Expansins facilitate cell expansion by mediating pH-dependent cell wall (CW) loosening. However, the role of expansins in controlling CW biomechanical properties in specific tissues and organs remains elusive. We monitored hormonal responsiveness and spatial specificity of expression and localization of expansins predicted to be the direct targets of cytokinin signaling in Arabidopsis (Arabidopsis thaliana). We found EXPANSIN1 (EXPA1) homogenously distributed throughout the CW of columella/lateral root cap, while EXPA10 and EXPA14 localized predominantly at 3-cell boundaries in the epidermis/cortex in various root zones. EXPA15 revealed cell-type-specific combination of homogenous vs. 3-cell boundaries localization. By comparing Brillouin frequency shift and AFM-measured Young's modulus, we demonstrated Brillouin light scattering (BLS) as a tool suitable for non-invasive in vivo quantitative assessment of CW viscoelasticity. Using both BLS and AFM, we showed that EXPA1 overexpression upregulated CW stiffness in the root transition zone (TZ). The dexamethasone-controlled EXPA1 overexpression induced fast changes in the transcription of numerous CW-associated genes, including several EXPAs and XYLOGLUCAN:XYLOGLUCOSYL TRANSFERASEs (XTHs), and associated with rapid pectin methylesterification determined by in situ Fourier-transform infrared spectroscopy in the root TZ. The EXPA1-induced CW remodeling is associated with the shortening of the root apical meristem, leading to root growth arrest. Based on our results, we propose that expansins control root growth by a delicate orchestration of CW biomechanical properties, possibly regulating both CW loosening and CW remodeling.

Biomechanical properties of the cell wall (CW) are important for many developmental and adaptive responses in plants. Expansins were shown to mediate pH-dependent CW enlargement via a process called CW loosening. Here, we provide a brief overview of expansin occurrence in plant and non-plant species, their structure and mode of action including the role of hormone-regulated CW acidification in the control of expansin activity. We depict the historical as well as recent CW models, discuss the role of expansins in the CW biomechanics and address the developmental importance of expansin-regulated CW loosening in cell elongation and new primordia formation. We summarise the data published so far on the role of expansins in the abiotic stress response as well as the rather scarce evidence and hypotheses on the possible mechanisms underlying expansin-mediated abiotic stress resistance. Finally, we wrap it up by highlighting possible future directions in expansin research.

Understanding how plants adapt their physiology to overcome severe and often multifactorial stress conditions in nature is vital in light of the climate crisis. This remains a challenge given the complex nature of the underlying molecular mechanisms. To provide a comprehensive picture of stress-mitigation mechanisms, an exhaustive analysis of publicly available stress-related transcriptomic data has been conducted. We combine a meta-analysis with an unsupervised machine-learning algorithm to identify a core of stress-related genes active at 1-6 h and 12-24 h of exposure in Arabidopsis thaliana shoots and roots. To ensure robustness and biological significance of the output, often lacking in meta-analyses, a triple validation is incorporated. We present a ‘stress gene core’: a set of key genes involved in plant tolerance to ten adverse environmental conditions and ethylene-precursor supplementation rather than individual conditions. Notably, ethylene plays a key regulatory role in this core, influencing gene expression and acting as a critical factor in stress tolerance. Additionally, the analysis provides insights into previously uncharacterized genes, key genes within large families, and gene expression dynamics, which are used to create biologically validated databases that can guide further abiotic stress research. These findings establish a strong framework for advancing multi-stress-resilient crops, paving the way for sustainable agriculture in the face of climate challenges.

Expansins are cell wall-modifying proteins implicated in plant growth and stress responses. In this study, we explored the differential localization of expansins in Arabidopsis thaliana shoots, with a focus on EXPA1, EXPA10, EXPA14, and EXPA15 utilizing pEXPA::EXPA translational fusion lines. Employing the chemically inducible system pOp6/LhGR for EXPA1 overexpression and high-throughput automatic phenotyping we evaluated the drought response and photosynthetic efficiency under stress conditions. We observed distinct expression patterns of expansins, with EXPA1 primarily localized in stomatal guard cells, while EXPA10 and EXPA15 showed strong cell wall (CW) localization in epidermal and other tissues. Overexpression of EXPA1 resulted in pronounced changes in CW-related gene expression, particularly during early stages of induction, including the upregulation of other expansins and CW-modifying enzymes. The induced EXPA1 line also displayed significant morphological changes in shoots, including smaller plant size, delayed senescence, and structural alterations in vascular tissues. Additionally, EXPA1 overexpression conferred drought tolerance, as evidenced by enhanced photosynthetic efficiency (F_v/F_M), and low steady-state non-photochemical quenching (NPQ) values under drought stress. These findings highlight the critical role of EXPA1 in regulating plant growth, development, and stress response, with potential applications in improving drought tolerance in crops.

Expansins facilitate cell expansion via mediating pH-dependent cell wall (CW) loosening. However, the role of expansins in the control of biomechanical CW properties in the tissue and organ context remains elusive. We determined hormonal responsiveness and specificity of expression and localization of expansins predicted to be direct targets of cytokinin signalling. We found EXPA1 homogenously distributed throughout the CW of columella/ lateral root cap, while EXPA10 and EXPA14 localized predominantly at the three-cell boundaries of epidermis/cortex in various root zones. Cell type-specific localization of EXPA15 overlaps with higher CW stiffness measured via Brillouin light scattering microscopy. As indicated by both Brillouin frequency shift and AFM-measured Young’s modulus, EXPA1 overexpression upregulated CW stiffness, associated with shortening of the root apical meristem and root growth arrest. We propose that root growth in Arabidopsis requires delicate orchestration of biomechanical CW properties via tight regulation of various expansins’ localization to specific cell types and extracellular domains.