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Histone acetylation plays critical roles in chromatin remodeling, DNA repair, and epigenetic regulation of gene expression, but the underlying mechanisms are unclear. Proteasomes usually catalyze ATP- and polyubiquitin-dependent proteolysis. Here, we show that the proteasomes containing the activator PA200 catalyze the polyubiquitin-independent degradation of histones. Most proteasomes in mammalian testes ("spermatoproteasomes") contain a spermatid/sperm-specific α subunit α4 s/PSMA8 and/or the catalytic ß subunits of immunoproteasomes in addition to PA200. Deletion of PA200 in mice abolishes acetylation-dependent degradation of somatic core histones during DNA double-strand breaks and delays core histone disappearance in elongated spermatids. Purified PA200 greatly promotes ATP-independent proteasomal degradation of the acetylated core histones, but not polyubiquitinated proteins. Furthermore, acetylation on histones is required for their binding to the bromodomain-like regions in PA200 and its yeast ortholog, Blm10. Thus, PA200/Blm10 specifically targets the core histones for acetylation-mediated degradation by proteasomes, providing mechanisms by which acetylation regulates histone degradation, DNA repair, and spermatogenesis.
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Reparación del ADN , Histonas/metabolismo , Proteínas Nucleares/metabolismo , Complejo de la Endopetidasa Proteasomal/metabolismo , Espermatogénesis , Testículo/metabolismo , Acetilación , Secuencia de Aminoácidos , Animales , Roturas del ADN de Doble Cadena , Humanos , Masculino , Ratones , Datos de Secuencia Molecular , Proteínas Nucleares/química , Estructura Terciaria de Proteína , Saccharomyces cerevisiae/metabolismo , Proteínas de Saccharomyces cerevisiae/metabolismo , Alineación de SecuenciaRESUMEN
DExD/H-box helicases are crucial regulators of RNA metabolism and antiviral innate immune responses; however, their role in bacteria-induced inflammation remains unclear. Here, we report that DDX5 interacts with METTL3 and METTL14 to form an m6A writing complex, which adds N6-methyladenosine to transcripts of toll-like receptor (TLR) 2 and TLR4, promoting their decay via YTHDF2-mediated RNA degradation, resulting in reduced expression of TLR2/4. Upon bacterial infection, DDX5 is recruited to Hrd1 at the endoplasmic reticulum in an MyD88-dependent manner and is degraded by the ubiquitin-proteasome pathway. This process disrupts the DDX5 m6A writing complex and halts m6A modification as well as degradation of TLR2/4 mRNAs, thereby promoting the expression of TLR2 and TLR4 and downstream NF-κB activation. The role of DDX5 in regulating inflammation is also validated in vivo, as DDX5- and METTL3-KO mice exhibit enhanced expression of inflammatory cytokines. Our findings show that DDX5 acts as a molecular switch to regulate inflammation during bacterial infection and shed light on mechanisms of quiescent inflammation during homeostasis.
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Adenina , Infecciones Bacterianas , Receptor Toll-Like 2 , Animales , Ratones , Adenina/análogos & derivados , Inflamación/genética , Metiltransferasas/genética , Receptor Toll-Like 2/genética , Receptor Toll-Like 4/genéticaRESUMEN
In this work, we hybridize an air cavity reflector and a nanopatterned sapphire substrate (NPSS) for making an inclined-sidewall-shaped deep ultraviolet micro light-emitting diode (DUV micro-LED) array to enhance the light extraction efficiency (LEE). A cost-effective hybrid photolithography process involving positive and negative photoresist (PR) is explored to fabricate air-cavity reflectors. The experimental results demonstrate a 9.88% increase in the optical power for the DUV micro-LED array with a bottom air-cavity reflector when compared with the conventional DUV micro-LED array with only a sidewall metal reflector. The bottom air-cavity reflector significantly contributes to the reduction of the light absorption and provides more escape paths for light, which in turn increases the LEE. Our investigations also report that such a designed air-cavity reflector exhibits a more pronounced impact on small-size micro-LED arrays, because more photons can propagate into escape cones by experiencing fewer scattering events from the air-cavity structure. Furthermore, the NPSS can enlarge the escape cone and serve as scattering centers to eliminate the waveguiding effect, which further enables the improved LEE for the DUV micro-LED array with an air-cavity reflector.
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It is known that light extraction efficiency (LEE) for AlGaN-based deep ultraviolet (DUV) light-emitting diodes (LEDs) can be enhanced by using an inclined sidewall of mesa. However, the reported optimal inclined angles are different. In this work, to explore the origin for enhancing the LEE of DUV LED by using inclined sidewalls, we investigate the effect of an inclined sidewall angle on the LEE for AlGaN-based DUV LEDs with different mesa diameters by using ray tracing. It is found that when compared to large-size DUV LEDs with inclined sidewall, the LEE of small-size DUV LEDs with inclined sidewall is enhanced from both the bottom and side surfaces due to the reduced scattering length and material absorption. Additionally, the optimal inclined sidewall angle is recommended within the range of 25°-65°, and the optimal angle for DUV LEDs decreases as the chip size increases. It can be attributed to the fact that there are two scattering mechanisms for the inclined sidewall. For smaller chip sizes, most of the light is directly scattered into escape cones by the inclined sidewall, resulting in a larger optimal angle. For larger chip sizes, the light firstly experiences total internal reflections by the out-light plane and then is scattered into escape cones by the inclined sidewalls, leading to a smaller optimal angle.
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In this Letter, an omni-directional reflector (ODR) with a thin hybrid dielectric layer (hybrid-ODR) is proposed to enhance the light extraction efficiency (LEE) for inclined-sidewall-shaped AlGaN-based deep ultraviolet light-emitting diode (DUV LED) by inserting a thin diamond with high refraction index into a conventional Al/Al2O3-based ODR. The three-dimensional finite-difference time-domain (3D FDTD) simulation results show that the LEE of TM-polarized light for the DUV LED with hybrid-ODR is enhanced by 18.5% compared with Al/Al2O3-based ODR. It is because the diamond can transform the evanescent wave in Al2O3 into the propagating light wave in diamond, thereby preventing effective excitation of the surface plasmon polariton (SPP) on the surface of the metal Al. Moreover, the Brewster's angle effect causes the TM-polarized light in diamond to propagate effectively into AlGaN. Furthermore, decreasing the total thickness of the dielectric layer also improves the scattering effect of the inclined sidewall. However, the utilization of hybrid-ODR results in a slight reduction in the LEE for transverse electric (TE) polarized light because the light is confined to the diamond layer and eventually absorbed by the metal Al.
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In this work, by using three-dimensional finite-difference time-domain (3D FDTD) method, the effect of conventional nano-patterned sapphire substrate (NPSS) on the optical crosstalk and the light extraction efficiency (LEE) for InGaN/GaN-based flip-chip micro light-emitting diodes (µ-LEDs) are systematically studied. We find that the conventional NPSS is not suitable for µ-LEDs. It is because the inclined mesa sidewall for µ-LEDs possesses a good scattering effect for µ-LEDs, but the introduced conventional NPSS causes part of the light be off escape cone between sapphire and air and become the guided light. To suppress the guided light and improve the optical crosstalk, a thick air layer between the n-GaN layer and the sapphire substrate can be used as a light filter to prevent the guided light from propagating into the sapphire. However, in reality, it is challenging to make the aforementioned air layer from point of fabrication view. Therefore, we propose the air-cavity patterned sapphire substrate (AC-PSS) as the light filter. Our results show that the crosstalk ratio can be decreased to the value even lower than 10%. The LEE can also be enhanced simultaneously due to combination effects of the filtering effect of the AC-PSS and the scattering effect of the inclined mesa sidewall.
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In this Letter, beveled mesas for 30 × 30â µm2 GaN-based micro-light-emitting diodes (µLEDs) with different inclination angles are designed, fabricated, and measured. We find that µLED with a mesa inclination angle of 28° has the lowest internal quantum efficiency (IQE) and the highest injection current density at which the peak IQE is obtained. This is due to the increased quantum confined Stark effect (QCSE) at the mesa edge. The increased QCSE results from the strong electric field coupling effect. Instead of radiative recombination, more nonradiative recombination and leakage current will be generated in the sidewall regions. Besides, the smallest angle (28°) also produces the lowest light extraction efficiency (LEE), which arises from the optical loss caused by the sidewall reflection at the beveled surface sides. Therefore, the inclination angle for the beveled mesa has to be increased to 52° and 61° by using Ni and SiO2 as hard masks, respectively. Experimental and numerical results show that the external quantum efficiency (EQE) and the optical power can be enhanced for the fabricated devices. Meanwhile, the reduced surface recombination rate also decreases the leakage current.
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In this paper, by using advanced numerical models, we investigate the impact of the AlN/GaN distributed Bragg reflector (DBR) and AlInN/GaN DBR on stimulated radiative recombination for GaN-based vertical-cavity-surface-emitting lasers (VCSELs). According to our results, when compared with the VCSEL with AlN/GaN DBR, we find that the VCSEL with AlInN/GaN DBR decreases the polarization-induced electric field in the active region, and this helps to increase the electron-hole radiative recombination. However, we also find that the AlInN/GaN DBR has a reduced reflectivity when compared with the AlN/GaN DBR with the same number of pairs. Furthermore, this paper suggests that more pairs of AlInN/GaN DBR will be set, which helps to even further increase the laser power. Hence, the 3 dB frequency can be increased for the proposed device. In spite of the increased laser power, the smaller thermal conductivity for AlInN than AlN results in the earlier thermal droop in the laser power for the proposed VCSEL.
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OBJECTIVE: To examine the influence of widely used protein affinity tags and the tobacco PR1a signal peptide (SP) on detection, purification and bioactivity analyses of the small oomycete apoplastic effector SCR96 in planta. RESULTS: Through agroinfiltration, the phytotoxic effector SCR96 of Phytophthora cactorum was expressed in Nicotiana benthamiana leaf apoplast as a fusion protein carrying single affinity tag (His, HA or FLAG) at either C- or N-terminus. Leaf necrosis caused by different affinity-tagged SCR96 varied among tags and replicates. All of tagged proteins can be detected by antibodies against SCR96. All of SCR96 fusions except N-terminally fused 6His-tagged protein were detected using tag antibodies, indicating that 6His tag may be degraded when fused at N-terminus. Interestingly, C-terminal His- and FLAG-tagged SCR96 maintained the biological activity after purification. In the substitution assay of SCR96 SP, we observed that PR1a SP can lead chimeric SCR96 expression in N. benthamiana, but the replacement totally disrupted its bioactivity. CONCLUSION: C-terminal His or FLAG tag, along with its original SP, is efficient enough to enable detection and purification of functional SCR96 from N. benthamiana leaf apoplast, which would facilitate plant-pathogen interaction studies.
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Nicotiana , Phytophthora , Nicotiana/genética , Nicotiana/metabolismo , Señales de Clasificación de Proteína/genética , Proteínas/metabolismo , Phytophthora/genética , Phytophthora/metabolismo , Anticuerpos/metabolismo , Cromatografía de AfinidadRESUMEN
Meiosis, which produces haploid progeny, is critical to ensuring both faithful genome transmission and genetic diversity. Proteasomes play critical roles at various stages of spermatogenesis, including meiosis, but the underlying mechanisms remain unclear. The atypical proteasomes, which contain the activator PA200, catalyze the acetylation-dependent degradation of the core histones in elongated spermatids and DNA repair in somatic cells. We show here that the testis-specific proteasome subunit α4s/PSMA8 is essential for male fertility by promoting proper formation of spermatoproteasomes, which harbor both PA200 and constitutive catalytic subunits. Immunostaining of a spermatocyte marker, SYCP3, indicated that meiosis was halted at the stage of spermatocytes in the α4s-deficient testes. α4s stimulated the in vitro degradation of the acetylated core histones, instead of nonacetylated histones, by the PA200-proteasome. Deletion of α4s blocked degradation of the core histones at DNA damage loci in spermatocytes, leading to meiotic arrest at metaphase I. Thus, α4s is required for histone degradation at meiotic DNA damage loci, proper progression of meiosis, and fertility in males by promoting proper formation of spermatoproteasomes. These results are important for understanding male infertility and might provide potential targets for male contraception or treatment of male infertility.
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Reparación del ADN , Histonas/metabolismo , Infertilidad Masculina/patología , Meiosis , Complejo de la Endopetidasa Proteasomal/metabolismo , Espermatocitos/citología , Espermatogénesis , Animales , Daño del ADN , Infertilidad Masculina/etiología , Infertilidad Masculina/metabolismo , Masculino , Ratones , Ratones Endogámicos C57BL , Complejo de la Endopetidasa Proteasomal/genética , Espermátides , Espermatocitos/metabolismoRESUMEN
In this report, the impact of different mesa designs on the optical and electrical characteristics for GaN-based micro-light emitting diodes (µLEDs) has been systematically and numerically investigated by using TCAD simulation tools. Our results show that an enhanced light extraction efficiency can be obtained by using beveled mesas. The inclined mesa angles can more effectively reflect the photons to the substrate, and this helps to extract the photons to free air for flip-chip µLEDs. However, it is found that the current injection is influenced by inclination angles for the investigated µLEDs, such that the beveled mesas make stronger charge-coupling effect and increase the electric field magnitude in the multiple quantum wells at the mesa edge, so that the carriers cannot be effective consumed by radiative recombination. As a result, this gives rise to stronger defect-induced nonradiative recombination at mesa surfaces. Therefore, there are tradeoffs between the LEEs and IQEs when changing the beveled angle, to maximize external quantum efficiency for GaN-based µLEDs, the beveled mesa angle shall be carefully designed and optimized.
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Low light extraction efficiency (LEE), high forward voltage and severe self-heating effect greatly affect the performance for AlGaN-based deep ultraviolet light-emitting diodes (DUV LEDs). In this work, surface-textured Ga-face n-AlGaN is fabricated low-costly using self-assembled SiO2 nanosphere as hard mask. The experimental results manifest that when compared with conventional DUV LEDs, the optical power, the forward voltage and the thermal characteristics for the DUV LEDs with surface-textured Ga-face n-AlGaN are improved obviously. It is because the surface-textured Ga-face n-AlGaN between mesa and the n-electrode can be used as the scattering center for trapped light, and this leads to the enhanced LEE. Furthermore, thanks to the surface-textured n-AlGaN under the n-electrode, the n-type ohmic contact area can be increased effectively. Therefore, the n-type ohmic contact resistance can be reduced and the better heat dissipation can be attained for the proposed flip-chip DUV LED.
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In this report, a p+-GaN/SiO2/Ni tunnel junction with a local SiO2 insulation layer is designed to manage the current distribution for commercially structured AlGaN-based deep-ultraviolet light-emitting diodes (DUV LEDs) with a thin p-GaN layer. The experimental and calculated results prove that, besides the increased hole injection at the p+-GaN/SiO2/Ni tunnel junction, the local SiO2 layer produces an in-plane unbalanced energy band in the p-GaN layer for the proposed DUV LEDs, thus modulating the carrier transport paths and increasing the spread of holes. Enhanced optical power is obtained when compared to conventional DUV LEDs. In addition, the influence of the position of the SiO2 insulation layer on the current distribution is also investigated in this work. Placing the SiO2 insulation layer in the middle position of the p+-GaN layer is most helpful for increasing the hole injection efficiency for commercially structured DUV LEDs.
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In this work, a 280-nm-wavelength deep-ultraviolet light-emitting diode (DUV LED) with a p+-GaN/SiO2/ITO tunnel junction is fabricated and investigated. Due to the decreased tunnel region width and enhanced electric field intensity in the 1-nm-thick SiO2 layer, the interband tunneling efficiency and the corresponding hole injection efficiency are promoted. Therefore, the external quantum efficiency (EQE) for the proposed device is increased when compared with a traditional DUV LED. In addition, an improved current spreading effect is observed for our proposed device. As a result, improved wall-plug efficiency (WPE) is obtained owing to the increased optical power and decreased forward operating voltage. Meanwhile, the enhanced electric field intensity in the SiO2 layer reduces the voltage drop in the p-n junction region for the proposed device, and thus the leakage current is reduced.
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In this work, we have proposed and fabricated a metal/Ga2O3/GaN hybrid structure metal-semiconductor-metal ultraviolet photodetector with low dark current and high responsivity. The Schottky contact of Ni/Ga2O3 makes the Ga2O3 layer fully depleted. The strong electric field in the Ga2O3 depletion region can push the photo-induced electrons from the Ga2O3 layer into the GaN layer for more efficient carrier transport. Therefore, the hybrid structure simultaneously utilizes the advantage of the absorption to solar-blind ultraviolet light by the Ga2O3 layer and the high electron mobility of the GaN layer. Thus, the dark current and the photocurrent for the proposed device can be greatly improved. As a result, an extremely high photo-to-dark-current ratio of 1.46 × 106 can be achieved. Furthermore, quick rise and fall times of 0.213â s and 0.027â s at the applied bias of 6â V are also obtained, respectively.
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Small cysteine-rich (SCR) proteins, including fungal avirulence proteins, play important roles in pathogen-plant interactions. SCR protein-encoding genes have been discovered in the genomes of Phytophthora pathogens but their functions during pathogenesis remain obscure. Here, we report the characterization of one Phytophthora capsici SCR protein (namely, SCR82) with similarity to Phytophthora cactorum phytotoxic protein PcF. The scr82 gene has 10 allelic sequences in the P. capsici population. Homologs of SCR82 were not identified in fungi or other organisms but in Phytophthora relative species. Initially, scr82 was weakly expressed during the mycelium, sporangium, and zoospore stages but quickly upregulated when the infection initiated. Both ectopic expression of SCR82 and recombinant yeast-expressed protein (rSCR82) caused cell death on tomato leaves. Upon treatment, rSCR82 induced plant defense responses, including the induction of defense gene expression, reactive oxygen species burst, and callose deposition. Knockout of scr82 in P. capsici by CRISPR/Cas9 severely impaired its virulence on host plants and significantly reduced its resistance against oxidative stress. Inversely, its overexpression increased the pathogen's virulence and tolerance to oxidative stress. Our results collectively demonstrate that SCR82 functions as both an important virulence factor and plant defense elicitor, which is conserved across Phytophthora spp.[Formula: see text] Copyright © 2021 The Author(s). This is an open access article distributed under the CC BY-NC-ND 4.0 International license.
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Phytophthora , Solanum lycopersicum , Cisteína , Enfermedades de las Plantas , Factores de Virulencia/genéticaRESUMEN
In this report, we investigate the impact of a thin p-GaN layer on the efficiency for AlGaN-based deep-ultraviolet light-emitting diodes (DUV LEDs). According to our results, the light extraction efficiency (LEE) becomes higher with the decrease of the p-GaN layer thickness, which can be ascribed to the decreased absorption of DUV emission by the thin p-GaN layer. Moreover, we also find that the variation trend of external quantum efficiency (EQE) is consistent with that of LEE. Therefore, we can speculate that high-efficiency DUV LEDs can be achieved by using thin p-GaN layer to increase the LEE. However, a thin p-GaN layer can also cause severe current crowding effect and the internal quantum efficiency (IQE) will be correspondingly reduced, which will restrict the improvement of EQE. In this work, we find that the adoption of a current spreading layer for such DUV LED with very thin p-GaN layer can facilitate the current spreading effect. For the purpose of demonstration, we then utilize a well-known p-AlGaN/n-AlGaN/p-AlGaN (PNP-AlGaN) structured current spreading layer. Our experimental and numerical results show that, as long as the current crowding effect can be suppressed, the DUV LED with thin p-GaN layer can significantly increase the EQE and the optical power thanks to the enhanced LEE.
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It is known that light extraction efficiency (LEE) for AlGaN-based deep ultraviolet light-emitting didoes (DUV LEDs) can be enhanced by using truncated cone arrays with inclined sidewalls. In this work, the air-cavity-shaped inclined sidewall is applied and the p-GaN layer at the top of the truncated cone is laterally over-etched so that more light escape paths are generated for AlGaN-based DUV LEDs. The experimental results manifest that when compared with DUV LEDs only having the air-cavity-shaped inclined sidewall, the optical power for the DUV LEDs with laterally over-etched p-GaN at the top of the truncated cone is enhanced by 30% without sacrificing the forward bias. It is because the over-etched p-GaN makes little effect on the carrier injection and does not affect the ohmic contact resistance. Moreover, the simulation results show that the truncated cone with laterally over-etched p-GaN layer can enhance the LEE because the reduced p-GaN area can suppress the optical absorption and supplies additional light paths for DUV photos. Then, more light will be reflected into escape cones at the sapphire side.
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Due to the increased surface-to-volume ratio, the surface recombination caused by sidewall defects is a key obstacle that limits the external quantum efficiency (EQE) for GaN-based micro-light-emitting diodes (µLEDs). In this work, we propose selectively removing the periphery p+-GaN layer so that the an artificially formed resistive ITO/p-GaN junction can be formed at the mesa edge. Three types of LEDs with different device dimensions of 30 × 30 µm2, 60 × 60 µm2 and 100 × 100 µm2 are investigated, respectively. We find that such resistive ITO/p-GaN junction can effectively prevent the holes from reaching the sidewalls for µLEDs with smaller size. Furthermore, such confinement of injection current also facilitates the hole injection into the active region for µLEDs. Therefore, the surface-defect-caused nonradiative recombination in the edge of mesa can be suppressed. Meantime, a reduction of current leakage caused by the sidewall defects can also be obtained. As a result, the measured and calculated external quantum efficiency (EQE) and optical output power for the proposed LED with small sizes are increased.
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In this work, a 280 nm AlGaN-based deep ultraviolet light-emitting diode (DUV LED) with a metal-insulator-semiconductor (MIS) structured n-electrode is fabricated and studied. The SiO2 insulator layer is adopted to form the MIS structure by using an atomic layer deposition system. After adopting the MIS-structured n-electrode, the SiO2 intermediate layer enables electron affinity for the contact metal to be higher than the conduction band of the n-AlGaN layer, which favors the electrons to be injected into the n-AlGaN layer by intraband tunneling rather than thermionic emission. Moreover, the thin SiO2 insulator can share the applied bias, which makes the n-AlGaN layer surface less depleted and thus further facilitates the electron injection. The improved electron injection capability at the metal-semiconductor interface helps reduce the contact resistance and increase electron concentration in the active region, which then improves external quantum efficiency and wall-plug efficiency for the proposed DUV LED.