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, F 7xTME3fam2061–36086951525 (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, F 7xTMNovel8015–929558426NilNilNilNilUnique 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, F Novel 15171–15653 44 160 Nil Nil Nil Nil Unique to EEHV4
E4B, F Novel15712–1657856288NilNilNilNilUnique to EEHV4
E4C, F Novel16779–1723454151NilNilNilNilUnique to EEHV4
E6A, CE27ex1Novel17811–1739857137E27EE20NilEE2035% (44%) match to E27
E6B, F Novel17978–182564992NilNilNilNilUnique to EEHV4
E6, C 7xTME6fam19679–188556027433 (89)EE43EE43EE43
Nil, CvCXCL2?E7ANilNilNilAbsent in EEHV1B/4/5
E7, C 7xTME6fam20663–199565523532 (84)EE42EE42EE42
E7B, C 7xTM Novel 21647–20871 26 258 Nil Nil Nil Nil Unique to EEHV4
Nil 7xTM E6fam E8 EE41 EE41 EE41 Absent in EEHV4
E9, C 7xTM E6fam 22653–21757 27 298 34 (18) EE40 Frag EE40 Match to C-term EE40 only (esp EEHV5)
E9A, F vOGT AcTransf Novel 22993–24237 42 414 Nil Nil Nil Nil Unique to EEHV4 c
E9B, C (Truncated) Novel 24664–24281 23 127 Nil Nil Nil EE39/40 Match to EEHV5 N-term16-aa EE39/40 only
E9C, C Novel 25233–24718 27 171 Nil Nil Nil Nil Unique to EEHV4
E10A, C 7xTME6fam26078–2519761 NilNilNil25 (46)Matches central EE40(EEHV5) only
Nil7xTME6famE10EE39EE39EE39Absent in EEHV4
E11, C 7xTME6fam27127–263636125436 (96)EE38EE38EE38
E12, C 7xTME6fam28314–274576028534 (77)EE37EE37EE37
E12A, C Novel28547–282816088NilNilNilNilUnique to EEHV4
E13, C 7xTME6fam29780–289655827152 (88)EE36EE36EE36
E14.1, C 7xTME14fam31024–302155926927 (91)NilNilNilDuplication of E14
E14.2, C 7xTME14fam32191–312805230324 (77)NilNilNilDuplication of E14
E14, C 7xTME14fam33228–324195726926 (79)EE35EE35EE35
E15, C vGPCR47xTME15fam34435–334255833634 (86)EE34EE34EE3426% (49%) match to Lox C-5-C
E16, C 7xTME14fam35571–347685726740 (97)EE33EE33EE33
NilNovelE16CNilNilNilConserved in EEHV1 and EEHV5
NilNovelE16A/BNilNilNilSpliced; unique to EEHV1A/B
E16D, C vECTL Novel 36141–35584 45 185 Nil Nil Nil Nil
E17, F E27ex1 Novel 36491–36826 45 111 37 (99) EE32 EE32 EE32 Match to E27, 30% (50%)
E17A, F Novel36944–3725263102NilNilNilNilUnique to EEHV4
NilNovelNilNilNilEE32AUnique to EEHV5
E18, F 7xTME18fam37215–380185726732 (70)EE31EE31EE31Related to E28 by 30% (51%)
NilNovelE18BEE30AEE30AEE30AAbsent in EEHV4
NilNovelE18AEE30EE30EE30Absent in EEHV4
E18C, F Novel38433–387205994NilNilNilNilUnique to EEHV4
E19, FORF-F2U54.5fam39303–410426857952 (89)EE29EE29EE2925% (77%) to ORF-F1
E20, CvGPCR4A7xTME15fam43188–421545734446 (84)EE28EE28EE2823% (67%) match to Lox RAIP3
E20B, C Novel45323–4490465139NilNilNilNilUnique to EEHV4
E20A, F Novel45338–456646110843 (33)EE27EE27EE27
E21, CvGPCR4B7xTME15fam46925–458105937135 (76)EE26EE26EE2627% (35%) match to Lox RAIP3
E22, F Novel47646–47921499148 (96)EE25EE25EE25
E22A, F Novel48621–48730537946 (48)EE24EE24EE24
E23B, C Novel48993–4932557110NilNilNilNilUnique 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, F 7xTME18fam53115–538615424844 (90)EE19EE19EE19Related to E18 by 30% (51%)
E29, F 7xTMNovel54092–547815522942 (91)EE18EE18EE18
E30, C Novel55453–5491155180<15EE17EE17EE17Acidic similarity only
E30A, C E30Aex2 Novel 55883–55534 42 133 Nil Nil Nil Nil Unique to EEHV4
E30Aex1 Novel 56146–56095 40 Nil Nil Nil Nil Unique to EEHV4
NilE31EE16EE16EE16Absent in EEHV4
E31A, C Novel56923–563215620035 (46)EE15EE15EE15Only N-term cons
E31B, C Novel57113–56610NilNilNilNilUnique to EEHV4
E31C, CE31CexNovel57646–571826113665 (12)EE14EE14EE14No ATG, splice to E32?
E32, CU14.5βδ?60473–577176091845 (82)EE13EE13EE13
Nil, F E33EE12AFragNilUnique 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, FMTAex187495–87627 65 (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, C Core108154–1065386353869 (89)U36U36U36
U35, F Core108266–108553489569 (98)U35U35U35
U34, F Core108717–1096195330063 (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, F gN Core 131800–132105 49 101 63 (53) U46 U46 U46 Env glycoprotein N
U47, C gO (ORF-D) Βδ 132837–132187 52 216 34 (94) U47 U47 U47 Env glycoprotein O
U48, C gH Core 135086–132816 49 756 47 (96) U48 U48 U48 Env glycoprotein H
U48.5, CTK (ORF-E)αγδ136101–1350525634950 (87)EE7EE7EE7Thymidine kinase
U49, F Core136100–1368015723348 (85)U49U49U49
U50, FPAC2Core136620–1383475757564 (99)U50U50U50Packaging
U51, FvGPCR17xTMβδ138430–1396475640542 (95)U51U51U51e
U52, C Core140605–1398325325765 (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, F βγδ152794–1541526245248 (79)U59U59U59
U60, CTERex3Core155504–1543775766092 (99)U60U60U60Terminase subunit 1
U62, F βγδ155733–156005549057 (97)U62U62U62
U63, F βγδ155944–1565465120067 (72)U63U63U63
U64, FPAC1Core156527–1585526454148 (63)U64U64U64Packaging
U65, F Core158055–1590715933848 (98)U65U65U65
U66, CTERex2Novel159272–15915390 (100)U66U66U66Terminase subunit 1
TERex1Core160224–1594905389 (99)U66U66U66Terminase subunit 1
U67, F βγδ160612–1617425837661 (98)U67U67U67
U68, F Core161739–1621045112166 (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, C Core174625–1738136327054 (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, C ORF-Oex3 Novel 186570–186007 41 709 37 (95) EE5 EE5 EE5
ORF-Oex2 Novel 186897–186677 40 <15 EE5 EE5 EE5
ORF-Oex1 Novel 188411–187067 41 <15 EE5 EE5 EE5
Nil ORF-Pex2 Novel E38 EE4 EE4 EE4 Absent in EEHV4
ORF-Pex1 Novel E38 EE4 EE4 EE4 Absent in EEHV4
Nil ORF-Qex2 Novel E39 EE3 EE3 Nil Absent in EEHV2, -4, -5
ORF-Qex1 Novel E39 EE3 EE3 Nil Absent in EEHV2, -4, -5
E39A, C ORF-Rex2 Novel 189482–188506 40 367 Nil Nil Nil Nil Unique to EEHV4
ORF-Rex1 Novel 189690–189564 32 Nil Nil Nil Nil Unique 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.