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Free, publicly-accessible full text available July 1, 2027
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SUMMARY Stomata are pores at the leaf surface that enable gas exchange and transpiration. The signaling pathways that regulate the differentiation of stomatal guard cells and the mechanisms of stomatal pore formation have been characterized inArabidopsis thaliana. However, the process by which stomatal complexes develop after pore formation into fully mature complexes is poorly understood. We tracked the morphogenesis of young stomatal complexes over time to establish characteristic geometric milestones along the path of stomatal maturation. Using 3D‐nanoindentation coupled with finite element modeling of young and mature stomata, we found that despite having thicker cell walls than young guard cells, mature guard cells are more energy efficient with respect to stomatal opening, potentially attributable to the increased mechanical anisotropy of their cell walls and smaller changes in turgor pressure between the closed and open states. Comparing geometric changes in young and mature guard cells of wild‐type and cellulose‐deficient plants revealed that although cellulose is required for normal stomatal maturation, mechanical anisotropy appears to be achieved by the collective influence of cellulose and additional wall components. Together, these data elucidate the dynamic geometric and biomechanical mechanisms underlying the development process of stomatal maturation.more » « less
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Bayer, Edward (Ed.)Abstract Stomatal function in plants is regulated by the nanoscale architecture of the cell wall and turgor pressure, which together control stomatal pore size to facilitate gas exchange and photosynthesis. The mechanical properties of the cell wall and cell geometry are critical determinants of stomatal dynamics. However, the specific biomechanical functions of wall constituents, for example, cellulose and pectins, and their impact on the work required to open or close the stomatal pore are unclear. Here, we use nanoindentation in normal and lateral directions, computational modeling, and microscopic imaging of cells from the model plant Arabidopsis thaliana to investigate the precise influences of wall architecture and turgor pressure on stomatal biomechanics. This approach allows us to quantify and compare the unique anisotropic properties of guard cells with normal composition, lower cellulose content, or alterations in pectin molecular weight. Using these data to calculate the work required to open the stomata reveals that the wild type, with a circumferential-to-longitudinal modulus ratio of 3:1, is the most energy-efficient of those studied. In addition, the tested genotypes displayed similar changes in their pore size despite large differences in wall thickness and biomechanical properties. These findings imply that homeostasis in stomatal function is maintained in the face of varying wall compositions and biomechanics by tuning wall thickness.more » « less
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Additive manufacturing (AM) often results in high strength but poor ductility in titanium alloys. Hybrid AM is a solution capable of improving both ductility and strength. In this study, hybrid AM of Ti-6Al-4 V was achieved by coupling directed energy deposition with interlayer machining. The microstructure, residual stress, and microhardness were examined to explain how interlayer machining caused a 63% improvement in ductility while retaining an equivalent strength to as-printed samples. Interlayer machining introduced recurrent interruptions in printing that allowed for slow cooling-induced coarsening of acicular α laths at the machined interfaces. The coarse α laths on the selectively machined layers increased dislocation motion under tensile loads and improved bulk ductility. The results highlighted in this publication demonstrate the feasibility of hybrid AM to enhance the toughness of titanium alloys.more » « less
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Chen, Tsu-Wei; Long, Stephen P (Ed.)Abstract Highly polarized cotton fibre cells that develop from the seed coat surface are the foundation of a multi-billion-dollar international textile industry. The unicellular trichoblast emerges as a hemispherical bulge that is efficiently converted to a narrower and elongated shape that extends for about 2 weeks before transitioning into a cellulose-generating machine. The polarized elongation phase employs an evolutionarily conserved microtubule-cellulose synthase control module that patterns the cell wall and enables highly anisotropic diffuse growth. As the multi-scale interactions and feedback controls among cytoskeletal systems, morphologically potent cell wall properties, and a changing cell geometry are uncovered, opportunities emerge to engineer architectural traits. However, in cotton, such efforts are hampered by insufficient knowledge about the underlying control mechanisms. For example, fibre diameter is an important trait that is determined during the earliest stages of development, but the basic growth mode and the mechanisms by which cytoskeletal and cell wall systems mediate fibre tapering are not known. This paper combines multiparametric and multiscale fibre phenotyping and finite element computational modelling of a growing cell to discover an evolutionarily conserved tapering mechanism. The actin network interconverts between two distinct longitudinal organizations that broadly distributes organelles and likely enables matrix secretion patterns that maintain cell wall thickness during growth. Based on plausible finite element models and quantitative analyses of the microtubule cytoskeleton, tapering and anisotropic growth is programmed by a constricting apical microtubule depletion zone and highly aligned microtubules along the fibre shaft. The finite element model points to a central role for tensile forces in the cell wall to dictate the densities and orientations of morphologically potent microtubules that pattern the cell wall.more » « less
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