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The ability of tree stems to recover from damage is critical for tree survival and may explain species distributions across disturbance regimes. Two primary responses to stem damage, decay compartmentalization and wound closure, act in concert to limit decay and pathogen spread. A previous study demonstrated a tradeoff between wound responses that varied with anatomical traits, but its wide taxonomic range made it hard to analyze responses in an evolutionary context. Here, we tested the stem wound responses of 13 species of Quercus inhabiting three habitats across a gradient of fire intensity. We also quantified anatomical and structural traits and phylogenetic position, in order to assess the relative contributions of ecological adaptation and phylogenetic history in determining traits. Xylem anatomical traits were phylogenetically constrained, while phloem traits and damage responses varied with habitat. Across habitats, hammock and sandhill species closed bark wounds effectively, whereas scrub species limited the spread of xylem decay. There was a tradeoff between wound closure and decay compartmentalization within the white+live oaks. The fact that some wound response traits are phylogenetically constrained while others respond to ecological pressures suggests that damage responses integrate mechanisms operating at several levels within plants.

The hydraulic properties of xylem and phloem differ but the magnitude and functional consequences of the differences are not well understood. Phloem and xylem functional areas, hydraulic conduit diameters and conduit frequency along the stems of Picea abies trees were measured and expressed as allometric functions of stem diameter and distance from stem apex. Conductivities of phloem and xylem were estimated from these scaling relations. Compared with xylem, phloem conduits were smaller and occupied a slightly larger fraction of conducting tissue area. Ten times more xylem than phloem was annually produced along the stem. Scaling of the conduit diameters and cross‐sectional areas with stem diameter were very similar in phloem and xylem. Phloem and xylem conduits scaled also similarly with distance from stem apex; widening downwards from the tree top, and reaching a plateau near the base of the living crown. Phloem conductivity was estimated to scale similarly to the conductivity of the outermost xylem ring, with the ratio of phloem to xylem conductivity being c. 2%. However, xylem conductivity was estimated to increase more than phloem conductivity with increasing tree dimensions as a result of accumulation of xylem sapwood. Phloem partly compensated for its smaller conducting area and narrower conduits by having a slightly higher conduit frequency.

The rare pit hypothesis predicts that the extensive inter-vessel pitting in large early-wood vessels of ring-porous trees should render many of these vessels extremely vulnerable to cavitation by air-seeding. This prediction was tested in Quercus gambelii. Cavitation was assessed from native hydraulic conductivity at field sap tension and in dehydrated branches. Single-vessel air injections gave air-seeding pressures through vessel files; these data were used to estimate air-seeding pressures for inter-vessel walls and pits. Extensive cavitation occurred at xylem sap tensions below 1 MPa. Refilling occurred below 0.5 MPa and was inhibited by phloem girdling. Remaining vessels cavitated over a wide range to above 4 MPa. Similarly, 40% of injected vessel files air-seeded below 1.0 MPa, whereas the remainder seeded over a wide range exceeding 5 MPa. Inter-vessel walls averaged 1.02 MPa air-seeding pressure, similar and opposite to the mean cavitation tension of 1.22 MPa. Consistent with the rare pit hypothesis, only 7% of inter-vessel pits were estimated to air-seed by 1.22 MPa. The results confirm the rare pit prediction that a significant fraction of large vessels in Q. gambelii experience high probability of failure by air-seeding.

The vascular network of leaves, comprising xylem and phloem, is a highly optimized system for the delivery of water, nutrients, and sugars. The design rules for these naturally occurring networks have been studied since the time of Leonardo da Vinci, who constructed a local rule for comparing the widths of in- and outgoing veins at branch points. Recently, physical models have been developed that seek to explain the full morphogenesis of leaf venial networks in which veins grow in response to local hydrodynamic feedback. Although these models go beyond simple local rules, they are challenging to compare to experimental data. Here, we extend these hydrodynamic models to a state where the direct comparison with images of full leaves becomes possible on the level of individual veins. We present a dataset of the venial networks of leaves that maintain full network topology and use this to discuss the benefits and drawbacks of such a direct comparison. We apply our approach to the direct estimation of a sink fluctuation parameter, demonstrating consistency within distinct leaf species. Finally, we utilize the ability of the model to run on full leaves to define and calculate exponents for a Murray’s law that applies to reticulate venation networks.

In the sieve elements (SEs) of the phloem, carbohydrates are transported throughout the whole plant from their site of production to sites of consumption or storage. SE structure, especially of the pore-rich end walls, has a direct effect on translocation efficiency. Differences in pore size and other features were interpreted as an evolutionary trend towards reduced hydraulic resistance. However, this has never been confirmed. Anatomical data of 447 species of woody angiosperms and gymnosperms were used for a phylogenetic analysis of end wall types, calculation of hydraulic resistance and correlation analysis with morphological and physiological variables. end wall types were defined according to pore arrangement: either grouped into a single area (simple) or into multiple areas along the end wall (compound). Convergent evolution of end wall types was demonstrated in woody angiosperms. In addition, an optimization of end wall resistance with plant height was discovered, but found to be independent of end wall type. While physiological factors also showed no correlation with end wall types, the number of sieve areas per end wall was found to scale with SE length. The results exclude the minimization of hydraulic resistance as evolutionary driver of different end wall types, contradicting this long-standing assumption. Instead, end wall type might depend on SE length.

Transport of carbohydrates and water are essential aspects of plant function. The aim of this study was to develop and test the methods for mechanistic modelling of quasi-stationary coupled phloem/xylem transport in the context of functional-structural plant modelling. The novelty of this approach is in combining analytical and computational methods. The plant structure is modelled at a metamer level with the internodes represented by conduit elements and the lateral organs represented by sources and sinks. Transport equations are solved analytically for each internode and then the solutions are adjusted and 'sewn' together using an iterative computational procedure taking into account concentration-dependent sinks and sources. The model is implemented in L-studio and uses the aspect-oriented modelling approach for phloem/xylem coupling. To our knowledge, this is the first transport model that provides continuous distributions of the system variables in a complex developing structure. The model takes into account non-linear dependence of phloem resistance and osmotic potential on the local carbohydrate concentration. The model solutions show excellent agreement with the existing results of other analytical and numerical models. These comparisons confirm the validity of the approximations made in the model. Combining analytical and computational methods made it possible to take into account continuous sink/source distribution within internodes without much increase in the complexity of the computational procedure, because the necessary changes in the model were mostly in the analytical part. The results emphasize sensitivity of phloem flux and lateral xylem flux to the presence of distributed sinks and sources along the transport system. The presented approach provides a new insight into mechanistic modelling of phloem/xylem transport in growing plants. It will be useful for both fine-scale modelling of carbohydrate dynamics and for creating simpler models at a growth unit level.

Maintaining phloem transport under water stress is expected to be crucial to whole-plant drought tolerance, but the traits that benefit phloem function under drought are poorly understood. Nearly half of surveyed angiosperm species, including important crops, use sucrose transporter proteins to actively load sugar into the phloem. Plants can alter transporter abundance in response to stress, providing a potential mechanism for active-loading species to closely regulate phloem loading rates to avoid drought-induced reductions or failures in phloem transport. We developed an integrated xylem-phloem-stomatal model to test this hypothesis by quantifying the joint impacts of transporter kinetics, phloem anatomy, and plant water status on sucrose export to sinks. We parameterized the model with phloem hydraulic resistances and sucrose transporter kinetic parameters compiled from the literature, and simulated loading regulation by allowing loading rates to decline exponentially with phloem pressure to prevent excessive sucrose concentrations from inducing viscosity limitations. In the absence of loading regulation, where loading rates were independent of phloem pressure, most resistance values produced unrealistic phloem pressures owing to viscosity effects, even under well-watered conditions. Conversely, pressure-regulated loading helped to control viscosity buildup and improved export to sinks for both lower and higher resistant phloem pathways, while maintaining realistic phloem pressures. Regulation also allowed for rapid loading and export in wet conditions while maintaining export and viable phloem pressures during drought. Therefore, we expect feedbacks between phloem pressure and loading to be critical to carbon transport in active-loading species, especially under drought, and for transporter kinetics to be strongly coordinated with phloem architecture and plant water status. This work provides an important and underexplored physiological framework to understand the ecophysiology of phloem transport under drought and to enhance the genetic engineering of crop plants.

We report that wall ingrowth deposition in phloem parenchyma (PP) transfer cells (TCs) in leaf veins of Arabidopsis (Arabidopsis thaliana) represents a novel trait of heteroblasty. Development of PP TCs involves extensive deposition of wall ingrowths adjacent to cells of the sieve element/companion cell complex. These PP TCs potentially facilitate phloem loading by enhancing efflux of symplasmic Suc for subsequent active uptake into cells of the sieve element/companion cell complex. PP TCs with extensive wall ingrowths are ubiquitous in mature cotyledons and juvenile leaves, but dramatically less so in mature adult leaves, an observation consistent with PP TC development reflecting vegetative phase change (VPC) in Arabidopsis. Consistent with this conclusion, the abundance of PP TCs with extensive wall ingrowths varied across rosette development in three ecotypes displaying differing durations of juvenile phase, and extensive deposition of wall ingrowths was observed in rejuvenated leaves following prolonged defoliation. PP TC development across juvenile, transition, and adult leaves correlated positively with levels of miR156, a major regulator of VPC in plants, and corresponding changes in wall ingrowth deposition were observed when miR156 was overexpressed or its activity suppressed by target mimicry. Analysis of plants carrying miR156-resistant forms of SQUAMOSA PROMOTER BINDING PROTEIN LIKE (SPL) genes showed that wall ingrowth deposition was increased in SPL9-group but not SPL3-group genes, indicating that SPL9-group genes may function as negative regulators of wall ingrowth deposition in PP TCs. Collectively, our results point to wall ingrowth deposition in PP TCs being under control of the genetic program regulating VPC.

Key messageOverexpression of CiNPR4 enhanced resistance of transgenic citrus plants to Huanglongbing by perceiving the salicylic acid and jasmonic acid signals and up-regulating the transcriptional activities of plant–pathogen interaction genes.Developing transgenic citrus plants with enhanced immunity is an efficient strategy to control citrus Huanglongbing (HLB). Here, a nonexpressor of pathogenesis-related gene 1 (NPR1) like gene from HLB-tolerant ‘Jackson’ grapefruit (Citrus paradisi Macf.), CiNPR4, was introduced into ‘Wanjincheng’ orange (Citrus sinensis Obseck). CiNPR4 expression was determined in transgenic citrus plants using quantitative real-time PCR analyses. The Candidatus Liberibacter asiaticus (CLas) pathogen of HLB was successfully transmitted to transgenic citrus plants by grafting infected buds. HLB symptoms developed in transgenic and wild-type (WT) plants by 9 months after inoculation. A CLas population analysis showed that 26.9% of transgenic lines exhibited significantly lower CLas titer levels compared with the CLas-infected WT plants at 21 months after inoculation. Lower starch contents and anatomical aberration levels in the phloem were observed in transgenic lines having enhanced resistance compared with CLas-infected WT plants. CiNPR4 overexpression changed the jasmonic acid, but not salicylic acid, level. Additionally, the jasmonic acid and salicylic acid levels increased after CLas infection. Transcriptome analyses revealed that the enhanced resistance of transgenic plants to HLB resulted from the up-regulated transcriptional activities of plant–pathogen interaction-related genes.

While tracheid size of conifers is often a good proxy of water transport efficiency, correlations between conifer wood structure and transport safety remain poorly understood. It is hypothesized that at least some of the variation in bordered pit and tracheid structure is associated with both transport efficiency and embolism resistance. Stem and root samples from three boreal Pinaceae species were collected to test this hypothesis. Tracheid and pit anatomy were studied using light microscopy as well as scanning and transmission electron microscopy. While tracheid size explained at least 90% of the variation in specific conductivity for stem and root samples, the strongest correlations with embolism resistance occurred at the pit level. Both torus thickness and depth of the pit chamber showed a linear increase with greater vulnerability to cavitation. Greater embolism resistance was correlated with increasing wood density and tracheid wall reinforcement. A thinner torus may be more flexible and better able to seal the pit aperture. The pit chamber depth is proportional to the distance that the margo needs to deflect for pit aspiration.