Ribosomal protein

A ribosomal protein (r-protein or rProtein[1][2][3]) is any of the proteins that, in conjunction with rRNA, make up the ribosomal subunits involved in the cellular process of translation. A large part of the knowledge about these organic molecules has come from the study of E. coli ribosomes. All ribosomal proteins have been isolated and many specific antibodies have been produced. These, together with electronic microscopy and the use of certain reactives, have allowed for the determination of the topography of the proteins in the ribosome. E. coli, other bacteria and Archaea have a 30S small subunit and a 50S large subunit, whereas humans and yeasts have a 40S small subunit and a 60S large subunit.[4] Equivalent subunits are frequently numbered differently between bacteria, Archaea, yeasts and humans.[5] More recently, a near-complete (near)atomic picture of the ribosomal proteins is emerging from the latest high-resolution cryo-EM data (including PDB ID: 5AFI).

Mitochondrial ribosomal protein L31
Identifiers
Symbol	L31
Pfam	PF09784
InterPro	IPR016340
PROSITE	PDOC00880
SCOPe	1m90 / SUPFAM
Available protein structures: Pfam   structures / ECOD   PDB RCSB PDB; PDBe; PDBj PDBsum structure summary

Conservation

A 2016 tree of life using 16 universally-conserved ribosomal protein sequences[6]

Ribosomal proteins are among the most highly conserved proteins across all life forms.[5] Among the 40 proteins found in various small ribosomal subunits (RPSs), 15 subunits are universally conserved across prokaryotes and eukaryotes. However, 7 subunits are only found in bacteria (S21, S6, S16, S18, S20, S21, and THX), while 17 subunits are only found in archea and eukaryotes.[5] Typically 22 proteins are found in bacterial small subunits and 32 in yeast, human and most likely most other eukaryotic species. Twenty-seven (out of 32) proteins of the eukaryotic small ribosomal subunit proteins are also present in archaea (no ribosomal protein is exclusively found in archaea), confirming that they are more closely related to eukaryotes than to eubacteria.[5]

Among the large ribosomal subunit (RPLs), 18 proteins are universal, i.e. found in both bacteria, eukaryotes, and archea. 14 proteins are only found in bacteria, while 27 proteins are only found in archea and eukaryotes. Again, archea have no proteins unique to them[5]

Essentiality

Despite their high conservation over billions of years of evolution, the absence of several ribosomal proteins in certain species shows that ribosomal subunits have been added and lost over the course of evolution. This is also reflected by the fact that several ribosomal proteins do not appear to be essential when deleted.[7] For instance, in E. coli nine ribosomal proteins (L15, L21, L24, L27, L29, L30, L34, S9, and S17) are nonessential for survival when deleted. Taken together with previous results, 22 of the 54 E. coli ribosomal protein genes can be individually deleted from the genome.[8] Similarly, 16 ribosomal proteins (L1, L9, L15, L22, L23, L28, L29, L32, L33.1, L33.2, L34, L35, L36, S6, S20, and S21) were successfully deleted in Bacillus subtilis. In conjunction with previous reports, 22 ribosomal proteins have been shown to be nonessential in B. subtilis, at least for cell proliferation.[9]

Assembly

In E. coli

The ribosome of E. coli has about 22 proteins in the small subunit (labelled S1 to S22) and 33 proteins in the large subunit (somewhat counter-intuitively called L1 to L36). All of them are different with three exceptions: one protein is found in both subunits (S20 and L26), L7 and L12 are acetylated and methylated forms of the same protein, and L8 is a complex of L7/L12 and L10. In addition, L31 is known to exist in two forms, the full length at 7.9 kilodaltons (kDa) and fragmented at 7.0 kDa. This is why the number of proteins in a ribosome is of 56. Except for S1 (with a molecular weight of 61.2 kDa), the other proteins range in weight between 4.4 and 29.7 kDa.[10]

Recent 'de novo' proteomics experiments where the authors characterized in vivo ribosome-assembly intermediates and associated assembly factors from wild-type Escherichia coli cells using a general quantitative mass spectrometry (qMS) approach have confirmed the presence of all the known small and large subunit components and have identified a total of 21 known and potentially new ribosome-assembly-factors that co-localise with various ribosomal particles.[11]

Disposition in the small ribosomal subunit

In the small (30S) subunit of E. coli ribosomes, the proteins denoted S4, S7, S8, S15, S17, S20 bind independently to 16S rRNA. After assembly of these primary binding proteins, S5, S6, S9, S12, S13, S16, S18, and S19 bind to the growing ribosome. These proteins also potentiate the addition of S2, S3, S10, S11, S14, and S21. Protein binding to helical junctions is important for initiating the correct tertiary fold of RNA and to organize the overall structure. Nearly all the proteins contain one or more globular domains. Moreover, nearly all contain long extensions that can contact the RNA in far-reaching regions. Additional stabilization results from the proteins' basic residues, as these neutralize the charge repulsion of the RNA backbone. Protein–protein interactions also exist to hold structure together by electrostatic and hydrogen bonding interactions. Theoretical investigations pointed to correlated effects of protein-binding onto binding affinities during the assembly process[12]

In eukaryotes

Ribosomes, which synthesize the proteome of cells, are complex ribonucleoproteins that, in eukaryotes, contain 79–80 proteins and four ribosomal RNAs(rRNAs). General or specialized chaperones solubilize the ribosomal proteins and facilitate their import into the nucleus. Assembly of the eukaryotic ribosome appears to be driven by the ribosomal proteins in vivo when assembly is also aided by chaperones. Most ribosomal proteins assemble with rRNA co-transcriptionally, becoming associated more stably as assembly proceeds, and the active sites of both subunits are constructed last.[5]

Table of ribosomal proteins

E.coli small 30S subunit

Subunit No.	Subunit name	E.coli protein	Pfam family with E.coli protein	Uniprot accession of E.coli protein
1	S1	RS1_ECOLI	PF00575	P0AG67
2	S2	RS2_ECOLI	PF00318	P0A7V0
3	S3	RS3_ECOLI	PF00189, PF07650	P0A7V3
4	S4	RS4_ECOLI	PF00163, PF01479	P0A7V8
5	S5	RS5_ECOLI	PF00333, PF03719	P0A7W1
6	S6	RS6_ECOLI	PF01250	P02358
7	S7	RS7_ECOLI	PF00177	P02359
8	S8	RS8_ECOLI	PF00410	P0A7W7
9	S9	RS9_ECOLI	PF00380	P0A7X3
10	S10	RS10_ECOLI	PF00338	P0A7R5
11	S11	RS11_ECOLI	PF00411	P0A7R9
12	S12	RS12_ECOLI	PF00164	P0A7S3
13	S13	RS13_ECOLI	PF00416	P0A7S9
14	S14	RS14_ECOLI	PF00253	P0AG59
15	S15	RS15_ECOLI	PF00312	P0ADZ4
16	S16	RS16_ECOLI	PF00886	P0A7T3
27	S17	RS17_ECOLI	PF00366	P0AG63
28	S18	RS18_ECOLI	PF01084	P0A7T7
29	S19	RS19_ECOLI	PF00203	P0A7U3
20	S20	RS20_ECOLI	PF01649	P0A7U7
21	S21	RS21_ECOLI	PF01165	P68681
22	S22	RS22_ECOLI	PF08136	C8U8F3

E.coli large 50S subunit

Subunit No.	Subunit name	E.coli protein	Pfam family with E.coli protein	Uniprot accession of E.coli protein
1	50S ribosomal protein L1	RL1_ECOLI	PF00687 Ribosomal protein L1p/L10e family	P0A7L0
2	50S ribosomal protein L2	RL2_ECOLI	PF03947 Ribosomal Proteins L2, C-terminal domain	P60422
2	50S ribosomal protein L2	RL2_ECOLI	PF00181 Ribosomal Proteins L2, RNA binding domain	P60422
3	50S ribosomal protein L3	RL3_ECOLI	PF00297 Ribosomal protein L3	P60438
4	50S ribosomal protein L4	RL4_ECOLI	PF00573 Ribosomal protein L4/L1 family	P60723
5	50S ribosomal protein L5	RL5_ECOLI	PF00281 Ribosomal protein L5	P62399
5	50S ribosomal protein L5	RL5_ECOLI	PF00673 ribosomal L5P family C-terminus	P62399
6	50S ribosomal protein L6	RL6_ECOLI	PF00347 Ribosomal protein L6	P0AG55
7/12	50S ribosomal protein L7/L12	RL7_ECOLI	PF16320 Ribosomal protein L7/L12 dimerisation domain	P0A7K2
8	50S ribosomal protein L7/L12	RL7_ECOLI	PF00542 Ribosomal protein L7/L12 C-terminal domain	P0A7K2
9	50S ribosomal protein L9	RL9_ECOLI	PF03948 Ribosomal protein L9, C-terminal domain	P0A7R1
9	50S ribosomal protein L9	RL9_ECOLI	PF01281 Ribosomal protein L9, N-terminal domain	P0A7R1
10	50S ribosomal protein L10	RL10_ECOLI	PF00466 Ribosomal protein L10	P0A7J3
11	50S ribosomal protein L11	RL11_ECOLI	PF03946 Ribosomal protein L11, N-terminal domain	P0A7J7
11	50S ribosomal protein L11	RL11_ECOLI	PF00298 Ribosomal protein L11, RNA binding domain	P0A7J7
13	50S ribosomal protein L13	RL13_ECOLI	PF00572 Ribosomal protein L13	P0AA10
14	50S ribosomal protein L14	RL14_ECOLI	PF00238 Ribosomal protein L14p/L23e	P0ADY3
15	50S ribosomal protein L15	RL15_ECOLI	PF00828 Ribosomal protein L18e/L15	P02413
16	50S ribosomal protein L16	RL16_ECOLI	PF00252 Ribosomal protein L16p/L10e	P0ADY7
17	50S ribosomal protein L17	RL17_ECOLI	PF01196 Ribosomal protein L17	P0AG44
18	50S ribosomal protein L18	RL18_ECOLI	PF00861 Ribosomal L18p/L5e family	P0C018
19	50S ribosomal protein L19	RL19_ECOLI	PF01245 Ribosomal protein L19	B1LPB3
20	50S ribosomal protein L20	RL20_ECOLI	PF00453 Ribosomal protein L20	P0A7L3
21	50S ribosomal protein L21	RL21_ECOLI	PF00829 Ribosomal prokaryotic L21 protein	P0AG48
22	50S ribosomal protein L22	RL22_ECOLI	PF00237 Ribosomal protein L22p/L17e	P61175
23	50S ribosomal protein L23	RL23_ECOLI	PF00276 Ribosomal protein L23	P0ADZ0
24	50S ribosomal protein L24	RL24_ECOLI	PF00467 KOW motif	P60624
25	50S ribosomal protein L25	RL25_ECOLI	PF01386 Ribosomal L25p family	P68919
27	50S ribosomal protein L27	RL27_ECOLI	PF01016 Ribosomal L27 protein	P0A7M0
28	50S ribosomal protein L28	RL28_ECOLI	PF00830 Ribosomal L28 family	P0A7M2
29	50S ribosomal protein L29	RL29_ECOLI	PF00831 Ribosomal L29 protein	P0A7M6
30	50S ribosomal protein L30	RL30_ECOLI	PF00327 Ribosomal protein L30p/L7e	P0AG51
31	50S ribosomal protein L31	RL31_ECOLI	PF01197 Ribosomal protein L31	P0A7M9
31B	50S ribosomal protein L31 type B	RL31B_ECOLI	PF01197 Ribosomal protein L31	P0A7M9
32	50S ribosomal protein L32	RL32_ECOLI	PF01783 Ribosomal L32p protein family	C4ZS29
33	50S ribosomal protein L33	RL33_ECOLI	PF00471 Ribosomal protein L33	P0A7N9
34	50S ribosomal protein L34	RL34_ECOLI	PF00468 Ribosomal protein L34	P0A7P6
35	50S ribosomal protein L35	RL35_ECOLI	PF01632 Ribosomal protein L35	P0A7Q2
36	50S ribosomal protein L36	RL36_ECOLI	PF00444 Ribosomal protein L36	P0A7Q7

Human small 40S subunit

Subunit No.	Subunit name	Human protein	Pfam family with Human protein
2	40S ribosomal protein S2	RS2_HUMAN	PF03719 Ribosomal protein S5, C-terminal domain
2	40S ribosomal protein S2	RS2_HUMAN	PF00333 Ribosomal protein S5, N-terminal domain
3	40S ribosomal protein S3	RS3_HUMAN	PF00189 Ribosomal protein S3, C-terminal domain
3A	40S ribosomal protein S3a	RS3A_HUMAN	PF01015 Ribosomal S3Ae family
4	40S ribosomal protein S4, X isoform	RS4X_HUMAN	PF08071 RS4NT (NUC023) domain
4	40S ribosomal protein S4, X isoform	RS4X_HUMAN	PF01479 S4 domain
4	40S ribosomal protein S4, X isoform	RS4X_HUMAN	PF00900 Ribosomal family S4e
4	40S ribosomal protein S4, X isoform	RS4X_HUMAN	PF16121 40S ribosomal protein S4 C-terminus
4	40S ribosomal protein S4, Y isoform 1	RS4Y1_HUMAN	PF08071 RS4NT (NUC023) domain
4	40S ribosomal protein S4, Y isoform 1	RS4Y1_HUMAN	PF00900 Ribosomal family S4e
4	40S ribosomal protein S4, Y isoform 1	RS4Y1_HUMAN	PF16121 40S ribosomal protein S4 C-terminus
4	40S ribosomal protein S4, Y isoform 2	RS4Y2_HUMAN	PF08071 RS4NT (NUC023) domain
4	40S ribosomal protein S4, Y isoform 2	RS4Y2_HUMAN	PF00900 Ribosomal family S4e
4	40S ribosomal protein S4, Y isoform 2	RS4Y2_HUMAN	PF16121 40S ribosomal protein S4 C-terminus
5	40S ribosomal protein S5	RS5_HUMAN	PF00177 Ribosomal protein S7p/S5e
6	40S ribosomal protein S6	RS6_HUMAN	PF01092 Ribosomal protein S6e
7	40S ribosomal protein S7	RS7_HUMAN	PF01251 Ribosomal protein S7e
8	40S ribosomal protein S8	RS8_HUMAN	PF01201 Ribosomal protein S8e
9	40S ribosomal protein S9	RS9_HUMAN	PF01479 S4 domain
9	40S ribosomal protein S9	RS9_HUMAN	PF00163 Ribosomal protein S4/S9 N-terminal domain
10	40S ribosomal protein S10	RS10_HUMAN	PF03501 Plectin/S10 domain
11	40S ribosomal protein S11	RS11_HUMAN	PF16205 Ribosomal_S17 N-terminal
11	40S ribosomal protein S11	RS11_HUMAN	PF00366 Ribosomal protein S17
12	40S ribosomal protein S12	RS12_HUMAN	PF01248 Ribosomal protein L7Ae/L30e/S12e/Gadd45 family
13	40S ribosomal protein S13	RS13_HUMAN	PF08069 Ribosomal S13/S15 N-terminal domain
13	40S ribosomal protein S13	RS13_HUMAN	PF00312 Ribosomal protein S15
14	40S ribosomal protein S14	RS14_HUMAN	PF00411 Ribosomal protein S11
15	40S ribosomal protein S15	RS15_HUMAN	PF00203 Ribosomal protein S19
15A	40S ribosomal protein S15a	RS15A_HUMAN	PF00410 Ribosomal protein S8
16	40S ribosomal protein S16	RS16_HUMAN	PF00380 Ribosomal protein S9/S16
17	40S ribosomal protein S17	RS17_HUMAN	PF00833 Ribosomal S17
18	40S ribosomal protein S18	RS18_HUMAN	PF00416 Ribosomal protein S13/S18
19	40S ribosomal protein S19	RS19_HUMAN	PF01090 Ribosomal protein S19e
20	40S ribosomal protein S20	RS20_HUMAN	PF00338 Ribosomal protein S10p/S20e
21	40S ribosomal protein S21	RS21_HUMAN	PF01249 Ribosomal protein S21e
23	40S ribosomal protein S23	RS23_HUMAN	PF00164 Ribosomal protein S12/S23
24	40S ribosomal protein S24	RS24_HUMAN	PF01282 Ribosomal protein S24e
25	40S ribosomal protein S25	RS25_HUMAN	PF03297 S25 ribosomal protein
26	40S ribosomal protein S26	RS26_HUMAN	PF01283 Ribosomal protein S26e
27	40S ribosomal protein S27	RS27_HUMAN	PF01667 Ribosomal protein S27
28	40S ribosomal protein S28	RS28_HUMAN	PF01200 Ribosomal protein S28e
29	40S ribosomal protein S29	RS29_HUMAN	PF00253 Ribosomal protein S14p/S29e
30	40S ribosomal protein S30	RS30_HUMAN	PF04758 Ribosomal protein S30
A	40S ribosomal protein SA	RSSA_HUMAN	PF16122 40S ribosomal protein SA C-terminus
A	40S ribosomal protein SA	RSSA_HUMAN	PF00318 Ribosomal protein S2
	RACK1/GNB2L1	RACK1_HUMAN	PF00400 Receptor for activated C kinase 1

Human large 60S subunit

Subunit No.	Subunit name	Human protein	Pfam family with Human protein
3	60S ribosomal protein L3	RL3_HUMAN	PF00297 Ribosomal protein L3
4	60S ribosomal protein L4	RL4_HUMAN	PF00573 Ribosomal protein L4/L1 family
4	60S ribosomal protein L4	RL4_HUMAN	PF14374 60S ribosomal protein L4
5	60S ribosomal protein L5	RL5_HUMAN	PF00861 Ribosomal L18p/L5e family
5	60S ribosomal protein L5	RL5_HUMAN	PF14204 Ribosomal L18 C-terminal region
6	60S ribosomal protein L6	RL6_HUMAN	PF01159 Ribosomal protein L6e
6	60S ribosomal protein L6	RL6_HUMAN	PF03868 Ribosomal protein L6, N-terminal
7A	60S ribosomal protein L7a	RL7A_HUMAN	PF01248 Ribosomal protein L7Ae/L30e/S12e/Gadd45 family
7	60S ribosomal protein L7	RL7_HUMAN	PF00327 Ribosomal protein L30p/L7e
7	60S ribosomal protein L7	RL7_HUMAN	PF08079 Ribosomal L30 N-terminal domain
8	60S ribosomal protein L8	RL8_HUMAN	PF00181 Ribosomal Proteins L2, RNA-binding domain
8	60S ribosomal protein L8	RL8_HUMAN	PF03947 Ribosomal Proteins L2, C-terminal
9	60S ribosomal protein L9	RL9_HUMAN	PF00347 Ribosomal protein L6
9	60S ribosomal protein L9	RL9_HUMAN	PF00347 Ribosomal protein L6
10A	60S ribosomal protein L10a	RL10A_HUMAN	PF00687 Ribosomal protein L1p/L10e family
10	60S ribosomal protein L10	RL10_HUMAN	PF00252 Ribosomal protein L16p/L10e
11	60S ribosomal protein L11	RL11_HUMAN	PF00281 Ribosomal protein L5
11	60S ribosomal protein L11	RL11_HUMAN	PF00673 ribosomal L5P family C-terminus
12	60S ribosomal protein L12	RL12_HUMAN	PF00298 Ribosomal protein L11, RNA-binding domain
12	60S ribosomal protein L12	RL12_HUMAN	PF03946 Ribosomal protein L11, N-terminal
13A	60S ribosomal protein L13a	RL13A_HUMAN	PF00572 Ribosomal protein L13
13	60S ribosomal protein L13	RL13_HUMAN	PF01294 Ribosomal protein L13e
14	60S ribosomal protein L14	RL14_HUMAN	PF01929 Ribosomal protein L14
15	60S ribosomal protein L15	RL15_HUMAN	PF00827 Ribosomal L15
17	60S ribosomal protein L17	RL17_HUMAN	PF00237 Ribosomal protein L22p/L17e
18A	60S ribosomal protein L18a	RL18A_HUMAN	PF01775 Ribosomal L18ae/LX protein domain
18	60S ribosomal protein L18	RL18_HUMAN	PF00828 Ribosomal_L18e
19	60S ribosomal protein L19	RL19_HUMAN	PF01280 Ribosomal protein L19e
21	60S ribosomal protein L21	RL21_HUMAN	PF01157 Ribosomal protein L21e
22	60S ribosomal protein L22	RL22_HUMAN	PF01776 Ribosomal L22e protein family
23A	60S ribosomal protein L23a	RL23A_HUMAN	PF00276 Ribosomal protein L23
23A	60S ribosomal protein L23a	RL23A_HUMAN	PF03939 Ribosomal protein L23, N-terminal
23	60S ribosomal protein L23	RL23_HUMAN	PF00238 Ribosomal protein L14p/L23e
24	60S ribosomal protein L24	RL24_HUMAN	PF01246 Ribosomal protein L24e
26	60S ribosomal protein L26	RL26_HUMAN	PF16906 Ribosomal proteins L26 eukaryotic, L24P archaeal
27A	60S ribosomal protein L27a	RL27A_HUMAN	PF00828 Ribosomal protein L18e/L15
27	60S ribosomal protein L27	RL27_HUMAN	PF01777 Ribosomal L27e protein family
28	60S ribosomal protein L28	RL28_HUMAN	PF01778 Ribosomal L28e protein family
29	60S ribosomal protein L29	RL29_HUMAN	PF01779 Ribosomal L29e protein family
30	60S ribosomal protein L30	RL30_HUMAN	PF01248 Ribosomal protein L7Ae/L30e/S12e/Gadd45 family
31	60S ribosomal protein L31	RL31_HUMAN	PF01198 Ribosomal protein L31e
32	60S ribosomal protein L32	RL32_HUMAN	PF01655 Ribosomal protein L32
34	60S ribosomal protein L34	RL34_HUMAN	PF01199 Ribosomal protein L34e
35A	60S ribosomal protein L35a	RL35A_HUMAN	PF01247 Ribosomal protein L35Ae
35	60S ribosomal protein L35	RL35_HUMAN	PF00831 Ribosomal L29 protein
36A	60S ribosomal protein L36a	RL36A_HUMAN	PF00935 Ribosomal protein L44
36	60S ribosomal protein L36	RL36_HUMAN	PF01158 Ribosomal protein L36e
37A	60S ribosomal protein L37a	RL37A_HUMAN	PF01780 Ribosomal L37ae protein family
37	60S ribosomal protein L37	RL37_HUMAN	PF01907 Ribosomal protein L37e
38	60S ribosomal protein L38	RL38_HUMAN	PF01781 Ribosomal L38e protein family
39	60S ribosomal protein L39	RL39_HUMAN	PF00832 Ribosomal L39 proteiin
40	Ubiquitin-60S ribosomal protein L40	RL40_HUMAN	PF01020 Ribosomal L40e family
41	60S ribosomal protein L41	RL41_HUMAN	PF05162 Ribosomal protein L41
P0	60S acidic ribosomal protein P0	RLA0_HUMAN	PF00466 Ribosomal protein L10
P0	60S acidic ribosomal protein P0	RLA0_HUMAN	PF00428 60s Acidic ribosomal protein
P1	60S acidic ribosomal protein P1	RLA1_HUMAN	PF00428 60s Acidic ribosomal protein
P2	60S acidic ribosomal protein P2	RLA2_HUMAN	PF00428 60s Acidic ribosomal protein

See also

• Alpha operon ribosome binding site
• Ribosomal protein L20 leader
• Mitochondrial ribosome, for a list of its protein subunits

References

1. Salini Konikkat: Dynamic Remodeling Events Drive the Removal of the ITS2 Spacer Sequence During Assembly of 60S Ribosomal Subunits in S. cerevisiae. Carnegie Mellon University Dissertations, Feb. 2016.
2. Elmar W. Weiler, Lutz Nover (2008), [, p. 532, at Google Books Allgemeine und molekulare Botanik] Check |url= value (help) (in German), Stuttgart: Georg Thieme Verlag, p. 532, ISBN 978-3-13-152791-2
3. Jesus de la Cruz, Katrin Karbstein, John L. Woolford, Jr. (2015), "Functions of Ribosomal Proteins in Assembly of Eukaryotic Ribosomes In Vivo", Annual review of biochemistry (in German), 84, pp. 93–129, doi:10.1146/annurev-biochem-060614-033917, PMC 4772166, PMID 25706898CS1 maint: multiple names: authors list (link)
4. Rodnina MV, Wintermeyer W (April 2011). "The ribosome as a molecular machine: the mechanism of tRNA-mRNA movement in translocation". Biochemical Society Transactions. 39 (2): 658–62. doi:10.1042/BST0390658. PMID 21428957.
5. Ban N, Beckmann R, Cate JH, Dinman JD, Dragon F, Ellis SR, Lafontaine DL, Lindahl L, Liljas A, Lipton JM, McAlear MA, Moore PB, Noller HF, Ortega J, Panse VG, Ramakrishnan V, Spahn CM, Steitz TA, Tchorzewski M, Tollervey D, Warren AJ, Williamson JR, Wilson D, Yonath A, Yusupov M (February 2014). "A new system for naming ribosomal proteins". Current Opinion in Structural Biology. 24: 165–9. doi:10.1016/j.sbi.2014.01.002. PMC 4358319. PMID 24524803.
6. Hug, Laura A.; Baker, Brett J.; Anantharaman, Karthik; Brown, Christopher T.; Probst, Alexander J.; Castelle, Cindy J.; Butterfield, Cristina N.; Hernsdorf, Alex W.; Amano, Yuki; Ise, Kotaro; Suzuki, Yohey; Dudek, Natasha; Relman, David A.; Finstad, Kari M.; Amundson, Ronald; Thomas, Brian C.; Banfield, Jillian F. (11 April 2016). "A new view of the tree of life". Nature Microbiology. 1 (5): 16048. doi:10.1038/nmicrobiol.2016.48. PMID 27572647.
7. Gao, F; Luo, H; Zhang, C. T.; Zhang, R (2015). "Gene Essentiality Analysis Based on DEG 10, an Updated Database of Essential Genes". Gene Essentiality. Methods in Molecular Biology. 1279. pp. 219–33. doi:10.1007/978-1-4939-2398-4_14. ISBN 978-1-4939-2397-7. PMID 25636622.
8. Shoji S, Dambacher CM, Shajani Z, Williamson JR, Schultz PG (2011). "Systematic chromosomal deletion of bacterial ribosomal protein genes". J. Mol. Biol. 413 (4): 751–61. doi:10.1016/j.jmb.2011.09.004. PMC 3694390. PMID 21945294.
9. Akanuma G, Nanamiya H, Natori Y, Yano K, Suzuki S, Omata S, Ishizuka M, Sekine Y, Kawamura F (2012). "Inactivation of ribosomal protein genes in Bacillus subtilis reveals importance of each ribosomal protein for cell proliferation and cell differentiation". J. Bacteriol. 194 (22): 6282–91. doi:10.1128/JB.01544-12. PMC 3486396. PMID 23002217.
10. Arnold RJ, Reilly JP (April 1999). "Observation of Escherichia coli ribosomal proteins and their posttranslational modifications by mass spectrometry". Analytical Biochemistry. 269 (1): 105–12. doi:10.1006/abio.1998.3077. PMID 10094780.
11. Chen SS, Williamson JR (February 2013). "Characterization of the ribosome biogenesis landscape in E. coli using quantitative mass spectrometry". Journal of Molecular Biology. 425 (4): 767–79. doi:10.1016/j.jmb.2012.11.040. PMC 3568210. PMID 23228329.
12. Hamacher K, Trylska J, McCammon JA (February 2006). "Dependency map of proteins in the small ribosomal subunit". PLoS Computational Biology. 2 (2): e10. Bibcode:2006PLSCB...2...10H. doi:10.1371/journal.pcbi.0020010. PMC 1364506. PMID 16485038.

Further reading

• Korobeinikova AV, Garber MB, Gongadze GM (June 2012). "Ribosomal proteins: structure, function, and evolution". Biochemistry (Moscow). 77 (6): 562–74. doi:10.1134/S0006297912060028. PMID 22817455.
• Armache, JP; Anger, AM; Márquez, V; Franckenberg, S; Fröhlich, T; Villa, E; Berninghausen, O; Thomm, M; Arnold, GJ; Beckmann, R; Wilson, DN (January 2013). "Promiscuous behaviour of archaeal ribosomal proteins: implications for eukaryotic ribosome evolution". Nucleic Acids Research. 41 (2): 1284–93. doi:10.1093/nar/gks1259. PMC 3553981. PMID 23222135.
• Ban N, Beckmann R, Cate JH, Dinman JD, Dragon F, Ellis SR, Lafontaine DL, Lindahl L, Liljas A, Lipton JM, McAlear MA, Moore PB, Noller HF, Ortega J, Panse VG, Ramakrishnan V, Spahn CM, Steitz TA, Tchorzewski M, Tollervey D, Warren AJ, Williamson JR, Wilson D, Yonath A, Yusupov M (February 2014). "A new system for naming ribosomal proteins". Current Opinion in Structural Biology. 24: 165–9. doi:10.1016/j.sbi.2014.01.002. PMC 4358319. PMID 24524803.

External links

• 30S Ribosomal proteins at biochem.umd.edu
• Ribosomal+Protein at the US National Library of Medicine Medical Subject Headings (MeSH)