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Animacy is an important framework through which humans view and categorize the world, but many objects do not easily fit within this scale. Plants are unique because they are very familiar to humans, yet the features and traits relevant for placement within the animacy scale are generally poorly understood by the public. Animacy occurs at three levels, with the inherent attributes of the object (biology), how they are perceived (cognition), and how they are expressed in languages (linguistics). Animacy is dependent on qualification and perception as alive, mobile, and intentional. In the absence of visible movement, classification is dependent on featural attributes indicating mobility or placement in a group recognized as animate (animalness). Plants have complicated bodies whose forms and structures are frequently clear representations of their life history and function (plantness), more than many animals, yet these signs of movement and activity are rarely recognized. The animacy scale may be more closely based on human similarity (humanness) with humans as the peak of life, mobility, and intentionality. As humans, we can have an anthropocentric viewpoint, rendering plants as scenery or utility, or use anthropomorphic interaction to better understand and recognize the dynamic lives of plants. The goal of this review is to compare the current evidence on the placement of plants within animacy and adjacent scales with the biology and habit of land plants, to better understand human perception and behaviour, and work with this process to educate and inform people about the complex lives of plants. [ABSTRACT FROM AUTHOR]

Finding a suitable support is a key process in the life history of climbing plants. Those that find a suitable support have greater performance and fitness than those that remain prostrate. Numerous studies on climbing plant behavior have elucidated the mechanistic details of support-searching and attachment. Far fewer studies have addressed the ecological significance of support-searching behavior and the factors that affect it. Among these, the diameter of supports influences their suitability. When the support diameter increases beyond some point, climbing plants are unable to maintain tensional forces and therefore lose attachment to the trellis. Here, we further investigate this issue by placing pea plants (Pisum sativum L.) in the situation of choosing between supports of different diameters while their movement was recorded by means of a three-dimensional motion analysis system. The results indicate that the way pea plants move can vary depending on whether they are presented with one or two potential supports. Furthermore, when presented with a choice between thin and thick supports, the plants showed a distinct preference for the former than the latter. The present findings shed further light on how climbing plants make decisions regarding support-searching and provide evidence that plants adopt one of several alternative plastic responses in a way that optimally corresponds to environmental scenarios. [ABSTRACT FROM AUTHOR]

Climbing plants require an external support to grow vertically and enhance light acquisition. Climbers that find a suitable support demonstrate greater performance and fitness than those that remain prostrate. Support search is characterized by oscillatory movements (i.e., circumnutation), in which plants rotate around a central axis during their growth. Numerous studies have elucidated the mechanistic details of circumnutation, but how this phenomenon is controlled during support searching remains unclear. To fill this gap, here we tested whether simulation-based machine learning methods can capture differences in movement patterns nested in actual kinematical data. We compared machine learning classifiers with the aim of generating models that learn to discriminate between circumnutation patterns related to the presence/absence of a support in the environment. Results indicate that there is a difference in the pattern of circumnutation, depending on the presence of a support, that can be learned and classified rather accurately. We also identify distinctive kinematic features at the level of the junction underneath the tendrils that seems to be a superior indicator for discerning the presence/absence of the support by the plant. Overall, machine learning approaches appear to be powerful tools for understanding the movement of plants. [ABSTRACT FROM AUTHOR]

Abstract Background In recent years, there has been an increase of interest in plant behaviour as represented by growth-driven responses. These are generally classified into nastic (internally driven) and tropic (environmentally driven) movements. Nastic movements include circumnutations, a circular movement of plant organs commonly associated with search and exploration, while tropisms refer to the directed growth of plant organs toward or away from environmental stimuli, such as light and gravity. Tracking these movements is therefore fundamental for the study of plant behaviour. Convolutional neural networks, as used for human and animal pose estimation, offer an interesting avenue for plant tracking. Here we adopted the Social LEAP Estimates Animal Poses (SLEAP) framework for plant tracking. We evaluated it on time-lapse videos of cases spanning a variety of parameters, such as: (i) organ types and imaging angles (e.g., top-view crown leaves vs. side-view shoots and roots), (ii) lighting conditions (full spectrum vs. IR), (iii) plant morphologies and scales (100 μm-scale Arabidopsis seedlings vs. cm-scale sunflowers and beans), and (iv) movement types (circumnutations, tropisms and twining). Results Overall, we found SLEAP to be accurate in tracking side views of shoots and roots, requiring only a low number of user-labelled frames for training. Top views of plant crowns made up of multiple leaves were found to be more challenging, due to the changing 2D morphology of leaves, and the occlusions of overlapping leaves. This required a larger number of labelled frames, and the choice of labelling “skeleton” had great impact on prediction accuracy, i.e., a more complex skeleton with fewer individuals (tracking individual plants) provided better results than a simpler skeleton with more individuals (tracking individual leaves). Conclusions In all, these results suggest SLEAP is a robust and versatile tool for high-throughput automated tracking of plants, presenting a new avenue for research focusing on plant dynamics.

Vertical leaf angles and their variation through time are directly related to several ecophysiological processes and properties. However, there is no efficient method for tracking leaf angles of plant canopies under field conditions.Here, we present AngleCam, a deep learning‐based approach to predict leaf angle distributions from horizontal photographs acquired with low‐cost timelapse cameras. AngleCam is based on pattern recognition with convolutional neural networks and trained with leaf angle distributions obtained from visual interpretation of more than 2500 plant photographs across different species and scene conditions.Leaf angle predictions were evaluated over a wide range of species and scene conditions using independent samples from visual interpretation (R2 = 0.84) and compared to leaf angle estimates obtained from terrestrial laser scanning (R2 = 0.75). AngleCam was tested for the long‐term monitoring of leaf angles for two broadleaf tree species in a temperate forest. The plausibility of the predicted leaf angle time series was underlined by its close relationship with environmental variables related to transpiration.The evaluations confirm that AngleCam is a robust and efficient method to track leaf angles under field conditions. The output of AngleCam is compatible with a range of applications, including functional‐structural plant modelling, Earth system modelling or radiative transfer modelling of plant canopies. AngleCam may also be used to predict leaf angle distributions for existing data, for instance from PhenoCam networks citizen science projects. [ABSTRACT FROM AUTHOR]

Pine cones show functionally highly resilient, hygroscopically actuated opening and closing movements, which are repeatable and function even in millions of years old, coalified cones. Although the functional morphology and biomechanics behind the individual seed scale motions are well understood, the initial opening of the cone, which is often accompanied by an audible cracking noise, is not. We therefore investigated the initial opening events of mature fresh cones of Scots pine (Pinus sylvestris) and their subsequent motion patterns. Using high-speed and time lapse videography, 3D digital image correlation techniques, force measurements, thermographic and chemical-rheological resin analyses, we are able to draw a holistic picture of the initial opening process involving the rupture of resin seals and very fast seed scale motion in the millisecond regime. The rapid cone opening was not accompanied by immediate seed release in our experiments and, therefore, cannot be assigned to ballistochory. As the involved passive hydraulic-elastic processes in cracking are very fine-tuned, we hypothesize that they are under tight mechanical-structural control to ensure an ecologically optimized seed release upon environmental conditions suitable for wind dispersal. In this context, we propose an interplay of humidity and temperature to be the external "drivers" for the initial cone opening, in which resin works as a crucial chemical-mechanical latch system. [ABSTRACT FROM AUTHOR]

Previous studies on the kinematics of pea plants' ascent and attach behavior have demonstrated that the signature of their movement varies depending on the kind of support. So far, these studies have been confined to artificial supports (e.g. wooden sticks). Little is known regarding the conditions under which pea plants could rely on biological supports (e.g. neighboring plants) for climbing toward the light. In this study, we capitalize on the 3D kinematic analysis of movement to ascertain whether pea plants scale their kinematics differently depending on whether they aim for artificial or biological support. Results suggest that biological support determines a smoother and more accurate behavior than that elicited by the artificial one. These results shed light on pea plants' ability to detect and classify the properties of objects and implement a movement plan attuned to the very nature of the support. We contend that such differences depend on the augmented multisensory experience elicited by the biological support.

Pine cones show functionally highly resilient, hygroscopically actuated opening and closing movements, which are repeatable and function even in millions of years old, coalified cones. Although the functional morphology and biomechanics behind the individual seed scale motions are well understood, the initial opening of the cone, which is often accompanied by an audible cracking noise, is not. We therefore investigated the initial opening events of mature fresh cones of Scots pine (Pinus sylvestris) and their subsequent motion patterns. Using high-speed and time lapse videography, 3D digital image correlation techniques, force measurements, thermographic and chemical-rheological resin analyses, we are able to draw a holistic picture of the initial opening process involving the rupture of resin seals and very fast seed scale motion in the millisecond regime. The rapid cone opening was not accompanied by immediate seed release in our experiments and, therefore, cannot be assigned to ballistochory. As the involved passive hydraulic-elastic processes in cracking are very fine-tuned, we hypothesize that they are under tight mechanical-structural control to ensure an ecologically optimized seed release upon environmental conditions suitable for wind dispersal. In this context, we propose an interplay of humidity and temperature to be the external “drivers” for the initial cone opening, in which resin works as a crucial chemical-mechanical latch system.

The evolutionary roots of carnivory in the Venus flytrap (Dionaea muscipula) stem from a defense response to plant injury caused by, e.g., herbivores. Dionaea muscipula aka. Darwin's most wonderful plant underwent extensive modification of leaves into snap-traps specialized for prey capture. Even the tiny seedlings of the Venus flytrap already produce fully functional, millimetersized traps. The trap size increases as the plant matures, enabling capture of larger prey. The movement of snap-traps is very fast (~100-300 ms) and is actuated by a combination of changes in the hydrostatic pressure of the leaf tissue with the release of prestress (embedded energy), triggering a snapthrough of the trap lobes. This instability phenomenon is facilitated by the double curvature of the trap lobes. In contrast, trap reopening is a slower process dependent on trap size and morphology, heavily reliant on turgor and/or cell growth. Once a prey item is caught, the trap reconfigures its shape, seals itself off and forms a digestive cavity allowing the plant to release an enzymatic cocktail to draw nutrition from its captive. Interestingly, a failed attempt to capture prey can come at a heavy cost: the trap can break during reopening, thus losing its functionality. In this mini-review, we provide a detailed account of morphological adaptations and biomechanical processes involved in the trap movement during D. muscipula hunting cycle, and discuss possible reasons for and consequences of trap breakage. We also provide a brief introduction to the biological aspects underlying plant motion and their evolutionary background. [ABSTRACT FROM AUTHOR]

Fast snapping in the carnivorous Venus flytrap (Dionaea muscipula) involves trap lobe bending and abrupt curvature inversion (snap‐buckling), but how do these traps reopen? Here, the trap reopening mechanics in two different D. muscipula clones, producing normal‐sized (N traps, max. ≈3 cm in length) and large traps (L traps, max. ≈4.5 cm in length) are investigated. Time‐lapse experiments reveal that both N and L traps can reopen by smooth and continuous outward lobe bending, but only L traps can undergo smooth bending followed by a much faster snap‐through of the lobes. Additionally, L traps can reopen asynchronously, with one of the lobes moving before the other. This study challenges the current consensus on trap reopening, which describes it as a slow, smooth process driven by hydraulics and cell growth and/or expansion. Based on the results gained via three‐dimensional digital image correlation (3D‐DIC), morphological and mechanical investigations, the differences in trap reopening are proposed to stem from a combination of size and slenderness of individual traps. This study elucidates trap reopening processes in the (in)famous Dionaea snap traps – unique shape‐shifting structures of great interest for plant biomechanics, functional morphology, and applications in biomimetics, i.e., soft robotics. [ABSTRACT FROM AUTHOR]

The evolutionary roots of carnivory in the Venus flytrap (Dionaea muscipula) stem from a defense response to plant injury caused by, e.g., herbivores. Dionaea muscipula aka. Darwin’s most wonderful plant underwent extensive modification of leaves into snap-traps specialized for prey capture. Even the tiny seedlings of the Venus flytrap already produce fully functional, millimeter-sized traps. The trap size increases as the plant matures, enabling capture of larger prey. The movement of snap-traps is very fast (~100–300 ms) and is actuated by a combination of changes in the hydrostatic pressure of the leaf tissue with the release of prestress (embedded energy), triggering a snap-through of the trap lobes. This instability phenomenon is facilitated by the double curvature of the trap lobes. In contrast, trap reopening is a slower process dependent on trap size and morphology, heavily reliant on turgor and/or cell growth. Once a prey item is caught, the trap reconfigures its shape, seals itself off and forms a digestive cavity allowing the plant to release an enzymatic cocktail to draw nutrition from its captive. Interestingly, a failed attempt to capture prey can come at a heavy cost: the trap can break during reopening, thus losing its functionality. In this mini-review, we provide a detailed account of morphological adaptations and biomechanical processes involved in the trap movement during D. muscipula hunting cycle, and discuss possible reasons for and consequences of trap breakage. We also provide a brief introduction to the biological aspects underlying plant motion and their evolutionary background.

Finding a suitable support is a key process in the life history of climbing plants. Those that find a suitable support have greater performance and fitness than those that remain prostrate. Numerous studies on climbing plant behavior have elucidated the mechanistic details of support-searching and attachment. Far fewer studies have addressed the ecological significance of support-searching behavior and the factors that affect it. Among these, the diameter of supports influences their suitability. When the support diameter increases beyond some point, climbing plants are unable to maintain tensional forces and therefore lose attachment to the trellis. Here, we further investigate this issue by placing pea plants (Pisum sativum L.) in the situation of choosing between supports of different diameters while their movement was recorded by means of a three-dimensional motion analysis system. The results indicate that the way pea plants move can vary depending on whether they are presented with one or two potential supports. Furthermore, when presented with a choice between thin and thick supports, the plants showed a distinct preference for the former than the latter. The present findings shed further light on how climbing plants make decisions regarding support-searching and provide evidence that plants adopt one of several alternative plastic responses in a way that optimally corresponds to environmental scenarios.

The celestial mechanics of the Sun, Moon, and Earth dominate the variations in gravitational force that all matter, live or inert, experiences on Earth. Expressed as gravimetric tides, these variations are pervasive and have forever been part of the physical ecology with which organisms evolved. Here, we first offer a brief review of previously proposed explanations that gravimetric tides constitute a tangible and potent force shaping the rhythmic activities of organisms. Through meta-analysis, we then interrogate data from three study cases and show the close association between the omnipresent gravimetric tides and cyclic activity. As exemplified by free-running cyclic locomotor activity in isopods, reproductive effort in coral, and modulation of growth in seedlings, biological rhythms coincide with temporal patterns of the local gravimetric tide. These data reveal that, in the presumed absence of rhythmic cues such as light and temperature, local gravimetric tide is sufficient to entrain cyclic behaviour. The present evidence thus questions the phenomenological significance of so-called free-run experiments. [ABSTRACT FROM AUTHOR]

Background: In recent years, there has been an increase of interest in plant behaviour as represented by growth-driven responses. These are generally classified into nastic (internally driven) and tropic (environmentally driven) movements. Nastic movements include circumnutations, a circular movement of plant organs commonly associated with search and exploration, while tropisms refer to the directed growth of plant organs toward or away from environmental stimuli, such as light and gravity. Tracking these movements is therefore fundamental for the study of plant behaviour. Convolutional neural networks, as used for human and animal pose estimation, offer an interesting avenue for plant tracking. Here we adopted the Social LEAP Estimates Animal Poses (SLEAP) framework for plant tracking. We evaluated it on time-lapse videos of cases spanning a variety of parameters, such as: (i) organ types and imaging angles (e.g., top-view crown leaves vs. side-view shoots and roots), (ii) lighting conditions (full spectrum vs. IR), (iii) plant morphologies and scales (100 μm-scale Arabidopsis seedlings vs. cm-scale sunflowers and beans), and (iv) movement types (circumnutations, tropisms and twining). Results: Overall, we found SLEAP to be accurate in tracking side views of shoots and roots, requiring only a low number of user-labelled frames for training. Top views of plant crowns made up of multiple leaves were found to be more challenging, due to the changing 2D morphology of leaves, and the occlusions of overlapping leaves. This required a larger number of labelled frames, and the choice of labelling "skeleton" had great impact on prediction accuracy, i.e., a more complex skeleton with fewer individuals (tracking individual plants) provided better results than a simpler skeleton with more individuals (tracking individual leaves). Conclusions: In all, these results suggest SLEAP is a robust and versatile tool for high-throughput automated tracking of plants, presenting a new avenue for research focusing on plant dynamics. [ABSTRACT FROM AUTHOR]

Climbing plants require an external support to grow vertically and enhance light acquisition. Climbers that find a suitable support demonstrate greater performance and fitness than those that remain prostrate. Support search is characterized by oscillatory movements (i.e., circumnutation), in which plants rotate around a central axis during their growth. Numerous studies have elucidated the mechanistic details of circumnutation, but how this phenomenon is controlled during support searching remains unclear. To fill this gap, here we tested whether simulation-based machine learning methods can capture differences in movement patterns nested in actual kinematical data. We compared machine learning classifiers with the aim of generating models that learn to discriminate between circumnutation patterns related to the presence/absence of a support in the environment. Results indicate that there is a difference in the pattern of circumnutation, depending on the presence of a support, that can be learned and classified rather accurately. We also identify distinctive kinematic features at the level of the junction underneath the tendrils that seems to be a superior indicator for discerning the presence/absence of the support by the plant. Overall, machine learning approaches appear to be powerful tools for understanding the movement of plants.

Abstract Diurnal branch movements in woody plants have only recently been described in detail. While previously only vegetative and reproductive structures have been known to move on hourly timescales, imaging technologies such as terrestrial laser scanning and near‐surface repeat digital photography provide a means of remotely monitoring plant movements at high enough temporal and spatial resolution to capture rhythmic movements of woody material. Virtually, nothing is known about the range of species and ecosystems in which woody movements might occur or what causes these movements. We report that diurnal woody branch movements occur in a number of tree and shrub species across a broad range of abiotic conditions. We examined detailed branch movements in one species, creosote (Larrea tridentata), and found that branch movements were highly correlated with humidity, air temperature, vapor pressure deficit, and stem water potential: all factors related to plant water status. We also found that live and dead branch movements were distinct in the timing of their movements and in the abiotic conditions with which they were most correlated. Changes in dead branch position were most correlated with humidity, with these movements consistently lagging 1–2 h behind changes in humidity. Live branch movements were also highly correlated with vapor pressure deficit and humidity but went from lagging 1–2 h behind changes in these abiotic conditions in summer to being nearly in sync in winter. We believe that this is the first study that (1) documents diurnal branch movements in creosote, (2) differentiates between the movements of live and dead branches, and (3) relates environmental data to these movements. We hope these findings encourage other researchers to more closely examine imagery from their sites for evidence of branch movements, which may provide deeper insights into water and solute movements in plants and physiological responses to water stress.

Diurnal branch movements in woody plants have only recently been described in detail. While previously only vegetative and reproductive structures have been known to move on hourly timescales, imaging technologies such as terrestrial laser scanning and near‐surface repeat digital photography provide a means of remotely monitoring plant movements at high enough temporal and spatial resolution to capture rhythmic movements of woody material. Virtually, nothing is known about the range of species and ecosystems in which woody movements might occur or what causes these movements. We report that diurnal woody branch movements occur in a number of tree and shrub species across a broad range of abiotic conditions. We examined detailed branch movements in one species, creosote (Larrea tridentata), and found that branch movements were highly correlated with humidity, air temperature, vapor pressure deficit, and stem water potential: all factors related to plant water status. We also found that live and dead branch movements were distinct in the timing of their movements and in the abiotic conditions with which they were most correlated. Changes in dead branch position were most correlated with humidity, with these movements consistently lagging 1–2 h behind changes in humidity. Live branch movements were also highly correlated with vapor pressure deficit and humidity but went from lagging 1–2 h behind changes in these abiotic conditions in summer to being nearly in sync in winter. We believe that this is the first study that (1) documents diurnal branch movements in creosote, (2) differentiates between the movements of live and dead branches, and (3) relates environmental data to these movements. We hope these findings encourage other researchers to more closely examine imagery from their sites for evidence of branch movements, which may provide deeper insights into water and solute movements in plants and physiological responses to water stress. [ABSTRACT FROM AUTHOR]

Solar tracking in the common sunflower, Helianthus annuus, is a dramatic example of a diurnal rhythm in plants. During the day, the shoot apex continuously reorients, following the sun's relative position so that the developing heads track from east to west. At night, the reverse happens, and the heads return and face east in anticipation of dawn. This daily cycle dampens and eventually stops at anthesis, after which the sunflower head maintains an easterly orientation. Although shoot apical heliotropism has long been the subject of physiological studies in sunflower, the underlying developmental, cellular, and molecular mechanisms that drive the directional growth and curvature of the stem in response to extrinsic and perhaps intrinsic cues are not known. Furthermore, the ecological functions of solar tracking and the easterly orientation of mature heads have been the subject of significant but unresolved speculation. In this review, we discuss the current state of knowledge about this complex, dynamic trait. Candidate mechanisms that may contribute to daytime and nighttime movement are highlighted, including light signaling, hormonal action, and circadian regulation of growth pathways. The merits of the diverse hypotheses advanced to explain the adaptive significance of heliotropism in sunflower are also considered.

The article focuses on complexity and diversity of motion amplification and control strategies in motile carnivorous plant traps. Topics include the plants have evolved mechanisms for elastic energy storage and release to power and control rapid motion, the lack of consistent terminology and conceptual frameworks describing elastically powered motion in both groups, and the iconic examples of fast movements can be found in carnivorous plants.

Background and Aims In angiosperms, many species disperse their seeds autonomously by rapid movement of the pericarp. The fruits of these species often have long rod- or long plate-shaped pericarps, which are suitable for ejecting seeds during fruit dehiscence by bending or coiling. However, here we show that fruit with a completely different shape can also rely on pericarp movement to disperse seeds explosively, as in Orixa japonica. Methods Fruit morphology was observed by hard tissue sectioning, scanning electron microscopy and micro-computed tomography, and the seed dispersal process was analysed using a high-speed camera. Comparisons were made of the geometric characteristics of pericarps before and after fruit dehiscence, and the mechanical process of pericarp movement was simulated with the aid of the finite element model. Key Results During fruit dehydration, the water drop-shaped endocarp of O. japonica with sandwich structure produced two-way bending deformation and cracking, and its width increased more than three-fold before opening. Meanwhile the same shaped exocarp with uniform structure could only produce small passive deformation under relatively large external forces. The endocarp forced the exocarp to open by hygroscopic movement before seed launching, and the exocarp provided the acceleration for seed launching through a reaction force. Conclusions Two layers of water drop-shaped pericarp in O. japonica form a structure similar to a slingshot, which launches the seed at high speed during fruit dehiscence. The results suggest that plants with explosive seed dispersal appear to have a wide variety of fruit morphology, and through a combination of different external shapes and internal structures, they are able to move rapidly using many sophisticated mechanisms. [ABSTRACT FROM AUTHOR]

The goal of the study was to quantify and identify patterns in circadian movements of small-leaved lime (Tillia cordata) saplings with the help of terrestrial laser scanning (TLS). The movements were monitored every 60 min 24 h a day and every 30 min in the hour of sunrise and sunset. In order to exclude wind effects the monitored saplings were indoors. The resulting point clouds were used in creating a time series of branch and foliage movements with high precision. The circadian vertical movement of saplings was evaluated through target points, which has a potential of capturing the point-wise movement more accurately. Our results clearly show that small saplings move their branches and leaves during 24 h in complex ways and that is difficult to identify general patterns. Since we worked with small saplings and our movement threshold was 5 mm, we detected random fluctuation–oscillation as the most common movement in monitored saplings. The results highlight the potential of TLS measurements in support of chronobiology and the possibilities to analyze circadian movements of saplings in controlled environment.

Background and Aims The endangered aquatic carnivorous waterwheel plant (Aldrovanda vesiculosa) catches prey with 3–5-mm-long underwater snap-traps. Trapping lasts 10–20 ms, which is 10-fold faster than in its famous sister, the terrestrial Venus flytrap (Dionaea muscipula). After successful capture, the trap narrows further and forms a 'stomach' for the digestion of prey, the so-called 'sickle-shaped cavity'. To date, knowledge is very scarce regarding the deformation process during narrowing and consequent functional morphology of the trap. Methods We performed comparative analyses of virtual 3D histology using computed tomography (CT) and conventional 2D histology. For 3D histology we established a contrasting agent-based preparation protocol tailored for delicate underwater plant tissues. Key Results Our analyses reveal new structural insights into the adaptive architecture of the complex A. vesiculosa snap-trap. In particular, we discuss in detail the arrangement of sensitive trigger hairs inside the trap and present actual 3D representations of traps with prey. In addition, we provide trap volume calculations at different narrowing stages. Furthermore, the motile zone close to the trap midrib, which is thought to promote not only the fast trap closure by hydraulics but also the subsequent trap narrowing and trap reopening, is described and discussed for the first time in its entirety. Conclusions Our research contributes to the understanding of a complex, fast and reversible underwater plant movement and supplements preparation protocols for CT analyses of other non-lignified and sensitive plant structures. [ABSTRACT FROM AUTHOR]

The rachis of most growing compound leaves observed in nature exhibits a stereotypical hook shape. In this study, we focus on the canonical case of Averrhoa carambola. Combining kinematics and mechanical investigation, we characterize this hook shape and shed light on its establishment and maintenance. We show quantitatively that the hook shape is a conserved bent zone propagating at constant velocity and constant distance from the apex throughout development. A simple mechanical test reveals non-zero intrinsic curvature profiles for the rachis during its growth, indicating that the hook shape is actively regulated. We show a robust spatial organization of growth, curvature, rigidity, and lignification, and their interplay. Regulatory processes appear to be specifically localized: in particular, differential growth occurs where the elongation rate drops. Finally, impairing the graviception of the leaf on a clinostat led to reduced hook curvature but not to its loss. Altogether, our results suggest a role for proprioception in the regulation of the leaf hook shape, likely mediated via mechanical strain. [ABSTRACT FROM AUTHOR]

SUMMARY: Circadian organ movements are ubiquitous in plants. These rhythmic outputs are thought to be regulated by the circadian clock and auxin signalling, but the underlying mechanisms have not been clarified. Flowers of Nicotiana attenuata change their orientation during the daytime through a 140° arc to balance the need for pollinators and the protection of their reproductive organs. This rhythmic trait is under the control of the circadian clock and results from bending and re‐straightening movements of the pedicel, stems that connect flowers to the inflorescence. Using an explant system that allowed pedicel growth and curvature responses to be characterized with high spatial and temporal resolution, we demonstrated that this movement is organ autonomous and mediated by auxin. Changes in the growth curvature of the pedicel are accompanied by an auxin gradient and dorsiventral asymmetry in auxin‐dependent transcriptional responses; application of auxin transport inhibitors influenced the normal movements of this organ. Silencing the expression of the circadian clock component ZEITLUPE (ZTL) arrested changes in the growth curvature of the pedicel and altered auxin signalling and responses. IAA19‐like, an Aux/IAA transcriptional repressor that is circadian regulated and differentially expressed between opposite tissues of the pedicel, and therefore possibly involved in the regulation of changes in organ curvature, physically interacted with ZTL. Together, these results are consistent with a direct link between the circadian clock and the auxin signalling pathway in the regulation of this rhythmic floral movement. Significance Statement: Circadian organ movements are widespread among plants. Yet, the molecular components that operate in these movements have remained unknown. Here, we demonstrate that the growth movements of the pedicel, the organ responsible for the circadian changes in orientation of Nicotiana attenuata flowers, are under the control of the circadian clock and auxin signalling; a result that opens potential avenues to investigate the relationship among different types of movements in plants. [ABSTRACT FROM AUTHOR]

The goal of the study was to quantify and identify patterns in circadian movements of small-leaved lime (Tillia cordata) saplings with the help of terrestrial laser scanning (TLS). The movements were monitored every 60 min 24 h a day and every 30 min in the hour of sunrise and sunset. In order to exclude wind effects the monitored saplings were indoors. The resulting point clouds were used in creating a time series of branch and foliage movements with high precision. The circadian vertical movement of saplings was evaluated through target points, which has a potential of capturing the point-wise movement more accurately. Our results clearly show that small saplings move their branches and leaves during 24 h in complex ways and that is difficult to identify general patterns. Since we worked with small saplings and our movement threshold was 5 mm, we detected random fluctuation–oscillation as the most common movement in monitored saplings. The results highlight the potential of TLS measurements in support of chronobiology and the possibilities to analyze circadian movements of saplings in controlled environment. [ABSTRACT FROM AUTHOR]

We developed biomimetic hygro-responsive composite polymer scales inspired by the reversible shape-changes of Bhutan pine (Pinus wallichiana) cone seed scales. The synthetic kinematic response is made possible through novel four-dimensional (4D) printing techniques with anisotropic material use, namely copolymers with embedded cellulose fibrils and ABS polymer. Multi-phase motion like the subsequent transversal and longitudinal bending deformation during desiccation of a natural pinecone scale can be structurally programmed into such printed hygromorphs. Both the natural concept generator (Bhutan pinecone scale) and the biomimetic technical structure (4D printed scale) were comparatively investigated as to their displacement and strain over time via three-dimensional digital image correlation methods. Our bioinspired prototypes can be the basis for tailored autonomous and selfsufficient flap and scale structures performing complex consecutive motions for technical applications, e.g. in architecture and soft robotics. [ABSTRACT FROM AUTHOR]

Background and Aims General anaesthetics are compounds that induce loss of responsiveness to environmental stimuli in animals and humans. The primary site of action of general anaesthetics is the nervous system, where anaesthetics inhibit neuronal transmission. Although plants do not have neurons, they generate electrical signals in response to biotic and abiotic stresses. Here, we investigated the effect of the general volatile anaesthetic diethyl ether on the ability to sense potential prey or herbivore attacks in the carnivorous plant Venus flytrap (Dionaea muscipula). Methods We monitored trap movement, electrical signalling, phytohormone accumulation and gene expression in response to the mechanical stimulation of trigger hairs and wounding under diethyl ether treatment. Key Results Diethyl ether completely inhibited the generation of action potentials and trap closing reactions, which were easily and rapidly restored when the anaesthetic was removed. Diethyl ether also inhibited the later response: jasmonic acid (JA) accumulation and expression of JA-responsive genes (cysteine protease dionain and type I chitinase). However, external application of JA bypassed the inhibited action potentials and restored gene expression under diethyl ether anaesthesia, indicating that downstream reactions from JA are not inhibited. Conclusions The Venus flytrap cannot sense prey or a herbivore attack under diethyl ether treatment caused by inhibited action potentials, and the JA signalling pathway as a consequence. [ABSTRACT FROM AUTHOR]

In this article we adapt a methodology customarily used to investigate movement in animals to study the movement of plants. The targeted movement is circumnutation, a helical organ movement widespread among plants. It is variable due to a different magnitude of the trajectory (amplitude) exhibited by the organ tip, duration of one cycle (period), circular, elliptical, pendulum-like or irregular shape and the clockwise and counterclockwise direction of rotation. The acquisition setup consists of two cameras used to obtain a stereoscopic vision for each plant. Cameras switch to infrared recording mode for low light level conditions, allowing continuous motion acquisition during the night. A dedicated software enables semi-automatic tracking of key points of the plant and reconstructs the 3D trajectory of each point along the whole movement. Three-dimensional trajectories for different points undergo a specific processing to compute those features suitable to describe circumnutation (e.g., maximum speed, circumnutation center, circumnutation length, etc.). By applying our method to the approach-to-grasp movement exhibited by climbing plants (Pisum sativum L.) it appears clear that the plants scale movement kinematics according to the features of the support in ways that are adaptive, flexible, anticipatory and goal-directed, reminiscent of how animals would act.

Despite the fundamental importance of plants to our very survival, student interest in plant biology is in decline as technology draws us further away from nature. Here we introduce Plant Tracer (http://www.planttracer.com), a Matlab-based program, which can quantify time-lapse videos of plant movement. We demonstrate that Plant Tracer can be used to distinguish altered movement qualities in the inflorescence (flowering) stem in the Arabidopsis pgm-1 (phosphoglucomutase) mutant when compared to wildtype, providing a genetic platform for students to evaluate how plants sense and respond to gravity and circumnutation (the back-and-forth swaying of plant organs). We show that both gravitropism and circumnutation is diminished in the pgm-1 mutant when compared to wildtype. In this way, Plant Tracer is a promising instructional tool for biology labs to quantify the genetics of plant movement using smartphones. [ABSTRACT FROM AUTHOR]

Nutations of plant organs are significantly affected by the circatidal modulation in the gravitational force exerted by the Moon and Sun (lunisolar tidal acceleration, Etide). In a previous study on nutational rotations of stem apices, we observed abrupt alterations in their direction and irregularities of the recorded trajectories. Such transitions have not yet been analysed in detail.Peppermint plants were continuously recorded with time‐lapse photography and aligned with contemporaneous time courses of the Etide estimates. Each nutational stem tip movement path was assigned to one of two groups, depending on its geometry, as: (i) regular elliptical movements and (ii) irregular movements (with a random type of trajectory). Analyses of the correlation between the plant nutation trajectory parameters and Etide, as well as of the trajectory geometry of the individual plants were performed.The trajectory geometry of young mint stem apices was related to the velocity of the apex rotation and significantly affected by the gravitational force estimated from the Etide. A low velocity of nutational movement, associated with the random character of the trajectory, usually occurred simultaneously with local minima or maxima of Etide. As the mint plant ages, the transitions in the stem tip trajectory were limited; no correspondence with Etide dynamics was observed.The results indicate that the plant tip geometry path transitions with respect to the changing gradient of lunisolar tidal acceleration could be interpreted as manifestation of a continuous accommodation of the shoot apical part to the state of minimum energy dissipation. [ABSTRACT FROM AUTHOR]

The fast motion of the snap-traps of the terrestrial Venus flytrap (Dionaea muscipula) have been intensively studied, in contrast to the tenfold faster underwater snap-traps of its phylogenetic sister, the waterwheel plant (Aldrovanda vesiculosa). Based on biomechanical and functional--morphological analyses and on a reverse biomimetic approach via mechanical modelling and computer simulations, we identify a combination of hydraulic turgor change and the release of prestress stored in the trap as essential for actuation. Our study is the first to identify and analyse in detail the motion principle of Aldrovanda, which not only leads to a deepened understanding of fast plant movements in general, but also contributes to the question of how snap-traps may have evolved and also allows for the development of novel biomimetic compliant mechanisms. [ABSTRACT FROM AUTHOR]

Summary: Plants, although sessile organisms, are nonetheless able to move their body parts; for example, during root contraction of geophytes or in the gravitropic reaction by woody stems. One of the major mechanisms enabling these movements is the development of specialized structures that possess contractile properties. Quite unlike animal muscles, for which the action is driven by protein–protein interactions in the protoplasma, the action of plant ‘muscles’ is polysaccharide‐based and located in the uniquely designed, highly cellulosic cell wall that is deposited specifically in fibers. This review describes the development of such cell walls as a widespread phenomenon in the plant kingdom, gives reasons why it should be considered as a tertiary cell wall, and discusses the mechanism of action of the ‘muscles’. The origin of the contractile properties lies in the tension of the axially oriented cellulose microfibrils due to entrapment of rhamnogalacturonan‐I aggregates that limits the lateral interaction of microfibrils. Long side chains of the nascent rhamnogalacturonan‐I are trimmed off during cell wall maturation leading to tension development. Similarities in the tertiary cell wall design in fibers of different plant origin indicate that the basic principles of tension creation may be universal in various ecophysiological situations. [ABSTRACT FROM AUTHOR]

In the September 2017 issue of Carnivorous Plant Newsletter, the term 'carnitropism' was proposed to name triggered movements occurring in Dionaea, Drosera, and Pinguicula genera. However, those phenomena were misconceived and their interpretation lacked scientific background, misleading to a flawed conclusion. Therefore, the present paper recommends to avoid the use of that term and recalls both the mechanisms behind plant movements and their associated names. [ABSTRACT FROM AUTHOR]

Action potentials (APs), i.e., long-distance electrical signals, and circumnutations (CN), i.e., endogenous plant organ movements, are shaped by ion fluxes and content in excitable and motor tissues. The appearance of APs and CN as well as growth parameters in seedlings and 3-week old plants of Helianthus annuus treated with osmotic and salt stress (0–500 mOsm) were studied. Time-lapse photography and extracellular measurements of electrical potential changes were performed. The hypocotyl length was strongly reduced by the osmotic and salt stress. CN intensity declined due to the osmotic but not salt stress. The period of CN in mild salt stress was similar to the control (~164 min) and increased to more than 200 min in osmotic stress. In sunflower seedlings growing in a hydroponic medium, spontaneous APs (SAPs) propagating basipetally and acropetally with a velocity of 12–20 cm min−1 were observed. The number of SAPs increased 2–3 times (7–10 SAPs 24 h−1plant−1) in the mild salt stress (160 mOsm NaCl and KCl), compared to the control and strong salt stress (3–4 SAPs 24 h−1 plant−1 in the control and 300 mOsm KCl and NaCl). Glutamate-induced series of APs were inhibited in the strong salt stress-treated seedlings but not at the mild salt stress and osmotic stress. Additionally, in 3-week old plants, the injection of the hypo- or hyperosmotic solution at the base of the sunflower stem evoked series of APs (3–24 APs) transmitted along the stem. It has been shown that osmotic and salt stresses modulate differently hypocotyl growth and CN and have an effect on spontaneous and evoked APs in sunflower seedlings. We suggested that potassium, sodium, and chloride ions at stress concentrations in the nutrient medium modulate sunflower excitability and CN.

Action potentials (APs), i.e., long-distance electrical signals, and circumnutations (CN), i.e., endogenous plant organ movements, are shaped by ion fluxes and content in excitable and motor tissues. The appearance of APs and CN as well as growth parameters in seedlings and 3-week old plants of Helianthus annuus treated with osmotic and salt stress (0–500 mOsm) were studied. Time-lapse photography and extracellular measurements of electrical potential changes were performed. The hypocotyl length was strongly reduced by the osmotic and salt stress. CN intensity declined due to the osmotic but not salt stress. The period of CN in mild salt stress was similar to the control (~164 min) and increased to more than 200 min in osmotic stress. In sunflower seedlings growing in a hydroponic medium, spontaneous APs (SAPs) propagating basipetally and acropetally with a velocity of 12–20 cm min−1 were observed. The number of SAPs increased 2–3 times (7–10 SAPs 24 h−1plant−1) in the mild salt stress (160 mOsm NaCl and KCl), compared to the control and strong salt stress (3–4 SAPs 24 h−1 plant−1 in the control and 300 mOsm KCl and NaCl). Glutamate-induced series of APs were inhibited in the strong salt stress-treated seedlings but not at the mild salt stress and osmotic stress. Additionally, in 3-week old plants, the injection of the hypo- or hyperosmotic solution at the base of the sunflower stem evoked series of APs (3–24 APs) transmitted along the stem. It has been shown that osmotic and salt stresses modulate differently hypocotyl growth and CN and have an effect on spontaneous and evoked APs in sunflower seedlings. We suggested that potassium, sodium, and chloride ions at stress concentrations in the nutrient medium modulate sunflower excitability and CN. [ABSTRACT FROM AUTHOR]

In modern urban existence, the complex lives of plants are often reduced to simple categories, which resonate with human utility. These categories speak little of the central role plants play in the ecological fabric of life on Earth. Plants have the ability to sense and respond to stimuli across various timescales and the time zones they inhabit are multi‐faceted. This complexity presents difficulties when the unfamiliar characteristics of "plantness" are revealed to everyday observers who may perceive plants as operating in a slow lane outside of their perception. This editorial draws attention to the rich time assemblages in which plants exist, and highlights the need for diverse representations with which to engage human attention to the botanical world. [ABSTRACT FROM AUTHOR]

Problems with understanding concepts and mechanisms connected to plant movements have been diagnosed among biology students. Alternative conceptions in understanding these phenomena are marginally studied. The diagnosis was based on a sample survey of university students and their lecturers, which was quantitatively and qualitatively exploratory in nature (via a questionnaire). The research was performed in two stages, before and after the lectures and laboratory on plant movements. We diagnosed eight alternative conceptions before the academic training started. After the classes, most were not been verified, and in addition, 12 new conceptions were diagnosed. Additionally, we report that teachers are not aware of students’ possible misunderstandings. They do not perceive students’ troubles with switching between levels of representations, nor their alternative conceptions. A case of “curse of knowledge” was observed and academic teacher training is recommended. Additionally, the need for metacognition as a crucial element in laboratory activities seems supported by our presented results. Such metacognition refers to students as well as teachers, which leads to the conclusion that teachers should be aware of students’ way of thinking and the development of knowledge in one’s own mind.

This paper presents a discovery: the change of the outline shape of mistletoe (Viscum album ssp. album) berries in vivo and in situ during ripening. It was found that a plant organ that is usually considered to merely increase in size actually changes shape in a specific rhythmic fashion. We introduce a new approach to chronobiological research on a macro-phenotypic scale to trace changes over long periods of time (with a resolution from hours to months) by using a dynamic form-determining parameter called Lambda (λ). λ is known in projective geometry as a measure for pertinent features of the outline shapes of egg-like forms, so called path curves. Ascertained circadian changes of form were analyzed for their correlation with environmental factors such as light, temperature, and other weather influences. Certain weather conditions such as sky cover, i.e., sunshine minutes per hour, have an impact on the amplitude of the daily change in form. The present paper suggests a possible supplement to established methods in chronobiology, as in this case the dynamic of form-change becomes a measurable feature, displaying a convincing accordance between mathematical rule and plant shape. [ABSTRACT FROM AUTHOR]

The goal of the study was to determine circadian movements of silver birch (Petula Bendula) branches and foliage detected with terrestrial laser scanning (TLS). The study consisted of two geographically separate experiments conducted in Finland and in Austria. Both experiments were carried out at the same time of the year and under similar outdoor conditions. Experiments consisted of 14 (Finland) and 77 (Austria) individual laser scans taken between sunset and sunrise. The resulting point clouds were used in creating a time series of branch movements. In the Finnish data, the vertical movement of the whole tree crown was monitored due to low volumetric point density. In the Austrian data, movements of manually selected representative points on branches were monitored. The movements were monitored from dusk until morning hours in order to avoid daytime wind effects. The results indicated that height deciles of the Finnish birch crown had vertical movements between -10.0 and 5.0cm compared to the situation at sunset. In the Austrian data, the maximum detected representative point movement was 10.0 cm. The temporal development of the movements followed a highly similar pattern in both experiments, with the maximum movements occurring about an hour and a half before (Austria) or around (Finland) sunrise. The results demonstrate the potential of terrestrial laser scanning measurements in support of chronobiology. [ABSTRACT FROM AUTHOR]

Background One of the best-known plant movements, phototropic solar tracking in sunflower (Helianthus annuus), has not yet been fully characterized. Two questions are still a matter of debate. (1) Is the adaptive significance solely an optimization of photosynthesis via the exposure of the leaves to the sun? (2) Is shade avoidance involved in this process? In this study, these concepts are discussed from a historical perspective and novel insights are provided. Scope and Methods Results from the primary literature on heliotropic growth movements led to the conclusion that these responses cease before anthesis, so that the flowering heads point to the East. Based on observations on 10-week-old plants, the diurnal East-West oscillations of the upper fifth of the growing stem and leaves in relation to the position of the sun (inclusive of nocturnal re-orientation) were documented, and photon fluence rates on the leaf surfaces on clear, cloudy and rainy days were determined. In addition, the light-response curve of net CO2 assimilation was determined on the upper leaves of the same batch of plants, and evidence for the occurrence of shade-avoidance responses in growing sunflower plants is summarized. Conclusions. Only elongating, vegetative sunflower shoots and the upper leaves perform phototropic solar tracking. Photon fluence response and CO2 assimilation measurements cast doubt on the 'photosynthesis-optimization hypothesis' as the sole explanation for the evolution of these plant movements. We suggest that the shadeavoidance response, which maximizes light-driven CO2 assimilation, plays a major role in solar tracking populations of competing sunflower plants, and an integrative scheme of these growth movements is provided. [ABSTRACT FROM AUTHOR]

Plants use rapid movements to disperse seed, spores, or pollen and catch animal prey. Most rapid-release mechanisms only work once and, if repeatable, regaining the prerelease state is a slow and costly process. We present an encompassing mechanism for a rapid, repeatable, passive-dynamic motion used by a carnivorous pitcher plant to catch prey. Nepenthes gracilis uses the impact of rain drops to catapult insects from the underside of the canopy-like pitcher lid into the fluid-filled trap below. High-speed video and laser vibrom-etry revealed that the lid acts as a torsional spring system, driven by rain drops. During the initial downstroke, the tip of the lid reached peak velocities similar to fast animal motions and an order of magnitude faster than the snap traps of Venus flytraps and catapulting tentacles of the sundew Drosera glanduligera. In contrast to these active movements, the N. gracilis lid oscillation requires neither mechanical preloading nor metabolic energy, and its repeatability is only limited by the intensity and duration of rainfall. The underside of the lid is coated with friction-reducing wax crystals, making insects more vulnerable to perturbations. We show that the trapping success of N. gracilis relies on the combination of material stiffness adapted for momentum transfer and the antiadhesive properties of the wax crystal surface. The impact-driven oscillation of the N. gracilis lid represents a new kind of rapid plant movement with adaptive function. Our findings establish the existence of a continuum between active and passive trapping mechanisms in carnivorous plants. [ABSTRACT FROM AUTHOR]