Early spring flowers rely on xylem hydration but are not limited by stem xylem conductivity.

Many woody plants produce large floral displays early in the spring when xylem transport can be variable and often reduced. To determine whether stem hydraulics impact floral water use, we quantified floral transpiration and tested whether it was correlated with stem xylem conductivity in five temperate woody species that flower before producing leaves. We measured inflorescence gas exchange, examined the relationship between diffusive conductance and inflorescence morphology, and estimated the amount of water supplied to an inflorescence by the phloem. We also tested for correlation between transpiration and native stem xylem conductivity for branches with leaves and branches with flowers. The flowers of our study species obtain most of their water from the xylem. Diffusive conductance was higher in small inflorescences, but water content and daily transpiration rates were greater for larger inflorescences. We found no correlation between floral transpiration per branch and stem xylem conductivity within species. The data suggest that inflorescence water loss during anthesis is not limited by the xylem in our study species. We highlight the impact of floral morphology on hydraulic traits and encourage exploration into temporal shifts in floral hydration.

Leaf out time correlates with wood anatomy across large geographic scales and within local communities.

There is a long-standing idea that the timing of leaf production in seasonally cold climates is linked to xylem anatomy, specifically vessel diameter because of the hydraulic requirements of expanding leaves. We tested for a relationship between the timing of leaf out and vessel diameter in 220 plants in three common gardens accounting for species' phylogenetic relationships. We investigated how vessel diameter related to wood porosity, plant height and leaf length. We also used dye perfusion tests to determine whether plants relied on xylem produced during the previous growing season at the time of leaf out. In all three gardens, there was later leaf out in species with wider vessels. Ring-porous species had the widest vessels, exhibited latest leaf out and relied less on xylem made during the previous growing season than diffuse-porous species. Wood anatomy and leaf phenology did not exhibit a phylogenetic signal. The timing of leaf out is correlated with wood anatomy across species regardless of species' geographic origin and phylogenetic relationships. This correlation could be a result of developmental and physiological links between leaves and wood or tied to a larger safety efficiency trade-off.

Coordination of spring vascular and organ phenology in deciduous angiosperms growing in seasonally cold climates.

In seasonally cold climates, many woody plants tolerate chilling and freezing temperatures by ceasing growth, shedding leaves and entering dormancy. At the same time, transport within these plants often decreases as the vascular system exhibits reduced functionality. As spring growth requires water and nutrients, we ask the question: how much does bud, leaf and flower development depend on the vasculature in spring? In this review, we present what is known about leaf, flower and vascular phenology to sort out this question. In early stages of bud development, buds rely on internal resources and do not appear to require vascular support. The situation changes during organ expansion, after leaves and flowers reconnect to the stem vascular system. However, there are major gaps in our understanding of the timing of vascular development, especially regarding the phloem, as well as the synchronization among leaves, flowers, stem and root vasculature. We believe these gaps are mainly the outcome of research completed in silo and urge future work to take a more integrative approach. We highlight current challenges and propose future directions to make rapid progress on this important topic in upcoming years.

Ontogenetic scaling of phloem sieve tube anatomy and hydraulic resistance with tree height in Quercus rubra.

PREMISE: The dimensions of phloem sieve elements have been shown to vary as a function of tree height, decreasing hydraulic resistance as the transport pathway lengthens. However, little is known about ontogenetic patterns of sieve element scaling. Here we examine within a single species (Quercus rubra) how decreases in hydraulic resistance with distance from the plant apex are mediated by overall plant size. METHODS: We sampled and imaged phloem tissue at multiple heights along the main stem and in the live crown of four size classes of trees using fluorescence and scanning electron microscopy. Sieve element length and radius, the number of sieve areas per compound plate, pore number, and pore radius were used to calculate total hydraulic resistance at each sampling location. RESULTS: Sieve element length varied with tree size, while sieve element radius, sieve pore radius, and the number of sieve areas per compound plate varied with sampling position. When data from all size classes were aggregated, all four variables followed a power-law trend with distance from the top of the tree. The net effect of these ontogenetic scalings was to make total hydraulic sieve tube resistance independent of tree height from 0.5 to over 20 m. CONCLUSIONS: Sieve element development responded to two pieces of information, tree size and distance from the apex, in a manner that conserved total sieve tube resistance across size classes. A further differentiated response between the phloem in the live crown and in the main stem is also suggested.

A temporal shift in resource allocation facilitates flowering before leaf out and spring vessel maturation in precocious species.

PREMISE OF THE STUDY: New growth in the spring requires resource mobilization in the vascular system at a time when xylem and phloem function are often reduced in seasonally cold climates. As a result, the timing of leaf out and/or flowering could depend on when the vascular system resumes normal function in the spring. This study investigated whether flowering time is influenced by vascular phenology in plants that flower precociously before they have leaves. METHODS: Flower, leaf, and vascular phenology were monitored in pairs of precocious and non-precocious congeners. Differences in resource allocation were quantified by measuring bud dry mass and water content throughout the year, floral hydration was modelled, and a girdling treatment completed on branches in the field. KEY RESULTS: Precocious flowering species invested more in floral buds the year before flowering than did their non-precocious congeners, thus mobilizing less water in the spring, which allowed flowering before new vessel maturation. CONCLUSIONS: A shift in the timing of resource allocation in precocious flowering plants allowed them to flower before the production of mature vessels and minimized the significance of seasonal changes in vascular function to their flowering phenology. The low investment required to complete floral development in the spring when the plant vascular system is often compromised could explain why flowers can emerge before leaf out.

An experimental test of fitness variation across a hydrologic gradient predicts willow and poplar species distributions.

Environmental filtering is an important community assembly process influencing species distributions. Contrasting species abundance patterns along environmental gradients are commonly used to provide evidence for environmental filtering. However, the same abundance patterns may result from alternative or concurrent assembly processes. Experimental tests are an important means to decipher whether species fitness varies with environment, in the absence of dispersal constraints and biotic interactions, and to draw conclusions about the importance of environmental filtering in community assembly. We performed an experimental test of environmental filtering in 14 closely related willow and poplar species (family Salicaceae) by transplanting cuttings of each species into 40 common gardens established along a natural hydrologic gradient in the field, where competition was minimized and herbivory was controlled. We analyzed species fitness responses to the hydrologic environment based on cumulative growth and survival over two years using aster fitness models. We also examined variation in nine drought and flooding tolerance traits expected to contribute to performance based on a priori understanding of plant function in relation to water availability and stress. We found substantial evidence that environmental filtering along the hydrologic gradient played a critical role in determining species distributions. Fitness variation of each species in the field experiment was used to model their water table depth optima. These optima predicted 68% of the variation in species realized hydrologic niches based on peak abundance in naturally assembled communities in the surrounding region. Multiple traits associated with water transport efficiency and water stress tolerance were correlated with species hydrologic niches, but they did not necessarily covary with each other. As a consequence, species occupying similar hydrologic niches had different combinations of trait values. Moreover, individual traits were less phylogenetically conserved than species hydrologic niches and integrated water stress tolerance as determined by multiple traits. We conclude that differential fitness among species along the hydrologic gradient was the consequence of multiple traits associated with water transport and water stress tolerance, expressed in different combinations by different species. Varying environmental tolerance, in turn, played a critical role in driving niche segregation among close relatives along the hydrologic gradient.

Allocation, stress tolerance and carbon transport in plants: how does phloem physiology affect plant ecology?

Despite the crucial role of carbon transport in whole plant physiology and its impact on plant-environment interactions and ecosystem function, relatively little research has tried to examine how phloem physiology impacts plant ecology. In this review, we highlight several areas of active research where inquiry into phloem physiology has increased our understanding of whole plant function and ecological processes. We consider how xylem-phloem interactions impact plant drought tolerance and reproduction, how phloem transport influences carbon allocation in trees and carbon cycling in ecosystems and how phloem function mediates plant relations with insects, pests, microbes and symbiotes. We argue that in spite of challenges that exist in studying phloem physiology, it is critical that we consider the role of this dynamic vascular system when examining the relationship between plants and their biotic and abiotic environment.

Plant hydraulics as a central hub integrating plant and ecosystem function: meeting report for 'Emerging Frontiers in Plant Hydraulics' (Washington, DC, May 2015).

Water plays a central role in plant biology and the efficiency of water transport throughout the plant affects both photosynthetic rate and growth, an influence that scales up deterministically to the productivity of terrestrial ecosystems. Moreover, hydraulic traits mediate the ways in which plants interact with their abiotic and biotic environment. At landscape to global scale, plant hydraulic traits are important in describing the function of ecological communities and ecosystems. Plant hydraulics is increasingly recognized as a central hub within a network by which plant biology is connected to palaeobiology, agronomy, climatology, forestry, community and ecosystem ecology and earth-system science. Such grand challenges as anticipating and mitigating the impacts of climate change, and improving the security and sustainability of our food supply rely on our fundamental knowledge of how water behaves in the cells, tissues, organs, bodies and diverse communities of plants. A workshop, 'Emerging Frontiers in Plant Hydraulics' supported by the National Science Foundation, was held in Washington DC, 2015 to promote open discussion of new ideas, controversies regarding measurements and analyses, and especially, the potential for expansion of up-scaled and down-scaled inter-disciplinary research, and the strengthening of connections between plant hydraulic research, allied fields and global modelling efforts.

The making of giant pumpkins: how selective breeding changed the phloem of Cucurbita maxima from source to sink.

Despite the success of breeding programmes focused on increasing fruit size, relatively little is known about the anatomical and physiological changes required to increase reproductive allocation. To address this gap in knowledge, we compared fruit/ovary anatomy, vascular structure and phloem transport of two varieties of giant pumpkins, and their smaller fruited progenitor under controlled environmental conditions. We also modelled carbon transport into the fruit of competitively grown plants using data collected in the field. There was no evidence that changes in leaf area or photosynthetic capacity impacted fruit size. Instead, giant varieties differed in their ovary morphology and contained more phloem on a cross-sectional area basis in their petioles and pedicels than the ancestral variety. These results suggest that sink activity is important in determining fruit size and that giant pumpkins have an enhanced capacity to transport carbon. The strong connection observed between carbon fixation, phloem structure and fruit growth in field-grown plants indicates that breeding for large fruit has led to changes throughout the carbon transport system that could have important implications for how we think about phloem transport velocity and carbon allocation.

Contrasting effects of plant species traits and moisture on the decomposition of multiple litter fractions.

Environmental variation in moisture directly influences plant litter decomposition through effects on microbial activity, and indirectly via plant species traits. Whether the effects of moisture and plant species traits are mutually reinforcing or counteracting during decomposition are unknown. To disentangle the effects of moisture from the effects of species traits that vary with moisture, we decomposed leaf litter from 12 plant species in the willow family (Salicaceae) with different native habitat moisture preferences in paired mesic and wetland plots. We fit litter mass loss data to an exponential decomposition model and estimated the decay rate of the rapidly cycling litter fraction and size of the remaining fraction that decays at a rate approaching zero. Litter traits that covaried with moisture in the species' native habitat significantly influenced the decomposition rate of the rapidly cycling litter fraction, but moisture in the decomposition environment did not. In contrast, for the slowly cycling litter fraction, litter traits that did not covary with moisture in the species' native habitat and moisture in the decomposition environment were significant. Overall, the effects of moisture and plant species traits on litter decomposition were somewhat reinforcing along a hydrologic gradient that spanned mesic upland to wetland (but not permanently surface-saturated) plots. In this system, plant trait and moisture effects may lead to greater in situ decomposition rates of wetland species compared to upland species; however, plant traits that do not covary with moisture will also influence decomposition of the slowest cycling litter fraction.

Phloem transport velocity varies over time and among vascular bundles during early cucumber seedling development.

We use a novel dye-tracing technique to measure in vivo phloem transport velocity in cucumber (Cucumis sativus) plants during early seedling development. We focus on seedlings because of their importance in plant establishment and because they provide a simple source and sink model of phloem transport. The dye-tracing method uses a photodiode to track the movement of a bleach front of fluorescent dye traveling in the phloem from the cotyledons (source) to the roots (sink). During early seedling development, phloem transport velocity in this direction can change 2-fold depending on vascular connectivity and the number of actively growing sinks. Prior to leaf expansion, vascular bundles attached to the first developing leaf demonstrate a decline in basipetal phloem transport that can be alleviated by the leaf's removal. At this stage, seedlings appear carbon limited and phloem transport velocity is correlated with cotyledon area, a pattern that is apparent both during cotyledon expansion and after source area manipulation. When the first leaf transitions to a carbon source, seedling growth rate increases and basipetal phloem transport velocity becomes more stable. Because bundles appear to operate autonomously, transport velocity can differ among vascular bundles. Together, these results demonstrate the dynamic and heterogeneous nature of phloem transport and underline the need for a better understanding of how changes in phloem physiology impact growth and allocation at this critical stage of development.

Phenological cues drive an apparent trade-off between freezing tolerance and growth in the family Salicaceae.

With increasing concern about the ecological consequences of global climate change, there has been renewed interest in understanding the processes that determine species range limits. We tested a long-hypothesized trade-off between freezing tolerance and growth rate that is often used to explain species range limits. We grew 24 willow and poplar species (family Salicaceae) collected from across North America in a greenhouse common garden under two climate treatments. Maximum entropy models were used to describe species distributions and to estimate species-specific climate parameters. A range of traits related to freezing tolerance, including senescence, budburst, and susceptibility to different temperature minima during and after acclimation were measured. As predicted, species from colder climates exhibited higher freezing tolerance and slower growth rates than species from warmer climates under certain environmental conditions. However, the average relative growth rate (millimeters per meter per day) of northern species markedly increased when a subset of species was grown under a long summer day length (20.5 h), indicating that genetically based day-length cues are required for growth regulation in these species. We conclude that the observed relationship between freezing tolerance and growth rate is not driven by differences in species' intrinsic growth capacity but by differences in the environmental cues that trigger growth. We propose that the coordinated evolution of freezing tolerance and growth phenology could be important in circumscribing willow and poplar range limits and may have important implications for species' current and future distributions.

A UK consensus statement on thromboprophylaxis for autologous breast reconstruction.

Microsurgical breast reconstruction accounts for 22% of breast reconstructions in the UK. Despite thromboprophylaxis, venous thromboembolism (VTE) occurs in up to 4% of cases. Using a Delphi process, this study established a UK consensus on VTE prophylaxis strategy, for patients undergoing autologous breast reconstruction using free-tissue transfer. It captured geographically divergent views, producing a guide that reflected the peer opinion and current evidence base. METHODS: Consensus was ascertained using a structured Delphi process. A specialist from each of the UK's 12 regions was invited to the expert panel. Commitment to three to four rounds of questions was sought at enrollment. Surveys were distributed electronically. An initial qualitative free-text survey was distributed to identify likely lines of consensus and dissensus. Each panelist was provided with full-text versions of key papers on the topic. Initial free-text responses were analyzed to develop a set of structured quantitative statements, which were refined via a second survey as a consensus was approached. RESULTS: The panel comprised 18 specialists: plastic surgeons and thrombosis experts from across the UK. Each specialist completed three rounds of surveys. Together, these plastic surgeons reported having performed more than 570 microsurgical breast reconstructions in the UK in 2019. A consensus was reached on 27 statements, detailing the assessment and delivery of VTE prophylaxis. CONCLUSION: To our knowledge, this is the first study to collate current practice, expert opinion from across the UK, and a literature review. The output was a practical guide for VTE prophylaxis for microsurgical breast reconstruction in any UK microsurgical breast reconstruction unit.

Seasonal changes in temperate woody plant phloem anatomy and physiology: implications for long-distance transport.

Seasonal changes in climate are accompanied by shifts in carbon allocation and phenological changes in woody angiosperms, the timing of which can have broad implications for species distributions, interactions and ecosystem processes. During critical transitions from autumn to winter and winter to spring, physiological and anatomical changes within the phloem could impose a physical limit on the ability of woody angiosperms to transport carbon and signals. There is a paucity of the literature that addresses tree (floral or foliar) phenology, seasonal phloem anatomy and seasonal phloem physiology together, so our knowledge of how carbon transport could fluctuate seasonally, especially in temperate climates is limited. We review phloem phenology focussing on how sieve element anatomy and phloem sap flow could affect carbon availability throughout the year with a focus on winter. To investigate whether flow is possible in the winter, we construct a simple model of phloem sap flow and investigate how changes to the sap concentration, pressure gradient and sieve plate pores could influence flow during the winter. Our model suggests that phloem transport in some species could occur year-round, even in winter, but current methods for measuring all the parameters surrounding phloem sap flow make it difficult to test this hypothesis. We highlight outstanding questions that remain about phloem functionality in the winter and emphasize the need for new methods to address gaps in our knowledge about phloem function.

Measuring Phloem Transport Velocity on a Tissue Level Using a Phloem-Mobile Dye.

Here we describe an in vivo dye-tracking method for measuring phloem transport velocity in seedlings, leaves and petioles and potentially other translucent plant tissues. The method requires measurement of the fluorescent signal of a phloem-mobile fluorescent dye using sensitive photo-sensors placed external to the plant. Following dye application, velocity is determined using laser fluorescence bleaching and measuring the time it takes for the bleach front to reach a light sensor. This method can be used to measure phloem transport velocity on intact plants with minimal disturbance and has a potential to be used under a variety of growth conditions and in the field. Because there are large differences among species in their anatomy, this method should be optimized to individual plants and tissue types.

Who operated on me? What patients understand about surgical trainees.

AIMS: To evaluate whether information delivered to patients about the level of training of the person operating on them is understandable and meaningful. To establish whether post-operative patients can identify when a doctor, undergoing training in surgery, has performed their surgery. To identify terms that are clear to patients, which can be used to improve understanding. METHOD: A 15-question survey was developed to evaluate the expectations, understanding and knowledge post-operative patients had about their recent surgery. Patients were questioned about the level of training of their surgeon, the current designations used and their knowledge about surgical training. RESULTS: Over a one-month period, 161 patients were requested to fill in surveys. Sixty-one percent of patients agreed that it is very important to know the level of training of their surgeon, but only 39% of patients could correctly identify this. Of the 62 patients who thought that a consultant had performed their procedure, 40% were correct. The terms we use to denote level of training are confusing to the majority of patients. CONCLUSION: The majority of patients are unable to recognise the level of training of the person who operated on them. There is a common public misconception that the consultant surgeon will be the operator in the majority procedures, however in teaching hospitals, a significant number of procedures are performed by doctors working under the supervision of the consultant surgeon. Many patients are not being adequately educated prior to surgery about the level of training of the person doing their procedure.

Impact of hemlock woolly adelgid (Adelges tsugae) infestation on xylem structure and function and leaf physiology in eastern hemlock (Tsuga canadensis).

Hemlock woolly adelgid (Adelges tsugae Annand) (HWA) is an invasive insect that feeds upon the foliage of eastern hemlock (Tsuga canadensis (L.) Carrière) trees, leading to a decline in health and often mortality. The exact mechanism leading to the demise of eastern hemlocks remains uncertain because little is known about how HWA infestation directly alters the host's physiology. To evaluate the physiological responses of eastern hemlock during early infestation of HWA, we measured needle loss, xylem hydraulic conductivity, vulnerability to cavitation, tracheid anatomy, leaf-level gas exchange, leaf water potential and foliar cation and nutrient levels on HWA-infested and noninfested even-aged trees in an experimental garden. HWA infestation resulted in higher xylem hydraulic conductivity correlated with an increase in average tracheid lumen area and no difference in vulnerability to cavitation, indicating that needle loss associated with HWA infestation could not be attributed to reduced xylem transport capacity. HWA-infested trees exhibited higher rates of net photosynthesis and significant changes in foliar nutrient partitioning, but showed no differences in branch increment growth rates compared with noninfested trees. This study suggests that HWA-induced decline in the health of eastern hemlock trees is not initially caused by compromised water relations or needle loss.

Maintenance of carbohydrate transport in tall trees.

Trees present a critical challenge to long-distance transport because as a tree grows in height and the transport pathway increases in length, the hydraulic resistance of the vascular tissue should increase. This has led many to question whether trees can rely on a passive transport mechanism to move carbohydrates from their leaves to their roots. Although species that actively load sugars into their phloem, such as vines and herbs, can increase the driving force for transport as they elongate, it is possible that many trees cannot generate high turgor pressures because they do not use transporters to load sugar into the phloem. Here, we examine how trees can maintain efficient carbohydrate transport as they grow taller by analysing sieve tube anatomy, including sieve plate geometry, using recently developed preparation and imaging techniques, and by measuring the turgor pressures in the leaves of a tall tree in situ. Across nine deciduous species, we find that hydraulic resistance in the phloem scales inversely with plant height because of a shift in sieve element structure along the length of individual trees. This scaling relationship seems robust across multiple species despite large differences in plate anatomy. The importance of this scaling becomes clear when phloem transport is modelled using turgor pressures measured in the leaves of a mature red oak tree. These pressures are of sufficient magnitude to drive phloem transport only in concert with structural changes in the phloem that reduce transport resistance. As a result, the key to the long-standing mystery of how trees maintain phloem transport as they increase in size lies in the structure of the phloem and its ability to change hydraulic properties with plant height.