480:
472:
structure of the human exosome has no enzymatically active protein. Despite this loss of catalytic activity, the structure of the core exosome is highly conserved from archaea to humans, suggesting that the complex performs a vital cellular function. In eukaryotes, the absence of the phosphorolytic activity is compensated by the presence of the hydrolytic enzymes, which are responsible for the ribonuclease activity of the exosome in such organisms.
225:
357:
694:(one of the RNase PH like proteins from the ring) and antibodies recognizing these proteins are found in approximately 30% of patients with the PM/Scl overlap syndrome. Although these two proteins are the main target of the autoantibodies, other exosome subunits and associated proteins (like C1D) can be targeted in these patients. At the current time, the most sensitive way to detect these antibodies is by using a
582:
436:
1265:
248:
244:-like proteins. In archaea there are two different PH-like proteins (called Rrp41 and Rrp42), each present three times in an alternating order. Eukaryotic exosome complexes have six different proteins that form the ring structure. Of these six eukaryotic proteins, three resemble the archaeal Rrp41 protein and the other three proteins are more similar to the archaeal Rrp42 protein.
40:
3568:
Wan, J.; Yourshaw, M.; Mamsa, H.; Rudnik-SchΓΆneborn, S.; Menezes, M. P.; Hong, J. E.; Leong, D. W.; Senderek, J.; Salman, M. S.; Chitayat, D.; Seeman, P.; Von Moers, A.; Graul-Neumann, L.; Kornberg, A. J.; Castro-Gago, M.; Sobrido, M. A. J. S.; Sanefuji, M.; Shieh, P. B.; Salamon, N.; Kim, R. C.;
475:
As stated above, the hydrolytic proteins Rrp6 and Rrp44 are associated with the exosome in yeast and in humans, besides Rrp6, two different proteins, Dis3 and Dis3L1 can be associated at the position of the yeast Rrp44 protein. Although originally the S1 domain proteins were thought to have 3'-5'
344:
exosome complexes and has a crucial role in the activity of the yeast exosome complex. While a human homologue of the protein exists, no evidence was found for a long time that its human homologue was associated with the human exosome complex. In 2010, however, it was discovered that humans have
443:
As stated above, the exosome complex contains many proteins with ribonuclease domains. The exact nature of these ribonuclease domains has changed across evolution from bacterial to archaeal to eukaryotic complexes as various activities have been gained and lost. The exosome is primarily a 3'-5'
295:
of some eukaryotic organisms, two RNase PH domains, and both an S1 and KH RNA binding domain are part of a single protein, which forms a trimeric complex that adopts a structure almost identical to that of the exosome. Because of this high similarity in both protein domains and structure, these
471:
In archaea, the Rrp41 subunit of the complex is a phosphorolytic exoribonuclease. Three copies of this protein are present in the ring and are responsible for the activity of the complex. In eukaryotes, none of the RNase PH subunits have retained this catalytic activity, meaning the core ring
398:
Apart from these two tightly bound protein subunits, many proteins interact with the exosome complex in both the cytoplasm and nucleus of cells. These loosely associated proteins may regulate the activity and specificity of the exosome complex. In the cytoplasm, the exosome interacts with
251:
Subunits and organisation of the archaeal (left) and eukaryotic (right) exosome complexes. Different proteins are numbered, showing that the archaeal exosome contains 4 different proteins, but the eukaryotic exosome contains nine different proteins. See the full
403:(ARE) binding proteins (e.g. KRSP and TTP), which can promote or prevent degradation of mRNAs. The nuclear exosome associates with RNA binding proteins (e.g. MPP6/Mpp6 and C1D/Rrp47 in humans/yeast) that are required for processing certain substrates.
452:. Exoribonucleases contained in exosome complexes are either phosphorolytic (the RNase PH-like proteins) or, in eukaryotes, hydrolytic (the RNase R and RNase D domain proteins). The phosphorolytic enzymes use inorganic phosphate to cleave the
340:(nucleases that use water to cleave the nucleotide bonds). In addition to being an exoribonucleolytic enzyme, Rrp44 also has endoribonucleolytic activity, which resides in a separate domain of the protein. In yeast, Rrp44 is associated with
702:, instead of complete proteins. By this method, autoantibodies are found in up to 55% of patients with the PM/Scl overlap syndrome, but they can also be detected in patients with either scleroderma, polymyositis, or dermatomyositis alone.
345:
three Rrp44 homologues and two of these can be associated with the exosome complex. These two proteins most likely degrade different RNA substrates due to their different cellular localization, with one being localized in the cytoplasm (
662:
from human cell extracts. In immunoprecipitation assays with sera from anti-exosome positive sera, a distinctive set of proteins is precipitated. Already years before the exosome complex was identified, this pattern was termed the
268:. In eukaryotes, three different "S1" proteins are bound to the ring, whereas in archaea either one or two different "S1" proteins can be part of the exosome (although there are always three S1 subunits attached to the complex).
483:
Schematic view of the archaeal (left) and eukaryotic (right) exosome complexes with the most common associated proteins. In color and marked with a star are the subunits of each complex that have catalytic activity. See
1809:
Walter, P; Klein, F; Lorentzen, E; Ilchmann, A; Klug, G; Evguenieva-Hackenberg, E (2006). "Characterization of native and reconstituted exosome complexes from the hyperthermophilic archaeon
Sulfolobus solfataricus".
605:(CUTs) that are produced from thousands of loci within the yeast genome. The importance of these unstable RNAs and their degradation are still unclear, but similar RNA species have also been detected in human cells.
589:
The exosome is a key complex in cellular RNA quality control. Unlike prokaryotes, eukaryotes possess highly active RNA surveillance systems that recognise unprocessed and mis-processed RNA-protein complexes (such as
1763:
Raijmakers, R; Egberts, WV; Van
Venrooij, WJ; Pruijn, GJ (2002). "Protein-protein interactions between human exosome components support the assembly of RNase PH-type subunits into a six-membered PNPase-like ring".
476:
hydrolytic exoribonuclease activity as well, the existence of this activity has recently been questioned and these proteins might have just a role in binding substrates prior to their degradation by the complex.
207:). Purification of this "PM/Scl complex" allowed the identification of more human exosome proteins and eventually the characterization of all components in the complex. In 2001, the increasing amount of
3267:
Raijmakers, R; Renz, M; Wiemann, C; Egberts, WV; Seelig, HP; Van
Venrooij, WJ; Pruijn, GJ (2004). "PM-Scl-75 is the main autoantigen in patients with the polymyositis/scleroderma overlap syndrome".
1243:
In humans, two different proteins can be associated in this position. In the cytoplasm of cells, Dis3L1 is associated with the exosome, whereas in the nucleus, Dis3 can bind to the core complex.
211:
that had become available allowed the prediction of exosome proteins in archaea, although it would take another 2 years before the first exosome complex from an archaeal organism was purified.
3048:
Neil H, Malabat C, d'Aubenton-Carafa Y, Xu Z, Steinmetz LM, Jacquier A (February 2009). "Widespread bidirectional promoters are the major source of cryptic transcripts in yeast".
1668:
Lorentzen, E; Walter, P; Fribourg, S; Evguenieva-Hackenberg, E; Klug, G; Conti, E (2005). "The archaeal exosome core is a hexameric ring structure with three catalytic subunits".
756:. In yeast cells treated with fluorouracil, defects were found in the processing of ribosomal RNA identical to those seen when the activity of the exosome was blocked by
390:
protein family. The protein PM/Scl-100 is most commonly part of exosome complexes in the nucleus of cells, but can form part of the cytoplasmic exosome complex as well.
509:(mRNA) molecules. The complex can degrade mRNA molecules that have been tagged for degradation because they contain errors, through interactions with proteins from the
3355:
Schilders, G; Egberts, WV; Raijmakers, R; Pruijn, GJ (2007). "C1D is a major autoantibody target in patients with the polymyositis-scleroderma overlap syndrome".
283:
processing, forms a hexameric ring consisting of six identical RNase PH proteins. In the case of PNPase, which is a phosphorolytic RNA-degrading protein found in
564:, growth stops and the cells eventually die. Both the core proteins of the exosome complex, as well as the two main associated proteins, are essential proteins.
1518:"Prediction of the archaeal exosome and its connections with the proteasome and the translation and transcription machineries by a comparative-genomic approach"
103:
The core of the exosome contains a six-membered ring structure to which other proteins are attached. In eukaryotic cells, the exosome complex is present in the
594:) prior to their exit from the nucleus. It is presumed that this system prevents aberrant complexes from interfering with important cellular processes such as
1896:"Crystal structure of the phosphorolytic exoribonuclease RNase PH from Bacillus subtilis and implications for its quaternary structure and tRNA binding"
195:. Not long after, in 1999, it was realized that the exosome was in fact the yeast equivalent of an already described complex in human cells called the
2921:
Carpousis AJ, AJ (2002). "The
Escherichia coli RNA degradosome: structure, function and relationship in other ribonucleolytic multienzyme complexes".
709:
of these patients can vary widely. The symptoms that are seen most often are the typical symptoms of the individual autoimmune diseases and include
2505:
Dziembowski, A; Lorentzen, E; Conti, E; SΓ©raphin, B (2007). "A single subunit, Dis3, is in essence responsible for yeast exosome core activity".
2334:
Raijmakers, R; Schilders, G; Pruijn, GJ (2004). "The exosome, a molecular machine for controlled RNA degradation in both nucleus and cytoplasm".
115:, although different proteins interact with the exosome complex in these compartments regulating the RNA degradation activity of the complex to
3656:
Houseley, J; Tollervey, D (2008). "The nuclear RNA surveillance machinery: the link between ncRNAs and genome structure in budding yeast?".
2236:
Tomecki, R; Kristiansen, MS; Lykke-Andersen, S; Chlebowski, A; Larsen, KM; Szczesny, RJ; Drazkowska, K; Pastula, A; et al. (2010).
725:. Treatment of these patients is symptomatic and is similar to treatment for the individual autoimmune disease, often involving either
328:
Besides these nine core exosome proteins, two other proteins often associate with the complex in eukaryotic organisms. One of these is
647:
3441:
Mahler, M; Raijmakers, R (2007). "Novel aspects of autoantibodies to the PM/Scl complex: Clinical, genetic and diagnostic insights".
1413:
Brouwer, R; Allmang, C; Raijmakers, R; Van
Aarssen, Y; Egberts, WV; Petfalski, E; Van Venrooij, WJ; Tollervey, D; Pruijn, GJ (2001).
3638:
1644:
680:
498:
2085:"The N-terminal PIN domain of the exosome subunit Rrp44 harbors endonuclease activity and tethers Rrp44 to the yeast core exosome"
3302:
Brouwer, R; Vree
Egberts, WT; Hengstman, GJ; Raijmakers, R; Van Engelen, BG; Seelig, HP; Renz, M; Mierau, R; et al. (2002).
414:(Ski2) and is involved in mRNA degradation. In the nucleus, the processing of rRNA and snoRNA by the exosome is mediated by the
3308:
601:
In addition to RNA processing, turnover and surveillance activities, the exosome is important for the degradation of so-called
537:
is the compartment where the majority of the exosome complexes are found. There it plays a role in the processing of the 5.8S
1945:"A duplicated fold is the structural basis for polynucleotide phosphorylase catalytic activity, processivity, and regulation"
304:. The RNase PH-like exosome proteins, PNPase and RNase PH all belong to the RNase PH family of RNases and are phosphorolytic
135:
in this case), and in eukaryotes also an endoribonucleolytic function, meaning it cleaves RNA at sites within the molecule.
2133:
3571:"Mutations in the RNA exosome component gene EXOSC3 cause pontocerebellar hypoplasia and spinal motor neuron degeneration"
760:
strategies. Lack of correct ribosomal RNA processing is lethal to cells, explaining the antimetabolic effect of the drug.
240:
The core of the complex has a ring structure consisting of six proteins that all belong to the same class of RNases, the
569:
568:
do not have an exosome complex; however, similar functions are performed by a simpler complex that includes the protein
276:
1986:"The PNPase, exosome and RNA helicases as the building components of evolutionarily-conserved RNA degradation machines"
705:
As the autobodies are found mainly in patients that have characteristics of several different autoimmune diseases, the
602:
2026:
671:
of cells, which sparked the suggestion that the antigen recognized by autoantibodies might be important in ribosome
3903:
3519:
Lum, PY; Armour, CD; Stepaniants, SB; Cavet, G; Wolf, MK; Butler, JS; Hinshaw, JC; Garnier, P; et al. (2004).
777:
406:
In addition to single proteins, other protein complexes interact with the exosome. One of those is the cytoplasmic
166:
2642:"Nonsense-mediated mRNA decay in mammalian cells involves decapping, deadenylating, and exonucleolytic activities"
3001:"Cryptic pol II transcripts are degraded by a nuclear quality control pathway involving a new poly(A) polymerase"
1234:
In archaea several exosome proteins are present in multiple copies, to form the full core of the exosome complex.
823:
2184:
Staals, RH; Bronkhorst, AW; Schilders, G; Slomovic, S; Schuster, G; Heck, AJ; Raijmakers, R; Pruijn, GJ (2010).
1464:
Chen, CY; Gherzi, R; Ong, SE; Chan, EL; Raijmakers, R; Pruijn, GJ; Stoecklin, G; Moroni, C; et al. (2001).
3728:
BΓΌttner, K; Wenig, K; Hopfner, KP (2006). "The exosome: a macromolecular cage for controlled RNA degradation".
2874:"Human cell growth requires a functional cytoplasmic exosome, which is involved in various mRNA decay pathways"
187:
116:
3621:
Schilders, G; Pruijn, GJ (2008). "Chapter 11 Biochemical
Studies of the Mammalian Exosome with Intact Cells".
56:
below. The channel through which RNA passes during degradation is visible at the center of the protein complex
710:
627:
510:
465:
521:. Several proteins that stabilize or destabilize mRNA molecules through binding to AU-rich elements in the
3898:
3888:
3737:
726:
3521:"Discovering modes of action for therapeutic compounds using a genome-wide screen of yeast heterozygotes"
2601:"Structural basis of 3' end RNA recognition and exoribonucleolytic cleavage by an exosome RNase PH core"
776:, cerebellar atrophy, progressive microcephaly and profound global developmental delay, consistent with
679:
exosome proteins have become available and these have been used to develop line immunoassays (LIAs) and
457:
229:
1315:"The Exosome: A Conserved Eukaryotic RNA Processing Complex Containing Multiple 3β²β5β² Exoribonucleases"
3112:
3057:
2732:"Localization of AU-rich element-containing mRNA in cytoplasmic granules containing exosome subunits"
2041:
1169:
773:
686:
In these diseases, antibodies are mainly directed against two of the proteins of the complex, called
676:
170:
3742:
2238:"The human core exosome interacts with differentially localized processive RNases: hDIS3 and hDIS3L"
585:
Two core subunits of the archaeal exosome (Rrp41 and Rrp42), bound to a small RNA molecule (in red).
1134:
1099:
1064:
1029:
994:
959:
924:
889:
854:
769:
659:
453:
309:
261:
162:
143:
131:. The exosome has an exoribonucleolytic function, meaning it degrades RNA starting at one end (the
3476:
Jablonska, S; Blaszczyk, M (1998). "Scleromyositis: a scleroderma/polymyositis overlap syndrome".
479:
439:
Reaction diagrams for both hydrolytic (left) and phosphorolytic (right) 3' end degradation of RNA.
3835:
3763:
3550:
3501:
3081:
3030:
2981:
2581:
2530:
2449:
2418:
LaCava, J; Houseley, J; Saveanu, C; Petfalski, E; Thompson, E; Jacquier, A; Tollervey, D (2005).
2065:
1835:
1693:
1495:
1346:
655:
623:
200:
3853:
3827:
3798:
3755:
3716:
3673:
3644:
3634:
3600:
3542:
3493:
3458:
3423:
3372:
3337:
3284:
3249:
3214:
3183:
Gelpi, C; AlguerΓ³, A; Angeles
Martinez, M; Vidal, S; Juarez, C; Rodriguez-Sanchez, JL (1991).
3165:
3130:
3073:
3022:
2973:
2938:
2903:
2854:
2805:
2753:
2712:
2663:
2622:
2573:
2522:
2441:
2400:
2351:
2316:
2267:
2215:
2163:
2114:
2057:
2007:
1966:
1925:
1876:
1827:
1791:
1745:
1685:
1650:
1640:
1598:
1547:
1487:
1446:
1395:
1338:
757:
595:
386:(in human). Like Rrp44, this protein is a hydrolytic exoribonuclease, but in this case of the
44:
31:
3819:
3788:
3747:
3706:
3698:
3665:
3626:
3590:
3582:
3532:
3485:
3450:
3413:
3403:
3364:
3327:
3317:
3276:
3241:
3204:
3196:
3157:
3120:
3065:
3012:
2965:
2930:
2893:
2885:
2844:
2836:
2795:
2787:
2743:
2702:
2694:
2653:
2612:
2563:
2514:
2482:
2431:
2390:
2382:
2343:
2306:
2298:
2257:
2249:
2205:
2197:
2153:
2145:
2104:
2096:
2049:
1997:
1956:
1915:
1907:
1866:
1819:
1781:
1773:
1735:
1727:
1677:
1632:
1588:
1580:
1537:
1529:
1477:
1436:
1426:
1385:
1377:
1328:
817:
706:
667:. Immunofluorescence using sera from these patients usually shows a typical staining of the
561:
224:
639:
557:
445:
419:
400:
337:
305:
301:
77:
3116:
3061:
2045:
3711:
3686:
3595:
3570:
3418:
3391:
3209:
3200:
3184:
2898:
2873:
2849:
2824:
2800:
2775:
2707:
2682:
2554:
2473:
2395:
2371:"Domain interactions within the Ski2/3/8 complex and between the Ski complex and Ski7p"
2370:
2262:
2237:
2210:
2185:
2158:
2109:
2084:
1920:
1895:
1740:
1715:
1593:
1566:
1364:
Allmang, C; Petfalski, E; Podtelejnikov, A; Mann, M; Tollervey, D; Mitchell, P (1999).
1319:
1270:
738:
619:
514:
356:
208:
192:
154:
151:
147:
139:
3868:
3863:
3858:
3630:
3537:
3520:
3304:"Autoantibodies directed to novel components of the PM/Scl complex, the human exosome"
2658:
2641:
2311:
2286:
1961:
1944:
1777:
1636:
1542:
1517:
1482:
1465:
1390:
1365:
1333:
1314:
581:
3882:
3751:
3569:
Vinters, H. V.; Chen, Z.; Zerres, K.; Ryan, M. M.; Nelson, S. F.; Jen, J. C. (2012).
3332:
3303:
3161:
2956:
Houseley J, LaCava J, Tollervey D (July 2006). "RNA-quality control by the exosome".
1823:
646:
of patients by a variety of assays. In the past, the most commonly used methods were
538:
506:
415:
124:
120:
3839:
3554:
3505:
2985:
2825:"MPP6 is an exosome-associated RNA-binding protein involved in 5.8S rRNA maturation"
2774:
Allmang, C; Kufel, J; Chanfreau, G; Mitchell, P; Petfalski, E; Tollervey, D (1999).
2585:
2534:
2453:
1839:
1565:
Evguenieva-Hackenberg, E; Walter, P; Hochleitner, E; Lottspeich, F; Klug, G (2003).
1499:
1350:
435:
3893:
3767:
3085:
3034:
2069:
1697:
1571:
1290:
745:
741:
672:
643:
635:
526:
292:
288:
108:
2469:"Erratum: Reconstitution, activities, and structure of the eukaryotic RNA exosome"
1366:"The yeast exosome and human PM-Scl are related complexes of 3' β 5' exonucleases"
367:
3669:
3454:
3232:
Targoff, IN; Reichlin, M (1985). "Nucleolar localization of the PM-Scl antigen".
2617:
2600:
2420:"RNA degradation by the exosome is promoted by a nuclear polyadenylation complex"
2149:
3101:"RNA exosome depletion reveals transcription upstream of active human promoters"
2698:
1855:"Crystal structure of the tRNA processing enzyme RNase PH from Aquifex aeolicus"
1286:
722:
631:
574:
407:
371:
247:
97:
3793:
3776:
3185:"Identification of protein components reactive with anti-PM/Scl autoantibodies"
3017:
3000:
2791:
2568:
2549:
2487:
2468:
2436:
2419:
1731:
1264:
132:
2002:
1985:
1584:
1279:
1260:
461:
85:
3702:
2302:
548:
Although most cells have other enzymes that can degrade RNA, either from the
3125:
3100:
2683:"A genomic screen in yeast reveals novel aspects of nonstop mRNA metabolism"
2347:
2134:"The exosome subunit Rrp44 plays a direct role in RNA substrate recognition"
1218:
1175:
1140:
1105:
1070:
1035:
1000:
965:
930:
895:
860:
714:
668:
591:
534:
502:
362:
297:
265:
161:
function by blocking the activity of the exosome. In addition, mutations in
128:
112:
104:
48:
39:
3864:
Structure of an archaeal exosome bound to RNA at the RCSB Protein Data Bank
3831:
3802:
3759:
3720:
3677:
3648:
3604:
3546:
3462:
3427:
3376:
3341:
3288:
3245:
3169:
3134:
3077:
3026:
2977:
2942:
2907:
2858:
2809:
2757:
2748:
2731:
2716:
2667:
2626:
2577:
2526:
2445:
2404:
2355:
2271:
2253:
2219:
2201:
2167:
2118:
2061:
2011:
1970:
1929:
1880:
1871:
1854:
1831:
1795:
1749:
1689:
1654:
1602:
1551:
1491:
1450:
1431:
1414:
1399:
1381:
553:
549:
522:
449:
313:
3623:
RNA Turnover in
Eukaryotes: Nucleases, Pathways and Analysis of mRNA Decay
3497:
3253:
3218:
2320:
2100:
1342:
360:"Ribbon view" of the partial structure of the yeast exosome subunit Rrp6,
3869:
Structure of the yeast exosome protein Rrp6 at the RCSB Protein Data Bank
3823:
3392:"Clinical evaluation of autoantibodies to a novel PM/Scl peptide antigen"
3390:
Mahler, M; Raijmakers, R; DΓ€hnrich, C; BlΓΌthner, M; Fritzler, MJ (2005).
2840:
2550:"Reconstitution, activities, and structure of the eukaryotic RNA exosome"
1466:"AU binding proteins recruit the exosome to degrade ARE-containing mRNAs"
718:
565:
556:
of the RNA, the exosome complex is essential for cell survival. When the
411:
284:
272:
241:
93:
3069:
2386:
2186:"Dis3-like 1: a novel exoribonuclease associated with the human exosome"
2053:
1911:
1313:
Mitchell, P; Petfalski, E; Shevchenko, A; Mann, M; Tollervey, D (1997).
3873:
3489:
2934:
2889:
1786:
1441:
695:
560:
of exosome proteins is artificially reduced or stopped, for example by
387:
333:
199:, which had been identified as an autoantigen in patients with certain
119:
specific to these cell compartments. Substrates of the exosome include
89:
73:
3368:
3280:
2287:"Comparative sequence analysis of ribonucleases HII, III, II PH and D"
1629:
Cell and molecular biology of the exosome: how to make or break an RNA
1533:
517:
pathways. In alternative fashion, mRNAs are degraded as part of their
2518:
1631:. International Review of Cytology. Vol. 251. pp. 159β208.
1513:
1211:
749:
651:
542:
346:
158:
17:
3586:
2969:
1681:
3777:"The Exosome and the Proteasome: Nano-Compartments for Degradation"
3408:
781:
3874:
3D macromolecular structures of exosomes at the EM Data Bank(EMDB)
3322:
3099:
Preker P, P; Nielsen, J; Kammler, S; et al. (December 2008).
753:
699:
630:, an autoimmune disease in which patients have symptoms from both
580:
530:
478:
434:
246:
223:
182:
84:(ribonucleic acid) molecules. Exosome complexes are found in both
38:
1207:
529:, the exosome is required for the correct processing of several
518:
350:
280:
2999:
Wyers F, F; Rougemaille, M; Badis, G; et al. (June 2005).
260:
Located on top of this ring are three proteins that have an S1
3859:
Structure of an archaeal exosome at the RCSB Protein Data Bank
2776:"Functions of the exosome in rRNA, snoRNA and snRNA synthesis"
541:(the first identified function of the exosome) and of several
317:
81:
2025:
Lebreton, A; Tomecki, R; Dziembowski, A; SΓ©raphin, B (2008).
1627:
Schilders, G; Van Dijk, E; Raijmakers, R; Pruijn, GJ (2006).
3854:
Structure of the human exosome at the RCSB Protein Data Bank
752:. It is one of the most successful drugs for treating solid
698:, derived from the PM/Scl-100 protein, as the antigen in an
271:
This ring structure is very similar to that of the proteins
2823:
Schilders, G; Raijmakers, R; Raats, JM; Pruijn, GJ (2005).
1252:
Contributes to the ribonucleolytic activity of the complex.
626:. These autoantibodies are mainly found in people with the
3625:. Methods in Enzymology. Vol. 448. pp. 211β226.
279:. In bacteria, the protein RNase PH, which is involved in
3148:
J.E. Pope, JE (2002). "Scleroderma overlap syndromes".
2083:
Schneider, C; Leung, E; Brown, J; Tollervey, D (2009).
1894:
Harlow, LS; Kadziola, A; Jensen, KF; Larsen, S (2004).
2027:"Endonucleolytic RNA cleavage by a eukaryotic exosome"
3687:"The exosome and RNA quality control in the nucleus"
1567:"An exosome-like complex in Sulfolobus solfataricus"
448:, meaning that it degrades RNA molecules from their
525:of mRNAs interact with the exosome complex. In the
1293:, that also contains an RNA binding ring structure
737:The exosome has been shown to be inhibited by the
138:Several proteins in the exosome are the target of
3810:Pruijn, GJ (2005). "Doughnuts dealing with RNA".
2132:Schneider, C; Anderson, JT; Tollervey, D (2007).
1716:"A view to a kill: structure of the RNA exosome"
2872:van Dijk, EL; Schilders, G; Pruijn, GJ (2007).
2500:
2498:
1282:, the main protein degrading machinery of cells
497:The exosome is involved in the degradation and
378:The second common associated protein is called
1709:
1707:
418:, which contains both RNA helicase (Mtr4) and
264:domain (RBD). Two proteins in addition have a
1415:"Three novel components of the human exosome"
505:of cells, it is involved in the turn-over of
8:
1984:Lin-Chao, S; Chiou, NT; Schuster, G (2007).
43:"Ribbon view" of the human exosome complex.
332:, a hydrolytic RNase, which belongs to the
232:of the human exosome complex. See the full
2681:Wilson, MA; Meaux, S; Van Hoof, A (2007).
2231:
2229:
1943:Symmons, MF; Jones, GH; Luisi, BF (2000).
1308:
1306:
791:
3812:Nature Structural & Molecular Biology
3792:
3741:
3710:
3594:
3536:
3417:
3407:
3331:
3321:
3208:
3124:
3016:
2897:
2848:
2799:
2747:
2706:
2657:
2616:
2567:
2507:Nature Structural & Molecular Biology
2486:
2435:
2394:
2310:
2261:
2209:
2179:
2177:
2157:
2108:
2001:
1960:
1919:
1870:
1853:Ishii, R; Nureki, O; Yokoyama, S (2003).
1785:
1739:
1670:Nature Structural & Molecular Biology
1592:
1541:
1481:
1440:
1430:
1389:
1332:
683:(ELISAs) for detecting these antibodies.
501:of a wide variety of RNA species. In the
2369:Wang, L; Lewis, MS; Johnson, AW (2005).
642:. Autoantibodies can be detected in the
355:
2769:
2767:
2548:Liu, Q; Greimann, JC; Lima, CD (2006).
2467:Liu, Q; Greimann, JC; Lima, CD (2007).
1302:
181:The exosome was first discovered as an
2640:LeJeune, F; Li, X; Maquat, LE (2003).
460:. The hydrolytic enzymes use water to
80:capable of degrading various types of
1622:
1620:
1618:
1616:
1614:
1612:
618:The exosome complex is the target of
485:
253:
233:
53:
7:
3189:Clinical and Experimental Immunology
2730:Lin, WJ; Duffy, A; Chen, CY (2007).
3201:10.1111/j.1365-2249.1990.tb05291.x
681:enzyme linked immunosorbent assays
25:
27:Protein complex that degrades RNA
3752:10.1111/j.1365-2958.2006.05331.x
3396:Arthritis Research & Therapy
3309:Arthritis Research & Therapy
3162:10.1097/00002281-200211000-00013
2336:European Journal of Cell Biology
1824:10.1111/j.1365-2958.2006.05393.x
1263:
1247:
1238:
1229:
349:) and the other in the nucleus (
3775:Lorentzen, E; Conti, E (2006).
3150:Current Opinion in Rheumatology
2736:Journal of Biological Chemistry
2599:Lorentzen, E; Conti, E (2005).
1859:Journal of Biological Chemistry
1516:; Wolf, YI; Aravind, L (2001).
1419:Journal of Biological Chemistry
690:(the RNase D like protein) and
312:to remove nucleotides from the
100:carries out similar functions.
3685:Vanacova, S; Stefl, R (2007).
1714:Shen, V; Kiledjian, M (2006).
1:
3631:10.1016/S0076-6879(08)02611-6
3538:10.1016/S0092-8674(03)01035-3
2659:10.1016/S1097-2765(03)00349-6
1990:Journal of Biomedical Science
1962:10.1016/S0969-2126(00)00521-9
1778:10.1016/S0022-2836(02)00947-6
1637:10.1016/S0074-7696(06)51005-8
1483:10.1016/S0092-8674(01)00578-5
1334:10.1016/S0092-8674(00)80432-8
185:in 1997 in the budding yeast
96:a simpler complex called the
3670:10.1016/j.bbagrm.2007.12.008
3455:10.1016/j.autrev.2007.01.013
2618:10.1016/j.molcel.2005.10.020
2150:10.1016/j.molcel.2007.06.006
1766:Journal of Molecular Biology
1182:
1147:
1112:
1077:
1042:
1007:
972:
937:
902:
867:
832:
603:cryptic unstable transcripts
533:RNA molecules. Finally, the
296:complexes are thought to be
204:
2699:10.1534/genetics.107.073205
729:or immunomodulating drugs.
3920:
3794:10.1016/j.cell.2006.05.002
3357:Arthritis & Rheumatism
3269:Arthritis & Rheumatism
3234:Arthritis & Rheumatism
3018:10.1016/j.cell.2005.04.030
2569:10.1016/j.cell.2006.10.037
2488:10.1016/j.cell.2007.09.019
2437:10.1016/j.cell.2005.04.029
1732:10.1016/j.cell.2006.11.035
778:pontocerebellar hypoplasia
167:pontocerebellar hypoplasia
142:in patients with specific
29:
2003:10.1007/s11373-007-9178-y
1585:10.1038/sj.embor.embor929
622:in patients with various
466:nucleotide monophosphates
228:Top and side view of the
3703:10.1038/sj.embor.7401005
2958:Nat. Rev. Mol. Cell Biol
2792:10.1093/emboj/18.19.5399
1289:, a complex involved in
464:these bonds β releasing
410:, which includes an RNA
308:, meaning that they use
188:Saccharomyces cerevisiae
68:, often just called the
30:Not to be confused with
3126:10.1126/science.1164096
2348:10.1078/0171-9335-00385
1370:Genes & Development
772:cause infantile spinal
628:PM/Scl overlap syndrome
511:nonsense mediated decay
458:nucleotide diphosphates
148:PM/Scl overlap syndrome
3730:Molecular Microbiology
3246:10.1002/art.1780280221
2829:Nucleic Acids Research
2749:10.1074/jbc.M702281200
2303:10.1093/nar/25.16.3187
2291:Nucleic Acids Research
2254:10.1038/emboj.2010.121
2202:10.1038/emboj.2010.122
2089:Nucleic Acids Research
1872:10.1074/jbc.M300639200
1812:Molecular Microbiology
1432:10.1074/jbc.M007603200
1382:10.1101/gad.13.16.2148
764:Neurological disorders
648:double immunodiffusion
586:
523:3' untranslated region
489:
440:
375:
266:K-homology (KH) domain
257:
237:
127:, and many species of
57:
3478:Clinical Rheumatology
744:, a drug used in the
584:
482:
438:
359:
336:family of hydrolytic
250:
227:
111:, and especially the
42:
3824:10.1038/nsmb0705-562
3658:Biochim Biophys Acta
3443:Autoimmunity Reviews
774:motor neuron disease
758:molecular biological
711:Raynaud's phenomenon
543:small nucleolar RNAs
454:phosphodiester bonds
171:motor neuron disease
3117:2008Sci...322.1851P
3070:10.1038/nature07747
3062:2009Natur.457.1038N
2923:Biochem. Soc. Trans
2387:10.1261/rna.2060405
2101:10.1093/nar/gkn1020
2054:10.1038/nature07480
2046:2008Natur.456..993L
1912:10.1110/ps.03477004
770:exosome component 3
660:immunoprecipitation
624:autoimmune diseases
394:Regulatory proteins
324:Associated proteins
310:inorganic phosphate
300:related and have a
203:years earlier (see
201:autoimmune diseases
163:exosome component 3
144:autoimmune diseases
3490:10.1007/BF01451281
2935:10.1042/BST0300150
2890:10.1261/rna.575107
2841:10.1093/nar/gki982
658:on HEp-2 cells or
656:immunofluorescence
587:
490:
488:for a full legend.
441:
431:Enzymatic function
376:
258:
238:
58:
3904:Protein complexes
3369:10.1002/art.22710
3281:10.1002/art.20056
3056:(7232): 1038β42.
2297:(16): 3187β3195.
2285:Mian, IS (1997).
1865:(34): 32397β404.
1534:10.1101/gr.162001
1225:
1224:
782:PCH1B; MIM 614678
727:immunosuppressive
707:clinical symptoms
675:. More recently,
596:protein synthesis
422:(Trf4) activity.
230:crystal structure
32:Exosome (vesicle)
16:(Redirected from
3911:
3843:
3806:
3796:
3771:
3745:
3736:(6): 1372β1379.
3724:
3714:
3681:
3652:
3609:
3608:
3598:
3565:
3559:
3558:
3540:
3516:
3510:
3509:
3473:
3467:
3466:
3438:
3432:
3431:
3421:
3411:
3387:
3381:
3380:
3352:
3346:
3345:
3335:
3325:
3299:
3293:
3292:
3264:
3258:
3257:
3229:
3223:
3222:
3212:
3180:
3174:
3173:
3145:
3139:
3138:
3128:
3111:(5909): 1851β4.
3096:
3090:
3089:
3045:
3039:
3038:
3020:
2996:
2990:
2989:
2953:
2947:
2946:
2918:
2912:
2911:
2901:
2869:
2863:
2862:
2852:
2835:(21): 6795β804.
2820:
2814:
2813:
2803:
2786:(19): 5399β410.
2771:
2762:
2761:
2751:
2742:(27): 19958β68.
2727:
2721:
2720:
2710:
2678:
2672:
2671:
2661:
2637:
2631:
2630:
2620:
2596:
2590:
2589:
2571:
2545:
2539:
2538:
2519:10.1038/nsmb1184
2502:
2493:
2492:
2490:
2464:
2458:
2457:
2439:
2415:
2409:
2408:
2398:
2366:
2360:
2359:
2331:
2325:
2324:
2314:
2282:
2276:
2275:
2265:
2242:The EMBO Journal
2233:
2224:
2223:
2213:
2190:The EMBO Journal
2181:
2172:
2171:
2161:
2129:
2123:
2122:
2112:
2080:
2074:
2073:
2031:
2022:
2016:
2015:
2005:
1981:
1975:
1974:
1964:
1940:
1934:
1933:
1923:
1891:
1885:
1884:
1874:
1850:
1844:
1843:
1806:
1800:
1799:
1789:
1760:
1754:
1753:
1743:
1711:
1702:
1701:
1665:
1659:
1658:
1624:
1607:
1606:
1596:
1562:
1556:
1555:
1545:
1510:
1504:
1503:
1485:
1461:
1455:
1454:
1444:
1434:
1410:
1404:
1403:
1393:
1361:
1355:
1354:
1336:
1310:
1273:
1268:
1267:
1251:
1242:
1233:
792:
788:List of subunits
733:Cancer treatment
562:RNA interference
365:
338:exoribonucleases
306:exoribonucleases
191:, an often-used
146:(especially the
51:
21:
3919:
3918:
3914:
3913:
3912:
3910:
3909:
3908:
3879:
3878:
3850:
3809:
3774:
3743:10.1.1.232.6756
3727:
3684:
3655:
3641:
3620:
3617:
3615:Further reading
3612:
3587:10.1038/ng.2254
3575:Nature Genetics
3567:
3566:
3562:
3518:
3517:
3513:
3475:
3474:
3470:
3440:
3439:
3435:
3389:
3388:
3384:
3354:
3353:
3349:
3301:
3300:
3296:
3266:
3265:
3261:
3231:
3230:
3226:
3182:
3181:
3177:
3147:
3146:
3142:
3098:
3097:
3093:
3047:
3046:
3042:
2998:
2997:
2993:
2970:10.1038/nrm1964
2955:
2954:
2950:
2920:
2919:
2915:
2871:
2870:
2866:
2822:
2821:
2817:
2773:
2772:
2765:
2729:
2728:
2724:
2680:
2679:
2675:
2639:
2638:
2634:
2598:
2597:
2593:
2547:
2546:
2542:
2504:
2503:
2496:
2466:
2465:
2461:
2417:
2416:
2412:
2381:(8): 1291β302.
2368:
2367:
2363:
2333:
2332:
2328:
2284:
2283:
2279:
2248:(14): 2342β57.
2235:
2234:
2227:
2196:(14): 2358β67.
2183:
2182:
2175:
2131:
2130:
2126:
2082:
2081:
2077:
2040:(7224): 993β6.
2029:
2024:
2023:
2019:
1983:
1982:
1978:
1955:(11): 1215β26.
1942:
1941:
1937:
1900:Protein Science
1893:
1892:
1888:
1852:
1851:
1847:
1808:
1807:
1803:
1762:
1761:
1757:
1713:
1712:
1705:
1682:10.1038/nsmb952
1667:
1666:
1662:
1647:
1626:
1625:
1610:
1564:
1563:
1559:
1522:Genome Research
1512:
1511:
1507:
1463:
1462:
1458:
1412:
1411:
1407:
1376:(16): 2148β58.
1363:
1362:
1358:
1312:
1311:
1304:
1300:
1269:
1262:
1259:
1246:
1237:
1228:
790:
766:
735:
640:dermatomyositis
616:
611:
519:normal turnover
495:
446:exoribonuclease
433:
428:
420:polyadenylation
401:AU-rich element
396:
361:
326:
302:common ancestor
222:
217:
179:
62:exosome complex
47:
35:
28:
23:
22:
15:
12:
11:
5:
3917:
3915:
3907:
3906:
3901:
3896:
3891:
3881:
3880:
3877:
3876:
3871:
3866:
3861:
3856:
3849:
3848:External links
3846:
3845:
3844:
3818:(7): 562β564.
3807:
3787:(4): 651β654.
3772:
3725:
3697:(7): 651β657.
3682:
3664:(4): 239β246.
3653:
3639:
3616:
3613:
3611:
3610:
3581:(6): 704β708.
3560:
3511:
3468:
3433:
3409:10.1186/ar1729
3402:(3): R704β13.
3382:
3363:(7): 2449β54.
3347:
3294:
3259:
3224:
3175:
3140:
3091:
3040:
2991:
2948:
2913:
2884:(7): 1027β35.
2864:
2815:
2763:
2722:
2673:
2646:Molecular Cell
2632:
2605:Molecular Cell
2591:
2562:(6): 1223β37.
2540:
2494:
2481:(1): 188β189.
2459:
2410:
2361:
2326:
2277:
2225:
2173:
2138:Molecular Cell
2124:
2095:(4): 1127β40.
2075:
2017:
1976:
1935:
1886:
1845:
1818:(4): 1076β89.
1801:
1755:
1703:
1660:
1645:
1608:
1557:
1505:
1456:
1425:(9): 6177β84.
1405:
1356:
1327:(4): 457β466.
1301:
1299:
1296:
1295:
1294:
1283:
1275:
1274:
1271:Biology portal
1258:
1255:
1254:
1253:
1244:
1235:
1223:
1222:
1215:
1205:
1202:
1199:
1196:
1190:
1187:
1184:
1180:
1179:
1172:
1167:
1164:
1161:
1158:
1155:
1152:
1149:
1145:
1144:
1137:
1132:
1129:
1126:
1123:
1120:
1117:
1114:
1110:
1109:
1102:
1097:
1094:
1091:
1088:
1085:
1082:
1079:
1075:
1074:
1067:
1062:
1059:
1056:
1053:
1050:
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1044:
1040:
1039:
1032:
1027:
1024:
1021:
1018:
1015:
1012:
1009:
1005:
1004:
997:
992:
989:
986:
983:
980:
977:
974:
970:
969:
962:
957:
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951:
948:
945:
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935:
934:
927:
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916:
913:
910:
907:
904:
900:
899:
892:
887:
884:
881:
878:
875:
872:
869:
865:
864:
857:
852:
849:
846:
843:
840:
837:
834:
830:
829:
826:
821:
815:
812:
805:
802:
799:
796:
789:
786:
765:
762:
739:antimetabolite
734:
731:
665:PM/Scl complex
620:autoantibodies
615:
612:
610:
607:
515:non-stop decay
494:
491:
432:
429:
427:
424:
395:
392:
382:(in yeast) or
325:
322:
298:evolutionarily
221:
218:
216:
213:
197:PM/Scl complex
193:model organism
178:
175:
155:chemotherapies
140:autoantibodies
76:intracellular
66:PM/Scl complex
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3916:
3905:
3902:
3900:
3899:Ribonucleases
3897:
3895:
3892:
3890:
3889:Nucleic acids
3887:
3886:
3884:
3875:
3872:
3870:
3867:
3865:
3862:
3860:
3857:
3855:
3852:
3851:
3847:
3841:
3837:
3833:
3829:
3825:
3821:
3817:
3813:
3808:
3804:
3800:
3795:
3790:
3786:
3782:
3778:
3773:
3769:
3765:
3761:
3757:
3753:
3749:
3744:
3739:
3735:
3731:
3726:
3722:
3718:
3713:
3708:
3704:
3700:
3696:
3692:
3688:
3683:
3679:
3675:
3671:
3667:
3663:
3659:
3654:
3650:
3646:
3642:
3640:9780123743787
3636:
3632:
3628:
3624:
3619:
3618:
3614:
3606:
3602:
3597:
3592:
3588:
3584:
3580:
3576:
3572:
3564:
3561:
3556:
3552:
3548:
3544:
3539:
3534:
3531:(1): 121β37.
3530:
3526:
3522:
3515:
3512:
3507:
3503:
3499:
3495:
3491:
3487:
3483:
3479:
3472:
3469:
3464:
3460:
3456:
3452:
3448:
3444:
3437:
3434:
3429:
3425:
3420:
3415:
3410:
3405:
3401:
3397:
3393:
3386:
3383:
3378:
3374:
3370:
3366:
3362:
3358:
3351:
3348:
3343:
3339:
3334:
3329:
3324:
3323:10.1186/ar389
3319:
3315:
3311:
3310:
3305:
3298:
3295:
3290:
3286:
3282:
3278:
3274:
3270:
3263:
3260:
3255:
3251:
3247:
3243:
3240:(2): 226β30.
3239:
3235:
3228:
3225:
3220:
3216:
3211:
3206:
3202:
3198:
3194:
3190:
3186:
3179:
3176:
3171:
3167:
3163:
3159:
3156:(6): 704β10.
3155:
3151:
3144:
3141:
3136:
3132:
3127:
3122:
3118:
3114:
3110:
3106:
3102:
3095:
3092:
3087:
3083:
3079:
3075:
3071:
3067:
3063:
3059:
3055:
3051:
3044:
3041:
3036:
3032:
3028:
3024:
3019:
3014:
3011:(5): 725β37.
3010:
3006:
3002:
2995:
2992:
2987:
2983:
2979:
2975:
2971:
2967:
2964:(7): 529β39.
2963:
2959:
2952:
2949:
2944:
2940:
2936:
2932:
2928:
2924:
2917:
2914:
2909:
2905:
2900:
2895:
2891:
2887:
2883:
2879:
2875:
2868:
2865:
2860:
2856:
2851:
2846:
2842:
2838:
2834:
2830:
2826:
2819:
2816:
2811:
2807:
2802:
2797:
2793:
2789:
2785:
2781:
2777:
2770:
2768:
2764:
2759:
2755:
2750:
2745:
2741:
2737:
2733:
2726:
2723:
2718:
2714:
2709:
2704:
2700:
2696:
2693:(2): 773β84.
2692:
2688:
2684:
2677:
2674:
2669:
2665:
2660:
2655:
2652:(3): 675β87.
2651:
2647:
2643:
2636:
2633:
2628:
2624:
2619:
2614:
2611:(3): 473β81.
2610:
2606:
2602:
2595:
2592:
2587:
2583:
2579:
2575:
2570:
2565:
2561:
2557:
2556:
2551:
2544:
2541:
2536:
2532:
2528:
2524:
2520:
2516:
2512:
2508:
2501:
2499:
2495:
2489:
2484:
2480:
2476:
2475:
2470:
2463:
2460:
2455:
2451:
2447:
2443:
2438:
2433:
2430:(5): 713β24.
2429:
2425:
2421:
2414:
2411:
2406:
2402:
2397:
2392:
2388:
2384:
2380:
2376:
2372:
2365:
2362:
2357:
2353:
2349:
2345:
2342:(5): 175β83.
2341:
2337:
2330:
2327:
2322:
2318:
2313:
2308:
2304:
2300:
2296:
2292:
2288:
2281:
2278:
2273:
2269:
2264:
2259:
2255:
2251:
2247:
2243:
2239:
2232:
2230:
2226:
2221:
2217:
2212:
2207:
2203:
2199:
2195:
2191:
2187:
2180:
2178:
2174:
2169:
2165:
2160:
2155:
2151:
2147:
2144:(2): 324β31.
2143:
2139:
2135:
2128:
2125:
2120:
2116:
2111:
2106:
2102:
2098:
2094:
2090:
2086:
2079:
2076:
2071:
2067:
2063:
2059:
2055:
2051:
2047:
2043:
2039:
2035:
2028:
2021:
2018:
2013:
2009:
2004:
1999:
1996:(4): 523β32.
1995:
1991:
1987:
1980:
1977:
1972:
1968:
1963:
1958:
1954:
1950:
1946:
1939:
1936:
1931:
1927:
1922:
1917:
1913:
1909:
1906:(3): 668β77.
1905:
1901:
1897:
1890:
1887:
1882:
1878:
1873:
1868:
1864:
1860:
1856:
1849:
1846:
1841:
1837:
1833:
1829:
1825:
1821:
1817:
1813:
1805:
1802:
1797:
1793:
1788:
1783:
1779:
1775:
1772:(4): 653β63.
1771:
1767:
1759:
1756:
1751:
1747:
1742:
1737:
1733:
1729:
1726:(6): 1093β5.
1725:
1721:
1717:
1710:
1708:
1704:
1699:
1695:
1691:
1687:
1683:
1679:
1676:(7): 575β81.
1675:
1671:
1664:
1661:
1656:
1652:
1648:
1646:9780123646552
1642:
1638:
1634:
1630:
1623:
1621:
1619:
1617:
1615:
1613:
1609:
1604:
1600:
1595:
1590:
1586:
1582:
1579:(9): 889β93.
1578:
1574:
1573:
1568:
1561:
1558:
1553:
1549:
1544:
1539:
1535:
1531:
1528:(2): 240β52.
1527:
1523:
1519:
1515:
1509:
1506:
1501:
1497:
1493:
1489:
1484:
1479:
1476:(4): 451β64.
1475:
1471:
1467:
1460:
1457:
1452:
1448:
1443:
1438:
1433:
1428:
1424:
1420:
1416:
1409:
1406:
1401:
1397:
1392:
1387:
1383:
1379:
1375:
1371:
1367:
1360:
1357:
1352:
1348:
1344:
1340:
1335:
1330:
1326:
1322:
1321:
1316:
1309:
1307:
1303:
1297:
1292:
1288:
1284:
1281:
1277:
1276:
1272:
1266:
1261:
1256:
1250:
1249:
1245:
1241:
1240:
1236:
1232:
1231:
1227:
1226:
1221:
1220:
1216:
1214:
1213:
1209:
1206:
1203:
1200:
1197:
1195:
1191:
1188:
1185:
1181:
1178:
1177:
1173:
1171:
1168:
1165:
1162:
1159:
1156:
1153:
1150:
1146:
1143:
1142:
1138:
1136:
1133:
1130:
1127:
1124:
1121:
1118:
1115:
1111:
1108:
1107:
1103:
1101:
1098:
1095:
1092:
1089:
1086:
1083:
1080:
1076:
1073:
1072:
1068:
1066:
1063:
1060:
1057:
1054:
1051:
1048:
1045:
1041:
1038:
1037:
1033:
1031:
1028:
1025:
1022:
1019:
1016:
1013:
1010:
1006:
1003:
1002:
998:
996:
993:
990:
987:
984:
981:
978:
975:
971:
968:
967:
963:
961:
958:
955:
952:
949:
946:
943:
940:
936:
933:
932:
928:
926:
923:
920:
917:
914:
911:
908:
905:
901:
898:
897:
893:
891:
888:
885:
882:
879:
876:
873:
870:
866:
863:
862:
858:
856:
853:
850:
847:
844:
841:
838:
835:
831:
827:
825:
822:
819:
816:
813:
810:
809:S. cerevisiae
806:
803:
800:
798:General name
797:
794:
793:
787:
785:
783:
779:
775:
771:
768:Mutations in
763:
761:
759:
755:
751:
747:
743:
740:
732:
730:
728:
724:
720:
716:
712:
708:
703:
701:
697:
693:
689:
684:
682:
678:
674:
670:
666:
661:
657:
653:
649:
645:
641:
637:
633:
629:
625:
621:
613:
608:
606:
604:
599:
597:
593:
583:
579:
577:
576:
572:, called the
571:
567:
563:
559:
555:
551:
546:
544:
540:
539:ribosomal RNA
536:
532:
531:small nuclear
528:
524:
520:
516:
512:
508:
507:messenger RNA
504:
500:
492:
487:
481:
477:
473:
469:
467:
463:
459:
455:
451:
447:
437:
430:
425:
423:
421:
417:
416:TRAMP complex
413:
409:
404:
402:
393:
391:
389:
385:
381:
373:
369:
364:
358:
354:
352:
348:
343:
339:
335:
331:
323:
321:
319:
315:
311:
307:
303:
299:
294:
290:
286:
282:
278:
274:
269:
267:
263:
255:
249:
245:
243:
235:
231:
226:
220:Core proteins
219:
214:
212:
210:
206:
202:
198:
194:
190:
189:
184:
176:
174:
172:
168:
164:
160:
156:
153:
152:antimetabolic
149:
145:
141:
136:
134:
130:
126:
125:ribosomal RNA
122:
121:messenger RNA
118:
114:
110:
106:
101:
99:
95:
91:
87:
83:
79:
75:
72:) is a multi-
71:
67:
63:
55:
50:
46:
41:
37:
33:
19:
3815:
3811:
3784:
3780:
3733:
3729:
3694:
3691:EMBO Reports
3690:
3661:
3657:
3622:
3578:
3574:
3563:
3528:
3524:
3514:
3484:(6): 465β7.
3481:
3477:
3471:
3449:(7): 432β7.
3446:
3442:
3436:
3399:
3395:
3385:
3360:
3356:
3350:
3316:(2): 134β8.
3313:
3307:
3297:
3275:(2): 565β9.
3272:
3268:
3262:
3237:
3233:
3227:
3195:(1): 59β64.
3192:
3188:
3178:
3153:
3149:
3143:
3108:
3104:
3094:
3053:
3049:
3043:
3008:
3004:
2994:
2961:
2957:
2951:
2929:(2): 150β5.
2926:
2922:
2916:
2881:
2877:
2867:
2832:
2828:
2818:
2783:
2780:EMBO Journal
2779:
2739:
2735:
2725:
2690:
2686:
2676:
2649:
2645:
2635:
2608:
2604:
2594:
2559:
2553:
2543:
2513:(1): 15β22.
2510:
2506:
2478:
2472:
2462:
2427:
2423:
2413:
2378:
2374:
2364:
2339:
2335:
2329:
2294:
2290:
2280:
2245:
2241:
2193:
2189:
2141:
2137:
2127:
2092:
2088:
2078:
2037:
2033:
2020:
1993:
1989:
1979:
1952:
1948:
1938:
1903:
1899:
1889:
1862:
1858:
1848:
1815:
1811:
1804:
1769:
1765:
1758:
1723:
1719:
1673:
1669:
1663:
1628:
1576:
1572:EMBO Reports
1570:
1560:
1525:
1521:
1508:
1473:
1469:
1459:
1422:
1418:
1408:
1373:
1369:
1359:
1324:
1318:
1291:RNA splicing
1248:
1239:
1230:
1217:
1210:
1198:Rrp44p/Dis3p
1193:
1174:
1139:
1104:
1069:
1034:
999:
964:
950:Rrp41p/Ski6p
929:
894:
859:
808:
767:
746:chemotherapy
742:fluorouracil
736:
704:
691:
687:
685:
664:
636:polymyositis
617:
614:Autoimmunity
600:
588:
573:
552:or from the
547:
496:
474:
470:
456:β releasing
442:
405:
397:
383:
379:
377:
341:
329:
327:
293:mitochondria
289:chloroplasts
270:
259:
239:
196:
186:
180:
137:
102:
69:
65:
61:
59:
36:
1787:2066/186665
1442:2066/186951
1287:spliceosome
845:Csl4p/Ski4p
828:Yeast gene
723:scleroderma
677:recombinant
650:using calf
634:and either
632:scleroderma
575:degradosome
408:Ski complex
370:in red and
320:molecules.
262:RNA binding
209:genome data
169:and spinal
150:) and some
98:degradosome
92:, while in
3883:Categories
1514:Koonin, EV
1298:References
1280:proteasome
1157:PM/Scl-100
824:Human gene
688:PM/Scl-100
654:extracts,
558:expression
499:processing
493:Substrates
384:PM/Scl-100
374:in yellow.
129:small RNAs
117:substrates
88:cells and
86:eukaryotic
3738:CiteSeerX
1949:Structure
1122:PM/Scl-75
909:S1/KH RBD
874:S1/KH RBD
780:type 1B (
715:arthritis
692:PM/Scl-75
673:synthesis
669:nucleolus
592:ribosomes
535:nucleolus
503:cytoplasm
462:hydrolyse
368:Ξ±-helices
215:Structure
177:Discovery
113:nucleolus
105:cytoplasm
3840:43218090
3832:15999107
3803:16713559
3760:16968219
3721:17603538
3678:18211833
3649:19111178
3605:22544365
3555:11922219
3547:14718172
3506:39237322
3463:17643929
3428:15899056
3377:17599775
3342:11879549
3289:14872500
3170:12410095
3135:19056938
3078:19169244
3027:15935759
2986:22499032
2978:16829983
2943:12035760
2908:17545563
2859:16396833
2810:10508172
2758:17470429
2717:17660569
2687:Genetics
2668:14527413
2627:16285928
2586:62785677
2578:17174896
2535:24691764
2527:17173052
2454:14898055
2446:15935758
2405:16043509
2356:15346807
2272:20531386
2220:20531389
2168:17643380
2119:19129231
2062:19060886
2012:17514363
1971:11080643
1930:14767080
1881:12746447
1840:27114625
1832:17078816
1796:12419256
1750:17174886
1690:15951817
1655:16939780
1603:12947419
1552:11157787
1500:14817671
1492:11719186
1451:11110791
1400:10465791
1351:16035676
1257:See also
1119:RNase PH
1084:RNase PH
1049:RNase PH
1014:RNase PH
979:RNase PH
944:RNase PH
814:Archaea
801:Domains
719:myositis
566:Bacteria
426:Function
412:helicase
372:Ξ²-sheets
287:and the
285:bacteria
273:RNase PH
242:RNase PH
94:bacteria
52:See the
3768:6872855
3712:1905902
3596:3366034
3498:9890673
3419:1174964
3254:3918546
3219:2199097
3210:1535032
3113:Bibcode
3105:Science
3086:4329373
3058:Bibcode
3035:1390706
2899:1894934
2850:1310903
2801:1171609
2708:2034642
2396:1370812
2321:9241229
2263:2910271
2211:2910272
2159:7610968
2110:2651783
2070:1808371
2042:Bibcode
1921:2286726
1741:1986773
1698:2003922
1594:1326366
1343:9390555
1219:YOL021C
1204:105β113
1194:Dis3L1
1189:RNase R
1176:YOR001W
1170:EXOSC10
1154:RNase D
1141:YDR280W
1128:(Rrp42)
1106:YCR035C
1093:(Rrp42)
1071:YDL111C
1036:YGR158C
1023:(Rrp41)
1001:YGR095C
988:(Rrp41)
966:YGR195W
931:YOL142W
896:YHR069C
861:YNL232W
807:Yeast (
795:Legend
696:peptide
609:Disease
527:nucleus
388:RNase D
334:RNase R
109:nucleus
90:archaea
78:complex
74:protein
70:exosome
3838:
3830:
3801:
3766:
3758:
3740:
3719:
3709:
3676:
3647:
3637:
3603:
3593:
3553:
3545:
3504:
3496:
3461:
3426:
3416:
3375:
3340:
3330:
3287:
3252:
3217:
3207:
3168:
3133:
3084:
3076:
3050:Nature
3033:
3025:
2984:
2976:
2941:
2906:
2896:
2857:
2847:
2808:
2798:
2756:
2715:
2705:
2666:
2625:
2584:
2576:
2533:
2525:
2452:
2444:
2403:
2393:
2354:
2319:
2312:146874
2309:
2270:
2260:
2218:
2208:
2166:
2156:
2117:
2107:
2068:
2060:
2034:Nature
2010:
1969:
1928:
1918:
1879:
1838:
1830:
1794:
1748:
1738:
1696:
1688:
1653:
1643:
1601:
1591:
1550:
1543:311015
1540:
1498:
1490:
1449:
1398:
1391:316947
1388:
1349:
1341:
1212:DIS3L1
1166:84β100
1135:EXOSC9
1125:Rrp45p
1100:EXOSC8
1090:Rrp43p
1065:EXOSC7
1055:Rrp42p
1052:hRrp42
1030:EXOSC6
995:EXOSC5
985:Rrp46p
982:hRrp46
960:EXOSC4
947:hRrp41
925:EXOSC3
918:(Rrp4)
915:Rrp40p
912:hRrp40
890:EXOSC2
855:EXOSC1
839:S1 RBD
804:Human
754:tumors
750:cancer
652:thymus
570:PNPase
554:5' end
450:3' end
347:DIS3L1
314:3' end
277:PNPase
256:below.
254:legend
236:below.
234:legend
165:cause
159:cancer
133:3β² end
54:legend
3836:S2CID
3764:S2CID
3551:S2CID
3502:S2CID
3333:83843
3082:S2CID
3031:S2CID
2982:S2CID
2582:S2CID
2531:S2CID
2450:S2CID
2066:S2CID
2030:(PDF)
1836:S2CID
1694:S2CID
1496:S2CID
1347:S2CID
1192:Dis3
1186:Rrp44
1160:Rrp6p
1131:34β49
1116:Rrp45
1096:30β44
1081:Rrp43
1061:29β32
1058:Rrp42
1046:Rrp42
1026:24β37
1020:Mtr3p
1017:hMtr3
991:25β28
976:Rrp46
956:26β28
953:Rrp41
941:Rrp41
921:27β32
906:Rrp40
886:28β39
880:Rrp4p
877:hRrp4
851:21β32
842:hCsl4
820:(kD)
700:ELISA
644:serum
486:below
366:with
330:Rrp44
205:below
183:RNase
18:Rrp44
3828:PMID
3799:PMID
3781:Cell
3756:PMID
3717:PMID
3674:PMID
3662:1779
3645:PMID
3635:ISBN
3601:PMID
3543:PMID
3525:Cell
3494:PMID
3459:PMID
3424:PMID
3373:PMID
3338:PMID
3285:PMID
3250:PMID
3215:PMID
3166:PMID
3131:PMID
3074:PMID
3023:PMID
3005:Cell
2974:PMID
2939:PMID
2904:PMID
2855:PMID
2806:PMID
2754:PMID
2713:PMID
2664:PMID
2623:PMID
2574:PMID
2555:Cell
2523:PMID
2474:Cell
2442:PMID
2424:Cell
2401:PMID
2352:PMID
2317:PMID
2268:PMID
2216:PMID
2164:PMID
2115:PMID
2058:PMID
2008:PMID
1967:PMID
1926:PMID
1877:PMID
1828:PMID
1792:PMID
1746:PMID
1720:Cell
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