. By providing an open source platform, the Pneumatron represents an easy, low-cost, and powerful tool for field measurements, which can significantly improve our understanding of plant-water relations and the mechanisms behind embolism.

C4 and crassulacean acid metabolism within a single leaf: deciphering key components behind a rare photosynthetic adaptation.

Although biochemically related, C4 and crassulacean acid metabolism (CAM) systems are expected to be incompatible. However, Portulaca species, including P. oleracea, operate C4 and CAM within a single leaf, and the mechanisms behind this unique photosynthetic arrangement remain largely unknown. Here, we employed RNA-seq to identify candidate genes involved exclusively or shared by C4 or CAM, and provided an in-depth characterization of their transcript abundance patterns during the drought-induced photosynthetic transitions in P. oleracea. Data revealed fewer candidate CAM-specific genes than those recruited to function in C4 . The putative CAM-specific genes were predominantly involved in night-time primary carboxylation reactions and malate movement across the tonoplast. Analysis of gene transcript-abundance regulation and photosynthetic physiology indicated that C4 and CAM coexist within a single P. oleracea leaf under mild drought conditions. Developmental and environmental cues were shown to regulate CAM expression in stems, whereas the shift from C4 to C4 -CAM hybrid photosynthesis in leaves was strictly under environmental control. Moreover, efficient starch turnover was identified as part of the metabolic adjustments required for CAM operation in both organs. These findings provide insights into C4 /CAM connectivity and compatibility, contributing to a deeper understanding of alternative ways to engineer CAM into C4 crop species.

Hurricane María tripled stem breaks and doubled tree mortality relative to other major storms.

Tropical cyclones are expected to intensify under a warming climate, with uncertain effects on tropical forests. One key challenge to predicting how more intense storms will influence these ecosystems is to attribute impacts specifically to storm meteorology rather than differences in forest characteristics. Here we compare tree damage data collected in the same forest in Puerto Rico after Hurricanes Hugo (1989, category 3), Georges (1998, category 3), and María (2017, category 4). María killed twice as many trees as Hugo, and for all but two species, broke 2- to 12-fold more stems than the other two storms. Species with high density wood were resistant to uprooting, hurricane-induced mortality, and were protected from breakage during Hugo but not María. Tree inventories and a wind exposure model allow us to attribute these differences in impacts to storm meteorology. A better understanding of risk factors associated with tree species susceptibility to severe storms is key to predicting the future of forest ecosystems under climate warming.

Can climate and soil conditions change the morpho-anatomy among individuals from different localities? A case study in Aldama grandiflora (Asteraceae) / Condições climáticas e de solo podem alterar a morfo-anatomia entre indivíduos de diferentes localidades? Um estudo de caso em Aldama grandiflora (Asteraceae)

Abstract Vegetative aerial organs are considerably more exposed to environmental conditions and can reflect the specific adaptations of plants to their local environment. Aldama grandiflora species are known to be widely distributed in Brazil; therefore, individuals from different populations of this species are thought to be exposed to different abiotic and biotic conditions. Several anatomical studies conducted on Brazilian Aldama species have mainly focused on the qualitative anatomical characters or traits of these species, but not on their quantitative traits. In this study, we evaluated whether climate and soil conditions can change the morphometry among individuals of A. grandiflora collected from six sites in the Goiás State, Brazil, by assessing their anatomical characters. Further, soil sampling was performed, and climate data were collected from all the six sites. The analysis indicated few statistical differences among the populations evaluated, showing that A. grandiflora presented consistent leaf and stem anatomical characteristics. The small morpho-anatomical differences found among individuals of the different populations evaluated, reflected the soil conditions in which these populations were grown. Therefore, environmental factors have a significant influence on the morpho-anatomy of Aldama grandiflora.

Resumo Os órgãos vegetativos aéreos estão consideravelmente mais expostos às condições ambientais e podem refletir as adaptações específicas das plantas ao seu habitat. A espécie Aldama grandiflora é amplamente distribuída no Brasil e, dessa forma, indivíduos de diferentes populações podem estar expostos a diferentes condições ambientais. Vários estudos anatômicos realizados com espécies brasileiras do gênero Aldama têm abordado, principalmente, as características anatômicas qualitativas dessas espécies, mas não em suas características quantitativas. Neste estudo avaliamos se as condições climáticas e do solo podem alterar a morfometria entre os indivíduos de A. grandiflora coletados em seis populações do Estado de Goiás. Foram avaliados os caracteres anatômicos foliares e caulinares, além da amostragem do solo e coleta de dados climáticos, para os seis locais. A análise indicou algumas diferenças estatísticas entre as populações avaliadas, mostrando que A. grandiflora apresentou características anatômicas foliares e caulinares bastante consistentes. As pequenas diferenças morfo-anatômicas encontradas entre indivíduos das diferentes populações avaliadas, refletiram as condições do solo nos quais essas populações se desenvolveram. Assim sendo, fatores ambientais relacionados ao clima e condições do solo têm uma influência significativa sobre a morfo-anatomia de Aldama grandiflora.

Living on borrowed time - Amazonian trees use decade-old storage carbon to survive for months after complete stem girdling.

Nonstructural carbon (NSC) reserves act as buffers to sustain tree activity during periods when carbon (C) assimilation does not meet C demand, but little is known about their age and accessibility; we designed a controlled girdling experiment in the Amazon to study tree survival on NSC reserves. We used bomb-radiocarbon (14 C) to monitor the time elapsed between C fixation and release ('age' of substrates). We simultaneously monitored how the mobilization of reserve C affected δ13 CO2 . Six ungirdled control trees relied almost exclusively on recent assimilates throughout the 17 months of measurement. The Δ14 C of CO2 emitted from the six girdled stems increased significantly over time after girdling, indicating substantial remobilization of storage NSC fixed up to 13-14 yr previously. This remobilization was not accompanied by a consistent change in observed δ13 CO2 . These trees have access to storage pools integrating C accumulated over more than a decade. Remobilization follows a very clear reverse chronological mobilization with younger reserve pools being mobilized first. The lack of a shift in the δ13 CO2 might indicate a constant contribution of starch hydrolysis to the soluble sugar pool even outside pronounced stress periods (regular mixing).

Carbon limitation, stem growth rate and the biomechanical cause of Corner's rules.

Background and aims: Corner's rules describe a global spectrum from large-leaved plants with thick, sparingly branched twigs with low-density stem tissues and thick piths to plants with thin, highly branched stems with high-density stem tissues and thin piths. The hypothesis was tested that, if similar crown areas fix similar amounts of carbon regardless of leaf size, then large-leaved species, with their distantly spaced leaves, require higher stem growth rates, lower stem tissue densities and stiffnesses, and therefore thicker twigs. Methods: Structural equation models were used to test the compatibility of this hypothesis with a dataset on leaf size, shoot tip spacing, stem growth rate and dimensions, and tissue density and mechanics, sampling 55 species drawn from across the angiosperm phylogeny from a morphologically diverse dry tropical community. Key results: Very good fit of structural equation models showed that the causal model is highly congruent with the data. Conclusions: Given similar amounts of carbon to allocate to stem growth, larger-leaved species require greater leaf spacing and therefore greater stem extension rates and longer stems, in turn requiring lower-density, more flexible, stem tissues than small-leaved species. A given stem can have high resistance to bending because it is thick (has high second moment of area I) or because its tissues are stiff (high Young's modulus E), the so-called E-I trade-off. Because of the E-I trade-off, large-leaved species have fast stem growth rates, low stem tissue density and tissue stiffness, and thick twigs with wide piths and thick bark. The agreement between hypothesis and data in structural equation analyses strongly suggests that Corner's rules emerge as the result of selection favouring the avoidance of self-shading in the context of broadly similar rates of carbon fixation per unit crown area across species.

Accelerated diversification and functional trait evolution in Velloziaceae reveal new insights into the origins of the campos rupestres' exceptional floristic richness.

Background and Aims: The greater diversity of plant clades in the Neotropics compared to their relatives in Africa is a pervasive pattern in biogeography. To better understand the causes of this imbalance, we studied the diversification dynamics of the monocot family Velloziaceae. In addition to being conspicuously richer in the Neotropics compared to the Palaeotropics, many species of Velloziaceae exhibit extreme desiccation tolerance (i.e. 'resurrection' behaviour), and other ecological specializations to life on rocky outcrops, poor sandy soils, open vegetation and seasonally dry climates. Velloziaceae is also ecologically dominant in the campos rupestres, a habitat having exceptionally high plant diversity and endemism in Brazil. Methods: We reconstructed a densely sampled time-calibrated molecular phylogeny and used state-dependent and state-independent models to estimate rates of lineage diversification in relation to continent-scale geographical occurrence and functional traits associated with desiccation tolerance and water storage capacity. Key Results: Independent shifts to faster diversification occurred within two Neotropical lineages, Vellozia and Barbacenia. The Vellozia radiation was associated with the presence of conspicuous aerial stems, and was followed by decreasing diversification rates during the Oligocene, a time of rising global temperatures and expanding open areas around the world. The Barbacenia radiation was faster and more recent, occurring during the cooling conditions of the Miocene, and associated with the acquisition of aquiferous parenchyma on the leaves. Conclusions: High species richness of Velloziaceae in South America has been driven by faster diversification in lineages predominantly occurring in the campos rupestres, putatively by the evolution of adaptive strategies in response to independent climatic events. The radiation of Vellozia in particular might have played a key role in the assembly of the campos rupestres vegetation.

Near-infrared spectrometry allows fast and extensive predictions of functional traits from dry leaves and branches.

The linking of individual functional traits to ecosystem processes is the basis for making generalizations in ecology, but the measurement of individual values is laborious and time consuming, preventing large-scale trait mapping. Also, in hyper-diverse systems, errors occur because identification is difficult, and species level values ignore intra-specific variation. To allow extensive trait mapping at the individual level, we evaluated the potential of Fourrier-Transformed Near Infra-Red Spectrometry (FT-NIR) to adequately describe 14 traits that are key for plant carbon, water, and nutrient balance. FT-NIR absorption spectra (1,000-2,500 nm) were obtained from dry leaves and branches of 1,324 trees of 432 species from a hyper-diverse Amazonian forest. FT-NIR spectra were related to measured traits for the same plants using partial least squares regressions. A further 80 plants were collected from a different site to evaluate model applicability across sites. Relative prediction error (RMSErel ) was calculated as the percentage of the trait value range represented by the final model RMSE. The key traits used in most functional trait studies; specific leaf area, leaf dry matter content, wood density and wood dry matter content can be well predicted by the model (R2  = 0.69-0.78, RMSErel  = 9-11%), while leaf density, xylem proportion, bark density and bark dry matter content can be moderately well predicted (R2  = 0.53-0.61, RMSErel  = 14-17%). Community-weighted means of all traits were well estimated with NIR, as did the shape of the frequency distribution of the community values for the above key traits. The model developed at the core site provided good estimations of the key traits of a different site. An evaluation of the sampling effort indicated that 400 or less individuals may be sufficient for establishing a good local model. We conclude that FT-NIR is an easy, fast and cheap method for the large-scale estimation of individual plant traits that was previously impossible. The ability to use dry intact leaves and branches unlocks the potential for using herbarium material to estimate functional traits; thus advancing our knowledge of community and ecosystem functioning from local to global scales.

Can climate and soil conditions change the morpho-anatomy among individuals from different localities? A case study in Aldama grandiflora (Asteraceae).

Vegetative aerial organs are considerably more exposed to environmental conditions and can reflect the specific adaptations of plants to their local environment. Aldama grandiflora species are known to be widely distributed in Brazil; therefore, individuals from different populations of this species are thought to be exposed to different abiotic and biotic conditions. Several anatomical studies conducted on Brazilian Aldama species have mainly focused on the qualitative anatomical characters or traits of these species, but not on their quantitative traits. In this study, we evaluated whether climate and soil conditions can change the morphometry among individuals of A. grandiflora collected from six sites in the Goiás State, Brazil, by assessing their anatomical characters. Further, soil sampling was performed, and climate data were collected from all the six sites. The analysis indicated few statistical differences among the populations evaluated, showing that A. grandiflora presented consistent leaf and stem anatomical characteristics. The small morpho-anatomical differences found among individuals of the different populations evaluated, reflected the soil conditions in which these populations were grown. Therefore, environmental factors have a significant influence on the morpho-anatomy of Aldama grandiflora.

Physiological differences between root suckers and saplings enlarge the regeneration niche in Eucryphia cordifolia Cav.

Many clonal plants produce vegetative recruits that remain connected to the parent plant. Such connections permit resource sharing among ramets, explaining the high survival rates of vegetative recruits during establishment under suboptimal conditions for sexual regeneration. We propose that differences in the regeneration niches of sexual and vegetative recruits reflect different physiological adjustments caused by parental supply of resources to the ramets. We conducted ecophysiological measurements in saplings and root suckers of Eucryphia cordifolia Cav., a tree species of the temperate rainforest of southern South America. We compared the following traits of saplings and suckers: gas exchange at the leaf level, crown architecture, daily crown carbon balance, biomass allocation to above-ground tissues (leaf-to-stem mass ratio, leaf mass area and leaf area ratio), xylem anatomy traits (lumen vessel fraction, vessel density and size) and stem ring width. We also correlated the growth rates of saplings and suckers with relevant environmental data (light and climate). Saplings showed morphological, architectural and physiological traits that enhance daily crown carbon balance and increase water-use efficiency, in order to supply their growth demands while minimizing water loss per unit of carbon gained. The radial growth of saplings diminished under dry conditions, which suggests a strong stomatal sensitivity to water availability. Suckers have low stomatal conductance, likely because the carbon supplied by the parent plant diminishes the necessity of high rates of photosynthesis. The low responsiveness of sucker growth to temporal changes in water availability also supports the existence of parental supply. The physiological differences between sexual and vegetative recruits satisfactorily explain the ecological niche of E. cordifolia, with saplings restricted to more closed and humid sites.

Role of tree size in moist tropical forest carbon cycling and water deficit responses.

Drought disproportionately affects larger trees in tropical forests, but implications for forest composition and carbon (C) cycling in relation to dry season intensity remain poorly understood. In order to characterize how C cycling is shaped by tree size and drought adaptations and how these patterns relate to spatial and temporal variation in water deficit, we analyze data from three forest dynamics plots spanning a moisture gradient in Panama that have experienced El Niño droughts. At all sites, aboveground C cycle contributions peaked below 50-cm stem diameter, with stems ≥ 50 cm accounting for on average 59% of live aboveground biomass, 45% of woody productivity and 49% of woody mortality. The dominance of drought-avoidance strategies increased interactively with stem diameter and dry season intensity. Although size-related C cycle contributions did not vary systematically across the moisture gradient under nondrought conditions, woody mortality of larger trees was disproportionately elevated under El Niño drought stress. Thus, large (> 50 cm) stems, which strongly mediate but do not necessarily dominate C cycling, have drought adaptations that compensate for their more challenging hydraulic environment, particularly in drier climates. However, these adaptations do not fully buffer the effects of severe drought, and increased large tree mortality dominates ecosystem-level drought responses.

Trait shifts associated with the subshrub life-history strategy in a tropical savanna.

Over the past 10 million years, tropical savanna environments have selected for small growth forms within woody plant lineages. The result has been the evolution of subshrubs (geoxyles), presumably as an adaptation to frequent fire. To evaluate the traits associated with the shift from tree to subshrub growth forms, we compared seed biomass, germination, survival, resprouting, biomass allocation, and photosynthesis between congeneric trees and subshrubs, and quantified phylogenetic conservatism. Despite large differences in adult morphology between trees and subshrub species, the differences are modest in seedlings, and most of the variation in traits was explained by genus, indicating considerable phylogenic conservatism. Regardless, tree seedlings invested more heavily in aboveground growth, compared to subshrubs, which is consistent with the adult strategy of savanna trees, which depend on a large resistant-fire stem. Subshrub seedlings also invest in greater non-structural carbohydrate reserves, likely as an adaptation to the high fire frequencies typical of tropical savannas. The modest differences as seedlings suggest that selective pressures during early development may not have contributed substantially to the evolution of the subshrub growth form and that the distinct allocation and life history must arise later in life. This is consistent with the interpretation that the subshrub growth form arose as a life-history strategy in which maturity is reached at a small stem size, allowing them to reproduce despite repeated fire-induced topkill. The convergent evolution of subshrubs within multiple tree lineages reaffirms the importance of fire in the origin and diversification of the flora of mesic savannas.

Methyl jasmonate and salicylic acid are able to modify cell wall but only salicylic acid alters biomass digestibility in the model grass Brachypodium distachyon.

In addition to playing a key role in the response to environmental changes, cell walls are also considered as a valuable feedstock for cellulosic ethanol. Here we explored the effects of the stress-response hormones, salicylic acid and methyl jasmonate, on cell wall biosynthesis and biomass digestibility in Brachypodium distachyon, a species recently considered as a suitable model for biomass conversion. We found that in response to salicylic acid or methyl jasmonate treatment, plant growth was reduced coupled with significant changes in cell wall composition. Cellulose content increased in response to methyl jasmonate whereas a reduction in lignin content was found after salicylic acid application. Moreover, hemicellulose composition was altered and increases in caffeic acid, ferulic acid and p-coumaric acid content were detected in response to both treatments. The hormonal profile and the expression pattern of genes involved in cell wall biosynthesis were also modified. Biomass digestibility was reduced in leaf tissue after salicylic acid treatment and was negatively correlated with ferulic acid and p-coumaric acid content. The results obtained here aid in our understanding of cell wall dynamics in response to stress and will enable the development of new strategies to improve cell wall digestibility in bioenergy feedstock.

Does water availability regulate biomass partitioning between trunk and branches?

The extent to which a vertical trunk is differentiated from its branches is a key trait for the architecture of trees and may affect interspecific relationships. In this study, we analysed the effect of soil water availability on biomass partitioning for Nothofagus pumilio by means of a nursery experiment. Juvenile trees were subject to three irrigation conditions: no irrigation, intermediate irrigation and high irrigation. Irrigation conditions emulated the mean precipitation of the most representative environments inhabited by N. pumilio. Changes in soil water availability modified the biomass partitioning patterns of trees. In comparison to the other two conditions, high irrigation caused: (i) a higher ratio of biomass partitioning to stems than roots; (ii) more trunk growth in relation to its branches; and (iii) more photosynthetic organs relative to the aboveground biomass. Trunk size relative to that of its most recent branches was not increased by water availability. Water availability may play a significant role in the capacity of N. pumilio for space occupation due to the effects on axis differentiation.

Cactus stem (Opuntia ficus-indica Mill): anatomy, physiology and chemical composition with emphasis on its biofunctional properties.

Cactus stem (Opuntia ficus-indica Mill) is native to Mesoamerica and marketed in different forms such as fresh, frozen or pre-cooked. Worldwide, this vegetable is recognized for its pharmaceutical actions, including its antioxidant, diuretic, anticarcinogenic, anti-inflammatory, anti-diabetic, and anti-hypercholesterolemic properties, as well as their antiviral and antispermatogenic effects. However, not all of these properties have been associated with its chemical composition; therefore, this review aims to present and integrate information available on the physiology and anatomy of cactus stem and its chemical composition, focusing on some of the many factors that determine its biofunctionality. © 2017 Society of Chemical Industry.

Exploring the bark thickness-stem diameter relationship: clues from lianas, successive cambia, monocots and gymnosperms.

Bark thickness is ecologically crucial, affecting functions from fire protection to photosynthesis. Bark thickness scales predictably with stem diameter, but there is little consensus on whether this scaling is a passive consequence of growth or an important adaptive phenomenon requiring explanation. With a comparative study across 913 species, we test the expectation that, if bark thickness-stem diameter scaling is adaptive, it should be possible to find ecological situations in which scaling is predictably altered, in this case between species with different types and deployments of phloem. 'Dicots' with successive cambia and monocots, which have phloem-free bark, had predictably thinner inner (mostly living) bark than plants with single cambia. Lianas, which supply large leaf areas with limited stem area, had much thicker inner bark than self-supporting plants. Gymnosperms had thicker outer bark than angiosperms. Inner bark probably scales with plant metabolic demands, for example with leaf area. Outer bark scales with stem diameter less predictably, probably reflecting diverse adaptive factors; for example, it tends to be thicker in fire-prone species and very thin when bark photosynthesis is favored. Predictable bark thickness-stem diameter scaling across plants with different photosynthate translocation demands and modes strongly supports the idea that this relationship is functionally important and adaptively significant.

The cost of avoiding freezing in stems: trade-off between xylem resistance to cavitation and supercooling capacity in woody plants.

Stems and leaves of Olea europaea L. (olive) avoid freezing damage by substantial supercooling during the winter season. Physiological changes during acclimation to low temperatures were studied in five olive cultivars. Water relations and hydraulic traits, ice nucleation temperature (INT) and temperatures resulting in 50% damage (LT50) were determined. All cultivars showed a gradual decrease in INT and LT50 from the dry and warm summer to the wet and cold winter in Patagonia, Argentina. During acclimation to low temperatures there was an increase in leaf cell wall rigidity and stomatal conductance (gs), as well as a decrease in leaf apoplastic water content, leaf water potential (Ψ), sap flow and stem hydraulic conductivity (ks). More negative Ψ as a consequence of high gs and detrimental effects of low temperatures on root activity resulted in a substantial loss of ks due to embolism formation. Seasonal stem INT decrease from summer to winter was directly related to the xylem resistance to cavitation, determined by the loss of ks across cultivars. Thus the loss of freezable water in xylem vessels by embolisms increased stem supercooling capacity and delayed ice propagation from stems to the leaves. For the first time, a trade-off between xylem resistance to cavitation and stem and leaf supercooling capacity was observed in plants that avoid extracellular freezing by permanent supercooling. The substantial loss of hydraulic function in olive cultivar stems by embolism formation with their high repair costs are compensated by avoiding plant damage at very low subzero temperatures.