Root and shoot phenology, architecture, and organ properties: an integrated trait network among 44 herbaceous wetland species.

Integrating traits across above- and belowground organs offers comprehensive insights into plant ecology, but their various functions also increase model complexity. This study aimed to illuminate the interspecific pattern of whole-plant trait correlations through a network lens, including a detailed analysis of the root system. Using a network algorithm that allows individual traits to belong to multiple modules, we characterize interrelations among 19 traits, spanning both shoot and root phenology, architecture, morphology, and tissue properties of 44 species, mostly herbaceous monocots from Northern Ontario wetlands, grown in a common garden. The resulting trait network shows three distinct yet partially overlapping modules. Two major trait modules indicate constraints of plant size and form, and resource economics, respectively. These modules highlight the interdependence between shoot size, root architecture and porosity, and a shoot-root coordination in phenology and dry-matter content. A third module depicts leaf biomechanical adaptations specific to wetland graminoids. All three modules overlap on shoot height, suggesting multifaceted constraints of plant stature. In the network, individual-level traits showed significantly higher centrality than tissue-level traits do, demonstrating a hierarchical trait integration. The presented whole-plant, integrated network suggests that trait covariation is essentially function-driven rather than organ-specific.

Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs.

The effects of plants on the biosphere, atmosphere and geosphere are key determinants of terrestrial ecosystem functioning. However, despite substantial progress made regarding plant belowground components, we are still only beginning to explore the complex relationships between root traits and functions. Drawing on the literature in plant physiology, ecophysiology, ecology, agronomy and soil science, we reviewed 24 aspects of plant and ecosystem functioning and their relationships with a number of root system traits, including aspects of architecture, physiology, morphology, anatomy, chemistry, biomechanics and biotic interactions. Based on this assessment, we critically evaluated the current strengths and gaps in our knowledge, and identify future research challenges in the field of root ecology. Most importantly, we found that belowground traits with the broadest importance in plant and ecosystem functioning are not those most commonly measured. Also, the estimation of trait relative importance for functioning requires us to consider a more comprehensive range of functionally relevant traits from a diverse range of species, across environments and over time series. We also advocate that establishing causal hierarchical links among root traits will provide a hypothesis-based framework to identify the most parsimonious sets of traits with the strongest links on functions, and to link genotypes to plant and ecosystem functioning.

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements.

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

Regional distribution patterns of wetland monocots with different root turnover strategies are associated with local variation in soil temperature.

Herbaceous perennial wetland monocots in Northern Ontario, Canada, show dichotomous root overwintering patterns, either with full senescence in autumn or survival over the winter, comparable to deciduous and evergreen leaf habits in trees. To test if these root strategies differ in their associations with growing season length, soil temperatures were recorded in autumn, winter and spring on 38 wetland sites with their dominant species being either of these two root overwintering strategies, altogether 19 monocot species. Traits associated with these strategies were assessed in garden experiments for a subset of these species. Sites with species with autumn-senescing roots were, on average, warmer than sites with species with overwintering roots. The 13 coldest sites were all sites of species with overwintering roots. Spring shoot growth in the field was delayed in species with autumn-senescing roots, despite their higher shoot relative growth rates in the garden, possibly due to the necessity to produce roots first. We conclude that species with autumn-senescing roots are more constrained by temperature than species with overwintering roots, limiting their occurrence in cool climates to locally warm soils and constraining their distribution towards the north. This is comparable to constraints on distribution of deciduous and evergreen trees.

Spring temperatures affect senescence and N uptake in autumn and N storage for winter in Rhynchospora alba (Cyperaceae).

Environmental and physiological factors underlying variation in timing of autumn senescence are not well known. We investigated how the time of the onset of the growth in spring affects senescence and its functional consequences for nitrogen (N) uptake in autumn and storage of N for the winter, in a species that each year develops its bulbils for storage and overwintering anew. Rhynchospora alba was grown outdoors with two treatments, identical except for a 3 week difference in the start of growth in May. Leaf and root growth and senescence, and N uptake were recorded from August to November. By August, late-starting plants had caught up in size and total N content, but had smaller bulbils. They had a higher δ 13C, indicating a higher stomatal conductance during growth. Leaf and root senescence were delayed, extending 15N tracer uptake by 4 weeks. Nevertheless, after senescence, plants with an early start had 55% more N in their overwintering bulbils, due to earlier and more efficient remobilization. We conclude that timing of senescence in R. alba is a result of an interplay between the status of winter storage and cold temperatures, constrained by a trade-off between prolonged nutrient uptake and efficient remobilization of nutrients.

Enclosed electronic system for force measurements in knee implants.

Total knee arthroplasty is a widely performed surgical technique. Soft tissue force balancing during the operation relies strongly on the experience of the surgeon in equilibrating tension in the collateral ligaments. Little information on the forces in the implanted prosthesis is available during surgery and post-operative treatment. This paper presents the design, fabrication and testing of an instrumented insert performing force measurements in a knee prosthesis. The insert contains a closed structure composed of printed circuit boards and incorporates a microfabricated polyimide thin-film piezoresistive strain sensor for each condylar compartment. The sensor is tested in a mechanical knee simulator that mimics in-vivo conditions. For characterization purposes, static and dynamic load patterns are applied to the instrumented insert. Results show that the sensors are able to measure forces up to 1.5 times body weight with a sensitivity fitting the requirements for the proposed use. Dynamic testing of the insert shows a good tracking of slow and fast changing forces in the knee prosthesis by the sensors.

Quasiuniversal connectedness percolation of polydisperse rod systems.

The connectedness percolation threshold (η(c)) and critical coordination number (Z(c)) of systems of penetrable spherocylinders characterized by a length polydispersity are studied by way of Monte Carlo simulations for several aspect ratio distributions. We find that (i) η(c) is a nearly universal function of the weight-averaged aspect ratio, with an approximate inverse dependence that extends to aspect ratios that are well below the slender rod limit and (ii) that percolation of impenetrable spherocylinders displays a similar quasiuniversal behavior. For systems with a sufficiently high degree of polydispersity, we find that Z(c) can become smaller than unity, in analogy with observations reported for generalized and complex networks.

Effects of combined drought and heavy metal stresses on xylem structure and hydraulic conductivity in red maple (Acer rubrum L.).

The effects of heavy metal stress, drought stress, and their combination on xylem structure in red maple (Acer rubrum) seedlings were investigated in an outdoor pot experiment. As metal-contaminated substrate, a mixture of 1.5% slag with sand was used, with Ni, Cu, Co, and Cr as the main contaminants. Plants grown on contaminated substrate had increased leaf metal concentrations. The two stresses reduced plant growth in an additive manner. The effects of metal and drought stresses on xylem characteristics were similar to each other, with a reduced proportion of xylem tissue, reduced conduit density in stems, and reduced conduit size in the roots. This resulted, in both stems and roots, in reductions in hydraulic conductance, xylem-specific conductivity, and leaf-specific conductivity. The similarity of the responses to the two stresses suggests that the plants' response to metals was actually a drought response, probably due to the reduced water uptake capacity of the metal-exposed roots. The only plant responses specific to metal stress were decreasing trends of stomatal density and chlorophyll content. In conclusion, the exposure to metals aggravates water stress in an additive manner, making the plants more vulnerable to drought.

Individual particle handling in a microfluidic system based on parallel laser trapping.

We present an optical trapping system combining individually addressable multiple laser traps with fluorescence spectroscopy. An in-line set of 64 near-IR laser diodes is used to create a line of individually addressable traps inside a microfluidic chip. This system is completed by an excitation/detection line for spectrally resolved fluorescence imaging of trapped particles. Highly parallel trapping in a constant flow (up to a few millimeters per second), fast particle handling rates (up to a few particles per second), and the possibility of recording fluorescence spectra of trapped objects lead to a performing bioanalytical platform, e.g., for highly parallel analysis and sorting.

Root Suberin Plays Important Roles in Reducing Water Loss and Sodium Uptake in Arabidopsis thaliana.

Suberin is a cell-wall-associated hetero-polymer deposited in specific plant tissues. The precise role of its composition and lamellae structure in protecting plants against abiotic stresses is unclear. In Arabidopsis thaliana, we tested the biochemical and physiological responses to water deficiency and NaCl treatment in mutants that are differentially affected in suberin composition and lamellae structure. Chronic drought stress increased suberin and suberin-associated waxes in wild-type plants. Suberin-deficient mutants were not more susceptible than the wild-type to the chronic drought stress imposed in this study. Nonetheless, the cyp86a1-1 cyp86b1-1 mutant, which had a severely altered suberin composition and lamellae structure, exhibited increased water loss through the root periderm. Cyp86a1-1 cyp86b1-1 also recorded lower relative water content in leaves. The abcg2-1 abcg6-1 abcg20-1 mutant, which has altered suberin composition and lamellae, was very sensitive to NaCl treatment. Furthermore, cyp86a1-1 cyp86b1-1 recorded a significant drop in the leaf K/Na ratio, indicating salt sensitivity. The far1-2 far4-1 far5-1 mutant, which did not show structural defects in the suberin lamellae, had similar responses to drought and NaCl treatments as the wild-type. Our results provide evidence that the suberin amount and lamellae structure are key features in the barrier function of suberin in reducing water loss and reducing sodium uptake through roots for better performance under drought and salt stresses.

Integrated LTCC pressure/flow/temperature multisensor for compressed air diagnostics.

We present a multisensor designed for industrial compressed air diagnostics and combining the measurement of pressure, flow, and temperature, integrated with the corresponding signal conditioning electronics in a single low-temperature co-fired ceramic (LTCC) package. The developed sensor may be soldered onto an integrated electro-fluidic platform by using standard surface mount device (SMD) technology, e.g., as a standard electronic component would be on a printed circuit board, obviating the need for both wires and tubes and thus paving the road towards low-cost integrated electro-fluidic systems. Several performance aspects of this device are presented and discussed, together with electronics design issues.

Optimal percolation of disordered segregated composites.

We evaluate the percolation threshold values for a realistic model of continuum segregated systems, where random spherical inclusions forbid the percolating objects, modeled by hardcore spherical particles surrounded by penetrable shells, to occupy large regions inside the composite. We find that the percolation threshold is generally a nonmonotonous function of segregation, and that an optimal (i.e., minimum) critical concentration exists well before maximum segregation is reached. We interpret this feature as originating from a competition between reduced available volume effects and enhanced concentrations needed to ensure percolation in the highly segregated regime. The relevance with existing segregated materials is discussed.

Determination of leaf fresh mass after storage between moist paper towels: constraints and reliability of the method.

To ensure comparability among leaf fresh mass measurements it is important to handle the leaves in a standardized manner. In the present work constraints of a commonly used method to achieve full turgor, storage between damp paper towels, were investigated. After overnight rehydration in a saturated atmosphere, the fresh mass of leaves of 14 species was measured, and the leaves were stored between paper towels (two treatments: moist and wet) at 4 degrees C. Their mass was measured after 24, 48, and 72 h. Leaf fresh mass increased during the first 24 h of storage between moist paper towels by an average of 1.8%, between wet towels by 3.3%. Among the species, the increase of leaf fresh mass between moist towels correlated with the species' desiccation propensity, indicating that it was rehydration from water loss during initial handling. On the other hand, between wet towels the fresh mass increase was associated with the species' leaf tissue structure, and it continued to increase beyond 24 h, indicating that the increase was a result of water penetration into the leaf air spaces. It is concluded that storage between moist paper towels results in reliable values of leaf fresh mass, and that desiccated leaves rehydrate well between moist towels. However, care has to be taken to avoid too wet conditions as they may lead to erroneously high fresh mass values, especially in species with large air spaces. Furthermore, exposure to unsaturated atmospheric conditions during handling has to be minimized.

Percolative properties of hard oblate ellipsoids of revolution with a soft shell.

We present an in-depth analysis of the geometrical percolation behavior in the continuum of random assemblies of hard oblate ellipsoids of revolution. Simulations were carried out by considering a broad range of aspect ratios, from spheres up to aspect-ratio-100 platelike objects, and with various limiting two-particle interaction distances, from 0.05 times the major axis up to 4.0 times the major axis. We confirm the widely reported trend of a consistent lowering of the hard particle critical volume fraction with increase of the aspect ratio. Moreover, by assimilating the limiting interaction distance to a shell of constant thickness surrounding the ellipsoids, we propose a simple relation based on the total excluded volume of these objects which allows us to estimate the critical concentration from a quantity that is quasi-invariant over a large spectrum of limiting interaction distances. Excluded volume and volume quantities are derived explicitly.

Chloroplast genomes of Lilium lancifolium, L. amabile, L. callosum, and L. philadelphicum: Molecular characterization and their use in phylogenetic analysis in the genus Lilium and other allied genera in the order Liliales.

Chloroplast (cp) genomes of Lilium amabile, L. callosum, L. lancifolium, and L. philadelphicum were fully sequenced. Using these four novel cp genome sequences and five other previously sequenced cp genomes, features of the cp genomes were characterized in detail among species in the genus Lilium and other related genera in the order Liliales. The lengths and nucleotide composition showed little variation. No structural variation was found among the cp genomes in Liliales. Gene contents were conserved among four newly sequenced cp genome in Lilium species, the only differences being in two pseudogenes. We identified 112 genes in 13 functional categories, 18 of which carried introns that were conserved among the species in Liliales. There were 16-21 SSR loci (>12 bp, >3 repeats) in the cp genomes in Lilium and the genomic locations of these loci were highly variable among the species. Average mutations were 15 SNPs per 1kb and 5 indels per 1kb, respectively, in the cp genomes of the newly sequenced four Lilium species. Phylogenetic classifications revealed some discrepancies between trees based on the cp genomes and previous classifications based on the morphology and geographic distributions.

Effects of heavy-metal-contaminated soil on growth, phenology and biomass turnover of Hieracium piloselloides.

The effects of low levels of heavy metals on plant growth, biomass turnover and reproduction were investigated for Hieracium pilosella. Plants were grown for 12 weeks on substrates with different concentrations of heavy metals obtained by diluting contaminated soils with silica sand. To minimize effects of other soil factors, the substrates were limed, fertilized, and well watered. The more metal-contaminated soil the substrate contained, the lower the leaf production rate and the plant mass were, and the more the phenological development was delayed. Flowering phenology was very sensitive to metals. Leaf life span was reduced at the highest and the lowest metal levels, the latter being a result of advanced seed ripening. Even if the effect of low metal levels on plant growth may be small, the delayed and reduced reproduction may have large effects at population, community and ecosystem level, and contribute to rapid evolution of metal tolerance.

Early Autumn Senescence in Red Maple (Acer rubrum L.) Is Associated with High Leaf Anthocyanin Content.

Several theories exist about the role of anthocyanins in senescing leaves. To elucidate factors contributing to variation in autumn leaf anthocyanin contents among individual trees, we analysed anthocyanins and other leaf traits in 27 individuals of red maple (Acer rubrum L.) over two growing seasons in the context of timing of leaf senescence. Red maple usually turns bright red in the autumn, but there is considerable variation among the trees. Leaf autumn anthocyanin contents were consistent between the two years of investigation. Autumn anthocyanin content strongly correlated with degree of chlorophyll degradation mid to late September, early senescing leaves having the highest concentrations of anthocyanins. It also correlated positively with leaf summer chlorophyll content and dry matter content, and negatively with specific leaf area. Time of leaf senescence and anthocyanin contents correlated with soil pH and with canopy openness. We conclude that the importance of anthocyanins in protection of leaf processes during senescence depends on the time of senescence. Rather than prolonging the growing season by enabling a delayed senescence, autumn anthocyanins in red maple in Ontario are important when senescence happens early, possibly due to the higher irradiance and greater danger of oxidative damage early in the season.

Root tissue structure is linked to ecological strategies of grasses.

The present study investigated to what extent there is a link between root tissue structure and ecological strategies of plant species; such a link is known for leaf tissue structure. We investigated experimentally root tissue mass density, root diameter and several characteristics of root anatomy in the axile roots of 19 perennial grass species from different habitats and related these parameters to the ecological behaviour of the species. Root characteristics were assessed in new roots produced by mature plants grown under standardized conditions. The ecological behaviour was characterized in terms of relative growth rate (RGR), plant height at maturity and ecological indicator values for nutrients, light and tolerance to mowing according to Ellenberg. We found a striking dichotomy between root anatomical characteristics associated with interspecific variation in RGR and those associated with variation in plant height. RGR correlated with anatomical characteristics that contribute to root robustness, whereas plant height correlated with characteristics associated with axile root hydraulic conductance. RGR correlated negatively with tissue mass density (TMDr ) in roots. Interspecific variation in TMDr was explained by the proportion of stele in the cross-sectional area (CSA) of the axile root and the proportion of cell wall in the CSA of the stele. For a given root diameter, slow growing species had smaller, albeit more numerous, xylem vessels, indicating a higher resistance to cavitation and protection against embolisms. Plant height correlated positively with root CSA, total xylem CSA and mean xylem vessel CSA, indicating a need for a high transport capacity in roots of species that attain a large size at maturity. TMDr correlated positively with dry matter content in leaves. The results emphasize the close relationship between tissue structure and growth characteristics at the whole-plant level.