Search

Highlight Arabidopsis actin-related protein ARP3 plays a role in amyloplast sedimentation and polar auxin redistribution during root gravitropism., Gravitropism is vital for shaping directional plant growth in response to the forces of gravity. Signals perceived in the gravity-sensing cells can be converted into biochemical signals and transmitted. Sedimentation of amyloplasts in the columella cells triggers asymmetric auxin redistribution in root tips, leading to downward root growth. The actin cytoskeleton is thought to play an important role in root gravitropism, although the molecular mechanism has not been resolved. DISTORTED1 (DIS1) encodes the ARP3 subunit of the Arabidopsis Actin-Related Protein 2/3 (ARP2/3) complex, and the ARP3/DIS1 mutant dis1-1 showed delayed root curvature after gravity stimulation. Microrheological analysis revealed that the high apparent viscosity within dis1-1 central columella cells is closely associated with abnormal movement trajectories of amyloplasts. Analysis using a sensitive auxin input reporter DII-VENUS showed that asymmetric auxin redistribution was reduced in the root tips of dis1-1, and the actin-disrupting drug Latrunculin B increased the asymmetric auxin redistribution. An uptake assay using the membrane-selective dye FM4-64 indicated that endocytosis was decelerated in dis1-1 root epidermal cells. Treatment and wash-out with Brefeldin A, which inhibits protein transport from the endoplasmic reticulum to the Golgi apparatus, showed that cycling of the auxin-transporter PIN-FORMED (PIN) proteins to the plasma membrane was also suppressed in dis1-1 roots. The results reveal that ARP3/DIS1 acts in root gravitropism by affecting amyloplast sedimentation and PIN-mediated polar auxin transport through regulation of PIN protein trafficking.

Gravitropism in plants is one of the most controversial issues. In the most wildly accepted starch-statolith hypothesis the sedimentation movement of amyloplasts in the gravisensing columella cells primarily triggers the asymmetric distribution of auxin which leads to the differential growth of the plant root. It has been gradually recognized that the inhomogeneous structures in statocytes arising from intracellular components such as cytoskeletons significantly affect the complex movements of amyloplasts and the final gravimorphogenesis. In this letter, we implement a diffusive dynamics measurement and inplanta microrheological analysis of amyloplasts in the wild-type plants and actin cytoskeleton mutants for the first time. We found that the intracellular environment of columella cells exhibits the spatial heterogeneity and the cage-confinement on amyloplasts which is the typically characteristics in colloidal suspensions. By comparing the distinct diffusive dynamics of amyloplasts in different types of plants with the behaviors of colloidal systems in different states, we quantitatively characterized the influence of the actin organization dominated intracellular envoronments on the amyloplast movements. Furthermore, the cage-confinement strength was measured by calculating the spatial fluctuation of local apparent viscosity within the columella cells. Finally, a linear association between the initial mechanical stimulation in the columella cells the subsequent intercellular signal transduction and the final gravity response was observed and a possible gravity sensing mechanism was suggested. It suggests the existence of a potential gravity-sensing mechanism that dictates a linear frustration effect of the actin cytoskeleton on the conversion of the mechanical stimulation of amyloplasts into gravitropic signals.

The self-diffusion problem of Brownian particles under the constraint of quasi-one-dimensional (q1D) channel has raised wide concern. The hydrodynamic interaction (HI) plays an important role in many practical problems and two-body interactions remain dominant under q1D constraint. We measure the diffusion coefficient of individual ellipsoid when two ellipsoidal particles are close to each other by video-microscopy measurement. Meanwhile, we obtain the numerical simulation results of diffusion coefficient using finite element software. We find that the self-diffusion coefficient of the ellipsoid decreases exponentially with the decrease of their mutual distance X when X < X 0 , where X 0 is the maximum distance of the ellipsoids to maintain their mutual influence, X 0 and the variation rate are related to the aspect ratio p=a/b. The mean squared displacement (MSD) of the ellipsoids indicates that the self-diffusion appears as a crossover region, in which the diffusion coefficient increases as the time increases in the intermediate time regime, which is proven to be caused by the spatial variations affected by the hydrodynamic interactions. These findings indicate that hydrodynamic interaction can significantly affect the self-diffusion behavior of adjacent particles and has important implications to the research of microfluidic problems in blood vessels and bones, drug delivery, and lab-on-chip.

We study the Brownian motion of a single ellipsoidal particle diffusing in a narrow channel by video-microscopy measurement. The experiments allow us to obtain the trajectories of ellipsoids and measure the diffusion coefficients. It is found that the channel constraints lead to suppression of the particle motion, especially the perpendicular motion to the channel, and the long axis of the particle tends to be parallel to the channel. A stable stratification phenomenon is observed, which is rarely discussed in studies of spherical particles. We also derive an approximate solution of theoretical prediction with the method of reflections, and obtain numerical simulation results using finite element software. They are proven to be effective by comparing them with the experimental results. All of these indicate that the aspect ratio and size of ellipsoid, the width of channel, and the transverse position distinctly affect the Brownian motion of ellipsoids.

Gravitropism in plants is key for orienting organs such as the Darwin’s description of the opposite growth direction of roots and shoots in his book The Power of Movement in Plants more than 100 years ago (Darwin, 1880). The gravitropic response of an oriented plant is divided into three sequential phases: gravity sensing, signal transmission, and the growth response. As the initial phase of gravitropism, gravity sensing (the conversion of the mechanical stimulus into a biochemical signal) has been intensively studied, but much is still unknown.