Comparative skeletal anatomy of salt marsh and western harvest mice in relation to locomotor ecology.

The salt marsh harvest mouse (Reithrodontomys raviventris) is an endangered species, endemic to the San Francisco Bay Estuary, that co-occurs with the more broadly distributed species, the western harvest mouse (Reithrodontomys megalotis). Despite their considerable external morphological similarities, the northern subspecies of salt marsh harvest mice have relatively longer and thicker tails than do western harvest mice, which may be related to their abilities to climb emergent marsh vegetation to avoid tidal inundation. We used micro-CT to compare post-cranial skeletal anatomy between the salt marsh and western harvest mouse, to examine whether the salt marsh harvest mouse's restriction to brackish marshes is associated with skeletal adaptations for scansorial locomotion. We found that salt marsh harvest mice exhibited a deeper 3rd caudal vertebra, a more caudally located longest tail vertebra, craniocaudally longer tail vertebrae, and a longer digit III proximal phalanx than western harvest mice. These phalangeal and vertebral characteristics are known to decrease body rotations during climbing, increase contact with substrates, and decrease fall susceptibility in arboreal mammals, suggesting that the salt marsh harvest mouse may be morphologically specialized for scansorial locomotion, adaptive for its dynamic wetland environment.

Trade-offs among transport, support, and storage in xylem from shrubs in a semiarid chaparral environment tested with structural equation modeling.

The xylem in plants is specialized to transport water, mechanically support the plant body, and store water and carbohydrates. Balancing these functions leads to trade-offs that are linked to xylem structure. We proposed a multivariate hypothesis regarding the main xylem functions and tested it using structural equation modeling. We sampled 29 native shrub species from field sites in semiarid Southern California. We quantified xylem water transport (embolism resistance and transport efficiency), mechanical strength, storage of water (capacitance) and starch, minimum hydrostatic pressures (Pmin), and proportions of fibers, vessels, and parenchyma, which were treated as a latent variable representing "cellular trade-offs." We found that xylem functions (transport, mechanical support, water storage, and starch storage) were independent, a result driven by PminPmin was strongly and directly or indirectly associated with all xylem functions as a hub trait. More negative Pmin was associated with increased embolism resistance and tissue strength and reduced capacitance and starch storage. We found strong support for a trade-off between embolism resistance and transport efficiency. Tissue strength was not directly associated with embolism resistance or transport efficiency, and any associations were indirect involving Pmin With Pmin removed from the model, cellular trade-offs were central and related to all other traits. We conclude that xylem traits are broadly governed by functional trade-offs and that the Pmin experienced by plants in the field exerts a strong influence over these relationships. Angiosperm xylem contains different cell types that contribute to different functions and that underpin trade-offs.

Xylem structure and hydraulic function in roots and stems of chaparral shrub species from high and low elevation in the Sierra Nevada, California.

Xylem structure and hydraulics were compared between individuals at lower and upper elevation distribution limits for five chaparral shrub species along a steep transect in the southern Sierra Nevada, California, USA. Higher-elevation plants experienced frequent winter freeze-thaw events and increased precipitation. We hypothesized that environmental differences would lead to xylem trait differences between high and low elevations, but predictions were complicated because both water stress (low elevation) and freeze-thaw events (high elevation) may select for similar traits, such as narrow vessel diameter. We found significant changes in the ratio of stem xylem area to leaf area (Huber value) between elevations, with more xylem area required to support leaves at low elevations. Co-occurring species significantly differed in their xylem traits, suggesting diverse strategies to cope with the highly seasonal environment of this Mediterranean-type climate region. Roots were more hydraulically efficient and more vulnerable to embolism relative to stems, potentially due to roots being buffered from freeze-thaw stress, which allows them to maintain wider diameter vessels. Knowledge of the structure and function of both roots and stems is likely important in understanding whole-plant response to environmental gradients.

Seasonal patterns of increases in stem girth, vessel development, and hydraulic function in deciduous tree species.

BACKGROUND AND AIMS: The onset of spring growth and vessel formation were examined within three deciduous woody plant species, Acer rubrum, Populus balsamifera ssp. trichocarpa and Quercus rubra. We were broadly interested in the lag between the onset of girth expansion and the formation of mature and hydraulically conductive vessels within the new xylem. METHODS: Dendrometers were installed on 20 trees (6-7 per species), and expansion of both bole and distal stems was monitored throughout the growing season in a common garden. For each species, four to six distal stems were harvested every other week for anatomical examination of vessel formation. Additionally, for Populus and Quercus, hydraulic conductivity measurements and active xylem staining were completed on all stem samples. KEY RESULTS: For all three species, the timing of girth expansion was similar. Expansion of distal branches occurred 12-37 d earlier than that of the bole. Vessel formation initiated several weeks prior to leaf-out, but no new earlywood vessels were mature at the time of bud break for Acer and Populus and only a few were present in Quercus. Initial stem girth expansion occurred 2 to >6 weeks before the maturation of the first current-year vessels, and there was an additional delay of up to 4 weeks before mature vessels became hydraulically functional. Hydraulic conductivity was strongly correlated with the number and diameter of stained vessels. CONCLUSIONS: Bud break and leaf expansion relied predominantly on water supplied by vessels formed during prior seasons. Early-season activity is likely affected by the function of older xylem vessels and the environmental factors that influence their structure and function. Understanding the functional lifespan of vessels and the varying contributions of new and older vessels to conductivity are critical to understanding of the phenology and vascular function of long-lived woody plants in response to changing climates.

Vegetation-type conversion of evergreen chaparral shrublands to savannahs dominated by exotic annual herbs: causes and consequences for ecosystem function.

Woody, evergreen shrublands are the archetypal community in mediterranean-type ecosystems, and these communities are profoundly changed when they undergo vegetation-type conversion (VTC) to become annual, herb-dominated communities. Recently, VTC has occurred throughout southern California chaparral shrublands, likely with changes in important ecosystem functions. The mechanisms that lead to VTC and subsequent changes to ecosystem processes are important to understand as they have regional and global implications for ecosystem services, climate change, land management, and policy. The main drivers of VTC are altered fire regimes, aridity, and anthropogenic disturbance. Some changes to ecosystem function are certain to occur with VTC, but their magnitudes are unclear, whereas other changes are unpredictable. I present two hypotheses: (1) VTC leads to warming that creates a positive feedback promoting additional VTC, and (2) altered nitrogen dynamics create negative feedbacks and promote an alternative stable state in which communities are dominated by herbs. The patterns described for California are mostly relevant to the other mediterranean-type shrublands of the globe, which are biodiversity hotspots and threatened by VTC. This review examines the extent and causes of VTC, ecosystem effects, and future research priorities.

Xylem biomechanics, water storage, and density within roots and shoots of an angiosperm tree species.

Xylem is a complex tissue that forms the bulk of tree bodies and has several functions, including to conduct water, store water and nutrients, and biomechanically support the plant body. We examined how xylem functional traits varied at different positions within 9-year-old Populus balsamifera subsp. trichocarpa. Whole trees were excavated, and xylem samples were collected at 1-m increments along the main root-to-shoot axis of six trees, from root tip to shoot tip. We examined biomechanical and water-storage traits of the xylem, including using a non-invasive imaging technique to examine water content within long, intact branches (high-resolution computed tomography; microCT). Xylem density, strength, and stiffness were greater in shoots than roots. Along the main root-to-shoot axis, xylem strength and stiffness were greatest at shoot tips, and the tissue became linearly weaker and less stiff down the plant and through the root. Roots had greater water storage with lower biomechanical support, and shoots had biomechanically stronger and stiffer xylem with lower water storage. These findings support trade-offs among xylem functions between roots and shoots. Understanding how xylem functions differ throughout tree bodies is important in understanding whole-tree functioning and how terrestrial plants endure numerous environmental challenges over decades of growth.

Starch storage capacity of sapwood is related to dehydration avoidance during drought.

PREMISE: The xylem tissue of plants performs three principal functions: transport of water, support of the plant body, and nutrient storage. Tradeoffs may arise because different structural requirements are associated with different functions or because suites of traits are under selection that relate to resource acquisition, use, and turnover. The structural and functional basis of xylem storage is not well established. We hypothesized that greater starch storage would be associated with greater sapwood parenchyma and reduced fibers, which would compromise resistance to xylem tensions during dehydration. METHODS: We measured cavitation resistance, minimum water potential, starch content, and sapwood parenchyma and fiber area in 30 species of southern California chaparral shrubs (evergreen and deciduous). RESULTS: We found that species storing greater starch within their xylem tended to avoid dehydration and were less cavitation resistant, and this was supported by phylogenetic independent contrasts. Greater sapwood starch was associated with greater parenchyma area and reduced fiber area. For species without living fibers, the associations with parenchyma were stronger, suggesting that living fibers may expand starch storage capacity while also contributing to the support function of the vascular tissue. Drought-deciduous species were associated with greater dehydration avoidance than evergreens. CONCLUSIONS: Evolutionary forces have led to an association between starch storage and dehydration resistance as part of an adaptive suite of traits. We found evidence for a tradeoff between tissue mechanical traits and starch storage; moreover, the evolution of novel strategies, such as starch-storing living fibers, may mitigate the strength of this tradeoff.

Factors controlling drought resistance in grapevine (Vitis vinifera, chardonnay): application of a new microCT method to assess functional embolism resistance.

PREMISE: Quantifying resistance to embolism in woody plants is important for understanding their drought response. Methods to accurately quantify resistance to embolism continue to be debated. METHODS: We used a new microCT-based approach that quantifies embolized conduits and also analyzes conductive conduits by using an x-ray-dense, iodine-rich tracer that moves though the vascular system and can easily be observed in microCT images. Many previous microCT studies assumed that all conduits were initially conductive, which may not be the case if there are developing or occluded conduits. We compared microCT results to a standard benchtop dehydration method and a centrifuge method. During dehydration, we measured gas exchange and quantified water potential at mortality. RESULTS: Our microCT curves agreed with previously published microCT curves from the same greenhouse-grown cultivar. We found a significant difference in embolism estimates if we assumed that all water-filled conduits were functional rather than only those containing tracer. Embolism estimates from microCT differed from both the benchtop and centrifuge methods. The benchtop and centrifuge methods did not differ from one another. CONCLUSIONS: The new microCT method presented here is valuable in sampling species that may contain nonconductive conduits. Disagreement between microCT and two other methods was likely due to differences in the ways they quantify embolism. MicroCT assess the theoretical effect of embolism, whereas benchtop and centrifuge methods directly measure hydraulic conductivity. The theoretical approach does not fully account for the resistances of flow through a complex 3D vascular network.

A Lysine-Targeted Affinity Label for Serine-ß-Lactamase Also Covalently Modifies New Delhi Metallo-ß-lactamase-1 (NDM-1).

The divergent sequences, protein structures, and catalytic mechanisms of serine- and metallo-ß-lactamases hamper the development of wide-spectrum ß-lactamase inhibitors that can block both types of enzymes. The O-aryloxycarbonyl hydroxamate inactivators of Enterobacter cloacae P99 class C serine-ß-lactamase are unusual covalent inhibitors in that they target both active-site Ser and Lys residues, resulting in a cross-link consisting of only two atoms. Many clinically relevant metallo-ß-lactamases have an analogous active-site Lys residue used to bind ß-lactam substrates, suggesting a common site to target with covalent inhibitors. Here, we demonstrate that an O-aryloxycarbonyl hydroxamate inactivator of serine-ß-lactamases can also serve as a classical affinity label for New Delhi metallo-ß-lactamase-1 (NDM-1). Rapid dilution assays, site-directed mutagenesis, and global kinetic fitting are used to map covalent modification at Lys211 and determine KI (140 µM) and kinact (0.045 min-1) values. Mass spectrometry of the intact protein and the use of ultraviolet photodissociation for extensive fragmentation confirm stoichiometric covalent labeling that occurs specifically at Lys211. A 2.0 Å resolution X-ray crystal structure of inactivated NDM-1 reveals that the covalent adduct is bound at the substrate-binding site but is not directly coordinated to the active-site zinc cluster. These results indicate that Lys-targeted affinity labels might be a successful strategy for developing compounds that can inactivate both serine- and metallo-ß-lactamases.

High-resolution computed tomography reveals dynamics of desiccation and rehydration in fern petioles of a desiccation-tolerant fern.

Desiccation-tolerant (DT) plants can dry past -100 MPa and subsequently recover function upon rehydration. Vascular DT plants face the unique challenges of desiccating and rehydrating complex tissues without causing structural damage. However, these dynamics have not been studied in intact DT plants. We used high resolution micro-computed tomography (microCT), light microscopy, and fluorescence microscopy to characterize the dynamics of tissue desiccation and rehydration in petioles (stipes) of intact DT ferns. During desiccation, xylem conduits in stipes embolized before cellular dehydration of living tissues within the vascular cylinder. During resurrection, the chlorenchyma and phloem within the stipe vascular cylinder rehydrated before xylem refilling. We identified unique stipe traits that may facilitate desiccation and resurrection of the vascular system, including xylem conduits containing pectin (which may confer flexibility and wettability); chloroplasts within the vascular cylinder; and an endodermal layer impregnated with hydrophobic substances that impede apoplastic leakage while facilitating the upward flow of water within the vascular cylinder. Resurrection ferns are a novel system for studying extreme dehydration recovery and embolism repair in the petioles of intact plants. The unique anatomical traits identified here may contribute to the spatial and temporal dynamics of water movement observed during desiccation and resurrection.

Wood structure and function change with maturity: Age of the vascular cambium is associated with xylem changes in current-year growth.

Xylem vessel structure changes as trees grow and mature. Age- and development-related changes in xylem structure are likely related to changes in hydraulic function. We examined whether hydraulic function, including hydraulic conductivity and vulnerability to water-stress-induced xylem embolism, changed over the course of cambial development in the stems of 17 tree species. We compared current-year growth of young (1-4 years), intermediate (2-7 years), and older (3-10 years) stems occurring in series along branches. Diffuse and ring porous species were examined, but nearly all species produced only diffuse porous xylem in the distal branches that were examined irrespective of their mature xylem porosity type. Vessel diameter and length increased with cambial age. Xylem became both more conductive and more cavitation resistant with cambial age. Ring porous species had longer and wider vessels and xylem that had higher conductivity and was more vulnerable to cavitation; however, these differences between porosity types were not present in young stem samples. Understanding plant hydraulic function and architecture requires the sampling of multiple-aged tissues because plants may vary considerably in their xylem structural and functional traits throughout the plant body, even over relatively short distances and closely aged tissues.

Direct comparison of four methods to construct xylem vulnerability curves: Differences among techniques are linked to vessel network characteristics.

During periods of dehydration, water transport through xylem conduits can become blocked by embolism formation. Xylem embolism compromises water supply to leaves and may lead to losses in productivity or plant death. Vulnerability curves (VCs) characterize plant losses in conductivity as xylem pressures decrease. VCs are widely used to characterize and predict plant water use at different levels of water availability. Several methodologies for constructing VCs exist and sometimes produce different results for the same plant material. We directly compared four VC construction methods on stems of black cottonwood (Populus trichocarpa), a model tree species: dehydration, centrifuge, X-ray-computed microtomography (microCT), and optical. MicroCT VC was the most resistant, dehydration and centrifuge VCs were intermediate, and optical VC was the most vulnerable. Differences among VCs were not associated with how cavitation was induced but were related to how losses in conductivity were evaluated: measured hydraulically (dehydration and centrifuge) versus evaluated from visual information (microCT and optical). Understanding how and why methods differ in estimating vulnerability to xylem embolism is important for advancing knowledge in plant ecophysiology, interpreting literature data, and using accurate VCs in water flux models for predicting plant responses to drought.

Specificity of extended O-aryloxycarbonyl hydroxamates as inhibitors of a class C ß-lactamase.

Class C ß-lactamases have previously been shown to be efficiently inactivated by O-aryloxycarbonyl hydroxamates. O-Phenoxycarbonyl-N-benzyloxycarbonylhydroxylamine (1) and O-phenoxycarbonyl-N-(R)-[(4-amino-4-carboxy-1-butyl)oxycarbonyl]hydroxylamine (2), for example, were found to be effective inactivators. The present paper describes a structure-activity study of these molecules to better define the important structural elements for high inhibitory activity. The results show that a well-positioned hydrophobic element (which may interact with the Tyr221 residue of the enzyme) and a negatively charged element, e.g. a carboxylate group (which may interact with Arg204), are required for high reactivity with the enzyme. The new compounds were found to inactivate by forming a carbonyl cross-linked enzyme (probably Ser64OCONHLys 315) as for 1 rather than the inert hydroxamoyl derivative observed with 2.

Kinetic Evidence for a Second Ligand Binding Site on Streptococcus pneumoniae Penicillin-Binding Protein 2x.

High molecular mass penicillin-binding proteins (PBPs, DD-peptidases) of class B, such as Streptococcus pneumoniae PBP2x, catalyze the cross-linking of peptidoglycan in bacterial cell wall biosynthesis and are thus important antibiotic targets. Despite their importance in this regard, structure-function studies of ligands of these enzymes have been impeded by the absence of useful substrates. In vitro, these enzymes do not catalyze peptide hydrolysis or aminolysis, their in vivo reaction, but some, such as PBP2x, do catalyze these reactions of certain thioesters such as PhCH2CONHCH2COSCH(D-Me)CO2- (2). We have now prepared several peptidoglycan-mimetic thioesters that we expected to more closely resemble the natural substrates of these enzymes. To our surprise, however, these compounds, although indeed substrates of PBP2x, did not, unlike 2, appear to form an acyl-enzyme intermediate during hydrolysis, and their turnover was inhibited by certain peptides and N-acylamino acids much more weakly than that of 2. An inhibitor of this type, N-benzyloxycarbonyl-d-glutamic acid, also quenched the fluorescence of PBP2x that had been labeled at the DD-peptidase active site by 6-dansylamidopenicillanic acid. These results were interpreted in terms of a model where the peptidoglycan-mimetic thioesters preferentially bound to and hydrolyzed at a site other than the classical DD-peptidase active site. This second site is likely to represent part of an extended binding site that accommodates a peptidoglycan substrate or regulator in vivo. Such a site may be a target for future inhibitor/antibiotic design.

Extensive drought-associated plant mortality as an agent of type-conversion in chaparral shrublands.

Contents Summary 498 I. Introduction 498 II. Ecological drought and vegetation type-conversion 499 III. Chaparral mortality during extreme drought events 501 IV. Some species survive drought and others do not 501 V. Recovery potential 502 VI. Conclusions 503 Acknowledgements 503 References 503 SUMMARY: California experienced an intense drought from 2012 to 2015, with southern California remaining in drought to the present. Widespread chaparral shrub mortality was observed during the peak of the drought in 2014. Some species were more impacted than others and shallow-rooted shrub species were the most vulnerable to drought-associated mortality. This type of drought represents what is termed an 'ecological drought' during which an ecosystem is driven beyond thresholds of vulnerability, triggering impairment of ecosystem services and feedbacks that may result in long-term type-conversion of natural communities. The ability of shrublands to recover will depend on the timing, intensity and seasonality of future extreme climate events, post-fire recruitment potential of species with obligate fire-associated recruitment, and interactions with other stresses.

Functional lifespans of xylem vessels: Development, hydraulic function, and post-function of vessels in several species of woody plants.

PREMISE OF THE STUDY: Xylem vessels transition through different stages during their functional lifespan, including expansion and development of vessel elements, transition to vessel hydraulic functionality, and eventual transition to post-functionality. We used information on vessel development and function to develop a model of vessel lifespan for woody plants. METHODS: We examined vessel functional lifespan using repeated anatomical sampling throughout the growing season, combined with active-xylem staining to evaluate vessel hydraulic transport functionality. These data were combined with a literature review. The transitions between vessel functional lifespans for several species are illustrated, including grapevine (Vitis vinifera L., Vitaceae), English oak (Quercus robur L., Fagaceae), American chestnut [Castanea dentata (Marshall) Borkh.; Fagaceae], and several arid and semi-arid shrub species. KEY RESULTS: In intact woody plants, development and maturation of vessel elements may be gradual. Once hydraulically functional, vessel elements connect to form a vessel network that is responsible for bulk hydraulic flow through the xylem. Vessels become nonfunctional due to the formation of gas emboli. In some species and under some conditions, vessel functionality of embolized conduits may be restored through refilling. Blockages, such as tyloses, gels, or gums, indicate permanent losses in hydraulic functional capacity; however, there may be some interesting exceptions to permanent loss of functionality for gel-based blockages. CONCLUSIONS: The gradual development and maturation of vessel elements in woody plants, variation in the onset of functionality between different populations of vessels throughout the growing season, and differences in the timing of vessel transitions to post-functionality are important aspects of plant hydraulic function.

Early warning biomarkers in major depressive disorder: a strategic approach to a testing question.

PURPOSE: Identification of biomarkers in major depressive disorder (MDD) has proceeded in an extemporised manner. No single biomarker has been identified with utility in screening, diagnosis, prognosis, or monitoring, and screening tests have different characteristics than the other functions. Using chaos, bifurcation, and perturbation (CBP) theories, the aim is to identify biomarkers to aid clinicians in screening for MDD. MATERIALS AND METHODS: MDD is a complex disorder; consequently, a reductionist approach to characterize the complex system changes found in MDD will be inchoate and unreliable. A holistic approach is used to identify biomarkers reflecting the tipping points seen before the catastrophic bifurcation that results in MDD. RESULTS: Applying CBP theories revealed skew, resistance to change, flickering, increased variance and autocorrelation as patterns of biomarkers. Integrals and differentials of extracellular and intracellular biomarkers were identified, specifically focussed on hypothalamo-pituitary axis (HPA) dysfunction, metabolic dysfunction, inflammation and mitochondrial oxidative stress, and tryptophan metabolism. CONCLUSIONS: Applying CBP theories to the dysfunctional complex biological systems in MDD led to development of integrals and differentials of biomarkers that can be used in screening for MDD and planning future biomarker research, targeting intracellular and extracellular inflammation, HPA axis dysfunction, and tryptophan metabolism.

Global convergence in the vulnerability of forests to drought.

Shifts in rainfall patterns and increasing temperatures associated with climate change are likely to cause widespread forest decline in regions where droughts are predicted to increase in duration and severity. One primary cause of productivity loss and plant mortality during drought is hydraulic failure. Drought stress creates trapped gas emboli in the water transport system, which reduces the ability of plants to supply water to leaves for photosynthetic gas exchange and can ultimately result in desiccation and mortality. At present we lack a clear picture of how thresholds to hydraulic failure vary across a broad range of species and environments, despite many individual experiments. Here we draw together published and unpublished data on the vulnerability of the transport system to drought-induced embolism for a large number of woody species, with a view to examining the likely consequences of climate change for forest biomes. We show that 70% of 226 forest species from 81 sites worldwide operate with narrow (<1 megapascal) hydraulic safety margins against injurious levels of drought stress and therefore potentially face long-term reductions in productivity and survival if temperature and aridity increase as predicted for many regions across the globe. Safety margins are largely independent of mean annual precipitation, showing that there is global convergence in the vulnerability of forests to drought, with all forest biomes equally vulnerable to hydraulic failure regardless of their current rainfall environment. These findings provide insight into why drought-induced forest decline is occurring not only in arid regions but also in wet forests not normally considered at drought risk.

Clinical Development Strategies and Considerations for Zika Vaccine Licensure.

The Zika outbreak that began in 2015 has spread from Brazil to countries across the Western Hemisphere including the United States, presenting global public health challenges that call for the expedited development and availability of preventive vaccines to protect against Zika virus disease. While the general principles guiding the nonclinical and clinical development for Zika vaccines are the same as those of other preventive vaccines, unique considerations apply, in particular if development occurs during a public health emergency. Furthermore, incomplete information about the pathogenesis of Zika virus disease and the mechanism by which candidate preventive vaccines potentially may confer protection presents additional challenges to their clinical development. Nevertheless, definition of clinical development strategies to enable sound regulatory assessment, with a goal toward licensure is critical for these products. This article will provide an overview of the regulatory considerations for the clinical development and licensure of Zika vaccine candidates including a discussion of clinical study designs, approaches to demonstrate vaccine effectiveness, and regulatory pathways to licensure.

Conflicting demands on angiosperm xylem: Tradeoffs among storage, transport and biomechanics.

The secondary xylem of woody plants transports water mechanically supports the plant body and stores resources. These three functions are interdependent giving rise to tradeoffs in function. Understanding the relationships among these functions and their structural basis forms the context in which to interpret xylem evolution. The tradeoff between xylem transport efficiency and safety from cavitation has been carefully examined with less focus on other functions, particularly storage. Here, we synthesize data on all three xylem functions in angiosperm branch xylem in the context of tradeoffs. Species that have low safety and efficiency, examined from a resource economics perspective, are predicted to be adapted for slow resource acquisition and turnover as characterizes some environments. Tradeoffs with water storage primarily arise because of differences in fibre traits, while tradeoffs in carbohydrate storage are driven by parenchyma content of tissue. We find support for a tradeoff between safety from cavitation and storage of both water and starch in branch xylem tissue and between water storage capacity and mechanical strength. Living fibres may facilitate carbohydrate storage without compromising mechanical strength. The division of labour between different xylem cell types allows for considerable functional and structural diversity at multiple scales.