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KEGG pathway analysis of these interactions can be found here
Genes from the HSV-1 genome are listed below with links to the ENTREZ gene database. Their interactions with host genes or proteins are also described. Host genes associated with Alzheimer's disease are in bold type.
This supplementary table relates to the publication:- "Interactions between the products of the Herpes simplex genome and Alzheimer's disease susceptibility genes: Relevance to pathological signalling cascades; C.J.Carter, " Neurochem.Int 2008
Other publications:
The Herpes
simplex (HSV-1) life cycle. Enhanced links to genes are provided by
An excellent review of the Herpes simplex life cycle is available at Dr Wagner’s website at http://darwin.bio.uci.edu/~faculty/wagner/index.html . This brief summary is derived from this site. The virus is composed of three concentric layers comprising a lipid envelope containing glycoproteins responsible for cell attachment and fusion, the tegument and the viral capsid containing the viral DNA. Following entry, the capsid, and certain tegument proteins travel along the microtubule network and dock at the nuclear pore, liberating the viral genome. The virus uses its own and the host’s genomic and transcription apparatus to synthesise the proteins necessary for subsequent viral copying and egress. These proteins are translated within the host cytoplasm, necessitating RNA export and the nuclear reimport of virally encoded proteins. These are assembled to a capsid within the nucleus, which is exported by fusion, with the aid of viral consituents inserted into the nuclear membrane. The outer glycoprotein layer is synthesised and processed by glycosylation in the rough endoplasmic reticulum and Golgi apparatus and inserted into the nuclear or cell membrane. The capsid, associated with tegument proteins is re-enveloped by budding into exocytotic vesicles. These fuse with the virally altered membrane allowing the escape of a mature infectious virus.
| Herpes simplex Virus 1 Genes |
Protein |
Host interactions |
| UL27 type 1 envelope glycoprotein B GeneID : 2703455 |
NP_044629 |
Binds to heparan sulphate proteoglycans and other unknown cellular effectors 13. Cellular entry is cholesterol dependent and blocked by the HMGCR inhibitor nystatin 14. Possesses an antigenic region with homology to APOE 11. Binds to host transcription factor YY1 88. Host defensins inhibit viral infection by binding to glycoprotein B 55 . Binds to perk (EIF2AK3) and prevents its activation by endoplasmic reticulum stress 93. |
US5 type 1 envelope glycoprotein D GeneID: 2703406 |
NP_044668 |
Ligand for the nectin receptor PVRL1 118 Accquires Mannose-6-phosphate residues and binds to Mannose 6 phosphate receptors 24 and to heparan sulfates 106. Binds to the host transcription factor yin-yang-1 (YY1) 48,78,88. Protects against FAS-mediated apoptosis via an NF-kappa-b mediated process 87. US3, US5 and US12 inhibit the cytotoxic effects of T-lymphocytes 7 |
UL44 type 1 envelope glycoprotein CGeneID : 2703410 |
NP_044646 |
Binds to coagulation factor X (F10) 80, which in activated form converts prothrombin to thrombin. Binds to heparin, heparan sulphate and complement 3B110. Factor X also degrades APP 50. |
| US8 type 1 envelope glycoprotein E GeneID: 2703448 |
NP_044670 |
Part of a HSV Fc epsilon receptor mimic 60 (cf FCER1G). Important for the localisation of virus in the trans-golgi network and subsequent delivery to cell junctions to promote intercellular spread 35,43,86. Phosphorylated by casein kinase II 90. |
| US8A protein US8A GeneID: 2703450 |
NP_044671 |
None found |
| US4type 1 envelope glycoprotein G GeneID: 2703404 |
NP_044666 |
None found |
UL22 type 1 envelope glycoprotein H GeneID : 2703373 |
NP_044623 |
Binds to alphavbeta3 integrins 981. Glycoproteins D, H, L and B interact with membrane lipids and allow entry into cells via membrane fusion 119. |
| US7 type 1 envelope glycoprotein I Other Aliases: HHV1gp098 GeneID: 2703446 |
NP_044669 |
Part of a HSV Fc epsilon reptor mimic 60. Antibodies neutralise infectivity in the presence of Complement 120. |
| UL53type 3 envelope glycoprotein K GeneID : 2703425 |
NP_044656 |
Regulates membrane fusion and promotes egress of viral particles 57,105 |
UL1 envelope glycoprotein LGeneID : 2703393 |
NP_044602 |
Glycoproteins B,D.H and L can promote cell fusion 23 |
| UL10 type 3 envelope glycoprotein M GeneID : 2703379 |
NP_044611 |
Induces clathrin mediated endocytosis of other viral proteins and host nectin receptors 31 |
| UL49A type 1 envelope glycoprotein N GeneID : 2703419 |
NP_044652 |
None found |
| UL20 type 3 envelope protein UL20 GeneID : 2703371 |
NP_044621 |
Involved in viral egress allowing escape of the virus to the extracellular space10. |
| UL43type 3 envelope protein UL43 GeneID : 2703408 |
NP_044645 |
None found |
| UL45type 2 membrane protein UL45 GeneID : 2703412 |
NP_044647 |
None found |
| UL56type 2 membrane protein UL56 GeneID : 2703428 |
NP_044659 |
Interacts with kinesin motor protein KIF1A71(HSV2) |
| US9 type 2 membrane protein US9 GeneID: 2703452 |
NP_044672 |
Interacts with the ubiquitin dependent proteasomal pathway of protein degradation 21 |
|
capsid triplex subunit 2 GeneID : 2703366 |
None found |
|
| UL19 major capsid protein GeneID : 2703368 |
NP_044620 |
Binds to host transcription factor YY1 88 |
|
capsid maturation protease GeneID : 2703453 |
NP_044627 |
None found |
|
capsid scaffold protein GeneID : 2703454 |
NP_044628 |
None found |
UL35small capsid protein VP26GeneID : 2703356 |
NP_044637 |
Binds to the dynein components MAPRE3, DYNLT1 and DYNLL1 37 |
|
capsid triplex subunit 1 GeneID : 2703359 |
NP_044640 |
UL38 blocks apoptosis by blocking caspase 3 and poly (ADP-ribose) polymerase cleavage (CMV) 124. Interacts with human hTAF(II)70101. Promoter region binds to DNA-dependent protein kinase 102. IL6 binds to IL6 response elements in the LAT (ICP27) and ICPO viral genes72. |
|
capsid portal protein GeneID : 2703429 |
NP_044607 |
None found |
| UL11 myristylated tegument protein GeneID : 2703380 |
NP_044612 |
Associates with nuclear and cytoplasmic membranease and the golgi apparatus and may play a role in egress from cells 8,9,20,82,83 |
UL13 tegument serine/threonine protein kinaseGeneID : 2703383 |
NP_044614 |
CDC2 mimic that phosphorylates casein kinase II beta and eukaryotic elongation factor 1delta 64.Casein kinase II phosphorylates APP, APOE and PSEN2. 46,104,127.Activates the host cyclin dependendent kinase CDC2 2-4 which in turn phosphorylates herpes simplex proteins including ICP0 and potentially 26 other HSV proteins that contain cdc2 concensusphosphorylation sites 2. CDC2 phosphorylates beta-amyloid (APP) 89 and tau (MAPT) and also phosphorylates and stabilises the sterol regulatory element-binding protein (SREBF1) 17. |
| UL14 tegument protein UL14 GeneID : 2703384 |
NP_044615 |
Antiapoptotic properties related to its heatshock protein like properties 131,132 |
| UL16tegument protein UL16 GeneID : 2703386 |
NP_044617 |
None found |
| UL17DNA packaging tegument protein UL17 GeneID : 2703388 |
NP_044618 |
None found |
| UL21tegument protein UL21 GeneID : 2703372 |
NP_044622 |
Has a certain homology to tau (MAPT), and like tau promotes the formation of longprocesses when transfected into non-neuronal cells. Both tau and UL21 promote the assembly of micotubules 121 |
| UL25DNA packaging tegument protein UL25 GeneID : 2703377 |
NP_044626 |
None found |
| UL36large tegument protein GeneID : 2703357 |
NP_044638 |
Possesses deubiquitinating activity (ubiquitin specific protease)63,114. |
| UL37tegument protein UL37 GeneID : 2703358 |
NP_044639 |
Binds to host transcription factor YY1 88 |
| UL41 tegument host shutoff protein GeneID : 2703365 |
NP_044643 |
Degrades host mRNA including GAPDH74, Downregulates mRNA for the TNF receptor TNFR1 76 |
UL46tegument protein VP11/12 GeneID: 2703413 |
NP_044648 |
None found |
| UL47tegument protein VP13/14 GeneID : 2703415 |
NP_044649 |
RNA-binding protein capable of shuttling between nucleus and cytoplasm 36 |
UL48 transactivating tegument protein VP16GeneID : 2703416 |
NP_044650 |
The homeo domain of the Oct-1 transcription factor (POU2F1)directs formation of a multiprotein-DNA complex containing Oct-1, the herpes simplex virus (HSV) trans-activator VP16, and a second host cell factor (HCF).117. Binds to host transcription factor YY1 78 The herpes simplex proteins VP16 and ICP4 both bind to host TATA binding proteins and the general transcription factor GTF2B, one of the ubiquitous transcription factors required for transcription initiation by RNA polymerase II 79,116.VP16 binds to arc92 (MED25) a mediator of RNA polymerase II transcription 133. |
| UL49tegument protein VP22 GeneID : 2703417 |
NP_044651 |
Interacts with the actin-associated motor protein nonmuscle myosin IIA (NMIIA)125. Interacts with Kinesin heavy chain uKHC 34. Phosphorylated by casein kinase II 40. Results in the hyperacetylation of microtubules 39. This property is shared by unphosphorylated tau (MAPT) 28 and both tau and VP22 are able to stabilise microtubules via this mechanism. |
| UL51 tegument protein UL51 GeneID : 2703422 |
NP_044654 |
Involved in the nuclear egress of viral particles 95 |
| UL7tegument protein UL7 GeneID : 2703431 |
NP_044608 |
None found |
| US11tegument protein US11 GeneID: 2703439 |
NP_044674 |
Interacts with Kinesin heavy chain (KIF5B) 34. Binds to pat1 (APPBP2) a binding partner of the cytoplasmic domain of APP 12. US11 also binds to and is phosphorylated by HIPK2 and US11 prevents the growth-arrest promoted by HIPK2 in Hela cells 47. HIPK2 binds to p53 and p73 (TP73) and is involved in the apoptotic pathways of these tumor proteins 68. Inhibits 2'-5'-oligoadenylate synthetase 112. Several studies have shown that the activity of this enzyme is markedly reduced in fibroblasts isolated from Alzheimer’s disease patients 6,66,100. Prevents PKR activation by inhibiting the PKR activator PACT (PRKRA) 92. |
| UL24nuclear protein UL24 GeneID : 2703375 |
NP_044625 |
None found |
| UL3nuclear protein UL3 GeneID : 2703461 |
NP_044604 |
None found |
| UL4nuclear protein UL4 GeneID : 2703362 |
NP_044605 |
Promoter contains a cyclic AMP concensus sequence that binds to CREB 61 |
| UL55nuclear protein UL55 GeneID : 2703427 |
NP_044658 |
Phosphorylated by casein kinase II (CMV) 58. |
| UL31nuclear egress lamina protein GeneID : 2703350 |
NP_044633 |
The HSV-1, U(L)31 and U(L)34 proteins are dependent on each other for proper targeting to the nuclear membrane and are required for efficient envelopment of nucleocapsids at the inner nuclear membrane. Bind to Lamin A/C (LMNA) 108 |
| UL34nuclear egress type 2 membrane protein GeneID : 2703355 |
NP_044636 |
Disrupts the nuclear lamina during infection Expression of either UL34 or US3 proteins alone disrupted lamin A/C and lamin B localization. Bind to Lamin A/C (LMNA) 108 109,115Interacts with a cytoplasmic dynein intermediate chain Dync1i1134 |
Miscellaneous viral genes with major effects on host physiology
| Herpes simplex Virus 1 Genes |
Protein |
Host interactions |
|
Record to support submission of GeneRIFs for a gene not in Entrez Gene GeneID: 2828259 |
Includes ICP8, ICP27 |
ICP8 binds to proteins involved in DNA repair, replication and recombination and to LMNA and PARP1 (mouse two-hybrid) 123. Viral ICP8, ICP22, ICP27 and ICP28 associate with host RNA polymerase II 32,75,138 ICP4 and ICP27 are anvolved in the suppression of TNF synthesis invoked by HSV to combat host inflammatory reponses 91. ICP27 stabilises the mRNA of IEX-1 (IER3) 30a protein that blocks both FAS and TNF mediated apoptosis 129. ICP27 also contains an element coding for a microrna that downregulates TGFB1 and SMAD3 expression 49. Phosphorylated by host CK2 and protein kinase A 137. ICP27 binds to PABG, EIF3 and EIF4G 44 and to HNRPKand CSNK2B 126. ICP27 activates JNK and P38 kinases 53. ICP27 Binds to RNA polymerase II32 ICP27 binds to and stimulates human casein kinase II 70 Casein kinase II phosphorylates APP, APOE and PSEN2. 46,104,127. HSV-1 infection results in an early activation of the transcription factor nuclear factor kappa beta (NFKB1) an effect mediated via the viral proteins ICP4 and ICP27 54. Herpes simplex infection has been shown to increase the phosphorylation of the host transcription factor SP1, an effect involving ICP4 and ICP27, that reduce its transcriptional efficacy67. |
| UL12 deoxyribonuclease GeneID : 2703382 |
NP_044613 |
A truncated form of UL12 (UL12.5) localises to mitochondria and deletes host mitochondrial DNA 111. |
US1regulatory protein ICP22 GeneID: 2703435 |
NP_044663 |
Activates the host cyclin dependendent kinase CDC2 2-4 which in turn phosphorylates herpes simplex proteins including ICP0 and potentially 26 other HSV proteins that contain CDC2 concensus phosphorylation sites 2. CDC2 phosphorylates beta-amyloid (APP) 89 and tau (MAPT) and also phosphorylates and stabilises the sterol regulatory element-binding protein (SREBF1) 17. UL13 and ICP22 mediate the stabilization of CDC2 and the replacement of its cellular partner, cyclin B, with the viral DNA polymerase processivity factor U(L)42.Binds to GLTSCR2 25 and MCRS126. Viral ICP8, ICP22, ICP27 and ICP28 associate with host RNA polymerase II 32,75,138. |
| US3 serine/threonine protein kinase US3 GeneID: 2703401 |
NP_044665 |
US3, US5 and US12 inhibit the cytotoxic effects of T-lymphocytes 7. Involved in prevention of FAS-mediated apoptosis 59. Inhibits mitochondrial electron transport.33 Prevents caspase dependent cleavage and activation of BAD 15. Activates potein kinase A and has similar kinase properties 16. Reverses cleavage of BID by granzyme B 27.Phosphorylates cytokeratin 17 94. Blocks histone deacetylation (HDAC1) 103 Phosphorylates UL34, ICP22, and Us9 and the cellular protein Bad 62. Herpes viral infection of arterial smooth muscle cells leads to the accumulation of cholesterol 52, partly due to a decrease in cholesteryl ester hydrolysis mediated by decreases in the activities of acid (lysosomal) cholesteryl ester hydrolase (LIPA) and neutral (cytoplasmic) cholesteryl ester hydrolase. Infection of these cells also increased the internalisation of low-density lipoprotein and increases the mRNA expression of the lipoprotein receptor LDLR. The activity and expression of the cholesterol synthetic enzymeHMGCR were also increased by HSV infection. In uninfected cells, mevalonic acid and 25-hydroxycholesterol (the product of cholesterol-25 hydroxylase: CH25H) reduced the activity of HMGCR but this effect was lost in the HSV-infected cells suggesting that the virus may interfere with the repressive effects of oxysterols on cholesterol synthesis. These effects were mediated by the viral kinase vpk1 (US3) 56. |
| US12 virion protein US10 Other Designations: TAP transporter inhibitor ICP47 GeneID: 2703441 |
NP_044675 |
Inhibits TAP2 5,45. US3, US5 and US12 inhibit the cytotoxic effects of T-lymphocytes 7 |
| RS1 transcriptional regulator ICP4 GeneID: 2703392 and 2703391 |
NP_044662 and NP_044676 |
CDC2 substrate 1. PKA substrate 130. Herpes simplex infection has been shown to increase the phosphorylation
of the host transcription factor SP1, an effect involving ICP4 and ICP27,
that reduces its transcriptional efficacy67. HSV-1 infection results
in an early activation of the transcription factor nuclear factor kappa
beta (NFKB1) an effect mediated via the viral proteins ICP4 and ICP27
54. Involved in the suppression of TNF synthesis
invoked by HSV to combat host inflammatory reponses (production of
IFN- |
UL9 DNA replication origin-binding helicaseGeneID : 2703434 |
NP_044610 |
Binds to NFB42 (FBXO2) 42 a component of the SCF (SKP1-cullin-F-box) ubiquitin ligasecomplex. FBXO2 is involved in the nuclear export of viral UL9 41 and in its degradation 42.FBXO2 recognises sugar chains (N-glycans), and ubiquitinates N-glycosylated proteins,including the T-cell receptor and the alpha-2-HS-glycoprotein (AHSG) 135It is involved in the endoplasmic reticulum associated degradation pathway (ERAD) 136. |
RL2 ubiquitin E3 ligase ICP0GeneID : 2703390 |
NP_044660 |
Binds to and ubiquitinates TP53 19. Binds to YY148. Blocks histone deacetylation (HDAC1) 103. Binds to GLTSCR2 25 The ICP0 protein of HSV1 interacts with certain histone deacetylases (HDAC4,5 and 9) via an amino terminal domain responsible for suppressing the activity of myocyte enhance factor MEF2A by nuclear sequestration. ICP0 overcomes the HDAC mediated suppression of MEF2A 81. Implicated as an important trigger of the apoptotic response involving PARP1 and caspase-3 cleavage but the pathway involved remains to be defined 113. IL6 binds to IL6 response elements in the LAT (ICP27) and ICPO viral genes72. ICP0 contains two E3 ubiquitin ligase domains, one of which binds to the host ubiquitin conjugating enzyme CDC34 and another that binds to host UbcH5a (UBE2D1)- and UbcH6 (UBE2E1) E2-conjugating enzymes51. Both these E2 conjugating enzymes are known to complex with the ubiquitin ligase e6ap (UBE3A) 96,97,which in turn binds to both ubiquilins (UBQLN1, 2) 69. UBQLN1 binds to both presenilins (PSEN1, PSEN2) and also regulates PEN2 and NCSTN protein levels, thus controlling the degradation of the gamma-secretase complex 84,85. ICP0 also interacts with a cellular ubiquitin-specific protease USP7 (also known as HAUSP) 18,51 a deubiquitylating enzyme that stabilises p53 (TP53) and MDM2. Binds to the adaptor protein CIN85 and forms a tripartite complex including CIN85 and the cbl ubiquitin ligase (CBL) 77. CIN 85 binds to numerous components of the tyrosine kinase signalling cascade including GRB2, SOS1 and PIK3R1128 while CBL ubiquitinates tyrosine kinase receptors, (including the lymphocyte tyrosine kinase LCK) 107targeting them for destruction. Binds to and stabilises cyclin D3 65. |
RL1 neurovirulence protein ICP34.5 GeneID : 2703396GeneID : 2703395 |
NP_044661 NP_044600 |
PPP1R15A (gadd34) mimic that dephosphorylates eif2alpha, the stress kinase phosphorylated by PKR 99,122 Complexes with proliferating cell nuclear antigen (PCNA) 22 |
US12virion protein US10 GeneID: 2703441 |
NP_044673 |
US3, US5 and US12 inhibit the cytotoxic effects of T-lymphocytes 7 |
| UL54 multifunctional expression regulator GeneID : 2703426 |
NP_044657 |
Interacts with HNRPK and casein kinase II 126 |
| UL2 uracil-DNA glycosylase GeneID: 2703370 |
NP_044603 |
None found |
| UL23 thymidine kinase GeneID : 2703374 |
NP_044624 |
Contains concensus sites for host transcription factors Spl, CTF, and TFIID73 |
| UL39 ribonucleotide reductase subunit 1 Other Aliases : HHV1gp057 GeneID : 2703361 |
NP_044641 |
Phosphorylated by casein kinase II 29 |
| UL40ribonucleotide reductase subunit 2 GeneID : 2703364 |
NP_044642 |
None found |
| UL50 : deoxyuridine triphosphatase GeneID : 2703421 |
NP_044653 |
Catalyzes the hydrolysis of dUTP to dUMP and pyrophosphate. |
| UL52 helicase-primase primase subunit GeneID : 2703423 |
NP_044655 |
None found |
| UL15DNA packaging terminase subunit 1 GeneID : 2703385 |
NP_044616 |
None found |
| UL28DNA packaging terminase subunit 2 GeneID : 2703457 |
NP_044630 |
None found |
| UL29 single-stranded DNA-binding protein GeneID : 2703458 |
NP_044631 |
None found |
| UL30DNA polymerase catalytic subunit GeneID : 2703462 |
NP_044632 |
None found |
| UL32DNA packaging protein UL32 GeneID : 2703352 |
NP_044634 |
cleavage and packaging of herpesvirus DNA into preassembled capsids |
| UL33DNA packaging protein UL33 GeneID : 2703353 |
NP_044635 |
None found |
| UL42DNA polymerase processivity subunit GeneID : 2703407 |
NP_044644 |
None found |
| UL5helicase-primase helicase subunit GeneID : 2703420 |
NP_044606 |
None found |
| UL8helicase-primase subunit GeneID : 2703432 |
NP_044609 |
None found |
| US2 virion protein US2 GeneID: 2703399 |
NP_044664 |
MHC class I binding protein. |
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TNFA, IL6, IL-12, and RANTES).