Mechanical forces have emerged as coordinating signals for most cell functions. Yet, because forces are invisible, mapping tensile stress patterns in tissues remains a major challenge in all ...kingdoms. Here we take advantage of the adhesion defects in the
mutant
to deduce stress patterns in tissues. By reducing the water potential and epidermal tension
, we rescued the adhesion defects in
, formally associating gaping and tensile stress patterns in the mutant. Using suboptimal water potential conditions, we revealed the relative contributions of shape- and growth-derived stress in prescribing maximal tension directions in aerial tissues. Consistently, the tension patterns deduced from the gaping patterns in
matched the pattern of cortical microtubules, which are thought to align with maximal tension, in wild-type organs. Conversely, loss of epidermis continuity in the
mutant hampered supracellular microtubule alignments, revealing that coordination through tensile stress requires cell-cell adhesion.
Sessile plants evolve diverse structures in response to complex environmental cues. These factors, in essence, involve mechanical stimuli, which must be sensed and coordinated properly by the plants ...to ensure effective growth and development. While we have accumulated substantial knowledge on plant mechanobiology, how plants translate mechanical information into three-dimensional structures is still an open question. In this review, we summarize our current understanding of plant mechanosensing at different levels, particularly using Arabidopsis as a model plant system. We also attempt to abstract the mechanosensing process and link the gaps from mechanical cues to the generation of complex plant structures. Here we review the recent advancements on mechanical response and transduction in plant morphogenesis, and we also raise several questions that interest us in different sections.
The above-ground organs in plants display a rich diversity, yet they grow to characteristic sizes and shapes. Organ morphogenesis progresses through a sequence of key events, which are robustly ...executed spatiotemporally as an emerging property of intrinsic molecular networks while adapting to various environmental cues. This Review focuses on the multiscale control of leaf morphogenesis. Beyond the list of known genetic determinants underlying leaf growth and shape, we focus instead on the emerging novel mechanisms of metabolic and biomechanical regulations that coordinate plant cell growth non-cell-autonomously. This reveals how metabolism and mechanics are not solely passive outcomes of genetic regulation but play instructive roles in leaf morphogenesis. Such an integrative view also extends to fluctuating environmental cues and evolutionary adaptation. This synthesis calls for a more balanced view on morphogenesis, where shapes are considered from the standpoints of geometry, genetics, energy and mechanics, and as emerging properties of the cellular expression of these different properties.
Plant cells cannot rearrange their positions; therefore, sharp tissue boundaries must be accurately programmed. Movement of the cell fate regulator SHORT-ROOT from the stele to the ground tissue has ...been associated with transferring positional information across tissue boundaries. The zinc finger BIRD protein JACKDAW has been shown to constrain SHORT-ROOT movement to a single layer, and other BIRD family proteins were postulated to counteract JACKDAW’s role in restricting SHORT-ROOT action range. Here, we report that regulation of SHORT-ROOT movement requires additional BIRD proteins whose action is critical for the establishment and maintenance of the boundary between stele and ground tissue. We show that BIRD proteins act in concert and not in opposition. The exploitation of asymmetric redundancies allows the separation of two BIRD functions: constraining SHORTROOT spread through nuclear retention and transcriptional regulation of key downstream SHORT-ROOT targets, including SCARECROW and CYCLIND6. Our data indicate that BIRD proteins promote formative divisions and tissue specification in the Arabidopsis thaliana root meristem ground tissue by tethering and regulating transcriptional competence of SHORT-ROOT complexes. As a result, a tissue boundary is not “locked in” after initial patterning like in many animal systems, but possesses considerable developmental plasticity due to continuous reliance on mobile transcription factors.
In plants, where cells cannot migrate, asymmetric cell divisions (ACDs) must be confined to the appropriate spatial context. We investigate tissue-generating asymmetric divisions in a stem cell ...daughter within the Arabidopsis root. Spatial restriction of these divisions requires physical binding of the stem cell regulator SCARECROW (SCR) by the RETINOBLASTOMA-RELATED (RBR) protein. In the stem cell niche, SCR activity is counteracted by phosphorylation of RBR through a cyclinD6;1-CDK complex. This cyclin is itself under transcriptional control of SCR and its partner SHORT ROOT (SHR), creating a robust bistable circuit with either high or low SHR-SCR complex activity. Auxin biases this circuit by promoting CYCD6;1 transcription. Mathematical modeling shows that ACDs are only switched on after integration of radial and longitudinal information, determined by SHR and auxin distribution, respectively. Coupling of cell-cycle progression to protein degradation resets the circuit, resulting in a “flip flop” that constrains asymmetric cell division to the stem cell region.
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► SCR binds its partner SHR to stimulate together with auxin CyclinD6 transcription ► CyclinD6 inactivates RBR, but RBR binds and inactivates SCR, blocking CyclinD6 ► This bistable switch interprets multiple gradients to trigger stem cell division ► Protein degradation is used to bring daughter cells back to an “off” state
Stem cells in the root meristem integrate radial and longitudinal gradients of tissue patterning determinants to ensure that only specific stem cells undergo asymmetric cell divisions.
The shoot apical meristem (SAM) gives rise to all aerial plant organs. Cell walls are thought to play a central role in this process, translating molecular regulation into dynamic changes in growth ...rate and direction, although their precise role in morphogenesis during organ formation is poorly understood. Here, we investigated the role of xyloglucans (XyGs), a major, yet functionally poorly characterized, wall component in the SAM of Arabidopsis (
). Using immunolabeling, biochemical analysis, genetic approaches, microindentation, laser ablation, and live imaging, we showed that XyGs are important for meristem shape and phyllotaxis. No difference in the Young's modulus (i.e. an indicator of wall stiffness) of the cell walls was observed when XyGs were perturbed. Mutations in enzymes required for XyG synthesis also affect other cell wall components such as cellulose content and pectin methylation status. Interestingly, control of cortical microtubule dynamics by the severing enzyme KATANIN became vital when XyGs were perturbed or absent. This suggests that the cytoskeleton plays an active role in compensating for altered cell wall composition.
Intercellular signaling through trafficking of regulatory proteins is a widespread phenomenon in plants and can deliver positional information for the determination of cell fate. In the Arabidopsis ...root meristem, the cell fate determinant SHORT‐ROOT (SHR), a GRAS domain transcription factor, acts as a signaling molecule from the stele to the adjacent layer to specify endodermal cell fate. Upon exiting the stele, SHR activates another GRAS domain transcription factor, SCARCROW (SCR), which, together with several BIRD/INDETERMINATE DOMAIN proteins, restricts movement of SHR to define a single cell layer of endodermis. Here we report that endodermal cell fate also requires the joint activity of both SCR and its closest homologue SCARECROW‐LIKE23 (SCL23). We show that SCL23 protein moves with zonation‐dependent directionality. Within the meristem, SCL23 exhibits short‐ranged movement from ground tissue to vasculature. Away from the meristem, SCL23 displays long‐range rootward movement into meristematic vasculature and a bidirectional radial spread, respectively. As a known target of SHR and SCR, SCL23 also interacts with SCR and SHR and can restrict intercellular outspread of SHR without relying on nuclear retention as SCR does. Collectively, our data show that SCL23 is a mobile protein that controls movement of SHR and acts redundantly with SCR to specify endodermal fate in the root meristem.
CuAlO2 was synthesized by a hydrothermal method, in which the Cu–O dimers were incorporated by simply altering the ratio of the reactants and the temperature. The incorporation process increases the ...grain size in CuAlO2, and modulates the work function and binding energies for CuAlO2 due to the partial substitution of Cu+ 3d10 with Cu2+ 3d9 orbitals in the valence band maximum by alloying non-isovalent Cu–O with a CuAlO2 host. Based on the ZnO nanorod arrays (NRs) ultraviolet photodetector, CuAlO2/Cu–O fabricated by the low-cost drop-coating method was used as the p-type hole transport layer. The incorporation of the Cu–O clusters into CuAlO2 lattice to enhance the conductivity of CuAlO2 is an effective way for improving ZnO NRs/CuAlO2 device performance. The photodetectors exhibit significant diode behavior, with a rectification ratio approaching 30 at ±1 V, and a dark saturation current density 0.81 mA cm−2. The responsivity of the ZnO-NRs-based UV photodetector increases from 13.2 to 91.3 mA/W at 0 V bias, with an increase in the detectivity from 2.35 × 1010 to 1.71 × 1011 Jones. Furthermore, the ZnO NRs/CuAlO2/Cu–O photodetector exhibits a maximum responsivity of 5002 mA/W at 1.5 V bias under 375 nm UV illumination.
The way in which interactions between mechanics and biochemistry lead to the emergence of complex cell and tissue organization is an old question that has recently attracted renewed interest from ...biologists, physicists, mathematicians and computer scientists. Rapid advances in optical physics, microscopy and computational image analysis have greatly enhanced our ability to observe and quantify spatiotemporal patterns of signalling, force generation, deformation, and flow in living cells and tissues. Powerful new tools for genetic, biophysical and optogenetic manipulation are allowing us to perturb the underlying machinery that generates these patterns in increasingly sophisticated ways. Rapid advances in theory and computing have made it possible to construct predictive models that describe how cell and tissue organization and dynamics emerge from the local coupling of biochemistry and mechanics. Together, these advances have opened up a wealth of new opportunities to explore how mechanochemical patterning shapes organismal development. In this roadmap, we present a series of forward-looking case studies on mechanochemical patterning in development, written by scientists working at the interface between the physical and biological sciences, and covering a wide range of spatial and temporal scales, organisms, and modes of development. Together, these contributions highlight the many ways in which the dynamic coupling of mechanics and biochemistry shapes biological dynamics: from mechanoenzymes that sense force to tune their activity and motor output, to collectives of cells in tissues that flow and redistribute biochemical signals during development.
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