Although it is a central question in biology, how cell shape controls intracellular dynamics largely remains an open question. Here, we show that the shape of Arabidopsis pavement cells creates a stress pattern that controls microtubule orientation, which then guides cell wall reinforcement. Live-imaging, combined with modeling of cell mechanics, shows that microtubules align along the maximal tensile stress direction within the cells, and atomic force microscopy demonstrates that this leads to reinforcement of the cell wall parallel to the microtubules. This feedback loop is regulated: cell-shape derived stresses could be overridden by imposed tissue level stresses, showing how competition between subcellular and supracellular cues control microtubule behavior. Furthermore, at the microtubule level, we identified an amplification mechanism in which mechanical stress promotes the microtubule response to stress by increasing severing activity. These multiscale feedbacks likely contribute to the robustness of microtubule behavior in plant epidermis.

Wavy walls built by nanofilaments: In the model plant Arabidopsis , pavement cells fit together with the lobes and curves of jigsaw puzzle pieces. Such complex cell shapes, in plants, were generally thought to be driven by turgor pressure. Haas et al. now show that the extracellular cell wall can actively shape the cell it contains without relying on turgor pressure. Nanofilaments of pectin homogalacturonan in the cell wall shift between crystalline and anisotropic phases according to whether they are methylated. The shift in form drives changes in cell wall shape that stand independent of turgor pressure. Science , this issue p. 1003

Significance: How does a cell decide when to divide or initiate DNA replication? How does it regulate its own growth? These fundamental questions are not well understood in most organisms; this lack of understanding is particularly true for multicellular eukaryotes. Following classical studies in yeast, we have quantified the key aspects of cell growth and division dynamics in the Arabidopsis apical stem cell niche. Our results disprove various theories for plant stem cell size/cell cycle regulation, such as that cell cycle progression is triggered when a prefixed critical size is attained, and constitute the necessary first step in the development of integrative mechanistic theories for the coordinated regulation of cell cycle progression, cell growth, and cell size in plants.

Cell cycle regulation: The hormone cytokinin regulates various aspects of plant development and physiology, largely by managing cell proliferation. Yang et al. show that cytokinin promotes nuclear localization of the transcription factor MYB3R4, which activates the expression of two importins and genes that tip the cell into the next phase of the cell cycle. The importins facilitate further MYB3R4 accumulation within the nucleus, accelerating the progression into mitosis. MYB3R4 and the importins dissipate when the nuclear membrane dissolves at prometaphase, so there is only one round of mitotic activation per cell cycle. Science , this issue p. 1350