scholarly journals The PyPIF5-PymiR156a-PySPL9-PyMYB114/MYB10 module regulates light-induced anthocyanin biosynthesis in red pear

2021 ◽  
Vol 1 (1) ◽  
Author(s):  
Hainan Liu ◽  
Qun Shu ◽  
Kui Lin-Wang ◽  
Andrew C. Allan ◽  
Richard V. Espley ◽  
...  
Keyword(s):  
Small Rna ◽  
Environmental Conditions ◽  
Anthocyanin Biosynthesis ◽  
Inhibitory Effect ◽  
Blocking Effect ◽  
Anthocyanin Accumulation ◽  
Rna Seq ◽  
Fruit Skin ◽  
Regulation Network ◽  
The Difference

AbstractSome cultivars of pear (Pyrus L.) show attractive red fruit skin due to anthocyanin accumulation. This pigmentation can be affected by environmental conditions, especially light. To explore the light-induced regulation network for anthocyanin biosynthesis and fruit coloration in pear, small RNA libraries and mRNA libraries from fruit skins of ‘Yunhongyihao’ pear were constructed to compare the difference between bagging and debagging treatments. Analysis of RNA-seq of fruit skins with limited light (bagged) and exposed to light (debagged), showed that PyPIF5 was down-regulated after bag removal. PymiR156a was also differentially expressed between bagged and debagged fruit skins. We found that PyPIF5 negatively regulated PymiR156a expression in bagged fruits by directly binding to the G-box motif in its promoter. In addition, PymiR156a overexpression promoted anthocyanin accumulation in both pear skin and apple calli. We confirmed that PymiR156a mediated the cleavage of PySPL9, and that the target PySPL9 protein could form heterodimers with two key anthocyanin regulators (PyMYB114/PyMYB10). We proposed a new module of PyPIF5-PymiR156a-PySPL9-PyMYB114/MYB10. When the bagged fruits were re-exposed to light, PyPIF5 was down-regulated and its inhibitory effect on PymiR156a was weakened, which leads to degradation of the target PySPL, thus eliminating the blocking effect of PySPL on the formation of the regulatory MYB complexes. Ultimately, this promotes anthocyanin biosynthesis in pear skin.

scholarly journals MdBBX21, a B-Box Protein, Positively Regulates Light-Induced Anthocyanin Accumulation in Apple Peel

2021 ◽  
Vol 12 ◽  
Author(s):  
Bo Zhang ◽  
Zhen-Zhen Zhu ◽  
Dong Qu ◽  
Bo-Chen Wang ◽  
Ni-Ni Hao ◽  
...  
Keyword(s):  
Anthocyanin Biosynthesis ◽  
Apple Fruit ◽  
Bimolecular Fluorescence Complementation ◽  
Anthocyanin Accumulation ◽  
Rna Seq ◽  
Yeast Two Hybrid ◽  
Fruit Peel ◽  
Apple Peel ◽  
Its Gene ◽  
Two Hybrid

The red coloration of apple (Malus × domestica Borkh.) is due to the accumulation of anthocyanins in the fruit peel. Light is essential for anthocyanin biosynthesis in apple. In this study, we performed a transcriptome sequencing (RNA-seq) analysis of apple fruit exposed to light after unbagging. The identified differentially expressed genes included MdBBX21, which is homologous to Arabidopsis BBX21, suggesting it may be involved in light-induced anthocyanin biosynthesis. Additionally, MdBBX21 was localized in the nucleus and its gene was expressed earlier than MdMYB1 in apple peel treated with light. Overexpressing MdBBX21 in Arabidopsis and apple calli under light increased anthocyanin accumulation. Dual-luciferase and yeast one-hybrid assays confirmed that MdBBX21 binds to the MdHY5, MdBBX20, and MdBBX22-1/2 promoters and induces expression. At the same time, MdHY5 can also activate the expression of MdBBX21. Furthermore, bimolecular fluorescence complementation and yeast two-hybrid assays demonstrated that MdBBX21 can interact with MdHY5. This interaction can significantly enhance MdMYB1 promoter activity. These findings clarify the molecular mechanism by which MdBBX21 positively regulates light-induced anthocyanin accumulation in apple.

scholarly journals The role of UV-B light on small RNA activity during grapevine berry development

2018 ◽  
Author(s):  
Sukumaran Sunitha ◽  
Rodrigo Loyola ◽  
José Antonio Alcalde ◽  
Patricio Arce-Johnson ◽  
José Tomás Matus ◽  
...  
Keyword(s):  
Small Rna ◽  
Anthocyanin Biosynthesis ◽  
Transcriptional Gene Silencing ◽  
Berry Development ◽  
Anthocyanin Accumulation ◽  
High Fluence ◽  
Gene Expressions ◽  
Time Points ◽  
Grapevine Berry

AbstractUV-B regulation of anthocyanin biosynthesis in vegetative and grapevine berry tissues has been extensively described. However, its relation with UV-B-regulated microRNAs (miRNAs) has not been addressed before in this species. We explored by deep sequencing of small RNA libraries the developmental dynamics and UV-B effects on miRNAs and associated phased small interfering RNA (phasi-RNAs)-producing loci abundances inin vitro-grown plantlets, in field-grown berry skins of cv. Cabernet Sauvignon, and low- and high UV-B fluence treatments of greenhouse-grown berries at several time points around veraison. We observed by RNA blotting a differential effect of low-versus high-fluence UV-B on miR828 abundances (an effector of anthocyanins and UV-absorbing polyphenolics) across berry development, and identified other miRNAs that correlated with miR828 dynamics. The functional significance of the observed UV-coordinated miRNA responses to UV was supported by degradome evidences of AGO-programmed slicing of mRNAs. Inverse co-expression of the up-regulated miRNAs miR156, miR482, miR530, and miR828 with cognate target gene expressions in response to high fluence UV-B measured by quantitative real-time PCR. These UV-response relationships were also corroborated by analyzing three published transcriptome datasets (berries subjected to UV-C for 1 hr [at pre-veraison], UV-B for five weeks post-veraison, and five red-skinned varieties across four berry development time points). Based on observed significant changes by UV-B on miRNA and derivative phasi-RNA abundances, we propose a regulatory network model of UV responses impacting anti-oxidant and stress-associated polyphenolic compound biosynthesis. In this model high-fluence UV-B increases miR168 (validated in a UV-B small RNA-derived degradome library to targetARGONAUTE1, which spawns phasi-RNAs) and miR530 (targets a novel Plus-3 domain mRNA), while decreasing miR403 abundances (validated to targetARGONAUTE2), thereby coordinating post-transcriptional gene silencing activities by different AGOs. Up-regulation of miR3627/4376 (validated to target Ca2+-transporting ATPase10 that spawns phasi-RNAs) could facilitate anthocyanin accumulation. miR395 and miR399, induced by sulfur and phosphorus starvation in other species (conditions known to trigger anthocyanin accumulation) respond positively to UV-B radiation and are shown to slice cognate targets in grapevine. miR156/miR535 is shown to targetSQUAMOSA PROMOTER-BINDINGtranscription factor genes that potentially regulate the activities of MYB-bHLH-WD40 complexes and thereby anthocyanin biosynthesis. Increases in MYB-bHLH-WD40 TFs could also contribute to the observed up-regulation of miR828 via the conserved and degradome-validated auto-regulatory loop involving miR828/TAS4abcto regulateMYBA6/A7/A5-MYB113-likelevels and thereby anthocyanin levels. These results and meta-analysis provide a basis for systems approaches to better understand non-coding RNA functions in response to UV.

MYB_SH[AL]QKY[RF] transcription factors MdLUX and MdPCL-like promote anthocyanin accumulation through DNA hypomethylation and MdF3H activation in apple

2020 ◽  
Author(s):  
Wen-Fang Li ◽  
Gai-Xing Ning ◽  
Cun-Wu Zuo ◽  
Ming-Yu Chu ◽  
Shi-Jin Yang ◽  
...  
Keyword(s):  
Dna Methylation ◽  
Transcription Factors ◽  
Promoter Region ◽  
Anthocyanin Biosynthesis ◽  
Apple Fruit ◽  
Anthocyanin Synthesis ◽  
Mrna Levels ◽  
Anthocyanin Accumulation ◽  
Dna Hypomethylation ◽  
Fruit Skin

Abstract Heritable DNA methylation is a highly conserved epigenetic mark that is important for many biological processes. In a previous transcriptomic study on the fruit skin pigmentation of apple (Malus domestica Borkh.) cv. ‘Red Delicious’ (G0) and its four continuous-generation bud sport mutants including ‘Starking Red’ (G1), ‘Starkrimson’ (G2), ‘Campbell Redchief’ (G3) and ‘Vallee spur’ (G4), we identified MYB transcription factors (TFs) MdLUX and MdPCL-like involved in regulating anthocyanin synthesis. However, how these TFs ultimately determine the fruit skin colour traits remain elusive. Here, bioinformatics analysis revealed that MdLUX and MdPCL-like contained a well-conserved motif SH[AL]QKY[RF] in their C-terminal region and were located in the nucleus of onion epidermal cells. Overexpression of MdLUX and MdPCL-like in ‘Golden Delicious’ fruits, ‘Gala’ calli and Arabidopsis thaliana promoted the accumulation of anthocyanin, whereas MdLUX and MdPCL-like suppression inhibited anthocyanin accumulation in ‘Red Fuji’ apple fruit skin. Yeast one-hybrid assays revealed that MdLUX and MdPCL-like may bind to the promoter region of the anthocyanin biosynthesis gene MdF3H. Dual-luciferase assays indicated that MdLUX and MdPCL-like activated MdF3H. The whole-genome DNA methylation study revealed that the methylation levels of the mCG context at the upstream (i.e., promoter region) of MdLUX and MdPCL-like were inversely correlated with their mRNA levels and anthocyanin accumulation. Hence, the data suggest that MYB_SH[AL]QKY[RF] TFs MdLUX and MdPCL-like promote anthocyanin biosynthesis in apple fruit skins through the DNA hypomethylation of their promoter regions and the activation of the structural flavonoid gene MdF3H.

scholarly journals REVEILLE Transcription Factors Contribute to the Nighttime Accumulation of Anthocyanins in ‘Red Zaosu’ (Pyrus Bretschneideri Rehd.) Pear Fruit Skin

2020 ◽  
Vol 21 (5) ◽  
pp. 1634
Author(s):  
Xieyu Li ◽  
Ting Wu ◽  
Hanting Liu ◽  
Rui Zhai ◽  
Yao Wen ◽  
...  
Keyword(s):  
Transcription Factors ◽  
Anthocyanin Biosynthesis ◽  
Expression Patterns ◽  
Luciferase Reporter ◽  
Anthocyanin Accumulation ◽  
Pear Fruit ◽  
Biosynthetic Genes ◽  
Expression Levels ◽  
Fruit Skin ◽  
Genes Encoding

Anthocyanin biosynthesis exhibits a rhythmic oscillation pattern in some plants. To investigate the correlation between the oscillatory regulatory network and anthocyanin biosynthesis in pear, the anthocyanin accumulation and the expression patterns of anthocyanin late biosynthetic genes (ALBGs) were investigated in fruit skin of ‘Red Zaosu’ (Pyrus bretschneideri Rehd.). The anthocyanin accumulated mainly during the night over three continuous days in the fruit skin, and the ALBGs’ expression patterns in ‘Red Zaosu’ fruit skin were oscillatory. However, the expression levels of typical anthocyanin-related transcription factors did not follow this pattern. Here, we found that the expression patterns of four PbREVEILLEs (PbRVEs), members of a class of atypical anthocyanin-regulated MYBs, were consistent with those of ALBGs in ‘Red Zaosu’ fruit skin over three continuous days. Additionally, transient expression assays indicated that the four PbRVEs promoted anthocyanin biosynthesis by regulating the expression of the anthocyanin biosynthetic genes encoding dihydroflavonol-4-reductase (DFR) and anthocyanidin synthase (ANS) in red pear fruit skin, which was verified using a dual-luciferase reporter assay. Moreover, a yeast one-hybrid assay indicated that PbRVE1a, 1b and 7 directly bound to PbDFR and PbANS promoters. Thus, PbRVEs promote anthocyanin accumulation at night by up-regulating the expression levels of PbDFR and PbANS in ‘Red Zaosu’ fruit skin.

scholarly journals Transcriptomic profiling of purple broccoli reveals light-induced anthocyanin biosynthetic signaling and structural genes

PeerJ ◽  
2020 ◽  
Vol 8 ◽  
pp. e8870 ◽  
Author(s):  
Chunqing Liu ◽  
Xueqin Yao ◽  
Guangqing Li ◽  
Lei Huang ◽  
Zhujie Xie
Keyword(s):  
Molecular Mechanisms ◽  
De Novo ◽  
Anthocyanin Biosynthesis ◽  
Digital Gene Expression ◽  
Flower Head ◽  
Anthocyanin Accumulation ◽  
Rna Seq ◽  
Light Conditions ◽  
Structural Genes ◽  
Rna Sequences

Purple Broccoli (Brassica oleracea L. var italica) attracts growing attention as a functional food. Its purple coloration is due to high anthocyanin amounts. Light represents a critical parameter affecting anthocyanins biosynthesis. In this study, ‘Purple Broccoli’, a light-responding pigmentation cultivar, was assessed for exploring the mechanism underlying light-induced anthocyanin biosynthesis by RNA-Seq. Cyanidin, delphinidin and malvidin derivatives were detected in broccoli head samples. Shading assays and RNA-seq analysis identified the flower head as more critical organ compared with leaves. Anthocyanin levels were assessed at 0, 7 and 11 days, respectively, with further valuation by RNA-seq under head-shading and light conditions. RNA sequences were de novo assembled into 50,329 unigenes, of which 38,701 were annotated against four public protein databases. Cluster analysis demonstrated that anthocyanin/phenylpropanoid biosynthesis, photosynthesis, and flavonoid biosynthesis in cluster 8 were the main metabolic pathways regulated by light and had showed associations with flower head growth. A total of 2,400 unigenes showed differential expression between the light and head-shading groups in cluster 8, including 650 co-expressed, 373 specifically expressed under shading conditions and 1,377 specifically expressed under normal light. Digital gene expression (DGE) analysis demonstrated that light perception and the signal transducers CRY3 and HY5 may control anthocyanin accumulation. Following shading, 15 structural genes involved in anthocyanin biosynthesis were downregulated, including PAL, C4H, 4CL, CHS, CHI, F3H and DFR. Moreover, six BoMYB genes (BoMYB6-1, BoMYB6-2, BoMYB6-3, BoMYB6-4, BoMYBL2-1 and BoMYBL2-2) and three BobHLH genes (BoTT8_5-1, BoTT8_5-2 and BoEGL5-3) were critical transcription factors controlling anthocyanin accumulation under light conditions. Based on these data, a light-associated anthocyanin biosynthesis pathway in Broccoli was proposed. This information could help improve broccoli properties, providing novel insights into the molecular mechanisms underpinning light-associated anthocyanin production in purple vegetables.

scholarly journals MicroRNAs May Play an Important Role in Sexual Reversal Process of Chinese Soft-Shelled Turtle, Pelodiscus sinensis

Genes ◽  
2021 ◽  
Vol 12 (11) ◽  
pp. 1696
Author(s):  
Tong Zhou ◽  
Hang Sha ◽  
Meng Chen ◽  
Guobin Chen ◽  
Guiwei Zou ◽  
...  
Keyword(s):  
Small Rna ◽  
Target Genes ◽  
Gonadal Development ◽  
Rna Seq ◽  
Pelodiscus Sinensis ◽  
Female Phenotype ◽  
Reversal Process ◽  
Differentially Expressed Mirnas ◽  
The Difference ◽  
Male To Female

The Chinese soft-shelled (Pelodiscus sinensis) turtle exhibits obvious sex dimorphism, which leads to the higher economic and nutritional value of male individuals. Exogenous hormones can cause the transformation from male to female phenotype during gonadal differentiation. However, the molecular mechanism related to the sexual reversal process is unclear. In this study, we compared the difference between the small RNAs of male, female, and pseudo-female turtles by small RNA-seq to understand the sexual reversal process of Chinese soft-shelled turtles. A certain dose of estrogen can cause the transformation of Chinese soft-shelled turtles from male to female, which are called pseudo-female individuals. The result of small RNA-seq has revealed that the characteristics of pseudo-females are very similar to females, but are strikingly different from males. The number of the microRNAs (miRNAs) of male individuals was significantly less than the number of female individuals or pseudo-female individuals, while the expression level of miRNAs of male individuals were significantly higher than the other two types. Furthermore, we found 533 differentially expressed miRNAs, including 173 up-regulated miRNAs and 360 down-regulated miRNAs, in the process of transformation from male to female phenotype. Cluster analysis of the total 602 differential miRNAs among females, males, and pseudo-females showed that miRNAs played a crucial role during the sexual differentiation. Among these differential miRNAs, we found 12 miRNAs related to gonadal development and verified their expression by qPCR. The TR-qPCR results confirmed the differential expression of 6 of the 12 miRNAs: miR-26a-5p, miR-212-5p, miR-202-5p, miR-301a, miR-181b-3p and miR-96-5p were involved in sexual reversal process, which was consistent with the results of omics. Using these six miRNAs and some of their target genes, we constructed a network diagram related to gonadal development. We suggest that these miRNAs may play an important role in the process of effective sex reversal, which would contribute to the breeding of all male strains of Chinese soft-shelled turtles.

Information support for expert evaluation of the states of the external environment

2020 ◽  
pp. 71-76
Author(s):  
Rasol Murtadha Najah
Keyword(s):  
Decision Making ◽  
Environmental Conditions ◽  
Fuzzy Inference ◽  
Environmental Parameters ◽  
Information Support ◽  
Expert Assessment ◽  
Fuzzy Variables ◽  
Decision Making Models ◽  
The Difference ◽  
State Of The Environment

This article discusses the application of methods to enhance the knowledge of experts to build a decision-making model based on the processing of physical data on the real state of the environment. Environmental parameters determine its ecological state. To carry out research in the field of expert assessment of environmental conditions, the analysis of known works in this field is carried out. The results of the analysis made it possible to justify the relevance of the application of analytical, stochastic models and models based on methods of enhancing the knowledge of experts — experts. It is concluded that the results of using analytical and stochastic objects are inaccurate, due to the complexity and poor mathematical description of the objects. The relevance of developing information support for an expert assessment of environmental conditions is substantiated. The difference of this article is that based on the analysis of the application of expert methods for assessing the state of the environment, a fuzzy logic adoption model and information support for assessing the environmental state of the environment are proposed. The formalization of the parameters of decision-making models using linguistic and fuzzy variables is considered. The formalization of parameters of decision-making models using linguistic and fuzzy variables was considered. The model’s description of fuzzy inference is given. The use of information support for environment state assessment is shown on the example of experts assessing of the land desertification stage.

scholarly journals The Purple Leaf (pl6) Mutation Regulates Leaf Color by Altering the Anthocyanin and Chlorophyll Contents in Rice

Plants ◽  
2020 ◽  
Vol 9 (11) ◽  
pp. 1477
Author(s):  
Asadullah Khan ◽  
Sanaullah Jalil ◽  
Huan Cao ◽  
Yohannes Tsago ◽  
Mustapha Sunusi ◽  
...  
Keyword(s):  
Gamma Ray ◽  
Anthocyanin Biosynthesis ◽  
Light Attenuation ◽  
Transcript Level ◽  
Underlying Mechanism ◽  
Anthocyanin Accumulation ◽  
Morphological Marker ◽  
Chlorophyll Contents ◽  
Rice Mutant ◽  
Purple Coloration

The anthocyanin biosynthesis attracts strong interest due to the potential antioxidant value and as an important morphological marker. However, the underlying mechanism of anthocyanin accumulation in plant tissues is not clearly understood. Here, a rice mutant with a purple color in the leaf blade, named pl6, was developed from wild type (WT), Zhenong 41, with gamma ray treatment. By map-based cloning, the OsPL6 gene was located on the short arm of chromosome 6. The multiple mutations, such as single nucleotide polymorphism (SNP) at −702, −598, −450, an insertion at −119 in the promoter, three SNPs and one 6-bp deletion in the 5′-UTR region, were identified, which could upregulate the expression of OsPL6 to accumulate anthocyanin. Subsequently, the transcript level of structural genes in the anthocyanin biosynthesis pathway, including OsCHS, OsPAL, OsF3H and OsF3′H, was elevated significantly. Histological analysis revealed that the light attenuation feature of anthocyanin has degraded the grana and stroma thylakoids, which resulted in poor photosynthetic efficiency of purple leaves. Despite this, the photoabatement and antioxidative activity of anthocyanin have better equipped the pl6 mutant to minimize the oxidative damage. Moreover, the contents of abscisic acid (ABA) and cytokanin (CK) were elevated along with anthocyanin accumulation in the pl6 mutant. In conclusion, our results demonstrate that activation of OsPL6 could be responsible for the purple coloration in leaves by accumulating excessive anthocyanin and further reveal that anthocyanin acts as a strong antioxidant to scavenge reactive oxygen species (ROS) and thus play an important role in tissue maintenance.

scholarly journals Characteristics of Extratropical Cyclones That Cause Tornadoes in Italy: A Preliminary Study

Atmosphere ◽  
2021 ◽  
Vol 12 (2) ◽  
pp. 180
Author(s):  
Eigo Tochimoto ◽  
Mario Marcello Miglietta ◽  
Leonardo Bagaglini ◽  
Roberto Ingrosso ◽  
Hiroshi Niino
Keyword(s):  
Environmental Conditions ◽  
Convective Available Potential Energy ◽  
Static Stability ◽  
Extratropical Cyclones ◽  
Cool Season ◽  
Southeastern Us ◽  
Different Seasons ◽  
The Difference ◽  
Upper Level ◽  
Strong Evaporation

Characteristics of extratropical cyclones that cause tornadoes in Italy are investigated. Tornadoes between 2007 and 2016 are analyzed, and statistical analysis of the associated cyclone structures and environments is performed using the JRA-55 reanalysis. Tornadoes are distributed sporadically around the cyclone location within a window of 10° × 10°. The difference in the cyclone tracks partially explains the seasonal variability in the distribution of tornadoes. The highest number of tornadoes occur south of the cyclone centers, mainly in the warm sector, while a few are observed along the cold front. Composite mesoscale parameters are examined to identify the environmental conditions associated with tornadoes in different seasons. Potential instability is favorable to tornado development in autumn. The highest convective available potential energy (CAPE) in this season is associated with relatively high-temperature and humidity at low-levels, mainly due to the strong evaporation over the warm Mediterranean Sea. Upper-level potential vorticity (PV) anomalies and the associated cold air reduce the static stability above the cyclone center, mainly in spring and winter. On average, the values of CAPE are lower than for US tornadoes and comparable with those occurring in Japan, while storm relative helicity (SREH) is comparable with US tornadoes and higher than Japanese tornadoes, indicating that the environmental conditions for Italian tornadoes have peculiar characteristics. Overall, the conditions emerging in this study are close to the high-shear, low-CAPE environments typical of cool-season tornadoes in the Southeastern US.

scholarly journals Small RNA-Sequencing: Approaches and Considerations for miRNA Analysis

Diagnostics ◽  
2021 ◽  
Vol 11 (6) ◽  
pp. 964
Author(s):  
Sarka Benesova ◽  
Mikael Kubista ◽  
Lukas Valihrach
Keyword(s):  
Rna Sequencing ◽  
Small Rna ◽  
High Sensitivity ◽  
Small Rna Sequencing ◽  
Rna Seq ◽  
Liquid Biopsies ◽  
Comprehensive Overview ◽  
Rna Molecules ◽  
Novel Mirna ◽  
The Many

MicroRNAs (miRNAs) are a class of small RNA molecules that have an important regulatory role in multiple physiological and pathological processes. Their disease-specific profiles and presence in biofluids are properties that enable miRNAs to be employed as non-invasive biomarkers. In the past decades, several methods have been developed for miRNA analysis, including small RNA sequencing (RNA-seq). Small RNA-seq enables genome-wide profiling and analysis of known, as well as novel, miRNA variants. Moreover, its high sensitivity allows for profiling of low input samples such as liquid biopsies, which have now found applications in diagnostics and prognostics. Still, due to technical bias and the limited ability to capture the true miRNA representation, its potential remains unfulfilled. The introduction of many new small RNA-seq approaches that tried to minimize this bias, has led to the existence of the many small RNA-seq protocols seen today. Here, we review all current approaches to cDNA library construction used during the small RNA-seq workflow, with particular focus on their implementation in commercially available protocols. We provide an overview of each protocol and discuss their applicability. We also review recent benchmarking studies comparing each protocol’s performance and summarize the major conclusions that can be gathered from their usage. The result documents variable performance of the protocols and highlights their different applications in miRNA research. Taken together, our review provides a comprehensive overview of all the current small RNA-seq approaches, summarizes their strengths and weaknesses, and provides guidelines for their applications in miRNA research.

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