A 2D model to study how secondary growth affects the self-supporting behaviour of climbing plants.

Climbing plants exhibit specialized shoots, called "searchers", to cross spaces and alternate between spatially discontinuous supports in their natural habitats. To achieve this task, searcher shoots combine both primary and secondary growth processes of their stems in order to support, orientate and explore their extensional growth into the environment. Currently, there is an increasing interest in developing models to describe plant growth and posture. However, the interactions between the sensing activity (e.g. photo-, gravi-, proprioceptive sensing) and the elastic responses are not yet fully understood. Here, we aim to model the extension and rigidification of searcher shoots. Our model defines variations in the radius (and consequently in mass distribution) along the shoot based on experimental data collected in natural habitats of two climbing species: Trachelospermum jasminoides (Lindl.) Lem. and Condylocarpon guianense Desf.. Using this framework, we predicted the sensory aspect of a plant, that is, the plant's response to external stimuli, and the plant's proprioception, that is, the plant's "self-awareness". The results suggest that the inclusion of the secondary growth in a model is fundamental to predict the postural development and self-supporting growth phase of shoots in climbing plants.

Microspines in tropical climbing plants: a small-scale fix for life in an obstacle course.

Many climbing plants have microspines on their stems, which facilitate attachment and prevent slipping and falling from host plant supports. Extending via growth through complex environments and anchoring stems to substrates with minimal contact forces are key benefits for climbing plants. Microspines are also highly desirable features for new technologies and applications in soft robotics. Using a novel sled-like device, we investigated static and sliding attachment forces generated by stems in 10 species of tropical climber from French Guiana differing in size and climbing habit. Eight species showed higher static and sliding forces when their stems were pulled in the basal direction against a standard surface than in the apical direction. This anisotropic behaviour suggests that tropical climbers have evolved different ratchet-like mechanisms that allow easy sliding forwards but are resistant to slipping downwards. The presence of a downwards 'stick-and-slip' phenomenon, where static attachment is not significantly stronger than maximal sliding attachment, was present in most species apart from three showing relatively weak attachment by microspines. This indicates that diverse microspine attachment strategies exist in climbing plants. This diversity of functional properties offers a range of potential design specifications for climbing strategies on different substrates for artificial climbing artefacts.

Secondary metabolites have more influence than morphophysiological traits on litter decomposability across genotypes of Arabidopsis thaliana.

Although interspecific variation in plant phenotype is recognised to impact afterlife processes such as litter decomposability, it is still unclear which traits and selection pressures explain these relationships. Examining intraspecific variation is crucial to identify and compare trait effects on decomposability, and investigate the potential role of natural selection. We studied the genetic variability and relationships between decomposability, plant traits typically related to decomposability at species level (morphophysiological traits), and leaf metabolites among a set of genotypes of Arabidopsis thaliana grown under controlled conditions. We also investigated correlations between decomposability and environmental variables at genotypes collection site. We investigated the genetic architecture of decomposability with genome-wide association studies (GWAS). There was large genetic variability in decomposability that was correlated with precipitation. Morphophysiological traits had a minor effect, while secondary metabolites, especially glucosinolates, were correlated with decomposability. Consistently, GWAS suggested that genes and metabolites related to the composition of cell membranes and envelopes control the variation of decomposability across genotypes. Our study suggests that decomposability varies within species as a result of metabolic adaptation to climate. Our findings highlight that subtle variations of defence-related metabolites like glucosinolates may strongly influence after-life processes such as decomposability.

Mechanical traits of fine roots as a function of topology and anatomy.

Background and Aims: Root mechanical traits, including tensile strength (Tr), tensile strain (εr) and modulus of elasticity (Er), are key functional traits that help characterize plant anchorage and the physical contribution of vegetation to landslides and erosion. The variability in these traits is high among tree fine roots and is poorly understood. Here, we explore the variation in root mechanical traits as well as their underlying links with morphological (diameter), architectural (topological order) and anatomical (stele and cortex sizes) traits. Methods: We investigated the four tropical tree species Pometia tomentosa, Barringtonia fusicarpa, Baccaurea ramiflora and Pittosporopsis kerrii in Xishuangbanna, Yunnan, China. For each species, we excavated intact, fresh, fine roots and measured mechanical and anatomical traits for each branching order. Key Results: Mechanical traits varied enormously among the four species within a narrow range of diameters (<2 mm): <0.1-65 MPa for Tr, 4-1135 MPa for Er and 0.4-37 % for εr. Across species, Tr and Er were strongly correlated with stele area ratio, which was also better correlated with topological order than with root diameter, especially at interspecific levels. Conclusions: Root topological order plays an important role in explaining variability in fine-root mechanical traits due to its reflection of root tissue development. Accounting for topological order when measuring fine-root traits therefore leads to greater empirical understanding of plant functions (e.g. anchorage) within and across species.

Embolism and mechanical resistances play a key role in dehydration tolerance of a perennial grass Dactylis glomerata L.

Background and Aims: More intense droughts under climate change threaten species resilience. Hydraulic strategies determine drought survival in woody plants but have been hardly studied in herbaceous species. We explored the intraspecific variability of hydraulic and morphological traits as indicators of dehydration tolerance in a perennial grass, cocksfoot (Dactylis glomerata), which has a large biogeographical distribution in Europe. Methods: Twelve populations of cocksfoot originating from Mediterranean, Temperate and Northern European areas were grown in a controlled environment in pots. Dehydration tolerance, leaf and stem anatomical traits and xylem pressure associated with 88 or 50 % loss of xylem conductance (P88, P50) were measured. Key Results: Across the 12 populations of cocksfoot, P50 ranged from -3.06 to - 6.36 MPa, while P88 ranged from -5.06 to -11.6 MPa. This large intraspecific variability of embolism thresholds corresponded with the biogeographical distribution and some key traits of the populations. In particular, P88 was correlated with dehydration tolerance (r = -0.79). The dehydration-sensitive Temperate populations exhibited the highest P88 (-6.1 MPa). The most dehydration-tolerant Mediterranean populations had the greatest leaf dry matter content and leaf fracture toughness, and the lowest P88 (-10.4 MPa). The Northern populations displayed intermediate trait values, potentially attributable to frost resistance. The thickness of metaxylem vessel walls in stems was highly correlated with P50 (r = -0.92), but no trade-off with stem lignification was observed. The relevance of the linkage between hydraulic and stomatal traits is discussed for drought survival in perennial grasses. Conclusions: Compared with woody species, the large intraspecific variability in dehydration tolerance and embolism resistance within cocksfoot has consequences for its sensitivity to climate change. To better understand adaptive strategies of herbaceous species to increasing drought and frost requires further exploration of the role of hydraulic and mechanical traits using a larger inter- and intraspecific range of species.

Major trends in stem anatomy and growth forms in the perianth-bearing Piperales, with special focus on Aristolochia.

BACKGROUND AND AIMS: The order Piperales has the highest diversity of growth forms among the earliest angiosperm lineages, including trees, shrubs, climbers and herbs. However, within the perianth-bearing Piperales (Asarum, Saruma, Lactoris, Hydnora, Prosopanche, Thottea and Aristolochia), climbing species only occur in the most species-rich genus Aristolochia. This study traces anatomical and morphological traits among these lineages, to detect trends in growth form evolution and developmental processes. METHODS: Transverse stem sections of different developmental stages of representatives of Asarum, Saruma, Lactoris, Hydnora, Thottea and Aristolochia were compared and anatomical traits were linked to growth form evolution. Biomechanical properties of representative climbers were determined in three-point bending tests and are discussed based on the anatomical observations. Growth form evolution of the perianth-bearing Piperales was reconstructed by ancestral character state reconstruction using Mesquite. KEY RESULTS: While species of Asarum and Saruma are exclusively herbaceous, species of the remaining genera show a higher diversity of growth habit and anatomy. This growth form diversity is accompanied by a more complex stem anatomy and appropriate biomechanical properties. The ancestral growth form of the perianth-bearing Piperales is reconstructed with either a shrub-like or herbaceous character state, while the following three backbone nodes in the reconstruction show a shrub-like character state. Accordingly, the climbing habit most probably evolved in the ancestor of Aristolochia. CONCLUSIONS: Since the ancestor of the perianth-bearing Piperales has been reconstructed with a herb- or shrub-like habit, it is proposed that the climbing habit is a derived growth form, which evolved with the diversification of Aristolochia, and might have been a key feature for its diversification. Observed anatomical synapomorphies, such as the perivascular fibres in Lactoris, Thottea and Aristolochia, support the phylogenetic relationship of several lineages within the perianth-bearing Piperales. In addition, the hypothesis that the vegetative organs of the holoparasitic Hydnoraceae are most probably rhizomes is confirmed.

A growing soft robot with climbing plant-inspired adaptive behaviors for navigation in unstructured environments.

Self-growing robots are an emerging solution in soft robotics for navigating, exploring, and colonizing unstructured environments. However, their ability to grow and move in heterogeneous three-dimensional (3D) spaces, comparable with real-world conditions, is still developing. We present an autonomous growing robot that draws inspiration from the behavioral adaptive strategies of climbing plants to navigate unstructured environments. The robot mimics climbing plants' apical shoot to sense and coordinate additive adaptive growth via an embedded additive manufacturing mechanism and a sensorized tip. Growth orientation, comparable with tropisms in real plants, is dictated by external stimuli, including gravity, light, and shade. These are incorporated within a vector field method to implement the preferred adaptive behavior for a given environment and task, such as growth toward light and/or against gravity. We demonstrate the robot's ability to navigate through growth in relation to voids, potential supports, and thoroughfares in otherwise complex habitats. Adaptive twining around vertical supports can provide an escape from mechanical stress due to self-support, reduce energy expenditure for construction costs, and develop an anchorage point to support further growth and crossing gaps. The robot adapts its material printing parameters to develop a light body and fast growth to twine on supports or a tougher body to enable self-support and cross gaps. These features, typical of climbing plants, highlight a potential for adaptive robots and their on-demand manufacturing. They are especially promising for applications in exploring, monitoring, and interacting with unstructured environments or in the autonomous construction of complex infrastructures.

Failure without Tears: Two-Step Attachment in a Climbing Cactus.

Climbing plants can be extremely adaptable to diverse habitats and capable of colonising perturbed, unstructured, and even moving environments. The timing of the attachment process, whether instantaneous (e.g., a pre-formed hook) or slow (growth process), crucially depends on the environmental context and the evolutionary history of the group concerned. We observed how spines and adhesive roots develop and tested their mechanical strength in the climbing cactus Selenicereus setaceus (Cactaceae) in its natural habitat. Spines are formed on the edges of the triangular cross-section of the climbing stem and originate in soft axillary buds (areoles). Roots are formed in the inner hard core of the stem (wood cylinder) and grow via tunnelling through soft tissue, emerging from the outer skin. We measured maximal spine strength and root strength via simple tensile tests using a field measuring Instron device. Spine and root strengths differ, and this has a biological significance for the support of the stem. Our measurements indicate that the measured mean strength of a single spine could theoretically support an average force of 2.8 N. This corresponds to an equivalent stem length of 2.62 m (mass of 285 g). The measured mean strength of root could theoretically support an average of 13.71 N. This corresponds to a stem length of 12.91 m (mass of 1398 g). We introduce the notion of two-step attachment in climbing plants. In this cactus, the first step deploys hooks that attach to a substrate; this process is instantaneous and is highly adapted for moving environments. The second step involves more solid root attachment to the substrate involving slower growth processes. We discuss how initial fast hook attachment can steady the plant on supports allowing for the slower root attachment. This is likely to be important in wind-prone and moving environmental conditions. We also explore how two-step anchoring mechanisms are of interest for technical applications, particularly for soft-bodied artefacts, which must safely deploy hard and stiff materials originating from a soft compliant body.

Escaping the lianoid habit: evolution of shrub-like growth forms in Aristolochia subgenus Isotrema (Aristolochiaceae).

PREMISE OF THE STUDY: A large range of growth forms is a notable aspect of angiosperm diversity and arguably a key element of their success. However, few studies within a phylogenetic context have explored how anatomical, developmental, and biomechanical traits are linked with growth form evolution. Aristolochia (∼500 species) consists predominantly of climbers, but a handful of shrub-like species are known from Aristolochia subgenus Isotrema (hereafter, shortened to Isotrema). We test hypotheses proposing that the establishment of functional traits linked to lianescence might limit the ability to evolve structurally diverse growth forms, particularly self-supporting forms. • METHODS: We focus on the origin of the shrub habit in Isotrema, from which we sampled representatives from climbing to self-supporting forms. Morphological, anatomical, and biomechanical characters are optimized on a chloroplast- and nuclear-derived phylogeny. • KEY RESULTS: Character-state reconstructions revealed that the climbing habit is plesiomorphic in Isotrema and shrub-like forms are derived from climbers. However, shrubs do not constitute a monophyletic group. Both shrubs and climbers show large multiseriate rays, but differ in terms of vessel size and proportion of fibers and soft tissues. • CONCLUSION: We suggest that while shrub-like species might have partly escaped from the constraints of life as lianas; their height size and stability are not typical of self-supporting shrubs and trees. Shrubs retained lianoid stem characters that are known to promote flexibility such as ray parenchyma. The transitions to a shrub-like form likely involved relatively simple, developmental changes that may be attributed to heterochronic processes.

Trellis-forming stems of a tropical liana Condylocarpon guianense (Apocynaceae): A plant-made safety net constructed by simple "start-stop" development.

Tropical vines and lianas have evolved mechanisms to avoid mechanical damage during their climbing life histories. We explore the mechanical properties and stem development of a tropical climber that develops trellises in tropical rain forest canopies. We measured the young stems of Condylocarpon guianensis (Apocynaceae) that construct complex trellises via self-supporting shoots, attached stems, and unattached pendulous stems. The results suggest that, in this species, there is a size (stem diameter) and developmental threshold at which plant shoots will make the developmental transition from stiff young shoots to later flexible stem properties. Shoots that do not find a support remain stiff, becoming pendulous and retaining numerous leaves. The formation of a second TYPE II (lianoid) wood is triggered by attachment, guaranteeing increased flexibility of light-structured shoots that transition from self-supporting searchers to inter-connected net-like trellis components. The results suggest that this species shows a "hard-wired" development that limits self-supporting growth among the slender stems that make up a liana trellis. The strategy is linked to a stem-twining climbing mode and promotes a rapid transition to flexible trellis elements in cluttered densely branched tropical forest habitats. These are situations that are prone to mechanical perturbation via wind action, tree falls, and branch movements. The findings suggest that some twining lianas are mechanically fine-tuned to produce trellises in specific habitats. Trellis building is carried out by young shoots that can perform very different functions via subtle development changes to ensure a safe space occupation of the liana canopy.

Effect of mechanical perturbation on the biomechanics, primary growth and secondary tissue development of inflorescence stems of Arabidopsis thaliana.

BACKGROUND AND AIMS: Mechanical perturbation is known to inhibit elongation of the inflorescence stem of Arabidopsis thaliana. The phenomenon has been reported widely for both herbaceous and woody plants, and has implications for how plants adjust their size and form to survive in mechanically perturbed environments. While this response is an important aspect of the plant's architecture, little is known about how mechanical properties of the inflorescence stem are modified or how its primary and secondary tissues respond to mechanical perturbation. METHODS: Plants of the Columbia-0 ecotype were exposed to controlled brushing treatments and then submitted to three-point bending tests to determine stem rigidity and stiffness. Contributions of different tissues to the inflorescence stem geometry were analysed. KEY RESULTS: Perturbed plants showed little difference in stem diameter, were 50 % shorter, 75 % less rigid and 70 % less stiff than controls. Changes in mechanical properties were linked to significant changes in tissue geometry - size and position of the pith, lignified interfascicular tissue and cortex - as well as a reduction in density of lignified cells. Stem mechanical properties were modified by changes in primary tissues and thus differ from changes observed in most woody plants tested with indeterminate growth - even though a vascular cambium is present in the inflorescence axis. CONCLUSIONS: The study suggests that delayed development of key primary developmental features of the stem in this ecotype of Arabidopsis results in a 'short and flexible' rather than a 'short and rigid' strategy for maintaining upright axes in conditions of severe mechanical perturbation. The mechanism is comparable with more general phenomena in plants where changes in developmental rate can significantly affect the overall growth form of the plant in both ecological and evolutionary contexts.

Structural performance of a climbing cactus: making the most of softness.

Climbing plants must reach supports and navigate gaps to colonize trees. This requires a structural organization ensuring the rigidity of so-called 'searcher' stems. Cacti have succulent stems adapted for water storage in dry habitats. We investigate how a climbing cactus Selenicereus setaceus develops its stem structure and succulent tissues for climbing. We applied a 'wide scale' approach combining field-based bending, tensile and swellability tests with fine-scale rheological, compression and anatomical analyses in laboratory conditions. Gap-spanning 'searcher' stems rely significantly on the soft cortex and outer skin of the stem for rigidity in bending (60-94%). A woody core contributes significantly to axial and radial compressive strength (80%). Rheological tests indicated that storage moduli were consistently higher than loss moduli indicating that the mucilaginous cortical tissue behaved like a viscoelastic solid with properties similar to physical or chemical hydrogels. Rheological and compression properties of the soft tissue changed from young to old stages. The hydrogel-skin composite is a multi-functional structure contributing to rigidity in searcher stems but also imparting compliance and benign failure in environmental situations when stems must fail. Soft tissue composites changing in function via changes in development and turgescence have a great potential for exploring candidate materials for technical applications.

Mechanosensing of stem bending and its interspecific variability in five neotropical rainforest species.

BACKGROUND AND AIMS: In rain forests, sapling survival is highly dependent on the regulation of trunk slenderness (height/diameter ratio): shade-intolerant species have to grow in height as fast as possible to reach the canopy but also have to withstand mechanical loadings (wind and their own weight) to avoid buckling. Recent studies suggest that mechanosensing is essential to control tree dimensions and stability-related morphogenesis. Differences in species slenderness have been observed among rainforest trees; the present study thus investigates whether species with different slenderness and growth habits exhibit differences in mechanosensitivity. METHODS: Recent studies have led to a model of mechanosensing (sum-of-strains model) that predicts a quantitative relationship between the applied sum of longitudinal strains and the plant's responses in the case of a single bending. Saplings of five different neotropical species (Eperua falcata, E. grandiflora, Tachigali melinonii, Symphonia globulifera and Bauhinia guianensis) were subjected to a regimen of controlled mechanical loading phases (bending) alternating with still phases over a period of 2 months. Mechanical loading was controlled in terms of strains and the five species were subjected to the same range of sum of strains. The application of the sum-of-strain model led to a dose-response curve for each species. Dose-response curves were then compared between tested species. KEY RESULTS: The model of mechanosensing (sum-of-strain model) applied in the case of multiple bending as long as the bending frequency was low. A comparison of dose-response curves for each species demonstrated differences in the stimulus threshold, suggesting two groups of responses among the species. Interestingly, the liana species B. guianensis exhibited a higher threshold than other Leguminosae species tested. CONCLUSIONS: This study provides a conceptual framework to study variability in plant mechanosensing and demonstrated interspecific variability in mechanosensing.

Mechanical Innovations of a Climbing Cactus: Functional Insights for a New Generation of Growing Robots.

Climbing plants are being increasingly viewed as models for bioinspired growing robots capable of spanning voids and attaching to diverse substrates. We explore the functional traits of the climbing cactus Selenicereus setaceus (Cactaceae) from the Atlantic forest of Brazil and discuss the potential of these traits for robotics applications. The plant is capable of growing through highly unstructured habitats and attaching to variable substrates including soil, leaf litter, tree surfaces, rocks, and fine branches of tree canopies in wind-blown conditions. Stems develop highly variable cross-sectional geometries at different stages of growth. They include cylindrical basal stems, triangular climbing stems and apical star-shaped stems searching for supports. Searcher stems develop relatively rigid properties for a given cross-sectional area and are capable of spanning voids of up to 1 m. Optimization of rigidity in searcher stems provide some potential design ideas for additive engineering technologies where climbing robotic artifacts must limit materials and mass for curbing bending moments and buckling while climbing and searching. A two-step attachment mechanism involves deployment of recurved, multi-angled spines that grapple on to wide ranging surfaces holding the stem in place for more solid attachment via root growth from the stem. The cactus is an instructive example of how light mass searchers with a winged profile and two step attachment strategies can facilitate traversing voids and making reliable attachment to a wide range of supports and surfaces.

Developmental plasticity and biomechanics of treelets and lianas in Manihot aff. quinquepartita (Euphorbiaceae): a branch-angle climber of French Guiana.

BACKGROUND AND AIMS: Most tropical lianas have specialized organs of attachment such as twining stems, hooks or tendrils but some do not. Many climbers also have an early self-supporting phase of growth and in some species this can produce treelet-sized individuals. This study focuses on how a liana can climb without specialized attachment organs and how biomechanical properties of the stem are modulated between self-supporting treelets and canopy-climbing lianas. METHODS: Biomechanics and stem development were investigated in self-supporting to climbing individuals of Manihot aff. quinquepartita (Euphorbiaceae) from tropical rain forest at Saül, central French Guiana. Bending tests were carried out close to the site of growth. Mechanical properties, including Young's elastic modulus, were observed with reference to habit type and changes in stem anatomy during development. KEY RESULTS: This liana species can show a remarkably long phase of self-supporting growth as treelets with stiff, juvenile wood characterizing the branches and main stem. During the early phase of climbing, stiff but unstable stem segments are loosely held in a vertical position to host plants via petiole bases. The stiffest stems--those having the highest values of Young's modulus measured in bending--belonged to young, leaning and climbing stems. Only when climbing stems are securely anchored into the surrounding vegetation by a system of wide-angled branches, does the plant develop highly flexible stem properties. As in many specialized lianas, the change in stiffness is linked to the development of wood with numerous large vessels and thin-walled fibres. CONCLUSIONS: Some angiosperms can develop highly effective climbing behaviour and specialized flexible stems without highly specialized organs of attachment. This is linked to a high degree of developmental plasticity in early stages of growth. Young individuals in either open or closed marginal forest conditions can grow as substantial treelets or as leaning/climbing plants, depending on the availability of host supports. The species of liana studied differs both in terms of development and biomechanics from many other lianas that climb via twining, tendrils or other specialized attachment organs.

Leaf economics and slow-fast adaptation across the geographic range of Arabidopsis thaliana.

Life history strategies of most organisms are constrained by resource allocation patterns that follow a 'slow-fast continuum'. It opposes slow growing and long-lived organisms with late investment in reproduction to those that grow faster, have earlier and larger reproductive effort and a short longevity. In plants, the Leaf Economics Spectrum (LES) depicts a leaf-level trade-off between the rate of carbon assimilation and leaf lifespan, as stressed in functional ecology from interspecific comparative studies. However, it is still unclear how the LES is connected to the slow-fast syndrome. Interspecific comparisons also impede a deep exploration of the linkage between LES variation and adaptation to climate. Here, we measured growth, morpho-physiological and life-history traits, at both the leaf and whole-plant levels, in 378 natural accessions of Arabidopsis thaliana. We found that the LES is tightly linked to variation in whole-plant functioning, and aligns with the slow-fast continuum. A genetic analysis further suggested that phenotypic differentiation results from the selection of different slow-fast strategies in contrasted climates. Slow growing and long-lived plants were preferentially found in cold and arid habitats while fast growing and short-lived ones in more favorable habitats. Our findings shed light on the role of the slow-fast continuum for plant adaptation to climate. More broadly, they encourage future studies to bridge functional ecology, genetics and evolutionary biology to improve our understanding of plant adaptation to environmental changes.

Lianas.

Rowe introduces the beautiful and fascinating plants classified as lianas.

Geometry, Allometry and Biomechanics of Fern Leaf Petioles: Their Significance for the Evolution of Functional and Ecological Diversity Within the Pteridaceae.

Herbaceous plants rely on a combination of turgor, ground tissues and geometry for mechanical support of leaves and stems. Unlike most angiosperms however, ferns employ a sub-dermal layer of fibers, known as a hypodermal sterome, for support of their leaves. The sterome is nearly ubiquitous in ferns, but nothing is known about its role in leaf biomechanics. The goal of this research was to characterize sterome attributes in ferns that experience a broad range of mechanical stresses, as imposed by their aquatic, xeric, epiphytic, and terrestrial niches. Members of the Pteridaceae meet this criteria well. The anatomical and functional morphometrics along with published values of tissue moduli were used to model petiole flexural rigidity and susceptibility to buckling in 20 species of the Pteridaceae. Strong allometric relationships were observed between sterome thickness and leaf size, with the sterome contributing over 97% to petiole flexural rigidity. Surprisingly, the small-statured cheilanthoid ferns allocated the highest fraction of their petiole to the sterome, while large leaves exploited aspects of geometry (second moment of area) to achieve bending resistance. This pattern also revealed an economy of function in which increasing sterome thickness was associated with decreasing fiber cell reinforcement, and fiber wall fraction. Lastly, strong petioles were associated with durable leaves, as approximated by specific leaf area. This study reveals meaningful patterns in fern leaf biomechanics that align with species leaf size, sterome attributes and life-history strategy.

Function of epidermal surfaces in the trapping efficiency of Nepenthes alata pitchers.

• Several epidermal microstructures characterize surfaces of pitcher plants and are presumably involved in their trapping function. Here we report the effects of Nepenthes alata surfaces on insect locomotion and trapping efficiency. • The architectural designs of pitcher surfaces were characterized using scanning electron microscopy. Two insect species - fruitfly (Drosophila melanogaster) and ant (Iridomyrmex humilis) - were tested for their ability to remain and walk on them. The relative contributions of various epidermal structures to trapping ability were quantified. • Pitchers were very effective traps for both insect species. They were slightly more efficient in capturing the ants, but slightly more effective in retaining captured flies. Trapping efficiency was attributed to the combined effects of several surfaces displaying different functions. The waxy zone played a key role in the slippery syndrome: in addition to the wax itself, the subjacent layer of convex lunate cells interfered considerably with insect locomotion. The unsubmersed glandular zone displayed an important retentive effect and secretions of the digestive glands are suspected to be adhesive. • Pad performances of the hairy and smooth system of attachment are discussed to explain the differences between the two insect species. This study aims to encourage biomechanical studies of plant-insect surface mechanisms.