Simplexvirus

VIRION

Enveloped, spherical to pleomorphic, 150-200 nm in diameter, T=16 icosahedral symmetry. The capsid consists of 162 capsomers and is surrounded by an amorphous tegument.

GENOME

Monopartite, linear, dsDNA genome of 152 kb. The genome contains terminal and internal reiterated sequences.

GENE EXPRESSION

All genes are transcribed by the host RNA polymerase II. and most mRNAs are unspliced. There are three temporal classes of genes: immediate-early (alpha), early (beta) and late (gamma). The immediate-early genes are transcribed immediately after infection to take control of cell defense and to activate early genes. These encode the proteins necessary for the viral DNA replication. The late genes mostly encode structural proteins. Latent genes can stop the replicative process at the early step. Certain proteins are downregulated translationaly by a leaky scanning from an upstream ORF.

REPLICATION

NUCLEAR

Lytic replication:

  1. Attachment of the viral gB, gC, gD and gH proteins to host receptors mediates endocytosis of the virus into the host cell.
  2. Fusion with the plasma membrane to release the core and the tegument proteins into the host cytoplasm.
  3. The capsid is transported to the nuclear pore where viral DNA is released into the nucleus.
  4. Transcription of immediate early genes which promote transcription of early genes and protect the virus against innate host immunity.
  5. Transcription of early viral mRNA by host polymerase II, encoding proteins involved in replication of the viral DNA.
  6. A first round of circular genome amplification occurs by bidirectional replication
  7. Synthesis of linear concatemer copies of viral DNA by rolling circle.
  8. Transcription of late mRNAs by host polymerase II, encoding structural proteins.
  9. Assembly of the virus in nuclear viral factories and budding through the inner lamella of the nuclear membrane which has been modified by the insertion of herpes glycoproteins, throughout the Golgi and final release at the plasma membrane.

Latent replication : replication of circular viral episome in tandem with the host cell DNA using the host cell replication machinery.

Host-virus interaction

Adaptive immune response inhibition

Herpes simplex protein US12 binds specifically to transporters associated with antigen processing (TAP), blocking peptide-binding to TAP and subsequent loading of peptides onto MHC class I molecules . Herpes simplex virus 1 glycoprotein B and US3 collaborate to inhibit CD1d antigen presentation and NKT cell function .

Apoptosis modulation

The viral kinase US3 modulates host apoptosis in different ways. It activates host protein kinase A/PKA to block apoptosis. Additionally, it also interacts with programmed cell death protein 4 (PDCD4) and retains it in the nucleus thereby inhibiting host apoptosis .

Autophagy modulation

Herpes simplex virus ICP34.5 confers neurovirulence by targeting the Beclin 1 autophagy protein and thus blocking host autophagy process .

Cell-cycle modulation

The viral proteins ICP27 and vhs act jointly to repress host gene transcription and blocks the cell cycle at the G1 phase . In addition the UL24 protein induces a cell cycle arrest at G2/M transition through inactivation of the host cyclinB/cdc2 complex .

Innate immune response inhibition

Herpes simplex virus inhibits the cascade leading to production of interferon-beta by targeting different cellular proteins. ICP34.5 forms a complex with host TBK1 and thus disrupts the interaction of TBK1-containing complexes with IRF3 . Another viral protein, ICP0, prevents IRF3 activation and subsequent induction of IFN-beta expression .

Host splicing inhibition

HSV-1 ICP27 is an alternative splicing regulator of host mRNA. It has been shown to act as a splicing silencer at the 3' splice site of the PML intron 7a .

Matching UniProtKB/Swiss-Prot entries

230 entries grouped by strain (browse by keywords)

All proteins are shown (view only 146 complete proteome entries)

73 entries

Human herpesvirus 1 (strain 17) (HHV-1) (Human herpes simplex virus 1) reference strain

Select_allDeselect_all
AN_HHV11 Alkaline nuclease (EC 3.1.-.-)
CEP2_HHV11 Cytoplasmic envelopment protein 2
CEP3_HHV11 Cytoplasmic envelopment protein 3
CEP1_HHV11 Cytoplasmic envelopment protein 1
CVC1_HHV11 Capsid vertex component 1
CVC2_HHV11 Capsid vertex component 2
DNBI_HHV11 Major DNA-binding protein
DUT_HHV11 Deoxyuridine 5'-triphosphate nucleotidohydrolase (dUTPase) (EC 3.6.1.23) (dUTP pyrophosphatase)
DPOL_HHV11 DNA polymerase catalytic subunit (EC 2.7.7.7) (EC 3.1.26.4)
EV45_HHV11 Envelope protein UL45 (18 kDa protein)
GH_HHV11 Envelope glycoprotein H (gH)
GI_HHV11 Envelope glycoprotein I (gI)
GE_HHV11 Envelope glycoprotein E (gE) (gE-1)
GD_HHV11 Envelope glycoprotein D (gD)
GJ_HHV11 Envelope glycoprotein J
GB_HHV11 Envelope glycoprotein B (gB)
GC_HHV11 Envelope glycoprotein C
GG_HHV11 Envelope glycoprotein G (gG) (gG-1)
GK_HHV11 Envelope glycoprotein K (Syncytial protein)
GL_HHV11 Envelope glycoprotein L (gL)
GM_HHV11 Envelope glycoprotein M (gM)
GN_HHV11 Envelope glycoprotein N
HELI_HHV11 DNA replication helicase (EC 3.6.4.-)
HEPA_HHV11 DNA helicase/primase complex-associated protein (HEPA) (Primase-associated factor)
MB43_HHV11 Membrane protein UL43
LTP_HHV11 Large tegument protein deneddylase (EC 3.4.19.12) (EC 3.4.22.-)
OBP_HHV11 Replication origin-binding protein (OBP) (OriBP)
PAP_HHV11 DNA polymerase processivity factor (DNA-binding protein UL42) (Polymerase accessory protein) (PAP)
ICP0_HHV11 E3 ubiquitin-protein ligase ICP0 (EC 2.3.2.27) (Alpha-0 protein) (Immediate-early protein IE110) ...
ICP47_HHV11 ICP47 protein (Immediate-early protein IE12) (Immediate-early-5) (Infected cell protein 47) (US12 ...
ICP4_HHV11 Major viral transcription factor ICP4 (Alpha-4 protein) (Infected cell protein 4) (Transcriptional ...
ICP27_HHV11 mRNA export factor (Immediate-early protein IE63) (Infected cell protein 27) (ICP27) (VMW63)
ICP34_HHV11 Neurovirulence factor ICP34.5 (Infected cell protein 34.5) (protein gamma(1)34.5)
ICP22_HHV11 Transcriptional regulator ICP22 (Immediate-early protein IE68) (Infected cell protein 22) (ICP22)
ITP_HHV11 Inner tegument protein
KITH_HHV11 Thymidine kinase (EC 2.7.1.21)
MCP_HHV11 Major capsid protein (MCP)
NEC1_HHV11 Nuclear egress protein 1
NEC2_HHV11 Nuclear egress protein 2
NP03_HHV11 Nuclear phosphoprotein UL3
NP04_HHV11 Nuclear protein UL4
PORTL_HHV11 Portal protein
PRIM_HHV11 DNA primase (EC 2.7.7.-)
RIR2_HHV11 Ribonucleoside-diphosphate reductase small subunit (EC 1.17.4.1) (Ribonucleotide reductase small ...
RIR1_HHV11 Ribonucleoside-diphosphate reductase large subunit (R1) (EC 1.17.4.1) (Ribonucleotide reductase ...
RNB_HHV11 RNA-binding protein (Vmw21)
SCP_HHV11 Small capsomere-interacting protein
SHUT_HHV11 Virion host shutoff protein (Vhs) (EC 3.1.27.-)
SCAF_HHV11 Capsid scaffolding protein (Capsid protein P40) (Protease precursor) (pPR) (Virion structural ...
TEG3_HHV11 Tegument protein UL14
TEG4_HHV11 Tegument protein UL21
TEG6_HHV11 Tegument protein UL55
TEG5_HHV11 Tegument protein UL47 (82/81 kDa tegument protein) (VMW82/81) (VP13/14)
TEG7_HHV11 Tegument protein UL51
TEG1_HHV11 Tegument protein UL46 (Tegument protein VP11/12)
TRM1_HHV11 Tripartite terminase subunit 1
TRM2_HHV11 Tripartite terminase subunit 2
TRM3_HHV11 Tripartite terminase subunit 3 (EC 3.1.-.-) (Terminase large subunit)
UL32_HHV11 Packaging protein UL32
UL56_HHV11 Protein UL56
UNG_HHV11 Uracil-DNA glycosylase (UDG) (EC 3.2.2.27) (UNG)
UL20_HHV11 Protein UL20
TRX1_HHV11 Triplex capsid protein 1
TRX2_HHV11 Triplex capsid protein 2
US03_HHV11 Serine/threonine-protein kinase US3 (EC 2.7.11.1)
UL24_HHV11 Protein UL24
US8A_HHV11 Protein US8.5
UL13_HHV11 Serine/threonine-protein kinase UL13 (EC 2.7.11.1) (Virion protein VMW57)
VP16_HHV11 Tegument protein VP16 (Alpha trans-inducing protein) (Alpha-TIF) (ICP25) (Vmw65)
US10_HHV11 Virion protein US10
VP22_HHV11 Tegument protein VP22
US02_HHV11 Protein US2
US9_HHV11 Envelope protein US9 (10 kDa protein)

72 entries

Human herpesvirus 2 (strain HG52) (HHV-2) (Human herpes simplex virus 2) reference strain

Select_allDeselect_all
CEP2_HHV2H Cytoplasmic envelopment protein 2
CEP3_HHV2H Cytoplasmic envelopment protein 3
CEP1_HHV2H Cytoplasmic envelopment protein 1
CVC2_HHV2H Capsid vertex component 2
CVC1_HHV2H Capsid vertex component 1
DPOL_HHV2H DNA polymerase catalytic subunit (EC 2.7.7.7) (EC 3.1.26.4)
DNBI_HHV2H Major DNA-binding protein
DUT_HHV2H Deoxyuridine 5'-triphosphate nucleotidohydrolase (dUTPase) (EC 3.6.1.23) (dUTP pyrophosphatase)
EV45_HHV2H Envelope protein UL45 (18 kDa protein)
GM_HHV2H Envelope glycoprotein M (gM)
GN_HHV2H Envelope glycoprotein N
GC_HHV2H Envelope glycoprotein C
GG_HHV2H Envelope glycoprotein G (gG) (gG-2)
GK_HHV2H Envelope glycoprotein K (Syncytial protein)
GB_HHV2H Envelope glycoprotein B (gB)
GE_HHV2H Envelope glycoprotein E (gE) (gE-2)
GD_HHV2H Envelope glycoprotein D (gD)
GH_HHV2H Envelope glycoprotein H (gH)
GI_HHV2H Envelope glycoprotein I (gI)
GJ_HHV2H Envelope glycoprotein J
GL_HHV2H Envelope glycoprotein L (gL)
HELI_HHV2H DNA replication helicase (EC 3.6.4.-)
HEPA_HHV2H DNA helicase/primase complex-associated protein (HEPA) (Primase-associated factor)
LTP_HHV2H Large tegument protein deneddylase (EC 3.4.19.12) (EC 3.4.22.-)
MB43_HHV2H Membrane protein UL43
PAP_HHV2H DNA polymerase processivity factor (DNA-binding protein UL42) (Polymerase accessory protein) (PAP)
ICP27_HHV2H mRNA export factor (Immediate-early protein IE63) (Infected cell protein 27) (ICP27) (VMW63)
ICP0_HHV2H E3 ubiquitin-protein ligase ICP0 (EC 2.3.2.27) (RING-type E3 ubiquitin transferaseICP0) (VMW118 ...
ICP22_HHV2H Transcriptional regulator ICP22 (Immediate-early protein IE68) (Infected cell protein 22) (ICP22)
ICP34_HHV2H Neurovirulence factor ICP34.5 (Infected cell protein 34.5) (protein gamma(1)34.5)
ICP47_HHV2H ICP47 protein (Immediate-early protein IE12) (Immediate-early-5) (Infected cell protein 47) (US12 ...
ICP4_HHV2H Major viral transcription factor ICP4 homolog (Alpha-4 protein) (Infected cell protein 4) ...
ITP_HHV2H Inner tegument protein
KITH_HHV2H Thymidine kinase (EC 2.7.1.21)
MCP_HHV2H Major capsid protein (MCP)
NEC1_HHV2H Nuclear egress protein 1
NEC2_HHV2H Nuclear egress protein 2
NP04_HHV2H Nuclear protein UL4
NP03_HHV2H Nuclear phosphoprotein UL3
OBP_HHV2H Replication origin-binding protein (OBP) (OriBP)
PRIM_HHV2H DNA primase (EC 2.7.7.-)
PORTL_HHV2H Portal protein
RIR2_HHV2H Ribonucleoside-diphosphate reductase small subunit (EC 1.17.4.1) (Ribonucleotide reductase small ...
RIR1_HHV2H Ribonucleoside-diphosphate reductase large subunit (R1) (EC 1.17.4.1) (Ribonucleotide reductase ...
RNB_HHV2H Probable RNA-binding protein
SCAF_HHV2H Capsid scaffolding protein (Capsid protein P40) (Protease precursor) (pPR) (Virion structural ...
SCP_HHV2H Small capsomere-interacting protein
SHUT_HHV2H Virion host shutoff protein (Vhs) (EC 3.1.27.-)
TEG4_HHV2H Tegument protein UL21
TEG3_HHV2H Tegument protein UL14
TEG5_HHV2H Tegument protein UL47 (82/81 kDa tegument protein) (VMW82/81) (VP13/14)
TEG7_HHV2H Tegument protein UL51
TEG1_HHV2H Tegument protein UL46 (Tegument protein VP11/12)
TEG6_HHV2H Tegument protein UL55
TRM1_HHV2H Tripartite terminase subunit 1
TRM2_HHV2H Tripartite terminase subunit 2
TRM3_HHV2H Tripartite terminase subunit 3 (EC 3.1.-.-) (Terminase large subunit)
UL20_HHV2H Protein UL20
TRX1_HHV2H Triplex capsid protein 1
TRX2_HHV2H Triplex capsid protein 2
UL32_HHV2H Packaging protein UL32
UL56_HHV2H Protein UL56
UNG_HHV2H Uracil-DNA glycosylase (UDG) (EC 3.2.2.27) (UNG)
US02_HHV2H Protein US2
US10_HHV2H Virion protein US10
US03_HHV2H Serine/threonine-protein kinase US3 (EC 2.7.11.1)
US8A_HHV2H Protein US8.5
VP16_HHV2H Tegument protein VP16 (Alpha trans-inducing protein) (Alpha-TIF) (ICP25) (Vmw65)
UL13_HHV2H Serine/threonine-protein kinase UL13 (EC 2.7.11.1)
UL24_HHV2H Protein UL24
US9_HHV2H Envelope protein US9 (10 kDa protein)
VP22_HHV2H Tegument protein VP22

1 entry

Cercopithecine herpesvirus 1 (CeHV-1) (Simian herpes B virus) reference strain

GE_CHV1 Envelope glycoprotein E (gE)

15 entries

Human herpesvirus 1 (strain F) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
DNBI_HHV1F Major DNA-binding protein
GE_HHV1F Envelope glycoprotein E (gE) (gE-1)
GB_HHV1F Envelope glycoprotein B (gB)
GD_HHV1F Envelope glycoprotein D (gD)
GH_HHV1F Envelope glycoprotein H (gH)
LRP1_HHV1F Latency-related protein 1
ICP34_HHV1F Neurovirulence factor ICP34.5 (Infected cell protein 34.5) (protein gamma(1)34.5)
LRP2_HHV1F Latency-related protein 2
NP03_HHV1F Nuclear phosphoprotein UL3
TEG1_HHV1F Tegument protein UL46 (Tegument protein VP11/12)
TEG5_HHV1F Tegument protein UL47 (82/81 kDa tegument protein) (VMW82/81) (VP13/14)
TRM1_HHV1F Tripartite terminase subunit 1
US8A_HHV1F US8.5 protein
US10_HHV1F Virion protein US10
VP16_HHV1F Tegument protein VP16 (Alpha trans-inducing protein) (Alpha-TIF) (ICP25) (Vmw65)

12 entries

Human herpesvirus 1 (strain KOS) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
CEP3_HHV1K Cytoplasmic envelopment protein 3
DNBI_HHV1K Major DNA-binding protein
DPOL_HHV1K DNA polymerase catalytic subunit (EC 2.7.7.7) (EC 3.1.26.4)
EV45_HHV1K Envelope protein UL45 (18 kDa protein)
GB_HHV1K Envelope glycoprotein B (gB)
GC_HHV1K Envelope glycoprotein C
GK_HHV1K Envelope glycoprotein K (Syncytial protein)
GD_HHV1K Envelope glycoprotein D (gD)
GL_HHV1K Envelope glycoprotein L (gL)
KITH_HHV1K Thymidine kinase (EC 2.7.1.21)
RIR2_HHV1K Ribonucleoside-diphosphate reductase small subunit (EC 1.17.4.1) (Ribonucleotide reductase small ...
SHUT_HHV1K Virion host shutoff protein (Vhs) (EC 3.1.27.-)

12 entries

Human herpesvirus 2 (strain 333) (HHV-2) (Human herpes simplex virus 2)

Select_allDeselect_all
EV45_HHV23 Envelope protein UL45 (18 kDa protein)
GC_HHV23 Envelope glycoprotein C
GI_HHV23 Envelope glycoprotein I (gI)
GB_HHV23 Envelope glycoprotein B (gB)
GD_HHV23 Envelope glycoprotein D (gD)
KITH_HHV23 Thymidine kinase (EC 2.7.1.21)
NP03_HHV23 Nuclear phosphoprotein UL3
RIR1_HHV23 Ribonucleoside-diphosphate reductase large subunit (R1) (EC 1.17.4.1) (Ribonucleotide reductase ...
RIR2_HHV23 Ribonucleoside-diphosphate reductase small subunit (EC 1.17.4.1) (Ribonucleotide reductase small ...
SHUT_HHV23 Virion host shutoff protein (Vhs) (EC 3.1.27.-)
UNG_HHV23 Uracil-DNA glycosylase (UDG) (EC 3.2.2.27) (UNG)
VP16_HHV23 Tegument protein VP16 (Alpha trans-inducing protein) (Alpha-TIF) (ICP25) (Vmw65)

6 entries

Human herpesvirus 1 (strain HFEM) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
GH_HHV1E Envelope glycoprotein H (gH)
ICP27_HHV1E mRNA export factor (Immediate-early protein IE63) (Infected cell protein 27) (ICP27)
KITH_HHV1E Thymidine kinase (EC 2.7.1.21)
TEG4_HHV1E Tegument protein UL21
TEG6_HHV1E Tegument protein UL55
UL56_HHV1E Protein UL56

3 entries

Cercopithecine herpesvirus 1 (CeHV-1) (Simian herpes B virus)

Select_allDeselect_all
GD_CHV1 Envelope glycoprotein D (gD)
US9_CHV1 Envelope protein US9 homolog (10 kDa protein)
VGLJ_CHV1 Envelope glycoprotein J

3 entries

Bovine herpesvirus 2 (strain BMV) (BoHV-2) (Bovine mammillitis virus)

Select_allDeselect_all
DNBI_BHV2B Major DNA-binding protein
GB_BHV2B Envelope glycoprotein B (gB)
TRM1_BHV2B Tripartite terminase subunit 1

3 entries

Human herpesvirus 1 (strain Angelotti) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
DPOL_HHV1A DNA polymerase catalytic subunit (EC 2.7.7.7) (EC 3.1.26.4)
GD_HHV1A Envelope glycoprotein D (gD)
TRM1_HHV1A Tripartite terminase subunit 1

3 entries

Human herpesvirus 1 (strain MP) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
EV45_HHV1M Envelope protein UL45 (18 kDa protein)
GK_HHV1M Envelope glycoprotein K (Syncytial protein)
RNB_HHV1M RNA-binding protein (Vmw21)

3 entries

Human herpesvirus 2 (strain G) (HHV-2) (Human herpes simplex virus 2)

Select_allDeselect_all
GC_HHV2G Envelope glycoprotein C (Glycoprotein F)
SHUT_HHV2G Virion host shutoff protein (Vhs) (EC 3.1.27.-)
TRX1_HHV2G Triplex capsid protein 1

2 entries

Herpes simplex virus type 2 (strain SA8) (Simian agent 8)

Select_allDeselect_all
GB_HSV2S Envelope glycoprotein B (gB)
ICP47_HSV2S ICP47 protein (Immediate-early protein IE12) (Immediate-early-5) (Infected cell protein 47) (US12 ...

2 entries

Human herpesvirus 1 (strain Patton) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
GD_HHV1P Envelope glycoprotein D (gD)
GB_HHV1P Envelope glycoprotein B (gB)

2 entries

Human herpesvirus 1 (strain SC16) (HHV-1) (Human herpes simplex virus 1)

Select_allDeselect_all
DPOL_HHV1S DNA polymerase catalytic subunit (EC 2.7.7.7) (EC 3.1.26.4)
KITH_HHV1S Thymidine kinase (EC 2.7.1.21)

2 entries

Human herpesvirus 2 (HHV-2) (Human herpes simplex virus 2)

Select_allDeselect_all
AN_HHV2 Alkaline nuclease (EC 3.1.-.-)
DNBI_HHV2 Major DNA-binding protein

2 entries

Saimiriine herpesvirus 1 (strain MV-5-4-PSL) (SaHV-1) (Marmoset herpesvirus)

Select_allDeselect_all
GB_SHV1 Envelope glycoprotein B (gB)
KITH_SHV1 Thymidine kinase (EC 2.7.1.21)

1 entry

Bovine herpesvirus 2 (strain BHM-1) (BoHV-2) (Bovine mammillitis virus)

KITH_BHV2H Thymidine kinase (EC 2.7.1.21)

1 entry

Cercopithecine herpesvirus 1 (strain E2490) (CeHV-1) (Simian herpes B virus)

ICP47_CHV1E ICP47 protein (Immediate-early protein IE12) (Immediate-early-5) (Infected cell protein 47) (US12 ...

1 entry

Cercopithecine herpesvirus 16 (CeHV-16) (Herpesvirus papio 2)

ICP47_CHV16 ICP47 protein (Immediate-early protein IE12) (Immediate-early-5) (Infected cell protein 47) (US12 ...

1 entry

Herpesvirus ateles type 1 (strain Lennette)

GB_HSVA1 Envelope glycoprotein B (gB)

1 entry

Human herpesvirus 1 (HHV-1) (Human herpes simplex virus 1)

KITH_HHV1 Thymidine kinase (EC 2.7.1.21)

1 entry

Human herpesvirus 1 (strain A44) (HHV-1) (Human herpes simplex virus 1)

TRX1_HHV14 Triplex capsid protein 1

1 entry

Human herpesvirus 1 (strain CL101) (HHV-1) (Human herpes simplex virus 1)

KITH_HHV1C Thymidine kinase (EC 2.7.1.21)

1 entry

Human herpesvirus 1 (strain CVG-2) (HHV-1) (Human herpes simplex virus 1)

ICP34_HHV1D Neurovirulence factor ICP34.5 (Infected cell protein 34.5) (protein gamma(1)34.5)

1 entry

Human herpesvirus 1 (strain HZT) (HHV-1) (Human herpes simplex virus 1)

GD_HHV1H Envelope glycoprotein D (gD)

1 entry

Human herpesvirus 1 (strain MGH-10) (HHV-1) (Human herpes simplex virus 1)

ICP34_HHV1N Neurovirulence factor ICP34.5 (Infected cell protein 34.5) (protein gamma(1)34.5)

1 entry

Human herpesvirus 1 (strain R15) (HHV-1) (Human herpes simplex virus 1)

GK_HHV1R Envelope glycoprotein K (Syncytial protein)

1 entry

Human herpesvirus 1 (strain R19) (HHV-1) (Human herpes simplex virus 1)

GK_HHV19 Envelope glycoprotein K (Syncytial protein)

1 entry

Human herpesvirus 2 (strain 186) (HHV-2) (Human herpes simplex virus 2)

DPOL_HHV21 DNA polymerase catalytic subunit (EC 2.7.7.7) (EC 3.1.26.4)

1 entry

Human herpesvirus 2 (strain B4327UR) (HHV-2) (Human herpes simplex virus 2)

GG_HHV2B Envelope glycoprotein G (gG) (gG-2)