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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 selfsupporting growth phase of shoots in climbing plants.

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.

Cacti are of interest for new bio-inspired technologies because of their remarkable adaptations to extreme environments. Recently, they have inspired functional designs from nano fibres to optimised buildings and architectures. We investigate the diversity of cactus skin properties in terms of toughness and resistance to cutting damage. Cacti are well known for their extreme adaptations to harsh environments, with soft, fleshy stems that expand and contract with water uptake and storage. This functioning is made possible by an extendable outer envelope (skin) and a fluted 3-dimensional structure of the stem. We explore the mechanical toughness and underlying structural organisation of the cactus skin in four species of cactus showing different growth forms. The toughness properties of the cactus skin is only one part of a multi-functional structure for surviving in extreme environments. The study suggests that survival involves a relatively “light” investment of tough materials in the outer envelope instead of a rigid “defensive” layer. This is capable of elastic deformation and enables water storage in challenging, arid environments. The main purpose of this article is to demonstrate the diversity of skin toughness and underlying structures in the biological world as providing potential new designs for technical envelopes.

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.

Climbing plants need to reach supports and position their leaves for light capture. Vines and lianas develop a large diversity of self-supporting shoots among diverse species and different kinds of attachment. A searcher's reach is a crucial trait for colonising supports in complex three-dimensional spaces. We explore the reach capacity and diversity of searcher shoots among representative temperate and tropical climbing plants. We investigate the overall range of variation between short-and long-reach searchers; the mechanical and anatomical organisations underlying reach capacities; how searcher architectures are linked to different climbing strategies such as stem twining, tendril climbing, root climbing, and branch-angle-hook climbing. We investigated reach and mechanical and anatomical organisations (stem rigidity and stiffness, stem and tissue geometry) in 29 climbing plant species from temperate and tropical habitats. Searchers show a wide range of maximal reach per species from 0.1 to 2.5 m. Flexural rigidity (EI) at the base of searchers increased with reach length; overall this increase was proportional although some longest-reaching shoots develop proportionally thinner searcher bases with higher stiffness [structural Young's modulus (E str)] than shorter-reach shoots. Bases of short-reach searchers rely more on primary tissues compared to long-reach shoots, which rely more on wood production. We identified different mechanical architectures for a given reach capacity across all species. These are linked to different kinds of attachment mechanisms, support foraging, and possibly leaf display. Plants attaching by twining of the main stem showed a wide range of reach capacity. They also developed lighter, more slender, less rigid, but generally relatively stiff (higher E str) shoots compared with tendril climbers and branch-angle-hook climbers. Differences in the mechanical architecture of searcher shoots in climbing plants are informative for understanding how diverse climbing plant species explore and colonise different kinds of three-dimensional spaces. This is a key feature that distinguishes different habitat preferences. We discuss how such knowledge is not only important for understanding functional biology and ecology of climbing plants but is also of interest for developing new technologies in soft robotics that mimic climbing plants that can navigate through unstructured environments.

Climbing palms in the Arecoideae (Desmoncus) and Calamoideae (rattan palms) both evolved cirrate leaves armed with hooks and grapnels for climbing. Some species of Calamoideae develop a different climbing organ known as the flagellum, which also bears hooks. The present study indicates that geometry and mechanical properties of the cirrus vary between species. Cirrate leaves are constructed to optimize bending and torsion in relation to the deployment of recurved hooks. Hook development, size, and strength vary along cirri and flagella and are consistent with observations of these attachment organs functioning as a ratchet mechanism: hooks increase in strength toward the base of attachment organs and always fail before the axis in strength tests. Hook size and strength differ between species and are related to body size and ecological preference. Larger species produce larger hooks, but smaller climbing palms of the understory deploy fine sharp hooks that are effective on small diameter supports as well as large branches and trunks. The ephemeral nature of climbing organs in palms provides a challenge to their life-history development, particularly in terms of mechanical constraints and remaining attached to the host vegetation; these differ significantly from many vines and lianas having more perennial modes of attachment.

Domestication can influence many functional traits in plants, from overall life-history and growth form to wood density and cell wall ultrastructure. Such changes can increase fitness of the domesticate in agricultural environments but may negatively affect survival in the wild. We studied effects of domestication on stem biomechanics in manioc by comparing domesticated and ancestral wild taxa from two different regions of greater Amazonia. We compared mechanical properties, tissue organisation and wood characteristics including microfibril angles in both wild and domesticated plants, each growing in two different habitats (forest or savannah) and varying in growth form (shrub or liana). Wild taxa grew as shrubs in open savannah but as lianas in overgrown and forested habitats. Growth form plasticity was retained in domesticated manioc. However, stems of the domesticate showed brittle failure. Wild plants differed in mechanical architecture between shrub and liana phenotypes, a difference that diminished between shrubs and lianas of the domesticate. Stems of wild plants were generally stiffer, failed at higher bending stresses and were less prone to brittle fracture compared with shrub and liana phenotypes of the domesticate. Biomechanical differences between stems of wild and domesticated plants were mainly due to changes in wood density and cellulose microfibril angle rather than changes in secondary growth or tissue geometry. Domestication did not significantly modify \"large-scale\" trait development or growth form plasticity, since both wild and domesticated manioc can develop as shrubs or lianas. However, \"finer-scale\" developmental traits crucial to mechanical stability and thus ecological success of the plant were significantly modified. This profoundly influenced the likelihood of brittle failure, particularly in long climbing stems, thereby also influencing the survival of the domesticate in natural situations vulnerable to mechanical perturbation. We discuss the different selective pressures that could explain evolutionary modifications of stem biomechanical properties under domestication in manioc.

A striking feature of early angiosperm lineages is the variety of life forms and growth forms, which ranges from herbs, aquatic herbs, climbers, and epiphytes to woody shrubs and trees. This morphological and anatomical diversity is arguably one of the factors explaining how angiosperms dominate many ecosystems worldwide. However, just how such a wide spectrum of growth forms has evolved in angiosperms remains unclear. In this review, we investigate patterns of growth form diversification in Piperales, an early-diverging lineage (with stem age estimated at 201–128 Myr ago) and the most morphologically diverse clade among magnoliids. We outline patterns of growth form diversity and architecture as well as the biomechanical significance of developmental characters, such the organization, loss, and gain of woodiness. Asaroideae and Saururaceae are terrestrial as well as semiaquatic to aquatic herbaceous perennials bearing rhizomes. The Aristolochioideae and Piperaceae show higher levels of growth form diversity and biomechanical organization, with complex patterns of increasing or decreasing woodiness and architectural organization. The climbing habit has probably evolved independently in the Aristolochiaceae and Piperaceae, while mechanically unstable shrubs and, less frequently, treelets have evolved several times within these two most species-rich clades. A key developmental character underlying diversity in most Piperales—with the exception of the herbaceousSaruma(Asaroideae)—is the conserved development of the wood cylinder, in which fusiform initials are limited to fascicular cambial initials. The resulting large fraction of raylike tissue in the stem—a highly characteristic feature of woody species in the Piperales—potentially introduced mechanical constraints on the diversification of self-supporting architectures. This was possibly circumvented by the architectural development of repeated, large-diameter meristems in some shrublike habits via sympodial growth. Patterns of growth form evolution within Piperales potentially mirror some of the overall trends observed among early-diverging angiosperms as a whole as well as angiosperms in general. These include profound changes in life form and growth form linked to large-scale transitions in woodiness, diversity of mechanical organization, and shifts in architectural development.

• 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.