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1. Lipid Signaling Requires ROS Production to Elicit Actin Cytoskeleton Remodeling during Plant Innate Immunity

   https://doi.org/10.3390/ijms23052447  

   Cao, Lingyan; Wang, Wenyi; Zhang, Weiwei; Staiger, Christopher J. (March 2022, International Journal of Molecular Sciences)

   In terrestrial plants a basal innate immune system, pattern-triggered immunity (PTI), has evolved to limit infection by diverse microbes. The remodeling of actin cytoskeletal arrays is now recognized as a key hallmark event during the rapid host cellular responses to pathogen attack. Several actin binding proteins have been demonstrated to fine tune the dynamics of actin filaments during this process. However, the upstream signals that stimulate actin remodeling during PTI signaling remain poorly characterized. Two second messengers, reactive oxygen species (ROS) and phosphatidic acid (PA), are elevated following pathogen perception or microbe-associated molecular pattern (MAMP) treatment, and the timing of signaling fluxes roughly correlates with actin cytoskeletal rearrangements. Here, we combined genetic analysis, chemical complementation experiments, and quantitative live-cell imaging experiments to test the role of these second messengers in actin remodeling and to order the signaling events during plant immunity. We demonstrated that PHOSPHOLIPASE Dβ (PLDβ) isoforms are necessary to elicit actin accumulation in response to flg22-associated PTI. Further, bacterial growth experiments and MAMP-induced apoplastic ROS production measurements revealed that PLDβ-generated PA acts upstream of ROS signaling to trigger actin remodeling through inhibition of CAPPING PROTEIN (CP) activity. Collectively, our results provide compelling evidence that PLDβ/PA functions upstream of RBOHD-mediated ROS production to elicit actin rearrangements during the innate immune response in Arabidopsis. 

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2. Nuclear Entanglement: New Insights Into the Role of Cytoskeleton and Nucleoskeleton in Plant Nuclear Function

   https://doi.org/10.1002/cm.22048  

   Groves, Norman R; Amstutz, Katelyn; Schumacher, Lily A; Meier, Iris (June 2025, Cytoskeleton)

   ABSTRACT Of the three types of cytoskeleton known in animals—actin, microtubules, and intermediate filaments—only actin and microtubules exist in plants. Both play important roles in cellular shaping, organelle movement, organization of the endomembrane system, and cell signaling. An emerging, but often overlooked role of the plant cytoskeleton is its dynamic and mutually influential interaction with the nucleus. Here, we summarize recent advances in understanding the role of the cytoskeleton in plant nuclear movement in different biological contexts, a role for nuclear envelope‐associated proteins in reorganizing the actin and microtubule cytoskeleton, and the molecular nature of the nucleus‐cytoskeleton interface and specific proteins contributing to it. In animals, the nucleoskeleton consists of the nuclear lamina, an intermediate‐filament meshwork underlying the nuclear envelope. Plants have evolved an equivalent of this structure, built by different types of proteins. Here, we highlight recent advances in understanding its filamentous organization, newly discovered protein interactions connecting it to nuclear pores, and exciting new evidence that—just like the animal lamina—the plant lamina is involved in chromatin reorganization and epigenetic changes. Together, these new developments create new opportunities toward a deeper understanding of this important regulatory connection between the cytoskeleton and the cell's largest organelle. 

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3. WASP family proteins regulate the mobility of the B cell receptor during signaling activation

   https://doi.org/10.1038/s41467-020-14335-8  

   Rey-Suarez, Ivan; Wheatley, Brittany A.; Koo, Peter; Bhanja, Anshuman; Shu, Zhou; Mochrie, Simon; Song, Wenxia; Shroff, Hari; Upadhyaya, Arpita (January 2020, Nature Communications)

   Abstract Regulation of membrane receptor mobility tunes cellular response to external signals, such as in binding of B cell receptors (BCR) to antigen, which initiates signaling. However, whether BCR signaling is regulated by BCR mobility, and what factors mediate this regulation, are not well understood. Here we use single molecule imaging to examine BCR movement during signaling activation and a novel machine learning method to classify BCR trajectories into distinct diffusive states. Inhibition of actin dynamics downstream of the actin nucleating factors, Arp2/3 and formin, decreases BCR mobility. Constitutive loss or acute inhibition of the Arp2/3 regulator, N-WASP, which is associated with enhanced signaling, increases the proportion of BCR trajectories with lower diffusivity. Furthermore, loss of N-WASP reduces the diffusivity of CD19, a stimulatory co-receptor, but not that of FcγRIIB, an inhibitory co-receptor. Our results implicate a dynamic actin network in fine-tuning receptor mobility and receptor-ligand interactions for modulating B cell signaling. 

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4. Plasticity of cell migration resulting from mechanochemical coupling

   https://doi.org/10.7554/eLife.48478  

   Cao, Yuansheng; Ghabache, Elisabeth; Rappel, Wouter-Jan (October 2019, eLife)

   Eukaryotic cells can migrate using different modes, ranging from amoeboid-like, during which actin filled protrusions come and go, to keratocyte-like, characterized by a stable morphology and persistent motion. How cells can switch between these modes is not well understood but waves of signaling events are thought to play an important role in these transitions. Here we present a simple two-component biochemical reaction-diffusion model based on relaxation oscillators and couple this to a model for the mechanics of cell deformations. Different migration modes, including amoeboid-like and keratocyte-like, naturally emerge through transitions determined by interactions between biochemical traveling waves, cell mechanics and morphology. The model predictions are explicitly verified by systematically reducing the protrusive force of the actin network in experiments using Dictyostelium discoideum cells. Our results indicate the importance of coupling signaling events to cell mechanics and morphology and may be applicable in a wide variety of cell motility systems. 

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5. A hybrid stochastic–deterministic mechanochemical model of cell polarization

   https://doi.org/10.1091/mbc.E19-09-0549  

   Copos, Calina; Mogilner, Alex (July 2020, Molecular Biology of the Cell)

   Edelstein-Keshet, Leah (Ed.)

   Polarization is a crucial component in cell differentiation, development, and motility, but its details are not yet well understood. At the onset of cell locomotion, cells break symmetry to form well-defined cell fronts and rears. This polarity establishment varies across cell types: in Dictyostelium discoideum cells, it is mediated by biochemical signaling pathways and can function in the absence of a cytoskeleton, while in keratocytes, it is tightly connected to cytoskeletal dynamics and mechanics. Theoretical models that have been developed to understand the onset of polarization have explored either signaling or mechanical pathways, yet few have explored mechanochemical mechanisms. However, many motile cells rely on both signaling modules and actin cytoskeleton to break symmetry and achieve a stable polarized state. We propose a general mechanochemical polarization model based on coupling between a stochastic model for the segregation of signaling molecules and a simplified mechanical model for actin cytoskeleton network competition. We find that local linear coupling between minimally nonlinear signaling and cytoskeletal systems, separately not supporting stable polarization, yields a robustly polarized cell state. The model captures the essence of spontaneous polarization of neutrophils, which has been proposed to emerge due to the competition between frontness and backness pathways. 

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