A number of hormones work together to control plant cell growth. Rapid Alkalinization Factor 1 (RALF1), a plant-derived small regulatory peptide, inhibits cell elongation through suppression of rhizosphere acidification in plants. Although a receptor-like kinase, FERONIA (FER), has been shown to act as a receptor for RALF1, the signaling mechanism remains unknown. In this study, we identified a receptor-like cytoplasmic kinase (RPM1-induced protein kinase, RIPK), a plasma membrane-associated member of the RLCK-VII subfamily, that is recruited to the receptor complex through interacting with FER in response to RALF1. RALF1 triggers the phosphorylation of both FER and RIPK in a mutually dependent manner. Genetic analysis of the fer-4 and ripk mutants reveals RIPK, as well as FER, to be required for RALF1 response in roots. The RALF1-FER-RIPK interactions may thus represent a mechanism for peptide signaling in plants.plant hormone | feronia | phosphorylation
FERONIA (FER), a plasma membrane receptor-like kinase, is a central regulator of cell growth that integrates environmental and endogenous signals. A peptide ligand rapid alkalinization factor 1 (RALF1) binds to FER and triggers a series of downstream events, including inhibition of Arabidopsis H+-ATPase 2 activity at the cell surface and regulation of gene expression in the nucleus. We report here that, upon RALF1 binding, FER first promotes ErbB3-binding protein 1 (EBP1) mRNA translation and then interacts with and phosphorylates the EBP1 protein, leading to EBP1 accumulation in the nucleus. There, EBP1 associates with the promoters of previously identified RALF1-regulated genes, such as CML38, and regulates gene transcription in response to RALF1 signaling. EBP1 appears to inhibit the RALF1 peptide response, thus forming a transcription–translation feedback loop (TTFL) similar to that found in circadian rhythm control. The plant RALF1-FER-EBP1 axis is reminiscent of animal epidermal growth factor receptor (EGFR) signaling, in which EGF peptide induces EGFR to interact with and phosphorylate EBP1, promoting EBP1 nuclear accumulation to control cell growth. Thus, we suggest that in response to peptide signals, plant FER and animal EGFR use the conserved key regulator EBP1 to control cell growth in the nucleus.
The molecular links between extracellular signals and the regulation of localized protein synthesis in plant cells are poorly understood. Here, we show that in Arabidopsis thaliana, the extracellular peptide RALF1 and its receptor, the FERONIA receptor kinase, promote root hair (RH) tip growth by modulating protein synthesis. We found that RALF1 promotes FERONIA-mediated phosphorylation of eIF4E1, a eukaryotic translation initiation factor that plays a crucial role in the control of mRNA translation rate. Phosphorylated eIF4E1 increases mRNA affinity and modulates mRNA translation and, thus, protein synthesis. The mRNAs targeted by the RALF1-FERONIA-eIF4E1 module include ROP2 and RSL4, which are important regulators of RH cell polarity and growth. RALF1 and FERONIA are expressed in a polar manner in RHs, which facilitate eIF4E1 polar localization and thus may control local ROP2 translation. Moreover, we demonstrated that high-level accumulation of RSL4 exerts negative-feedback regulation of RALF1 expression by directly binding the RALF1 gene promoter, determining the final RH size. Our study reveals that the link between RALF1-FERONIA signaling and protein synthesis constitutes a novel component regulating cell expansion in these polar growing cells.
Home Digital Voice Assistants (HDVAs) are getting popular in recent years. Users can control smart devices and get living assistance through those HDVAs (e.g., Amazon Alexa, Google Home) using voice. In this work, we study the insecurity of HDVA service by using Amazon Alexa as a case study. We disclose three security vulnerabilities which root in the insecure access control of Alexa services. We then exploit them to devise two proof-of-concept attacks, home burglary and fake order, where the adversary can remotely command the victim's Alexa device to open a door or place an order from Amazon.com. The insecure access control is that the Alexa device not only relies on a single-factor authentication but also takes voice commands even if no people are around. We thus argue that HDVAs should have another authentication factor, a physical presence based access control; that is, they can accept voice commands only when any person is detected nearby. To this end, we devise a Virtual Security Button (VSButton), which leverages the WiFi technology to detect indoor human motions. Once any indoor human motion is detected, the Alexa device is enabled to accept voice commands. Our evaluation results show that it can effectively differentiate indoor motions from the cases of no motion and outdoor motions in both the laboratory and real world settings.• This paper is officially published at CNS 2018 [5].
The environmentally responsive signaling pathways that link global transcriptomic changes through alternative splicing (AS) to plant fitness remain unclear. Here, we found that the interaction of the extracellular rapid alkalinization FACTOR 1 (RALF1) peptide with its receptor FERONIA (FER) triggered a rapid and massive RNA AS response by interacting with and phosphorylating glycine-rich RNA binding protein7 (GRP7) to elevate GRP7 nuclear accumulation in Arabidopsis thaliana. FER-dependent GRP7 phosphorylation enhanced its mRNA binding ability and its association with the spliceosome component U1-70K to enable splice site selection, modulating dynamic AS. Genetic reversal of a RALF1-FER–dependent splicing target partly rescued mutants deficient in GRP7. AS of GRP7 itself induced nonsense-mediated decay feedback to the RALF1-FER-GRP7 module, fine-tuning stress responses, and cell growth. The RALF1-FER-GRP7 module provides a paradigm for regulatory mechanisms of RNA splicing in response to external stimuli.
Precise and dynamic imaging of extracellular pH is one crucial yet challenging task for studying cell physiological and pathological processes. Here, we construct a DNA tweezer to dynamically monitor pH changes of cellular microenvironments. The DNA tweezer contains three key elements: a three-strand ssDNA-frame labeled with cholesterol to anchor it on the cell membrane, a pH-sensitive i-motif sequence in the middle to dynamically control the switch between the "open" and "closed" states of the DNA tweezer, and a pair of FRET fluorophores (rhodamine green and rhodamine red) on the two arms of the tweezer to reflect its state. With cholesterol, a natural component of cell membranes, as an anchoring element, the sensor exhibited high cell-membrane-insertion efficiency and low cytotoxicity. Using the i-motif as a sensing element, it can quickly and reversibly respond to extracellular pH in the pH range of 5.0−7.5 and further perform real-time imaging of cell-surface-pH changes with excellent spatial and temporal resolution. Moreover, apoplastic-pH change during the alkalization process of plant roots caused by rapid-alkalinization factor (RALF1) was directly detected by the sensor, demonstrating the potential applications of the sensor in cell biology, biomedical research, and plant-tissue engineering.
Rate adaptation (RA) has been used to achieve high goodput. In this work, we explore to use RA for energy efficiency in 802.11n NICs. We show that current MIMO RA algorithms are not energy efficient for NICs despite ensuring high throughput. The fundamental problem is that, the high-throughput setting is not equivalent to the energy-efficient one. Marginal throughput gain may be realized at high energy cost. We propose EERA, an energy-based RA solution that trades off goodput for energy savings at NICs. Our experiments have confirmed its energy savings at NICs while keeping the cost at the device level and across clients acceptable.
Cell expansion is coordinated by several cues, but available energy is the major factor determining growth. Receptor protein kinase FERONIA (FER) is a master regulator of cell expansion, but the details of its control mechanisms are not clear. Here we show that FER interacts with cytosolic enzyme glyceraldehyde-3-phosphate dehydrogenase (GAPDH, GAPC1 and GAPC2), that catalyzes a key reaction in glycolysis, which contributes to energy production. When there is an FER deficiency, there are corresponding decreases in the enzyme activity of GAPDH and increased amounts of starch. More importantly, gapc1/2 mutants mimic fer4 mutants. These data indicate that FER regulated starch content is an evolutionarily conserved function in plants that connects the cell expansion and energy metabolism pathways.
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