A unifying modeling of plant shoot gravitropism with an explicit account of the effects of growth

Gravitropism, the slow reorientation of plant growth in response to gravity, is a major determinant of the form and posture of land plants. Recently a universal model of shoot gravitropism, the AC model, was presented, in which the dynamics of the tropic movement is only determined by the conflicting controls of (1) graviception that tends to curve the plants toward the vertical, and (2) proprioception that tends to keep the stem straight. This model was found to be valid for many species and over two orders of magnitude of organ size. However, the motor of the movement, the elongation, was purposely neglected in the AC model. If growth effects are to be taken into account, it is necessary to consider the material derivative, i.e., the rate of change of curvature bound to expanding and convected organ elements. Here we show that it is possible to rewrite the material equation of curvature in a compact simplified form that directly expresses the curvature variation as a function of the median elongation and of the distribution of the differential growth. By using this extended model, called the ACĖ model, growth is found to have two main destabilizing effects on the tropic movement: (1) passive orientation drift, which occurs when a curved element elongates without differential growth, and (2) fixed curvature, when an element leaves the elongation zone and is no longer able to actively change its curvature. By comparing the AC and ACĖ models to experiments, these two effects are found to be negligible. Our results show that the simplified AC mode can be used to analyze gravitropism and posture control in actively elongating plant organs without significant information loss

La biomécanique des plantes ou " Comment les plantes tiennent debout ? "

La biomécanique des plantes est née d'une approche interdisciplinaire. Elle regroupe ainsi des biologistes, des forestiers ou des agronomes, et des mécaniciens (des matériaux et des structures et plus récemment des fluides). Elle s'intéresse à la façon dont les plantes se portent et croissent. En particulier comment les plantes terrestres croissent en hauteur " contre " la gravité et dans le vent ; mais aussi comment les plantes aquatiques et les algues se développent dans les courants

Inclination not force is sensed by plants during shoot gravitropism

International audienceGravity perception plays a key role in how plants develop and adapt to environmental changes. However, more than a century after the pioneering work of Darwin, little is known on the sensing mechanism. Using a centrifugal device combined with growth kinematics imaging, we show that shoot gravitropic responses to steady levels of gravity in four representative angiosperm species is independent of gravity intensity. All gravitropic responses tested are dependent only on the angle of inclination from the direction of gravity. We thus demonstrate that shoot gravitropism is stimulated by sensing inclination not gravitational force or acceleration as previously believed. This contrasts with the otolith system in the internal ear of vertebrates and explains the robustness of the control of growth direction by plants despite perturbations like wind shaking. Our results will help retarget the search for the molecular mechanism linking shifting statoliths to signal transduction

Monitoring the regulation of gene expression in a growing organ using a fluid mechanics formalism

<p>Abstract</p> <p>Background</p> <p>Technological advances have enabled the accurate quantification of gene expression, even within single cell types. While transcriptome analyses are routinely performed, most experimental designs only provide snapshots of gene expression. Molecular mechanisms underlying cell fate or positional signalling have been revealed through these discontinuous datasets. However, in developing multicellular structures, temporal and spatial cues, known to directly influence transcriptional networks, get entangled as the cells are displaced and expand. Access to an unbiased view of the spatiotemporal regulation of gene expression occurring during development requires a specific framework that properly quantifies the rate of change of a property in a moving and expanding element, such as a cell or an organ segment.</p> <p>Results</p> <p>We show how the rate of change in gene expression can be quantified by combining kinematics and real-time polymerase chain reaction data in a mechanistic model which considers any organ as a continuum. This framework was applied in order to assess the developmental regulation of the two reference genes <it>Actin11 </it>and <it>Elongation Factor 1-β </it>in the apex of poplar root. The growth field was determined by time-lapse photography and transcript density was obtained at high spatial resolution. The net accumulation rates of the transcripts of the two genes were found to display highly contrasted developmental profiles. <it>Actin11 </it>showed pulses of up and down regulation in the accelerating and decelerating parts of the growth zone while the dynamic of <it>EF1β </it>were much slower. This framework provides key information about gene regulation in a developing organ, such as the location, the duration and the intensity of gene induction/repression.</p> <p>Conclusions</p> <p>We demonstrated that gene expression patterns can be monitored using the continuity equation without using mutants or reporter constructions. Given the rise of imaging technologies, this framework in our view opens a new way to dissect the molecular basis of growth regulation, even in non-model species or complex structures.</p

Measuring local and global vibration modes in model plants

International audienceModel plants are extensively used in biological studies, and their mechanical behaviour needs to be better understood, in relation to studies in mechanoperception for instance. We present here the first approach to derive experimentally the modal parameters of two of these plants, Arabidopsis thaliana and Populus tremula alba. A classical sinusoidal sweep excitation is used, with a measurement of displacements based on LKT optical flow tracking, followed by a Bi-Orthogonal decomposition (BOD). This allows to estimate several modal frequencies for each plant, as well as the corresponding spatial localizations of deformation. Analyzing the modal frequencies, we show that global and local modes correspond to distinct ranges of frequencies and depend differently on plant size. Possible phenotyping applications are then discussed

Caractérisation vibratoire d'une plante : expériences et calculs

La dynamique d'une plante est caractérisée par stimulation mécanique à l'aide d'un pot vibrant par balayages entre 0 et 40Hz. On analyse les films obtenus pour obtenir le champ de vitesse, que l'on traite par Décomposition Bi Orthogonale. On détermine ainsi l'existence d'un mode dans la plage de fréquence considérée et la déformée modale associée. D'autre part, un calcul éléments finis des modes à partir des caractéristiques réelles de la plante est effectué et les résultats sont comparés aux expériences. On observe des modes localisés et organisés en modes de tige, de feuilles et d'apex

Velocimetric third-harmonic generation microscopy: micrometer-scale quantification of morphogenetic movements in unstained embryos

International audienceWe demonstrate the association of third-harmonic generation (THG) microscopy and particle image velocimetry (PIV) analysis as a novel functional imaging technique for automated micrometer-scale characterization of morphogenetic movements in developing embryos. Using a combined two-photon-excited fluorescence and THG microscope, we characterize the optical properties of Drosophila embryos and show that sustained THG imaging does not perturb sensitive developmental dynamics. Velocimetric THG imaging provides a quantitative description of the dynamics of internal structures in unstained wild-type and mutant embryos

DORN1/P2K1 and purino-calcium signalling in plants: making waves with extracellular ATP.

BACKGROUND AND AIMS:Extracellular ATP governs a range of plant functions, including cell viability, adaptation and cross-kingdom interactions. Key functions of extracellular ATP in leaves and roots may involve an increase in cytosolic free calcium as a second messenger ('calcium signature'). The main aim here was to determine to what extent leaf and root calcium responses require the DORN1/P2K1 extracellular ATP receptor in Arabidopsis thaliana. The second aim was to test whether extracellular ATP can generate a calcium wave in the root. METHODS:Leaf and root responses to extracellular ATP were reviewed for their possible links to calcium signalling and DORN1/P2K1. Leaves and roots of wild type and dorn1 plants were tested for cytosolic calcium increase in response to ATP, using aequorin. The spatial abundance of DORN1/P2K1 in the root was estimated using green fluorescent protein. Wild type roots expressing GCaMP3 were used to determine the spatial variation of cytosolic calcium increase in response to extracellular ATP. KEY RESULTS:Leaf and root ATP-induced calcium signatures differed markedly. The leaf signature was only partially dependent on DORN1/P2K1, while the root signature was fully dependent. The distribution of DORN1/P2K1 in the root supports a key role in the generation of the apical calcium signature. Root apical and sub-apical calcium signatures may operate independently of each other but an apical calcium increase can drive a sub-apical increase, consistent with a calcium wave. CONCLUSION:DORN1 could underpin several calcium-related responses but it may not be the only receptor for extracellular ATP in Arabidopsis. The root has the capacity for a calcium wave, triggered by extracellular ATP at the apex

The zinc finger protein PtaZFP2 negatively controls stem growth and gene expression responsiveness to external mechanical loads in poplar

Mechanical cues are essential signals regulating plant growth and development. In response to wind, trees develop a thigmomorphogenetic response characterized by a reduction in longitudinal growth, an increase in diameter growth, and changes in mechanical properties. The molecular mechanisms behind these processes are poorly understood. In poplar, PtaZFP2, a C2H2 transcription factor, is rapidly up-regulated after stem bending. To investigate the function of PtaZFP2, we analyzed PtaZFP2-overexpressing poplars (Populus tremula 9 Populus alba). To unravel the genes downstream PtaZFP2, a transcriptomic analysis was performed. PtaZFP2-overexpressing poplars showed longitudinal and cambial growth reductions together with an increase in the tangent and hardening plastic moduli. The regulation level of mechanoresponsive genes was much weaker after stem bending in PtaZFP2-overexpressing poplars than in wild-type plants, showing that PtaZFP2 negatively modulates plant responsiveness to mechanical stimulation. Microarray analysis revealed a high proportion of down-regulated genes in PtaZFP2-overexpressing poplars. Among these genes, several were also shown to be regulated by mechanical stimulation. Our results confirmed the important role of PtaZFP2 during plant acclimation to mechanical load, in particular through a negative control of plant molecular responsiveness. This desensitization process could modulate the amplitude and duration of the plant response during recurrent stimuli