T4SS ID	19
Strain	Agrobacterium tumefaciens str. C58
Replicon	plasmid Ti [Browse all T4SS(s) in this replicon]
Accession	NC_003065
Location	183877..201962
Name	VirB
Function	effector translocation and conjugation
Classification	Type IVA; Type P
Experimental investigation has been performed on this T4SS.

Information of structure: VirB8

T4SS components

Component	VirB1	VirB2	VirB3	VirB4	VirB5	VirB6	VirB7	VirB8	VirB9
Number	1	1	1	1	1	1	1	1	1
Component	VirB10	VirB11	VirD4
Number	1	1	1

The information of T4SS components from NC_003065
#	Locus tag (Gene)	Coordinates [+/-], size (bp)	Protein GI	Product	Component
1	Atu6164 (tzs)	179262..179993 [-], 732	41223351	trans-zeatin secretion protein
2	Atu6165	180279..180470 [+], 192	159161997	hypothetical protein
3	Atu6166 (virA)	180831..183332 [+], 2502	16119781	two-component VirA-like sensor kinase
4	Atu6167 (virB1)	183877..184614 [+], 738	41223307	type IV secretion system lytic transglycosylase VirB1	VirB1
5	Atu6168 (virB2)	184614..184979 [+], 366	16119783	type IV secretion system pilin subunit VirB2	VirB2
6	Atu6169 (virB3)	184979..185305 [+], 327	16119784	type IV secretion system protein VirB3	VirB3
7	Atu6170 (virB4)	185305..187674 [+], 2370	16119786	type IV secretion system protein VirB4	VirB4
8	Atu6171 (virB5)	187691..188353 [+], 663	16119787	type IV secretion system protein VirB5	VirB5
9	Atu6172 (virB6)	188454..189341 [+], 888	41223308	type IV secretion system protein VirB6	VirB6
10	Atu6173 (virB7)	189377..189544 [+], 168	159161998	type IV secretion system lipoprotein VirB7	VirB7
11	Atu6174 (virB8)	189531..190244 [+], 714	41223309	type IV secretion system protein VirB8	VirB8
12	Atu6175 (virB9)	190241..191122 [+], 882	16119791	type IV secretion system protein VirB9	VirB9
13	Atu6176 (virB10)	191119..192252 [+], 1134	16119792	type IV secretion system protein VirB10	VirB10
14	Atu6177 (virB11)	192293..193327 [+], 1035	41223310	type IV secretion system protein VirB11	VirB11
15	Atu6178 (virG)	193546..194307 [+], 762	16119794	two-component response regulator VirG
16	Atu6179 (virC2)	194449..195057 [-], 609	159161999	putative crown gall tumor protein VirC2
17	Atu6180 (virC1)	195060..195755 [-], 696	16119796	putative crown gall tumor protein VirC1
18	Atu6181 (virD1)	196026..196469 [+], 444	41223311	type IV secretion system T-DNA border endonuclease VirD1
19	Atu6182 (virD2)	196503..197846 [+], 1344	16119798	type IV secretion system T-DNA border endonuclease VirD2
20	Atu6183 (virD3)	197861..199882 [+], 2022	159162000	virA/G regulated protein
21	Atu6184 (virD4)	199956..201962 [+], 2007	41223312	type IV secretion system protein VirD4	VirD4
22	Atu6185 (virD5)	202074..204596 [+], 2523	16119801	virA/G regulated protein
23	Atu6186 (virE3)	204666..205232 [+], 567	159162001	virA/G regulated protein
24	Atu6188 (virE0)	205684..205935 [+], 252	159162002	virA/G regulated protein
25	Atu6189 (virE1)	206051..206242 [+], 192	16119804	type IV secretion system chaperone VirE1

 This T4SS contains information of accessory protein.

Genes in the 5-Kb flanking regions if available, or non-essential genes in the T4SS gene cluster if any.

Download FASTA format files

Proteins        Genes

This system delivers both oncogenic DNA and proteins into plant cells. The transferred DNA (T-DNA, NC_003065 REGION: 1..24836) is processed by VirD1, VirD2, and VirC1 (Dtr proteins) and covalently bound to VirD2 to form a VirD2-T-strand transfer intermediate. The T-DNA complex is then transferred by this system into target cells. In addition, the T4SS independently translocates four additional effector proteins: VirE2, VirE3, VirF and VirD5.

Effectors

VirD2, VirD5, VirE2, VirE3, VirF

The information of protein effectors
#	Locus tag (Gene)	Coordinates [+/-], size (bp)	Protein GI	Product	*
1	Atu6182 (virD2)	196503..197846 [+], 1344	16119798	type IV secretion system T-DNA border endonuclease VirD2	VirD2
2	Atu6185 (virD5)	202074..204596 [+], 2523	16119801	virA/G regulated protein	VirD5
3	Atu6190 (virE2)	206256..207926 [+], 1671	16119805	type IV secretion system single-stranded DNA binding protein VirE2	VirE2
4	Atu6191 (virE3)	207993..210047 [+], 2055	16119806	virA/G regulated protein	VirE3
5	Atu6154 (virF)	172489..173427 [+], 939	16119971	exported virulence protein	VirF

 T4SE derived from experimental literature.

Download FASTA format files

Proteins        Genes

The information on structure of this T4ss

#	Name	Image	Resource	Detail	Reference
1	VirB8		PDB (2CC3)	Structure of Agrobacterium tumefaciens VirB8 protein	(1) PubMed: 16481621

(1) Bailey S; Ward D; Middleton R; Grossmann JG; Zambryski PC (2006). Agrobacterium tumefaciens VirB8 structure reveals potential protein-protein interaction sites. Proc Natl Acad Sci U S A. 103(8):2582-7. [PudMed:16481621]

The information on requirements for T4SS substrate-channel docking

Certain T4SS substrates require secretion chaperones for translocation. These chaperones often possess physical properties (small size of 15 kDa, acidic pI, and amphipathic helices) resembling those of chaperones associated with the type III secretion systems, a family of macromolecular translocation systems ancestrally related to bacterial flagella.

#	Accessory protein(GI)	motif(s)	Substrate(s)	Function	Reference
1	VirC1 (16119796)	ND	VirD2	VirC1 pairwise interacts with the processing factors VirD2 relaxase, VirC2, and VirD1 to stimulate generation of multiple copies per cell of the T-DNA substrate. VirC1 promotes conjugative T-DNA transfer.	(1) PubMed: 17505518
2	VirE1 chaperone (16119804)	CT50aa sufficient (interacts with the VirD4 T4CP. CT100aa(VirE2) sufficient for the T4CP interation)	VirE2	Translocation of the VirE2 effector through the A. tumefaciens VirB/VirD4 T4SS requires VirE1.	(2) PubMed: 19946141

Tips:
1.Substrate(s): For the conjugation systems, the listed proteins are relaxases that bind a cognate T4CP and are delivered to recipient cells. For the effector translocator systems, the listed proteins are effectors that play a role in the infection processes of the bacterial pathogen.
2.motif(s):The motifs listed are required for substrate translocation. In some cases, the protein or its C-terminal fragment (CT) is sufficient to mediate translocation to target cells, as shown by fusion to a reporter protein such as Cre recombinase or adenylate cyclase. Amino acids (aa) at positions listed relative to the C-terminal fragment (subscript) are required for translocation, as shown by mutational analysis. ND, not determined. Parentheses indicate that the interaction between a protein substrate and a cognate T4CP has been experimentally shown.
3.Accessory protein: Accessory factors required for T4SS channel docking or translocation. The proposed function in mediating substrate-T4SS channel docking is shown in parentheses.PubMed:19946141

(1) Atmakuri K; Cascales E; Burton OT; Banta LM; Christie PJ (2007). Agrobacterium ParA/MinD-like VirC1 spatially coordinates early conjugative DNA transfer reactions. EMBO J. 26(10):2540-51. [PudMed:17505518]

(2) Alvarez-Martinez CE; Christie PJ (2009). Biological diversity of prokaryotic type IV secretion systems. Microbiol Mol Biol Rev. 73(4):775-808. [PudMed:19946141]

(1) Aguilar J; Cameron TA; Zupan J; Zambryski P (2011). Membrane and core periplasmic Agrobacterium tumefaciens virulence Type IV secretion system components localize to multiple sites around the bacterial perimeter during lateral attachment to plant cells. MBio. 2(6):e00218-11. [PudMed:22027007]

(2) Sivanesan D; Baron C (2011). The dimer interface of Agrobacterium tumefaciens VirB8 is important for type IV secretion system function, stability, and association of VirB2 with the core complex. J Bacteriol. 193(9):2097-106. [PudMed:21398549]

(3) Mossey P; Hudacek A; Das A (2010). Agrobacterium tumefaciens type IV secretion protein VirB3 is an inner membrane protein and requires VirB4, VirB7, and VirB8 for stabilization. J Bacteriol. 192(11):2830-8. [PudMed:20348257]

(4) Aguilar J; Zupan J; Cameron TA; Zambryski PC (2010). Agrobacterium type IV secretion system and its substrates form helical arrays around the circumference of virulence-induced cells. Proc Natl Acad Sci U S A. 107(8):3758-63. [PudMed:20133577]

(5) Zupan J; Hackworth CA; Aguilar J; Ward D; Zambryski P (2007). VirB1* promotes T-pilus formation in the vir-Type IV secretion system of Agrobacterium tumefaciens. J Bacteriol. 189(18):6551-63. [PudMed:17631630]

(6) Draper O; Middleton R; Doucleff M; Zambryski PC (2006). Topology of the VirB4 C terminus in the Agrobacterium tumefaciens VirB/D4 type IV secretion system. J Biol Chem. 281(49):37628-35. [PudMed:17038312]

(7) Wessel M; Klusener S; Godeke J; Fritz C; Hacker S; Narberhaus F (2006). Virulence of Agrobacterium tumefaciens requires phosphatidylcholine in the bacterial membrane. Mol Microbiol. 62(3):906-15. [PudMed:17010159]

(8) McCullen CA; Binns AN (2006). Agrobacterium tumefaciens and plant cell interactions and activities required for interkingdom macromolecular transfer. Annu Rev Cell Dev Biol. 22:101-27. [PudMed:16709150]

(9) Bailey S; Ward D; Middleton R; Grossmann JG; Zambryski PC (2006). Agrobacterium tumefaciens VirB8 structure reveals potential protein-protein interaction sites. Proc Natl Acad Sci U S A. 103(8):2582-7. [PudMed:16481621]

(10) Middleton R; Sjolander K; Krishnamurthy N; Foley J; Zambryski P (2005). Predicted hexameric structure of the Agrobacterium VirB4 C terminus suggests VirB4 acts as a docking site during type IV secretion. Proc Natl Acad Sci U S A. 102(5):1685-90. [PudMed:15668378]

(11) Ward DV; Draper O; Zupan JR; Zambryski PC (2002). Peptide linkage mapping of the Agrobacterium tumefaciens vir-encoded type IV secretion system reveals protein subassemblies. Proc Natl Acad Sci U S A. 99(17):11493-500. [PudMed:12177441]

(12) Hapfelmeier S; Domke N; Zambryski PC; Baron C (2000). VirB6 is required for stabilization of VirB5 and VirB3 and formation of VirB7 homodimers in Agrobacterium tumefaciens. J Bacteriol. 182(16):4505-11. [PudMed:10913084]

This literature contains experimental investigation

This literature contains bioinformatics investigation