Alphapapillomavirus (taxid:333750)

VIRION

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Non-enveloped. Small, icosahedral, about 60 nm in size. A single molecule of circular dsDNA is contained within the T=7 icosahedral capsid, which is composed of 72 pentamers.
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GENOME

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Circular dsDNA, about 8 kb in size, associated with cellular histones.
On rare non-specific recombination, the viral genome can be integrated in host chromosome. This inactivates the integrated virus but can gives the host cell a replicative advantage sometimes leading to malignant tumours.

GENE EXPRESSION

Only one strand of the genome is transcribed and yield two classes of proteins expressed by alternative splicing :
a) Early Proteins: non-structural regulatory proteins (E1-E7).
b) Late Proteins: the structural proteins L1 and L2. E7 can be expressed by leaky scanning from the E6 mRNA.

REPLICATION

NUCLEAR

Replication is divided in two distinct steps that are linked to the differentiation state of the host epithelial cell:

a) The plasmid replication takes place in the basal squamous epithelial cells. It corresponds to viral DNA replication in synchrony with the host cell chromosome in order to ensures an average of one viral genome per basal cell.

  1. Attachment of the viral proteins to host receptors mediates endocytosis into vesicles in the basal squamous epithelial cell.
  2. Transport to the nucleus and uncoating of the viral DNA.
  3. Early-region transcription and translation of the early proteins.
  4. Steady-state viral DNA nuclear replication. Requires the origin of DNA replication in cis and the viral E1 and E2 proteins in trans.

b) The vegetative replication, which occurs in differentiated keratinocytes. In these cells, which no longer undergo cellular DNA synthesis, there is a burst of viral DNA synthesis with active production of virions.

  1. Vegetative viral DNA synthesis.
  2. Transcription of the late region.
  3. Capsid proteins L1 and L2 synthesis.
  4. Nuclear capsid assembly and release of viruses.

Host-virus interaction

Adaptive immune response inhibition

The E5 protein in papillomaviruses plays a role in the inhibition of host adaptive response by down-regulating the cell surface expression of MHC class I .

Apoptosis modulation

The HPV16-E6 protein has the ability to prevent apoptosis of infected cells through its binding to FADD and caspase 8 .

Cell-cycle modulation

The E7 protein from the high-risk HPV type 18 forms a complex with cyclin E and regulates transit from G1 to S. E7 binds also cyclin A, which regulates transit through the S and G2/M phases of the cell cycle .

Innate immune response inhibition

Papillomaviruses inhibits the cascade leading to production of interferon-beta by targeting the IRF3 cellular protein. Indeed, The viral E6 protein is able to interact with host IRF3 and inhibits its transcriptional activity .

Matching UniProtKB/Swiss-Prot entries

(all links/actions below point to uniprot.org website)

303 entries grouped by protein

40 entries

Replication protein E1 (EC 3.6.4.12) (ATP-dependent helicase E1)

Select_allDeselect_all
VE1_HPV07
Human papillomavirus 7 reference strain
VE1_HPV10
Human papillomavirus type 10 reference strain
VE1_HPV11
Human papillomavirus 11 reference strain
VE1_HPV16
Human papillomavirus type 16 reference strain
VE1_HPV18
Human papillomavirus type 18 reference strain
VE1_HPV26
Human papillomavirus type 26 reference strain
VE1_HPV27
Human papillomavirus 27 reference strain
VE1_HPV29
Human papillomavirus 29 reference strain
VE1_HPV31
Human papillomavirus 31 reference strain
VE1_HPV32
Human papillomavirus type 32 reference strain
VE1_HPV35
Human papillomavirus 35 reference strain
VE1_HPV44
Human papillomavirus 44 reference strain
VE1_HPV61
Human papillomavirus type 61 reference strain
VE1_HPV70
Human papillomavirus type 70 reference strain
VE1_HPV73
Human papillomavirus 73 reference strain
VE1_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VE1_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VE1_HPV03
Human papillomavirus 3
VE1_HPV13
Human papillomavirus 13
VE1_HPV28
Human papillomavirus 28
VE1_HPV2A
Human papillomavirus type 2a
VE1_HPV30
Human papillomavirus 30
VE1_HPV33
Human papillomavirus 33
VE1_HPV34
Human papillomavirus type 34
VE1_HPV39
Human papillomavirus 39
VE1_HPV40
Human papillomavirus 40
VE1_HPV42
Human papillomavirus 42
VE1_HPV45
Human papillomavirus 45
VE1_HPV51
Human papillomavirus 51
VE1_HPV52
Human papillomavirus 52
VE1_HPV54
Human papillomavirus type 54
VE1_HPV57
Human papillomavirus 57
VE1_HPV58
Human papillomavirus 58
VE1_HPV59
Human papillomavirus 59
VE1_HPV66
Human papillomavirus 66
VE1_HPV69
Human papillomavirus 69
VE1_HPV6A
Human papillomavirus type 6a
VE1_HPV6B
Human papillomavirus type 6b
VE1_HPV72
Human papillomavirus 72
VE1_PCPVC
Pygmy chimpanzee papillomavirus type 1C (PCPV-1C)

39 entries

Regulatory protein E2

Select_allDeselect_all
VE2_HPV07
Human papillomavirus 7 reference strain
VE2_HPV10
Human papillomavirus type 10 reference strain
VE2_HPV11
Human papillomavirus 11 reference strain
VE2_HPV16
Human papillomavirus type 16 reference strain
VE2_HPV18
Human papillomavirus type 18 reference strain
VE2_HPV26
Human papillomavirus type 26 reference strain
VE2_HPV27
Human papillomavirus 27 reference strain
VE2_HPV29
Human papillomavirus 29 reference strain
VE2_HPV31
Human papillomavirus 31 reference strain
VE2_HPV32
Human papillomavirus type 32 reference strain
VE2_HPV35
Human papillomavirus 35 reference strain
VE2_HPV44
Human papillomavirus 44 reference strain
VE2_HPV53
Human papillomavirus type 53 reference strain
VE2_HPV56
Human papillomavirus 56 reference strain
VE2_HPV61
Human papillomavirus type 61 reference strain
VE2_HPV70
Human papillomavirus type 70 reference strain
VE2_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VE2_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VE2_HPV03
Human papillomavirus 3
VE2_HPV13
Human papillomavirus 13
VE2_HPV28
Human papillomavirus 28
VE2_HPV2A
Human papillomavirus type 2a
VE2_HPV30
Human papillomavirus 30
VE2_HPV33
Human papillomavirus 33
VE2_HPV34
Human papillomavirus type 34
VE2_HPV39
Human papillomavirus 39
VE2_HPV40
Human papillomavirus 40
VE2_HPV42
Human papillomavirus 42
VE2_HPV45
Human papillomavirus 45
VE2_HPV51
Human papillomavirus 51
VE2_HPV52
Human papillomavirus 52
VE2_HPV54
Human papillomavirus type 54
VE2_HPV55
Human papillomavirus type 55
VE2_HPV57
Human papillomavirus 57
VE2_HPV58
Human papillomavirus 58
VE2_HPV66
Human papillomavirus 66
VE2_HPV69
Human papillomavirus 69
VE2_HPV6A
Human papillomavirus type 6a
VE2_HPV6B
Human papillomavirus type 6b

21 entries

Protein E4

Select_allDeselect_all
VE4_HPV11
Human papillomavirus 11 reference strain
VE4_HPV16
Human papillomavirus type 16 reference strain

Protein E4 (E1^E4)

VE4_HPV18
Human papillomavirus type 18 reference strain
VE4_HPV29
Human papillomavirus 29 reference strain
VE4_HPV31
Human papillomavirus 31 reference strain
VE4_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain

Probable protein E4

VE4_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain

Probable protein E4

VE4_HPV13
Human papillomavirus 13
VE4_HPV28
Human papillomavirus 28
VE4_HPV2A
Human papillomavirus type 2a
VE4_HPV33
Human papillomavirus 33

Protein E4 (E1^E4)

VE4_HPV35
Human papillomavirus 35
VE4_HPV39
Human papillomavirus 39
VE4_HPV42
Human papillomavirus 42
VE4_HPV51
Human papillomavirus 51
VE4_HPV54
Human papillomavirus type 54
VE4_HPV57
Human papillomavirus 57
VE4_HPV58
Human papillomavirus 58
VE4_HPV6A
Human papillomavirus type 6a
VE4_HPV6B
Human papillomavirus type 6b

Protein E4 (E1^E4)

VE4_HPV6C
Human papillomavirus type 6c

Probable protein E4

13 entries

Probable protein E5

Select_allDeselect_all
VE5_HPV16
Human papillomavirus type 16 reference strain
VE5_HPV18
Human papillomavirus type 18 reference strain
VE5_HPV31
Human papillomavirus 31 reference strain
VE5_HPV70
Human papillomavirus type 70 reference strain
VE5_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VE5_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VE5_HPV13
Human papillomavirus 13
VE5_HPV33
Human papillomavirus 33
VE5_HPV35
Human papillomavirus 35
VE5_HPV39
Human papillomavirus 39
VE5_HPV42
Human papillomavirus 42
VE5_HPV51
Human papillomavirus 51
VE5_HPV58
Human papillomavirus 58

4 entries

Probable protein E5A

Select_allDeselect_all
VE5A_HPV11
Human papillomavirus 11 reference strain
VE5A_HPV6A
Human papillomavirus type 6a
VE5A_HPV6B
Human papillomavirus type 6b
VE5A_HPV6C
Human papillomavirus type 6c

3 entries

Probable protein E5B

Select_allDeselect_all
VE5B_HPV11
Human papillomavirus 11 reference strain
VE5B_HPV6B
Human papillomavirus type 6b
VE5B_HPV6C
Human papillomavirus type 6c

41 entries

Protein E6

Select_allDeselect_all
VE6_HPV07
Human papillomavirus 7 reference strain
VE6_HPV10
Human papillomavirus type 10 reference strain
VE6_HPV11
Human papillomavirus 11 reference strain
VE6_HPV16
Human papillomavirus type 16 reference strain
VE6_HPV18
Human papillomavirus type 18 reference strain
VE6_HPV26
Human papillomavirus type 26 reference strain
VE6_HPV27
Human papillomavirus 27 reference strain
VE6_HPV29
Human papillomavirus 29 reference strain
VE6_HPV31
Human papillomavirus 31 reference strain
VE6_HPV32
Human papillomavirus type 32 reference strain
VE6_HPV35
Human papillomavirus 35 reference strain
VE6_HPV44
Human papillomavirus 44 reference strain
VE6_HPV53
Human papillomavirus type 53 reference strain
VE6_HPV56
Human papillomavirus 56 reference strain
VE6_HPV61
Human papillomavirus type 61 reference strain
VE6_HPV70
Human papillomavirus type 70 reference strain
VE6_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VE6_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VE6_HPV03
Human papillomavirus 3
VE6_HPV13
Human papillomavirus 13
VE6_HPV28
Human papillomavirus 28
VE6_HPV2A
Human papillomavirus type 2a
VE6_HPV30
Human papillomavirus 30
VE6_HPV33
Human papillomavirus 33
VE6_HPV34
Human papillomavirus type 34
VE6_HPV39
Human papillomavirus 39
VE6_HPV40
Human papillomavirus 40
VE6_HPV42
Human papillomavirus 42
VE6_HPV43
Human papillomavirus 43
VE6_HPV45
Human papillomavirus 45
VE6_HPV51
Human papillomavirus 51
VE6_HPV52
Human papillomavirus 52
VE6_HPV54
Human papillomavirus type 54
VE6_HPV57
Human papillomavirus 57
VE6_HPV58
Human papillomavirus 58
VE6_HPV66
Human papillomavirus 66
VE6_HPV68
Human papillomavirus 68
VE6_HPV69
Human papillomavirus 69
VE6_HPV6A
Human papillomavirus type 6a
VE6_HPV6B
Human papillomavirus type 6b
VE6_HPV94
Human papillomavirus type 94

40 entries

Protein E7

Select_allDeselect_all
VE7_HPV07
Human papillomavirus 7 reference strain
VE7_HPV10
Human papillomavirus type 10 reference strain
VE7_HPV11
Human papillomavirus 11 reference strain
VE7_HPV16
Human papillomavirus type 16 reference strain
VE7_HPV18
Human papillomavirus type 18 reference strain
VE7_HPV26
Human papillomavirus type 26 reference strain
VE7_HPV27
Human papillomavirus 27 reference strain
VE7_HPV29
Human papillomavirus 29 reference strain
VE7_HPV31
Human papillomavirus 31 reference strain
VE7_HPV32
Human papillomavirus type 32 reference strain
VE7_HPV35
Human papillomavirus 35 reference strain
VE7_HPV44
Human papillomavirus 44 reference strain
VE7_HPV53
Human papillomavirus type 53 reference strain
VE7_HPV56
Human papillomavirus 56 reference strain
VE7_HPV61
Human papillomavirus type 61 reference strain
VE7_HPV70
Human papillomavirus type 70 reference strain
VE7_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VE7_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VE7_HPV13
Human papillomavirus 13
VE7_HPV28
Human papillomavirus 28
VE7_HPV2A
Human papillomavirus type 2a
VE7_HPV30
Human papillomavirus 30
VE7_HPV33
Human papillomavirus 33
VE7_HPV34
Human papillomavirus type 34
VE7_HPV39
Human papillomavirus 39
VE7_HPV40
Human papillomavirus 40
VE7_HPV42
Human papillomavirus 42
VE7_HPV45
Human papillomavirus 45
VE7_HPV51
Human papillomavirus 51
VE7_HPV52
Human papillomavirus 52
VE7_HPV54
Human papillomavirus type 54
VE7_HPV55
Human papillomavirus type 55
VE7_HPV57
Human papillomavirus 57
VE7_HPV58
Human papillomavirus 58
VE7_HPV66
Human papillomavirus 66
VE7_HPV68
Human papillomavirus 68
VE7_HPV69
Human papillomavirus 69
VE7_HPV6A
Human papillomavirus type 6a
VE7_HPV6B
Human papillomavirus type 6b
VE7_HPV94
Human papillomavirus type 94

15 entries

Protein E8^E2C

Select_allDeselect_all
VE8E2_HPV11
Human papillomavirus 11 reference strain
VE8E2_HPV16
Human papillomavirus type 16 reference strain
VE8E2_HPV18
Human papillomavirus type 18 reference strain
VE8E2_HPV29
Human papillomavirus 29 reference strain
VE8E2_HPV31
Human papillomavirus 31 reference strain
VE8E2_HPV13
Human papillomavirus 13
VE8E2_HPV28
Human papillomavirus 28
VE8E2_HPV33
Human papillomavirus 33
VE8E2_HPV35
Human papillomavirus 35
VE8E2_HPV39
Human papillomavirus 39
VE8E2_HPV42
Human papillomavirus 42
VE8E2_HPV51
Human papillomavirus 51
VE8E2_HPV54
Human papillomavirus type 54
VE8E2_HPV57
Human papillomavirus 57
VE8E2_HPV58
Human papillomavirus 58

44 entries

Major capsid protein L1

Select_allDeselect_all
VL1_HPV07
Human papillomavirus 7 reference strain
VL1_HPV10
Human papillomavirus type 10 reference strain
VL1_HPV11
Human papillomavirus 11 reference strain
VL1_HPV16
Human papillomavirus type 16 reference strain
VL1_HPV18
Human papillomavirus type 18 reference strain
VL1_HPV26
Human papillomavirus type 26 reference strain
VL1_HPV27
Human papillomavirus 27 reference strain
VL1_HPV29
Human papillomavirus 29 reference strain
VL1_HPV31
Human papillomavirus 31 reference strain
VL1_HPV32
Human papillomavirus type 32 reference strain
VL1_HPV35
Human papillomavirus 35 reference strain
VL1_HPV44
Human papillomavirus 44 reference strain
VL1_HPV53
Human papillomavirus type 53 reference strain
VL1_HPV56
Human papillomavirus 56 reference strain
VL1_HPV61
Human papillomavirus type 61 reference strain
VL1_HPV62
Human papillomavirus 62 reference strain
VL1_HPV70
Human papillomavirus type 70 reference strain
VL1_HPV82
Human papillomavirus 82 reference strain
VL1_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VL1_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VL1_HPV03
Human papillomavirus 3
VL1_HPV13
Human papillomavirus 13
VL1_HPV28
Human papillomavirus 28
VL1_HPV2A
Human papillomavirus type 2a
VL1_HPV30
Human papillomavirus 30
VL1_HPV33
Human papillomavirus 33
VL1_HPV34
Human papillomavirus type 34
VL1_HPV39
Human papillomavirus 39
VL1_HPV40
Human papillomavirus 40
VL1_HPV42
Human papillomavirus 42
VL1_HPV43
Human papillomavirus 43
VL1_HPV45
Human papillomavirus 45
VL1_HPV51
Human papillomavirus 51
VL1_HPV52
Human papillomavirus 52
VL1_HPV54
Human papillomavirus type 54
VL1_HPV55
Human papillomavirus type 55
VL1_HPV57
Human papillomavirus 57
VL1_HPV58
Human papillomavirus 58
VL1_HPV66
Human papillomavirus 66
VL1_HPV67
Human papillomavirus 67
VL1_HPV68
Human papillomavirus 68
VL1_HPV69
Human papillomavirus 69
VL1_HPV6A
Human papillomavirus type 6a
VL1_HPV6B
Human papillomavirus type 6b

43 entries

Minor capsid protein L2

Select_allDeselect_all
VL2_HPV07
Human papillomavirus 7 reference strain
VL2_HPV10
Human papillomavirus type 10 reference strain
VL2_HPV11
Human papillomavirus 11 reference strain
VL2_HPV16
Human papillomavirus type 16 reference strain
VL2_HPV18
Human papillomavirus type 18 reference strain
VL2_HPV26
Human papillomavirus type 26 reference strain
VL2_HPV27
Human papillomavirus 27 reference strain
VL2_HPV29
Human papillomavirus 29 reference strain
VL2_HPV31
Human papillomavirus 31 reference strain
VL2_HPV32
Human papillomavirus type 32 reference strain
VL2_HPV35
Human papillomavirus 35 reference strain
VL2_HPV44
Human papillomavirus 44 reference strain
VL2_HPV53
Human papillomavirus type 53 reference strain
VL2_HPV56
Human papillomavirus 56 reference strain
VL2_HPV61
Human papillomavirus type 61 reference strain
VL2_HPV62
Human papillomavirus 62 reference strain
VL2_HPV70
Human papillomavirus type 70 reference strain
VL2_HPV82
Human papillomavirus 82 reference strain
VL2_MMPV1
Macaca mulata papillomavirus 1 (Rhpv 1) (Rhesus papillomavirus type 1) reference strain
VL2_PCPV1
Pygmy chimpanzee papillomavirus type 1 (PCPV-1) reference strain
VL2_HPV03
Human papillomavirus 3
VL2_HPV13
Human papillomavirus 13
VL2_HPV28
Human papillomavirus 28
VL2_HPV2A
Human papillomavirus type 2a
VL2_HPV30
Human papillomavirus 30
VL2_HPV33
Human papillomavirus 33
VL2_HPV34
Human papillomavirus type 34
VL2_HPV39
Human papillomavirus 39
VL2_HPV40
Human papillomavirus 40
VL2_HPV42
Human papillomavirus 42
VL2_HPV43
Human papillomavirus 43
VL2_HPV45
Human papillomavirus 45
VL2_HPV51
Human papillomavirus 51
VL2_HPV52
Human papillomavirus 52
VL2_HPV54
Human papillomavirus type 54
VL2_HPV55
Human papillomavirus type 55
VL2_HPV57
Human papillomavirus 57
VL2_HPV58
Human papillomavirus 58
VL2_HPV66
Human papillomavirus 66
VL2_HPV67
Human papillomavirus 67
VL2_HPV69
Human papillomavirus 69
VL2_HPV6A
Human papillomavirus type 6a
VL2_HPV6B
Human papillomavirus type 6b
These structures were created with the help of Colabfold by Jason Nomburg, Nathan Price and Jennifer Doudna (ModelArchive).

Alphapapillomavirus 9 taxid:337041

Protein ModelArchive
E5 ma-jd-viral-59228
Major capsid protein L1 ma-jd-viral-67296
Minor capsid protein L2 ma-jd-viral-44935
Protein E7 ma-jd-viral-13716
Regulatory protein E2 ma-jd-viral-61179
Replication protein E1 (EC 3.6.4.12) (ATP-dependent helicase E1) ma-jd-viral-21239

Human papillomavirus type 16 taxid:333760

Protein ModelArchive
E7 ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
E7 protein ma-jd-viral-13716
Major capsid protein L1 ma-jd-viral-67296
Major capsid protein L1 ma-jd-viral-67296
Major capsid protein L1 ma-jd-viral-67296
Minor capsid protein L2 ma-jd-viral-44935
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Oncoprotein E7 ma-jd-viral-13716
Probable protein E5 ma-jd-viral-59228
Probable protein E5 ma-jd-viral-59228
Probable protein E5 ma-jd-viral-59228
Protein E4 ma-jd-viral-55570
Protein E4 (E1^E4) ma-jd-viral-55570
Protein E6 ma-jd-viral-00866
Protein E6 ma-jd-viral-00866
Protein E6 ma-jd-viral-00866
Protein E6 ma-jd-viral-00866
Protein E6 ma-jd-viral-00866
Protein E6 ma-jd-viral-00866
Protein E7 ma-jd-viral-13716
Protein E7 ma-jd-viral-13716
Protein E7 ma-jd-viral-13716
Regulatory protein E2 ma-jd-viral-61179
Regulatory protein E2 ma-jd-viral-61179
Regulatory protein E2 ma-jd-viral-61179
Replication protein E1 (EC 3.6.4.12) (ATP-dependent helicase E1) ma-jd-viral-21239
Replication protein E1 (EC 3.6.4.12) (ATP-dependent helicase E1) ma-jd-viral-21239

Papio hamadryas papillomavirus 1 taxid:990303

Protein ModelArchive
Early protein E4 ma-jd-viral-54834
Early protein E5 ma-jd-viral-66451
Major capsid protein L1 ma-jd-viral-67269
Minor capsid protein L2 ma-jd-viral-44943
Protein E6 ma-jd-viral-00859
Protein E7 ma-jd-viral-13719
Regulatory protein E2 ma-jd-viral-61115
Replication protein E1 (EC 3.6.4.12) (ATP-dependent helicase E1) ma-jd-viral-21321