Table 1 Proteins present in the SSU processome
Protein
Nop1
Nop56
Alias
Gene
YDL014w
YLR197w
MW(kDa)
34.4
56.7
Nop5/58
YOR310c
56.8
Snu13
YEL026w
13.4
Sof1
YLL011w
56.8
Yes
Mpp10
YJR002w
66.8
Yes
Imp3
YHR148w
21.8
Yes
NP_060755 (54%)
Imp4
YNL075w
33.5
Yes
NNP77
Dhr1
YMR128w
144.8
Yes
CAB45191 (28%)
Rrp9
Rrp5
YPR137w
YMR229c
64.9
193.0
Yes
Yes
Sik1
Essential?
Homolog
Yes
NP_001427 (67%);FBL
Yes
CAB43677 (24%);
NOP56
Yes
AAD27610 (47%);
NOP5/58
Yes
NP_004999 (71%);
NHP2L1/NHPX
NP_056235 (46%);
NNP34
CAA67120 (30%);
MPP10
Utp1
Pwp2
YCR057c
103.9
Yes
Utp2
Nop14
YDL148c
94.3
Yes
NP_004695 (28%)
AAA77669 (27%);
RRP5
NP_005040 (43%);
NNP62
BAA19121 (25%)
Utp3
Sas10
YDL153c
70.1
Yes
NP_065101 (22%)
Utp4
YDR324c
87.8
Yes
Utp5
YDR398w
71.9
Yes
CAB04569 (21%) CE
AAF49526 (22%) DM
NNP53
Utp6
YDR449c
52.3
Yes
NP_06898 (32%)
Utp7
YER082c
62.3
Yes
THC513304 (44%);
NNP24
Utp8
Utp9
Utp10
YGR128c
YHR196w
YJL109c
80.1
65.1
199.9
Yes
Yes
Yes
Utp11
YKL099c
30.3
Yes
YLR129w
106.3
Yes
Utp13
Utp14
YLR222c
YML093w
91.0
102.9
Yes
Yes
Utp15
YMR093w
57.5
Yes
No; coldsensitive
Yes
Utp12
Dip2
Utp16
Bud21
YOR078w
24.2
Utp17
Nan1
YPL126w
101.1
NP_060542 (27%);
NNP71
THC497115 (32%);
NNP13
NP_006775 (37%);
NNP63
NNP43
BAA13396 (29%)
AAF60457 (23%) CE
AAF45765 (32%) DM
CAB11248 (21%) SP
Motif/Comment
methyltransferase
KKE/D
Citations
1-4
5,6
KKE/D
5,7,8
binds box C/D
snoRNAs, U4
snRNA
WD rpts.
9-11
coiled-coils;
Imp3, Imp4,
Utp3-interacting
S4 RBD; Mpp10interacting
RNA binding
superfamily;
Mpp10interacting
DEAH box
helicase
WD rpts.
S1 RBD;
TPR rpts
WD rpts.
13,14
coiled-coils
NAP family;
silencing; Mpp10interacting
WD rpts.
WD rpts.; Utp15interacting
crooked neck-like
TPR
WD rpts.;
adenylate binding
site
9,12
9,15
9,15,16
17
18,19
9,20
This work;
9,21,22
This work;
23,24
This work;
25
This work.
This work; 9
This work.
This work; 9
coiled-coils
HEAT rpts.
This work.
This work.
This work; 9
coiled-coils
This work; 9
WD rpts.
This work;
9,26
This work; 9
This work.
WD rpts.
coiled-coils;
ATP/GTP binding
site (P-loop)
WD rpts.;
Utp5-interacting
Coiled-coils
WD rpts.; exit
from mitosis
This work.
This work;
27
This work;
28
Information was compiled using the sources and computer analysis software listed in the Methods
except that the disruption phenotype of UTP16 was tested directly. The homologs are from humans
unless otherwise listed. CE, C. elegans; DM, D. melanogaster; SP, S. pombe. U3 processome
purification was carried out twice. This list along with the 5 ribosomal proteins and the 3
contaminants mentioned in the text represent all of the proteins analyzed from the first purification
plus all of the proteins present as more than one peptide from the second purification.
1. Henriquez, R. G., Blobel, G. & Aris, J. P. Isolation and sequencing of NOP1. A yeast gene encoding a nucleolar
protein homologous to a human autoimmune antigen. Journal of Biological Chemistry 265, 2209-2215 (1990).
2. Schimmang, T., Tollervey, D., Kern, H., Frank, R. & Hurt, E. C. A yeast nucleolar protein related to mammalian
fibrillarin is associated with small nucleolar RNA and is essential for viability. EMBO Journal 8, 4015-4024 (1989).
3. Tollervey, D., Lehtonen, H., Carmo-Fonseca, M. & Hurt, E. C. The small nucleolar RNP protein NOP1 (fibrillarin) is
required for pre-rRNA processing in yeast. EMBO Journal 10, 573-583 (1991).
4. Aris, J. P. & Blobel, G. cDNA cloning and sequencing of human fibrillarin, a conserved nucleolar protein recognized
by autoimmune antisera. Proc Natl Acad Sci U S A 88, 931-935 (1991).
5. Gautier, T., Berges, T., Tollervey, D. & Hurt, E. Nucleolar KKE/D repeat proteins Nop56p and Nop58p interact with
Nop1p and are required for ribosome biogenesis. Mol Cell Biol 17, 7088-7098 (1997).
6. Morin, J. P., Downs, J. A., Snodgrass, A. M. & Gilmore, T. D. Genetic analysis of growth inhibition by GAL4-IB-
in Saccharomyces cerevisiae. Cell Growth Differ. 6, 789-798 (1995).
7. Lyman, S. K., Gerace, L. & Baserga, S. J. Human Nop5/Nop58 is a component common to the box C/D small
nucleolar ribonucleoproteins. RNA 5, 1597-1604 (1999).
8. Wu, P., Brockenbrough, J. S., Metcalfe, A. C., Chen, S. & Aris, J. P. Nop5p is a small nucleolar ribonucleoprotein
component required for pre-18 S rRNA processing in yeast. J Biol Chem 273, 16453-16463 (1998).
9. Andersen, J. S. et al. Directed proteomic analysis of the human nucleolus. Curr. Biol. 12, 1-11 (2002).
10.Saito, H., Fujiwara, T., Shin, S., Okui, K. & Nakamura, Y. Cloning and mapping of a human novel cDNA
(NHP2L1) that encodes a protein highly homologous to yeast nuclear protein NHP2. Cytogenet. Cell. Genet. 72, 191193 (1996).
11.Watkins, N. J. et al. A common core RNP structure shared between the small nucleolar box C/D RNPs and the
spliceosomal U4 snRNP. Cell 103, 457-466 (2000).
12.Jansen, R., Tollervey, D. & Hurt, E. C. A U3 snoRNP protein with homology to splicing factor PRP4 and G beta
domains is required for ribosomal RNA processing. EMBO Journal 12, 2549-2558 (1993).
13.Dunbar, D. A., Wormsley, S., Agentis, T. M. & Baserga, S. J. Mpp10p, a U3 small nucleolar ribonucleoprotein
component required for pre-18S rRNA processing in yeast. Mol Cell Biol 17, 5803-5812 (1997).
14.Westendorf, J. M. et al. M phase phosphoprotein 10 is a human U3 small nucleolar ribonucleoprotein component.
Mol Biol Cell 9, 437-449 (1998).
15.Lee, S. J. & Baserga, S. J. Imp3p and Imp4p: two specific components of the U3 small nucleolar ribonucleoprotein
that are essential for pre-18S rRNA processing. Molecular and Cellular Biology 19, 5441-5452 (1999).
16.Wehner, K. A. & Baserga, S. J. The 70-like motif: a eukaryotic RNA binding domain unique to a superfamily of
proteins required for ribosome biogenesis. Mol. Cell. 9, 329-339 (2002).
17.Colley, A., Beggs, J. D., Tollervey, D. & Lafontaine, D. L. Dhr1p, a putative DEAH-Box RNA helicase, is
associated with the box C+D snoRNP U3. Mol. Cell. Biol. 20, 7238-7246 (2000).
18.Pluk, H., Soffner, J., Luhrmann, R. & van Venrooij, W. J. cDNA cloning and characterization of the human U3 small
nucleolar ribonucleoprotein complex-associated 55-kilodalton protein. Mol Cell Biol 18, 488-498 (1998).
19.Venema, J., Vos, H. R., Faber, A. W., van Venrooij, W. J. & Raue, H. A. Yeast Rrp9p is an evolutionarily conserved
U3 snoRNP protein essential for the early pre-rRNA processing cleavages and requires box C for its association. RNA
6, 1660-1671 (2000).
20.Venema, J. & Tollervey, D. Rrp5 is required for formation of both 18S and 5.8S rRNA in yeast. EMBO J. 15, 57015714 (1996).
21.Shafaatian, R., Payton, M. A. & Reid, J. D. PWP2, a member of the WD-repeat family of proteins, is an essential
Saccharomyces cerevisiae gene involved in cell separation. Mol Gen Genet 252, 101-114. (1996).
22.Lalioti, M. D. et al. Cloning the cDNA of human PWP2, which encodes a protein with WD repeats and maps to
21q22.3. Genomics 35, 321-327. (1996).
23.Rout, M. P. et al. The yeast nuclear pore complex: composition, architecture, and transport mechanism. J. Cell Biol.
148, 635-651 (2000).
24.Liu, P. C. & Thiele, D. J. Novel stress-responsive genes EMG1 and NOP14 encode conserved, interacting proteins
required for 40S ribosome biogenesis. Mol. Biol. Cell 12, 3644-3657 (2001).
25.Kamakaka, R. & Rine, J. Sir- and silencer-independent disruption of silencing in Saccharomyces by Sas10p.
Genetics 149, 903-914 (1998).
26.Storozhenko, S., Inze, D., Van Montagu, M. & Kushnir, S. Arabidopsis coactivator ALY-like proteins, DIP1 and
DIP2, interact physically with the DNA-binding domain of the Zn-finger poly(ADP-ribose) polymerase. J. Exp. Bot.
53, 1375-1380 (2001).
27.Ni, L. & Snyder, M. A genomic study of the bipolar bud site selection pattern in Saccharomyces cerevisiae. Mol Biol
Cell 12, 2147-2170. (2001).
28.Shou, W. et al. Exit from mitosis is triggered by Tem1-dependent release of the protein phosphatase Cdc14 from
nucleolar RENT complex. Cell 97, 233-244 (1999).