TABLE 1

Gene content and major features of the complete 205,896-bp EEHV4(Baylor) genomeh,i

Gene name and orientationProtein nameTypeFamily or statusPosition coordinates% GC contentProtein size (aa)% amino acid identity (% length matched) to:Note or comment
EEHV1A KimbaEEHV1A RamanEEHV1B EmeliaEEHV5 Vijay
TR3.5× 22-bp340–420Related to multimerized 17-bp repeats in TR of EEHV1A/1B/5
TRRegulatory motifs1070–1410All have a cluster of 6–9× palindromic (8-bp) CREB-binding sites
NilvFUT9NovelE47EE63EE63EE63Absent in EEHV4a
Nil7xTMNovelNilNilNilEE62BUnique to EEHV5
NilIgFamNilNilNilEE62AUnique to EEHV5
NilvGPCR77xTME3famE48EE62NilEE62Absent in EEHV4
NilE49famE49EE61NilFragAbsent in EEHV4
NilvIgF1NovelE50EE60NilNilAbsent in EEHV4, -5 outside the probosciviruses
NilvGPCR87xTME3famFragFragEE59NilAbsent in EEHV4
NilE49famE51EE58EE58NilAbsent in EEHV4
NilvIgF2IgFamE52EE57FragNilAbsent in EEHB4
NilE49famNilNilEE56NilAbsent in EEHV4
NilIgFamNilNilEE55NilAbsent in EEHV4
NilIgFamNilNilEE54NilAbsent in EEHV4
NilvIgF2.4IgFamFragEE53EE53NilAbsent in EEHV4
NilvIgF2.5IgFamE53EE52EE52EE52Absent in EEHV4
NilvOX2-1NovelE54EE51EE51EE51Absent in EEHV4
NilvIgF3IgFamE55EE50EE50EE50Absent in EEHV4
NilvCD48IgFamNilNilNilEE49DUnique to EEHV5
NilIgFamNilNilNilEE49CUnique to EEHV5
NilvCD48IgFamNilNilNilEE49BUnique to EEHV5
NilIgFamNilNilNilEE49AUnique to EEHV5
E1, F7xTME3fam2061–36086951525 (45)EE49EE49EE49N-term S/T extended
NilCys-rich7xTME3famE2EE48EE48EE48Absent in EEHV4
E3, FvGPCR67xTME3fam3981–49286831530 (70)EE47EE47EE47Match to RAIP3 or C-5-Afam
E3.1, F vGPCR6.17xTME3fam4991–61696239228 (60)EE45EE45EE45Unique to EEHV4
E3.2, F vGPCR6.27xTME3fam6747–77276732830 (45)EE45EE45EE45Unique to EEHV4
E2A, F7xTMNovel8015–929558426NilNilNilNilUnique to EEHV4; S/T dom
E3.3, FvGPCR6.37xTME3fam9714–107215033534 (41)EE45EE45EE45Unique to EEHV4
E3.4, FvGPCR6.47xTME3fam11637–126745634538 (69)EE45EE45EE45Unique to EEHV4
E4, FvGCNT1AcTransfNovel13026–146426353861 (68)EE46EE46EE46b
NilvGPCR57xTME3famE5EE45EE45EE45Absent in EEHV4
NilNovelE5AEE44EE44NilShort, memb, absent EEHV4
NilvCD48IgFamNilNilNilEE44AUnique to EEHV5
E4A, FNovel15171–1565344160NilNilNilNilUnique to EEHV4
E4B, FNovel15712–1657856288NilNilNilNilUnique to EEHV4
E4C, FNovel16779–1723454151NilNilNilNilUnique to EEHV4
E6A, CE27ex1Novel17811–1739857137E27EE20NilEE2035% (44%) match to E27
E6B, FNovel17978–182564992NilNilNilNilUnique to EEHV4
E6, C7xTME6fam19679–188556027433 (89)EE43EE43EE43
Nil, CvCXCL2?E7ANilNilNilAbsent in EEHV1B/4/5
E7, C7xTME6fam20663–199565523532 (84)EE42EE42EE42
E7B, C7xTMNovel21647–2087126258NilNilNilNilUnique to EEHV4
Nil7xTME6famE8EE41EE41EE41Absent in EEHV4
E9, C7xTME6fam22653–217572729834 (18)EE40FragEE40Match to C-term EE40 only (esp EEHV5)
E9A, FvOGTAcTransfNovel22993–2423742414NilNilNilNilUnique to EEHV4c
E9B, C(Truncated)Novel24664–2428123127NilNilNilEE39/40Match to EEHV5 N-term16-aa EE39/40 only
E9C, CNovel25233–2471827171NilNilNilNilUnique to EEHV4
E10A, C7xTME6fam26078–2519761NilNilNil25 (46)Matches central EE40(EEHV5) only
Nil7xTME6famE10EE39EE39EE39Absent in EEHV4
E11, C7xTME6fam27127–263636125436 (96)EE38EE38EE38
E12, C7xTME6fam28314–274576028534 (77)EE37EE37EE37
E12A, CNovel28547–282816088NilNilNilNilUnique to EEHV4
E13, C7xTME6fam29780–289655827152 (88)EE36EE36EE36
E14.1, C7xTME14fam31024–302155926927 (91)NilNilNilDuplication of E14
E14.2, C7xTME14fam32191–312805230324 (77)NilNilNilDuplication of E14
E14, C7xTME14fam33228–324195726926 (79)EE35EE35EE35
E15, C vGPCR47xTME15fam34435–334255833634 (86)EE34EE34EE3426% (49%) match to Lox C-5-C
E16, C7xTME14fam35571–347685726740 (97)EE33EE33EE33
NilNovelE16CNilNilNilConserved in EEHV1 and EEHV5
NilNovelE16A/BNilNilNilSpliced; unique to EEHV1A/B
E16D, CvECTLNovel36141–3558445185NilNilNilNil
E17, FE27ex1Novel36491–368264511137 (99)EE32EE32EE32Match to E27, 30% (50%)
E17A, FNovel36944–3725263102NilNilNilNilUnique to EEHV4
NilNovelNilNilNilEE32AUnique to EEHV5
E18, F7xTME18fam37215–380185726732 (70)EE31EE31EE31Related to E28 by 30% (51%)
NilNovelE18BEE30AEE30AEE30AAbsent in EEHV4
NilNovelE18AEE30EE30EE30Absent in EEHV4
E18C, FNovel38433–387205994NilNilNilNilUnique to EEHV4
E19, FORF-F2U54.5fam39303–410426857952 (89)EE29EE29EE2925% (77%) to ORF-F1
E20, CvGPCR4A7xTME15fam43188–421545734446 (84)EE28EE28EE2823% (67%) match to Lox RAIP3
E20B, CNovel45323–4490465139NilNilNilNilUnique to EEHV4
E20A, FNovel45338–456646110843 (33)EE27EE27EE27
E21, CvGPCR4B7xTME15fam46925–458105937135 (76)EE26EE26EE2627% (35%) match to Lox RAIP3
E22, FNovel47646–47921499148 (96)EE25EE25EE25
E22A, FNovel48621–48730537946 (48)EE24EE24EE24
E23B, CNovel48993–4932557110NilNilNilNilUnique to EEHV4
E24B, CvOX2-Bex2Novel49596–4925056132E54EE51EE51EE51
vOX2-Bex1Novel50001–49950Short first exon
NilvOX2-3E24EE23EE23EE23Absent in EEHV4
NilvOX2-V (E23A)NilNilNilEE22AUnique to EEHV5
NilvOX2-2E25EE22EE22EE22Absent in EEHV4
E26, CvGPCR37xTME3fam51287–504185028942 (92)EE21EE21EE21 Match to ChemR C-5-C
E27, FE27ex1E27Novel52138–526065524557 (58)EE20ex1EE20ex1EE20ex1Related to E6A, E17
E27ex2Novel52785–5305359NilNilNilNilUnrelated to EE20ex2
E28, F7xTME18fam53115–538615424844 (90)EE19EE19EE19Related to E18 by 30% (51%)
E29, F7xTMNovel54092–547815522942 (91)EE18EE18EE18
E30, CNovel55453–5491155180<15EE17EE17EE17Acidic similarity only
E30A, CE30Aex2Novel55883–5553442133NilNilNilNilUnique to EEHV4
E30Aex1Novel56146–5609540NilNilNilNilUnique to EEHV4
NilE31EE16EE16EE16Absent in EEHV4
E31A, CNovel56923–563215620035 (46)EE15EE15EE15Only N-term cons
E31B, CNovel57113–56610NilNilNilNilUnique to EEHV4
E31C, CE31CexNovel57646–571826113665 (12)EE14EE14EE14No ATG, splice to E32?
E32, CU14.5βδ?60473–577176091845 (82)EE13EE13EE13
Nil, FE33EE12AFragNilUnique to EEHV1A
E33ANovel60991–61236508137 (59)EE12EE12EE12
U14, CU14βδ63078–614085955637 (75)U14U14U14
U13.5, CU13.5βδ64742–634445243276 (54)UL34UL34UL34
U12, CvGPCR2ex27xTMβδ67327–650605778350 (53)U12U12U12
vGPCR2ex167537–6745456 (100)U12U12U12Short first exon
E34, FORF-CNovel68025–74276592,08342 (16)U11U11U11Only N-term cons in EEHV1, and -5
U4, FU4U4βδ74389–760536155458 (94)U4U4U424% (38%) HHV6 U4
U4.5, FORF-BU4βδ76687–784446358559 (93)EE11EE11EE1124% (30%) U4
E35, FORF-ANovel79137–816596384051 (46)EE10EE10EE10
U44, CU44Core82518–837295540377 (23)U44U44U44Only C-term cons in EEHV1 and -5
U43, FPRICore83629–87234591,20151 (84)U43U43U43Primase subunit
U42, FMTAex187495–8762765 (51)U42U42U42Short first exon
MTAex2Core87948–92020641,40152 (24)U42U42U42Posttranscriptional regulator
Ori-Lyt92346–93325d
U41, FMDBPCore93861–97376611,17163 (99)U41U41U41SS DNA binding protein
U40, FTER2Core97498–995915969773 (98)U40U40U40
U39, FgBCore99533–1021215786264 (94)U39U39U39Env glycoprotein B
U38, FPOLCore102273–105527621,08465 (99)U38U38U38DNA polymerase
U37, CDOCCore106545–1057405926864 (97)U37U37U37Docking protein
U36, CCore108154–1065386353869 (89)U36U36U36
U35, FCore108266–108553489569 (98)U35U35U35
U34, FCore108717–1096195330063 (99)U34U34U34
U33, FCRPβγδ109914–1115186453449 (91)U33U33U33Cys-rich protein
U32, FSCPCore111409–111672588746 (67)U32U32U32Small capsid protein
U31, CTEG-LCore118856–111873652,32144 (89)U31U31U31Large tegument
U30, CTEG-SCore124420–119289611,71345 (53)U30U30U30Small tegument
U29, FTRI1Core124423–1253136029660 (98)U29U29U29Capsid triplex 1
U28, FRRACore125563–1280736183668 (66)U28U28U28Ribonucleotide reductase A
U27.5, FRRB (ORF-H)αγδ128212–1291175330175 (99)EE9EE9EE9Ribonucleotide reductase B
U27, FPPFCore129702–1310236443752 (65)U27U27U27Pol processivity factor
U45.7, FORF-JNovel131035–1317845821644 (33)EE8EE8EE8
U46, FgNCore131800–1321054910163 (53)U46U46U46Env glycoprotein N
U47, CgO (ORF-D)Βδ132837–1321875221634 (94)U47U47U47Env glycoprotein O
U48, CgHCore135086–1328164975647 (96)U48U48U48Env glycoprotein H
U48.5, CTK (ORF-E)αγδ136101–1350525634950 (87)EE7EE7EE7Thymidine kinase
U49, FCore136100–1368015723348 (85)U49U49U49
U50, FPAC2Core136620–1383475757564 (99)U50U50U50Packaging
U51, FvGPCR17xTMβδ138430–1396475640542 (95)U51U51U51e
U52, CCore140605–1398325325765 (98)U52U52U52
U53, FSCA/PROCore140698–1424856059549 (91)U53U53U53Scaffold protease
U54.5, CORF-F1U54.5fam144154–1427156147938 (99)U54U54U5427% (95%) match to ORF-F2
U56, CTRI2Core145344–1444455929968 (99)U56U56U56Capsid triplex 2
U5, CMCPCore149563–145511631,35071 (99)U57U57U57Major capsid protein
U58, FvTBPβγδ150117–153134611,00563 (87)U58U58U58TATA-binding protein
U59, Fβγδ152794–1541526245248 (79)U59U59U59
U60, CTERex3Core155504–1543775766092 (99)U60U60U60Terminase subunit 1
U62, Fβγδ155733–156005549057 (97)U62U62U62
U63, Fβγδ155944–1565465120067 (72)U63U63U63
U64, FPAC1Core156527–1585526454148 (63)U64U64U64Packaging
U65, FCore158055–1590715933848 (98)U65U65U65
U66, CTERex2Novel159272–15915390 (100)U66U66U66Terminase subunit 1
TERex1Core160224–1594905389 (99)U66U66U66Terminase subunit 1
U67, Fβγδ160612–1617425837661 (98)U67U67U67
U68, FCore161739–1621045112166 (98)U68U68U68
U69, FCPKCore162613–1642325953957 (96)U69U69U69Conserved protein kinase
U70, FEXOCore164529–1660946152153 (97)U70U70U70Exonuclease
U71, FMyrTegCore166031–1663425610341 (67)U71U71U71Myristylated tegument
U72, CgMCore167655–1665375537266 (93)U72U72U72Envelope glycoprotein M
U73, FOBP (ORF-G)αδ168094–171645611,18365 (68)U73U73U73Origin-binding protein
U74, FPAFCore171659–1738306372361 (91)U74U74U74Pol-associated factor
U75, CCore174625–1738136327054 (84)U75U75U75
U76, CPORCore176750–1745796372374 (78)U76U76U76Portal protein
U77, FHELCore176701–1795746395779 (79)U77U77U77Helicase subunit
E36, FORF-MNovel180833–1838056599062 (19)U79U79U79Env glycoprotein M (only N-term cons)
NilORF-N, vCXCL1NovelE36AEE6NilEE6Chemokine-like, absent in 1B, 4
U81, CUDGCore185284–1842776333568 (68)U81U81U81Uracil DNA glycosylase
U82, CgLCore186080–1852534827537 (94)U82U82U82Env glycoprotein L
E37, CORF-Oex3Novel186570–1860074170937 (95)EE5EE5EE5
ORF-Oex2Novel186897–18667740<15EE5EE5EE5
ORF-Oex1Novel188411–18706741<15EE5EE5EE5
NilORF-Pex2NovelE38EE4EE4EE4Absent in EEHV4
ORF-Pex1NovelE38EE4EE4EE4Absent in EEHV4
NilORF-Qex2NovelE39EE3EE3NilAbsent in EEHV2, -4, -5
ORF-Qex1NovelE39EE3EE3NilAbsent in EEHV2, -4, -5
E39A, CORF-Rex2Novel189482–18850640367NilNilNilNilUnique to EEHV4
ORF-Rex1Novel189690–18956432NilNilNilNilUnique to EEHV4
E40, CORF-KNovel194372–189975661,46572 (16)EE2EE2EE2Only C-term cons
E44A, CORF-SNovel200795–1998096432836 (84)EE1AEE1AEE1AOverlaps ORF-L
E44, CORF-L IE-likeNovel201282–195226652,01852 (11)EE1EE1EE1Transcriptional regulator
TRPalindrome202971–20301445-bp hairpin
TRPackaging motifs205612–205665f
TRPackaging motifs205784–205896g
  • a Fucosyl transferase 9 = EC 2.4.1.152.

  • b Acetylglucosamine transferase 1 = EC 2.4.1.1.

  • c UDP-β-Gal N-acetylglucosamine transferase 3, also known as O-linked N-acetylglucosamine transferase = EC 2.4.1.255.

  • d Complex dyad symmetry. Resemblance to alphaherpesvirus Ori-L and Ori-S as well as HHV6 Ori-Lyt, but not to cytomegalovirus Ori-Lyt, much larger than EEHV1 and EEHV5 versions, 3× 90-bp and other dyad symmetry elements with 5× OBP-binding site motifs plus 35× 20-bp AT-rich tandem repeats.

  • e No matches to other betaherpesvirus vGPCRs.

  • f 83% DNA match over 54 bp to terminal repeat motifs at 2852 to 2905 and 180311 to 180358 in EEHV1B(Emelia).

  • g 72% DNA match over 112 bp to terminal repeat motifs present in all three copies of the “a” sequence of HSV-1(KOS).

  • h The six clusters of genes or exons with unusually low GC content are shown in bold.

  • i Abbreviations: TR, tandem repeat; N-term, N terminal; dom, domain; memb, membrane; C term, C terminal; esp, especially; cons, conservation; SS, single stranded; UDG, uracil DNA glycosylase; Frag, fragmented; F, forward strand; C, complementary strand.