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Plant roots are considered highly efficient soil explorers. As opposed to the push‐driven penetration strategy commonly used by many digging organisms, roots penetrate by growing, adding new cells at the tip, and elongating over a well‐defined growth zone. However, a comprehensive understanding of the mechanical aspects associated with root penetration is currently lacking. We perform penetration experiments following Arabidopsis thaliana roots growing into an agar gel environment, and a needle of similar dimensions pushed into the same agar. We measure and compare the environmental deformations in both cases by following the displacement of fluorescent beads embedded within the gel, combining confocal microscopy and Digital Volume Correlation (DVC) analysis. We find that deformations are generally smaller for growing roots. To better understand the mechanical differences between the two penetration strategies, we develop a computational model informed by experiments. Simulations show that, compared to push‐driven penetration, grow‐driven penetration reduces frictional forces and mechanical work, with lower propagation of displacements in the surrounding medium. These findings shed light on the complex interaction of plant roots with their environment, providing a quantitative understanding based on a comparative approach. Summary Statement: Inspired by the exceptional penetration abilities of plant roots, here we perform an experimental and computational analysis of root‐soil mechanical interactions. We find that, compared to pushing, a grow‐driven strategy reduces frictional forces and mechanical work, with lower propagation of displacements in the surrounding medium. [ABSTRACT FROM AUTHOR]

This study sought to better understand how time of day (ToD) or turgor pressure might affect the flexural stiffness of sweet sorghum stalks and potentially regulate stalk lodging resistance. Stalk flexural stiffness measured across a 48 h period in 2019 showed a significant diurnal association with leaf water potential and stalk flexural stiffness. While the correlation between stalk flexural stiffness and this proxy for internal turgor status was statistically significant, it only accounted for roughly 2% of the overall variance in stiffness. Given that turgor status is a dynamic rather than fixed physiological variable like the cellular structure, these data suggest that internal turgor plays a small yet significant role in influencing the flexural stiffness of fully mature stalks prior to a stalk lodging event. The association was assessed at earlier developmental stages across three distinct cultivars and found not to be significant. Panicle weight and stalk basal weight, but not stalk Brix or water content, were found to be better predictors of stalk flexural stiffness than either ToD or turgor status. Observation across three cultivars and four distinct developmental stages ranging from the vegetative to the hard-dough stages suggests that stalk flexural stiffness changes significantly as a function of time. However, neither ToD nor turgor status appear to meaningfully contribute to observed variations in stalk flexural stiffness in either individual stalks or across larger populations. As turgor status was not found to meaningfully influence stalk strength or flexural stiffness at any developmental time point examined in any of the three sweet sorghum cultivars under study, turgor pressure likely offers only inconsequential contributions to the biomechanics underlying sweet sorghum stalk lodging resistance. [ABSTRACT FROM AUTHOR]

About forty years after its first application, digital image correlation (DIC) has become an established method for measuring surface displacements and deformations of objects under stress. To date, DIC has been used in a variety of in vitro and in vivo studies to biomechanically characterise biological samples in order to reveal biomimetic principles. However, when surfaces of samples strongly deform or twist, they cannot be thoroughly traced. To overcome this challenge, different DIC setups have been developed to provide additional sensor perspectives and, thus, capture larger parts of an object's surface. Herein, we discuss current solutions for this multi-perspective DIC, and we present our own approach to a 360° DIC system based on a single stereo-camera setup. Using this setup, we are able to characterise the desiccation-driven opening mechanism of two woody Hakea fruits over their entire surfaces. Both the breaking mechanism and the actuation of the two valves in predominantly dead plant material are models for smart materials. Based on these results, an evaluation of the setup for 360° DIC regarding its use in deducing biomimetic principles is given. Furthermore, we propose a way to improve and apply the method for future measurements. [ABSTRACT FROM AUTHOR]

This study sought to better understand how time of day (ToD) or turgor pressure might affect the flexural stiffness of sweet sorghum stalks and potentially regulate stalk lodging resistance. Stalk flexural stiffness measured across a 48 h period in 2019 showed a significant diurnal association with leaf water potential and stalk flexural stiffness. While the correlation between stalk flexural stiffness and this proxy for internal turgor status was statistically significant, it only accounted for roughly 2% of the overall variance in stiffness. Given that turgor status is a dynamic rather than fixed physiological variable like the cellular structure, these data suggest that internal turgor plays a small yet significant role in influencing the flexural stiffness of fully mature stalks prior to a stalk lodging event. The association was assessed at earlier developmental stages across three distinct cultivars and found not to be significant. Panicle weight and stalk basal weight, but not stalk Brix or water content, were found to be better predictors of stalk flexural stiffness than either ToD or turgor status. Observation across three cultivars and four distinct developmental stages ranging from the vegetative to the hard-dough stages suggests that stalk flexural stiffness changes significantly as a function of time. However, neither ToD nor turgor status appear to meaningfully contribute to observed variations in stalk flexural stiffness in either individual stalks or across larger populations. As turgor status was not found to meaningfully influence stalk strength or flexural stiffness at any developmental time point examined in any of the three sweet sorghum cultivars under study, turgor pressure likely offers only inconsequential contributions to the biomechanics underlying sweet sorghum stalk lodging resistance.

Tendrils of climbing plants coil along their length, thus forming a striking helical spring and generating tensional forces. It is found that, for tendrils of the passion flower Passiflora caerulea, the generated force lies in the range of 6–140 mN, which is sufficient to lash the plant tightly to its substrate. Further, it is revealed that the generated force strongly correlates with the water status of the plant. Based on a combination of in situ force measurements with anatomical investigations and dehydration‐rehydration experiments on both entire tendril segments and isolated lignified tissues, a two‐phasic mechanism for spring formation is proposed. First, during the free coiling phase, the center of the tendril begins to lignify unilaterally. At this stage, both the generated tension and the stability of the form of the spring still depend on turgor pressure. The unilateral contraction of a bilayer as being the possible driving force for the tendril coiling motion is discussed. Second, in a stabilization phase, the entire center of the coiled tendril lignifies, stiffening the spring and securing its function irrespective of its hydration status. [ABSTRACT FROM AUTHOR]

About forty years after its first application, digital image correlation (DIC) has become an established method for measuring surface displacements and deformations of objects under stress. To date, DIC has been used in a variety of in vitro and in vivo studies to biomechanically characterise biological samples in order to reveal biomimetic principles. However, when surfaces of samples strongly deform or twist, they cannot be thoroughly traced. To overcome this challenge, different DIC setups have been developed to provide additional sensor perspectives and, thus, capture larger parts of an object’s surface. Herein, we discuss current solutions for this multi-perspective DIC, and we present our own approach to a 360° DIC system based on a single stereo-camera setup. Using this setup, we are able to characterise the desiccation-driven opening mechanism of two woody Hakea fruits over their entire surfaces. Both the breaking mechanism and the actuation of the two valves in predominantly dead plant material are models for smart materials. Based on these results, an evaluation of the setup for 360° DIC regarding its use in deducing biomimetic principles is given. Furthermore, we propose a way to improve and apply the method for future measurements.

Abstract Tendrils of climbing plants coil along their length, thus forming a striking helical spring and generating tensional forces. It is found that, for tendrils of the passion flower Passiflora caerulea, the generated force lies in the range of 6–140 mN, which is sufficient to lash the plant tightly to its substrate. Further, it is revealed that the generated force strongly correlates with the water status of the plant. Based on a combination of in situ force measurements with anatomical investigations and dehydration‐rehydration experiments on both entire tendril segments and isolated lignified tissues, a two‐phasic mechanism for spring formation is proposed. First, during the free coiling phase, the center of the tendril begins to lignify unilaterally. At this stage, both the generated tension and the stability of the form of the spring still depend on turgor pressure. The unilateral contraction of a bilayer as being the possible driving force for the tendril coiling motion is discussed. Second, in a stabilization phase, the entire center of the coiled tendril lignifies, stiffening the spring and securing its function irrespective of its hydration status.

ABSTRACT Lianas are woody climbers and their stems and leaves deal with different environmental pressures such as resistance to mechanical damage and dehydration. The damage resistance of plants can be biomechanically evaluated by their stiffness, bending and toughness. Despite the well-known relationship between physical resistance and moisture of plant organs in woody plants, this relationship is uncertain and has not been previously evaluated in lianas. Thus, this study investigated experimentally the effect of stems and leaf dehydration on the structural Young’s modulus in the stem and fracture toughness in leaves across time in the liana Amphilophium crucigerum (Bignoniaceae). Ten stem and leaf samples were collected and assigned to two distinct conditions: (i) samples kept moist and (ii) samples underwent gradual dehydration with natural moisture loss by air exposition. Successive measures of structural Young’s modulus and fracture toughness were taken every 4 hours during a 48-hour period for both conditions. Stem and leaf samples which underwent gradual dehydration showed greater bending stiffness and fracture toughness, respectively, while the samples kept moist presented no changes in any studied biomechanical features during the entire experiment. We concluded that the moisture of both stem and leaf samples are critical factors to estimate the biomechanical properties of lianas stem and leaves.

Many plant tissues exhibit the property of frost resistance. This is mainly due to two factors: one is related to metabolic effects, while the other stems from structural properties of plants leading to dehydration of their cells. The present contribution aims at assessing the impact of ice formation on frost-resistant plant tissues with a focus on structural properties specifically applied to Equisetum hyemale. In this particular case, there is an extracellular ice formation in so-called vallecular canals and the pith cavity, which leads to a dehydration of the tissue cells to avoid intracellular ice formation, what would be fatal for the cells and subsequently for the whole plant. To address the underlying phenomena in the plant, a coupled thermo-hydro-mechanical model based on the Theory of Porous Media is introduced as the modelling framework. The dehydration of the tissue cells is referred to as of quasi-double-porosity nature, since the water is mobile within the intercellular space, but confined to the cells in the intracellular space and consequently kinematically coupled to them. However, the mass exchange of water across the cell wall is considered. The presented numerical example shows the strong coupling of the underlying processes as well as the quasi-double-porosity feature. Finally, it supports the experimental finding of the vallecular canals as the main location of ice formation. [ABSTRACT FROM AUTHOR]

Plants have evolved different mechanisms to disperse from parent plants and improve germination to sustain their survival. The study of seed dispersal mechanisms, with the related structural and functional characteristics, is an active research topic for ecology, plant diversity, climate change, as well as for its relevance for material science and engineering. The natural mechanisms of seed dispersal show a rich source of robust, highly adaptive, mass and energy efficient mechanisms for optimized passive flying, landing, crawling and drilling. The secret of seeds mobility is embodied in the structural features and anatomical characteristics of their tissues, which are designed to be selectively responsive to changes in the environmental conditions, and which make seeds one of the most fascinating examples of morphological computation in Nature. Particularly clever for their spatial mobility performance, are those seeds that use their morphology and structural characteristics to be carried by the wind and dispersed over great distances (i.e. “winged” and “parachute” seeds), and seeds able to move and penetrate in soil with a self-burial mechanism driven by their hygromorphic properties and morphological features. By looking at their motion mechanisms, new design principles can be extracted and used as inspiration for smart artificial systems endowed with embodied intelligence. This mini-review systematically collects, for the first time together, the morphological, structural, biomechanical and aerodynamic information from selected plant seeds relevant to take inspiration for engineering design of soft robots, and discusses potential future developments in the field across material science, plant biology, robotics and embodied intelligence.

Flax is one of the oldest cultivated plants and one of the most important fiber crops in the world. Thousands of years of selection have resulted in the unique biomechanics of this plant, the principles of which are not yet fully understood. Detailed anatomical analysis of flax stems at different heights and at different stages of plant development (herringbone, fast growth, flowering) allowed us to follow the dynamics of primary and secondary growth of these plants. The evaluation of the mechanical properties of the stems and their individual parts at the same stages made it possible to identify the tissues that take on the main mechanical load at different periods of the plant's life. In the younger parts of the stems, this tissue is the fiber-enriched phloem, whose fibers rapidly acquire a considerable elastic modulus. Xylem, which takes longer to develop, provides the main mechanical load in more mature parts of the stems and at later stages of plant development. Fast growth of flax stems occurs against the background of delayed xylem development. The onset of cell wall thickening in phloem fibers (the position of the snap point) is not determined by the mass of the plant part above it. The results obtained highlight the contribution of different tissues to the functioning of the plant as a holistic biomechanical system. • The contributions of phloem fibers and xylem into stem strength change with time. • Younger stem parts mechanically rely more on phloem fibers, older parts – on xylem. • Maximum plant elongation occurs with temporary reduction of secondary growth. • The position of the snap point is independent of the mass of the plant part above it. • The area of stem cavity correlates with the area of bast fiber tertiary cell walls. [ABSTRACT FROM AUTHOR]

This paper explores the links between genotype, plant development, plant structure and plant material properties. The barley husk has two organs, the lemma and the palea, which protect the grain. When the husk is exposed to mechanical stress, such as during harvesting, it can be damaged or detached. This is known as grain skinning, which is detrimental to grain quality and has a significant economic impact on industry. This study focused on the lemma, the husk organ which is most susceptible to grain skinning. This study tested three hypotheses: (1) genotype and plant development determine lemma structure, (2) lemma structure influences the material properties of the lemma, and (3) the material properties of the lemma determine grain skinning risk. The effect of genotype was investigated by using plant material from four malting barley varieties: two with a high risk of grain skinning, two with a low risk. Plant material was assessed at two stages of plant development (anthesis, GS 65; grain filling, GS 77). Structure was assessed using light microscopy to measure three physiological features: thickness, vasculature and cell area. Material properties were approximated using a controlled impact assay and by analyzing fragmentation behavior. Genotype had a significant effect on lemma structure and material properties from anthesis. This indicates that differences between genotypes were established during floral development. The lemma was significantly thinner in high risk genotypes, compared to low risk genotypes. Consequently, in high risk genotypes, the lemma was significantly more likely to fragment. This indicates a relationship between reduced lemma thickness and increased fragmentation. Traditionally, a thin husk has been considered beneficial for malting quality, due to an association with malt extract. However, this study finds a thin lemma is less resistant to mechanical stress. This may explain the differences in grain skinning risk in the genotypes studied.

This paper explores the links between genotype, plant development, plant structure and plant material properties. The barley husk has two organs, the lemma and the palea, which protect the grain. When the husk is exposed to mechanical stress, such as during harvesting, it can be damaged or detached. This is known as grain skinning , which is detrimental to grain quality and has a significant economic impact on industry. This study focused on the lemma, the husk organ which is most susceptible to grain skinning. This study tested three hypotheses: (1) genotype and plant development determine lemma structure, (2) lemma structure influences the material properties of the lemma, and (3) the material properties of the lemma determine grain skinning risk. The effect of genotype was investigated by using plant material from four malting barley varieties: two with a high risk of grain skinning, two with a low risk. Plant material was assessed at two stages of plant development (anthesis, GS 65; grain filling, GS 77). Structure was assessed using light microscopy to measure three physiological features: thickness, vasculature and cell area. Material properties were approximated using a controlled impact assay and by analyzing fragmentation behavior. Genotype had a significant effect on lemma structure and material properties from anthesis. This indicates that differences between genotypes were established during floral development. The lemma was significantly thinner in high risk genotypes, compared to low risk genotypes. Consequently, in high risk genotypes, the lemma was significantly more likely to fragment. This indicates a relationship between reduced lemma thickness and increased fragmentation. Traditionally, a thin husk has been considered beneficial for malting quality, due to an association with malt extract. However, this study finds a thin lemma is less resistant to mechanical stress. This may explain the differences in grain skinning risk in the genotypes studied. [ABSTRACT FROM AUTHOR]

Plant shoot gravitropism is a complex phenomenon resulting from gravity sensing, curvature sensing (proprioception), the ability to uphold self-weight and growth. Although recent data analysis and modelling have revealed the detailed morphology of shoot bending, the relative contribution of bending force (derived from the gravi-proprioceptive response) and stretching force (derived from shoot axial growth) behind gravitropism remains poorly understood. To address this gap, we combined morphological data with a theoretical model to analyze shoot bending in wild-type and lazy1-like 1 mutant Arabidopsis thaliana. Using data from actual bending events, we searched for model parameters that minimized discrepancies between the data and mathematical model. The resulting model suggests that both the bending force and the stretching force differ significantly between the wild type and mutant. We discuss the implications of the mechanical forces associated with differential cell growth and present a plausible mechanical explanation of shoot gravitropism.

Featured Application: The featured application of this work is the optimization of breeding techniques for plants susceptible to being damaged by wind. Stem lodging is the bending or breakage of stems in the wind that result in negative economic impacts to producers and processors of small grain crops. To address this issue, plant breeders attempt to quantify lodging using proxy traits such as stem structure and biomechanics. Stem lodging is a function of both stem strength and elasticity. In this paper, we explore the biomechanics of stems approaching the lodging, or permanent bending, condition. Oat, wheat, and two types of barley varying in lodging resistance were exposed to standard growing conditions over the course of a season. Their capability of returning from a bent to unbent state was characterized using a push force meter that measured resistant force and displacement over time. Changes in stem energy and power were then calculated using displacement and force measurements. Lodging susceptibility could be differentiated by stem strength, displacement and change in power measurements depending on small grain species without damaging the plant. These measurements could be used by small cereal grain breeding programs as proxy traits to determine lodging susceptibility without destructively testing or waiting for storm events, thus saving time and resources. [ABSTRACT FROM AUTHOR]

Summary: Aquatic bladderworts (Utricularia gibba and U. australis) capture zooplankton in mechanically triggered underwater traps. With characteristic dimensions less than 1 mm, the trapping structures are among the smallest known to capture prey by suction, a mechanism that is not effective in the creeping‐flow regime where viscous forces prevent the generation of fast and energy‐efficient suction flows.To understand what makes suction feeding possible on the small scale of bladderwort traps, we characterised their suction flows experimentally (using particle image velocimetry) and mathematically (using computational fluid dynamics and analytical mathematical models).We show that bladderwort traps avoid the adverse effects of creeping flow by generating strong, fast‐onset suction pressures. Our findings suggest that traps use three morphological adaptations: the trap walls' fast release of elastic energy ensures strong and constant suction pressure; the trap door's fast opening ensures effectively instantaneous onset of suction; the short channel leading into the trap ensures undeveloped flow, which maintains a wide effective channel diameter.Bladderwort traps generate much stronger suction flows than larval fish with similar gape sizes because of the traps' considerably stronger suction pressures. However, bladderworts' ability to generate strong suction flows comes at considerable energetic expense. [ABSTRACT FROM AUTHOR]

Like alternative splicing, leaky ribosomal scanning (LRS), which occurs at suboptimal translational initiation codons, increases the physiological flexibility of the genome by allowing alternative translation. Comprehensive analysis of 22 208 human mRNAs indicates that, although the most important positions relative to the first nucleotide of the initiation codon, -3 and +4, are usually such that support initiation (A-3 = 42%, G-3 = 36% and G+4 = 47%), only 37.4% of the genes adhere to the purine (R)-3/G+4 rule at both positions simultaneously, suggesting that LRS may occur in some of the remaining (62.6%) genes. Moreover, 12.5% of the genes lack both R-3 and G+4, potentially leading to sLRS. Compared with 11 genes known to undergo LRS, 10 genes with experimental evidence for high fidelity A+1T+2G+3 initiation codons adhered much more strongly to the R-3/G+4 rule. Among the intron-less histone genes, only the H3 genes adhere to the R-3/G+4 rule, while the H1, H2A, H2B and H4 genes usually lack either R-3 or G+4. To address in vivo the significance of the previously described LRS of H4 mRNAs, which results in alternative translation of the osteogenic growth peptide, transgenic mice were engineered that ubiquitously and constitutively express a mutant H4 mRNA with an A+1T+1 mutation. These transgenic mice, in particular the females, have a high bone mass phenotype, attributable to increased bone formation. These data suggest that many genes may fulfill cryptic functions by LRS.

Summary: Transitions from animal to wind pollination have occurred repeatedly in flowering plants, driven by structural and biomechanical modifications to flowers. But the initial changes promoting wind pollination are poorly understood, especially those required to release pollen into airflows – the critical first stage of wind pollination.Using a wind tunnel, we performed a comparative study of pollen release biomechanics in 36 species of animal‐ and wind‐pollinated Thalictrum. We quantified pollination syndromes and stamen natural frequency (fn), a key vibration parameter, to determine if floral traits reliably predicted pollen release probability. We then investigated if pollen release was caused by wind‐induced resonance vibration of stamens.We detected wind‐induced stamen resonance in 91% of species and a strong effect of stamen acceleration on pollen release, inversely driven by fn. However, unlike fn, pollination syndromes did not reliably predict the probability of pollen release among species.Our results directly link fn to the capacity of stamens to release pollen by wind and suggest that structural mechanisms reducing fn are likely to be important for initiating transitions from animal to wind pollination. Our inability to predict the probability of pollen release based on pollination syndromes suggests diverse phenotypic trajectories from animal to wind pollination. [ABSTRACT FROM AUTHOR]

Plants are dynamic. They adjust their shape for feeding, defence, and reproduction. Such plant movements are critical for their survival. We present selected examples covering a range of movements from single cell to tissue level and over a range of time scales. We focus on reversible turgor-driven shape changes. Recent insights into the mechanisms of stomata, bladderwort, the waterwheel, and the Venus flytrap are presented. The underlying physical principles (turgor, osmosis, membrane permeability, wall stress, snap buckling, and elastic instability) are highlighted, and advances in our understanding of these processes are summarized. [ABSTRACT FROM AUTHOR]

Wind pollination has evolved from insect pollination in numerous angiosperm lineages and is associated with a characteristic syndrome of morphological traits. The traits initiating transitions to wind pollination and the ecological drivers involved are poorly understood. Here, we examine this problem in Thalictrum pubescens, an ambophilous (insect and wind pollination) species that probably represents a transitional state in the evolution of wind pollination. We investigated wind-induced pollen release by forced harmonic motion by measuring stamen natural frequency ( fn), a key vibration parameter, and its variability among nine populations. We assessed the repeatability of fn over consecutive growing seasons, the effect of this parameter on pollen release in a wind tunnel, and male reproductive success in the field using experimental manipulation of the presence or absence of pollinators. We found significant differences among populations and high repeatability within genotypes in fn. The wind tunnel assay revealed a strong negative correlation between fn and pollen release. Siring success was greatest for plants with lower fn when pollinators were absent, but this advantage diminished when pollinators were present. Our biomechanical analysis of the wind-flower interface has identified fn as a key trait for understanding early stages in the transition from insect to wind pollination. [ABSTRACT FROM AUTHOR]

Insects fall prey to the Venus flytrap (Dionaea muscipula) when they touch the sensory hairs located on the flytrap lobes, causing sudden trap closure. The mechanical stimulus imparted by the touch produces an electrical response in the sensory cells of the trigger hair. These cells are found in a constriction near the hair base, where a notch appears around the hair’s periphery. There are mechanosensitive ion channels (MSCs) in the sensory cells that open due to a change in membrane tension; however, the kinematics behind this process is unclear. In this study, we investigate how the stimulus acts on the sensory cells by building a multi-scale hair model, using morphometric data obtained from μ-CT scans. We simulated a single-touch stimulus and evaluated the resulting cell wall stretch. Interestingly, the model showed that high stretch values are diverted away from the notch periphery and, instead, localized in the interior regions of the cell wall. We repeated our simulations for different cell shape variants to elucidate how the morphology influences the location of these high-stretch regions. Our results suggest that there is likely a higher mechanotransduction activity in these ’hotspots’, which may provide new insights into the arrangement and functioning of MSCs in the flytrap.

Individual plant cells are the building blocks for all plantae and artificially constructed plant biomaterials, like biocomposites. Secondary cell walls (SCWs) are a key component for mediating mechanical strength and stiffness in both living vascular plants and biocomposite materials. In this paper, we study the structure and biomechanics of cultured plant cells during the cellular developmental stages associated with SCW formation. We use a model culture system that induces transdifferentiation of Arabidopsis thaliana cells to xylem vessel elements, upon treatment with dexamethasone (DEX). We group the transdifferentiation process into three distinct stages, based on morphological observations of the cell walls. The first stage includes cells with only a primary cell wall (PCW), the second covers cells that have formed a SCW, and the third stage includes cells with a ruptured tonoplast and partially or fully degraded PCW. We adopt a multi-scale approach to study the mechanical properties of cells in these three stages. We perform large-scale indentations with a micro-compression system in three different osmotic conditions. Atomic force microscopy (AFM) nanoscale indentations in water allow us to isolate the cell wall response. We propose a spring-based model to deconvolve the competing stiffness contributions from turgor pressure, PCW, SCW and cytoplasm in the stiffness of differentiating cells. Prior to triggering differentiation, cells in hypotonic pressure conditions are significantly stiffer than cells in isotonic or hypertonic conditions, highlighting the dominant role of turgor pressure. Plasmolyzed cells with a SCW reach similar levels of stiffness as cells with maximum turgor pressure. The stiffness of the PCW in all of these conditions is lower than the stiffness of the fully-formed SCW. Our results provide the first experimental characterization of the mechanics of SCW formation at single cell level.

Stem lodging is the bending or breakage of stems in the wind that result in negative economic impacts to producers and processors of small grain crops. To address this issue, plant breeders attempt to quantify lodging using proxy traits such as stem structure and biomechanics. Stem lodging is a function of both stem strength and elasticity. In this paper, we explore the biomechanics of stems approaching the lodging, or permanent bending, condition. Oat, wheat, and two types of barley varying in lodging resistance were exposed to standard growing conditions over the course of a season. Their capability of returning from a bent to unbent state was characterized using a push force meter that measured resistant force and displacement over time. Changes in stem energy and power were then calculated using displacement and force measurements. Lodging susceptibility could be differentiated by stem strength, displacement and change in power measurements depending on small grain species without damaging the plant. These measurements could be used by small cereal grain breeding programs as proxy traits to determine lodging susceptibility without destructively testing or waiting for storm events, thus saving time and resources.

Resurrection plants have fascinated scientists since centuries as they can fully recover from cellular water contents below 10%, concomitantly showing remarkable leaf folding motions. While physiological adaptations have been meticulously investigated, the understanding of structural and mechanical adaptations of this phenomenon is scarce. Using imaging and bending techniques during dehydration-rehydration experiments, morphological, anatomical, and biomechanical properties of desiccation-tolerant Ramonda myconi are examined, and selected structural adaptations are compared to those of homoiohydrous Monophyllaea horsfieldii (both Gesneriaceae). At low water availability, intact and cut-off R. myconi leaves undergo considerable morphological alterations, which are fully and repeatedly reversible upon rehydration. Furthermore, their petioles show a triphasic mechanical behavior having a turgor-based structural stability at high (Phase 1), a flexible mechanically state at intermediate (Phase 2) and a material-based stability at low water contents (Phase 3). Lastly, manipulation experiments with cut-off plant parts revealed that both the shape alterations of individual structures, as well as, the general leaf kinematics largely rely on passive swelling and shrinking processes. Taken together, R. myconi possesses structural and mechanical adaptations to desiccation (in addition to physiological adaptations), which may mainly be passively driven by its water status influenced by the water fluctuations in its surroundings.

Resurrection plants have fascinated scientists since centuries as they can fully recover from cellular water contents below 10%, concomitantly showing remarkable leaf folding motions. While physiological adaptations have been meticulously investigated, the understanding of structural and mechanical adaptations of this phenomenon is scarce. Using imaging and bending techniques during dehydration-rehydration experiments, morphological, anatomical, and biomechanical properties of desiccation-tolerant Ramonda myconi are examined, and selected structural adaptations are compared to those of homoiohydrous Monophyllaea horsfieldii (both Gesneriaceae). At low water availability, intact and cut-off R. myconi leaves undergo considerable morphological alterations, which are fully and repeatedly reversible upon rehydration. Furthermore, their petioles show a triphasic mechanical behavior having a turgor-based structural stability at high (Phase 1), a flexible mechanically state at intermediate (Phase 2) and a material-based stability at low water contents (Phase 3). Lastly, manipulation experiments with cut-off plant parts revealed that both the shape alterations of individual structures, as well as, the general leaf kinematics largely rely on passive swelling and shrinking processes. Taken together, R. myconi possesses structural and mechanical adaptations to desiccation (in addition to physiological adaptations), which may mainly be passively driven by its water status influenced by the water fluctuations in its surroundings. [ABSTRACT FROM AUTHOR]

Agar is widely used in studies of root growth since it can be mixed at different concentrations to impact mechanical impedance. At high concentrations (1.2–1.5%), growth of Arabidopsis roots has been found to be wavy, but little research has explored this behavior based on a quantitative understanding of mechanical behavior. To this end, agar media with concentration ranging from 0.5% to 1.2% were prepared to produce gradient resistance during root penetration, and Young's moduli and penetrometer resistance were tested. Arabidopsis roots were then cultivated in these agar media with gradient stiffness. The result showed that Young's modulus increased linearly with the increase of concentration of agar media. For Arabidopsis primary roots, it was preferred to develop a helical pattern in agar media with concentration from 0.5% to 1.0%. As stiffness of agar increased, the percentage of helical roots and helix diameters in each agar medium declined; root lengths and auxin distributions showed variety. We demonstrate that the size of helical deformation decreases with agar stiffness as expected by theoretical analysis based on a combination of growth-induced mechanical buckling. In conclusion, the resistance from agar media impacts the properties of root helix, and helical roots growth is driven by growth force. Growth force and external mechanical forces contribute to root phenotypes in Arabidopsis . [ABSTRACT FROM AUTHOR]

Plants are able to sense and respond to minute tilt from the vertical direction of the gravity, which is key to maintain their upright posture during development. However, gravisensing in plants relies on a peculiar sensor made of microsize starch-filled grains (statoliths) that sediment and form tiny granular piles at the bottom of the cell. How such a sensor can detect inclination is unclear, as granular materials like sand are known to display flow threshold and finite avalanche angle due to friction and interparticle jamming. Here, we address this issue by combining direct visualization of statolith avalanches in plant cells and experiments in biomimetic cells made of microfluidic cavities filled with a suspension of heavy Brownian particles. We show that, despite their granular nature, statoliths move and respond to the weakest angle, as a liquid clinometer would do. Comparison between the biological and biomimetic systems reveals that this liquid-like behavior comes from the cell activity, which agitates statoliths with an apparent temperature one order of magnitude larger than actual temperature. Our results shed light on the key role of active fluctuations of statoliths for explaining the remarkable sensitivity of plants to inclination. Our study also provides support to a recent scenario of gravity perception in plants, by bridging the active granular rheology of statoliths at the microscopic level to the macroscopic gravitropic response of the plant. [ABSTRACT FROM AUTHOR]

Plants constantly undergo external mechanical loads such as wind or touch and respond to these stimuli by acclimating their growth processes. A fascinating feature of this mechanicalinduced growth response is that it can occur rapidly and at long distance from the initial site of stimulation, suggesting the existence of a fast signal that propagates across the whole plant. The nature and origin of the signal is still not understood, but it has been recently suggested that it could be purely mechanical and originate from the coupling between the local deformation of the tissues (bending) and the water pressure in the plant vascular system. Here, we address the physical origin of this hydromechanical coupling using a biomimetic strategy. We designed soft artificial branches perforated with longitudinal liquid-filled channels that mimic the basic features of natural stems and branches. In response to bending, a strong overpressure is generated in the channels that varies quadratically with the bending curvature. A model based on a mechanism analogous to the ovalization of hollow tubes enables us to predict quantitatively this nonlinear poroelastic response and identify the key physical parameters that control the generation of the pressure pulse. Further experiments conducted on natural tree branches reveal the same phenomenology. Once rescaled by the model prediction, both the biomimetic and natural branches fall on the same master curve, enlightening the universality of our poroelastic mechanism for the generation of hydraulic signals in plants. [ABSTRACT FROM AUTHOR]

Trees are the largest living beings that have ever lived on Earth, with more than one hundred meters high, masses of several thousand tons and ages of several thousand years. In order to achieve this, among other functional conditions, trees have to cope with all kinds of mechanical loads. Therefore, to meet these structural requirements, trees have wood, an exceptional material from a mechanical point of view, and offer a number of design strategies ranging from ensuring structural stability to provide adequate safety factor using a minimum amount of material, with consequent metabolic efficiency. In that sense, this paper presents the most important strategies of growth, anatomy and morphology that use the leading arborescent plants families, gymnosperms and angiosperms, highlighting the features of natural plant materials, which have a hierarchical organization, and in most cases are fiber-reinforced composite materials, materials with cellular structure, or both, such as wood. [ABSTRACT FROM AUTHOR]

Main conclusion : The order of the internodes, and their geometry and mechanical characteristics influence the capability of the Equisetum stem to vibrate, potentially stimulating spore liberation at the optimum stress setting along the stem. Equisetum hyemale L. plants represent a special example of cellular solid construction with mechanical stability achieved by a high second moment of area and relatively high resistance against local buckling. We proposed the hypothesis that the order of E. hyemale L. stem internodes, their geometry and mechanical characteristics influence the capability of the stem to vibrate, stimulating spore liberation at the minimum stress setting value along the stem. An analysis of apex vibration was done based on videos presenting the behavior of an Equisetum clump filmed in a wind tunnel and also as a result of excitation by bending the stem by 20°. We compared these data with the vibrations of stems of the same size but deprived of the three topmost internodes. Also, we created a finite element model (FEM), upon which we have based the 'natural' stem vibration as a copy of the real object, 'random' with reshuffled internodes and 'uniform', created as one tube with the characters averaged from all internodes. The natural internode arrangement influences the frequency and amplitude of the apex vibration, maintaining an equal stress distribution in the stem, which may influence the capability for efficient spore spreading. [ABSTRACT FROM AUTHOR]

Energetic trade-offs are particularly pertinent to bio-ballistic systems which impart energy to projectiles exclusively during launch. We investigated such trade-offs in the spring-propelled seeds of Loropetalum chinense , Hamamelis virginiana and Fortunearia sinensis . Using similar seed-shooting mechanisms, fruits of these confamilial plants (Hamamelidaceae) span an order of magnitude in spring and seed mass. We expected that as seed mass increases, launch speed decreases. Instead, launch speed was relatively constant regardless of seed mass. We tested if fruits shoot larger seeds by storing more elastic potential energy (PE). Spring mass and PE increased as seed mass increased (in order of increasing seed mass: L. chinense , H. virginiana , F. sinensis ). As seed mass to spring mass ratio increased (ratios: H. virginiana = 0.50, F. sinensis = 0.65, L. chinense = 0.84), mass-specific PE storage increased. The conversion efficiency of PE to seed kinetic energy (KE) decreased with increasing fruit mass. Therefore, similar launch speeds across scales occurred because (i) larger fruits stored more PE and (ii) smaller fruits had higher mass-specific PE storage and improved PE to KE conversion. By examining integrated spring and projectile mechanics in our focal species, we revealed diverse, energetic scaling strategies relevant to spring-propelled systems navigating energetic trade-offs.

This article is a Commentary on Clair et al.,221: 209–217. [ABSTRACT FROM AUTHOR]

Plants must meet mechanical as well as physiological and reproductive requirements for survival. Management of internal and external stresses is achieved through their unique hierarchical architecture. Stiffness is determined by a combination of morphological (geometrical) and compositional variables that vary across multiple length scales ranging from the whole plant to organ, tissue, cell and cell wall levels. These parameters include, among others, organ diameter, tissue organization, cell size, density and turgor pressure, and the thickness and composition of cell walls. These structural parameters and their consequences on plant stiffness are reviewed in the context of work on stems of the genetic reference plant Arabidopsis thaliana (Arabidopsis), and the suitability of Arabidopsis as a model system for consistent investigation of factors controlling plant stiffness is put forward. Moving beyond Arabidopsis, the presence of morphological parameters causing stiffness gradients across length-scales leads to beneficial emergent properties such as increased load-bearing capacity and reversible actuation. Tailoring of plant stiffness for old and new purposes in agriculture and forestry can be achieved through bioengineering based on the knowledge of the morphological and compositional parameters of plant stiffness in combination with gene identification through the use of genetics. [ABSTRACT FROM AUTHOR]

• Background and Aims Plant design refers to the construction of the plant body or its constituent parts in terms of form and function. Although neighbourhood structure is recognized as a factor that limits plant survival and species coexistence, its relative importance in plant design is not well understood. We conducted field research to analyse how the surrounding environment of neighbourhood structure and related effects on light availability are associated with changes in plant design in two understorey plants (Palicourea padifolia and Psychotria elata) within two successional stages of a cloud forest in Costa Rica. • Methods Features of plant neighbourhood physical structure and light availability, estimated using hemispherical photographs, were used as variables that reflect the surrounding environment. Measures of plant biomechanics, allometry, branching and plant slenderness were used as functional plant attributes that reflect plant design. We propose a framework using a partial least squares path model and used it to test this association. • Key Results The multidimensional response of plant design of these species suggests that decreases in the heightbased factor of safety and increases in mechanical load and developmental stability are influenced by increases in maximum height of neighbours and a distance-dependence interference index more than neighbourhood plant density or neighbour aggregation. Changes in plant branching and slenderness are associated positively with light availability and negatively with canopy cover. • Conclusions Although it has been proposed that plant design varies according to plant density and light availability, we found that neighbour size and distance-dependence interference are associated with changes in biomechanics, allometry and branching, and they must be considered as key factors that contribute to the adaptation and coexistence of these plants in this highly diverse forest community. [ABSTRACT FROM AUTHOR]

Several species of the plant family Martyniaceae are characterised by unique lignified capsules with hook-shaped extensions that interlock with hooves and ankles of large mammals to disperse the seeds. The arrangement of fruit endocarp fibre tissues is exceptional and intriguing among plants. Structure–function-relationships of these slender, curved, but mechanically highly stressed fruit extensions are of particular interest that may inspire advanced biomimetic composite materials. In the present study, we analyse mechanical properties and fracture behaviour of the hook-shaped fruit extensions under different load conditions. The results are correlated with calculated stress distributions, the specific cell wall structure, and chemical composition, providing a detailed interpretation of the complex fruit tissue microstructure. At the cell wall level, both a large microfibril angle and greater strain rates resulted in Young’s moduli of 4–9 GPa, leading to structural plasticity. Longitudinally arranged fibre bundles contribute to a great tensile strength. At the tissue level, transversely oriented fibres absorb radial stresses upon bending, whereas cells encompass and pervade longitudinal fibre bundles, thus, stabilise them against buckling. During bending and torsion, microcracks between axial fibre bundles are probably spanned analogous to a circular anchor. Our study fathoms a highly specialized plant structure, substantiating former assumptions about epizoochory as dispersal mode. While the increased flexibility allows for proper attachment of fruits during dynamical locomotion, the high strength and stability prevent a premature failure due to heavy loads exerted by the animal. [ABSTRACT FROM AUTHOR]

Main conclusion: Trichomes are involved in petiole movement and likely function as a part of the plant biomechanical system serving as an additional reservoir of hydrostatic pressure. The large, non-glandular trichomes on Cucurbita petioles occur across collenchyma strands. Time-lapse imaging was used to study the leaf reorientation of Cucurbita maxima 'Bambino' plants placed in horizontal position. The experiment comprised four variants of the large non-glandular petiole trichomes: (1) intact, (2) mechanically removed, (3) dehydrated, and (4) intact but with longitudinally injured petioles. Isolated strands of collenchyma with intact epidermis or epidermis mechanically removed from the abaxial and adaxial sides of the petiole were subjected to breaking test. The stiffness of the non-isolated tissue with intact epidermis was measured using the micro-indentation method. Petioles without trichomes did not exhibit tropic response, and the dehydration of trichomes slowed and prevented complete leaf reorientation. Isolated strands of collenchyma showed no correlation between strength values and position on the petiole. However, strands of collenchyma with epidermis exhibited a significantly greater strength regardless of their position on the petiole. The indentation test showed that non-isolated collenchyma is stiffer on the abaxial side of the petiole. Trichomes from the abaxial side of the petiole were larger at their base. The application of the 'tensile triangles method' revealed that these trichomes had a biomechanically optimized shape in comparison to the adaxial side. We conclude that trichomes can be involved in plant biomechanical system and serve as an additional reservoir of hydrostatic pressure that is necessary for maintaining petioles in the prestressed state. [ABSTRACT FROM AUTHOR]

Experimental studies show that plant root morphologies can vary widely from straight gravity-aligned primary roots to fractal-like root architectures. However, the opaqueness of soil makes it difficult to observe how environmental factors modulate these patterns. Here, we combine a transparent hydrogel growth medium with a custom built 3D laser scanner to directly image the morphology of Medicago truncatula primary roots. In our experiments, root growth is obstructed by an inclined plane in the growth medium. As the tilt of this rigid barrier is varied, we find Medicago transitions between randomly directed root coiling, sinusoidal root waving, and normal gravity-aligned morphologies. Although these root phenotypes appear morphologically distinct, our analysis demonstrates the divisions are less well defined, and instead, can be viewed as a 2D biased random walk that seeks the path of steepest decent along the inclined plane. Features of this growth response are remarkably similar to the widely known run-and-tumble chemotactic behavior of Escherichia coli bacteria, where biased random walks are used as optimal strategies for nutrient uptake. [ABSTRACT FROM AUTHOR]

Organisms living in highly variable environments have to display integrated strategies to deal with both systematic and random variation occurring at different temporal and spatial scales. Two predictions were tested by analysing geographic-scale patterns of seed size and seed retention (serotiny) in Chorizanthe rigida, a strict winter desert annual that delays seed dispersal and releases propagules after rainfall events: (i) adaptation to systematic environmental cues occurs by means of changes in morphology, and (ii) within-individual variation in seed size allows a differential response to rainfall cues: while some seeds germinate rapidly others are retained for future rainfall events., We quantified morphological variation and performed germination experiments on C. rigida propagules (involucres + achenes) from six populations distributed throughout the Mojave and Sonoran deserts covering: (i) a systematic, west-to-east, winter-to-bi-seasonal (summer and winter) precipitation gradient and (ii) a winter-rain unpredictability gradient inferred from long-term climatic data., The propagule retention structure (i.e. base area of the pedicel) of C. rigida individuals experiencing bi-seasonal rainfall is double the size of those that have evolved under a strict winter rainfall regime, showing that populations living in bi-seasonal environments have higher seed retention, which allows them to avoid releasing seeds to a summer rainfall cue., Within-individual variance of propagule size varied significantly between populations and was correlated with winter rainfall variability in each site., Germination varied as a function of propagule size; smaller seeds germinated more readily than larger seeds. Increased variability in propagule size might result in a more variable germination response., Under common experimental conditions germination varied significantly among sites and was negatively correlated with mean winter effective precipitation, suggesting that propagules from populations in drier sites have lower germination moisture thresholds., Synthesis. C. rigida propagules have larger bases in deserts with biseasonal rainfall, which allows them to avoid seed release during summer rainfall cues, and display within-individual seed variance associated to rainfall unpredictability, a trait often interpreted as a bet-hedging strategy. Our study provides empirical evidence of an integrated strategy that allows to cope with both random and systematic rainfall variation. [ABSTRACT FROM AUTHOR]