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Extracellular RNA (also known as exRNA or exosomal RNA) describes [[RNA]] species present outside of the cells from which they were transcribed. In homo sapiens, exRNAs have been discovered in bodily fluids such as venous blood, saliva, breast milk, urine, semen, menstrual blood, and vaginal fluid <!citations!>. Although their biological function is not fully understood, exRNAs have been proposed to play a role in a variety of biolgical processes including [[Syntrophy|syntrophy]] <!citation!>, intercellular communication, and cell regulation <!citations!>. Due to their potential biological and clinical significance, the [[National Insitutes of Health]] published a Request for Applications (RFA) for investigating extracellular RNA biology<ref>{{cite web|last=NIH|first=USA|title=NIH Common Fund RFA for exRNA Communication|url=http://commonfund.nih.gov/exrna/|accessdate=7 November 2012}}</ref>.
Extracellular RNA (also known as exRNA or exosomal RNA) describes [[RNA]] species present outside of the cells from which they were transcribed. In homo sapiens, exRNAs have been discovered in bodily fluids such as venous blood, saliva, breast milk, urine, semen, menstrual blood, and vaginal fluid<ref>{{cite journal|last=Chen|first=X|coauthors=Ba, Y; Ma, L; Cai, X; Yin, Y; Wang, K; Guo, J; Zhang, Y; Chen, J; Guo, X; Li, Q; Li, X; Wang, W; Zhang, Y; Wang, J; Jiang, X; Xiang, Y; Xu, C; Zheng, P; Zhang, J; Li, R; Zhang, H; Shang, X; Gong, T; Ning, G; Wang, J; Zen, K; Zhang, J; Zhang, CY|title=Characterization of microRNAs in serum: a novel class of biomarkers for diagnosis of cancer and other diseases.|journal=Cell research|date=2008 Oct|year=2008|volume=18|issue=10|pages=997-1006|pmid=18766170}}</ref><ref>{{cite journal|last=Michael|first=A|coauthors=Bajracharya, SD; Yuen, PS; Zhou, H; Star, RA; Illei, GG; Alevizos, I|title=Exosomes from human saliva as a source of microRNA biomarkers.|journal=Oral diseases|date=2010 Jan|volume=16|issue=1|pages=34-8|pmid=19627513}}</ref><ref>{{cite journal|last=Kosaka|first=N|coauthors=Izumi, H; Sekine, K; Ochiya, T|title=microRNA as a new immune-regulatory agent in breast milk.|journal=Silence|date=2010 Mar 1|volume=1|issue=1|pages=7|pmid=20226005}}</ref><ref>{{cite journal|last=Menke|first=TB|coauthors=Warnecke, JM|title=Improved conditions for isolation and quantification of RNA in urine specimens.|journal=Annals of the New York Academy of Sciences|date=2004 Jun|volume=1022|pages=185-9|pmid=15251958}}</ref><ref>{{cite journal|last=Zubakov|first=D|coauthors=Boersma, AW; Choi, Y; van Kuijk, PF; Wiemer, EA; Kayser, M|title=MicroRNA markers for forensic body fluid identification obtained from microarray screening and quantitative RT-PCR confirmation.|journal=International journal of legal medicine|date=2010 May|volume=124|issue=3|pages=217-26|pmid=20145944}}</ref><ref>{{cite journal|last=Hanson|first=EK|coauthors=Lubenow, H; Ballantyne, J|title=Identification of forensically relevant body fluids using a panel of differentially expressed microRNAs.|journal=Analytical biochemistry|date=2009 Apr 15|volume=387|issue=2|pages=303-14|pmid=19454234}}</ref>. Although their biological function is not fully understood, exRNAs have been proposed to play a role in a variety of biolgical processes including [[Syntrophy|syntrophy]] <!citation!>, intercellular communication, and cell regulation <!citations!>. Due to their potential biological and clinical significance, the [[National Insitutes of Health]] published a Request for Applications (RFA) for investigating extracellular RNA biology<ref>{{cite web|last=NIH|first=USA|title=NIH Common Fund RFA for exRNA Communication|url=http://commonfund.nih.gov/exrna/|accessdate=7 November 2012}}</ref>.


==Background==
==Background==

Revision as of 01:52, 4 February 2013

This sandbox is in the article namespace. Either move this page into your userspace, or remove the {{User sandbox}} template. Extracellular RNA (also known as exRNA or exosomal RNA) describes RNA species present outside of the cells from which they were transcribed. In homo sapiens, exRNAs have been discovered in bodily fluids such as venous blood, saliva, breast milk, urine, semen, menstrual blood, and vaginal fluid[1][2][3][4][5][6]. Although their biological function is not fully understood, exRNAs have been proposed to play a role in a variety of biolgical processes including syntrophy <!citation!>, intercellular communication, and cell regulation <!citations!>. Due to their potential biological and clinical significance, the National Insitutes of Health published a Request for Applications (RFA) for investigating extracellular RNA biology[7].

Background

A cartoon representation of the environments in which extracellular RNAs have been discovered.

Both prokaryotic and eukaryotic cells are known to secrete RNA <!citations!>. The Endosomal Sorting Complex Required for Transport (ESCRT) machinery was previously considered as a possible mechanism for RNA secretion from the cell, but more recently research studying microRNA secretion in human embryonic kidney cells and Cercopithecus aethiops kidney cells identified neutral sphingomyelinase 2 (nSMase2), an enzyme involved in ceramide biosynthesis, as a regulator of microRNA secretion levels. ExRNAs are often found packaged within vesicles such as exosomes, ectosomes, prostasomes, microvesicles, and apoptotic bodies <!citations!>. Although RNAs can be excreted from the cell without an enveloping container, the presence of ribonucleases <!link!> in extracellular environments would degrade the molecule.

Types of ExRNA

Extracellular RNA should not be viewed as a category describing a set of RNAs with a specific biological function or belonging to a particular RNA family. Similar to the term "non-coding RNA" <!link!>, "extracellular RNA" defines a group of several types of RNAs <!link!> whose functions are diverse, yet they share a common attribute which, in the case of exRNAs, is existence in an extracellular environment. The following types of RNA have been found outside the cell:

  • Messenger RNA (mRNA)
  • Transfer RNA (tRNA)
  • MicroRNA (miRNA)
  • Small interfering RNA (siRNA)
  • Long non-coding RNA (lncRNA)

Though prevalent inside of the cell, ribosomal RNA (rRNA)<!link!> does not seem to be a common exRNA. Efforts by Valadi et al. to characterize exosomal RNA using the Agilent Bioanalyzer technology showed little to no trace of 18S and 28S rRNA in exosomes secreted by MC/9 murine mast cells<!citations!>, and similar conclusions were made by Skog et al. for rRNA in gliobastoma microvesicle <!citations!>.

Function

To successfully function in extracellular environments, exRNA is often enclosed within a vesicular body to prevent its digestion by RNases. In some cases such as its use in prokaryotic syntrophy, exRNA is not packaged because the recipient cells use the ribnonuclease-digested nucleotides <!citation!>. The use of extracellular vesicles to protect exRNA from degradation is believed to be linked with the use of these containers as a way to transport RNA between cells. Biochemical evidence supports the idea that exRNA uptake is a common process, suggesting new pathways for intercellular communication. As a result, the presence, absence, and relative abundance of certain exRNAs can be correlated with changes in cellular signaling and may indicate specific disease states.[1]

Despite a limited understanding of exRNA biology, current research has shown the role of exRNAs to be multi-faceted [9][1][2][3][4]. Extracellular miRNAs are capable of targeting messenger RNAs in the recipient cell through RNA interference pathways.[7][8] In vitro experiments have shown the transfer of specific exRNAs into recipient cells inhibiting protein expression and preventing cancer cell growth <!PMC2878508!>. In addition to mRNAs being regulated by exRNAs, mRNAs can act as exRNAs to carry genetic information between cells. Messenger RNA contained in microvesicles secreted from glioblastomal cells were shown to generate a functional protein in recipient (human brain microvascular endothelial) cells in vitro. In another study of extracellular mRNAs, mRNAs transported by microvesicles from endothelial progenitor cells (EPCs) to human microvascular and macrovascular endothelial cells triggered angiogenesis in both the in vitro and in vivo setting. <!citations!>. Work by Hunter et al. used Ingenuity Pathway Analysis (IPA) <!link!> software that associated exRNAs found in human blood microvesicles with pathways involved in blood cell differentiation, metabolism, and immune function <!citation!>. These experimental and bioinformatics analyses favor the hypothesis that exRNAs play a role in numerous biological processes.

ExRNA Detection

Several methods have been developed or adapted to detect, characterize, and quantify exRNA from biological samples. RT-PCR, cDNA microarrays, and RNA sequencing are common techniques for RNA analysis. Applying these methods to study exRNAs mainly differs from cellular RNA experiments in the RNA isolation and/or extraction steps.

RT-PCR

For known exRNA nucleotide sequences, RT-PCR can be applied to detect their presence within a sample as well as quantify their abundance. This is done through first reverse transcribing the RNA sequencing into cDNA. The cDNA then serves as a template for PCR amplification. The major benefits of using RT-PCR are its quantitative accuracy in a dynamic range and increased sensitivity compared to methods such as RNase protection assays and dot blot hybridization. The disadvantage to RT-PCR is the requirement of costly supplies, and the necessity of sound experimental design and an in-depth understanding of normalization techniques in order to obtain accurate results and conclusions. http://www.biotechniques.com/BiotechniquesJournal/2005/July/Real-time-PCR-for-mRNA-quantitation/biotechniques-45505.html

Microfluidics

Microfluidic platforms such as the Agilent Bioanalyzer <!link!> are useful in assessing the quality of exRNA samples. With the Agilent Bioanalyzer, a lab-on-chip technology <!link!> that uses a sample of isolated RNA measures the length and quantity of RNA in the sample, and the results of the experiment can be represented as a digital electrophoresis gel image or an electropherogram <!link!>. Because a diverse range of RNAs can be detected by this technology, it is an effective method for more generally determining what types of RNAs are present in exRNAs samples through using size characterization.

cDNA Microarrays

Microarrays allow for larger-scale exRNA characterization and quantification. Microarrays used for RNA studies first generate different cDNA oligonucleotides (probes) that are attached to the microarray chip. An RNA sample can then be added to the chip, and RNAs with sequence complementarity to the cDNA probe will bind and generate a fluorescent signal that can be quantified. Micro RNA arrays have been used in exRNA studies to generate miRNA profiles of bodily fluids. http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3167594/ http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3505968/

RNA Sequencing

The advent of massively parallel sequencing (next-generation sequencing) lead to variations in DNA sequencing that allowed for high-throughput analyses of many genomic properties. Among these DNA sequencing-derived methods is RNA sequencing. The main advantage of RNA sequencing over other methods for exRNA detection and quantification is its high-throughput capabilities. Unlike microarrays, RNA sequencing is not constrained by factors such as oligonucleotide generation, and the number of probes that can be added to a chip. Indirect RNA sequencing of exRNA samples involves generating a cDNA library from the exRNAs followed by PCR amplification and sequencing. In 2009, Helicos Biosciences published a method for directly sequencing RNA molecules called Direct RNA sequencing (DRS™). (Nature 461, 814-818.). Regardless of the RNA sequencing platform, inherent biases exist at various steps in the experiment, but methods have been proposed to correct for these biases with promising results.

Clinical Significance

As growing evidence supports the function of exRNAs as intercellular communicators, research efforts are investigating the possibility of utilizing exRNAs in disease diagnosis, prognosis, and therapeutics <!citation!>

Biomarkers

The potential of extracellular RNAs to serve as biomarkers is significant not only because of their role in intercellular signaling but also due to developments in next generation sequencing that enable high throughput profiling.[10][11] The simplest form of an exRNA biomarker is the presence (or absence) of a specific extracellular RNA. These biological signatures have been discovered in exRNA studies of cancer, diabetes, arthritis, and prion-related diseases. <!citations!>

Cancer

A major research area of interest for exRNA has been its role in cancer. The table below (adapted from Kosaka et al.<!CITATION!>) lists several types of cancer in which exRNAs have been shown to be associated:

Cancer Type Biomarker Candidate
Diffuse large B-cell lymphoma (DLBCL) Expression levels of miR-155, miR-210 and miR-21 were higher in DLBCL patient sera compared to control sera; High miR-21 expression was associated with relapse-free survival
Prostate cancer Serum levels of miR-141 can distinguish patients with prostate cancer from healthy controls
Ovarian cancer The levels of the 8 specific miRNAs were similar between cellular and exosomal miRNAs. Exosomal miRNA from ovarian cancer patients exhibited similar profiles, which were significantly distinct from profiles observed in benign disease; miR-21, miR-92, miR-93, miR-126 and miR-29a were significantly overexpressed in

the serum from cancer patients compared to controls

Non small cell lung cancer Eleven serum miRNAs were found to be altered more than 5-fold between longer-survival and shorter-survival groups, and levels of four miRNAs were significantly associated with overall survival
Acute myeloid leukemia and acute lymphoblastic leukemia miR-92a decreased in the plasmas of acute leukemia patients
Breast cancer Increased miR-195 levels in patients were reflected in tumors, and circulating levels of miR-195 and let-7a decreased in cancer patients postoperatively, to levels comparable with control subjects; miR-155 was differentially expressed in the serum of women with hormone-sensitive compared to women with hormone-insensitive

breast cancer

Gastric cancer The plasma concentrations of miR-17-5p, miR-21, miR-106a, and miR-106b were significantly higher in patients than controls,whereas let-7a was lower in patients
Pancreatic cancer Circulating miR-210 levels are elevated in pancreatic cancer patients
Pancreatic ductal adenocarcinoma The combined analyses of four miRNAs (miR-21, miR-210, miR-155, and miR-196a) in plasma can discriminate patients from normal healthy individuals
Squamous cell carcinoma (SCC) of tongue Plasma miR-184 levels were significantly higher in tongue SCC patients in comparison with normal individuals, and the levels were significantly reduced after surgical removal of the primary tumors
Colorectal cancer Both miR-17-3p and miR-92 were significantly elevated in the patients, and the plasma levels of these miRNAs were reduced after surgery
Hepatocellular carcinoma (HCC) An increased amount of miR-500 was found in the sera of the HCC patients, and its levels in sera returned to normal after the surgical treatment

Forensic Application

$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$$ Extracellular RNAs (also known as exRNAs or exosomal RNAs) are RNAs found outside of the cell where they were transcribed. These RNAs are often found packaged within microvesicles, exosomes or ectosomes.

Function

Specific molecular mechanisms are required to protect RNA from degradation in the extracellular environment.[8][9][10][11] There is now biochemical evidence that extracellular RNA uptake is common, suggesting new pathways for intercellular communication.[12][13] Indeed, extracellular miRNAs (microRNAs) have been shown to be capable of targeting messenger RNAs in the recipient cell through RNA interference pathways.[14][15] It is likely that many of the RNAs identified in the extracellular environment are functional.[16][8][9][10][11] As a result, the presence, absence, and relative abundance of certain extracellular RNAs can be correlated with changes in cellular signaling, and may indicate specific disease states.[8]

The potential of extracellular RNAs to serve as biomarkers is significant not only because of their role in intercellular signaling, but also due to developments in next generation sequencing that enable high throughput profiling.[17][18] The simplest form of an exRNA biomarker is the presence (or absence) of a specific extracellular RNA. For example, a recent profile of prion infected neuronal cells revealed that their exosomes contained increased levels of let-7b, let-7i, miR-128a, miR-21, miR-222, miR-29b, miR-342-3p and miR-424. However, miR-146 is found at a lower-level when compared to non-infected cells.[8]

In 2012 the US National Institutes of Health (NIH) recently published a Request for Applications (RFA) to study extracellular RNA Communication.[19]

See also

References

  1. ^ Chen, X (2008 Oct). "Characterization of microRNAs in serum: a novel class of biomarkers for diagnosis of cancer and other diseases". Cell research. 18 (10): 997–1006. PMID 18766170. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: date and year (link)
  2. ^ Michael, A (2010 Jan). "Exosomes from human saliva as a source of microRNA biomarkers". Oral diseases. 16 (1): 34–8. PMID 19627513. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  3. ^ Kosaka, N (2010 Mar 1). "microRNA as a new immune-regulatory agent in breast milk". Silence. 1 (1): 7. PMID 20226005. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  4. ^ Menke, TB (2004 Jun). "Improved conditions for isolation and quantification of RNA in urine specimens". Annals of the New York Academy of Sciences. 1022: 185–9. PMID 15251958. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  5. ^ Zubakov, D (2010 May). "MicroRNA markers for forensic body fluid identification obtained from microarray screening and quantitative RT-PCR confirmation". International journal of legal medicine. 124 (3): 217–26. PMID 20145944. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  6. ^ Hanson, EK (2009 Apr 15). "Identification of forensically relevant body fluids using a panel of differentially expressed microRNAs". Analytical biochemistry. 387 (2): 303–14. PMID 19454234. {{cite journal}}: Check date values in: |date= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  7. ^ NIH, USA. "NIH Common Fund RFA for exRNA Communication". Retrieved 7 November 2012.
  8. ^ a b c d Bellingham SA, Coleman BM, Hill AF (2012). "Small RNA deep sequencing reveals a distinct miRNA signature released in exosomes from prion-infected neuronal cells". Nucleic Acids Res. PMID 22965126.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ a b Bellingham SA, Guo BB, Coleman BM, Hill AF (2012). "Exosomes: vehicles for the transfer of toxic proteins associated with neurodegenerative diseases?". Front Physiol. 3: 124. PMID 22563321.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  10. ^ a b Coleman BM, Hanssen E, Lawson VA, Hill AF (2012). "Prion-infected cells regulate the release of exosomes with distinct ultrastructural features". FASEB J. 26 (10): 4160–73. PMID 22767229.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ a b Hessvik NP, Phuyal S, Brech A, Sandvig K, Llorente A (2012). "Profiling of microRNAs in exosomes released from PC-3 prostate cancer cells". Biochim Biophys Acta. 1819 (11–12): 1154–1163. PMID 22982408.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ^ Valadi H, Ekström K, Bossios A, Sjöstrand M, Lee JJ, Lötvall JO (2007). "Exosome-mediated transfer of mRNAs and microRNAs is a novel mechanism of genetic exchange between cells". Nat Cell Biol. 9 (6): 654–9. PMID 17486113.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  13. ^ Lässer C, Alikhani VS, Ekström K, Eldh M, Paredes PT, Bossios A, Sjöstrand M, Gabrielsson S, Lötvall J, Valadi H (2011). "Human saliva, plasma and breast milk exosomes contain RNA: uptake by macrophages". J Transl Med. 9: 9. PMID 21235781.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  14. ^ Iguchi H, Kosaka N, Ochiya T (2010). "Secretory microRNAs as a versatile communication tool". Commun Integr Biol. 3 (5): 478–81. PMID 21057646.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  15. ^ Kosaka N, Iguchi H, Ochiya T (2010). "Circulating microRNA in body fluid: a new potential biomarker for cancer diagnosis and prognosis". Cancer Sci. 101 (10): 2087–92. PMID 20624164.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  16. ^ Iguchi H, Kosaka N, Ochiya T (2010). "Versatile applications of microRNA in anti-cancer drug discovery: from therapeutics to biomarkers". Curr Drug Discov Technol. 7 (2): 95–105. PMID 20836759.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  17. ^ Cloonan N, Xu Q, Faulkner GJ, Taylor DF, Tang DT, Kolle G, Grimmond SM (2009). "RNA-MATE: a recursive mapping strategy for high-throughput RNA-sequencing data". Bioinformatics. 25 (19): 2615–6. PMID 19648138.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  18. ^ Majewski J, Pastinen T (2011). "The study of eQTL variations by RNA-seq: from SNPs to phenotypes". Trends Genet. 27 (2): 72–9. PMID 21122937.
  19. ^ NIH, USA. "NIH Common Fund RFA for exRNA Communication". Retrieved 7 November 2012.

Further reading

  1. ^ Gudipati RK, Xu Z, Lebreton A, Séraphin B, Steinmetz LM, Jacquier A, Libri D (2012). "Extensive Degradation of RNA Precursors by the Exosome in Wild-Type Cells". Mol Cell. PMID 23000176.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  2. ^ Wan J, Yourshaw M, Mamsa H, Rudnik-Schöneborn S, Menezes MP, Hong JE, Leong DW, Senderek J, Salman MS, Chitayat D, Seeman P, von Moers A, Graul-Neumann L, Kornberg AJ, Castro-Gago M, Sobrido MJ, Sanefuji M, Shieh PB, Salamon N, Kim RC, Vinters HV, Chen Z, Zerres K, Ryan MM, Nelson SF, Jen JC (2012). "Mutations in the RNA exosome component gene EXOSC3 cause pontocerebellar hypoplasia and spinal motor neuron degeneration". Nat Genet. 44 (6): 704–8. PMID 22544365.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  3. ^ Xue Z, Yuan H, Guo J, Liu Y (2012). "Reconstitution of an Argonaute-dependent small RNA biogenesis pathway reveals a handover mechanism involving the RNA exosome and the exonuclease QIP". Mol Cell. 46 (3): 299–310. PMID 22516970.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  4. ^ Lässer C (2012). "Exosomal RNA as biomarkers and the therapeutic potential of exosome vectors". Expert Opin Biol Ther. 12 Suppl 1: S189-97. PMID 22506888.
  5. ^ Witharana C, Roppelt V, Lochnit G, Klug G, Evguenieva-Hackenberg E (2012). "Heterogeneous complexes of the RNA exosome in Sulfolobus solfataricus". Biochimie. 94 (7): 1578–87. PMID 22503705.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  6. ^ Holub P, Lalakova J, Cerna H, Pasulka J, Sarazova M, Hrazdilova K, Arce MS, Hobor F, Stefl R, Vanacova S (2012). "Air2p is critical for the assembly and RNA-binding of the TRAMP complex and the KOW domain of Mtr4p is crucial for exosome activation". Nucleic Acids Res. 40 (12): 5679–93. PMID 22402490.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. ^ Nag A, Steitz JA (2012). "Tri-snRNP-associated proteins interact with subunits of the TRAMP and nuclear exosome complexes, linking RNA decay and pre-mRNA splicing". RNA Biol. 9 (3): 334–42. PMID 22336707.
  8. ^ Ramírez-Garrastacho M, Esteban R (2011). "Yeast RNA viruses as indicators of exosome activity: human exosome hCsl4p participates in RNA degradation in Saccharomyces cerevisiae'". Yeast. 28 (12): 821–32. PMID 22068837.
  9. ^ Lange H, Gagliardi D (2011). "The exosome and 3'-5' RNA degradation in plants". Adv Exp Med Biol. 702: 50–62. PMID 21713677.
  10. ^ Lange H, Sement FM, Gagliardi D (2011). "MTR4, a putative RNA helicase and exosome co-factor, is required for proper rRNA biogenesis and development in Arabidopsis thaliana". Plant J. 68 (1): 51–63. PMID 21682783.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  11. ^ Flynn RA, Almada AE, Zamudio JR, Sharp PA (2011). "Antisense RNA polymerase II divergent transcripts are P-TEFb dependent and substrates for the RNA exosome". Proc Natl Acad Sci U S A. 108 (26): 10460–5. PMID 21670248.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  12. ^ Malet H, Lorentzen E. "Mechanisms of RNA recruitment by the exosome". RNA Biol. 8 (3): 398–403. PMID 21558791.
  13. ^ Taylor DD, Zacharias W, Gercel-Taylor C (2011). "Exosome isolation for proteomic analyses and RNA profiling". Methods Mol Biol. 728: 235–46. PMID 21468952.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  14. ^ Lykke-Andersen S, Tomecki R, Jensen TH, Dziembowski A. "The eukaryotic RNA exosome: same scaffold but variable catalytic subunits". RNA Biol. 8 (1): 61–6. PMID 21289487.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  15. ^ Silverstein RA, González de Valdivia E, Visa N (2011). "The incorporation of 5-fluorouracil into RNA affects the ribonucleolytic activity of the exosome subunit Rrp6". Mol Cancer Res. 9 (3): 332–40. PMID 21289297.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  16. ^ Niederberger T, Hartung S, Hopfner KP, Tresch A. "Processive RNA decay by the exosome: merits of a quantitative Bayesian sampling approach". RNA Biol. 8 (1): 55–60. PMID 21282980.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  17. ^ Basu U, Meng FL, Keim C, Grinstein V, Pefanis E, Eccleston J, Zhang T, Myers D, Wasserman CR, Wesemann DR, Januszyk K, Gregory RI, Deng H, Lima CD, Alt FW (2011). "The RNA exosome targets the AID cytidine deaminase to both strands of transcribed duplex DNA substrates". Cell. 144 (3): 353–63. PMID 21255825.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  18. ^ Lardenois A, Liu Y, Walther T, Chalmel F, Evrard B, Granovskaia M, Chu A, Davis RW, Steinmetz LM, Primig M (2011). "Execution of the meiotic noncoding RNA expression program and the onset of gametogenesis in yeast require the conserved exosome subunit Rrp6". Proc Natl Acad Sci U S A. 108 (3): 1058–63. PMID 21149693.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  19. ^ Costello JL, Stead JA, Feigenbutz M, Jones RM, Mitchell P (2011). "The C-terminal region of the exosome-associated protein Rrp47 is specifically required for box C/D small nucleolar RNA 3'-maturation". J Biol Chem. 286 (6): 4535–43. PMID 21135092.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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Category:RNA

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