| |
α
|
β
|
γ
| |
α
|
β
|
γ
|
|---|
| |
HPV16
|
HPV18
|
HPV6
|
HPV8
|
HPV4
| |
HPV16
|
HPV18
|
HPV6
|
HPV8
|
HPV4
|
|---|
|
AbaA
|
+
|
+
| |
+
|
+
|
HFH-8
|
+
|
+
|
+
|
+
|
+
|
|
AhR/Arnt
|
+
| |
+
| | |
HLF
|
+
| | | | |
|
AP-1
|
+
|
+
|
+
|
+
|
+
|
HNF-3beta
|
+
| |
+
|
+
|
+
|
|
AREB6
|
+
|
+
|
+
|
+
|
+
|
lk-1
| | | | |
+
|
|
Arnt
|
+
| |
+
| |
+
|
Lmo2 complex
| |
+
|
+
|
+
|
+
|
|
Athb-1
|
+
| | | |
+
|
Mat1-Mc
|
+
|
+
|
+
|
+
| |
|
Bcd
| |
+
|
+
|
+
|
+
|
MATa1
| | | |
+
| |
|
Brc- Z1
| | | |
+
| |
Max
|
+
| | | | |
|
BR-c Z4
|
+
|
+
|
+
|
+
| |
MCM1
|
+
| | | | |
|
Brn-2
|
+
| |
+
|
+
|
+
|
MYB.Ph3
|
+
| | | |
+
|
|
CCAAT box
| | |
+
| | |
NF-E2
| | | |
+
| |
|
C/EBP
| |
+
| |
+
| |
NF-Y
| |
+
|
+
|
+
|
+
|
|
CDP CR3+HD
| |
+
| |
+
| |
Nkx2-5
|
+
| |
+
| | |
|
c-Ets-1(p54)
|
+
|
+
|
+
| |
+
|
N-Myc
|
+
| | | | |
|
CF2-II
|
+
|
+
|
+
|
+
|
+
|
NRF-2
| | | | |
+
|
|
CHOP-C
| |
+
|
+
|
+
| |
oct-1
|
+
|
+
|
+
|
+
|
+
|
|
c-Myb
|
+
|
+
|
+
|
+
|
+
|
OCT-x
| | | | |
+
|
|
c-Myc/Max
|
+
| | | | |
PacC
| | |
+
| |
+
|
|
Croc
|
+
| | |
+
| |
PHO4
|
+
| |
+
| |
+
|
|
dl
| |
+
| | |
+
|
RAP1
| |
+
| | | |
|
E2
|
+
|
+
|
+
|
+
|
+
|
RFX1
| |
+
| | | |
|
E2F
| |
+
| | | |
RORalpha1
|
+
| |
+
| | |
|
E47
| |
+
|
+
|
+
|
+
|
S8
|
+
|
+
|
+
| |
+
|
|
E74A
| | | | |
+
|
SBF-1
| |
+
|
+
|
+
|
+
|
|
Elf-1
|
+
|
+
|
+
|
+
|
+
|
Skn-1
|
+
|
+
|
+
|
+
|
+
|
|
Elk-1
|
+
| | | |
+
|
Sn
| |
+
|
+
|
+
| |
|
ER
|
+
| | | |
+
|
Sox-5
|
+
|
+
|
+
|
+
|
+
|
|
FOXD3
|
+
| |
+
|
+
|
+
|
SOX-9
|
+
|
+
|
+
|
+
|
+
|
|
FOXJ2
|
+
|
+
|
+
|
+
| |
STATx
| | | |
+
|
+
|
|
Freac-2
|
+
| |
+
|
+
| |
StuAp
| |
+
|
+
| | |
|
Freac-7
|
+
|
+
|
+
|
+
|
+
|
Su(H)
| |
+
| | | |
|
GATA-1
| |
+
|
+
|
+
|
+
|
TATA
|
+
|
+
|
+
| |
+
|
|
GATA-2
| |
+
|
+
| |
+
|
TCF11
| | | |
+
| |
|
GATA-3
| |
+
| | | |
TGIF
|
+
| | | |
+
|
|
GATA-x
| |
+
|
+
|
+
|
+
|
USF
|
+
| |
+
| | |
|
GBP
|
+
| |
+
| |
+
|
VBP
| | |
+
| | |
|
GCN4
|
+
| |
+
|
+
|
+
|
v-ErbA
| | |
+
| | |
|
Gfi-1
|
+
| | | | |
v-Maf
|
+
| | | | |
|
Hand1/E47
| | | |
+
|
+
|
v-Myb
|
+
|
+
|
+
|
+
|
+
|
|
Hairy
| | |
+
| | |
XFD-1
|
+
| | |
+
| |
|
HAP2/3/4
| | | | |
+
|
XFD-2
|
+
| |
+
|
+
| |
|
HFH-1
|
+
| |
+
|
+
| |
YY1
|
+
| | | | |
|
HFH-3
|
+
|
+
|
+
|
+
|
+
|
Zeste
| | | |
+
|
+
|
- Transcription factor binding sites were predicted with MATCH, using the nucleotide matricial description of each site as compiled in TRANSFAC. The coincidence levels between the binding site sequence and the sequence in the URR were fixed to optimise simultaneously the number of false positives and false negatives. It is obvious that different papillomaviruses contain different transcription factor binding sites in their URR. Some of them are common to all papillomaviruses, such as AP-1, E2, HFH-3 or Oct-1. Other TFBS are type-exclusive, genus-exclusive, or are shared by papillomaviruses that infect the same host. The high dimensionality of these results makes it necessary to analyse them with information reduction techniques, such as principal component analysis or genetic algorithms.
