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2.
Phys Chem Chem Phys ; 22(4): 2188-2192, 2020 Jan 29.
Artigo em Inglês | MEDLINE | ID: mdl-31912828

RESUMO

The interplay between multiple chromophores in nucleic acids and photosynthetic proteins gives rise to complex electronic phenomena and largely governs the de-excitation dynamics. Electronic coupling between bases in the excited states of single strands of DNA and RNA may extend over several bases and likely protects nucleic acids from harmful UV damage. Here we report on the coupling between bases in single RNA strands of cytosine and find that the excited state is delocalized over up to five bases at neutral pH, where all bases are non-protonated (i.e. neutral). Delocalization is over four bases at 278 nm excitation, while it involves five bases at shorter wavelengths of 188 nm and 201 nm. This is in contrast to only nearest-neighbour interactions for corresponding DNA strands as previously reported. The current results seemingly corroborate earlier findings of larger spatial communication in RNA than in DNA strands of adenine, but there is no obvious link between the overall structure of strands and delocalization lengths. RNA cytosine strands form a tight helix, while comparatively, adenine strands show less tight packing, also compared to their DNA counterparts, and yet exhibit even higher delocalisation.


Assuntos
Citosina/química , Citosina/efeitos da radiação , RNA/química , RNA/efeitos da radiação , Fenômenos Eletromagnéticos , Conformação de Ácido Nucleico/efeitos da radiação , Raios Ultravioleta
3.
Chem Commun (Camb) ; 56(9): 1317-1324, 2020 Jan 28.
Artigo em Inglês | MEDLINE | ID: mdl-31904034

RESUMO

G-quadruplexes are nucleic acids secondary structures that can be formed in guanine-rich sequences. More than 30 years ago, their formation was first observed in telomeric DNA. Since then, a number of other sequences capable of forming G-quadruplex structures have been described and increasing evidence supporting their formation in the context of living cells has been accumulated. To fully underpin the biological significance of G-quadruplexes and their potential as therapeutic targets, several chemical-biology tools and methods have been developed to map and visualise these nucleic acids secondary structures in human cells. In this review, we critically present the most relevant methods developed to investigate G-quadruplex prevalence in human cells and to study their biological functions, presenting the next key chemical-biology challenges that need to be addressed to fully unravel G-quadruplex mediated biology and their therapeutic potential.


Assuntos
DNA/química , Quadruplex G , Genoma Humano , Sondas Moleculares/química , RNA/química , DNA/genética , Humanos , Ligações de Hidrogênio , RNA/genética
4.
Chem Commun (Camb) ; 56(9): 1365-1368, 2020 Jan 28.
Artigo em Inglês | MEDLINE | ID: mdl-31909398

RESUMO

New methods to control local RNA methylation are needed to elucidate the function of individual m6A. Here, fusion proteins between the programmable RNA binding protein PUF and the m6A demethylase FTO or METTL14 methyltransferase were designed. FTO-PUFs and METTL14-PUFs showed sequence-specific RNA demethylation and methylation activities, respectively.


Assuntos
Desmetilação , Proteínas de Ligação a RNA/metabolismo , RNA/metabolismo , Metilação , RNA/química , Proteínas de Ligação a RNA/química
5.
Chemistry ; 26(1): 102-113, 2020 Jan 02.
Artigo em Inglês | MEDLINE | ID: mdl-31454110

RESUMO

Ribonucleic acid oligonucleotides (RNAs) play pivotal roles in cellular function (riboswitches), chemical biology applications (SELEX-derived aptamers), cell biology and biomedical applications (transcriptomics). Furthermore, a growing number of RNA forms (long non-coding RNAs, circular RNAs) but also RNA modifications are identified, showing the ever increasing functional diversity of RNAs. To describe and understand this functional diversity, structural studies of RNA are increasingly important. However, they are often more challenging than protein structural studies as RNAs are substantially more dynamic and their function is often linked to their structural transitions between alternative conformations. NMR is a prime technique to characterize these structural dynamics with atomic resolution. To extend the NMR size limitation and to characterize large RNAs and their complexes above 200 nucleotides, new NMR techniques have been developed. This Minireview reports on the development of NMR methods that utilize detection on low-γ nuclei (heteronuclei like 13 C or 15 N with lower gyromagnetic ratio than 1 H) to obtain unique structural and dynamic information for large RNA molecules in solution. Experiments involve through-bond correlations of nucleobases and the phosphodiester backbone of RNA for chemical shift assignment and make information on hydrogen bonding uniquely accessible. Previously unobservable NMR resonances of amino groups in RNA nucleobases are now detected in experiments involving conformational exchange-resistant double-quantum 1 H coherences, detected by 13 C NMR spectroscopy. Furthermore, 13 C and 15 N chemical shifts provide valuable information on conformations. All the covered aspects point to the advantages of low-γ nuclei detection experiments in RNA.


Assuntos
Ressonância Magnética Nuclear Biomolecular , RNA/química , Isótopos de Carbono/química , Marcação por Isótopo , Isótopos de Nitrogênio/química , Prótons
6.
Bioresour Technol ; 298: 122534, 2020 Feb.
Artigo em Inglês | MEDLINE | ID: mdl-31835200

RESUMO

Granular activated carbon (GAC) has been shown to mediate direct interspecies electron transfer (DIET) in anaerobic digestion. Adding GAC to up-flow anaerobic sludge bed reactor increased the total biomass slightly from 20.0 to 26.6 gVSS/reactor, and maximum organic removal capacity remarkably from 285 to 1660 mgCOD/L/d. Since GAC occupied 7% of reactor volume (denser than suspended sludge, settled to the reactor bottom), we used a spatial sampling strategy (sludge bed top/mid/bottom layers, and tightly attached GAC-biofilm) and DNA- and RNA-based community analyses. RNA-based analysis demonstrated significant community differences between the non-GAC and GAC-amended reactors (p < 0.05) based on ANOSIM statistical analysis. In comparison, DNA-based analysis showed little community difference between these reactors (p > 0.05). RNA-based analysis revealed active enrichments in GAC-biofilm, including bacteria Geobacter, Syntrophus, Desulfovibrio and Blvii28, and archaea Methanosaeta and Methanospirillum. These are potential electro-active syntrophic microorganisms related with DIET, which expand the previously defined list of DIET microorganisms.


Assuntos
Reatores Biológicos , Metano , Anaerobiose , Elétrons , RNA , Esgotos
7.
Arch Microbiol ; 202(1): 105-114, 2020 Jan.
Artigo em Inglês | MEDLINE | ID: mdl-31485711

RESUMO

Escherichia coli MazF is a toxin protein that cleaves RNA at ACA sequences. Its activation has been thought to cause growth inhibition, primarily through indiscriminate cleavage of RNA. To investigate responses following MazF activation, transcriptomic profiles of mazF-overexpressing and non-overexpressing E. coli K12 cells were compared. Analyses of differentially expressed genes demonstrated that the presence and the number of ACA trimers in RNA was unrelated to cellular RNA levels. Mapping differentially expressed genes onto the chromosome identified two chromosomal segments in which upregulated genes formed clusters, and these segments were absent in the chromosomes of E. coli strains other than K12. These results suggest that MazF regulates selective, rather than indiscriminate, categories of genes, and is involved in the regulation of horizontally acquired genes. We conclude that the primary role of MazF is not only cleaving RNA indiscriminately but also generating a specific cellular state.


Assuntos
Proteínas de Ligação a DNA/metabolismo , Endorribonucleases/metabolismo , Proteínas de Escherichia coli/genética , Proteínas de Escherichia coli/metabolismo , Escherichia coli/genética , Escherichia coli/metabolismo , Regulação Bacteriana da Expressão Gênica/genética , RNA/genética , Proteínas de Ligação a DNA/genética , Endorribonucleases/genética , Escherichia coli K12/genética , Escherichia coli K12/metabolismo , RNA/química
8.
Acta Neurochir Suppl ; 127: 105-119, 2020.
Artigo em Inglês | MEDLINE | ID: mdl-31407071

RESUMO

The protein kinase RNA-like endoplasmic reticulum kinase (PERK) pathway, which is a branch of the unfolded protein response, participates in a range of pathophysiological processes of neurological diseases. However, few studies have investigated the role of the PERK in intracerebral hemorrhage (ICH). The present study evaluated the role of the PERK pathway during the early phase of ICH-induced secondary brain injury (SBI) and its potential mechanisms. An autologous whole blood ICH model was established in rats, and cultured primary cortical neurons were treated with oxyhemoglobin to mimic ICH in vitro. We found that levels of phosphorylated alpha subunit of eukaryotic translation initiation factor 2 (p-eIF2α) and activating transcription factor 4 (ATF4) increased significantly and peaked at 12 h during the early phase of the ICH. To further elucidate the role of the PERK pathway, we assessed the effects of the PERK inhibitor, GSK2606414, and the eIF2α dephosphorylation antagonist, salubrinal, at 12 h after ICH both in vivo and in vitro. Inhibition of PERK with GSK2606414 suppressed the protein levels of p-eIF2α and ATF4, resulting in increase of transcriptional activator CCAAT/enhancer-binding protein homologous protein (CHOP) and caspase-12, which promoted apoptosis and reduced neuronal survival. Treatment with salubrinal yielded opposite results, which suggested that activation of the PERK pathway could promote neuronal survival and reduce apoptosis. In conclusion, the present study has demonstrated the neuroprotective effects of the PERK pathway during the early phase of ICH-induced SBI. These findings highlight the potential value of PERK pathway as a therapeutic target for ICH.


Assuntos
Lesões Encefálicas , Hemorragia Cerebral , RNA , eIF-2 Quinase , Animais , Lesões Encefálicas/metabolismo , Hemorragia Cerebral/metabolismo , Fator de Iniciação 2 em Eucariotos , Ratos , eIF-2 Quinase/metabolismo
9.
Gene ; 724: 144150, 2020 Jan 15.
Artigo em Inglês | MEDLINE | ID: mdl-31589961

RESUMO

Ovarian cancer (OC) is the deadliest form of gynecologic malignancy, with the majority of patients being diagnosed only once the disease reaches an advanced stage owing to a lack of available biomarkers capable of accurately detecting the disease. Stable circular RNAs (circRNAs) can be found at high levels in exosomes, and there is evidence to suggest that they may be viable diagnostic biomarkers for certain cancers. However, circRNAs in the serum of OC patients have rarely been evaluated to date. We therefore sought to investigate serum circRNA profiles of OC patients, and to explore whether these sorts of circRNAs could be used to detect early OC, serving as biomarkers of disease that may allow for the earlier treatment thereof. Second-generation sequencing was used to screen differentially expressed circRNAs in OC patient serum and also in the serum obtained from healthy controls, and circRNA expression was confirmed by qPCR. A bioinformatics-based approach was then used to assess what biological functions might be affected be the altered regulation of these RNA molecules. We further conducted GO, KEGG, and network analyses to further explore the expression of circRNAs. We detected 178 differentially expressed circRNAs in OC patient serum, of which 175 were up-regulated and 3 were down-regulated. We validated 5 of these identified circRNAs by qPCR to confirm their expression, and further found these RNAs to be closely linked with FC gamma R-mediated phagocytosis, VEGF signaling, Transcriptional misregulation in cancer, Chemokine signaling, ErbB signaling, and TNF signaling based on conducted analyses. This study provides a profile of circRNAs in OC patient serum, revealing a pattern of dysregulation of these RNAs associated with OC. Our bioinformatics analysis suggested that these circRNAs are likely related to OC development, and as such they may be viable novel OC biomarkers.


Assuntos
Regulação Neoplásica da Expressão Gênica , Neoplasias Ovarianas/genética , RNA/sangue , Sítios de Ligação , Biomarcadores Tumorais/sangue , Biomarcadores Tumorais/genética , Estudos de Casos e Controles , Biologia Computacional/métodos , Receptores ErbB/genética , Receptores ErbB/metabolismo , Feminino , Perfilação da Expressão Gênica , Ontologia Genética , Redes Reguladoras de Genes , Humanos , RNA/genética , RNA Mensageiro/metabolismo , Reação em Cadeia da Polimerase em Tempo Real , Reprodutibilidade dos Testes , Regulação para Cima , Fator A de Crescimento do Endotélio Vascular/genética , Fator A de Crescimento do Endotélio Vascular/metabolismo
10.
Gene ; 725: 144170, 2020 Jan 30.
Artigo em Inglês | MEDLINE | ID: mdl-31647996

RESUMO

Caragana korshinskii Kom. is a legume shrub that is widely distributed across desert habitats with gravely, sandy, and saline soils in Asia and Africa. C. korshinskii has highly developed roots and a strong tolerance to abiotic stress. At present, there are few genetic studies of C. korshinskii because of the limited availability of genomic resources. To understand the comprehensive mechanisms that are associated with drought tolerance, we used RNA-seq to survey the differentially expressed genes (DEGs) in comparisons of drought-treated and control plants. After analysing the sequencing results, we found 440 differentially expressed genes existing in drought-treated and control plants. Among the DEGs, 39 unigenes showed up-regulated expression after drought treatment, while 401 unigenes were down-regulated. We used the KEGG database to annotate these drought-induced genes; 126 unigenes were identified by KEGG pathway annotation, and approximately 28% of the unigenes with known function fell into categories related to fatty acid metabolism, starch, sucrose metabolism, and nitrogen metabolism, suggesting that these pathways or processes may be involved in the drought response. Finally, we confirmed that one gene has a potential function in drought tolerance. Our study is the first to provide transcriptomic resources for Caragana korshinskii and to determine its digital gene expression profile under conditions of drought stress using the assembled transcriptomic data for reference. These data provide a valuable resource for genetic and genomic studies of desert plants under abiotic stress conditions.


Assuntos
Caragana/genética , Perfilação da Expressão Gênica/métodos , Estresse Fisiológico/genética , Secas , Fabaceae/genética , Regulação da Expressão Gênica de Plantas/genética , Sequenciamento de Nucleotídeos em Larga Escala/métodos , Anotação de Sequência Molecular , RNA/genética , Transcriptoma/genética
12.
J Agric Food Chem ; 67(49): 13737-13750, 2019 Dec 11.
Artigo em Inglês | MEDLINE | ID: mdl-31789024

RESUMO

Genistein is abundant in animal feed. In this study, the side effects of high-dose genistein on intestinal health and hypothalamic RNA profile were evaluated. Chicks exposed to high-dose genistein by intraperitoneal injection (416 ± 21, 34.5 ± 2.5) and feed supplementation (308 ± 19, 27.2 ± 2.1) both showed a reduced body weight gain and feed intake in comparison with the control group (261 ± 16, 22.7 ± 1.6, P < 0.01). In comparison with the control (22.4 ± 0.5, 33.3 ± 2.4), serum levels of albumin and total protein were decreased after high-dose genistein injection (21.6 ± 0.5, 31.8 ± 1.6) and diet supplementation (20.6 ± 0.9, 29.9 ± 2.5, P < 0.001). Interestingly, the genistein diet presented the chick hypothalamus with downregulated expression of bitter receptors (TAS1R3, P < 0.05). Meanwhile, it upregulated the expressions of TAS2R1 (P < 0.05) and downstream genes (PLCB2 and IP3R3) in the ileum (P < 0.05). Accordingly, high-dose dietary genistein reduced villus height and the abundance of Lactobacillus, along with the increased abundance of pathogenic bacteria in the ileum (P < 0.05). Furthermore, transcriptomic analysis identified 348 differently expressed genes (168 upregulated and 224 downregulated) in the high-dose dietary genistein treated group in comparison with the control (P < 0.05, |log2FoldChange| > 0.585). Therefore, high-dose dietary genistein altered the hypothalamic RNA profile and signal processing. Cluster analysis further revealed that high-dose dietary genistein significantly influenced apoptosis, the immune process, and the whole synthesis of steroid hormones in the hypothalamus (P < 0.05). In conclusion, high-dose dietary genistein altered the hypothalamic RNA profile and intestinal health of female chicks.


Assuntos
Galinhas/metabolismo , Suplementos Nutricionais/efeitos adversos , Genisteína/efeitos adversos , Hipotálamo/metabolismo , RNA/genética , Ração Animal/efeitos adversos , Ração Animal/análise , Animais , Peso Corporal/efeitos dos fármacos , Galinhas/genética , Galinhas/crescimento & desenvolvimento , Galinhas/imunologia , Ingestão de Alimentos/efeitos dos fármacos , Feminino , Genisteína/análise , Hipotálamo/efeitos dos fármacos , Íleo/efeitos dos fármacos , Íleo/metabolismo , RNA/metabolismo , Esteroides/metabolismo
14.
Adv Exp Med Biol ; 1203: 113-132, 2019.
Artigo em Inglês | MEDLINE | ID: mdl-31811632

RESUMO

The RNA exosome is a highly conserved ribonuclease endowed with 3'-5' exonuclease and endonuclease activities. The multisubunit complex resides in both the nucleus and the cytoplasm, with varying compositions and activities between the two compartments. While the cytoplasmic exosome functions mostly in mRNA quality control pathways, the nuclear RNA exosome partakes in the 3'-end processing and complete decay of a wide variety of substrates, including virtually all types of noncoding (nc) RNAs. To handle these diverse tasks, the nuclear exosome engages with dedicated cofactors, some of which serve as activators by stimulating decay through oligoA addition and/or RNA helicase activities or, as adaptors, by recruiting RNA substrates through their RNA-binding capacities. Most nuclear exosome cofactors contain the essential RNA helicase Mtr4 (MTR4 in humans). However, apart from Mtr4, nuclear exosome cofactors have undergone significant evolutionary divergence. Here, we summarize biochemical and functional knowledge about the nuclear exosome and exemplify its cofactor variety by discussing the best understood model organisms-the budding yeast Saccharomyces cerevisiae, the fission yeast Schizosaccharomyces pombe, and human cells.


Assuntos
Coenzimas , Complexo Multienzimático de Ribonucleases do Exossomo , RNA Nuclear , Coenzimas/metabolismo , RNA Helicases DEAD-box/metabolismo , Complexo Multienzimático de Ribonucleases do Exossomo/metabolismo , Humanos , RNA/metabolismo , RNA Nuclear/metabolismo , RNA não Traduzido/metabolismo , Saccharomyces cerevisiae , Proteínas de Saccharomyces cerevisiae/metabolismo
15.
Nihon Yakurigaku Zasshi ; 154(6): 294-300, 2019.
Artigo em Japonês | MEDLINE | ID: mdl-31787679

RESUMO

The most common form of DNA is a right-handed helix, the B-form DNA. DNA can also adopt a variety of alternative conformations, termed non-B-form DNA secondary structures, including the G-quadruplex (G4). Furthermore, non-canonical RNA G4 secondary structures are also observed. Recent bioinformatics analysis revealed genomic positions of G4. In addition, G4 formation may be associated with various biological functions, including DNA replication, transcription, epigenetic modification, and RNA metabolism. In this review, we focus on G4 structures in neuronal functions, which may have important roles reveal mechanisms underlying neurological disorders. In addition, we discuss the potential of G4s as a therapeutic target for neurological diseases.


Assuntos
Quadruplex G , Terapia de Alvo Molecular , Doenças do Sistema Nervoso/tratamento farmacológico , DNA , Humanos , Conformação de Ácido Nucleico , RNA
16.
Adv Exp Med Biol ; 1185: 71-77, 2019.
Artigo em Inglês | MEDLINE | ID: mdl-31884591

RESUMO

Inherited retinal dystrophies (IRDs) are genetic diseases affecting 1 in every 3000 individuals worldwide. Nowadays, more than 250 genes have been associated with different forms of IRD. In the last decade, it has been shown that gene therapy is a promising approach to correct the genetic defects underlying IRD. In fact, voretigene neparvovec-rzyl (Luxturna™), the first commercialized gene therapy drug to treat RPE65-associated Leber congenital amaurosis, has opened new venues. However, IRDs are highly heterogeneous at genetic level making the design of novel strategies complicated. Unfortunately, the size of several frequently mutated genes is not suitable for the approved conventional therapeutic viral vectors; therefore, there is an urgent need for the development of alternatives, such as those targeting the pre-mRNA. In this mini-review, the potential of RNA-based strategies for IRDs is discussed.


Assuntos
Terapia Genética , RNA/uso terapêutico , Distrofias Retinianas/terapia , Vetores Genéticos , Humanos , Amaurose Congênita de Leber/terapia , Distrofias Retinianas/genética
17.
Adv Exp Med Biol ; 1185: 85-89, 2019.
Artigo em Inglês | MEDLINE | ID: mdl-31884593

RESUMO

Inherited retinal diseases (IRD) encompass a wide spectrum of hereditary blindness with significant genetic heterogeneity. Therapeutics regulating gene expression on an RNA level have significant promise for treating IRD. In this review, we review the molecular basis of oligonucleotide therapeutics such as ribozymes, RNA interference (RNAi), antisense oligonucleotides (ASO), CRISPRi/a, and their applications to treatments of IRD.


Assuntos
RNA/uso terapêutico , Doenças Retinianas/terapia , Repetições Palindrômicas Curtas Agrupadas e Regularmente Espaçadas , Humanos , Oligonucleotídeos Antissenso/uso terapêutico , Interferência de RNA , RNA Catalítico/uso terapêutico
18.
Sheng Wu Yi Xue Gong Cheng Xue Za Zhi ; 36(6): 1038-1042, 2019 Dec 25.
Artigo em Chinês | MEDLINE | ID: mdl-31875380

RESUMO

Circular RNA (circRNA) is a type of single-stranded RNA that binds in a closed loop structure by covalent bond. It is highly expressed and has diverse functions in the eukaryotic transcriptome, and it also has the potential to regulate the process of cell differentiation. Stem cells are important seed cells and common research tools in the field of tissue engineering, which have multi-directional differentiation potential and low immunogenicity. Its clinical application for the treatment of diseases has broad prospects, and the research on their differentiation mechanism has gradually penetrated to the molecular level. A number of studies have shown that circRNA participates in stem cell differentiation and plays a key role through a variety of pathways. This article focuses on the expression changes of circRNA during stem cell differentiation and its research advancement in regulating the differentiation mechanism of various stem cells. The review also prospects its possible role in tissue regeneration and repair, in order to further study the molecular mechanism of circRNA involved in stem cell differentiation and provide ideas for clinical practice of stem cells in biomedical engineering.


Assuntos
Diferenciação Celular , Engenharia Tecidual , RNA , Células-Tronco , Transcriptoma
19.
BMC Bioinformatics ; 20(Suppl 24): 596, 2019 Dec 22.
Artigo em Inglês | MEDLINE | ID: mdl-31861975

RESUMO

BACKGROUND: Adenosine-to-inosine RNA editing can markedly diversify the transcriptome, leading to a variety of critical molecular and biological processes in mammals. Over the past several years, researchers have developed several new pipelines and software packages to identify RNA editing sites with a focus on downstream statistical analysis and functional interpretation. RESULTS: Here, we developed a user-friendly public webserver named MIRIA that integrates statistics and visualization techniques to facilitate the comprehensive analysis of RNA editing sites data identified by the pipelines and software packages. MIRIA is unique in that provides several analytical functions, including RNA editing type statistics, genomic feature annotations, editing level statistics, genome-wide distribution of RNA editing sites, tissue-specific analysis and conservation analysis. We collected high-throughput RNA sequencing (RNA-seq) data from eight tissues across seven species as the experimental data for MIRIA and constructed an example result page. CONCLUSION: MIRIA provides both visualization and analysis of mammal RNA editing data for experimental biologists who are interested in revealing the functions of RNA editing sites. MIRIA is freely available at https://mammal.deepomics.org.


Assuntos
Mamíferos , Edição de RNA , Análise de Sequência de RNA , Transcriptoma , Animais , Genoma , Genômica , Sequenciamento de Nucleotídeos em Larga Escala/métodos , Humanos , Mamíferos/genética , RNA/genética , Análise de Sequência de RNA/métodos
20.
Mol Biol (Mosk) ; 53(6): 924-932, 2019.
Artigo em Russo | MEDLINE | ID: mdl-31876273

RESUMO

Mitochondria of many living species internalize nuclear DNA-encoded ribonucleic acids. The pools of imported RNA molecules, as well as fine mechanisms of these processes, are highly species-specific. To date, baker's yeast Saccharomyces cerevisiae are the best studied in this regard. Moreover, the processes of yeast RNA mitochondrial import have been the basis of modeling several gene therapy strategies aimed to palliate negative effects of pathogenic mutations in human mitochondrial DNA. In this review, we summarize our current knowledge about the molecular events taking place in course of yeast RNA import into mitochondria. Also, we describe how this process can be used for compensation of pathogenic mutations in mitochondrial genomes of humans.


Assuntos
Terapia Genética/tendências , Mitocôndrias/genética , Mitocôndrias/metabolismo , RNA/metabolismo , DNA Mitocondrial/genética , Humanos , Saccharomyces cerevisiae/citologia , Saccharomyces cerevisiae/genética , Saccharomyces cerevisiae/metabolismo
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