id sid tid token lemma pos cord-048487-kamvao4b 1 1 key key JJ cord-048487-kamvao4b 2 1 : : : cord-048487-kamvao4b 2 2 cord-048487-kamvao4b cord-048487-kamvao4b NN cord-048487-kamvao4b 2 3 authors author NNS cord-048487-kamvao4b 2 4 : : : cord-048487-kamvao4b 2 5 O'Brien O'Brien NNP cord-048487-kamvao4b 2 6 , , , cord-048487-kamvao4b 2 7 John John NNP cord-048487-kamvao4b 2 8 D D NNP cord-048487-kamvao4b 2 9 ; ; : cord-048487-kamvao4b 2 10 She -PRON- PRP cord-048487-kamvao4b 2 11 , , , cord-048487-kamvao4b 2 12 Zhen Zhen NNP cord-048487-kamvao4b 2 13 - - HYPH cord-048487-kamvao4b 2 14 Su Su NNP cord-048487-kamvao4b 2 15 ; ; : cord-048487-kamvao4b 2 16 Suchard Suchard NNP cord-048487-kamvao4b 2 17 , , , cord-048487-kamvao4b 2 18 Marc Marc NNP cord-048487-kamvao4b 2 19 A a DT cord-048487-kamvao4b 2 20 title title NN cord-048487-kamvao4b 2 21 : : : cord-048487-kamvao4b 2 22 Dating date VBG cord-048487-kamvao4b 2 23 the the DT cord-048487-kamvao4b 2 24 time time NN cord-048487-kamvao4b 2 25 of of IN cord-048487-kamvao4b 2 26 viral viral JJ cord-048487-kamvao4b 2 27 subtype subtype NN cord-048487-kamvao4b 2 28 divergence divergence NN cord-048487-kamvao4b 2 29 date date NN cord-048487-kamvao4b 2 30 : : : cord-048487-kamvao4b 2 31 2008 2008 CD cord-048487-kamvao4b 2 32 - - SYM cord-048487-kamvao4b 2 33 06 06 CD cord-048487-kamvao4b 2 34 - - HYPH cord-048487-kamvao4b 2 35 09 09 CD cord-048487-kamvao4b 2 36 journal journal NN cord-048487-kamvao4b 2 37 : : : cord-048487-kamvao4b 3 1 BMC BMC NNP cord-048487-kamvao4b 3 2 Evol Evol NNP cord-048487-kamvao4b 3 3 Biol Biol NNP cord-048487-kamvao4b 3 4 DOI DOI NNP cord-048487-kamvao4b 3 5 : : : cord-048487-kamvao4b 3 6 10.1186/1471 10.1186/1471 CD cord-048487-kamvao4b 3 7 - - HYPH cord-048487-kamvao4b 3 8 2148 2148 CD cord-048487-kamvao4b 3 9 - - HYPH cord-048487-kamvao4b 3 10 8 8 CD cord-048487-kamvao4b 3 11 - - SYM cord-048487-kamvao4b 3 12 172 172 CD cord-048487-kamvao4b 3 13 sha sha NNP cord-048487-kamvao4b 3 14 : : : cord-048487-kamvao4b 3 15 341a9a57e865081e81a9c60b97802773761ce8c4 341a9a57e865081e81a9c60b97802773761ce8c4 CD cord-048487-kamvao4b 3 16 doc_id doc_id CD cord-048487-kamvao4b 3 17 : : : cord-048487-kamvao4b 3 18 48487 48487 CD cord-048487-kamvao4b 3 19 cord_uid cord_uid NNS cord-048487-kamvao4b 3 20 : : : cord-048487-kamvao4b 4 1 kamvao4b kamvao4b NNS cord-048487-kamvao4b 5 1 Precise precise JJ cord-048487-kamvao4b 5 2 dating dating NN cord-048487-kamvao4b 5 3 of of IN cord-048487-kamvao4b 5 4 viral viral JJ cord-048487-kamvao4b 5 5 subtype subtype NN cord-048487-kamvao4b 5 6 divergence divergence NN cord-048487-kamvao4b 5 7 enables enable VBZ cord-048487-kamvao4b 5 8 researchers researcher NNS cord-048487-kamvao4b 5 9 to to TO cord-048487-kamvao4b 5 10 correlate correlate VB cord-048487-kamvao4b 5 11 divergence divergence NN cord-048487-kamvao4b 5 12 with with IN cord-048487-kamvao4b 5 13 geographic geographic JJ cord-048487-kamvao4b 5 14 and and CC cord-048487-kamvao4b 5 15 demographic demographic JJ cord-048487-kamvao4b 5 16 occurrences occurrence NNS cord-048487-kamvao4b 5 17 . . . cord-048487-kamvao4b 6 1 When when WRB cord-048487-kamvao4b 6 2 historical historical JJ cord-048487-kamvao4b 6 3 data datum NNS cord-048487-kamvao4b 6 4 are be VBP cord-048487-kamvao4b 6 5 absent absent JJ cord-048487-kamvao4b 6 6 ( ( -LRB- cord-048487-kamvao4b 6 7 that that RB cord-048487-kamvao4b 6 8 is is RB cord-048487-kamvao4b 6 9 , , , cord-048487-kamvao4b 6 10 the the DT cord-048487-kamvao4b 6 11 overwhelming overwhelming JJ cord-048487-kamvao4b 6 12 majority majority NN cord-048487-kamvao4b 6 13 ) ) -RRB- cord-048487-kamvao4b 6 14 , , , cord-048487-kamvao4b 6 15 viral viral JJ cord-048487-kamvao4b 6 16 sequence sequence NN cord-048487-kamvao4b 6 17 sampling sampling NN cord-048487-kamvao4b 6 18 on on IN cord-048487-kamvao4b 6 19 a a DT cord-048487-kamvao4b 6 20 time time NN cord-048487-kamvao4b 6 21 scale scale NN cord-048487-kamvao4b 6 22 commensurate commensurate NN cord-048487-kamvao4b 6 23 with with IN cord-048487-kamvao4b 6 24 the the DT cord-048487-kamvao4b 6 25 rate rate NN cord-048487-kamvao4b 6 26 of of IN cord-048487-kamvao4b 6 27 substitution substitution NN cord-048487-kamvao4b 6 28 permits permit VBZ cord-048487-kamvao4b 6 29 the the DT cord-048487-kamvao4b 6 30 inference inference NN cord-048487-kamvao4b 6 31 of of IN cord-048487-kamvao4b 6 32 the the DT cord-048487-kamvao4b 6 33 times time NNS cord-048487-kamvao4b 6 34 of of IN cord-048487-kamvao4b 6 35 subtype subtype NN cord-048487-kamvao4b 6 36 divergence divergence NN cord-048487-kamvao4b 6 37 . . . cord-048487-kamvao4b 7 1 Currently currently RB cord-048487-kamvao4b 7 2 , , , cord-048487-kamvao4b 7 3 researchers researcher NNS cord-048487-kamvao4b 7 4 use use VBP cord-048487-kamvao4b 7 5 two two CD cord-048487-kamvao4b 7 6 strategies strategy NNS cord-048487-kamvao4b 7 7 to to TO cord-048487-kamvao4b 7 8 approach approach VB cord-048487-kamvao4b 7 9 this this DT cord-048487-kamvao4b 7 10 task task NN cord-048487-kamvao4b 7 11 , , , cord-048487-kamvao4b 7 12 both both DT cord-048487-kamvao4b 7 13 requiring require VBG cord-048487-kamvao4b 7 14 strong strong JJ cord-048487-kamvao4b 7 15 conditions condition NNS cord-048487-kamvao4b 7 16 on on IN cord-048487-kamvao4b 7 17 the the DT cord-048487-kamvao4b 7 18 molecular molecular JJ cord-048487-kamvao4b 7 19 clock clock NN cord-048487-kamvao4b 7 20 assumption assumption NN cord-048487-kamvao4b 7 21 of of IN cord-048487-kamvao4b 7 22 substitution substitution NN cord-048487-kamvao4b 7 23 rate rate NN cord-048487-kamvao4b 7 24 . . . cord-048487-kamvao4b 8 1 As as IN cord-048487-kamvao4b 8 2 the the DT cord-048487-kamvao4b 8 3 underlying underlying JJ cord-048487-kamvao4b 8 4 structure structure NN cord-048487-kamvao4b 8 5 of of IN cord-048487-kamvao4b 8 6 the the DT cord-048487-kamvao4b 8 7 substitution substitution NN cord-048487-kamvao4b 8 8 rate rate NN cord-048487-kamvao4b 8 9 process process NN cord-048487-kamvao4b 8 10 at at IN cord-048487-kamvao4b 8 11 the the DT cord-048487-kamvao4b 8 12 time time NN cord-048487-kamvao4b 8 13 of of IN cord-048487-kamvao4b 8 14 subtype subtype NN cord-048487-kamvao4b 8 15 divergence divergence NN cord-048487-kamvao4b 8 16 is be VBZ cord-048487-kamvao4b 8 17 not not RB cord-048487-kamvao4b 8 18 understood understand VBN cord-048487-kamvao4b 8 19 and and CC cord-048487-kamvao4b 8 20 likely likely RB cord-048487-kamvao4b 8 21 highly highly RB cord-048487-kamvao4b 8 22 variable variable JJ cord-048487-kamvao4b 8 23 , , , cord-048487-kamvao4b 8 24 we -PRON- PRP cord-048487-kamvao4b 8 25 present present VBP cord-048487-kamvao4b 8 26 a a DT cord-048487-kamvao4b 8 27 simple simple JJ cord-048487-kamvao4b 8 28 method method NN cord-048487-kamvao4b 8 29 that that WDT cord-048487-kamvao4b 8 30 estimates estimate VBZ cord-048487-kamvao4b 8 31 rates rate NNS cord-048487-kamvao4b 8 32 of of IN cord-048487-kamvao4b 8 33 substitution substitution NN cord-048487-kamvao4b 8 34 , , , cord-048487-kamvao4b 8 35 and and CC cord-048487-kamvao4b 8 36 from from IN cord-048487-kamvao4b 8 37 there there RB cord-048487-kamvao4b 8 38 , , , cord-048487-kamvao4b 8 39 times time NNS cord-048487-kamvao4b 8 40 of of IN cord-048487-kamvao4b 8 41 divergence divergence NN cord-048487-kamvao4b 8 42 , , , cord-048487-kamvao4b 8 43 without without IN cord-048487-kamvao4b 8 44 use use NN cord-048487-kamvao4b 8 45 of of IN cord-048487-kamvao4b 8 46 an an DT cord-048487-kamvao4b 8 47 assumed assumed JJ cord-048487-kamvao4b 8 48 molecular molecular JJ cord-048487-kamvao4b 8 49 clock clock NN cord-048487-kamvao4b 8 50 . . . cord-048487-kamvao4b 9 1 We -PRON- PRP cord-048487-kamvao4b 9 2 accomplish accomplish VBP cord-048487-kamvao4b 9 3 this this DT cord-048487-kamvao4b 9 4 by by IN cord-048487-kamvao4b 9 5 blending blend VBG cord-048487-kamvao4b 9 6 estimates estimate NNS cord-048487-kamvao4b 9 7 of of IN cord-048487-kamvao4b 9 8 the the DT cord-048487-kamvao4b 9 9 substitution substitution NN cord-048487-kamvao4b 9 10 rate rate NN cord-048487-kamvao4b 9 11 for for IN cord-048487-kamvao4b 9 12 triplets triplet NNS cord-048487-kamvao4b 9 13 of of IN cord-048487-kamvao4b 9 14 dated date VBN cord-048487-kamvao4b 9 15 sequences sequence NNS cord-048487-kamvao4b 9 16 where where WRB cord-048487-kamvao4b 9 17 each each DT cord-048487-kamvao4b 9 18 sequence sequence NN cord-048487-kamvao4b 9 19 draws draw VBZ cord-048487-kamvao4b 9 20 from from IN cord-048487-kamvao4b 9 21 a a DT cord-048487-kamvao4b 9 22 distinct distinct JJ cord-048487-kamvao4b 9 23 viral viral JJ cord-048487-kamvao4b 9 24 subtype subtype NN cord-048487-kamvao4b 9 25 , , , cord-048487-kamvao4b 9 26 providing provide VBG cord-048487-kamvao4b 9 27 a a DT cord-048487-kamvao4b 9 28 zeroth zeroth JJ cord-048487-kamvao4b 9 29 - - HYPH cord-048487-kamvao4b 9 30 order order NN cord-048487-kamvao4b 9 31 approximation approximation NN cord-048487-kamvao4b 9 32 for for IN cord-048487-kamvao4b 9 33 the the DT cord-048487-kamvao4b 9 34 rate rate NN cord-048487-kamvao4b 9 35 between between IN cord-048487-kamvao4b 9 36 subtypes subtype NNS cord-048487-kamvao4b 9 37 . . . cord-048487-kamvao4b 10 1 As as IN cord-048487-kamvao4b 10 2 an an DT cord-048487-kamvao4b 10 3 example example NN cord-048487-kamvao4b 10 4 , , , cord-048487-kamvao4b 10 5 we -PRON- PRP cord-048487-kamvao4b 10 6 calculate calculate VBP cord-048487-kamvao4b 10 7 the the DT cord-048487-kamvao4b 10 8 time time NN cord-048487-kamvao4b 10 9 of of IN cord-048487-kamvao4b 10 10 divergence divergence NN cord-048487-kamvao4b 10 11 for for IN cord-048487-kamvao4b 10 12 three three CD cord-048487-kamvao4b 10 13 genes gene NNS cord-048487-kamvao4b 10 14 among among IN cord-048487-kamvao4b 10 15 influenza influenza NN cord-048487-kamvao4b 10 16 subtypes subtype NNS cord-048487-kamvao4b 10 17 A A NNP cord-048487-kamvao4b 10 18 - - : cord-048487-kamvao4b 10 19 H3N2 H3N2 NNP cord-048487-kamvao4b 10 20 and and CC cord-048487-kamvao4b 10 21 B b NN cord-048487-kamvao4b 10 22 using use VBG cord-048487-kamvao4b 10 23 subtype subtype NN cord-048487-kamvao4b 10 24 C c NN cord-048487-kamvao4b 10 25 as as IN cord-048487-kamvao4b 10 26 an an DT cord-048487-kamvao4b 10 27 outgroup outgroup NN cord-048487-kamvao4b 10 28 . . . cord-048487-kamvao4b 11 1 We -PRON- PRP cord-048487-kamvao4b 11 2 show show VBP cord-048487-kamvao4b 11 3 a a DT cord-048487-kamvao4b 11 4 time time NN cord-048487-kamvao4b 11 5 of of IN cord-048487-kamvao4b 11 6 divergence divergence NN cord-048487-kamvao4b 11 7 approximately approximately RB cord-048487-kamvao4b 11 8 100 100 CD cord-048487-kamvao4b 11 9 years year NNS cord-048487-kamvao4b 11 10 ago ago RB cord-048487-kamvao4b 11 11 , , , cord-048487-kamvao4b 11 12 substantially substantially RB cord-048487-kamvao4b 11 13 more more RBR cord-048487-kamvao4b 11 14 recent recent JJ cord-048487-kamvao4b 11 15 than than IN cord-048487-kamvao4b 11 16 previous previous JJ cord-048487-kamvao4b 11 17 estimates estimate NNS cord-048487-kamvao4b 11 18 which which WDT cord-048487-kamvao4b 11 19 range range VBP cord-048487-kamvao4b 11 20 from from IN cord-048487-kamvao4b 11 21 250 250 CD cord-048487-kamvao4b 11 22 to to TO cord-048487-kamvao4b 11 23 3800 3800 CD cord-048487-kamvao4b 11 24 years year NNS cord-048487-kamvao4b 11 25 ago ago RB cord-048487-kamvao4b 11 26 . . . cord-048487-kamvao4b 12 1 Precise precise JJ cord-048487-kamvao4b 12 2 estimates estimate NNS cord-048487-kamvao4b 12 3 are be VBP cord-048487-kamvao4b 12 4 sorely sorely RB cord-048487-kamvao4b 12 5 lacking lack VBG cord-048487-kamvao4b 12 6 for for IN cord-048487-kamvao4b 12 7 dating date VBG cord-048487-kamvao4b 12 8 the the DT cord-048487-kamvao4b 12 9 emergence emergence NN cord-048487-kamvao4b 12 10 and and CC cord-048487-kamvao4b 12 11 divergence divergence NN cord-048487-kamvao4b 12 12 of of IN cord-048487-kamvao4b 12 13 viral viral JJ cord-048487-kamvao4b 12 14 subtypes subtype NNS cord-048487-kamvao4b 12 15 . . . cord-048487-kamvao4b 13 1 Improved improve VBN cord-048487-kamvao4b 13 2 estimates estimate NNS cord-048487-kamvao4b 13 3 equip equip VBP cord-048487-kamvao4b 13 4 epidemiologists epidemiologist NNS cord-048487-kamvao4b 13 5 and and CC cord-048487-kamvao4b 13 6 virologists virologist NNS cord-048487-kamvao4b 13 7 to to TO cord-048487-kamvao4b 13 8 begin begin VB cord-048487-kamvao4b 13 9 to to TO cord-048487-kamvao4b 13 10 correlate correlate VB cord-048487-kamvao4b 13 11 these these DT cord-048487-kamvao4b 13 12 important important JJ cord-048487-kamvao4b 13 13 establishing establish VBG cord-048487-kamvao4b 13 14 events event NNS cord-048487-kamvao4b 13 15 with with IN cord-048487-kamvao4b 13 16 historical historical JJ cord-048487-kamvao4b 13 17 demographic demographic JJ cord-048487-kamvao4b 13 18 changes change NNS cord-048487-kamvao4b 13 19 , , , cord-048487-kamvao4b 13 20 geographical geographical JJ cord-048487-kamvao4b 13 21 invasions invasion NNS cord-048487-kamvao4b 13 22 and and CC cord-048487-kamvao4b 13 23 zoonoses zoonosis NNS cord-048487-kamvao4b 13 24 , , , cord-048487-kamvao4b 13 25 the the DT cord-048487-kamvao4b 13 26 transferring transferring NN cord-048487-kamvao4b 13 27 of of IN cord-048487-kamvao4b 13 28 a a DT cord-048487-kamvao4b 13 29 virus virus NN cord-048487-kamvao4b 13 30 from from IN cord-048487-kamvao4b 13 31 one one CD cord-048487-kamvao4b 13 32 host host NN cord-048487-kamvao4b 13 33 species specie NNS cord-048487-kamvao4b 13 34 to to IN cord-048487-kamvao4b 13 35 another another DT cord-048487-kamvao4b 13 36 [ [ -LRB- cord-048487-kamvao4b 13 37 7 7 CD cord-048487-kamvao4b 13 38 , , , cord-048487-kamvao4b 13 39 1 1 CD cord-048487-kamvao4b 13 40 , , , cord-048487-kamvao4b 13 41 25 25 CD cord-048487-kamvao4b 13 42 ] ] -RRB- cord-048487-kamvao4b 13 43 . . . cord-048487-kamvao4b 14 1 For for IN cord-048487-kamvao4b 14 2 example example NN cord-048487-kamvao4b 14 3 , , , cord-048487-kamvao4b 14 4 archeological archeological JJ cord-048487-kamvao4b 14 5 sequence sequence NN cord-048487-kamvao4b 14 6 data datum NNS cord-048487-kamvao4b 14 7 can can MD cord-048487-kamvao4b 14 8 furnish furnish VB cord-048487-kamvao4b 14 9 accurate accurate JJ cord-048487-kamvao4b 14 10 dates date NNS cord-048487-kamvao4b 14 11 and and CC cord-048487-kamvao4b 14 12 show show VBP cord-048487-kamvao4b 14 13 that that IN cord-048487-kamvao4b 14 14 substantial substantial JJ cord-048487-kamvao4b 14 15 genomic genomic JJ cord-048487-kamvao4b 14 16 changes change NNS cord-048487-kamvao4b 14 17 associate associate VBP cord-048487-kamvao4b 14 18 with with IN cord-048487-kamvao4b 14 19 geographical geographical JJ cord-048487-kamvao4b 14 20 invasion invasion NN cord-048487-kamvao4b 14 21 and and CC cord-048487-kamvao4b 14 22 zoonosis zoonosis NN cord-048487-kamvao4b 14 23 [ [ -LRB- cord-048487-kamvao4b 14 24 14 14 CD cord-048487-kamvao4b 14 25 , , , cord-048487-kamvao4b 14 26 17 17 CD cord-048487-kamvao4b 14 27 ] ] -RRB- cord-048487-kamvao4b 14 28 . . . cord-048487-kamvao4b 15 1 Further further RB cord-048487-kamvao4b 15 2 , , , cord-048487-kamvao4b 15 3 the the DT cord-048487-kamvao4b 15 4 recent recent JJ cord-048487-kamvao4b 15 5 availability availability NN cord-048487-kamvao4b 15 6 of of IN cord-048487-kamvao4b 15 7 viral viral JJ cord-048487-kamvao4b 15 8 gene gene NN cord-048487-kamvao4b 15 9 sequences sequence NNS cord-048487-kamvao4b 15 10 sampled sample VBN cord-048487-kamvao4b 15 11 at at IN cord-048487-kamvao4b 15 12 a a DT cord-048487-kamvao4b 15 13 pace pace NN cord-048487-kamvao4b 15 14 commensurate commensurate NN cord-048487-kamvao4b 15 15 with with IN cord-048487-kamvao4b 15 16 their -PRON- PRP$ cord-048487-kamvao4b 15 17 rate rate NN cord-048487-kamvao4b 15 18 of of IN cord-048487-kamvao4b 15 19 nucleotide nucleotide JJ cord-048487-kamvao4b 15 20 substitution substitution NN cord-048487-kamvao4b 15 21 vastly vastly RB cord-048487-kamvao4b 15 22 augments augment VBZ cord-048487-kamvao4b 15 23 the the DT cord-048487-kamvao4b 15 24 ability ability NN cord-048487-kamvao4b 15 25 to to TO cord-048487-kamvao4b 15 26 rigorously rigorously RB cord-048487-kamvao4b 15 27 infer infer VB cord-048487-kamvao4b 15 28 the the DT cord-048487-kamvao4b 15 29 time time NN cord-048487-kamvao4b 15 30 scale scale NN cord-048487-kamvao4b 15 31 of of IN cord-048487-kamvao4b 15 32 phylogenies phylogeny NNS cord-048487-kamvao4b 15 33 and and CC cord-048487-kamvao4b 15 34 hence hence RB cord-048487-kamvao4b 15 35 determine determine VB cord-048487-kamvao4b 15 36 the the DT cord-048487-kamvao4b 15 37 time time NN cord-048487-kamvao4b 15 38 of of IN cord-048487-kamvao4b 15 39 the the DT cord-048487-kamvao4b 15 40 most most RBS cord-048487-kamvao4b 15 41 recent recent JJ cord-048487-kamvao4b 15 42 common common JJ cord-048487-kamvao4b 15 43 ancestor ancestor NN cord-048487-kamvao4b 15 44 ( ( -LRB- cord-048487-kamvao4b 15 45 TMRCA TMRCA NNP cord-048487-kamvao4b 15 46 ) ) -RRB- cord-048487-kamvao4b 15 47 for for IN cord-048487-kamvao4b 15 48 different different JJ cord-048487-kamvao4b 15 49 viral viral JJ cord-048487-kamvao4b 15 50 types type NNS cord-048487-kamvao4b 15 51 [ [ -LRB- cord-048487-kamvao4b 15 52 18 18 CD cord-048487-kamvao4b 15 53 , , , cord-048487-kamvao4b 15 54 26 26 CD cord-048487-kamvao4b 15 55 , , , cord-048487-kamvao4b 15 56 6 6 CD cord-048487-kamvao4b 15 57 ] ] -RRB- cord-048487-kamvao4b 15 58 . . . cord-048487-kamvao4b 16 1 Systematic systematic JJ cord-048487-kamvao4b 16 2 studies study NNS cord-048487-kamvao4b 16 3 characterize characterize VBP cord-048487-kamvao4b 16 4 the the DT cord-048487-kamvao4b 16 5 substitution substitution NN cord-048487-kamvao4b 16 6 process process NN cord-048487-kamvao4b 16 7 and and CC cord-048487-kamvao4b 16 8 substitution substitution NN cord-048487-kamvao4b 16 9 rate rate NN cord-048487-kamvao4b 16 10 process process NN cord-048487-kamvao4b 16 11 of of IN cord-048487-kamvao4b 16 12 several several JJ cord-048487-kamvao4b 16 13 classes class NNS cord-048487-kamvao4b 16 14 of of IN cord-048487-kamvao4b 16 15 viral viral JJ cord-048487-kamvao4b 16 16 subtypes subtype NNS cord-048487-kamvao4b 16 17 in in IN cord-048487-kamvao4b 16 18 , , , cord-048487-kamvao4b 16 19 for for IN cord-048487-kamvao4b 16 20 example example NN cord-048487-kamvao4b 16 21 , , , cord-048487-kamvao4b 16 22 Dengue dengue NN cord-048487-kamvao4b 16 23 , , , cord-048487-kamvao4b 16 24 influenza influenza NN cord-048487-kamvao4b 16 25 subtype subtype NN cord-048487-kamvao4b 16 26 A A NNP cord-048487-kamvao4b 16 27 , , , cord-048487-kamvao4b 16 28 human human JJ cord-048487-kamvao4b 16 29 immunodeficiency immunodeficiency NN cord-048487-kamvao4b 16 30 virus virus NN cord-048487-kamvao4b 16 31 ( ( -LRB- cord-048487-kamvao4b 16 32 HIV HIV NNP cord-048487-kamvao4b 16 33 ) ) -RRB- cord-048487-kamvao4b 16 34 and and CC cord-048487-kamvao4b 16 35 the the DT cord-048487-kamvao4b 16 36 virus virus NN cord-048487-kamvao4b 16 37 responsible responsible JJ cord-048487-kamvao4b 16 38 for for IN cord-048487-kamvao4b 16 39 sudden sudden JJ cord-048487-kamvao4b 16 40 acute acute JJ cord-048487-kamvao4b 16 41 respiratory respiratory JJ cord-048487-kamvao4b 16 42 syndrome syndrome NN cord-048487-kamvao4b 16 43 ( ( -LRB- cord-048487-kamvao4b 16 44 SARS SARS NNP cord-048487-kamvao4b 16 45 ) ) -RRB- cord-048487-kamvao4b 16 46 . . . cord-048487-kamvao4b 17 1 For for IN cord-048487-kamvao4b 17 2 the the DT cord-048487-kamvao4b 17 3 last last JJ cord-048487-kamvao4b 17 4 three three CD cord-048487-kamvao4b 17 5 viruses virus NNS cord-048487-kamvao4b 17 6 , , , cord-048487-kamvao4b 17 7 a a DT cord-048487-kamvao4b 17 8 unique unique JJ cord-048487-kamvao4b 17 9 zoonotic zoonotic JJ cord-048487-kamvao4b 17 10 transfer transfer NN cord-048487-kamvao4b 17 11 appears appear VBZ cord-048487-kamvao4b 17 12 to to TO cord-048487-kamvao4b 17 13 co co VB cord-048487-kamvao4b 17 14 - - VB cord-048487-kamvao4b 17 15 occur occur VB cord-048487-kamvao4b 17 16 with with IN cord-048487-kamvao4b 17 17 substantial substantial JJ cord-048487-kamvao4b 17 18 changes change NNS cord-048487-kamvao4b 17 19 in in IN cord-048487-kamvao4b 17 20 both both CC cord-048487-kamvao4b 17 21 the the DT cord-048487-kamvao4b 17 22 composition composition NN cord-048487-kamvao4b 17 23 of of IN cord-048487-kamvao4b 17 24 nucleotides nucleotide NNS cord-048487-kamvao4b 17 25 and and CC cord-048487-kamvao4b 17 26 amino amino JJ cord-048487-kamvao4b 17 27 acids acid NNS cord-048487-kamvao4b 17 28 as as RB cord-048487-kamvao4b 17 29 well well RB cord-048487-kamvao4b 17 30 as as IN cord-048487-kamvao4b 17 31 alterations alteration NNS cord-048487-kamvao4b 17 32 in in IN cord-048487-kamvao4b 17 33 the the DT cord-048487-kamvao4b 17 34 rate rate NN cord-048487-kamvao4b 17 35 of of IN cord-048487-kamvao4b 17 36 nucleotide nucleotide JJ cord-048487-kamvao4b 17 37 substitution substitution NN cord-048487-kamvao4b 17 38 [ [ -LRB- cord-048487-kamvao4b 17 39 15 15 CD cord-048487-kamvao4b 17 40 , , , cord-048487-kamvao4b 17 41 14 14 CD cord-048487-kamvao4b 17 42 , , , cord-048487-kamvao4b 17 43 1 1 CD cord-048487-kamvao4b 17 44 ] ] -RRB- cord-048487-kamvao4b 17 45 . . . cord-048487-kamvao4b 18 1 In in IN cord-048487-kamvao4b 18 2 Dengue Dengue NNP cord-048487-kamvao4b 18 3 , , , cord-048487-kamvao4b 18 4 where where WRB cord-048487-kamvao4b 18 5 a a DT cord-048487-kamvao4b 18 6 single single JJ cord-048487-kamvao4b 18 7 subtype subtype NN cord-048487-kamvao4b 18 8 simultaneously simultaneously RB cord-048487-kamvao4b 18 9 inhabits inhabit VBZ cord-048487-kamvao4b 18 10 two two CD cord-048487-kamvao4b 18 11 hosts host NNS cord-048487-kamvao4b 18 12 ( ( -LRB- cord-048487-kamvao4b 18 13 humans human NNS cord-048487-kamvao4b 18 14 and and CC cord-048487-kamvao4b 18 15 Aedes Aedes NNP cord-048487-kamvao4b 18 16 aegypti aegypti NNS cord-048487-kamvao4b 18 17 ) ) -RRB- cord-048487-kamvao4b 18 18 in in IN cord-048487-kamvao4b 18 19 a a DT cord-048487-kamvao4b 18 20 persistent persistent JJ cord-048487-kamvao4b 18 21 zoonotic zoonotic JJ cord-048487-kamvao4b 18 22 process process NN cord-048487-kamvao4b 18 23 , , , cord-048487-kamvao4b 18 24 the the DT cord-048487-kamvao4b 18 25 introduction introduction NN cord-048487-kamvao4b 18 26 of of IN cord-048487-kamvao4b 18 27 the the DT cord-048487-kamvao4b 18 28 virus virus NN cord-048487-kamvao4b 18 29 to to IN cord-048487-kamvao4b 18 30 new new JJ cord-048487-kamvao4b 18 31 geographical geographical JJ cord-048487-kamvao4b 18 32 environments environment NNS cord-048487-kamvao4b 18 33 associates associate NNS cord-048487-kamvao4b 18 34 with with IN cord-048487-kamvao4b 18 35 a a DT cord-048487-kamvao4b 18 36 dramatic dramatic JJ cord-048487-kamvao4b 18 37 increase increase NN cord-048487-kamvao4b 18 38 in in IN cord-048487-kamvao4b 18 39 sequence sequence NN cord-048487-kamvao4b 18 40 diversity diversity NN cord-048487-kamvao4b 18 41 [ [ -LRB- cord-048487-kamvao4b 18 42 25 25 CD cord-048487-kamvao4b 18 43 ] ] -RRB- cord-048487-kamvao4b 18 44 . . . cord-048487-kamvao4b 19 1 Unfortunately unfortunately RB cord-048487-kamvao4b 19 2 , , , cord-048487-kamvao4b 19 3 no no DT cord-048487-kamvao4b 19 4 studies study NNS cord-048487-kamvao4b 19 5 thus thus RB cord-048487-kamvao4b 19 6 far far RB cord-048487-kamvao4b 19 7 analyze analyze VBP cord-048487-kamvao4b 19 8 the the DT cord-048487-kamvao4b 19 9 rate rate NN cord-048487-kamvao4b 19 10 of of IN cord-048487-kamvao4b 19 11 nucleotide nucleotide JJ cord-048487-kamvao4b 19 12 substitution substitution NN cord-048487-kamvao4b 19 13 during during IN cord-048487-kamvao4b 19 14 either either DT cord-048487-kamvao4b 19 15 geographical geographical JJ cord-048487-kamvao4b 19 16 invasion invasion NN cord-048487-kamvao4b 19 17 or or CC cord-048487-kamvao4b 19 18 zoonosis zoonosis NN cord-048487-kamvao4b 19 19 . . . cord-048487-kamvao4b 20 1 Consequently consequently RB cord-048487-kamvao4b 20 2 , , , cord-048487-kamvao4b 20 3 studies study NNS cord-048487-kamvao4b 20 4 of of IN cord-048487-kamvao4b 20 5 the the DT cord-048487-kamvao4b 20 6 date date NN cord-048487-kamvao4b 20 7 of of IN cord-048487-kamvao4b 20 8 origins origin NNS cord-048487-kamvao4b 20 9 of of IN cord-048487-kamvao4b 20 10 viral viral JJ cord-048487-kamvao4b 20 11 subtypes subtype NNS cord-048487-kamvao4b 20 12 must must MD cord-048487-kamvao4b 20 13 use use VB cord-048487-kamvao4b 20 14 strong strong JJ cord-048487-kamvao4b 20 15 a a DT cord-048487-kamvao4b 20 16 priori priori FW cord-048487-kamvao4b 20 17 assumptions assumption NNS cord-048487-kamvao4b 20 18 on on IN cord-048487-kamvao4b 20 19 the the DT cord-048487-kamvao4b 20 20 rate rate NN cord-048487-kamvao4b 20 21 structure structure NN cord-048487-kamvao4b 20 22 of of IN cord-048487-kamvao4b 20 23 nucleotide nucleotide JJ cord-048487-kamvao4b 20 24 substitution substitution NN cord-048487-kamvao4b 20 25 . . . cord-048487-kamvao4b 21 1 Two two CD cord-048487-kamvao4b 21 2 primary primary JJ cord-048487-kamvao4b 21 3 methods method NNS cord-048487-kamvao4b 21 4 find find VBP cord-048487-kamvao4b 21 5 use use NN cord-048487-kamvao4b 21 6 to to IN cord-048487-kamvao4b 21 7 date date NN cord-048487-kamvao4b 21 8 the the DT cord-048487-kamvao4b 21 9 time time NN cord-048487-kamvao4b 21 10 of of IN cord-048487-kamvao4b 21 11 viral viral JJ cord-048487-kamvao4b 21 12 subtype subtype NN cord-048487-kamvao4b 21 13 divergence divergence NN cord-048487-kamvao4b 21 14 . . . cord-048487-kamvao4b 22 1 The the DT cord-048487-kamvao4b 22 2 most most RBS cord-048487-kamvao4b 22 3 commonly commonly RB cord-048487-kamvao4b 22 4 employed employ VBN cord-048487-kamvao4b 22 5 approach approach NN cord-048487-kamvao4b 22 6 determines determine VBZ cord-048487-kamvao4b 22 7 the the DT cord-048487-kamvao4b 22 8 divergence divergence NN cord-048487-kamvao4b 22 9 time time NN cord-048487-kamvao4b 22 10 of of IN cord-048487-kamvao4b 22 11 subtypes subtype NNS cord-048487-kamvao4b 22 12 using use VBG cord-048487-kamvao4b 22 13 a a DT cord-048487-kamvao4b 22 14 molecular molecular JJ cord-048487-kamvao4b 22 15 clock clock NN cord-048487-kamvao4b 22 16 assumption assumption NN cord-048487-kamvao4b 22 17 ( ( -LRB- cord-048487-kamvao4b 22 18 MCA MCA NNP cord-048487-kamvao4b 22 19 ) ) -RRB- cord-048487-kamvao4b 22 20 over over IN cord-048487-kamvao4b 22 21 an an DT cord-048487-kamvao4b 22 22 entire entire JJ cord-048487-kamvao4b 22 23 phylogeny phylogeny NN cord-048487-kamvao4b 22 24 [ [ -LRB- cord-048487-kamvao4b 22 25 18 18 CD cord-048487-kamvao4b 22 26 , , , cord-048487-kamvao4b 22 27 21 21 CD cord-048487-kamvao4b 22 28 , , , cord-048487-kamvao4b 22 29 5 5 CD cord-048487-kamvao4b 22 30 , , , cord-048487-kamvao4b 22 31 26 26 CD cord-048487-kamvao4b 22 32 ] ] -RRB- cord-048487-kamvao4b 22 33 . . . cord-048487-kamvao4b 23 1 In in IN cord-048487-kamvao4b 23 2 its -PRON- PRP$ cord-048487-kamvao4b 23 3 strict strict JJ cord-048487-kamvao4b 23 4 formulation formulation NN cord-048487-kamvao4b 23 5 , , , cord-048487-kamvao4b 23 6 the the DT cord-048487-kamvao4b 23 7 MCA MCA NNP cord-048487-kamvao4b 23 8 posits posit VBZ cord-048487-kamvao4b 23 9 a a DT cord-048487-kamvao4b 23 10 proportional proportional JJ cord-048487-kamvao4b 23 11 relation relation NN cord-048487-kamvao4b 23 12 between between IN cord-048487-kamvao4b 23 13 the the DT cord-048487-kamvao4b 23 14 number number NN cord-048487-kamvao4b 23 15 of of IN cord-048487-kamvao4b 23 16 substitutions substitution NNS cord-048487-kamvao4b 23 17 and and CC cord-048487-kamvao4b 23 18 the the DT cord-048487-kamvao4b 23 19 intervening intervene VBG cord-048487-kamvao4b 23 20 time time NN cord-048487-kamvao4b 23 21 period period NN cord-048487-kamvao4b 23 22 over over IN cord-048487-kamvao4b 23 23 the the DT cord-048487-kamvao4b 23 24 entire entire JJ cord-048487-kamvao4b 23 25 phylogeny phylogeny NN cord-048487-kamvao4b 23 26 . . . cord-048487-kamvao4b 24 1 Looser loose JJR cord-048487-kamvao4b 24 2 forms form NNS cord-048487-kamvao4b 24 3 of of IN cord-048487-kamvao4b 24 4 MCAs mca NNS cord-048487-kamvao4b 24 5 require require VBP cord-048487-kamvao4b 24 6 only only RB cord-048487-kamvao4b 24 7 that that IN cord-048487-kamvao4b 24 8 the the DT cord-048487-kamvao4b 24 9 proportionality proportionality NN cord-048487-kamvao4b 24 10 hold hold NN cord-048487-kamvao4b 24 11 along along IN cord-048487-kamvao4b 24 12 individual individual JJ cord-048487-kamvao4b 24 13 branches branch NNS cord-048487-kamvao4b 24 14 , , , cord-048487-kamvao4b 24 15 with with IN cord-048487-kamvao4b 24 16 the the DT cord-048487-kamvao4b 24 17 rates rate NNS cord-048487-kamvao4b 24 18 across across IN cord-048487-kamvao4b 24 19 branches branch NNS cord-048487-kamvao4b 24 20 drawn draw VBN cord-048487-kamvao4b 24 21 from from IN cord-048487-kamvao4b 24 22 a a DT cord-048487-kamvao4b 24 23 pre pre JJ cord-048487-kamvao4b 24 24 - - JJ cord-048487-kamvao4b 24 25 specified specified JJ cord-048487-kamvao4b 24 26 distribution distribution NN cord-048487-kamvao4b 24 27 [ [ -LRB- cord-048487-kamvao4b 24 28 5 5 CD cord-048487-kamvao4b 24 29 ] ] -RRB- cord-048487-kamvao4b 24 30 . . . cord-048487-kamvao4b 25 1 Committed commit VBN cord-048487-kamvao4b 25 2 to to IN cord-048487-kamvao4b 25 3 some some DT cord-048487-kamvao4b 25 4 variant variant NN cord-048487-kamvao4b 25 5 of of IN cord-048487-kamvao4b 25 6 the the DT cord-048487-kamvao4b 25 7 MCA MCA NNP cord-048487-kamvao4b 25 8 , , , cord-048487-kamvao4b 25 9 current current JJ cord-048487-kamvao4b 25 10 algorithms algorithm NNS cord-048487-kamvao4b 25 11 then then RB cord-048487-kamvao4b 25 12 estimate estimate VBP cord-048487-kamvao4b 25 13 the the DT cord-048487-kamvao4b 25 14 rate rate NN cord-048487-kamvao4b 25 15 of of IN cord-048487-kamvao4b 25 16 nucleotide nucleotide JJ cord-048487-kamvao4b 25 17 substitution substitution NN cord-048487-kamvao4b 25 18 over over IN cord-048487-kamvao4b 25 19 all all DT cord-048487-kamvao4b 25 20 taxa taxon NNS cord-048487-kamvao4b 25 21 in in IN cord-048487-kamvao4b 25 22 a a DT cord-048487-kamvao4b 25 23 given give VBN cord-048487-kamvao4b 25 24 set set NN cord-048487-kamvao4b 25 25 . . . cord-048487-kamvao4b 26 1 Consequently consequently RB cord-048487-kamvao4b 26 2 , , , cord-048487-kamvao4b 26 3 these these DT cord-048487-kamvao4b 26 4 methods method NNS cord-048487-kamvao4b 26 5 provide provide VBP cord-048487-kamvao4b 26 6 inference inference NN cord-048487-kamvao4b 26 7 most most RBS cord-048487-kamvao4b 26 8 suitable suitable JJ cord-048487-kamvao4b 26 9 for for IN cord-048487-kamvao4b 26 10 situations situation NNS cord-048487-kamvao4b 26 11 where where WRB cord-048487-kamvao4b 26 12 sequence sequence NN cord-048487-kamvao4b 26 13 evolution evolution NN cord-048487-kamvao4b 26 14 follows follow VBZ cord-048487-kamvao4b 26 15 a a DT cord-048487-kamvao4b 26 16 MCA MCA NNP cord-048487-kamvao4b 26 17 ( ( -LRB- cord-048487-kamvao4b 26 18 e.g. e.g. RB cord-048487-kamvao4b 26 19 influenza influenza NN cord-048487-kamvao4b 26 20 A A NNP cord-048487-kamvao4b 26 21 - - : cord-048487-kamvao4b 26 22 H3N2 H3N2 NNP cord-048487-kamvao4b 26 23 in in IN cord-048487-kamvao4b 26 24 human human JJ cord-048487-kamvao4b 26 25 hosts host NNS cord-048487-kamvao4b 26 26 , , , cord-048487-kamvao4b 26 27 as as IN cord-048487-kamvao4b 26 28 in in IN cord-048487-kamvao4b 26 29 [ [ -LRB- cord-048487-kamvao4b 26 30 9 9 CD cord-048487-kamvao4b 26 31 ] ] -RRB- cord-048487-kamvao4b 26 32 ) ) -RRB- cord-048487-kamvao4b 26 33 or or CC cord-048487-kamvao4b 26 34 deviates deviate VBZ cord-048487-kamvao4b 26 35 from from IN cord-048487-kamvao4b 26 36 the the DT cord-048487-kamvao4b 26 37 MCA MCA NNP cord-048487-kamvao4b 26 38 homogenously homogenously RB cord-048487-kamvao4b 26 39 in in IN cord-048487-kamvao4b 26 40 time time NN cord-048487-kamvao4b 26 41 ( ( -LRB- cord-048487-kamvao4b 26 42 e.g. e.g. RB cord-048487-kamvao4b 26 43 perhaps perhaps RB cord-048487-kamvao4b 26 44 influenza influenza NN cord-048487-kamvao4b 26 45 A a NN cord-048487-kamvao4b 26 46 in in IN cord-048487-kamvao4b 26 47 wild wild JJ cord-048487-kamvao4b 26 48 fowl fowl NN cord-048487-kamvao4b 26 49 , , , cord-048487-kamvao4b 26 50 see see VB cord-048487-kamvao4b 26 51 [ [ -LRB- cord-048487-kamvao4b 26 52 3 3 CD cord-048487-kamvao4b 26 53 ] ] -RRB- cord-048487-kamvao4b 26 54 ) ) -RRB- cord-048487-kamvao4b 26 55 . . . cord-048487-kamvao4b 27 1 In in IN cord-048487-kamvao4b 27 2 considering consider VBG cord-048487-kamvao4b 27 3 divergence divergence NN cord-048487-kamvao4b 27 4 events event NNS cord-048487-kamvao4b 27 5 between between IN cord-048487-kamvao4b 27 6 viral viral JJ cord-048487-kamvao4b 27 7 subtypes subtype NNS cord-048487-kamvao4b 27 8 , , , cord-048487-kamvao4b 27 9 even even RB cord-048487-kamvao4b 27 10 when when WRB cord-048487-kamvao4b 27 11 the the DT cord-048487-kamvao4b 27 12 MCA MCA NNP cord-048487-kamvao4b 27 13 well well RB cord-048487-kamvao4b 27 14 - - HYPH cord-048487-kamvao4b 27 15 approximates approximate VBZ cord-048487-kamvao4b 27 16 nucleotide nucleotide JJ cord-048487-kamvao4b 27 17 substitution substitution NN cord-048487-kamvao4b 27 18 within within IN cord-048487-kamvao4b 27 19 a a DT cord-048487-kamvao4b 27 20 given give VBN cord-048487-kamvao4b 27 21 subtype subtype NN cord-048487-kamvao4b 27 22 , , , cord-048487-kamvao4b 27 23 the the DT cord-048487-kamvao4b 27 24 above above JJ cord-048487-kamvao4b 27 25 methods method NNS cord-048487-kamvao4b 27 26 may may MD cord-048487-kamvao4b 27 27 incorrectly incorrectly RB cord-048487-kamvao4b 27 28 infer infer VB cord-048487-kamvao4b 27 29 the the DT cord-048487-kamvao4b 27 30 time time NN cord-048487-kamvao4b 27 31 of of IN cord-048487-kamvao4b 27 32 divergence divergence NN cord-048487-kamvao4b 27 33 across across IN cord-048487-kamvao4b 27 34 subtypes subtype NNS cord-048487-kamvao4b 27 35 . . . cord-048487-kamvao4b 28 1 By by IN cord-048487-kamvao4b 28 2 either either CC cord-048487-kamvao4b 28 3 assuming assume VBG cord-048487-kamvao4b 28 4 that that IN cord-048487-kamvao4b 28 5 a a DT cord-048487-kamvao4b 28 6 single single JJ cord-048487-kamvao4b 28 7 rate rate NN cord-048487-kamvao4b 28 8 of of IN cord-048487-kamvao4b 28 9 nucleotide nucleotide JJ cord-048487-kamvao4b 28 10 substitution substitution NN cord-048487-kamvao4b 28 11 holds hold VBZ cord-048487-kamvao4b 28 12 for for IN cord-048487-kamvao4b 28 13 the the DT cord-048487-kamvao4b 28 14 region region NN cord-048487-kamvao4b 28 15 preceding precede VBG cord-048487-kamvao4b 28 16 the the DT cord-048487-kamvao4b 28 17 common common JJ cord-048487-kamvao4b 28 18 ancestor ancestor NN cord-048487-kamvao4b 28 19 of of IN cord-048487-kamvao4b 28 20 each each DT cord-048487-kamvao4b 28 21 subtype subtype NN cord-048487-kamvao4b 28 22 or or CC cord-048487-kamvao4b 28 23 by by IN cord-048487-kamvao4b 28 24 smoothing smooth VBG cord-048487-kamvao4b 28 25 the the DT cord-048487-kamvao4b 28 26 rate rate NN cord-048487-kamvao4b 28 27 of of IN cord-048487-kamvao4b 28 28 nucleotide nucleotide JJ cord-048487-kamvao4b 28 29 substitution substitution NN cord-048487-kamvao4b 28 30 over over IN cord-048487-kamvao4b 28 31 clades clade NNS cord-048487-kamvao4b 28 32 with with IN cord-048487-kamvao4b 28 33 different different JJ cord-048487-kamvao4b 28 34 numbers number NNS cord-048487-kamvao4b 28 35 of of IN cord-048487-kamvao4b 28 36 taxa taxon NNS cord-048487-kamvao4b 28 37 , , , cord-048487-kamvao4b 28 38 the the DT cord-048487-kamvao4b 28 39 adherence adherence NN cord-048487-kamvao4b 28 40 to to IN cord-048487-kamvao4b 28 41 a a DT cord-048487-kamvao4b 28 42 MCA MCA NNP cord-048487-kamvao4b 28 43 prevents prevent VBZ cord-048487-kamvao4b 28 44 direct direct JJ cord-048487-kamvao4b 28 45 inference inference NN cord-048487-kamvao4b 28 46 of of IN cord-048487-kamvao4b 28 47 the the DT cord-048487-kamvao4b 28 48 rate rate NN cord-048487-kamvao4b 28 49 during during IN cord-048487-kamvao4b 28 50 subtype subtype NN cord-048487-kamvao4b 28 51 divergence divergence NN cord-048487-kamvao4b 28 52 . . . cord-048487-kamvao4b 29 1 Suzuki Suzuki NNP cord-048487-kamvao4b 29 2 and and CC cord-048487-kamvao4b 29 3 Nei Nei NNP cord-048487-kamvao4b 29 4 ( ( -LRB- cord-048487-kamvao4b 29 5 2002 2002 CD cord-048487-kamvao4b 29 6 ) ) -RRB- cord-048487-kamvao4b 29 7 propose propose VB cord-048487-kamvao4b 29 8 an an DT cord-048487-kamvao4b 29 9 alternative alternative NN cord-048487-kamvao4b 29 10 , , , cord-048487-kamvao4b 29 11 more more RBR cord-048487-kamvao4b 29 12 heuristic heuristic JJ cord-048487-kamvao4b 29 13 method method NN cord-048487-kamvao4b 29 14 of of IN cord-048487-kamvao4b 29 15 estimation estimation NN cord-048487-kamvao4b 29 16 to to TO cord-048487-kamvao4b 29 17 counteract counteract VB cord-048487-kamvao4b 29 18 the the DT cord-048487-kamvao4b 29 19 problem problem NN cord-048487-kamvao4b 29 20 of of IN cord-048487-kamvao4b 29 21 differing differ VBG cord-048487-kamvao4b 29 22 rates rate NNS cord-048487-kamvao4b 29 23 of of IN cord-048487-kamvao4b 29 24 substitution substitution NN cord-048487-kamvao4b 29 25 before before RB cord-048487-kamvao4b 29 26 and and CC cord-048487-kamvao4b 29 27 after after IN cord-048487-kamvao4b 29 28 zoonotic zoonotic JJ cord-048487-kamvao4b 29 29 events event NNS cord-048487-kamvao4b 29 30 [ [ -LRB- cord-048487-kamvao4b 29 31 23 23 CD cord-048487-kamvao4b 29 32 , , , cord-048487-kamvao4b 29 33 25 25 CD cord-048487-kamvao4b 29 34 ] ] -RRB- cord-048487-kamvao4b 29 35 . . . cord-048487-kamvao4b 30 1 In in IN cord-048487-kamvao4b 30 2 these these DT cord-048487-kamvao4b 30 3 studies study NNS cord-048487-kamvao4b 30 4 , , , cord-048487-kamvao4b 30 5 the the DT cord-048487-kamvao4b 30 6 evolutionary evolutionary JJ cord-048487-kamvao4b 30 7 models model NNS cord-048487-kamvao4b 30 8 draw draw VBP cord-048487-kamvao4b 30 9 a a DT cord-048487-kamvao4b 30 10 distinction distinction NN cord-048487-kamvao4b 30 11 between between IN cord-048487-kamvao4b 30 12 the the DT cord-048487-kamvao4b 30 13 rate rate NN cord-048487-kamvao4b 30 14 of of IN cord-048487-kamvao4b 30 15 substitution substitution NN cord-048487-kamvao4b 30 16 within within IN cord-048487-kamvao4b 30 17 a a DT cord-048487-kamvao4b 30 18 given give VBN cord-048487-kamvao4b 30 19 subtype subtype NN cord-048487-kamvao4b 30 20 and and CC cord-048487-kamvao4b 30 21 the the DT cord-048487-kamvao4b 30 22 rate rate NN cord-048487-kamvao4b 30 23 of of IN cord-048487-kamvao4b 30 24 substitution substitution NN cord-048487-kamvao4b 30 25 between between IN cord-048487-kamvao4b 30 26 subtypes subtype NNS cord-048487-kamvao4b 30 27 . . . cord-048487-kamvao4b 31 1 However however RB cord-048487-kamvao4b 31 2 , , , cord-048487-kamvao4b 31 3 trouble trouble NN cord-048487-kamvao4b 31 4 arises arise VBZ cord-048487-kamvao4b 31 5 since since IN cord-048487-kamvao4b 31 6 there there EX cord-048487-kamvao4b 31 7 are be VBP cord-048487-kamvao4b 31 8 no no DT cord-048487-kamvao4b 31 9 methods method NNS cord-048487-kamvao4b 31 10 for for IN cord-048487-kamvao4b 31 11 estimating estimate VBG cord-048487-kamvao4b 31 12 the the DT cord-048487-kamvao4b 31 13 latter latter JJ cord-048487-kamvao4b 31 14 quantity quantity NN cord-048487-kamvao4b 31 15 . . . cord-048487-kamvao4b 32 1 Consequently consequently RB cord-048487-kamvao4b 32 2 , , , cord-048487-kamvao4b 32 3 the the DT cord-048487-kamvao4b 32 4 models model NNS cord-048487-kamvao4b 32 5 assume assume VBP cord-048487-kamvao4b 32 6 that that IN cord-048487-kamvao4b 32 7 the the DT cord-048487-kamvao4b 32 8 rate rate NN cord-048487-kamvao4b 32 9 of of IN cord-048487-kamvao4b 32 10 substitution substitution NN cord-048487-kamvao4b 32 11 for for IN cord-048487-kamvao4b 32 12 portions portion NNS cord-048487-kamvao4b 32 13 of of IN cord-048487-kamvao4b 32 14 the the DT cord-048487-kamvao4b 32 15 phylogeny phylogeny NN cord-048487-kamvao4b 32 16 between between IN cord-048487-kamvao4b 32 17 the the DT cord-048487-kamvao4b 32 18 subtypes subtype NNS cord-048487-kamvao4b 32 19 equals equal VBZ cord-048487-kamvao4b 32 20 the the DT cord-048487-kamvao4b 32 21 mean mean JJ cord-048487-kamvao4b 32 22 rate rate NN cord-048487-kamvao4b 32 23 in in IN cord-048487-kamvao4b 32 24 the the DT cord-048487-kamvao4b 32 25 initial initial JJ cord-048487-kamvao4b 32 26 host host NN cord-048487-kamvao4b 32 27 species specie VBZ cord-048487-kamvao4b 32 28 population population NN cord-048487-kamvao4b 32 29 . . . cord-048487-kamvao4b 33 1 For for IN cord-048487-kamvao4b 33 2 instance instance NN cord-048487-kamvao4b 33 3 , , , cord-048487-kamvao4b 33 4 in in IN cord-048487-kamvao4b 33 5 dating date VBG cord-048487-kamvao4b 33 6 the the DT cord-048487-kamvao4b 33 7 time time NN cord-048487-kamvao4b 33 8 of of IN cord-048487-kamvao4b 33 9 divergence divergence NN cord-048487-kamvao4b 33 10 between between IN cord-048487-kamvao4b 33 11 influenza influenza NN cord-048487-kamvao4b 33 12 B b NN cord-048487-kamvao4b 33 13 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 33 14 and and CC cord-048487-kamvao4b 33 15 influenza influenza NN cord-048487-kamvao4b 33 16 C c CD cord-048487-kamvao4b 33 17 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 33 18 - - HYPH cord-048487-kamvao4b 33 19 esterase esterase NN cord-048487-kamvao4b 33 20 , , , cord-048487-kamvao4b 33 21 Suzuki Suzuki NNP cord-048487-kamvao4b 33 22 and and CC cord-048487-kamvao4b 33 23 Nei Nei NNP cord-048487-kamvao4b 33 24 use use VBP cord-048487-kamvao4b 33 25 the the DT cord-048487-kamvao4b 33 26 rate rate NN cord-048487-kamvao4b 33 27 of of IN cord-048487-kamvao4b 33 28 amino amino NN cord-048487-kamvao4b 33 29 acid acid NN cord-048487-kamvao4b 33 30 substitution substitution NN cord-048487-kamvao4b 33 31 for for IN cord-048487-kamvao4b 33 32 water water NN cord-048487-kamvao4b 33 33 fowl fowl NN cord-048487-kamvao4b 33 34 for for IN cord-048487-kamvao4b 33 35 the the DT cord-048487-kamvao4b 33 36 portions portion NNS cord-048487-kamvao4b 33 37 of of IN cord-048487-kamvao4b 33 38 the the DT cord-048487-kamvao4b 33 39 phylogeny phylogeny NN cord-048487-kamvao4b 33 40 previous previous JJ cord-048487-kamvao4b 33 41 to to IN cord-048487-kamvao4b 33 42 the the DT cord-048487-kamvao4b 33 43 TMRCA TMRCA NNP cord-048487-kamvao4b 33 44 of of IN cord-048487-kamvao4b 33 45 these these DT cord-048487-kamvao4b 33 46 two two CD cord-048487-kamvao4b 33 47 proteins protein NNS cord-048487-kamvao4b 33 48 [ [ -LRB- cord-048487-kamvao4b 33 49 23 23 CD cord-048487-kamvao4b 33 50 ] ] -RRB- cord-048487-kamvao4b 33 51 . . . cord-048487-kamvao4b 34 1 While while IN cord-048487-kamvao4b 34 2 this this DT cord-048487-kamvao4b 34 3 method method NN cord-048487-kamvao4b 34 4 may may MD cord-048487-kamvao4b 34 5 accurately accurately RB cord-048487-kamvao4b 34 6 reflect reflect VB cord-048487-kamvao4b 34 7 the the DT cord-048487-kamvao4b 34 8 rate rate NN cord-048487-kamvao4b 34 9 within within IN cord-048487-kamvao4b 34 10 avian avian JJ cord-048487-kamvao4b 34 11 and and CC cord-048487-kamvao4b 34 12 human human JJ cord-048487-kamvao4b 34 13 hosts host NNS cord-048487-kamvao4b 34 14 , , , cord-048487-kamvao4b 34 15 it -PRON- PRP cord-048487-kamvao4b 34 16 neglects neglect VBZ cord-048487-kamvao4b 34 17 whatever whatever WDT cord-048487-kamvao4b 34 18 additional additional JJ cord-048487-kamvao4b 34 19 changes change NNS cord-048487-kamvao4b 34 20 in in IN cord-048487-kamvao4b 34 21 the the DT cord-048487-kamvao4b 34 22 rate rate NN cord-048487-kamvao4b 34 23 of of IN cord-048487-kamvao4b 34 24 substitution substitution NN cord-048487-kamvao4b 34 25 are be VBP cord-048487-kamvao4b 34 26 due due JJ cord-048487-kamvao4b 34 27 to to IN cord-048487-kamvao4b 34 28 the the DT cord-048487-kamvao4b 34 29 process process NN cord-048487-kamvao4b 34 30 of of IN cord-048487-kamvao4b 34 31 zoonotic zoonotic JJ cord-048487-kamvao4b 34 32 adaptation adaptation NN cord-048487-kamvao4b 34 33 , , , cord-048487-kamvao4b 34 34 likely likely RB cord-048487-kamvao4b 34 35 leading lead VBG cord-048487-kamvao4b 34 36 to to IN cord-048487-kamvao4b 34 37 a a DT cord-048487-kamvao4b 34 38 substantial substantial JJ cord-048487-kamvao4b 34 39 underestimation underestimation NN cord-048487-kamvao4b 34 40 of of IN cord-048487-kamvao4b 34 41 the the DT cord-048487-kamvao4b 34 42 date date NN cord-048487-kamvao4b 34 43 of of IN cord-048487-kamvao4b 34 44 the the DT cord-048487-kamvao4b 34 45 TMRCA TMRCA NNP cord-048487-kamvao4b 34 46 . . . cord-048487-kamvao4b 35 1 The the DT cord-048487-kamvao4b 35 2 study study NN cord-048487-kamvao4b 35 3 here here RB cord-048487-kamvao4b 35 4 focuses focus VBZ cord-048487-kamvao4b 35 5 on on IN cord-048487-kamvao4b 35 6 influenza influenza NN cord-048487-kamvao4b 35 7 , , , cord-048487-kamvao4b 35 8 although although IN cord-048487-kamvao4b 35 9 the the DT cord-048487-kamvao4b 35 10 techniques technique NNS cord-048487-kamvao4b 35 11 are be VBP cord-048487-kamvao4b 35 12 readily readily RB cord-048487-kamvao4b 35 13 applied apply VBN cord-048487-kamvao4b 35 14 to to IN cord-048487-kamvao4b 35 15 other other JJ cord-048487-kamvao4b 35 16 rapidly rapidly RB cord-048487-kamvao4b 35 17 evolving evolve VBG cord-048487-kamvao4b 35 18 organisms organism NNS cord-048487-kamvao4b 35 19 . . . cord-048487-kamvao4b 36 1 Influenza influenza NN cord-048487-kamvao4b 36 2 has have VBZ cord-048487-kamvao4b 36 3 three three CD cord-048487-kamvao4b 36 4 types type NNS cord-048487-kamvao4b 36 5 , , , cord-048487-kamvao4b 36 6 A A NNP cord-048487-kamvao4b 36 7 , , , cord-048487-kamvao4b 36 8 B b NN cord-048487-kamvao4b 36 9 and and CC cord-048487-kamvao4b 36 10 C C NNP cord-048487-kamvao4b 36 11 , , , cord-048487-kamvao4b 36 12 classified classify VBN cord-048487-kamvao4b 36 13 based base VBN cord-048487-kamvao4b 36 14 on on IN cord-048487-kamvao4b 36 15 serological serological JJ cord-048487-kamvao4b 36 16 analysis analysis NN cord-048487-kamvao4b 36 17 . . . cord-048487-kamvao4b 37 1 To to IN cord-048487-kamvao4b 37 2 date date NN cord-048487-kamvao4b 37 3 , , , cord-048487-kamvao4b 37 4 only only RB cord-048487-kamvao4b 37 5 type type NN cord-048487-kamvao4b 37 6 A a DT cord-048487-kamvao4b 37 7 sequences sequence NNS cord-048487-kamvao4b 37 8 have have VBP cord-048487-kamvao4b 37 9 been be VBN cord-048487-kamvao4b 37 10 demonstrably demonstrably RB cord-048487-kamvao4b 37 11 associated associate VBN cord-048487-kamvao4b 37 12 with with IN cord-048487-kamvao4b 37 13 global global JJ cord-048487-kamvao4b 37 14 pandemics pandemic NNS cord-048487-kamvao4b 37 15 [ [ -LRB- cord-048487-kamvao4b 37 16 4 4 CD cord-048487-kamvao4b 37 17 ] ] -RRB- cord-048487-kamvao4b 37 18 . . . cord-048487-kamvao4b 38 1 Since since IN cord-048487-kamvao4b 38 2 modern modern JJ cord-048487-kamvao4b 38 3 surveillance surveillance NN cord-048487-kamvao4b 38 4 began begin VBD cord-048487-kamvao4b 38 5 in in IN cord-048487-kamvao4b 38 6 the the DT cord-048487-kamvao4b 38 7 1930s 1930 NNS cord-048487-kamvao4b 38 8 , , , cord-048487-kamvao4b 38 9 type type NN cord-048487-kamvao4b 38 10 B b NN cord-048487-kamvao4b 38 11 has have VBZ cord-048487-kamvao4b 38 12 only only RB cord-048487-kamvao4b 38 13 been be VBN cord-048487-kamvao4b 38 14 responsible responsible JJ cord-048487-kamvao4b 38 15 for for IN cord-048487-kamvao4b 38 16 mild mild JJ cord-048487-kamvao4b 38 17 epidemics epidemic NNS cord-048487-kamvao4b 38 18 while while IN cord-048487-kamvao4b 38 19 type type NN cord-048487-kamvao4b 38 20 C C NNP cord-048487-kamvao4b 38 21 has have VBZ cord-048487-kamvao4b 38 22 been be VBN cord-048487-kamvao4b 38 23 nearly nearly RB cord-048487-kamvao4b 38 24 asymptomatic asymptomatic JJ cord-048487-kamvao4b 38 25 in in IN cord-048487-kamvao4b 38 26 human human JJ cord-048487-kamvao4b 38 27 infection infection NN cord-048487-kamvao4b 38 28 . . . cord-048487-kamvao4b 39 1 Several several JJ cord-048487-kamvao4b 39 2 subtypes subtype NNS cord-048487-kamvao4b 39 3 of of IN cord-048487-kamvao4b 39 4 A a DT cord-048487-kamvao4b 39 5 , , , cord-048487-kamvao4b 39 6 notably notably RB cord-048487-kamvao4b 39 7 H1N1 H1N1 NNP cord-048487-kamvao4b 39 8 and and CC cord-048487-kamvao4b 39 9 H3N2 H3N2 NNP cord-048487-kamvao4b 39 10 , , , cord-048487-kamvao4b 39 11 are be VBP cord-048487-kamvao4b 39 12 currently currently RB cord-048487-kamvao4b 39 13 co co NN cord-048487-kamvao4b 39 14 - - NN cord-048487-kamvao4b 39 15 circulating circulate VBG cord-048487-kamvao4b 39 16 in in IN cord-048487-kamvao4b 39 17 the the DT cord-048487-kamvao4b 39 18 human human JJ cord-048487-kamvao4b 39 19 population population NN cord-048487-kamvao4b 39 20 . . . cord-048487-kamvao4b 40 1 As as IN cord-048487-kamvao4b 40 2 the the DT cord-048487-kamvao4b 40 3 H1N1 H1N1 NNP cord-048487-kamvao4b 40 4 and and CC cord-048487-kamvao4b 40 5 H3N2 H3N2 NNP cord-048487-kamvao4b 40 6 subtypes subtype NNS cord-048487-kamvao4b 40 7 may may MD cord-048487-kamvao4b 40 8 be be VB cord-048487-kamvao4b 40 9 as as RB cord-048487-kamvao4b 40 10 divergent divergent JJ cord-048487-kamvao4b 40 11 from from IN cord-048487-kamvao4b 40 12 each each DT cord-048487-kamvao4b 40 13 other other JJ cord-048487-kamvao4b 40 14 as as IN cord-048487-kamvao4b 40 15 they -PRON- PRP cord-048487-kamvao4b 40 16 are be VBP cord-048487-kamvao4b 40 17 from from IN cord-048487-kamvao4b 40 18 types type NNS cord-048487-kamvao4b 40 19 B b NN cord-048487-kamvao4b 40 20 and and CC cord-048487-kamvao4b 40 21 C C NNP cord-048487-kamvao4b 40 22 , , , cord-048487-kamvao4b 40 23 we -PRON- PRP cord-048487-kamvao4b 40 24 will will MD cord-048487-kamvao4b 40 25 refer refer VB cord-048487-kamvao4b 40 26 to to IN cord-048487-kamvao4b 40 27 all all DT cord-048487-kamvao4b 40 28 types type NNS cord-048487-kamvao4b 40 29 and and CC cord-048487-kamvao4b 40 30 subtypes subtype NNS cord-048487-kamvao4b 40 31 simply simply RB cord-048487-kamvao4b 40 32 as as IN cord-048487-kamvao4b 40 33 subtypes subtype NNS cord-048487-kamvao4b 40 34 for for IN cord-048487-kamvao4b 40 35 the the DT cord-048487-kamvao4b 40 36 remainder remainder NN cord-048487-kamvao4b 40 37 of of IN cord-048487-kamvao4b 40 38 this this DT cord-048487-kamvao4b 40 39 paper paper NN cord-048487-kamvao4b 40 40 . . . cord-048487-kamvao4b 41 1 We -PRON- PRP cord-048487-kamvao4b 41 2 select select VBP cord-048487-kamvao4b 41 3 for for IN cord-048487-kamvao4b 41 4 this this DT cord-048487-kamvao4b 41 5 study study NN cord-048487-kamvao4b 41 6 three three CD cord-048487-kamvao4b 41 7 genes gene NNS cord-048487-kamvao4b 41 8 , , , cord-048487-kamvao4b 41 9 coding code VBG cord-048487-kamvao4b 41 10 for for IN cord-048487-kamvao4b 41 11 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 41 12 ( ( -LRB- cord-048487-kamvao4b 41 13 HA HA NNP cord-048487-kamvao4b 41 14 ) ) -RRB- cord-048487-kamvao4b 41 15 , , , cord-048487-kamvao4b 41 16 the the DT cord-048487-kamvao4b 41 17 matrix matrix NN cord-048487-kamvao4b 41 18 protein protein NN cord-048487-kamvao4b 41 19 ( ( -LRB- cord-048487-kamvao4b 41 20 MP MP NNP cord-048487-kamvao4b 41 21 ) ) -RRB- cord-048487-kamvao4b 41 22 and and CC cord-048487-kamvao4b 41 23 the the DT cord-048487-kamvao4b 41 24 non non JJ cord-048487-kamvao4b 41 25 - - JJ cord-048487-kamvao4b 41 26 structural structural JJ cord-048487-kamvao4b 41 27 protein protein NN cord-048487-kamvao4b 41 28 ( ( -LRB- cord-048487-kamvao4b 41 29 NS NS NNP cord-048487-kamvao4b 41 30 ) ) -RRB- cord-048487-kamvao4b 41 31 responsible responsible JJ cord-048487-kamvao4b 41 32 for for IN cord-048487-kamvao4b 41 33 interfering interfere VBG cord-048487-kamvao4b 41 34 with with IN cord-048487-kamvao4b 41 35 host host NN cord-048487-kamvao4b 41 36 immune immune JJ cord-048487-kamvao4b 41 37 response response NN cord-048487-kamvao4b 41 38 . . . cord-048487-kamvao4b 42 1 Subtype Subtype NNP cord-048487-kamvao4b 42 2 C C NNP cord-048487-kamvao4b 42 3 has have VBZ cord-048487-kamvao4b 42 4 a a DT cord-048487-kamvao4b 42 5 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 42 6 - - HYPH cord-048487-kamvao4b 42 7 esterase esterase NN cord-048487-kamvao4b 42 8 gene gene NN cord-048487-kamvao4b 42 9 that that WDT cord-048487-kamvao4b 42 10 is be VBZ cord-048487-kamvao4b 42 11 analogous analogous JJ cord-048487-kamvao4b 42 12 to to IN cord-048487-kamvao4b 42 13 the the DT cord-048487-kamvao4b 42 14 hemagglutin hemagglutin NNP cord-048487-kamvao4b 42 15 gene gene NN cord-048487-kamvao4b 42 16 in in IN cord-048487-kamvao4b 42 17 other other JJ cord-048487-kamvao4b 42 18 subtypes subtype NNS cord-048487-kamvao4b 42 19 [ [ -LRB- cord-048487-kamvao4b 42 20 1 1 CD cord-048487-kamvao4b 42 21 ] ] -RRB- cord-048487-kamvao4b 42 22 . . . cord-048487-kamvao4b 43 1 We -PRON- PRP cord-048487-kamvao4b 43 2 hence hence RB cord-048487-kamvao4b 43 3 refer refer VBP cord-048487-kamvao4b 43 4 to to IN cord-048487-kamvao4b 43 5 the the DT cord-048487-kamvao4b 43 6 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 43 7 gene gene NN cord-048487-kamvao4b 43 8 generally generally RB cord-048487-kamvao4b 43 9 and and CC cord-048487-kamvao4b 43 10 the the DT cord-048487-kamvao4b 43 11 hemagglutininesterase hemagglutininesterase NNP cord-048487-kamvao4b 43 12 gene gene NN cord-048487-kamvao4b 43 13 when when WRB cord-048487-kamvao4b 43 14 referring refer VBG cord-048487-kamvao4b 43 15 specifically specifically RB cord-048487-kamvao4b 43 16 to to IN cord-048487-kamvao4b 43 17 the the DT cord-048487-kamvao4b 43 18 subtype subtype NN cord-048487-kamvao4b 43 19 C c NN cord-048487-kamvao4b 43 20 sequences sequence NNS cord-048487-kamvao4b 43 21 . . . cord-048487-kamvao4b 44 1 We -PRON- PRP cord-048487-kamvao4b 44 2 present present VBP cord-048487-kamvao4b 44 3 a a DT cord-048487-kamvao4b 44 4 simple simple JJ cord-048487-kamvao4b 44 5 estimation estimation NN cord-048487-kamvao4b 44 6 tool tool NN cord-048487-kamvao4b 44 7 to to TO cord-048487-kamvao4b 44 8 determine determine VB cord-048487-kamvao4b 44 9 the the DT cord-048487-kamvao4b 44 10 date date NN cord-048487-kamvao4b 44 11 of of IN cord-048487-kamvao4b 44 12 divergence divergence NN cord-048487-kamvao4b 44 13 among among IN cord-048487-kamvao4b 44 14 viral viral JJ cord-048487-kamvao4b 44 15 subtypes subtype NNS cord-048487-kamvao4b 44 16 that that WDT cord-048487-kamvao4b 44 17 overcomes overcome VBZ cord-048487-kamvao4b 44 18 the the DT cord-048487-kamvao4b 44 19 difficulties difficulty NNS cord-048487-kamvao4b 44 20 encountered encounter VBN cord-048487-kamvao4b 44 21 with with IN cord-048487-kamvao4b 44 22 use use NN cord-048487-kamvao4b 44 23 of of IN cord-048487-kamvao4b 44 24 the the DT cord-048487-kamvao4b 44 25 MCA MCA NNP cord-048487-kamvao4b 44 26 by by IN cord-048487-kamvao4b 44 27 measuring measure VBG cord-048487-kamvao4b 44 28 the the DT cord-048487-kamvao4b 44 29 pairwise pairwise JJ cord-048487-kamvao4b 44 30 rate rate NN cord-048487-kamvao4b 44 31 of of IN cord-048487-kamvao4b 44 32 substitution substitution NN cord-048487-kamvao4b 44 33 between between IN cord-048487-kamvao4b 44 34 taxa taxon NNS cord-048487-kamvao4b 44 35 . . . cord-048487-kamvao4b 45 1 Our -PRON- PRP$ cord-048487-kamvao4b 45 2 estimator estimator NN cord-048487-kamvao4b 45 3 derives derive VBZ cord-048487-kamvao4b 45 4 from from IN cord-048487-kamvao4b 45 5 the the DT cord-048487-kamvao4b 45 6 triplet triplet NN cord-048487-kamvao4b 45 7 statistic statistic NN cord-048487-kamvao4b 45 8 developed develop VBN cord-048487-kamvao4b 45 9 in in IN cord-048487-kamvao4b 45 10 [ [ -LRB- cord-048487-kamvao4b 45 11 26 26 CD cord-048487-kamvao4b 45 12 , , , cord-048487-kamvao4b 45 13 22 22 CD cord-048487-kamvao4b 45 14 , , , cord-048487-kamvao4b 45 15 13 13 CD cord-048487-kamvao4b 45 16 ] ] -RRB- cord-048487-kamvao4b 45 17 , , , cord-048487-kamvao4b 45 18 where where WRB cord-048487-kamvao4b 45 19 each each DT cord-048487-kamvao4b 45 20 sequence sequence NN cord-048487-kamvao4b 45 21 member member NN cord-048487-kamvao4b 45 22 of of IN cord-048487-kamvao4b 45 23 the the DT cord-048487-kamvao4b 45 24 triplet triplet NN cord-048487-kamvao4b 45 25 draws draw VBZ cord-048487-kamvao4b 45 26 from from IN cord-048487-kamvao4b 45 27 a a DT cord-048487-kamvao4b 45 28 different different JJ cord-048487-kamvao4b 45 29 subtype subtype NN cord-048487-kamvao4b 45 30 . . . cord-048487-kamvao4b 46 1 In in IN cord-048487-kamvao4b 46 2 this this DT cord-048487-kamvao4b 46 3 manner manner NN cord-048487-kamvao4b 46 4 , , , cord-048487-kamvao4b 46 5 we -PRON- PRP cord-048487-kamvao4b 46 6 generate generate VBP cord-048487-kamvao4b 46 7 from from IN cord-048487-kamvao4b 46 8 each each DT cord-048487-kamvao4b 46 9 triplet triplet NN cord-048487-kamvao4b 46 10 an an DT cord-048487-kamvao4b 46 11 estimate estimate NN cord-048487-kamvao4b 46 12 of of IN cord-048487-kamvao4b 46 13 the the DT cord-048487-kamvao4b 46 14 rate rate NN cord-048487-kamvao4b 46 15 of of IN cord-048487-kamvao4b 46 16 nucleotide nucleotide JJ cord-048487-kamvao4b 46 17 substitution substitution NN cord-048487-kamvao4b 46 18 between between IN cord-048487-kamvao4b 46 19 the the DT cord-048487-kamvao4b 46 20 most most RBS cord-048487-kamvao4b 46 21 recently recently RB cord-048487-kamvao4b 46 22 diverged diverge VBN cord-048487-kamvao4b 46 23 subtypes subtype NNS cord-048487-kamvao4b 46 24 , , , cord-048487-kamvao4b 46 25 and and CC cord-048487-kamvao4b 46 26 consequently consequently RB cord-048487-kamvao4b 46 27 provide provide VB cord-048487-kamvao4b 46 28 an an DT cord-048487-kamvao4b 46 29 estimate estimate NN cord-048487-kamvao4b 46 30 of of IN cord-048487-kamvao4b 46 31 the the DT cord-048487-kamvao4b 46 32 TMRCA TMRCA NNP cord-048487-kamvao4b 46 33 . . . cord-048487-kamvao4b 47 1 This this DT cord-048487-kamvao4b 47 2 circumvents circumvent VBZ cord-048487-kamvao4b 47 3 the the DT cord-048487-kamvao4b 47 4 problems problem NNS cord-048487-kamvao4b 47 5 posed pose VBN cord-048487-kamvao4b 47 6 by by IN cord-048487-kamvao4b 47 7 earlier early JJR cord-048487-kamvao4b 47 8 methods method NNS cord-048487-kamvao4b 47 9 by by IN cord-048487-kamvao4b 47 10 directly directly RB cord-048487-kamvao4b 47 11 estimating estimate VBG cord-048487-kamvao4b 47 12 the the DT cord-048487-kamvao4b 47 13 pairwise pairwise JJ cord-048487-kamvao4b 47 14 rate rate NN cord-048487-kamvao4b 47 15 of of IN cord-048487-kamvao4b 47 16 nucleotide nucleotide JJ cord-048487-kamvao4b 47 17 substitution substitution NN cord-048487-kamvao4b 47 18 over over IN cord-048487-kamvao4b 47 19 the the DT cord-048487-kamvao4b 47 20 set set NN cord-048487-kamvao4b 47 21 of of IN cord-048487-kamvao4b 47 22 pairs pair NNS cord-048487-kamvao4b 47 23 of of IN cord-048487-kamvao4b 47 24 sequences sequence NNS cord-048487-kamvao4b 47 25 straddling straddle VBG cord-048487-kamvao4b 47 26 the the DT cord-048487-kamvao4b 47 27 subtype subtype NN cord-048487-kamvao4b 47 28 divergence divergence NN cord-048487-kamvao4b 47 29 without without IN cord-048487-kamvao4b 47 30 any any DT cord-048487-kamvao4b 47 31 further further JJ cord-048487-kamvao4b 47 32 rate rate NN cord-048487-kamvao4b 47 33 assumptions assumption NNS cord-048487-kamvao4b 47 34 other other JJ cord-048487-kamvao4b 47 35 than than IN cord-048487-kamvao4b 47 36 the the DT cord-048487-kamvao4b 47 37 existence existence NN cord-048487-kamvao4b 47 38 of of IN cord-048487-kamvao4b 47 39 a a DT cord-048487-kamvao4b 47 40 mean mean NN cord-048487-kamvao4b 47 41 . . . cord-048487-kamvao4b 48 1 However however RB cord-048487-kamvao4b 48 2 , , , cord-048487-kamvao4b 48 3 this this DT cord-048487-kamvao4b 48 4 method method NN cord-048487-kamvao4b 48 5 is be VBZ cord-048487-kamvao4b 48 6 only only RB cord-048487-kamvao4b 48 7 capable capable JJ cord-048487-kamvao4b 48 8 of of IN cord-048487-kamvao4b 48 9 determining determine VBG cord-048487-kamvao4b 48 10 the the DT cord-048487-kamvao4b 48 11 rate rate NN cord-048487-kamvao4b 48 12 between between IN cord-048487-kamvao4b 48 13 two two CD cord-048487-kamvao4b 48 14 subtypes subtype NNS cord-048487-kamvao4b 48 15 where where WRB cord-048487-kamvao4b 48 16 a a DT cord-048487-kamvao4b 48 17 third third NN cord-048487-kamvao4b 48 18 , , , cord-048487-kamvao4b 48 19 more more RBR cord-048487-kamvao4b 48 20 distantly distantly RB cord-048487-kamvao4b 48 21 related relate VBN cord-048487-kamvao4b 48 22 , , , cord-048487-kamvao4b 48 23 subtype subtype NN cord-048487-kamvao4b 48 24 functions function NNS cord-048487-kamvao4b 48 25 as as IN cord-048487-kamvao4b 48 26 an an DT cord-048487-kamvao4b 48 27 outgroup outgroup NN cord-048487-kamvao4b 48 28 . . . cord-048487-kamvao4b 49 1 This this DT cord-048487-kamvao4b 49 2 method method NN cord-048487-kamvao4b 49 3 thus thus RB cord-048487-kamvao4b 49 4 trades trade VBZ cord-048487-kamvao4b 49 5 the the DT cord-048487-kamvao4b 49 6 ad ad FW cord-048487-kamvao4b 49 7 hoc hoc FW cord-048487-kamvao4b 49 8 rate rate NN cord-048487-kamvao4b 49 9 assumptions assumption NNS cord-048487-kamvao4b 49 10 of of IN cord-048487-kamvao4b 49 11 the the DT cord-048487-kamvao4b 49 12 previous previous JJ cord-048487-kamvao4b 49 13 methods method NNS cord-048487-kamvao4b 49 14 with with IN cord-048487-kamvao4b 49 15 two two CD cord-048487-kamvao4b 49 16 implicit implicit JJ cord-048487-kamvao4b 49 17 conditions condition NNS cord-048487-kamvao4b 49 18 : : : cord-048487-kamvao4b 50 1 ( ( -LRB- cord-048487-kamvao4b 50 2 i i NN cord-048487-kamvao4b 50 3 ) ) -RRB- cord-048487-kamvao4b 50 4 that that IN cord-048487-kamvao4b 50 5 subtypes subtype NNS cord-048487-kamvao4b 50 6 have have VBP cord-048487-kamvao4b 50 7 a a DT cord-048487-kamvao4b 50 8 unique unique JJ cord-048487-kamvao4b 50 9 divergence divergence NN cord-048487-kamvao4b 50 10 and and CC cord-048487-kamvao4b 50 11 ( ( -LRB- cord-048487-kamvao4b 50 12 ii ii NN cord-048487-kamvao4b 50 13 ) ) -RRB- cord-048487-kamvao4b 51 1 a a DT cord-048487-kamvao4b 51 2 third third JJ cord-048487-kamvao4b 51 3 , , , cord-048487-kamvao4b 51 4 comparable comparable JJ cord-048487-kamvao4b 51 5 subtype subtype NN cord-048487-kamvao4b 51 6 is be VBZ cord-048487-kamvao4b 51 7 available available JJ cord-048487-kamvao4b 51 8 to to TO cord-048487-kamvao4b 51 9 serve serve VB cord-048487-kamvao4b 51 10 as as IN cord-048487-kamvao4b 51 11 an an DT cord-048487-kamvao4b 51 12 outgroup outgroup NN cord-048487-kamvao4b 51 13 . . . cord-048487-kamvao4b 52 1 In in IN cord-048487-kamvao4b 52 2 exchange exchange NN cord-048487-kamvao4b 52 3 , , , cord-048487-kamvao4b 52 4 we -PRON- PRP cord-048487-kamvao4b 52 5 arrive arrive VBP cord-048487-kamvao4b 52 6 at at IN cord-048487-kamvao4b 52 7 a a DT cord-048487-kamvao4b 52 8 precise precise JJ cord-048487-kamvao4b 52 9 statistical statistical JJ cord-048487-kamvao4b 52 10 measure measure NN cord-048487-kamvao4b 52 11 of of IN cord-048487-kamvao4b 52 12 the the DT cord-048487-kamvao4b 52 13 TMRCA TMRCA NNP cord-048487-kamvao4b 52 14 that that WDT cord-048487-kamvao4b 52 15 converges converge VBZ cord-048487-kamvao4b 52 16 as as IN cord-048487-kamvao4b 52 17 the the DT cord-048487-kamvao4b 52 18 number number NN cord-048487-kamvao4b 52 19 of of IN cord-048487-kamvao4b 52 20 taxa taxon NNS cord-048487-kamvao4b 52 21 increases increase NNS cord-048487-kamvao4b 52 22 and and CC cord-048487-kamvao4b 52 23 is be VBZ cord-048487-kamvao4b 52 24 robust robust JJ cord-048487-kamvao4b 52 25 to to IN cord-048487-kamvao4b 52 26 the the DT cord-048487-kamvao4b 52 27 balancing balancing NN cord-048487-kamvao4b 52 28 of of IN cord-048487-kamvao4b 52 29 the the DT cord-048487-kamvao4b 52 30 numbers number NNS cord-048487-kamvao4b 52 31 of of IN cord-048487-kamvao4b 52 32 taxa taxon NNS cord-048487-kamvao4b 52 33 between between IN cord-048487-kamvao4b 52 34 different different JJ cord-048487-kamvao4b 52 35 subtypes subtype NNS cord-048487-kamvao4b 52 36 . . . cord-048487-kamvao4b 53 1 We -PRON- PRP cord-048487-kamvao4b 53 2 show show VBP cord-048487-kamvao4b 53 3 that that IN cord-048487-kamvao4b 53 4 applying apply VBG cord-048487-kamvao4b 53 5 this this DT cord-048487-kamvao4b 53 6 method method NN cord-048487-kamvao4b 53 7 to to IN cord-048487-kamvao4b 53 8 dating date VBG cord-048487-kamvao4b 53 9 the the DT cord-048487-kamvao4b 53 10 divergence divergence NN cord-048487-kamvao4b 53 11 of of IN cord-048487-kamvao4b 53 12 influenza influenza NN cord-048487-kamvao4b 53 13 subtypes subtype NNS cord-048487-kamvao4b 53 14 A A NNP cord-048487-kamvao4b 53 15 - - : cord-048487-kamvao4b 53 16 H3N2 H3N2 NNP cord-048487-kamvao4b 53 17 and and CC cord-048487-kamvao4b 53 18 B B NNP cord-048487-kamvao4b 53 19 gives give VBZ cord-048487-kamvao4b 53 20 a a DT cord-048487-kamvao4b 53 21 time time NN cord-048487-kamvao4b 53 22 of of IN cord-048487-kamvao4b 53 23 diver diver NN cord-048487-kamvao4b 53 24 - - HYPH cord-048487-kamvao4b 53 25 gence gence NNP cord-048487-kamvao4b 53 26 approximately approximately RB cord-048487-kamvao4b 53 27 100 100 CD cord-048487-kamvao4b 53 28 years year NNS cord-048487-kamvao4b 53 29 before before IN cord-048487-kamvao4b 53 30 present present JJ cord-048487-kamvao4b 53 31 , , , cord-048487-kamvao4b 53 32 substantially substantially RB cord-048487-kamvao4b 53 33 more more RBR cord-048487-kamvao4b 53 34 recent recent JJ cord-048487-kamvao4b 53 35 than than IN cord-048487-kamvao4b 53 36 previous previous JJ cord-048487-kamvao4b 53 37 estimates estimate NNS cord-048487-kamvao4b 53 38 . . . cord-048487-kamvao4b 54 1 To to TO cord-048487-kamvao4b 54 2 calculate calculate VB cord-048487-kamvao4b 54 3 the the DT cord-048487-kamvao4b 54 4 rate rate NN cord-048487-kamvao4b 54 5 of of IN cord-048487-kamvao4b 54 6 nucleotide nucleotide JJ cord-048487-kamvao4b 54 7 substitution substitution NN cord-048487-kamvao4b 54 8 , , , cord-048487-kamvao4b 54 9 we -PRON- PRP cord-048487-kamvao4b 54 10 require require VBP cord-048487-kamvao4b 54 11 a a DT cord-048487-kamvao4b 54 12 measurement measurement NN cord-048487-kamvao4b 54 13 of of IN cord-048487-kamvao4b 54 14 the the DT cord-048487-kamvao4b 54 15 number number NN cord-048487-kamvao4b 54 16 of of IN cord-048487-kamvao4b 54 17 nucleotide nucleotide JJ cord-048487-kamvao4b 54 18 substitutions substitution NNS cord-048487-kamvao4b 54 19 occurring occur VBG cord-048487-kamvao4b 54 20 in in IN cord-048487-kamvao4b 54 21 a a DT cord-048487-kamvao4b 54 22 given give VBN cord-048487-kamvao4b 54 23 time time NN cord-048487-kamvao4b 54 24 interval interval NN cord-048487-kamvao4b 54 25 . . . cord-048487-kamvao4b 55 1 Starting start VBG cord-048487-kamvao4b 55 2 from from IN cord-048487-kamvao4b 55 3 a a DT cord-048487-kamvao4b 55 4 given give VBN cord-048487-kamvao4b 55 5 set set NN cord-048487-kamvao4b 55 6 of of IN cord-048487-kamvao4b 55 7 aligned align VBN cord-048487-kamvao4b 55 8 sequences sequence NNS cord-048487-kamvao4b 55 9 { { -LRB- cord-048487-kamvao4b 55 10 s s NN cord-048487-kamvao4b 55 11 1 1 CD cord-048487-kamvao4b 55 12 , , , cord-048487-kamvao4b 55 13 ... ... NFP cord-048487-kamvao4b 55 14 , , , cord-048487-kamvao4b 55 15 s s NN cord-048487-kamvao4b 55 16 n n NN cord-048487-kamvao4b 55 17 } } -RRB- cord-048487-kamvao4b 55 18 for for IN cord-048487-kamvao4b 55 19 n n CD cord-048487-kamvao4b 55 20 taxa taxon NNS cord-048487-kamvao4b 55 21 , , , cord-048487-kamvao4b 55 22 we -PRON- PRP cord-048487-kamvao4b 55 23 define define VBP cord-048487-kamvao4b 55 24 the the DT cord-048487-kamvao4b 55 25 pairwise pairwise JJ cord-048487-kamvao4b 55 26 distance distance NN cord-048487-kamvao4b 55 27 in in IN cord-048487-kamvao4b 55 28 number number NN cord-048487-kamvao4b 55 29 of of IN cord-048487-kamvao4b 55 30 substitutions substitution NNS cord-048487-kamvao4b 55 31 to to TO cord-048487-kamvao4b 55 32 be be VB cord-048487-kamvao4b 55 33 the the DT cord-048487-kamvao4b 55 34 estimates estimate NNS cord-048487-kamvao4b 55 35 { { -LRB- cord-048487-kamvao4b 55 36 K K NNP cord-048487-kamvao4b 55 37 ij ij NNP cord-048487-kamvao4b 55 38 } } -RRB- cord-048487-kamvao4b 55 39 under under IN cord-048487-kamvao4b 55 40 a a DT cord-048487-kamvao4b 55 41 given give VBN cord-048487-kamvao4b 55 42 model model NN cord-048487-kamvao4b 55 43 of of IN cord-048487-kamvao4b 55 44 nucleotide nucleotide JJ cord-048487-kamvao4b 55 45 substitution substitution NN cord-048487-kamvao4b 55 46 . . . cord-048487-kamvao4b 56 1 Naturally naturally RB cord-048487-kamvao4b 56 2 the the DT cord-048487-kamvao4b 56 3 unobservable unobservable JJ cord-048487-kamvao4b 56 4 true true JJ cord-048487-kamvao4b 56 5 values value NNS cord-048487-kamvao4b 56 6 { { -LRB- cord-048487-kamvao4b 56 7 D d NN cord-048487-kamvao4b 56 8 ij ij NN cord-048487-kamvao4b 56 9 } } -RRB- cord-048487-kamvao4b 56 10 of of IN cord-048487-kamvao4b 56 11 the the DT cord-048487-kamvao4b 56 12 pairwise pairwise JJ cord-048487-kamvao4b 56 13 distances distance NNS cord-048487-kamvao4b 56 14 differ differ VBP cord-048487-kamvao4b 56 15 from from IN cord-048487-kamvao4b 56 16 their -PRON- PRP$ cord-048487-kamvao4b 56 17 estimates estimate NNS cord-048487-kamvao4b 56 18 { { -LRB- cord-048487-kamvao4b 56 19 K K NNP cord-048487-kamvao4b 56 20 ij ij NNP cord-048487-kamvao4b 56 21 } } -RRB- cord-048487-kamvao4b 56 22 . . . cord-048487-kamvao4b 57 1 To to TO cord-048487-kamvao4b 57 2 understand understand VB cord-048487-kamvao4b 57 3 this this DT cord-048487-kamvao4b 57 4 difference difference NN cord-048487-kamvao4b 57 5 , , , cord-048487-kamvao4b 57 6 we -PRON- PRP cord-048487-kamvao4b 57 7 associate associate VBP cord-048487-kamvao4b 57 8 each each DT cord-048487-kamvao4b 57 9 D d NN cord-048487-kamvao4b 57 10 ij ij NN cord-048487-kamvao4b 57 11 with with IN cord-048487-kamvao4b 57 12 an an DT cord-048487-kamvao4b 57 13 error error NN cord-048487-kamvao4b 57 14 ε ε NN cord-048487-kamvao4b 57 15 ij ij NNP cord-048487-kamvao4b 57 16 and and CC cord-048487-kamvao4b 57 17 assume assume VB cord-048487-kamvao4b 57 18 that that IN cord-048487-kamvao4b 57 19 ε ε NNP cord-048487-kamvao4b 57 20 ij ij NNP cord-048487-kamvao4b 57 21 tends tend VBZ cord-048487-kamvao4b 57 22 to to IN cord-048487-kamvao4b 57 23 zero zero CD cord-048487-kamvao4b 57 24 as as IN cord-048487-kamvao4b 57 25 sequence sequence NN cord-048487-kamvao4b 57 26 lengths length NNS cord-048487-kamvao4b 57 27 increase increase VBP cord-048487-kamvao4b 57 28 without without IN cord-048487-kamvao4b 57 29 bound bind VBN cord-048487-kamvao4b 57 30 . . . cord-048487-kamvao4b 58 1 We -PRON- PRP cord-048487-kamvao4b 58 2 further further RB cord-048487-kamvao4b 58 3 assume assume VBP cord-048487-kamvao4b 58 4 that that IN cord-048487-kamvao4b 58 5 the the DT cord-048487-kamvao4b 58 6 covariance covariance NN cord-048487-kamvao4b 58 7 between between IN cord-048487-kamvao4b 58 8 errors error NNS cord-048487-kamvao4b 58 9 , , , cord-048487-kamvao4b 58 10 cov(ε cov(ε NNP cord-048487-kamvao4b 58 11 ij ij NNP cord-048487-kamvao4b 58 12 ; ; : cord-048487-kamvao4b 58 13 ε ε NNP cord-048487-kamvao4b 58 14 mn mn NNP cord-048487-kamvao4b 58 15 ) ) -RRB- cord-048487-kamvao4b 58 16 , , , cord-048487-kamvao4b 58 17 is be VBZ cord-048487-kamvao4b 58 18 bounded bound VBN cord-048487-kamvao4b 58 19 and and CC cord-048487-kamvao4b 58 20 known know VBN cord-048487-kamvao4b 58 21 . . . cord-048487-kamvao4b 59 1 For for IN cord-048487-kamvao4b 59 2 time time NN cord-048487-kamvao4b 59 3 measurements measurement NNS cord-048487-kamvao4b 59 4 , , , cord-048487-kamvao4b 59 5 we -PRON- PRP cord-048487-kamvao4b 59 6 assume assume VBP cord-048487-kamvao4b 59 7 that that IN cord-048487-kamvao4b 59 8 each each DT cord-048487-kamvao4b 59 9 sequence sequence NN cord-048487-kamvao4b 59 10 is be VBZ cord-048487-kamvao4b 59 11 labeled label VBN cord-048487-kamvao4b 59 12 by by IN cord-048487-kamvao4b 59 13 a a DT cord-048487-kamvao4b 59 14 sampling sampling NN cord-048487-kamvao4b 59 15 time time NN cord-048487-kamvao4b 59 16 t t NNP cord-048487-kamvao4b 59 17 i i PRP cord-048487-kamvao4b 59 18 given give VBN cord-048487-kamvao4b 59 19 in in IN cord-048487-kamvao4b 59 20 consistent consistent JJ cord-048487-kamvao4b 59 21 units unit NNS cord-048487-kamvao4b 59 22 . . . cord-048487-kamvao4b 60 1 Since since IN cord-048487-kamvao4b 60 2 we -PRON- PRP cord-048487-kamvao4b 60 3 know know VBP cord-048487-kamvao4b 60 4 only only RB cord-048487-kamvao4b 60 5 the the DT cord-048487-kamvao4b 60 6 sampling sampling NN cord-048487-kamvao4b 60 7 time time NN cord-048487-kamvao4b 60 8 of of IN cord-048487-kamvao4b 60 9 a a DT cord-048487-kamvao4b 60 10 given give VBN cord-048487-kamvao4b 60 11 sample sample NN cord-048487-kamvao4b 60 12 up up RP cord-048487-kamvao4b 60 13 to to IN cord-048487-kamvao4b 60 14 the the DT cord-048487-kamvao4b 60 15 unit unit NN cord-048487-kamvao4b 60 16 of of IN cord-048487-kamvao4b 60 17 time time NN cord-048487-kamvao4b 60 18 reported report VBN cord-048487-kamvao4b 60 19 ( ( -LRB- cord-048487-kamvao4b 60 20 day day NN cord-048487-kamvao4b 60 21 , , , cord-048487-kamvao4b 60 22 month month NN cord-048487-kamvao4b 60 23 , , , cord-048487-kamvao4b 60 24 year year NN cord-048487-kamvao4b 60 25 ) ) -RRB- cord-048487-kamvao4b 60 26 we -PRON- PRP cord-048487-kamvao4b 60 27 posit posit VBP cord-048487-kamvao4b 60 28 an an DT cord-048487-kamvao4b 60 29 uniform uniform JJ cord-048487-kamvao4b 60 30 error error NN cord-048487-kamvao4b 61 1 ν ν NNP cord-048487-kamvao4b 61 2 i i NN cord-048487-kamvao4b 61 3 ~ ~ NFP cord-048487-kamvao4b 61 4 U u NN cord-048487-kamvao4b 61 5 [ [ -LRB- cord-048487-kamvao4b 61 6 0 0 NFP cord-048487-kamvao4b 61 7 , , , cord-048487-kamvao4b 61 8 1 1 CD cord-048487-kamvao4b 61 9 ] ] -RRB- cord-048487-kamvao4b 61 10 underlying underlie VBG cord-048487-kamvao4b 61 11 each each DT cord-048487-kamvao4b 61 12 t t NN cord-048487-kamvao4b 61 13 i i PRP cord-048487-kamvao4b 61 14 over over IN cord-048487-kamvao4b 61 15 the the DT cord-048487-kamvao4b 61 16 unit unit NN cord-048487-kamvao4b 61 17 sampling sampling NN cord-048487-kamvao4b 61 18 interval interval NN cord-048487-kamvao4b 61 19 . . . cord-048487-kamvao4b 62 1 To to TO cord-048487-kamvao4b 62 2 complete complete VB cord-048487-kamvao4b 62 3 the the DT cord-048487-kamvao4b 62 4 error error NN cord-048487-kamvao4b 62 5 structure structure NN cord-048487-kamvao4b 62 6 specification specification NN cord-048487-kamvao4b 62 7 we -PRON- PRP cord-048487-kamvao4b 62 8 force force VBP cord-048487-kamvao4b 62 9 the the DT cord-048487-kamvao4b 62 10 two two CD cord-048487-kamvao4b 62 11 forms form NNS cord-048487-kamvao4b 62 12 of of IN cord-048487-kamvao4b 62 13 error error NN cord-048487-kamvao4b 62 14 ( ( -LRB- cord-048487-kamvao4b 62 15 ν ν NNP cord-048487-kamvao4b 62 16 i i NNP cord-048487-kamvao4b 62 17 and and CC cord-048487-kamvao4b 62 18 ε ε NNP cord-048487-kamvao4b 62 19 ij ij NNP cord-048487-kamvao4b 62 20 ) ) -RRB- cord-048487-kamvao4b 62 21 to to TO cord-048487-kamvao4b 62 22 be be VB cord-048487-kamvao4b 62 23 independent independent JJ cord-048487-kamvao4b 62 24 . . . cord-048487-kamvao4b 63 1 Finally finally RB cord-048487-kamvao4b 63 2 , , , cord-048487-kamvao4b 63 3 for for IN cord-048487-kamvao4b 63 4 a a DT cord-048487-kamvao4b 63 5 set set NN cord-048487-kamvao4b 63 6 of of IN cord-048487-kamvao4b 63 7 three three CD cord-048487-kamvao4b 63 8 sequences sequence NNS cord-048487-kamvao4b 63 9 ( ( -LRB- cord-048487-kamvao4b 63 10 s s NNP cord-048487-kamvao4b 63 11 i i NNP cord-048487-kamvao4b 63 12 , , , cord-048487-kamvao4b 63 13 s s NNP cord-048487-kamvao4b 63 14 j j NNP cord-048487-kamvao4b 63 15 , , , cord-048487-kamvao4b 63 16 s s NNP cord-048487-kamvao4b 63 17 k k NNP cord-048487-kamvao4b 63 18 ) ) -RRB- cord-048487-kamvao4b 63 19 and and CC cord-048487-kamvao4b 63 20 their -PRON- PRP$ cord-048487-kamvao4b 63 21 associated associate VBN cord-048487-kamvao4b 63 22 pairwise pairwise JJ cord-048487-kamvao4b 63 23 distances distance NNS cord-048487-kamvao4b 63 24 , , , cord-048487-kamvao4b 63 25 we -PRON- PRP cord-048487-kamvao4b 63 26 enforce enforce VBP cord-048487-kamvao4b 63 27 a a DT cord-048487-kamvao4b 63 28 fixed fix VBN cord-048487-kamvao4b 63 29 topology topology NN cord-048487-kamvao4b 63 30 among among IN cord-048487-kamvao4b 63 31 sequences sequence NNS cord-048487-kamvao4b 63 32 , , , cord-048487-kamvao4b 63 33 as as IN cord-048487-kamvao4b 63 34 shown show VBN cord-048487-kamvao4b 63 35 in in IN cord-048487-kamvao4b 63 36 Figure figure NN cord-048487-kamvao4b 63 37 1 1 CD cord-048487-kamvao4b 63 38 , , , cord-048487-kamvao4b 63 39 via via IN cord-048487-kamvao4b 63 40 methods method NNS cord-048487-kamvao4b 63 41 outlined outline VBN cord-048487-kamvao4b 63 42 in in IN cord-048487-kamvao4b 63 43 [ [ -LRB- cord-048487-kamvao4b 63 44 26 26 CD cord-048487-kamvao4b 63 45 ] ] -RRB- cord-048487-kamvao4b 63 46 . . . cord-048487-kamvao4b 64 1 We -PRON- PRP cord-048487-kamvao4b 64 2 augment augment VBP cord-048487-kamvao4b 64 3 the the DT cord-048487-kamvao4b 64 4 topology topology NN cord-048487-kamvao4b 64 5 with with IN cord-048487-kamvao4b 64 6 the the DT cord-048487-kamvao4b 64 7 observed observed JJ cord-048487-kamvao4b 64 8 sampling sampling NN cord-048487-kamvao4b 64 9 times time NNS cord-048487-kamvao4b 64 10 of of IN cord-048487-kamvao4b 64 11 the the DT cord-048487-kamvao4b 64 12 three three CD cord-048487-kamvao4b 64 13 sequences sequence NNS cord-048487-kamvao4b 64 14 , , , cord-048487-kamvao4b 64 15 α α NN cord-048487-kamvao4b 64 16 , , , cord-048487-kamvao4b 64 17 the the DT cord-048487-kamvao4b 64 18 divergence divergence NN cord-048487-kamvao4b 64 19 time time NN cord-048487-kamvao4b 64 20 between between IN cord-048487-kamvao4b 64 21 the the DT cord-048487-kamvao4b 64 22 two two CD cord-048487-kamvao4b 64 23 sequences sequence NNS cord-048487-kamvao4b 64 24 of of IN cord-048487-kamvao4b 64 25 interest interest NN cord-048487-kamvao4b 64 26 and and CC cord-048487-kamvao4b 64 27 β β NN cord-048487-kamvao4b 64 28 , , , cord-048487-kamvao4b 64 29 the the DT cord-048487-kamvao4b 64 30 divergence divergence NN cord-048487-kamvao4b 64 31 time time NN cord-048487-kamvao4b 64 32 of of IN cord-048487-kamvao4b 64 33 all all DT cord-048487-kamvao4b 64 34 sequences sequence NNS cord-048487-kamvao4b 64 35 . . . cord-048487-kamvao4b 65 1 When when WRB cord-048487-kamvao4b 65 2 necessary necessary JJ cord-048487-kamvao4b 65 3 for for IN cord-048487-kamvao4b 65 4 clarity clarity NN cord-048487-kamvao4b 65 5 , , , cord-048487-kamvao4b 65 6 we -PRON- PRP cord-048487-kamvao4b 65 7 write write VBP cord-048487-kamvao4b 65 8 α α NNP cord-048487-kamvao4b 65 9 ij ij NN cord-048487-kamvao4b 65 10 to to TO cord-048487-kamvao4b 65 11 indicate indicate VB cord-048487-kamvao4b 65 12 the the DT cord-048487-kamvao4b 65 13 true true JJ cord-048487-kamvao4b 65 14 time time NN cord-048487-kamvao4b 65 15 of of IN cord-048487-kamvao4b 65 16 divergence divergence NN cord-048487-kamvao4b 65 17 between between IN cord-048487-kamvao4b 65 18 sequences sequence NNS cord-048487-kamvao4b 66 1 i i PRP cord-048487-kamvao4b 66 2 and and CC cord-048487-kamvao4b 66 3 j. j. NNP cord-048487-kamvao4b 66 4 Under under IN cord-048487-kamvao4b 66 5 our -PRON- PRP$ cord-048487-kamvao4b 66 6 triplet triplet NN cord-048487-kamvao4b 66 7 method method NN cord-048487-kamvao4b 66 8 , , , cord-048487-kamvao4b 66 9 we -PRON- PRP cord-048487-kamvao4b 66 10 aim aim VBP cord-048487-kamvao4b 66 11 to to TO cord-048487-kamvao4b 66 12 estimate estimate VB cord-048487-kamvao4b 66 13 the the DT cord-048487-kamvao4b 66 14 true true JJ cord-048487-kamvao4b 66 15 rate rate NN cord-048487-kamvao4b 66 16 of of IN cord-048487-kamvao4b 66 17 nucleotide nucleotide JJ cord-048487-kamvao4b 66 18 substitution substitution NN cord-048487-kamvao4b 66 19 , , , cord-048487-kamvao4b 66 20 p p NN cord-048487-kamvao4b 66 21 ij ij NN cord-048487-kamvao4b 66 22 , , , cord-048487-kamvao4b 66 23 between between IN cord-048487-kamvao4b 66 24 sequences sequence NNS cord-048487-kamvao4b 66 25 s s NNP cord-048487-kamvao4b 66 26 i i NNP cord-048487-kamvao4b 66 27 and and CC cord-048487-kamvao4b 66 28 s s NNP cord-048487-kamvao4b 66 29 j j NNP cord-048487-kamvao4b 66 30 with with IN cord-048487-kamvao4b 66 31 an an DT cord-048487-kamvao4b 66 32 unobserved unobserved JJ cord-048487-kamvao4b 66 33 error error NN cord-048487-kamvao4b 66 34 δ δ NN cord-048487-kamvao4b 66 35 ij ij NN cord-048487-kamvao4b 66 36 . . . cord-048487-kamvao4b 67 1 With with IN cord-048487-kamvao4b 67 2 respect respect NN cord-048487-kamvao4b 67 3 to to IN cord-048487-kamvao4b 67 4 outgroup outgroup NN cord-048487-kamvao4b 67 5 sequence sequence NN cord-048487-kamvao4b 68 1 k k NNP cord-048487-kamvao4b 68 2 , , , cord-048487-kamvao4b 68 3 an an DT cord-048487-kamvao4b 68 4 unbiased unbiased JJ cord-048487-kamvao4b 68 5 estimate estimate NN cord-048487-kamvao4b 68 6 is be VBZ cord-048487-kamvao4b 68 7 where where WRB cord-048487-kamvao4b 68 8 the the DT cord-048487-kamvao4b 68 9 factor factor NN cord-048487-kamvao4b 68 10 corrects correct VBZ cord-048487-kamvao4b 68 11 for for IN cord-048487-kamvao4b 68 12 bias bias NN cord-048487-kamvao4b 68 13 resulting result VBG cord-048487-kamvao4b 68 14 from from IN cord-048487-kamvao4b 68 15 the the DT cord-048487-kamvao4b 68 16 time time NN cord-048487-kamvao4b 68 17 sampling sampling NN cord-048487-kamvao4b 68 18 error error NN cord-048487-kamvao4b 68 19 structure structure NN cord-048487-kamvao4b 68 20 ( ( -LRB- cord-048487-kamvao4b 68 21 see see VB cord-048487-kamvao4b 68 22 Appendix Appendix NNP cord-048487-kamvao4b 68 23 for for IN cord-048487-kamvao4b 68 24 derivation derivation NN cord-048487-kamvao4b 68 25 ) ) -RRB- cord-048487-kamvao4b 68 26 . . . cord-048487-kamvao4b 69 1 We -PRON- PRP cord-048487-kamvao4b 69 2 superscript superscript VBP cord-048487-kamvao4b 69 3 to to TO cord-048487-kamvao4b 69 4 denote denote VB cord-048487-kamvao4b 69 5 its -PRON- PRP$ cord-048487-kamvao4b 69 6 weak weak JJ cord-048487-kamvao4b 69 7 dependence dependence NN cord-048487-kamvao4b 69 8 on on IN cord-048487-kamvao4b 69 9 outgroup outgroup NN cord-048487-kamvao4b 69 10 sequence sequence NN cord-048487-kamvao4b 69 11 k. k. NNP cord-048487-kamvao4b 69 12 Dependence Dependence NNP cord-048487-kamvao4b 69 13 is be VBZ cord-048487-kamvao4b 69 14 weak weak JJ cord-048487-kamvao4b 69 15 as as IN cord-048487-kamvao4b 69 16 the the DT cord-048487-kamvao4b 69 17 path path NN cord-048487-kamvao4b 69 18 of of IN cord-048487-kamvao4b 69 19 evolution evolution NN cord-048487-kamvao4b 69 20 from from IN cord-048487-kamvao4b 69 21 t t NNP cord-048487-kamvao4b 69 22 k k NNP cord-048487-kamvao4b 69 23 to to IN cord-048487-kamvao4b 69 24 α α NNP cord-048487-kamvao4b 69 25 is be VBZ cord-048487-kamvao4b 69 26 shared share VBN cord-048487-kamvao4b 69 27 between between IN cord-048487-kamvao4b 69 28 the the DT cord-048487-kamvao4b 69 29 paths path NNS cord-048487-kamvao4b 69 30 from from IN cord-048487-kamvao4b 69 31 sequence sequence NN cord-048487-kamvao4b 69 32 k k NN cord-048487-kamvao4b 69 33 to to IN cord-048487-kamvao4b 69 34 both both DT cord-048487-kamvao4b 69 35 sequence sequence NN cord-048487-kamvao4b 69 36 i i PRP cord-048487-kamvao4b 69 37 and and CC cord-048487-kamvao4b 69 38 sequence sequence NN cord-048487-kamvao4b 69 39 j j NNP cord-048487-kamvao4b 69 40 and and CC cord-048487-kamvao4b 69 41 hence hence RB cord-048487-kamvao4b 69 42 largely largely RB cord-048487-kamvao4b 69 43 cancels cancel VBZ cord-048487-kamvao4b 69 44 out out RP cord-048487-kamvao4b 69 45 in in IN cord-048487-kamvao4b 69 46 Equation equation NN cord-048487-kamvao4b 69 47 1 1 CD cord-048487-kamvao4b 69 48 . . . cord-048487-kamvao4b 70 1 We -PRON- PRP cord-048487-kamvao4b 70 2 make make VBP cord-048487-kamvao4b 70 3 this this DT cord-048487-kamvao4b 70 4 transparent transparent JJ cord-048487-kamvao4b 70 5 in in IN cord-048487-kamvao4b 70 6 the the DT cord-048487-kamvao4b 70 7 following follow VBG cord-048487-kamvao4b 70 8 derivation derivation NN cord-048487-kamvao4b 70 9 . . . cord-048487-kamvao4b 71 1 For for IN cord-048487-kamvao4b 71 2 brevity brevity NN cord-048487-kamvao4b 71 3 , , , cord-048487-kamvao4b 71 4 we -PRON- PRP cord-048487-kamvao4b 71 5 consider consider VBP cord-048487-kamvao4b 71 6 only only RB cord-048487-kamvao4b 71 7 unobservable unobservable JJ cord-048487-kamvao4b 71 8 true true JJ cord-048487-kamvao4b 71 9 values value NNS cord-048487-kamvao4b 71 10 , , , cord-048487-kamvao4b 71 11 ignoring ignore VBG cord-048487-kamvao4b 71 12 error error NN cord-048487-kamvao4b 71 13 terms term NNS cord-048487-kamvao4b 71 14 . . . cord-048487-kamvao4b 72 1 Let let VB cord-048487-kamvao4b 72 2 u u PRP cord-048487-kamvao4b 72 3 be be VB cord-048487-kamvao4b 72 4 the the DT cord-048487-kamvao4b 72 5 location location NN cord-048487-kamvao4b 72 6 on on IN cord-048487-kamvao4b 72 7 the the DT cord-048487-kamvao4b 72 8 triplet triplet NN cord-048487-kamvao4b 72 9 in in IN cord-048487-kamvao4b 72 10 Figure figure NN cord-048487-kamvao4b 72 11 1 1 CD cord-048487-kamvao4b 72 12 corresponding correspond VBG cord-048487-kamvao4b 72 13 to to IN cord-048487-kamvao4b 72 14 time time NN cord-048487-kamvao4b 72 15 α α NN cord-048487-kamvao4b 72 16 and and CC cord-048487-kamvao4b 72 17 let let VB cord-048487-kamvao4b 72 18 p p NN cord-048487-kamvao4b 72 19 xy xy PRP cord-048487-kamvao4b 72 20 be be VB cord-048487-kamvao4b 72 21 the the DT cord-048487-kamvao4b 72 22 true true JJ cord-048487-kamvao4b 72 23 rate rate NN cord-048487-kamvao4b 72 24 along along IN cord-048487-kamvao4b 72 25 the the DT cord-048487-kamvao4b 72 26 path path NN cord-048487-kamvao4b 72 27 connecting connect VBG cord-048487-kamvao4b 72 28 locations location NNS cord-048487-kamvao4b 73 1 x x LS cord-048487-kamvao4b 74 1 and and CC cord-048487-kamvao4b 74 2 y. y. NNP cord-048487-kamvao4b 75 1 Then then RB cord-048487-kamvao4b 75 2 , , , cord-048487-kamvao4b 75 3 as as IN cord-048487-kamvao4b 75 4 distance distance NN cord-048487-kamvao4b 75 5 is be VBZ cord-048487-kamvao4b 75 6 rate rate NN cord-048487-kamvao4b 75 7 multiplied multiply VBN cord-048487-kamvao4b 75 8 by by IN cord-048487-kamvao4b 75 9 time time NN cord-048487-kamvao4b 75 10 , , , cord-048487-kamvao4b 75 11 we -PRON- PRP cord-048487-kamvao4b 75 12 have have VBP cord-048487-kamvao4b 75 13 Subtracting subtract VBG cord-048487-kamvao4b 75 14 the the DT cord-048487-kamvao4b 75 15 first first JJ cord-048487-kamvao4b 75 16 equation equation NN cord-048487-kamvao4b 75 17 from from IN cord-048487-kamvao4b 75 18 the the DT cord-048487-kamvao4b 75 19 second second JJ cord-048487-kamvao4b 75 20 equation equation NN cord-048487-kamvao4b 75 21 , , , cord-048487-kamvao4b 75 22 which which WDT cord-048487-kamvao4b 75 23 is be VBZ cord-048487-kamvao4b 75 24 equivalent equivalent JJ cord-048487-kamvao4b 75 25 to to IN cord-048487-kamvao4b 75 26 Equation equation NN cord-048487-kamvao4b 75 27 1 1 CD cord-048487-kamvao4b 75 28 . . . cord-048487-kamvao4b 76 1 This this DT cord-048487-kamvao4b 76 2 derivation derivation NN cord-048487-kamvao4b 76 3 makes make VBZ cord-048487-kamvao4b 76 4 clear clear JJ cord-048487-kamvao4b 76 5 that that IN cord-048487-kamvao4b 76 6 the the DT cord-048487-kamvao4b 76 7 estimator estimator NN cord-048487-kamvao4b 76 8 ( ( -LRB- cord-048487-kamvao4b 76 9 1 1 CD cord-048487-kamvao4b 76 10 ) ) -RRB- cord-048487-kamvao4b 76 11 measures measure VBZ cord-048487-kamvao4b 76 12 the the DT cord-048487-kamvao4b 76 13 rate rate NN cord-048487-kamvao4b 76 14 along along IN cord-048487-kamvao4b 76 15 the the DT cord-048487-kamvao4b 76 16 path path NN cord-048487-kamvao4b 76 17 from from IN cord-048487-kamvao4b 76 18 sequence sequence NN cord-048487-kamvao4b 76 19 i i PRP cord-048487-kamvao4b 76 20 to to TO cord-048487-kamvao4b 76 21 sequence sequence VB cord-048487-kamvao4b 76 22 j j NNP cord-048487-kamvao4b 76 23 , , , cord-048487-kamvao4b 76 24 with with IN cord-048487-kamvao4b 76 25 only only JJ cord-048487-kamvao4b 76 26 incidental incidental JJ cord-048487-kamvao4b 76 27 dependence dependence NN cord-048487-kamvao4b 76 28 on on IN cord-048487-kamvao4b 76 29 sequence sequence NN cord-048487-kamvao4b 77 1 k. k. NNP cord-048487-kamvao4b 78 1 The the DT cord-048487-kamvao4b 78 2 variance variance NN cord-048487-kamvao4b 78 3 for for IN cord-048487-kamvao4b 78 4 the the DT cord-048487-kamvao4b 78 5 estimator estimator NN cord-048487-kamvao4b 78 6 ( ( -LRB- cord-048487-kamvao4b 78 7 1 1 CD cord-048487-kamvao4b 78 8 ) ) -RRB- cord-048487-kamvao4b 78 9 is be VBZ cord-048487-kamvao4b 78 10 well well RB cord-048487-kamvao4b 78 11 approximated approximate VBN cord-048487-kamvao4b 78 12 by by IN cord-048487-kamvao4b 78 13 Further further RB cord-048487-kamvao4b 78 14 , , , cord-048487-kamvao4b 78 15 we -PRON- PRP cord-048487-kamvao4b 78 16 can can MD cord-048487-kamvao4b 78 17 estimate estimate VB cord-048487-kamvao4b 78 18 the the DT cord-048487-kamvao4b 78 19 time time NN cord-048487-kamvao4b 78 20 of of IN cord-048487-kamvao4b 78 21 subtype subtype NN cord-048487-kamvao4b 78 22 divergence divergence NN cord-048487-kamvao4b 78 23 α α NNP cord-048487-kamvao4b 78 24 ( ( -LRB- cord-048487-kamvao4b 78 25 Figure figure NN cord-048487-kamvao4b 78 26 1 1 CD cord-048487-kamvao4b 78 27 ) ) -RRB- cord-048487-kamvao4b 78 28 between between IN cord-048487-kamvao4b 78 29 sequences sequence NNS cord-048487-kamvao4b 78 30 via via IN cord-048487-kamvao4b 79 1 We -PRON- PRP cord-048487-kamvao4b 79 2 note note VBP cord-048487-kamvao4b 79 3 that that IN cord-048487-kamvao4b 79 4 the the DT cord-048487-kamvao4b 79 5 term term NN cord-048487-kamvao4b 79 6 t t NN cord-048487-kamvao4b 80 1 i i PRP cord-048487-kamvao4b 80 2 + + CC cord-048487-kamvao4b 81 1 t t NNP cord-048487-kamvao4b 81 2 j j NNP cord-048487-kamvao4b 81 3 -1 -1 NNP cord-048487-kamvao4b 81 4 is be VBZ cord-048487-kamvao4b 81 5 used use VBN cord-048487-kamvao4b 81 6 rather rather RB cord-048487-kamvao4b 81 7 than than IN cord-048487-kamvao4b 82 1 t t NNP cord-048487-kamvao4b 82 2 i i NNP cord-048487-kamvao4b 82 3 + + CC cord-048487-kamvao4b 82 4 t t NNP cord-048487-kamvao4b 82 5 j j NNP cord-048487-kamvao4b 82 6 to to TO cord-048487-kamvao4b 82 7 account account VB cord-048487-kamvao4b 82 8 for for IN cord-048487-kamvao4b 82 9 the the DT cord-048487-kamvao4b 82 10 expected expected JJ cord-048487-kamvao4b 82 11 error error NN cord-048487-kamvao4b 82 12 coming come VBG cord-048487-kamvao4b 82 13 from from IN cord-048487-kamvao4b 82 14 the the DT cord-048487-kamvao4b 82 15 uniformly uniformly RB cord-048487-kamvao4b 82 16 The the DT cord-048487-kamvao4b 82 17 phylogenetic phylogenetic JJ cord-048487-kamvao4b 82 18 relationships relationship NNS cord-048487-kamvao4b 82 19 between between IN cord-048487-kamvao4b 82 20 three three CD cord-048487-kamvao4b 82 21 sequences sequence NNS cord-048487-kamvao4b 82 22 s s NNP cord-048487-kamvao4b 82 23 i i NNP cord-048487-kamvao4b 82 24 , , , cord-048487-kamvao4b 82 25 s s NNP cord-048487-kamvao4b 82 26 j j NNP cord-048487-kamvao4b 82 27 and and CC cord-048487-kamvao4b 82 28 s s NNP cord-048487-kamvao4b 82 29 k k XX cord-048487-kamvao4b 82 30 , , , cord-048487-kamvao4b 82 31 sampled sample VBN cord-048487-kamvao4b 82 32 on on IN cord-048487-kamvao4b 82 33 dates date NNS cord-048487-kamvao4b 82 34 t t NNP cord-048487-kamvao4b 82 35 i i NNP cord-048487-kamvao4b 82 36 , , , cord-048487-kamvao4b 82 37 t t NNP cord-048487-kamvao4b 82 38 j j NNP cord-048487-kamvao4b 82 39 and and CC cord-048487-kamvao4b 82 40 t t NNP cord-048487-kamvao4b 82 41 k k NNP cord-048487-kamvao4b 82 42 respectively respectively RB cord-048487-kamvao4b 82 43 As as IN cord-048487-kamvao4b 82 44 nucleotide nucleotide JJ cord-048487-kamvao4b 82 45 data data NN cord-048487-kamvao4b 82 46 increases increase NNS cord-048487-kamvao4b 82 47 without without IN cord-048487-kamvao4b 82 48 bound bind VBN cord-048487-kamvao4b 82 49 , , , cord-048487-kamvao4b 82 50 K K NNP cord-048487-kamvao4b 82 51 ij ij NN cord-048487-kamvao4b 82 52 → → SYM cord-048487-kamvao4b 82 53 D d NN cord-048487-kamvao4b 82 54 ij ij NN cord-048487-kamvao4b 82 55 and and CC cord-048487-kamvao4b 82 56 → → SYM cord-048487-kamvao4b 82 57 p p NN cord-048487-kamvao4b 82 58 ij ij NN cord-048487-kamvao4b 82 59 , , , cord-048487-kamvao4b 82 60 ensuring ensure VBG cord-048487-kamvao4b 82 61 that that IN cord-048487-kamvao4b 82 62 → → SYM cord-048487-kamvao4b 82 63 α α NN cord-048487-kamvao4b 82 64 ij ij NN cord-048487-kamvao4b 82 65 . . . cord-048487-kamvao4b 83 1 For for IN cord-048487-kamvao4b 83 2 finite finite JJ cord-048487-kamvao4b 83 3 sequence sequence NN cord-048487-kamvao4b 83 4 lengths length NNS cord-048487-kamvao4b 83 5 , , , cord-048487-kamvao4b 83 6 this this DT cord-048487-kamvao4b 83 7 relation relation NN cord-048487-kamvao4b 83 8 ensures ensure VBZ cord-048487-kamvao4b 83 9 that that DT cord-048487-kamvao4b 83 10 . . . cord-048487-kamvao4b 84 1 To to TO cord-048487-kamvao4b 84 2 gain gain VB cord-048487-kamvao4b 84 3 an an DT cord-048487-kamvao4b 84 4 understanding understanding NN cord-048487-kamvao4b 84 5 of of IN cord-048487-kamvao4b 84 6 this this DT cord-048487-kamvao4b 84 7 estimator estimator NN cord-048487-kamvao4b 84 8 , , , cord-048487-kamvao4b 84 9 we -PRON- PRP cord-048487-kamvao4b 84 10 note note VBP cord-048487-kamvao4b 84 11 that that IN cord-048487-kamvao4b 84 12 with with IN cord-048487-kamvao4b 84 13 a a DT cord-048487-kamvao4b 84 14 standard standard JJ cord-048487-kamvao4b 84 15 model model NN cord-048487-kamvao4b 84 16 of of IN cord-048487-kamvao4b 84 17 substitution substitution NN cord-048487-kamvao4b 84 18 ( ( -LRB- cord-048487-kamvao4b 84 19 e.g. e.g. RB cord-048487-kamvao4b 84 20 JC69 JC69 NNP cord-048487-kamvao4b 84 21 , , , cord-048487-kamvao4b 84 22 HKY85 HKY85 NNP cord-048487-kamvao4b 84 23 ) ) -RRB- cord-048487-kamvao4b 84 24 , , , cord-048487-kamvao4b 84 25 a a DT cord-048487-kamvao4b 84 26 rate rate NN cord-048487-kamvao4b 84 27 of of IN cord-048487-kamvao4b 84 28 substitution substitution NN cord-048487-kamvao4b 84 29 of of IN cord-048487-kamvao4b 84 30 10 10 CD cord-048487-kamvao4b 84 31 -4 -4 NNS cord-048487-kamvao4b 84 32 ( ( -LRB- cord-048487-kamvao4b 84 33 s s NNP cord-048487-kamvao4b 84 34 / / SYM cord-048487-kamvao4b 84 35 s s NNP cord-048487-kamvao4b 84 36 / / SYM cord-048487-kamvao4b 84 37 yr yr NN cord-048487-kamvao4b 84 38 ) ) -RRB- cord-048487-kamvao4b 84 39 and and CC cord-048487-kamvao4b 84 40 a a DT cord-048487-kamvao4b 84 41 sequence sequence NN cord-048487-kamvao4b 84 42 of of IN cord-048487-kamvao4b 84 43 2000 2000 CD cord-048487-kamvao4b 84 44 nucleotides nucleotide NNS cord-048487-kamvao4b 84 45 , , , cord-048487-kamvao4b 84 46 the the DT cord-048487-kamvao4b 84 47 above above JJ cord-048487-kamvao4b 84 48 estimator estimator NN cord-048487-kamvao4b 84 49 yields yield VBZ cord-048487-kamvao4b 84 50 a a DT cord-048487-kamvao4b 84 51 standard standard JJ cord-048487-kamvao4b 84 52 error error NN cord-048487-kamvao4b 84 53 of of IN cord-048487-kamvao4b 84 54 approximately approximately RB cord-048487-kamvao4b 84 55 23 23 CD cord-048487-kamvao4b 84 56 years year NNS cord-048487-kamvao4b 84 57 [ [ -LRB- cord-048487-kamvao4b 84 58 20 20 CD cord-048487-kamvao4b 84 59 ] ] -RRB- cord-048487-kamvao4b 84 60 . . . cord-048487-kamvao4b 85 1 The the DT cord-048487-kamvao4b 85 2 above above JJ cord-048487-kamvao4b 85 3 derivations derivation NNS cord-048487-kamvao4b 85 4 express express VBP cord-048487-kamvao4b 85 5 our -PRON- PRP$ cord-048487-kamvao4b 85 6 rate rate NN cord-048487-kamvao4b 85 7 and and CC cord-048487-kamvao4b 85 8 time time NN cord-048487-kamvao4b 85 9 estimates estimate NNS cord-048487-kamvao4b 85 10 for for IN cord-048487-kamvao4b 85 11 a a DT cord-048487-kamvao4b 85 12 single single JJ cord-048487-kamvao4b 85 13 triplet triplet NN cord-048487-kamvao4b 85 14 of of IN cord-048487-kamvao4b 85 15 sequences sequence NNS cord-048487-kamvao4b 85 16 . . . cord-048487-kamvao4b 86 1 We -PRON- PRP cord-048487-kamvao4b 86 2 now now RB cord-048487-kamvao4b 86 3 consider consider VBP cord-048487-kamvao4b 86 4 estimates estimate NNS cord-048487-kamvao4b 86 5 that that WDT cord-048487-kamvao4b 86 6 combine combine VBP cord-048487-kamvao4b 86 7 information information NN cord-048487-kamvao4b 86 8 across across IN cord-048487-kamvao4b 86 9 multiple multiple JJ cord-048487-kamvao4b 86 10 representative representative JJ cord-048487-kamvao4b 86 11 sequences sequence NNS cord-048487-kamvao4b 86 12 from from IN cord-048487-kamvao4b 86 13 each each DT cord-048487-kamvao4b 86 14 subtype subtype NN cord-048487-kamvao4b 86 15 . . . cord-048487-kamvao4b 87 1 For for IN cord-048487-kamvao4b 87 2 discussion discussion NN cord-048487-kamvao4b 87 3 , , , cord-048487-kamvao4b 87 4 we -PRON- PRP cord-048487-kamvao4b 87 5 label label VBP cord-048487-kamvao4b 87 6 subtypes subtype VBZ cord-048487-kamvao4b 87 7 A A NNP cord-048487-kamvao4b 87 8 , , , cord-048487-kamvao4b 87 9 B b NN cord-048487-kamvao4b 87 10 and and CC cord-048487-kamvao4b 87 11 C C NNP cord-048487-kamvao4b 87 12 ( ( -LRB- cord-048487-kamvao4b 87 13 which which WDT cord-048487-kamvao4b 87 14 are be VBP cord-048487-kamvao4b 87 15 only only RB cord-048487-kamvao4b 87 16 incidentally incidentally RB cord-048487-kamvao4b 87 17 the the DT cord-048487-kamvao4b 87 18 same same JJ cord-048487-kamvao4b 87 19 as as IN cord-048487-kamvao4b 87 20 the the DT cord-048487-kamvao4b 87 21 labels label NNS cord-048487-kamvao4b 87 22 for for IN cord-048487-kamvao4b 87 23 influenza influenza NN cord-048487-kamvao4b 87 24 ) ) -RRB- cord-048487-kamvao4b 88 1 and and CC cord-048487-kamvao4b 88 2 we -PRON- PRP cord-048487-kamvao4b 88 3 assume assume VBP cord-048487-kamvao4b 88 4 the the DT cord-048487-kamvao4b 88 5 topology topology NN cord-048487-kamvao4b 88 6 in in IN cord-048487-kamvao4b 88 7 Figure figure NN cord-048487-kamvao4b 88 8 1 1 CD cord-048487-kamvao4b 88 9 for for IN cord-048487-kamvao4b 88 10 these these DT cord-048487-kamvao4b 88 11 groups group NNS cord-048487-kamvao4b 88 12 . . . cord-048487-kamvao4b 89 1 We -PRON- PRP cord-048487-kamvao4b 89 2 let let VBP cord-048487-kamvao4b 89 3 n n NN cord-048487-kamvao4b 89 4 r r NN cord-048487-kamvao4b 89 5 , , , cord-048487-kamvao4b 89 6 where where WRB cord-048487-kamvao4b 89 7 r r NN cord-048487-kamvao4b 89 8 ∈ ∈ NN cord-048487-kamvao4b 89 9 { { -LRB- cord-048487-kamvao4b 89 10 A a NN cord-048487-kamvao4b 89 11 , , , cord-048487-kamvao4b 89 12 where where WRB cord-048487-kamvao4b 89 13 P p NN cord-048487-kamvao4b 89 14 α α NN cord-048487-kamvao4b 89 15 is be VBZ cord-048487-kamvao4b 89 16 the the DT cord-048487-kamvao4b 89 17 sum sum NN cord-048487-kamvao4b 89 18 of of IN cord-048487-kamvao4b 89 19 the the DT cord-048487-kamvao4b 89 20 inverse inverse JJ cord-048487-kamvao4b 89 21 variance variance NN cord-048487-kamvao4b 89 22 of of IN cord-048487-kamvao4b 89 23 each each DT cord-048487-kamvao4b 89 24 estimate estimate NN cord-048487-kamvao4b 89 25 , , , cord-048487-kamvao4b 89 26 . . . cord-048487-kamvao4b 90 1 Having have VBG cord-048487-kamvao4b 90 2 found find VBN cord-048487-kamvao4b 90 3 , , , cord-048487-kamvao4b 90 4 we -PRON- PRP cord-048487-kamvao4b 90 5 estimate estimate VBP cord-048487-kamvao4b 90 6 its -PRON- PRP$ cord-048487-kamvao4b 90 7 variance variance NN cord-048487-kamvao4b 90 8 by by IN cord-048487-kamvao4b 90 9 a a DT cord-048487-kamvao4b 90 10 bootstrap bootstrap NN cord-048487-kamvao4b 90 11 resampling resampling NN cord-048487-kamvao4b 90 12 of of IN cord-048487-kamvao4b 90 13 sequences sequence NNS cord-048487-kamvao4b 90 14 from from IN cord-048487-kamvao4b 90 15 each each DT cord-048487-kamvao4b 90 16 subtype subtype NN cord-048487-kamvao4b 90 17 [ [ -LRB- cord-048487-kamvao4b 90 18 8 8 CD cord-048487-kamvao4b 90 19 ] ] -RRB- cord-048487-kamvao4b 90 20 . . . cord-048487-kamvao4b 91 1 The the DT cord-048487-kamvao4b 91 2 computational computational JJ cord-048487-kamvao4b 91 3 efficiency efficiency NN cord-048487-kamvao4b 91 4 of of IN cord-048487-kamvao4b 91 5 this this DT cord-048487-kamvao4b 91 6 estimator estimator NN cord-048487-kamvao4b 91 7 is be VBZ cord-048487-kamvao4b 91 8 on on IN cord-048487-kamvao4b 91 9 the the DT cord-048487-kamvao4b 91 10 order order NN cord-048487-kamvao4b 91 11 O(n o(n NN cord-048487-kamvao4b 91 12 3 3 CD cord-048487-kamvao4b 91 13 ) ) -RRB- cord-048487-kamvao4b 91 14 for for IN cord-048487-kamvao4b 91 15 a a DT cord-048487-kamvao4b 91 16 tree tree NN cord-048487-kamvao4b 91 17 of of IN cord-048487-kamvao4b 91 18 n n NN cord-048487-kamvao4b 91 19 taxa taxon NNS cord-048487-kamvao4b 91 20 . . . cord-048487-kamvao4b 92 1 This this DT cord-048487-kamvao4b 92 2 is be VBZ cord-048487-kamvao4b 92 3 natural natural JJ cord-048487-kamvao4b 92 4 as as IN cord-048487-kamvao4b 92 5 each each DT cord-048487-kamvao4b 92 6 of of IN cord-048487-kamvao4b 92 7 the the DT cord-048487-kamvao4b 92 8 initial initial JJ cord-048487-kamvao4b 92 9 rate rate NN cord-048487-kamvao4b 92 10 estimates estimate NNS cord-048487-kamvao4b 92 11 is be VBZ cord-048487-kamvao4b 92 12 composed compose VBN cord-048487-kamvao4b 92 13 of of IN cord-048487-kamvao4b 92 14 information information NN cord-048487-kamvao4b 92 15 concerning concern VBG cord-048487-kamvao4b 92 16 three three CD cord-048487-kamvao4b 92 17 taxa taxon NNS cord-048487-kamvao4b 92 18 . . . cord-048487-kamvao4b 93 1 While while IN cord-048487-kamvao4b 93 2 the the DT cord-048487-kamvao4b 93 3 growth growth NN cord-048487-kamvao4b 93 4 of of IN cord-048487-kamvao4b 93 5 computational computational JJ cord-048487-kamvao4b 93 6 expense expense NN cord-048487-kamvao4b 93 7 in in IN cord-048487-kamvao4b 93 8 the the DT cord-048487-kamvao4b 93 9 number number NN cord-048487-kamvao4b 93 10 of of IN cord-048487-kamvao4b 93 11 taxa taxon NNS cord-048487-kamvao4b 93 12 may may MD cord-048487-kamvao4b 93 13 appear appear VB cord-048487-kamvao4b 93 14 unpleasant unpleasant JJ cord-048487-kamvao4b 93 15 , , , cord-048487-kamvao4b 93 16 in in IN cord-048487-kamvao4b 93 17 practice practice NN cord-048487-kamvao4b 93 18 this this DT cord-048487-kamvao4b 93 19 algorithm algorithm NN cord-048487-kamvao4b 93 20 is be VBZ cord-048487-kamvao4b 93 21 both both CC cord-048487-kamvao4b 93 22 fast fast JJ cord-048487-kamvao4b 93 23 and and CC cord-048487-kamvao4b 93 24 stable stable JJ cord-048487-kamvao4b 93 25 , , , cord-048487-kamvao4b 93 26 owing owe VBG cord-048487-kamvao4b 93 27 to to IN cord-048487-kamvao4b 93 28 the the DT cord-048487-kamvao4b 93 29 absence absence NN cord-048487-kamvao4b 93 30 of of IN cord-048487-kamvao4b 93 31 costly costly JJ cord-048487-kamvao4b 93 32 optimization optimization NN cord-048487-kamvao4b 93 33 procedures procedure NNS cord-048487-kamvao4b 93 34 for for IN cord-048487-kamvao4b 93 35 parameter parameter NN cord-048487-kamvao4b 93 36 inference inference NN cord-048487-kamvao4b 93 37 , , , cord-048487-kamvao4b 93 38 and and CC cord-048487-kamvao4b 93 39 is be VBZ cord-048487-kamvao4b 93 40 able able JJ cord-048487-kamvao4b 93 41 to to TO cord-048487-kamvao4b 93 42 handle handle VB cord-048487-kamvao4b 93 43 data data NN cord-048487-kamvao4b 93 44 sets set NNS cord-048487-kamvao4b 93 45 of of IN cord-048487-kamvao4b 93 46 thousands thousand NNS cord-048487-kamvao4b 93 47 of of IN cord-048487-kamvao4b 93 48 taxa taxon NNS cord-048487-kamvao4b 93 49 . . . cord-048487-kamvao4b 94 1 The the DT cord-048487-kamvao4b 94 2 authors author NNS cord-048487-kamvao4b 94 3 detail detail VBP cord-048487-kamvao4b 94 4 the the DT cord-048487-kamvao4b 94 5 computational computational JJ cord-048487-kamvao4b 94 6 efficiency efficiency NN cord-048487-kamvao4b 94 7 of of IN cord-048487-kamvao4b 94 8 a a DT cord-048487-kamvao4b 94 9 similar similar JJ cord-048487-kamvao4b 94 10 statistic statistic NN cord-048487-kamvao4b 94 11 in in IN cord-048487-kamvao4b 94 12 [ [ -LRB- cord-048487-kamvao4b 94 13 26 26 CD cord-048487-kamvao4b 94 14 ] ] -RRB- cord-048487-kamvao4b 94 15 . . . cord-048487-kamvao4b 95 1 As as IN cord-048487-kamvao4b 95 2 an an DT cord-048487-kamvao4b 95 3 example example NN cord-048487-kamvao4b 95 4 , , , cord-048487-kamvao4b 95 5 for for IN cord-048487-kamvao4b 95 6 the the DT cord-048487-kamvao4b 95 7 data datum NNS cord-048487-kamvao4b 95 8 presented present VBN cord-048487-kamvao4b 95 9 below below IN cord-048487-kamvao4b 95 10 all all DT cord-048487-kamvao4b 95 11 computations computation NNS cord-048487-kamvao4b 95 12 required require VBN cord-048487-kamvao4b 95 13 only only RB cord-048487-kamvao4b 95 14 a a DT cord-048487-kamvao4b 95 15 few few JJ cord-048487-kamvao4b 95 16 seconds second NNS cord-048487-kamvao4b 95 17 on on IN cord-048487-kamvao4b 95 18 a a DT cord-048487-kamvao4b 95 19 desktop desktop NN cord-048487-kamvao4b 95 20 computer computer NN cord-048487-kamvao4b 95 21 . . . cord-048487-kamvao4b 96 1 We -PRON- PRP cord-048487-kamvao4b 96 2 demonstrate demonstrate VBP cord-048487-kamvao4b 96 3 the the DT cord-048487-kamvao4b 96 4 advantage advantage NN cord-048487-kamvao4b 96 5 of of IN cord-048487-kamvao4b 96 6 our -PRON- PRP$ cord-048487-kamvao4b 96 7 triplet triplet NN cord-048487-kamvao4b 96 8 estimator estimator NN cord-048487-kamvao4b 96 9 through through IN cord-048487-kamvao4b 96 10 analysis analysis NN cord-048487-kamvao4b 96 11 of of IN cord-048487-kamvao4b 96 12 influenza influenza NN cord-048487-kamvao4b 96 13 A A NNP cord-048487-kamvao4b 96 14 - - : cord-048487-kamvao4b 96 15 H3N2 H3N2 NNP cord-048487-kamvao4b 96 16 / / SYM cord-048487-kamvao4b 96 17 B b NN cord-048487-kamvao4b 96 18 subtype subtype NN cord-048487-kamvao4b 96 19 divergence divergence NN cord-048487-kamvao4b 96 20 using use VBG cord-048487-kamvao4b 96 21 the the DT cord-048487-kamvao4b 96 22 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 96 23 ( ( -LRB- cord-048487-kamvao4b 96 24 HA HA NNP cord-048487-kamvao4b 96 25 ) ) -RRB- cord-048487-kamvao4b 96 26 , , , cord-048487-kamvao4b 96 27 matrix matrix NN cord-048487-kamvao4b 96 28 protein protein NN cord-048487-kamvao4b 96 29 ( ( -LRB- cord-048487-kamvao4b 96 30 MP MP NNP cord-048487-kamvao4b 96 31 ) ) -RRB- cord-048487-kamvao4b 96 32 and and CC cord-048487-kamvao4b 96 33 non non JJ cord-048487-kamvao4b 96 34 - - JJ cord-048487-kamvao4b 96 35 structural structural JJ cord-048487-kamvao4b 96 36 ( ( -LRB- cord-048487-kamvao4b 96 37 NS NS NNP cord-048487-kamvao4b 96 38 ) ) -RRB- cord-048487-kamvao4b 96 39 genes gene NNS cord-048487-kamvao4b 96 40 . . . cord-048487-kamvao4b 97 1 Each each DT cord-048487-kamvao4b 97 2 analysis analysis NN cord-048487-kamvao4b 97 3 is be VBZ cord-048487-kamvao4b 97 4 performed perform VBN cord-048487-kamvao4b 97 5 on on IN cord-048487-kamvao4b 97 6 60 60 CD cord-048487-kamvao4b 97 7 gene gene NN cord-048487-kamvao4b 97 8 sequences sequence NNS cord-048487-kamvao4b 97 9 constructed construct VBN cord-048487-kamvao4b 97 10 from from IN cord-048487-kamvao4b 97 11 20 20 CD cord-048487-kamvao4b 97 12 genomes genome NNS cord-048487-kamvao4b 97 13 each each DT cord-048487-kamvao4b 97 14 drawn draw VBN cord-048487-kamvao4b 97 15 from from IN cord-048487-kamvao4b 97 16 influenza influenza NN cord-048487-kamvao4b 97 17 subtypes subtype NNS cord-048487-kamvao4b 98 1 A a LS cord-048487-kamvao4b 98 2 - - : cord-048487-kamvao4b 98 3 H3N2 H3N2 NNP cord-048487-kamvao4b 98 4 , , , cord-048487-kamvao4b 98 5 B B NNP cord-048487-kamvao4b 98 6 and and CC cord-048487-kamvao4b 98 7 C. C. NNP cord-048487-kamvao4b 99 1 We -PRON- PRP cord-048487-kamvao4b 99 2 download download VBP cord-048487-kamvao4b 99 3 these these DT cord-048487-kamvao4b 99 4 data datum NNS cord-048487-kamvao4b 99 5 along along IN cord-048487-kamvao4b 99 6 with with IN cord-048487-kamvao4b 99 7 their -PRON- PRP$ cord-048487-kamvao4b 99 8 dates date NNS cord-048487-kamvao4b 99 9 of of IN cord-048487-kamvao4b 99 10 sampling sampling NN cord-048487-kamvao4b 99 11 from from IN cord-048487-kamvao4b 99 12 the the DT cord-048487-kamvao4b 99 13 Los Los NNP cord-048487-kamvao4b 99 14 Alamos Alamos NNP cord-048487-kamvao4b 99 15 Influenza Influenza NNP cord-048487-kamvao4b 99 16 Database Database NNP cord-048487-kamvao4b 99 17 [ [ -LRB- cord-048487-kamvao4b 99 18 16 16 CD cord-048487-kamvao4b 99 19 ] ] -RRB- cord-048487-kamvao4b 99 20 . . . cord-048487-kamvao4b 100 1 We -PRON- PRP cord-048487-kamvao4b 100 2 perform perform VBP cord-048487-kamvao4b 100 3 sequence sequence NN cord-048487-kamvao4b 100 4 alignment alignment NN cord-048487-kamvao4b 100 5 using use VBG cord-048487-kamvao4b 100 6 ClustalX clustalx PRP cord-048487-kamvao4b 100 7 [ [ -LRB- cord-048487-kamvao4b 100 8 24 24 CD cord-048487-kamvao4b 100 9 , , , cord-048487-kamvao4b 100 10 version version NN cord-048487-kamvao4b 100 11 1.8 1.8 CD cord-048487-kamvao4b 100 12 ] ] -RRB- cord-048487-kamvao4b 100 13 . . . cord-048487-kamvao4b 101 1 For for IN cord-048487-kamvao4b 101 2 consistency consistency NN cord-048487-kamvao4b 101 3 with with IN cord-048487-kamvao4b 101 4 previous previous JJ cord-048487-kamvao4b 101 5 studies study NNS cord-048487-kamvao4b 101 6 of of IN cord-048487-kamvao4b 101 7 A A NNP cord-048487-kamvao4b 101 8 - - : cord-048487-kamvao4b 101 9 H3N2 H3N2 NNP cord-048487-kamvao4b 101 10 HA HA NNP cord-048487-kamvao4b 101 11 evolution evolution NN cord-048487-kamvao4b 101 12 , , , cord-048487-kamvao4b 101 13 we -PRON- PRP cord-048487-kamvao4b 101 14 use use VBP cord-048487-kamvao4b 101 15 the the DT cord-048487-kamvao4b 101 16 HKY HKY NNP cord-048487-kamvao4b 101 17 model model NN cord-048487-kamvao4b 101 18 of of IN cord-048487-kamvao4b 101 19 nucleotide nucleotide JJ cord-048487-kamvao4b 101 20 substitution substitution NN cord-048487-kamvao4b 101 21 [ [ -LRB- cord-048487-kamvao4b 101 22 10 10 CD cord-048487-kamvao4b 101 23 ] ] -RRB- cord-048487-kamvao4b 101 24 . . . cord-048487-kamvao4b 102 1 We -PRON- PRP cord-048487-kamvao4b 102 2 use use VBP cord-048487-kamvao4b 102 3 the the DT cord-048487-kamvao4b 102 4 TREBLE treble NN cord-048487-kamvao4b 102 5 algorithm algorithm NN cord-048487-kamvao4b 102 6 , , , cord-048487-kamvao4b 102 7 which which WDT cord-048487-kamvao4b 102 8 implements implement VBZ cord-048487-kamvao4b 102 9 a a DT cord-048487-kamvao4b 102 10 MCA MCA NNP cord-048487-kamvao4b 102 11 , , , cord-048487-kamvao4b 102 12 on on IN cord-048487-kamvao4b 102 13 sets set NNS cord-048487-kamvao4b 102 14 of of IN cord-048487-kamvao4b 102 15 sequences sequence NNS cord-048487-kamvao4b 102 16 solely solely RB cord-048487-kamvao4b 102 17 drawn draw VBN cord-048487-kamvao4b 102 18 from from IN cord-048487-kamvao4b 102 19 a a DT cord-048487-kamvao4b 102 20 single single JJ cord-048487-kamvao4b 102 21 subtype subtype NN cord-048487-kamvao4b 102 22 to to TO cord-048487-kamvao4b 102 23 derive derive VB cord-048487-kamvao4b 102 24 within within IN cord-048487-kamvao4b 102 25 - - HYPH cord-048487-kamvao4b 102 26 subtype subtype NN cord-048487-kamvao4b 102 27 rates rate NNS cord-048487-kamvao4b 102 28 . . . cord-048487-kamvao4b 103 1 The the DT cord-048487-kamvao4b 103 2 phylogenetic phylogenetic JJ cord-048487-kamvao4b 103 3 tree tree NN cord-048487-kamvao4b 103 4 , , , cord-048487-kamvao4b 103 5 generated generate VBN cord-048487-kamvao4b 103 6 by by IN cord-048487-kamvao4b 103 7 TREBLE TREBLE NNP cord-048487-kamvao4b 103 8 , , , cord-048487-kamvao4b 103 9 for for IN cord-048487-kamvao4b 103 10 the the DT cord-048487-kamvao4b 103 11 HA HA NNP cord-048487-kamvao4b 103 12 gene gene NN cord-048487-kamvao4b 103 13 is be VBZ cord-048487-kamvao4b 103 14 depicted depict VBN cord-048487-kamvao4b 103 15 in in IN cord-048487-kamvao4b 103 16 Figure figure NN cord-048487-kamvao4b 103 17 2 2 CD cord-048487-kamvao4b 103 18 ( ( -LRB- cord-048487-kamvao4b 103 19 ± ± CD cord-048487-kamvao4b 103 20 0.48 0.48 CD cord-048487-kamvao4b 103 21 ) ) -RRB- cord-048487-kamvao4b 103 22 × × NN cord-048487-kamvao4b 103 23 10 10 CD cord-048487-kamvao4b 104 1 -3 -3 NNP cord-048487-kamvao4b 105 1 s s NNP cord-048487-kamvao4b 105 2 / / SYM cord-048487-kamvao4b 105 3 s s NNP cord-048487-kamvao4b 105 4 / / SYM cord-048487-kamvao4b 105 5 yr yr NN cord-048487-kamvao4b 105 6 and and CC cord-048487-kamvao4b 105 7 the the DT cord-048487-kamvao4b 105 8 subtype subtype NN cord-048487-kamvao4b 105 9 C c NN cord-048487-kamvao4b 105 10 rate rate NN cord-048487-kamvao4b 105 11 is be VBZ cord-048487-kamvao4b 105 12 1.31 1.31 CD cord-048487-kamvao4b 105 13 ( ( -LRB- cord-048487-kamvao4b 105 14 ± ± CD cord-048487-kamvao4b 105 15 0.33 0.33 CD cord-048487-kamvao4b 105 16 ) ) -RRB- cord-048487-kamvao4b 105 17 × × NN cord-048487-kamvao4b 105 18 10 10 CD cord-048487-kamvao4b 106 1 -3 -3 NNP cord-048487-kamvao4b 107 1 s s NNP cord-048487-kamvao4b 107 2 / / SYM cord-048487-kamvao4b 107 3 s s NNP cord-048487-kamvao4b 107 4 / / SYM cord-048487-kamvao4b 107 5 yr yr NN cord-048487-kamvao4b 107 6 . . . cord-048487-kamvao4b 108 1 Lastly lastly RB cord-048487-kamvao4b 108 2 , , , cord-048487-kamvao4b 108 3 for for IN cord-048487-kamvao4b 108 4 the the DT cord-048487-kamvao4b 108 5 NS NS NNP cord-048487-kamvao4b 108 6 gene gene NN cord-048487-kamvao4b 108 7 , , , cord-048487-kamvao4b 108 8 the the DT cord-048487-kamvao4b 108 9 rates rate NNS cord-048487-kamvao4b 108 10 are be VBP cord-048487-kamvao4b 108 11 similar similar JJ cord-048487-kamvao4b 108 12 to to IN cord-048487-kamvao4b 108 13 those those DT cord-048487-kamvao4b 108 14 of of IN cord-048487-kamvao4b 108 15 the the DT cord-048487-kamvao4b 108 16 MP MP NNP cord-048487-kamvao4b 108 17 gene gene NN cord-048487-kamvao4b 108 18 . . . cord-048487-kamvao4b 109 1 The the DT cord-048487-kamvao4b 109 2 subtype subtype NN cord-048487-kamvao4b 109 3 A a NN cord-048487-kamvao4b 109 4 - - : cord-048487-kamvao4b 109 5 H3N2 H3N2 NNP cord-048487-kamvao4b 109 6 rate rate NN cord-048487-kamvao4b 109 7 is be VBZ cord-048487-kamvao4b 109 8 2.14 2.14 CD cord-048487-kamvao4b 109 9 ( ( -LRB- cord-048487-kamvao4b 109 10 ± ± CD cord-048487-kamvao4b 109 11 0.25 0.25 CD cord-048487-kamvao4b 109 12 ) ) -RRB- cord-048487-kamvao4b 109 13 × × NN cord-048487-kamvao4b 109 14 10 10 CD cord-048487-kamvao4b 110 1 -3 -3 NNP cord-048487-kamvao4b 111 1 s s NNP cord-048487-kamvao4b 111 2 / / SYM cord-048487-kamvao4b 111 3 s s NNP cord-048487-kamvao4b 111 4 / / SYM cord-048487-kamvao4b 111 5 yr yr NN cord-048487-kamvao4b 111 6 , , , cord-048487-kamvao4b 111 7 the the DT cord-048487-kamvao4b 111 8 subtype subtype NN cord-048487-kamvao4b 111 9 B b NN cord-048487-kamvao4b 111 10 rate rate NN cord-048487-kamvao4b 111 11 is be VBZ cord-048487-kamvao4b 111 12 1.92 1.92 CD cord-048487-kamvao4b 111 13 ( ( -LRB- cord-048487-kamvao4b 111 14 ± ± CD cord-048487-kamvao4b 111 15 0.20 0.20 CD cord-048487-kamvao4b 111 16 ) ) -RRB- cord-048487-kamvao4b 111 17 × × NN cord-048487-kamvao4b 111 18 10 10 CD cord-048487-kamvao4b 112 1 -3 -3 NNP cord-048487-kamvao4b 112 2 s s NNP cord-048487-kamvao4b 112 3 / / SYM cord-048487-kamvao4b 112 4 s s NNP cord-048487-kamvao4b 112 5 / / SYM cord-048487-kamvao4b 112 6 yr yr NN cord-048487-kamvao4b 112 7 , , , cord-048487-kamvao4b 112 8 and and CC cord-048487-kamvao4b 112 9 the the DT cord-048487-kamvao4b 112 10 subtype subtype NN cord-048487-kamvao4b 112 11 C c NN cord-048487-kamvao4b 112 12 rate rate NN cord-048487-kamvao4b 112 13 is be VBZ cord-048487-kamvao4b 112 14 1.68 1.68 CD cord-048487-kamvao4b 112 15 ( ( -LRB- cord-048487-kamvao4b 112 16 ± ± CD cord-048487-kamvao4b 112 17 0.51 0.51 CD cord-048487-kamvao4b 112 18 ) ) -RRB- cord-048487-kamvao4b 113 1 × × NN cord-048487-kamvao4b 113 2 10 10 CD cord-048487-kamvao4b 114 1 -3 -3 NNP cord-048487-kamvao4b 114 2 s s NNP cord-048487-kamvao4b 114 3 / / SYM cord-048487-kamvao4b 114 4 s s NNP cord-048487-kamvao4b 114 5 / / SYM cord-048487-kamvao4b 114 6 yr yr NN cord-048487-kamvao4b 114 7 . . . cord-048487-kamvao4b 115 1 Table table NN cord-048487-kamvao4b 115 2 1 1 CD cord-048487-kamvao4b 115 3 presents present VBZ cord-048487-kamvao4b 115 4 these these DT cord-048487-kamvao4b 115 5 results result NNS cord-048487-kamvao4b 115 6 . . . cord-048487-kamvao4b 116 1 Figure figure NN cord-048487-kamvao4b 116 2 3 3 CD cord-048487-kamvao4b 116 3 provides provide VBZ cord-048487-kamvao4b 116 4 histograms histogram NNS cord-048487-kamvao4b 116 5 of of IN cord-048487-kamvao4b 116 6 the the DT cord-048487-kamvao4b 116 7 bootstrap bootstrap NN cord-048487-kamvao4b 116 8 distributions distribution NNS cord-048487-kamvao4b 116 9 for for IN cord-048487-kamvao4b 116 10 all all DT cord-048487-kamvao4b 116 11 three three CD cord-048487-kamvao4b 116 12 genes gene NNS cord-048487-kamvao4b 116 13 and and CC cord-048487-kamvao4b 116 14 subtypes subtype NNS cord-048487-kamvao4b 116 15 . . . cord-048487-kamvao4b 117 1 Assuming assume VBG cord-048487-kamvao4b 117 2 a a DT cord-048487-kamvao4b 117 3 molecular molecular JJ cord-048487-kamvao4b 117 4 clock clock NN cord-048487-kamvao4b 117 5 within within IN cord-048487-kamvao4b 117 6 a a DT cord-048487-kamvao4b 117 7 subtype subtype NN cord-048487-kamvao4b 117 8 and and CC cord-048487-kamvao4b 117 9 with with IN cord-048487-kamvao4b 117 10 the the DT cord-048487-kamvao4b 117 11 rates rate NNS cord-048487-kamvao4b 117 12 above above RB cord-048487-kamvao4b 117 13 , , , cord-048487-kamvao4b 117 14 we -PRON- PRP cord-048487-kamvao4b 117 15 generated generate VBD cord-048487-kamvao4b 117 16 the the DT cord-048487-kamvao4b 117 17 corresponding corresponding JJ cord-048487-kamvao4b 117 18 dates date NNS cord-048487-kamvao4b 117 19 of of IN cord-048487-kamvao4b 117 20 the the DT cord-048487-kamvao4b 117 21 TMCRA TMCRA NNS cord-048487-kamvao4b 117 22 . . . cord-048487-kamvao4b 118 1 Figure figure NN cord-048487-kamvao4b 118 2 3 3 CD cord-048487-kamvao4b 118 3 shows show VBZ cord-048487-kamvao4b 118 4 histograms histogram NNS cord-048487-kamvao4b 118 5 of of IN cord-048487-kamvao4b 118 6 the the DT cord-048487-kamvao4b 118 7 TMRCA TMRCA NNP cord-048487-kamvao4b 118 8 estimates estimate NNS cord-048487-kamvao4b 118 9 for for IN cord-048487-kamvao4b 118 10 different different JJ cord-048487-kamvao4b 118 11 genes gene NNS cord-048487-kamvao4b 118 12 and and CC cord-048487-kamvao4b 118 13 subtypes subtype NNS cord-048487-kamvao4b 118 14 . . . cord-048487-kamvao4b 119 1 All all DT cord-048487-kamvao4b 119 2 genes gene NNS cord-048487-kamvao4b 119 3 are be VBP cord-048487-kamvao4b 119 4 similar similar JJ cord-048487-kamvao4b 119 5 in in IN cord-048487-kamvao4b 119 6 dating date VBG cord-048487-kamvao4b 119 7 the the DT cord-048487-kamvao4b 119 8 TMRCA TMRCA NNP cord-048487-kamvao4b 119 9 for for IN cord-048487-kamvao4b 119 10 A A NNP cord-048487-kamvao4b 119 11 - - : cord-048487-kamvao4b 119 12 H3N2 H3N2 NNP cord-048487-kamvao4b 119 13 to to IN cord-048487-kamvao4b 119 14 approximately approximately RB cord-048487-kamvao4b 119 15 1965 1965 CD cord-048487-kamvao4b 119 16 ( ( -LRB- cord-048487-kamvao4b 119 17 1964 1964 CD cord-048487-kamvao4b 119 18 , , , cord-048487-kamvao4b 119 19 1965 1965 CD cord-048487-kamvao4b 119 20 , , , cord-048487-kamvao4b 119 21 and and CC cord-048487-kamvao4b 119 22 1962 1962 CD cord-048487-kamvao4b 119 23 for for IN cord-048487-kamvao4b 119 24 HA HA NNP cord-048487-kamvao4b 119 25 , , , cord-048487-kamvao4b 119 26 MP MP NNP cord-048487-kamvao4b 119 27 and and CC cord-048487-kamvao4b 119 28 NS NS NNP cord-048487-kamvao4b 119 29 genes gene NNS cord-048487-kamvao4b 119 30 , , , cord-048487-kamvao4b 119 31 respectively respectively RB cord-048487-kamvao4b 119 32 ) ) -RRB- cord-048487-kamvao4b 119 33 . . . cord-048487-kamvao4b 120 1 These these DT cord-048487-kamvao4b 120 2 dates date NNS cord-048487-kamvao4b 120 3 are be VBP cord-048487-kamvao4b 120 4 consistent consistent JJ cord-048487-kamvao4b 120 5 with with IN cord-048487-kamvao4b 120 6 the the DT cord-048487-kamvao4b 120 7 emergence emergence NN cord-048487-kamvao4b 120 8 of of IN cord-048487-kamvao4b 120 9 the the DT cord-048487-kamvao4b 120 10 A A NNP cord-048487-kamvao4b 120 11 - - HYPH cord-048487-kamvao4b 120 12 H3N2 H3N2 NNP cord-048487-kamvao4b 120 13 subtype subtype NN cord-048487-kamvao4b 120 14 into into IN cord-048487-kamvao4b 120 15 global global JJ cord-048487-kamvao4b 120 16 circulation circulation NN cord-048487-kamvao4b 120 17 dur dur NN cord-048487-kamvao4b 120 18 - - : cord-048487-kamvao4b 120 19 ing e VBG cord-048487-kamvao4b 120 20 the the DT cord-048487-kamvao4b 120 21 1968 1968 CD cord-048487-kamvao4b 120 22 pandemic pandemic NN cord-048487-kamvao4b 120 23 [ [ -LRB- cord-048487-kamvao4b 120 24 1 1 CD cord-048487-kamvao4b 120 25 ] ] -RRB- cord-048487-kamvao4b 120 26 . . . cord-048487-kamvao4b 121 1 Both both DT cord-048487-kamvao4b 121 2 the the DT cord-048487-kamvao4b 121 3 MP MP NNP cord-048487-kamvao4b 121 4 and and CC cord-048487-kamvao4b 121 5 NS NS NNP cord-048487-kamvao4b 121 6 genes gene NNS cord-048487-kamvao4b 121 7 date date VBP cord-048487-kamvao4b 121 8 the the DT cord-048487-kamvao4b 121 9 TMRCA TMRCA NNP cord-048487-kamvao4b 121 10 of of IN cord-048487-kamvao4b 121 11 subtype subtype NN cord-048487-kamvao4b 121 12 B b NN cord-048487-kamvao4b 121 13 to to IN cord-048487-kamvao4b 121 14 1943 1943 CD cord-048487-kamvao4b 121 15 , , , cord-048487-kamvao4b 121 16 while while IN cord-048487-kamvao4b 121 17 the the DT cord-048487-kamvao4b 121 18 HA HA NNP cord-048487-kamvao4b 121 19 rate rate NN cord-048487-kamvao4b 121 20 places place NNS cord-048487-kamvao4b 121 21 the the DT cord-048487-kamvao4b 121 22 TMRCA TMRCA NNP cord-048487-kamvao4b 121 23 at at IN cord-048487-kamvao4b 121 24 1953 1953 CD cord-048487-kamvao4b 121 25 . . . cord-048487-kamvao4b 122 1 This this DT cord-048487-kamvao4b 122 2 latter latter JJ cord-048487-kamvao4b 122 3 value value NN cord-048487-kamvao4b 122 4 is be VBZ cord-048487-kamvao4b 122 5 inconsistent inconsistent JJ cord-048487-kamvao4b 122 6 with with IN cord-048487-kamvao4b 122 7 the the DT cord-048487-kamvao4b 122 8 influenza influenza NN cord-048487-kamvao4b 122 9 B b IN cord-048487-kamvao4b 122 10 sub sub NN cord-048487-kamvao4b 122 11 - - NNS cord-048487-kamvao4b 122 12 epidemics epidemic NNS cord-048487-kamvao4b 122 13 of of IN cord-048487-kamvao4b 122 14 1950 1950 CD cord-048487-kamvao4b 122 15 - - SYM cord-048487-kamvao4b 122 16 51 51 CD cord-048487-kamvao4b 122 17 but but CC cord-048487-kamvao4b 122 18 is be VBZ cord-048487-kamvao4b 122 19 consistent consistent JJ cord-048487-kamvao4b 122 20 with with IN cord-048487-kamvao4b 122 21 the the DT cord-048487-kamvao4b 122 22 emergence emergence NN cord-048487-kamvao4b 122 23 of of IN cord-048487-kamvao4b 122 24 the the DT cord-048487-kamvao4b 122 25 more more RBR cord-048487-kamvao4b 122 26 lethal lethal JJ cord-048487-kamvao4b 122 27 Victoria Victoria NNP cord-048487-kamvao4b 122 28 strain strain NN cord-048487-kamvao4b 122 29 of of IN cord-048487-kamvao4b 122 30 influenza influenza NN cord-048487-kamvao4b 122 31 B b NN cord-048487-kamvao4b 122 32 in in IN cord-048487-kamvao4b 122 33 1953 1953 CD cord-048487-kamvao4b 122 34 [ [ -LRB- cord-048487-kamvao4b 122 35 11 11 CD cord-048487-kamvao4b 122 36 ] ] -RRB- cord-048487-kamvao4b 122 37 . . . cord-048487-kamvao4b 123 1 Each each DT cord-048487-kamvao4b 123 2 of of IN cord-048487-kamvao4b 123 3 these these DT cord-048487-kamvao4b 123 4 estimates estimate NNS cord-048487-kamvao4b 123 5 has have VBZ cord-048487-kamvao4b 123 6 a a DT cord-048487-kamvao4b 123 7 standard standard JJ cord-048487-kamvao4b 123 8 error error NN cord-048487-kamvao4b 123 9 of of IN cord-048487-kamvao4b 123 10 approximately approximately RB cord-048487-kamvao4b 123 11 2 2 CD cord-048487-kamvao4b 123 12 years year NNS cord-048487-kamvao4b 123 13 and and CC cord-048487-kamvao4b 123 14 so so RB cord-048487-kamvao4b 123 15 these these DT cord-048487-kamvao4b 123 16 discrepancies discrepancy NNS cord-048487-kamvao4b 123 17 may may MD cord-048487-kamvao4b 123 18 be be VB cord-048487-kamvao4b 123 19 accounted account VBN cord-048487-kamvao4b 123 20 by by IN cord-048487-kamvao4b 123 21 measurement measurement NN cord-048487-kamvao4b 123 22 uncertainty uncertainty NN cord-048487-kamvao4b 123 23 . . . cord-048487-kamvao4b 124 1 The the DT cord-048487-kamvao4b 124 2 10 10 CD cord-048487-kamvao4b 124 3 year year NN cord-048487-kamvao4b 124 4 gap gap NN cord-048487-kamvao4b 124 5 between between IN cord-048487-kamvao4b 124 6 the the DT cord-048487-kamvao4b 124 7 TMCRA TMCRA NNP cord-048487-kamvao4b 124 8 suggested suggest VBN cord-048487-kamvao4b 124 9 by by IN cord-048487-kamvao4b 124 10 the the DT cord-048487-kamvao4b 124 11 different different JJ cord-048487-kamvao4b 124 12 genes gene NNS cord-048487-kamvao4b 124 13 can can MD cord-048487-kamvao4b 124 14 be be VB cord-048487-kamvao4b 124 15 explained explain VBN cord-048487-kamvao4b 124 16 by by IN cord-048487-kamvao4b 124 17 a a DT cord-048487-kamvao4b 124 18 reassortment reassortment JJ cord-048487-kamvao4b 124 19 event event NN cord-048487-kamvao4b 124 20 . . . cord-048487-kamvao4b 125 1 Finally finally RB cord-048487-kamvao4b 125 2 , , , cord-048487-kamvao4b 125 3 the the DT cord-048487-kamvao4b 125 4 TMRCA TMRCA NNP cord-048487-kamvao4b 125 5 of of IN cord-048487-kamvao4b 125 6 subtype subtype NN cord-048487-kamvao4b 125 7 C C NNP cord-048487-kamvao4b 125 8 is be VBZ cord-048487-kamvao4b 125 9 calculated calculate VBN cord-048487-kamvao4b 125 10 as as IN cord-048487-kamvao4b 125 11 1952 1952 CD cord-048487-kamvao4b 125 12 and and CC cord-048487-kamvao4b 125 13 1953 1953 CD cord-048487-kamvao4b 125 14 by by IN cord-048487-kamvao4b 125 15 the the DT cord-048487-kamvao4b 125 16 MP MP NNP cord-048487-kamvao4b 125 17 and and CC cord-048487-kamvao4b 125 18 NS NS NNP cord-048487-kamvao4b 125 19 genes gene NNS cord-048487-kamvao4b 125 20 , , , cord-048487-kamvao4b 125 21 respectively respectively RB cord-048487-kamvao4b 125 22 , , , cord-048487-kamvao4b 125 23 while while IN cord-048487-kamvao4b 125 24 the the DT cord-048487-kamvao4b 125 25 HA HA NNP cord-048487-kamvao4b 125 26 gene gene NN cord-048487-kamvao4b 125 27 places place VBZ cord-048487-kamvao4b 125 28 the the DT cord-048487-kamvao4b 125 29 TMCRA TMCRA NNP cord-048487-kamvao4b 125 30 at at IN cord-048487-kamvao4b 125 31 1906 1906 CD cord-048487-kamvao4b 125 32 . . . cord-048487-kamvao4b 126 1 This this DT cord-048487-kamvao4b 126 2 nearly nearly RB cord-048487-kamvao4b 126 3 half half JJ cord-048487-kamvao4b 126 4 century century NN cord-048487-kamvao4b 126 5 discrepancy discrepancy NN cord-048487-kamvao4b 126 6 suggests suggest VBZ cord-048487-kamvao4b 126 7 that that IN cord-048487-kamvao4b 126 8 the the DT cord-048487-kamvao4b 126 9 subtype subtype NN cord-048487-kamvao4b 126 10 C C NNP cord-048487-kamvao4b 126 11 HA HA NNP cord-048487-kamvao4b 126 12 gene gene NN cord-048487-kamvao4b 126 13 experienced experience VBD cord-048487-kamvao4b 126 14 a a DT cord-048487-kamvao4b 126 15 markedly markedly RB cord-048487-kamvao4b 126 16 different different JJ cord-048487-kamvao4b 126 17 evolutionary evolutionary JJ cord-048487-kamvao4b 126 18 history history NN cord-048487-kamvao4b 126 19 than than IN cord-048487-kamvao4b 126 20 either either CC cord-048487-kamvao4b 126 21 the the DT cord-048487-kamvao4b 126 22 MP MP NNP cord-048487-kamvao4b 126 23 or or CC cord-048487-kamvao4b 126 24 the the DT cord-048487-kamvao4b 126 25 NS NS NNP cord-048487-kamvao4b 126 26 gene gene NN cord-048487-kamvao4b 126 27 . . . cord-048487-kamvao4b 127 1 A a DT cord-048487-kamvao4b 127 2 biologically biologically RB cord-048487-kamvao4b 127 3 plausible plausible JJ cord-048487-kamvao4b 127 4 explanation explanation NN cord-048487-kamvao4b 127 5 would would MD cord-048487-kamvao4b 127 6 be be VB cord-048487-kamvao4b 127 7 a a DT cord-048487-kamvao4b 127 8 reassortment reassortment JJ cord-048487-kamvao4b 127 9 event event NN cord-048487-kamvao4b 127 10 . . . cord-048487-kamvao4b 128 1 Another another DT cord-048487-kamvao4b 128 2 possible possible JJ cord-048487-kamvao4b 128 3 explanation explanation NN cord-048487-kamvao4b 128 4 is be VBZ cord-048487-kamvao4b 128 5 that that IN cord-048487-kamvao4b 128 6 non non JJ cord-048487-kamvao4b 128 7 - - JJ cord-048487-kamvao4b 128 8 MCA mca JJ cord-048487-kamvao4b 128 9 rate rate NN cord-048487-kamvao4b 128 10 behavior behavior NN cord-048487-kamvao4b 128 11 has have VBZ cord-048487-kamvao4b 128 12 lead lead NN cord-048487-kamvao4b 128 13 to to IN cord-048487-kamvao4b 128 14 substantial substantial JJ cord-048487-kamvao4b 128 15 bias bias NN cord-048487-kamvao4b 128 16 in in IN cord-048487-kamvao4b 128 17 dating date VBG cord-048487-kamvao4b 128 18 the the DT cord-048487-kamvao4b 128 19 TMRCA TMRCA NNP cord-048487-kamvao4b 128 20 . . . cord-048487-kamvao4b 129 1 We -PRON- PRP cord-048487-kamvao4b 129 2 now now RB cord-048487-kamvao4b 129 3 compare compare VBP cord-048487-kamvao4b 129 4 the the DT cord-048487-kamvao4b 129 5 results result NNS cord-048487-kamvao4b 129 6 from from IN cord-048487-kamvao4b 129 7 pairwise pairwise JJ cord-048487-kamvao4b 129 8 rate rate NN cord-048487-kamvao4b 129 9 estimates estimate NNS cord-048487-kamvao4b 129 10 across across IN cord-048487-kamvao4b 129 11 subtypes subtype NNS cord-048487-kamvao4b 129 12 A A NNP cord-048487-kamvao4b 129 13 - - : cord-048487-kamvao4b 129 14 H3N2 H3N2 NNP cord-048487-kamvao4b 129 15 and and CC cord-048487-kamvao4b 129 16 B B NNP cord-048487-kamvao4b 129 17 with with IN cord-048487-kamvao4b 129 18 those those DT cord-048487-kamvao4b 129 19 from from IN cord-048487-kamvao4b 129 20 application application NN cord-048487-kamvao4b 129 21 of of IN cord-048487-kamvao4b 129 22 the the DT cord-048487-kamvao4b 129 23 MCA MCA NNP cord-048487-kamvao4b 129 24 to to IN cord-048487-kamvao4b 129 25 the the DT cord-048487-kamvao4b 129 26 same same JJ cord-048487-kamvao4b 129 27 data datum NNS cord-048487-kamvao4b 129 28 . . . cord-048487-kamvao4b 130 1 These these DT cord-048487-kamvao4b 130 2 results result NNS cord-048487-kamvao4b 130 3 are be VBP cord-048487-kamvao4b 130 4 summarized summarize VBN cord-048487-kamvao4b 130 5 in in IN cord-048487-kamvao4b 130 6 Table Table NNP cord-048487-kamvao4b 130 7 2 2 CD cord-048487-kamvao4b 130 8 and and CC cord-048487-kamvao4b 130 9 Histograms Histograms NNPS cord-048487-kamvao4b 130 10 of of IN cord-048487-kamvao4b 130 11 the the DT cord-048487-kamvao4b 130 12 time time NN cord-048487-kamvao4b 130 13 of of IN cord-048487-kamvao4b 130 14 most most RBS cord-048487-kamvao4b 130 15 recent recent JJ cord-048487-kamvao4b 130 16 common common JJ cord-048487-kamvao4b 130 17 ancestor ancestor NN cord-048487-kamvao4b 130 18 for for IN cord-048487-kamvao4b 130 19 subtypes subtype NNS cord-048487-kamvao4b 131 1 A a LS cord-048487-kamvao4b 131 2 - - : cord-048487-kamvao4b 131 3 H3N2 H3N2 NNP cord-048487-kamvao4b 131 4 , , , cord-048487-kamvao4b 131 5 B B NNP cord-048487-kamvao4b 131 6 and and CC cord-048487-kamvao4b 131 7 C C NNP cord-048487-kamvao4b 131 8 , , , cord-048487-kamvao4b 131 9 respectively respectively RB cord-048487-kamvao4b 131 10 , , , cord-048487-kamvao4b 131 11 derived derive VBN cord-048487-kamvao4b 131 12 from from IN cord-048487-kamvao4b 131 13 molecular molecular JJ cord-048487-kamvao4b 131 14 clock clock NN cord-048487-kamvao4b 131 15 estimates estimate NNS cord-048487-kamvao4b 131 16 on on IN cord-048487-kamvao4b 131 17 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 131 18 ( ( -LRB- cord-048487-kamvao4b 131 19 HA HA NNP cord-048487-kamvao4b 131 20 ) ) -RRB- cord-048487-kamvao4b 131 21 , , , cord-048487-kamvao4b 131 22 matrix matrix NN cord-048487-kamvao4b 131 23 ( ( -LRB- cord-048487-kamvao4b 131 24 MP MP NNP cord-048487-kamvao4b 131 25 ) ) -RRB- cord-048487-kamvao4b 131 26 and and CC cord-048487-kamvao4b 132 1 nonstructural nonstructural NNP cord-048487-kamvao4b 132 2 ( ( -LRB- cord-048487-kamvao4b 132 3 NS NS NNP cord-048487-kamvao4b 132 4 ) ) -RRB- cord-048487-kamvao4b 132 5 gene gene NN cord-048487-kamvao4b 132 6 sequences sequence NNS cord-048487-kamvao4b 133 1 This this DT cord-048487-kamvao4b 133 2 discrepancy discrepancy NN cord-048487-kamvao4b 133 3 between between IN cord-048487-kamvao4b 133 4 the the DT cord-048487-kamvao4b 133 5 two two CD cord-048487-kamvao4b 133 6 sets set NNS cord-048487-kamvao4b 133 7 of of IN cord-048487-kamvao4b 133 8 estimates estimate NNS cord-048487-kamvao4b 133 9 of of IN cord-048487-kamvao4b 133 10 the the DT cord-048487-kamvao4b 133 11 TMRCA TMRCA NNP cord-048487-kamvao4b 133 12 likely likely RB cord-048487-kamvao4b 133 13 owes owe VBZ cord-048487-kamvao4b 133 14 to to IN cord-048487-kamvao4b 133 15 the the DT cord-048487-kamvao4b 133 16 inability inability NN cord-048487-kamvao4b 133 17 of of IN cord-048487-kamvao4b 133 18 the the DT cord-048487-kamvao4b 133 19 MCA MCA NNP cord-048487-kamvao4b 133 20 to to TO cord-048487-kamvao4b 133 21 integrate integrate VB cord-048487-kamvao4b 133 22 information information NN cord-048487-kamvao4b 133 23 from from IN cord-048487-kamvao4b 133 24 the the DT cord-048487-kamvao4b 133 25 period period NN cord-048487-kamvao4b 133 26 of of IN cord-048487-kamvao4b 133 27 evolution evolution NN cord-048487-kamvao4b 133 28 between between IN cord-048487-kamvao4b 133 29 the the DT cord-048487-kamvao4b 133 30 two two CD cord-048487-kamvao4b 133 31 subtypes subtype NNS cord-048487-kamvao4b 133 32 , , , cord-048487-kamvao4b 133 33 leading lead VBG cord-048487-kamvao4b 133 34 to to IN cord-048487-kamvao4b 133 35 a a DT cord-048487-kamvao4b 133 36 substantial substantial JJ cord-048487-kamvao4b 133 37 underestimate underestimate NN cord-048487-kamvao4b 133 38 of of IN cord-048487-kamvao4b 133 39 the the DT cord-048487-kamvao4b 133 40 rate rate NN cord-048487-kamvao4b 133 41 of of IN cord-048487-kamvao4b 133 42 substitution substitution NN cord-048487-kamvao4b 133 43 , , , cord-048487-kamvao4b 133 44 and and CC cord-048487-kamvao4b 133 45 consequent consequent JJ cord-048487-kamvao4b 133 46 underestimation underestimation NN cord-048487-kamvao4b 133 47 of of IN cord-048487-kamvao4b 133 48 the the DT cord-048487-kamvao4b 133 49 date date NN cord-048487-kamvao4b 133 50 of of IN cord-048487-kamvao4b 133 51 the the DT cord-048487-kamvao4b 133 52 TMRCA TMRCA NNP cord-048487-kamvao4b 133 53 . . . cord-048487-kamvao4b 134 1 We -PRON- PRP cord-048487-kamvao4b 134 2 present present VBP cord-048487-kamvao4b 134 3 a a DT cord-048487-kamvao4b 134 4 new new JJ cord-048487-kamvao4b 134 5 method method NN cord-048487-kamvao4b 134 6 for for IN cord-048487-kamvao4b 134 7 ascertaining ascertain VBG cord-048487-kamvao4b 134 8 the the DT cord-048487-kamvao4b 134 9 rate rate NN cord-048487-kamvao4b 134 10 of of IN cord-048487-kamvao4b 134 11 nucleotide nucleotide JJ cord-048487-kamvao4b 134 12 substitution substitution NN cord-048487-kamvao4b 134 13 between between IN cord-048487-kamvao4b 134 14 subtypes subtype NNS cord-048487-kamvao4b 134 15 and and CC cord-048487-kamvao4b 134 16 apply apply VB cord-048487-kamvao4b 134 17 this this DT cord-048487-kamvao4b 134 18 method method NN cord-048487-kamvao4b 134 19 together together RB cord-048487-kamvao4b 134 20 with with IN cord-048487-kamvao4b 134 21 traditional traditional JJ cord-048487-kamvao4b 134 22 MCA MCA NNP cord-048487-kamvao4b 134 23 methods method NNS cord-048487-kamvao4b 134 24 to to IN cord-048487-kamvao4b 134 25 date date NN cord-048487-kamvao4b 134 26 the the DT cord-048487-kamvao4b 134 27 divergence divergence NN cord-048487-kamvao4b 134 28 of of IN cord-048487-kamvao4b 134 29 influenza influenza NN cord-048487-kamvao4b 134 30 subtypes subtype NNS cord-048487-kamvao4b 134 31 A A NNP cord-048487-kamvao4b 134 32 - - : cord-048487-kamvao4b 134 33 H3N2 H3N2 NNP cord-048487-kamvao4b 134 34 , , , cord-048487-kamvao4b 134 35 B B NNP cord-048487-kamvao4b 134 36 , , , cord-048487-kamvao4b 134 37 and and CC cord-048487-kamvao4b 134 38 C. C. NNP cord-048487-kamvao4b 135 1 We -PRON- PRP cord-048487-kamvao4b 135 2 use use VBP cord-048487-kamvao4b 135 3 three three CD cord-048487-kamvao4b 135 4 genes gene NNS cord-048487-kamvao4b 135 5 , , , cord-048487-kamvao4b 135 6 HA HA NNP cord-048487-kamvao4b 135 7 , , , cord-048487-kamvao4b 135 8 MP MP NNP cord-048487-kamvao4b 135 9 and and CC cord-048487-kamvao4b 135 10 NS NS NNP cord-048487-kamvao4b 135 11 , , , cord-048487-kamvao4b 135 12 to to IN cord-048487-kamvao4b 135 13 date date NN cord-048487-kamvao4b 135 14 two two CD cord-048487-kamvao4b 135 15 types type NNS cord-048487-kamvao4b 135 16 of of IN cord-048487-kamvao4b 135 17 divergence divergence NN cord-048487-kamvao4b 135 18 events event NNS cord-048487-kamvao4b 135 19 : : : cord-048487-kamvao4b 135 20 the the DT cord-048487-kamvao4b 135 21 time time NN cord-048487-kamvao4b 135 22 of of IN cord-048487-kamvao4b 135 23 the the DT cord-048487-kamvao4b 135 24 most most RBS cord-048487-kamvao4b 135 25 recent recent JJ cord-048487-kamvao4b 135 26 common common NN cord-048487-kamvao4b 135 27 of of IN cord-048487-kamvao4b 135 28 each each DT cord-048487-kamvao4b 135 29 subtype subtype NN cord-048487-kamvao4b 135 30 and and CC cord-048487-kamvao4b 135 31 the the DT cord-048487-kamvao4b 135 32 time time NN cord-048487-kamvao4b 135 33 of of IN cord-048487-kamvao4b 135 34 divergence divergence NN cord-048487-kamvao4b 135 35 between between IN cord-048487-kamvao4b 135 36 two two CD cord-048487-kamvao4b 135 37 subtypes subtype NNS cord-048487-kamvao4b 135 38 , , , cord-048487-kamvao4b 135 39 A A NNP cord-048487-kamvao4b 135 40 - - HYPH cord-048487-kamvao4b 135 41 H3N2 H3N2 NNP cord-048487-kamvao4b 135 42 and and CC cord-048487-kamvao4b 135 43 B. B. NNP cord-048487-kamvao4b 136 1 For for IN cord-048487-kamvao4b 136 2 the the DT cord-048487-kamvao4b 136 3 former former JJ cord-048487-kamvao4b 136 4 event event NN cord-048487-kamvao4b 136 5 type type NN cord-048487-kamvao4b 136 6 , , , cord-048487-kamvao4b 136 7 we -PRON- PRP cord-048487-kamvao4b 136 8 show show VBP cord-048487-kamvao4b 136 9 that that IN cord-048487-kamvao4b 136 10 the the DT cord-048487-kamvao4b 136 11 three three CD cord-048487-kamvao4b 136 12 genes gene NNS cord-048487-kamvao4b 136 13 are be VBP cord-048487-kamvao4b 136 14 loosely loosely RB cord-048487-kamvao4b 136 15 consistent consistent JJ cord-048487-kamvao4b 136 16 in in IN cord-048487-kamvao4b 136 17 their -PRON- PRP$ cord-048487-kamvao4b 136 18 dating dating NN cord-048487-kamvao4b 136 19 of of IN cord-048487-kamvao4b 136 20 the the DT cord-048487-kamvao4b 136 21 TMRCA TMRCA NNP cord-048487-kamvao4b 136 22 of of IN cord-048487-kamvao4b 136 23 the the DT cord-048487-kamvao4b 136 24 subtypes subtype NNS cord-048487-kamvao4b 136 25 , , , cord-048487-kamvao4b 136 26 with with IN cord-048487-kamvao4b 136 27 the the DT cord-048487-kamvao4b 136 28 notable notable JJ cord-048487-kamvao4b 136 29 exception exception NN cord-048487-kamvao4b 136 30 of of IN cord-048487-kamvao4b 136 31 the the DT cord-048487-kamvao4b 136 32 HA HA NNP cord-048487-kamvao4b 136 33 - - HYPH cord-048487-kamvao4b 136 34 derived derive VBN cord-048487-kamvao4b 136 35 estimate estimate NN cord-048487-kamvao4b 136 36 of of IN cord-048487-kamvao4b 136 37 subtype subtype NN cord-048487-kamvao4b 136 38 C C NNP cord-048487-kamvao4b 136 39 's 's POS cord-048487-kamvao4b 136 40 TMRCA TMRCA NNP cord-048487-kamvao4b 136 41 approximately approximately RB cord-048487-kamvao4b 136 42 50 50 CD cord-048487-kamvao4b 136 43 years year NNS cord-048487-kamvao4b 136 44 before before IN cord-048487-kamvao4b 136 45 the the DT cord-048487-kamvao4b 136 46 MP MP NNP cord-048487-kamvao4b 136 47 - - : cord-048487-kamvao4b 136 48 and and CC cord-048487-kamvao4b 136 49 NS NS NNP cord-048487-kamvao4b 136 50 - - HYPH cord-048487-kamvao4b 136 51 derived derive VBN cord-048487-kamvao4b 136 52 estimates estimate NNS cord-048487-kamvao4b 136 53 . . . cord-048487-kamvao4b 137 1 This this DT cord-048487-kamvao4b 137 2 discrepancy discrepancy NN cord-048487-kamvao4b 137 3 may may MD cord-048487-kamvao4b 137 4 indicate indicate VB cord-048487-kamvao4b 137 5 either either CC cord-048487-kamvao4b 137 6 that that IN cord-048487-kamvao4b 137 7 subtype subtype NN cord-048487-kamvao4b 137 8 C C NNP cord-048487-kamvao4b 137 9 's 's POS cord-048487-kamvao4b 137 10 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 137 11 - - HYPH cord-048487-kamvao4b 137 12 esterase esterase NN cord-048487-kamvao4b 137 13 gene gene NN cord-048487-kamvao4b 137 14 engaged engage VBN cord-048487-kamvao4b 137 15 in in IN cord-048487-kamvao4b 137 16 a a DT cord-048487-kamvao4b 137 17 biologically biologically RB cord-048487-kamvao4b 137 18 significant significant JJ cord-048487-kamvao4b 137 19 event event NN cord-048487-kamvao4b 137 20 , , , cord-048487-kamvao4b 137 21 such such JJ cord-048487-kamvao4b 137 22 as as IN cord-048487-kamvao4b 137 23 reassortment reassortment NN cord-048487-kamvao4b 137 24 , , , cord-048487-kamvao4b 137 25 or or CC cord-048487-kamvao4b 137 26 that that IN cord-048487-kamvao4b 137 27 MCA MCA NNP cord-048487-kamvao4b 137 28 estimation estimation NN cord-048487-kamvao4b 137 29 does do VBZ cord-048487-kamvao4b 137 30 not not RB cord-048487-kamvao4b 137 31 adequately adequately RB cord-048487-kamvao4b 137 32 model model VB cord-048487-kamvao4b 137 33 the the DT cord-048487-kamvao4b 137 34 evolution evolution NN cord-048487-kamvao4b 137 35 of of IN cord-048487-kamvao4b 137 36 the the DT cord-048487-kamvao4b 137 37 gene gene NN cord-048487-kamvao4b 137 38 . . . cord-048487-kamvao4b 138 1 For for IN cord-048487-kamvao4b 138 2 the the DT cord-048487-kamvao4b 138 3 divergence divergence NN cord-048487-kamvao4b 138 4 between between IN cord-048487-kamvao4b 138 5 subtypes subtype NNS cord-048487-kamvao4b 138 6 A A NNP cord-048487-kamvao4b 138 7 - - : cord-048487-kamvao4b 138 8 H3N2 H3N2 NNP cord-048487-kamvao4b 138 9 and and CC cord-048487-kamvao4b 138 10 B B NNP cord-048487-kamvao4b 138 11 , , , cord-048487-kamvao4b 138 12 previous previous JJ cord-048487-kamvao4b 138 13 studies study NNS cord-048487-kamvao4b 138 14 using use VBG cord-048487-kamvao4b 138 15 the the DT cord-048487-kamvao4b 138 16 MCA MCA NNP cord-048487-kamvao4b 138 17 generally generally RB cord-048487-kamvao4b 138 18 place place VBP cord-048487-kamvao4b 138 19 a a DT cord-048487-kamvao4b 138 20 time time NN cord-048487-kamvao4b 138 21 of of IN cord-048487-kamvao4b 138 22 divergence divergence NN cord-048487-kamvao4b 138 23 of of IN cord-048487-kamvao4b 138 24 several several JJ cord-048487-kamvao4b 138 25 hundred hundred CD cord-048487-kamvao4b 138 26 years year NNS cord-048487-kamvao4b 138 27 ago ago RB cord-048487-kamvao4b 138 28 , , , cord-048487-kamvao4b 138 29 ranging range VBG cord-048487-kamvao4b 138 30 from from IN cord-048487-kamvao4b 138 31 the the DT cord-048487-kamvao4b 138 32 16th 16th NN cord-048487-kamvao4b 138 33 to to IN cord-048487-kamvao4b 138 34 early early JJ cord-048487-kamvao4b 138 35 19th 19th JJ cord-048487-kamvao4b 138 36 centuries century NNS cord-048487-kamvao4b 138 37 . . . cord-048487-kamvao4b 139 1 Other other JJ cord-048487-kamvao4b 139 2 analysis analysis NN cord-048487-kamvao4b 139 3 have have VBP cord-048487-kamvao4b 139 4 yielded yield VBN cord-048487-kamvao4b 139 5 estimates estimate NNS cord-048487-kamvao4b 139 6 of of IN cord-048487-kamvao4b 139 7 3600 3600 CD cord-048487-kamvao4b 139 8 years year NNS cord-048487-kamvao4b 139 9 ago ago RB cord-048487-kamvao4b 139 10 [ [ -LRB- cord-048487-kamvao4b 139 11 23 23 CD cord-048487-kamvao4b 139 12 ] ] -RRB- cord-048487-kamvao4b 139 13 . . . cord-048487-kamvao4b 140 1 In in IN cord-048487-kamvao4b 140 2 the the DT cord-048487-kamvao4b 140 3 current current JJ cord-048487-kamvao4b 140 4 study study NN cord-048487-kamvao4b 140 5 , , , cord-048487-kamvao4b 140 6 application application NN cord-048487-kamvao4b 140 7 of of IN cord-048487-kamvao4b 140 8 the the DT cord-048487-kamvao4b 140 9 MCA MCA NNP cord-048487-kamvao4b 140 10 yielded yield VBD cord-048487-kamvao4b 140 11 estimates estimate NNS cord-048487-kamvao4b 140 12 in in IN cord-048487-kamvao4b 140 13 the the DT cord-048487-kamvao4b 140 14 last last JJ cord-048487-kamvao4b 140 15 half half NN cord-048487-kamvao4b 140 16 of of IN cord-048487-kamvao4b 140 17 the the DT cord-048487-kamvao4b 140 18 18th 18th JJ cord-048487-kamvao4b 140 19 century century NN cord-048487-kamvao4b 140 20 . . . cord-048487-kamvao4b 141 1 However however RB cord-048487-kamvao4b 141 2 , , , cord-048487-kamvao4b 141 3 applying apply VBG cord-048487-kamvao4b 141 4 the the DT cord-048487-kamvao4b 141 5 pairwise pairwise JJ cord-048487-kamvao4b 141 6 rate rate NN cord-048487-kamvao4b 141 7 estimate estimate NN cord-048487-kamvao4b 141 8 developed develop VBN cord-048487-kamvao4b 141 9 above above IN cord-048487-kamvao4b 141 10 we -PRON- PRP cord-048487-kamvao4b 141 11 find find VBP cord-048487-kamvao4b 141 12 uniformly uniformly RB cord-048487-kamvao4b 141 13 , , , cord-048487-kamvao4b 141 14 across across IN cord-048487-kamvao4b 141 15 genes gene NNS cord-048487-kamvao4b 141 16 , , , cord-048487-kamvao4b 141 17 that that IN cord-048487-kamvao4b 141 18 the the DT cord-048487-kamvao4b 141 19 divergence divergence NN cord-048487-kamvao4b 141 20 likely likely RB cord-048487-kamvao4b 141 21 occurred occur VBD cord-048487-kamvao4b 141 22 in in IN cord-048487-kamvao4b 141 23 the the DT cord-048487-kamvao4b 141 24 very very RB cord-048487-kamvao4b 141 25 early early JJ cord-048487-kamvao4b 141 26 20th 20th JJ cord-048487-kamvao4b 141 27 century century NN cord-048487-kamvao4b 141 28 . . . cord-048487-kamvao4b 142 1 The the DT cord-048487-kamvao4b 142 2 discrepancy discrepancy NN cord-048487-kamvao4b 142 3 between between IN cord-048487-kamvao4b 142 4 these these DT cord-048487-kamvao4b 142 5 two two CD cord-048487-kamvao4b 142 6 measures measure NNS cord-048487-kamvao4b 142 7 is be VBZ cord-048487-kamvao4b 142 8 likely likely JJ cord-048487-kamvao4b 142 9 due due IN cord-048487-kamvao4b 142 10 to to IN cord-048487-kamvao4b 142 11 the the DT cord-048487-kamvao4b 142 12 increased increase VBN cord-048487-kamvao4b 142 13 modeling modeling NN cord-048487-kamvao4b 142 14 flexibility flexibility NN cord-048487-kamvao4b 142 15 of of IN cord-048487-kamvao4b 142 16 the the DT cord-048487-kamvao4b 142 17 pairwise pairwise JJ cord-048487-kamvao4b 142 18 rate rate NN cord-048487-kamvao4b 142 19 estimate estimate NN cord-048487-kamvao4b 142 20 relative relative JJ cord-048487-kamvao4b 142 21 to to IN cord-048487-kamvao4b 142 22 the the DT cord-048487-kamvao4b 142 23 MCA MCA NNP cord-048487-kamvao4b 142 24 . . . cord-048487-kamvao4b 143 1 This this DT cord-048487-kamvao4b 143 2 discrepancy discrepancy NN cord-048487-kamvao4b 143 3 between between IN cord-048487-kamvao4b 143 4 the the DT cord-048487-kamvao4b 143 5 rates rate NNS cord-048487-kamvao4b 143 6 and and CC cord-048487-kamvao4b 143 7 corresponding correspond VBG cord-048487-kamvao4b 143 8 TMCRA TMCRA NNP cord-048487-kamvao4b 143 9 estimates estimate NNS cord-048487-kamvao4b 143 10 has have VBZ cord-048487-kamvao4b 143 11 important important JJ cord-048487-kamvao4b 143 12 biological biological JJ cord-048487-kamvao4b 143 13 consequence consequence NN cord-048487-kamvao4b 143 14 . . . cord-048487-kamvao4b 144 1 The the DT cord-048487-kamvao4b 144 2 phylogenetic phylogenetic JJ cord-048487-kamvao4b 144 3 divergence divergence NN cord-048487-kamvao4b 144 4 between between IN cord-048487-kamvao4b 144 5 subtype subtype NN cord-048487-kamvao4b 144 6 A a NN cord-048487-kamvao4b 144 7 - - : cord-048487-kamvao4b 144 8 H3N2 H3N2 NNP cord-048487-kamvao4b 144 9 and and CC cord-048487-kamvao4b 144 10 B B NNP cord-048487-kamvao4b 144 11 corresponds correspond VBZ cord-048487-kamvao4b 144 12 to to IN cord-048487-kamvao4b 144 13 a a DT cord-048487-kamvao4b 144 14 subspeciation subspeciation NN cord-048487-kamvao4b 144 15 event event NN cord-048487-kamvao4b 144 16 for for IN cord-048487-kamvao4b 144 17 the the DT cord-048487-kamvao4b 144 18 virus virus NN cord-048487-kamvao4b 144 19 . . . cord-048487-kamvao4b 145 1 The the DT cord-048487-kamvao4b 145 2 results result NNS cord-048487-kamvao4b 145 3 in in IN cord-048487-kamvao4b 145 4 this this DT cord-048487-kamvao4b 145 5 study study NN cord-048487-kamvao4b 145 6 indicate indicate VBP cord-048487-kamvao4b 145 7 that that IN cord-048487-kamvao4b 145 8 the the DT cord-048487-kamvao4b 145 9 process process NN cord-048487-kamvao4b 145 10 of of IN cord-048487-kamvao4b 145 11 speciation speciation NN cord-048487-kamvao4b 145 12 is be VBZ cord-048487-kamvao4b 145 13 not not RB cord-048487-kamvao4b 145 14 neutral neutral JJ cord-048487-kamvao4b 145 15 but but CC cord-048487-kamvao4b 145 16 instead instead RB cord-048487-kamvao4b 145 17 a a DT cord-048487-kamvao4b 145 18 period period NN cord-048487-kamvao4b 145 19 of of IN cord-048487-kamvao4b 145 20 rapid rapid JJ cord-048487-kamvao4b 145 21 and and CC cord-048487-kamvao4b 145 22 intense intense JJ cord-048487-kamvao4b 145 23 genetic genetic JJ cord-048487-kamvao4b 145 24 change change NN cord-048487-kamvao4b 145 25 . . . cord-048487-kamvao4b 146 1 The the DT cord-048487-kamvao4b 146 2 three three CD cord-048487-kamvao4b 146 3 genes gene NNS cord-048487-kamvao4b 146 4 studied study VBN cord-048487-kamvao4b 146 5 here here RB cord-048487-kamvao4b 146 6 consistently consistently RB cord-048487-kamvao4b 146 7 show show VBP cord-048487-kamvao4b 146 8 large large JJ cord-048487-kamvao4b 146 9 acceleration acceleration NN cord-048487-kamvao4b 146 10 in in IN cord-048487-kamvao4b 146 11 the the DT cord-048487-kamvao4b 146 12 rate rate NN cord-048487-kamvao4b 146 13 of of IN cord-048487-kamvao4b 146 14 nucleotide nucleotide JJ cord-048487-kamvao4b 146 15 substitution substitution NN cord-048487-kamvao4b 146 16 for for IN cord-048487-kamvao4b 146 17 the the DT cord-048487-kamvao4b 146 18 divergence divergence NN cord-048487-kamvao4b 146 19 period period NN cord-048487-kamvao4b 146 20 relative relative JJ cord-048487-kamvao4b 146 21 to to IN cord-048487-kamvao4b 146 22 the the DT cord-048487-kamvao4b 146 23 rates rate NNS cord-048487-kamvao4b 146 24 observed observe VBN cord-048487-kamvao4b 146 25 within within IN cord-048487-kamvao4b 146 26 a a DT cord-048487-kamvao4b 146 27 stable stable JJ cord-048487-kamvao4b 146 28 subtype subtype NN cord-048487-kamvao4b 146 29 . . . cord-048487-kamvao4b 147 1 This this DT cord-048487-kamvao4b 147 2 study study NN cord-048487-kamvao4b 147 3 gives give VBZ cord-048487-kamvao4b 147 4 strong strong JJ cord-048487-kamvao4b 147 5 evidence evidence NN cord-048487-kamvao4b 147 6 that that IN cord-048487-kamvao4b 147 7 , , , cord-048487-kamvao4b 147 8 at at IN cord-048487-kamvao4b 147 9 least least JJS cord-048487-kamvao4b 147 10 for for IN cord-048487-kamvao4b 147 11 influenza influenza JJ cord-048487-kamvao4b 147 12 viral viral JJ cord-048487-kamvao4b 147 13 subtype subtype NN cord-048487-kamvao4b 147 14 divergence divergence NN cord-048487-kamvao4b 147 15 , , , cord-048487-kamvao4b 147 16 the the DT cord-048487-kamvao4b 147 17 process process NN cord-048487-kamvao4b 147 18 of of IN cord-048487-kamvao4b 147 19 subspeciation subspeciation NN cord-048487-kamvao4b 147 20 is be VBZ cord-048487-kamvao4b 147 21 associated associate VBN cord-048487-kamvao4b 147 22 not not RB cord-048487-kamvao4b 147 23 just just RB cord-048487-kamvao4b 147 24 with with IN cord-048487-kamvao4b 147 25 large large JJ cord-048487-kamvao4b 147 26 genomic genomic JJ cord-048487-kamvao4b 147 27 changes change NNS cord-048487-kamvao4b 147 28 but but CC cord-048487-kamvao4b 147 29 also also RB cord-048487-kamvao4b 147 30 with with IN cord-048487-kamvao4b 147 31 an an DT cord-048487-kamvao4b 147 32 accelerated accelerated JJ cord-048487-kamvao4b 147 33 , , , cord-048487-kamvao4b 147 34 finite finite JJ cord-048487-kamvao4b 147 35 process process NN cord-048487-kamvao4b 147 36 of of IN cord-048487-kamvao4b 147 37 adaption adaption NN cord-048487-kamvao4b 147 38 . . . cord-048487-kamvao4b 148 1 Histograms Histograms NNP cord-048487-kamvao4b 148 2 of of IN cord-048487-kamvao4b 148 3 the the DT cord-048487-kamvao4b 148 4 time time NN cord-048487-kamvao4b 148 5 of of IN cord-048487-kamvao4b 148 6 most most RBS cord-048487-kamvao4b 148 7 recent recent JJ cord-048487-kamvao4b 148 8 common common JJ cord-048487-kamvao4b 148 9 ancestor ancestor NN cord-048487-kamvao4b 148 10 of of IN cord-048487-kamvao4b 148 11 subtypes subtype NNS cord-048487-kamvao4b 148 12 A A NNP cord-048487-kamvao4b 148 13 - - : cord-048487-kamvao4b 148 14 H3N2 H3N2 NNP cord-048487-kamvao4b 148 15 and and CC cord-048487-kamvao4b 148 16 B B NNP cord-048487-kamvao4b 148 17 , , , cord-048487-kamvao4b 148 18 derived derive VBN cord-048487-kamvao4b 148 19 from from IN cord-048487-kamvao4b 148 20 molecular molecular JJ cord-048487-kamvao4b 148 21 clock clock NN cord-048487-kamvao4b 148 22 estimates estimate NNS cord-048487-kamvao4b 148 23 ( ( -LRB- cord-048487-kamvao4b 148 24 light light JJ cord-048487-kamvao4b 148 25 grey grey NN cord-048487-kamvao4b 148 26 ) ) -RRB- cord-048487-kamvao4b 148 27 and and CC cord-048487-kamvao4b 148 28 pairwise pairwise JJ cord-048487-kamvao4b 148 29 estimates estimate NNS cord-048487-kamvao4b 148 30 ( ( -LRB- cord-048487-kamvao4b 148 31 dark dark NNP cord-048487-kamvao4b 148 32 grey grey NNP cord-048487-kamvao4b 148 33 ) ) -RRB- cord-048487-kamvao4b 148 34 on on IN cord-048487-kamvao4b 148 35 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 148 36 ( ( -LRB- cord-048487-kamvao4b 148 37 HA HA NNP cord-048487-kamvao4b 148 38 ) ) -RRB- cord-048487-kamvao4b 148 39 , , , cord-048487-kamvao4b 148 40 matrix matrix NN cord-048487-kamvao4b 148 41 ( ( -LRB- cord-048487-kamvao4b 148 42 MP MP NNP cord-048487-kamvao4b 148 43 ) ) -RRB- cord-048487-kamvao4b 148 44 and and CC cord-048487-kamvao4b 148 45 nonstructural nonstructural JJ cord-048487-kamvao4b 148 46 ( ( -LRB- cord-048487-kamvao4b 148 47 NS NS NNP cord-048487-kamvao4b 148 48 ) ) -RRB- cord-048487-kamvao4b 148 49 gene gene NN cord-048487-kamvao4b 148 50 sequences sequence NNS cord-048487-kamvao4b 148 51 Assuming assume VBG cord-048487-kamvao4b 148 52 that that IN cord-048487-kamvao4b 148 53 the the DT cord-048487-kamvao4b 148 54 more more RBR cord-048487-kamvao4b 148 55 recent recent JJ cord-048487-kamvao4b 148 56 estimate estimate NN cord-048487-kamvao4b 148 57 is be VBZ cord-048487-kamvao4b 148 58 correct correct JJ cord-048487-kamvao4b 148 59 , , , cord-048487-kamvao4b 148 60 a a DT cord-048487-kamvao4b 148 61 subsequent subsequent JJ cord-048487-kamvao4b 148 62 question question NN cord-048487-kamvao4b 148 63 is be VBZ cord-048487-kamvao4b 148 64 whether whether IN cord-048487-kamvao4b 148 65 or or CC cord-048487-kamvao4b 148 66 not not RB cord-048487-kamvao4b 148 67 a a DT cord-048487-kamvao4b 148 68 pandemic pandemic NN cord-048487-kamvao4b 148 69 or or CC cord-048487-kamvao4b 148 70 epidemic epidemic NN cord-048487-kamvao4b 148 71 associates associate NNS cord-048487-kamvao4b 148 72 with with IN cord-048487-kamvao4b 148 73 subtype subtype NN cord-048487-kamvao4b 148 74 A A NNP cord-048487-kamvao4b 148 75 - - : cord-048487-kamvao4b 148 76 H3N2 H3N2 NNP cord-048487-kamvao4b 148 77 / / SYM cord-048487-kamvao4b 148 78 B b NN cord-048487-kamvao4b 148 79 divergence divergence NN cord-048487-kamvao4b 148 80 . . . cord-048487-kamvao4b 149 1 In in IN cord-048487-kamvao4b 149 2 the the DT cord-048487-kamvao4b 149 3 twentieth twentieth JJ cord-048487-kamvao4b 149 4 century century NN cord-048487-kamvao4b 149 5 , , , cord-048487-kamvao4b 149 6 all all DT cord-048487-kamvao4b 149 7 influenza influenza NN cord-048487-kamvao4b 149 8 pandemics pandemic NNS cord-048487-kamvao4b 149 9 associate associate VBP cord-048487-kamvao4b 149 10 with with IN cord-048487-kamvao4b 149 11 the the DT cord-048487-kamvao4b 149 12 emergence emergence NN cord-048487-kamvao4b 149 13 or or CC cord-048487-kamvao4b 149 14 reemergence reemergence NN cord-048487-kamvao4b 149 15 of of IN cord-048487-kamvao4b 149 16 subtypes subtype NNS cord-048487-kamvao4b 149 17 ( ( -LRB- cord-048487-kamvao4b 149 18 A A NNP cord-048487-kamvao4b 149 19 - - HYPH cord-048487-kamvao4b 149 20 H1N1 H1N1 NNP cord-048487-kamvao4b 149 21 in in IN cord-048487-kamvao4b 149 22 1918 1918 CD cord-048487-kamvao4b 149 23 , , , cord-048487-kamvao4b 149 24 A A NNP cord-048487-kamvao4b 149 25 - - : cord-048487-kamvao4b 149 26 H2N2 H2N2 NNP cord-048487-kamvao4b 149 27 in in IN cord-048487-kamvao4b 149 28 1957 1957 CD cord-048487-kamvao4b 149 29 and and CC cord-048487-kamvao4b 149 30 A A NNP cord-048487-kamvao4b 149 31 - - : cord-048487-kamvao4b 149 32 H3N2 H3N2 NNP cord-048487-kamvao4b 149 33 in in IN cord-048487-kamvao4b 149 34 1968 1968 CD cord-048487-kamvao4b 149 35 ) ) -RRB- cord-048487-kamvao4b 149 36 . . . cord-048487-kamvao4b 150 1 Serological serological JJ cord-048487-kamvao4b 150 2 analysis analysis NN cord-048487-kamvao4b 150 3 indicates indicate VBZ cord-048487-kamvao4b 150 4 that that IN cord-048487-kamvao4b 150 5 the the DT cord-048487-kamvao4b 150 6 1897 1897 CD cord-048487-kamvao4b 150 7 pandemic pandemic NN cord-048487-kamvao4b 150 8 was be VBD cord-048487-kamvao4b 150 9 likely likely JJ cord-048487-kamvao4b 150 10 due due IN cord-048487-kamvao4b 150 11 to to IN cord-048487-kamvao4b 150 12 subtype subtype NN cord-048487-kamvao4b 150 13 A A NNP cord-048487-kamvao4b 150 14 - - : cord-048487-kamvao4b 150 15 H2N2 H2N2 NNP cord-048487-kamvao4b 150 16 . . . cord-048487-kamvao4b 151 1 However however RB cord-048487-kamvao4b 151 2 , , , cord-048487-kamvao4b 151 3 the the DT cord-048487-kamvao4b 151 4 pandemic pandemic NN cord-048487-kamvao4b 151 5 of of IN cord-048487-kamvao4b 151 6 1900 1900 CD cord-048487-kamvao4b 151 7 is be VBZ cord-048487-kamvao4b 151 8 of of IN cord-048487-kamvao4b 151 9 uncertain uncertain JJ cord-048487-kamvao4b 151 10 type type NN cord-048487-kamvao4b 151 11 , , , cord-048487-kamvao4b 151 12 although although IN cord-048487-kamvao4b 151 13 it -PRON- PRP cord-048487-kamvao4b 151 14 is be VBZ cord-048487-kamvao4b 151 15 commonly commonly RB cord-048487-kamvao4b 151 16 reported report VBN cord-048487-kamvao4b 151 17 in in IN cord-048487-kamvao4b 151 18 the the DT cord-048487-kamvao4b 151 19 literature literature NN cord-048487-kamvao4b 151 20 as as IN cord-048487-kamvao4b 151 21 being be VBG cord-048487-kamvao4b 151 22 due due JJ cord-048487-kamvao4b 151 23 to to IN cord-048487-kamvao4b 151 24 A a DT cord-048487-kamvao4b 151 25 - - : cord-048487-kamvao4b 151 26 H3N2 H3N2 NNP cord-048487-kamvao4b 151 27 [ [ -LRB- cord-048487-kamvao4b 151 28 4 4 CD cord-048487-kamvao4b 151 29 ] ] -RRB- cord-048487-kamvao4b 151 30 . . . cord-048487-kamvao4b 152 1 The the DT cord-048487-kamvao4b 152 2 above above JJ cord-048487-kamvao4b 152 3 analysis analysis NN cord-048487-kamvao4b 152 4 suggests suggest VBZ cord-048487-kamvao4b 152 5 that that IN cord-048487-kamvao4b 152 6 it -PRON- PRP cord-048487-kamvao4b 152 7 is be VBZ cord-048487-kamvao4b 152 8 possible possible JJ cord-048487-kamvao4b 152 9 to to TO cord-048487-kamvao4b 152 10 postulate postulate VB cord-048487-kamvao4b 152 11 that that IN cord-048487-kamvao4b 152 12 the the DT cord-048487-kamvao4b 152 13 cause cause NN cord-048487-kamvao4b 152 14 of of IN cord-048487-kamvao4b 152 15 this this DT cord-048487-kamvao4b 152 16 pandemic pandemic NN cord-048487-kamvao4b 152 17 is be VBZ cord-048487-kamvao4b 152 18 due due JJ cord-048487-kamvao4b 152 19 to to IN cord-048487-kamvao4b 152 20 the the DT cord-048487-kamvao4b 152 21 emergence emergence NN cord-048487-kamvao4b 152 22 of of IN cord-048487-kamvao4b 152 23 subtype subtype NN cord-048487-kamvao4b 152 24 A a NN cord-048487-kamvao4b 152 25 - - : cord-048487-kamvao4b 152 26 H3N2 H3N2 NNP cord-048487-kamvao4b 152 27 or or CC cord-048487-kamvao4b 152 28 B. B. NNP cord-048487-kamvao4b 153 1 As as IN cord-048487-kamvao4b 153 2 noted note VBN cord-048487-kamvao4b 153 3 above above RB cord-048487-kamvao4b 153 4 , , , cord-048487-kamvao4b 153 5 we -PRON- PRP cord-048487-kamvao4b 153 6 condition condition VBP cord-048487-kamvao4b 153 7 the the DT cord-048487-kamvao4b 153 8 results result NNS cord-048487-kamvao4b 153 9 presented present VBN cord-048487-kamvao4b 153 10 here here RB cord-048487-kamvao4b 153 11 on on IN cord-048487-kamvao4b 153 12 a a DT cord-048487-kamvao4b 153 13 specific specific JJ cord-048487-kamvao4b 153 14 sequence sequence NN cord-048487-kamvao4b 153 15 alignment alignment NN cord-048487-kamvao4b 153 16 . . . cord-048487-kamvao4b 154 1 As as IN cord-048487-kamvao4b 154 2 the the DT cord-048487-kamvao4b 154 3 question question NN cord-048487-kamvao4b 154 4 under under IN cord-048487-kamvao4b 154 5 consideration consideration NN cord-048487-kamvao4b 154 6 concerns concern VBZ cord-048487-kamvao4b 154 7 the the DT cord-048487-kamvao4b 154 8 divergence divergence NN cord-048487-kamvao4b 154 9 of of IN cord-048487-kamvao4b 154 10 specific specific JJ cord-048487-kamvao4b 154 11 genes gene NNS cord-048487-kamvao4b 154 12 and and CC cord-048487-kamvao4b 154 13 proteins protein NNS cord-048487-kamvao4b 154 14 over over IN cord-048487-kamvao4b 154 15 a a DT cord-048487-kamvao4b 154 16 ( ( -LRB- cord-048487-kamvao4b 154 17 presumably presumably RB cord-048487-kamvao4b 154 18 ) ) -RRB- cord-048487-kamvao4b 154 19 long long JJ cord-048487-kamvao4b 154 20 time time NN cord-048487-kamvao4b 154 21 scale scale NN cord-048487-kamvao4b 154 22 , , , cord-048487-kamvao4b 154 23 the the DT cord-048487-kamvao4b 154 24 capacity capacity NN cord-048487-kamvao4b 154 25 to to TO cord-048487-kamvao4b 154 26 generate generate VB cord-048487-kamvao4b 154 27 reasonable reasonable JJ cord-048487-kamvao4b 154 28 alignments alignment NNS cord-048487-kamvao4b 154 29 diminishes diminish VBZ cord-048487-kamvao4b 154 30 with with IN cord-048487-kamvao4b 154 31 increasing increase VBG cord-048487-kamvao4b 154 32 time time NN cord-048487-kamvao4b 154 33 of of IN cord-048487-kamvao4b 154 34 divergence divergence NN cord-048487-kamvao4b 154 35 between between IN cord-048487-kamvao4b 154 36 types type NNS cord-048487-kamvao4b 154 37 , , , cord-048487-kamvao4b 154 38 conditional conditional JJ cord-048487-kamvao4b 154 39 on on IN cord-048487-kamvao4b 154 40 the the DT cord-048487-kamvao4b 154 41 rate rate NN cord-048487-kamvao4b 154 42 of of IN cord-048487-kamvao4b 154 43 substitution substitution NN cord-048487-kamvao4b 154 44 . . . cord-048487-kamvao4b 155 1 We -PRON- PRP cord-048487-kamvao4b 155 2 find find VBP cord-048487-kamvao4b 155 3 that that IN cord-048487-kamvao4b 155 4 for for IN cord-048487-kamvao4b 155 5 the the DT cord-048487-kamvao4b 155 6 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 155 7 gene gene NN cord-048487-kamvao4b 155 8 , , , cord-048487-kamvao4b 155 9 a a DT cord-048487-kamvao4b 155 10 proportion proportion NN cord-048487-kamvao4b 155 11 of of IN cord-048487-kamvao4b 155 12 sequence sequence NN cord-048487-kamvao4b 155 13 alignments alignment NNS cord-048487-kamvao4b 155 14 support support VBP cord-048487-kamvao4b 155 15 the the DT cord-048487-kamvao4b 155 16 split split NN cord-048487-kamvao4b 155 17 of of IN cord-048487-kamvao4b 155 18 subtype subtype NN cord-048487-kamvao4b 155 19 B b NN cord-048487-kamvao4b 155 20 from from IN cord-048487-kamvao4b 155 21 subtype subtype NN cord-048487-kamvao4b 155 22 C c NN cord-048487-kamvao4b 155 23 after after IN cord-048487-kamvao4b 155 24 the the DT cord-048487-kamvao4b 155 25 split split NN cord-048487-kamvao4b 155 26 between between IN cord-048487-kamvao4b 155 27 subtypes subtype NNS cord-048487-kamvao4b 155 28 A A NNP cord-048487-kamvao4b 155 29 - - : cord-048487-kamvao4b 155 30 H3N2 H3N2 NNP cord-048487-kamvao4b 155 31 and and CC cord-048487-kamvao4b 155 32 B B NNP cord-048487-kamvao4b 155 33 , , , cord-048487-kamvao4b 155 34 in in IN cord-048487-kamvao4b 155 35 opposition opposition NN cord-048487-kamvao4b 155 36 to to IN cord-048487-kamvao4b 155 37 the the DT cord-048487-kamvao4b 155 38 topology topology NN cord-048487-kamvao4b 155 39 enforced enforce VBN cord-048487-kamvao4b 155 40 in in IN cord-048487-kamvao4b 155 41 our -PRON- PRP$ cord-048487-kamvao4b 155 42 analysis analysis NN cord-048487-kamvao4b 155 43 . . . cord-048487-kamvao4b 156 1 Hence hence RB cord-048487-kamvao4b 156 2 , , , cord-048487-kamvao4b 156 3 to to IN cord-048487-kamvao4b 156 4 some some DT cord-048487-kamvao4b 156 5 unknown unknown JJ cord-048487-kamvao4b 156 6 degree degree NN cord-048487-kamvao4b 156 7 , , , cord-048487-kamvao4b 156 8 our -PRON- PRP$ cord-048487-kamvao4b 156 9 analysis analysis NN cord-048487-kamvao4b 156 10 is be VBZ cord-048487-kamvao4b 156 11 necessarily necessarily RB cord-048487-kamvao4b 156 12 biased bias VBN cord-048487-kamvao4b 156 13 by by IN cord-048487-kamvao4b 156 14 the the DT cord-048487-kamvao4b 156 15 choice choice NN cord-048487-kamvao4b 156 16 of of IN cord-048487-kamvao4b 156 17 alignment alignment NN cord-048487-kamvao4b 156 18 . . . cord-048487-kamvao4b 157 1 This this DT cord-048487-kamvao4b 157 2 suggests suggest VBZ cord-048487-kamvao4b 157 3 that that IN cord-048487-kamvao4b 157 4 improved improve VBN cord-048487-kamvao4b 157 5 dating dating NN cord-048487-kamvao4b 157 6 can can MD cord-048487-kamvao4b 157 7 be be VB cord-048487-kamvao4b 157 8 found find VBN cord-048487-kamvao4b 157 9 by by IN cord-048487-kamvao4b 157 10 integrating integrate VBG cord-048487-kamvao4b 157 11 estimation estimation NN cord-048487-kamvao4b 157 12 procedures procedure NNS cord-048487-kamvao4b 157 13 over over IN cord-048487-kamvao4b 157 14 an an DT cord-048487-kamvao4b 157 15 ensemble ensemble NN cord-048487-kamvao4b 157 16 of of IN cord-048487-kamvao4b 157 17 alignments alignment NNS cord-048487-kamvao4b 157 18 [ [ -LRB- cord-048487-kamvao4b 157 19 19 19 CD cord-048487-kamvao4b 157 20 ] ] -RRB- cord-048487-kamvao4b 157 21 . . . cord-048487-kamvao4b 158 1 The the DT cord-048487-kamvao4b 158 2 pairwise pairwise JJ cord-048487-kamvao4b 158 3 estimate estimate NN cord-048487-kamvao4b 158 4 method method NN cord-048487-kamvao4b 158 5 presented present VBN cord-048487-kamvao4b 158 6 above above RB cord-048487-kamvao4b 158 7 is be VBZ cord-048487-kamvao4b 158 8 accurate accurate JJ cord-048487-kamvao4b 158 9 in in IN cord-048487-kamvao4b 158 10 the the DT cord-048487-kamvao4b 158 11 scale scale NN cord-048487-kamvao4b 158 12 where where WRB cord-048487-kamvao4b 158 13 is be VBZ cord-048487-kamvao4b 158 14 the the DT cord-048487-kamvao4b 158 15 total total JJ cord-048487-kamvao4b 158 16 time time NN cord-048487-kamvao4b 158 17 over over IN cord-048487-kamvao4b 158 18 the the DT cord-048487-kamvao4b 158 19 phylogeny phylogeny NN cord-048487-kamvao4b 158 20 and and CC cord-048487-kamvao4b 158 21 p p NN cord-048487-kamvao4b 158 22 is be VBZ cord-048487-kamvao4b 158 23 mean mean JJ cord-048487-kamvao4b 158 24 rate rate NN cord-048487-kamvao4b 158 25 over over IN cord-048487-kamvao4b 158 26 the the DT cord-048487-kamvao4b 158 27 phylogeny phylogeny NN cord-048487-kamvao4b 158 28 [ [ -LRB- cord-048487-kamvao4b 158 29 26 26 CD cord-048487-kamvao4b 158 30 ] ] -RRB- cord-048487-kamvao4b 158 31 . . . cord-048487-kamvao4b 159 1 This this DT cord-048487-kamvao4b 159 2 relation relation NN cord-048487-kamvao4b 159 3 dictates dictate VBZ cord-048487-kamvao4b 159 4 that that IN cord-048487-kamvao4b 159 5 as as IN cord-048487-kamvao4b 159 6 divergence divergence NN cord-048487-kamvao4b 159 7 events event NNS cord-048487-kamvao4b 159 8 become become VBP cord-048487-kamvao4b 159 9 more more RBR cord-048487-kamvao4b 159 10 remote remote JJ cord-048487-kamvao4b 159 11 the the DT cord-048487-kamvao4b 159 12 ability ability NN cord-048487-kamvao4b 159 13 of of IN cord-048487-kamvao4b 159 14 the the DT cord-048487-kamvao4b 159 15 triplet triplet NN cord-048487-kamvao4b 159 16 method method NN cord-048487-kamvao4b 159 17 to to TO cord-048487-kamvao4b 159 18 resolve resolve VB cord-048487-kamvao4b 159 19 the the DT cord-048487-kamvao4b 159 20 time time NN cord-048487-kamvao4b 159 21 of of IN cord-048487-kamvao4b 159 22 divergence divergence NN cord-048487-kamvao4b 159 23 diminishes diminish VBZ cord-048487-kamvao4b 159 24 . . . cord-048487-kamvao4b 160 1 While while IN cord-048487-kamvao4b 160 2 this this DT cord-048487-kamvao4b 160 3 limit limit NN cord-048487-kamvao4b 160 4 prohibits prohibit VBZ cord-048487-kamvao4b 160 5 the the DT cord-048487-kamvao4b 160 6 calculation calculation NN cord-048487-kamvao4b 160 7 of of IN cord-048487-kamvao4b 160 8 remote remote JJ cord-048487-kamvao4b 160 9 divergence divergence NN cord-048487-kamvao4b 160 10 events event NNS cord-048487-kamvao4b 160 11 , , , cord-048487-kamvao4b 160 12 the the DT cord-048487-kamvao4b 160 13 example example NN cord-048487-kamvao4b 160 14 presented present VBN cord-048487-kamvao4b 160 15 above above IN cord-048487-kamvao4b 160 16 lies lie NNS cord-048487-kamvao4b 160 17 within within IN cord-048487-kamvao4b 160 18 the the DT cord-048487-kamvao4b 160 19 appropriate appropriate JJ cord-048487-kamvao4b 160 20 scale scale NN cord-048487-kamvao4b 160 21 . . . cord-048487-kamvao4b 161 1 In in IN cord-048487-kamvao4b 161 2 place place NN cord-048487-kamvao4b 161 3 of of IN cord-048487-kamvao4b 161 4 a a DT cord-048487-kamvao4b 161 5 specific specific JJ cord-048487-kamvao4b 161 6 MCA MCA NNP cord-048487-kamvao4b 161 7 , , , cord-048487-kamvao4b 161 8 the the DT cord-048487-kamvao4b 161 9 estimates estimate NNS cord-048487-kamvao4b 161 10 presented present VBN cord-048487-kamvao4b 161 11 here here RB cord-048487-kamvao4b 161 12 directly directly RB cord-048487-kamvao4b 161 13 calculate calculate VB cord-048487-kamvao4b 161 14 the the DT cord-048487-kamvao4b 161 15 rate rate NN cord-048487-kamvao4b 161 16 of of IN cord-048487-kamvao4b 161 17 substitutions substitution NNS cord-048487-kamvao4b 161 18 between between IN cord-048487-kamvao4b 161 19 taxa taxon NNS cord-048487-kamvao4b 161 20 from from IN cord-048487-kamvao4b 161 21 different different JJ cord-048487-kamvao4b 161 22 viral viral JJ cord-048487-kamvao4b 161 23 subtypes subtype NNS cord-048487-kamvao4b 161 24 . . . cord-048487-kamvao4b 162 1 As as IN cord-048487-kamvao4b 162 2 such such JJ cord-048487-kamvao4b 162 3 estimates estimate NNS cord-048487-kamvao4b 162 4 span span VBP cord-048487-kamvao4b 162 5 paths path NNS cord-048487-kamvao4b 162 6 between between IN cord-048487-kamvao4b 162 7 subtypes subtype NNS cord-048487-kamvao4b 162 8 , , , cord-048487-kamvao4b 162 9 they -PRON- PRP cord-048487-kamvao4b 162 10 simultaneously simultaneously RB cord-048487-kamvao4b 162 11 capture capture VBP cord-048487-kamvao4b 162 12 the the DT cord-048487-kamvao4b 162 13 rate rate NN cord-048487-kamvao4b 162 14 evolution evolution NN cord-048487-kamvao4b 162 15 along along IN cord-048487-kamvao4b 162 16 branches branch NNS cord-048487-kamvao4b 162 17 both both CC cord-048487-kamvao4b 162 18 within within IN cord-048487-kamvao4b 162 19 and and CC cord-048487-kamvao4b 162 20 between between IN cord-048487-kamvao4b 162 21 subtypes subtype NNS cord-048487-kamvao4b 162 22 . . . cord-048487-kamvao4b 163 1 From from IN cord-048487-kamvao4b 163 2 these these DT cord-048487-kamvao4b 163 3 estimates estimate NNS cord-048487-kamvao4b 163 4 , , , cord-048487-kamvao4b 163 5 we -PRON- PRP cord-048487-kamvao4b 163 6 are be VBP cord-048487-kamvao4b 163 7 able able JJ cord-048487-kamvao4b 163 8 to to TO cord-048487-kamvao4b 163 9 directly directly RB cord-048487-kamvao4b 163 10 infer infer VB cord-048487-kamvao4b 163 11 the the DT cord-048487-kamvao4b 163 12 time time NN cord-048487-kamvao4b 163 13 of of IN cord-048487-kamvao4b 163 14 divergence divergence NN cord-048487-kamvao4b 163 15 between between IN cord-048487-kamvao4b 163 16 subtypes subtype NNS cord-048487-kamvao4b 163 17 . . . cord-048487-kamvao4b 164 1 As as IN cord-048487-kamvao4b 164 2 a a DT cord-048487-kamvao4b 164 3 trade trade NN cord-048487-kamvao4b 164 4 - - HYPH cord-048487-kamvao4b 164 5 off off NN cord-048487-kamvao4b 164 6 for for IN cord-048487-kamvao4b 164 7 limited limited JJ cord-048487-kamvao4b 164 8 MCAs mca NNS cord-048487-kamvao4b 164 9 , , , cord-048487-kamvao4b 164 10 the the DT cord-048487-kamvao4b 164 11 method method NN cord-048487-kamvao4b 164 12 requires require VBZ cord-048487-kamvao4b 164 13 an an DT cord-048487-kamvao4b 164 14 outgroup outgroup NN cord-048487-kamvao4b 164 15 subtype subtype NN cord-048487-kamvao4b 164 16 to to TO cord-048487-kamvao4b 164 17 function function VB cord-048487-kamvao4b 164 18 as as IN cord-048487-kamvao4b 164 19 an an DT cord-048487-kamvao4b 164 20 origin origin NN cord-048487-kamvao4b 164 21 relative relative JJ cord-048487-kamvao4b 164 22 to to IN cord-048487-kamvao4b 164 23 the the DT cord-048487-kamvao4b 164 24 subtypes subtype NNS cord-048487-kamvao4b 164 25 under under IN cord-048487-kamvao4b 164 26 consideration consideration NN cord-048487-kamvao4b 164 27 . . . cord-048487-kamvao4b 165 1 We -PRON- PRP cord-048487-kamvao4b 165 2 feel feel VBP cord-048487-kamvao4b 165 3 that that IN cord-048487-kamvao4b 165 4 the the DT cord-048487-kamvao4b 165 5 triplet triplet NN cord-048487-kamvao4b 165 6 method method NN cord-048487-kamvao4b 165 7 provides provide VBZ cord-048487-kamvao4b 165 8 a a DT cord-048487-kamvao4b 165 9 simple simple JJ cord-048487-kamvao4b 165 10 and and CC cord-048487-kamvao4b 165 11 widely widely RB cord-048487-kamvao4b 165 12 applicable applicable JJ cord-048487-kamvao4b 165 13 way way NN cord-048487-kamvao4b 165 14 to to TO cord-048487-kamvao4b 165 15 calculate calculate VB cord-048487-kamvao4b 165 16 the the DT cord-048487-kamvao4b 165 17 dates date NNS cord-048487-kamvao4b 165 18 of of IN cord-048487-kamvao4b 165 19 divergence divergence NN cord-048487-kamvao4b 165 20 of of IN cord-048487-kamvao4b 165 21 rapidly rapidly RB cord-048487-kamvao4b 165 22 evolving evolve VBG cord-048487-kamvao4b 165 23 organisms organism NNS cord-048487-kamvao4b 165 24 without without IN cord-048487-kamvao4b 165 25 the the DT cord-048487-kamvao4b 165 26 pitfalls pitfall NNS cord-048487-kamvao4b 165 27 of of IN cord-048487-kamvao4b 165 28 the the DT cord-048487-kamvao4b 165 29 MCA MCA NNP cord-048487-kamvao4b 165 30 . . . cord-048487-kamvao4b 166 1 We -PRON- PRP cord-048487-kamvao4b 166 2 present present VBP cord-048487-kamvao4b 166 3 a a DT cord-048487-kamvao4b 166 4 simple simple JJ cord-048487-kamvao4b 166 5 method method NN cord-048487-kamvao4b 166 6 for for IN cord-048487-kamvao4b 166 7 calculating calculate VBG cord-048487-kamvao4b 166 8 the the DT cord-048487-kamvao4b 166 9 time time NN cord-048487-kamvao4b 166 10 of of IN cord-048487-kamvao4b 166 11 viral viral JJ cord-048487-kamvao4b 166 12 subtype subtype NN cord-048487-kamvao4b 166 13 divergence divergence NN cord-048487-kamvao4b 166 14 that that WDT cord-048487-kamvao4b 166 15 does do VBZ cord-048487-kamvao4b 166 16 not not RB cord-048487-kamvao4b 166 17 assume assume VB cord-048487-kamvao4b 166 18 a a DT cord-048487-kamvao4b 166 19 molecular molecular JJ cord-048487-kamvao4b 166 20 clock clock NN cord-048487-kamvao4b 166 21 over over IN cord-048487-kamvao4b 166 22 the the DT cord-048487-kamvao4b 166 23 entire entire JJ cord-048487-kamvao4b 166 24 phylogeny phylogeny NN cord-048487-kamvao4b 166 25 . . . cord-048487-kamvao4b 167 1 Additionally additionally RB cord-048487-kamvao4b 167 2 , , , cord-048487-kamvao4b 167 3 the the DT cord-048487-kamvao4b 167 4 estimator estimator NN cord-048487-kamvao4b 167 5 of of IN cord-048487-kamvao4b 167 6 this this DT cord-048487-kamvao4b 167 7 method method NN cord-048487-kamvao4b 167 8 , , , cord-048487-kamvao4b 167 9 a a DT cord-048487-kamvao4b 167 10 weighted weighted JJ cord-048487-kamvao4b 167 11 sum sum NN cord-048487-kamvao4b 167 12 of of IN cord-048487-kamvao4b 167 13 pairwise pairwise JJ cord-048487-kamvao4b 167 14 estimates estimate NNS cord-048487-kamvao4b 167 15 , , , cord-048487-kamvao4b 167 16 furnishes furnish VBZ cord-048487-kamvao4b 167 17 a a DT cord-048487-kamvao4b 167 18 defined define VBN cord-048487-kamvao4b 167 19 variance variance NN cord-048487-kamvao4b 167 20 for for IN cord-048487-kamvao4b 167 21 the the DT cord-048487-kamvao4b 167 22 time time NN cord-048487-kamvao4b 167 23 of of IN cord-048487-kamvao4b 167 24 the the DT cord-048487-kamvao4b 167 25 most most RBS cord-048487-kamvao4b 167 26 common common JJ cord-048487-kamvao4b 167 27 ancestor ancestor NN cord-048487-kamvao4b 167 28 between between IN cord-048487-kamvao4b 167 29 subtypes subtype NNS cord-048487-kamvao4b 167 30 . . . cord-048487-kamvao4b 168 1 As as IN cord-048487-kamvao4b 168 2 a a DT cord-048487-kamvao4b 168 3 tradeoff tradeoff NN cord-048487-kamvao4b 168 4 for for IN cord-048487-kamvao4b 168 5 this this DT cord-048487-kamvao4b 168 6 increased increase VBN cord-048487-kamvao4b 168 7 precision precision NN cord-048487-kamvao4b 168 8 , , , cord-048487-kamvao4b 168 9 the the DT cord-048487-kamvao4b 168 10 structure structure NN cord-048487-kamvao4b 168 11 of of IN cord-048487-kamvao4b 168 12 the the DT cord-048487-kamvao4b 168 13 triplet triplet NN cord-048487-kamvao4b 168 14 statistic statistic NN cord-048487-kamvao4b 168 15 requires require VBZ cord-048487-kamvao4b 168 16 an an DT cord-048487-kamvao4b 168 17 outgroup outgroup NN cord-048487-kamvao4b 168 18 set set NN cord-048487-kamvao4b 168 19 of of IN cord-048487-kamvao4b 168 20 sequences sequence NNS cord-048487-kamvao4b 168 21 , , , cord-048487-kamvao4b 168 22 usually usually RB cord-048487-kamvao4b 168 23 a a DT cord-048487-kamvao4b 168 24 closely closely RB cord-048487-kamvao4b 168 25 related relate VBN cord-048487-kamvao4b 168 26 subtype subtype NN cord-048487-kamvao4b 168 27 . . . cord-048487-kamvao4b 169 1 We -PRON- PRP cord-048487-kamvao4b 169 2 apply apply VBP cord-048487-kamvao4b 169 3 this this DT cord-048487-kamvao4b 169 4 estimator estimator NN cord-048487-kamvao4b 169 5 to to IN cord-048487-kamvao4b 169 6 the the DT cord-048487-kamvao4b 169 7 case case NN cord-048487-kamvao4b 169 8 of of IN cord-048487-kamvao4b 169 9 influenza influenza NN cord-048487-kamvao4b 169 10 subtype subtype NN cord-048487-kamvao4b 169 11 divergence divergence NN cord-048487-kamvao4b 169 12 , , , cord-048487-kamvao4b 169 13 considering consider VBG cord-048487-kamvao4b 169 14 three three CD cord-048487-kamvao4b 169 15 genes gene NNS cord-048487-kamvao4b 169 16 . . . cord-048487-kamvao4b 170 1 We -PRON- PRP cord-048487-kamvao4b 170 2 show show VBP cord-048487-kamvao4b 170 3 that that IN cord-048487-kamvao4b 170 4 the the DT cord-048487-kamvao4b 170 5 estimated estimate VBN cord-048487-kamvao4b 170 6 divergence divergence NN cord-048487-kamvao4b 170 7 time time NN cord-048487-kamvao4b 170 8 of of IN cord-048487-kamvao4b 170 9 subtypes subtype NNS cord-048487-kamvao4b 171 1 A a LS cord-048487-kamvao4b 171 2 - - : cord-048487-kamvao4b 171 3 H3N2 H3N2 NNP cord-048487-kamvao4b 171 4 and and CC cord-048487-kamvao4b 171 5 B B NNP cord-048487-kamvao4b 171 6 is be VBZ cord-048487-kamvao4b 171 7 more more JJR cord-048487-kamvao4b 171 8 than than IN cord-048487-kamvao4b 171 9 a a DT cord-048487-kamvao4b 171 10 century century NN cord-048487-kamvao4b 171 11 later later RB cord-048487-kamvao4b 171 12 than than IN cord-048487-kamvao4b 171 13 those those DT cord-048487-kamvao4b 171 14 calculated calculate VBN cord-048487-kamvao4b 171 15 with with IN cord-048487-kamvao4b 171 16 a a DT cord-048487-kamvao4b 171 17 molecular molecular JJ cord-048487-kamvao4b 171 18 clock clock NN cord-048487-kamvao4b 171 19 . . . cord-048487-kamvao4b 172 1 Since since IN cord-048487-kamvao4b 172 2 we -PRON- PRP cord-048487-kamvao4b 172 3 assume assume VBP cord-048487-kamvao4b 172 4 that that IN cord-048487-kamvao4b 172 5 the the DT cord-048487-kamvao4b 172 6 ν ν NNP cord-048487-kamvao4b 172 7 and and CC cord-048487-kamvao4b 172 8 ε ε NNP cord-048487-kamvao4b 172 9 structures structure NNS cord-048487-kamvao4b 172 10 are be VBP cord-048487-kamvao4b 172 11 independent independent JJ cord-048487-kamvao4b 172 12 , , , cord-048487-kamvao4b 172 13 the the DT cord-048487-kamvao4b 172 14 right right JJ cord-048487-kamvao4b 172 15 side side NN cord-048487-kamvao4b 172 16 of of IN cord-048487-kamvao4b 172 17 the the DT cord-048487-kamvao4b 172 18 equation equation NN cord-048487-kamvao4b 172 19 can can MD cord-048487-kamvao4b 172 20 be be VB cord-048487-kamvao4b 172 21 further further RB cord-048487-kamvao4b 172 22 reduced reduce VBN cord-048487-kamvao4b 172 23 , , , cord-048487-kamvao4b 172 24 yielding yield VBG cord-048487-kamvao4b 172 25 Let let NN cord-048487-kamvao4b 173 1 Δt Δt NNS cord-048487-kamvao4b 173 2 = = NFP cord-048487-kamvao4b 174 1 t t NNP cord-048487-kamvao4b 174 2 i i NNP cord-048487-kamvao4b 174 3 -t -t : cord-048487-kamvao4b 174 4 j j NNP cord-048487-kamvao4b 174 5 . . . cord-048487-kamvao4b 175 1 The the DT cord-048487-kamvao4b 175 2 final final JJ cord-048487-kamvao4b 175 3 expectation expectation NN cord-048487-kamvao4b 175 4 on on IN cord-048487-kamvao4b 175 5 the the DT cord-048487-kamvao4b 175 6 right right JJ cord-048487-kamvao4b 175 7 hand hand NN cord-048487-kamvao4b 175 8 side side NN cord-048487-kamvao4b 175 9 resolves resolve VBZ cord-048487-kamvao4b 175 10 by by IN cord-048487-kamvao4b 175 11 direct direct JJ cord-048487-kamvao4b 175 12 integration integration NN cord-048487-kamvao4b 175 13 , , , cord-048487-kamvao4b 175 14 We -PRON- PRP cord-048487-kamvao4b 175 15 note note VBP cord-048487-kamvao4b 175 16 that that IN cord-048487-kamvao4b 175 17 as as IN cord-048487-kamvao4b 175 18 the the DT cord-048487-kamvao4b 175 19 sampling sampling NN cord-048487-kamvao4b 175 20 time time NN cord-048487-kamvao4b 175 21 is be VBZ cord-048487-kamvao4b 175 22 independent independent JJ cord-048487-kamvao4b 175 23 of of IN cord-048487-kamvao4b 175 24 the the DT cord-048487-kamvao4b 175 25 rate rate NN cord-048487-kamvao4b 175 26 of of IN cord-048487-kamvao4b 175 27 nucleotide nucleotide JJ cord-048487-kamvao4b 175 28 substitution substitution NN cord-048487-kamvao4b 175 29 , , , cord-048487-kamvao4b 175 30 the the DT cord-048487-kamvao4b 175 31 error error NN cord-048487-kamvao4b 175 32 increases increase VBZ cord-048487-kamvao4b 175 33 in in IN cord-048487-kamvao4b 175 34 proportion proportion NN cord-048487-kamvao4b 175 35 to to IN cord-048487-kamvao4b 175 36 the the DT cord-048487-kamvao4b 175 37 magnitude magnitude NN cord-048487-kamvao4b 175 38 of of IN cord-048487-kamvao4b 175 39 the the DT cord-048487-kamvao4b 175 40 initial initial JJ cord-048487-kamvao4b 175 41 statistic statistic NN cord-048487-kamvao4b 175 42 . . . cord-048487-kamvao4b 176 1 We -PRON- PRP cord-048487-kamvao4b 176 2 can can MD cord-048487-kamvao4b 176 3 then then RB cord-048487-kamvao4b 176 4 create create VB cord-048487-kamvao4b 176 5 a a DT cord-048487-kamvao4b 176 6 new new JJ cord-048487-kamvao4b 176 7 , , , cord-048487-kamvao4b 176 8 unbiased unbiased JJ cord-048487-kamvao4b 176 9 statistic statistic NN cord-048487-kamvao4b 176 10 by by IN cord-048487-kamvao4b 176 11 counterbalancing counterbalance VBG cord-048487-kamvao4b 176 12 the the DT cord-048487-kamvao4b 176 13 original original JJ cord-048487-kamvao4b 176 14 statistic statistic NN cord-048487-kamvao4b 176 15 with with IN cord-048487-kamvao4b 176 16 this this DT cord-048487-kamvao4b 176 17 factor factor NN cord-048487-kamvao4b 176 18 , , , cord-048487-kamvao4b 176 19 making make VBG cord-048487-kamvao4b 176 20 a a DT cord-048487-kamvao4b 176 21 new new JJ cord-048487-kamvao4b 176 22 statistic statistic NN cord-048487-kamvao4b 176 23 Influenza Influenza NNP cord-048487-kamvao4b 176 24 virus virus NN cord-048487-kamvao4b 176 25 genetics genetics NN cord-048487-kamvao4b 176 26 Epidemiology epidemiology NN cord-048487-kamvao4b 176 27 of of IN cord-048487-kamvao4b 176 28 influenza influenza NN cord-048487-kamvao4b 176 29 C C NNP cord-048487-kamvao4b 176 30 virus virus NN cord-048487-kamvao4b 176 31 in in IN cord-048487-kamvao4b 176 32 man man NN cord-048487-kamvao4b 176 33 : : : cord-048487-kamvao4b 176 34 multiple multiple JJ cord-048487-kamvao4b 176 35 evolutionary evolutionary JJ cord-048487-kamvao4b 176 36 lineages lineage NNS cord-048487-kamvao4b 176 37 and and CC cord-048487-kamvao4b 176 38 low low JJ cord-048487-kamvao4b 176 39 rate rate NN cord-048487-kamvao4b 176 40 of of IN cord-048487-kamvao4b 176 41 change change NN cord-048487-kamvao4b 176 42 Avian avian JJ cord-048487-kamvao4b 176 43 influenza influenza NN cord-048487-kamvao4b 176 44 virus virus NN cord-048487-kamvao4b 176 45 exhibits exhibit VBZ cord-048487-kamvao4b 176 46 rapid rapid JJ cord-048487-kamvao4b 176 47 evolutionary evolutionary JJ cord-048487-kamvao4b 176 48 dynamics dynamic NNS cord-048487-kamvao4b 177 1 Influenza influenza NN cord-048487-kamvao4b 177 2 pandemic pandemic NN cord-048487-kamvao4b 177 3 periodicity periodicity NN cord-048487-kamvao4b 177 4 , , , cord-048487-kamvao4b 177 5 virus virus NN cord-048487-kamvao4b 177 6 recycling recycling NN cord-048487-kamvao4b 177 7 , , , cord-048487-kamvao4b 177 8 and and CC cord-048487-kamvao4b 177 9 the the DT cord-048487-kamvao4b 177 10 art art NN cord-048487-kamvao4b 177 11 of of IN cord-048487-kamvao4b 177 12 risk risk NN cord-048487-kamvao4b 177 13 assessment assessment NN cord-048487-kamvao4b 177 14 . . . cord-048487-kamvao4b 178 1 Emerging emerge VBG cord-048487-kamvao4b 178 2 Infectious Infectious NNP cord-048487-kamvao4b 178 3 Diseases Diseases NNPS cord-048487-kamvao4b 179 1 Relaxed relaxed JJ cord-048487-kamvao4b 179 2 phylogenetics phylogenetic NNS cord-048487-kamvao4b 179 3 and and CC cord-048487-kamvao4b 179 4 dating date VBG cord-048487-kamvao4b 179 5 with with IN cord-048487-kamvao4b 179 6 confidence confidence NN cord-048487-kamvao4b 179 7 Estimating estimating NN cord-048487-kamvao4b 179 8 mutation mutation NN cord-048487-kamvao4b 179 9 parameters parameter NNS cord-048487-kamvao4b 179 10 , , , cord-048487-kamvao4b 179 11 population population NN cord-048487-kamvao4b 179 12 history history NN cord-048487-kamvao4b 179 13 and and CC cord-048487-kamvao4b 179 14 genealogy genealogy NN cord-048487-kamvao4b 179 15 simultaneously simultaneously RB cord-048487-kamvao4b 179 16 from from IN cord-048487-kamvao4b 179 17 temporally temporally RB cord-048487-kamvao4b 179 18 spaced space VBN cord-048487-kamvao4b 179 19 sequence sequence NN cord-048487-kamvao4b 179 20 data datum NNS cord-048487-kamvao4b 179 21 Measurably measurably RB cord-048487-kamvao4b 179 22 evolving evolve VBG cord-048487-kamvao4b 179 23 populations population NNS cord-048487-kamvao4b 179 24 Ecological ecological JJ cord-048487-kamvao4b 179 25 and and CC cord-048487-kamvao4b 179 26 immunologial immunologial JJ cord-048487-kamvao4b 179 27 determinants determinant NNS cord-048487-kamvao4b 179 28 of of IN cord-048487-kamvao4b 179 29 influenza influenza NN cord-048487-kamvao4b 179 30 evolution evolution NN cord-048487-kamvao4b 179 31 Dating date VBG cord-048487-kamvao4b 179 32 the the DT cord-048487-kamvao4b 179 33 human human JJ cord-048487-kamvao4b 179 34 - - HYPH cord-048487-kamvao4b 179 35 ape ape NN cord-048487-kamvao4b 179 36 splitting splitting NN cord-048487-kamvao4b 179 37 by by IN cord-048487-kamvao4b 179 38 a a DT cord-048487-kamvao4b 179 39 molecular molecular JJ cord-048487-kamvao4b 179 40 clock clock NN cord-048487-kamvao4b 179 41 of of IN cord-048487-kamvao4b 179 42 mitochondrial mitochondrial JJ cord-048487-kamvao4b 179 43 DNA dna NN cord-048487-kamvao4b 180 1 A a DT cord-048487-kamvao4b 180 2 twenty twenty CD cord-048487-kamvao4b 180 3 - - HYPH cord-048487-kamvao4b 180 4 one one CD cord-048487-kamvao4b 180 5 - - HYPH cord-048487-kamvao4b 180 6 year year NN cord-048487-kamvao4b 180 7 experience experience NN cord-048487-kamvao4b 180 8 with with IN cord-048487-kamvao4b 180 9 anitgenic anitgenic NNP cord-048487-kamvao4b 180 10 variation variation NN cord-048487-kamvao4b 180 11 among among IN cord-048487-kamvao4b 180 12 influenza influenza NN cord-048487-kamvao4b 180 13 B b NN cord-048487-kamvao4b 180 14 viruses virus NNS cord-048487-kamvao4b 181 1 Robust robust JJ cord-048487-kamvao4b 181 2 statistics statistic NNS cord-048487-kamvao4b 181 3 : : : cord-048487-kamvao4b 181 4 A a DT cord-048487-kamvao4b 181 5 review review NN cord-048487-kamvao4b 181 6 Statistical statistical JJ cord-048487-kamvao4b 181 7 estimation estimation NN cord-048487-kamvao4b 181 8 of of IN cord-048487-kamvao4b 181 9 parameters parameter NNS cord-048487-kamvao4b 181 10 in in IN cord-048487-kamvao4b 181 11 a a DT cord-048487-kamvao4b 181 12 phylogenetic phylogenetic JJ cord-048487-kamvao4b 181 13 tree tree NN cord-048487-kamvao4b 181 14 using use VBG cord-048487-kamvao4b 181 15 a a DT cord-048487-kamvao4b 181 16 dynamics dynamic NNS cord-048487-kamvao4b 181 17 model model NN cord-048487-kamvao4b 181 18 of of IN cord-048487-kamvao4b 181 19 the the DT cord-048487-kamvao4b 181 20 substitutional substitutional JJ cord-048487-kamvao4b 181 21 process process NN cord-048487-kamvao4b 181 22 Tracing trace VBG cord-048487-kamvao4b 181 23 the the DT cord-048487-kamvao4b 181 24 origin origin NN cord-048487-kamvao4b 181 25 and and CC cord-048487-kamvao4b 181 26 history history NN cord-048487-kamvao4b 181 27 of of IN cord-048487-kamvao4b 181 28 the the DT cord-048487-kamvao4b 181 29 HIV-2 HIV-2 NNP cord-048487-kamvao4b 181 30 epidemic epidemic NN cord-048487-kamvao4b 181 31 . . . cord-048487-kamvao4b 182 1 Proceeding proceed VBG cord-048487-kamvao4b 182 2 of of IN cord-048487-kamvao4b 182 3 the the DT cord-048487-kamvao4b 182 4 National National NNP cord-048487-kamvao4b 182 5 Academy Academy NNP cord-048487-kamvao4b 182 6 of of IN cord-048487-kamvao4b 182 7 Sciences Sciences NNPS cord-048487-kamvao4b 182 8 Date Date NNP cord-048487-kamvao4b 182 9 of of IN cord-048487-kamvao4b 182 10 origin origin NN cord-048487-kamvao4b 182 11 of of IN cord-048487-kamvao4b 182 12 the the DT cord-048487-kamvao4b 182 13 SARS SARS NNP cord-048487-kamvao4b 182 14 coronavirus coronavirus NN cord-048487-kamvao4b 182 15 strains strain NNS cord-048487-kamvao4b 183 1 The the DT cord-048487-kamvao4b 183 2 value value NN cord-048487-kamvao4b 183 3 of of IN cord-048487-kamvao4b 183 4 a a DT cord-048487-kamvao4b 183 5 database database NN cord-048487-kamvao4b 183 6 in in IN cord-048487-kamvao4b 183 7 surveillance surveillance NN cord-048487-kamvao4b 183 8 and and CC cord-048487-kamvao4b 183 9 vaccine vaccine NN cord-048487-kamvao4b 183 10 selection selection NN cord-048487-kamvao4b 183 11 Transmissibility transmissibility NN cord-048487-kamvao4b 183 12 of of IN cord-048487-kamvao4b 183 13 1918 1918 CD cord-048487-kamvao4b 183 14 pandemic pandemic NN cord-048487-kamvao4b 183 15 influenza influenza NN cord-048487-kamvao4b 183 16 Estimating estimate VBG cord-048487-kamvao4b 183 17 the the DT cord-048487-kamvao4b 183 18 rate rate NN cord-048487-kamvao4b 183 19 of of IN cord-048487-kamvao4b 183 20 molecular molecular JJ cord-048487-kamvao4b 183 21 evolution evolution NN cord-048487-kamvao4b 183 22 : : : cord-048487-kamvao4b 184 1 Incorporating incorporate VBG cord-048487-kamvao4b 184 2 non non JJ cord-048487-kamvao4b 184 3 - - JJ cord-048487-kamvao4b 184 4 contemporaneous contemporaneous JJ cord-048487-kamvao4b 184 5 sequences sequence NNS cord-048487-kamvao4b 184 6 into into IN cord-048487-kamvao4b 184 7 maximum maximum JJ cord-048487-kamvao4b 184 8 likelihood likelihood NN cord-048487-kamvao4b 184 9 phylogenies phylogeny NNS cord-048487-kamvao4b 184 10 Joint joint JJ cord-048487-kamvao4b 184 11 Bayesian bayesian JJ cord-048487-kamvao4b 184 12 estimation estimation NN cord-048487-kamvao4b 184 13 of of IN cord-048487-kamvao4b 184 14 alignment alignment NN cord-048487-kamvao4b 184 15 and and CC cord-048487-kamvao4b 184 16 phylogeny phylogeny NN cord-048487-kamvao4b 184 17 Tests test NNS cord-048487-kamvao4b 184 18 of of IN cord-048487-kamvao4b 184 19 applicability applicability NN cord-048487-kamvao4b 184 20 of of IN cord-048487-kamvao4b 184 21 several several JJ cord-048487-kamvao4b 184 22 substitution substitution NN cord-048487-kamvao4b 184 23 models model NNS cord-048487-kamvao4b 184 24 for for IN cord-048487-kamvao4b 184 25 DNA dna NN cord-048487-kamvao4b 184 26 sequence sequence NN cord-048487-kamvao4b 184 27 data data NN cord-048487-kamvao4b 184 28 r8s r8s CD cord-048487-kamvao4b 184 29 : : : cord-048487-kamvao4b 185 1 inferring infer VBG cord-048487-kamvao4b 185 2 absolute absolute JJ cord-048487-kamvao4b 185 3 rates rate NNS cord-048487-kamvao4b 185 4 of of IN cord-048487-kamvao4b 185 5 molecular molecular JJ cord-048487-kamvao4b 185 6 evolution evolution NN cord-048487-kamvao4b 185 7 and and CC cord-048487-kamvao4b 185 8 divergence divergence NN cord-048487-kamvao4b 185 9 times time NNS cord-048487-kamvao4b 185 10 in in IN cord-048487-kamvao4b 185 11 the the DT cord-048487-kamvao4b 185 12 absence absence NN cord-048487-kamvao4b 185 13 of of IN cord-048487-kamvao4b 185 14 a a DT cord-048487-kamvao4b 185 15 molecular molecular JJ cord-048487-kamvao4b 185 16 clock clock NN cord-048487-kamvao4b 185 17 A a DT cord-048487-kamvao4b 185 18 viral viral JJ cord-048487-kamvao4b 185 19 sampling sampling NN cord-048487-kamvao4b 185 20 design design NN cord-048487-kamvao4b 185 21 for for IN cord-048487-kamvao4b 185 22 testing test VBG cord-048487-kamvao4b 185 23 the the DT cord-048487-kamvao4b 185 24 molecular molecular JJ cord-048487-kamvao4b 185 25 clock clock NN cord-048487-kamvao4b 185 26 and and CC cord-048487-kamvao4b 185 27 for for IN cord-048487-kamvao4b 185 28 estimating estimate VBG cord-048487-kamvao4b 185 29 evolutionary evolutionary JJ cord-048487-kamvao4b 185 30 rates rate NNS cord-048487-kamvao4b 185 31 and and CC cord-048487-kamvao4b 185 32 divergence divergence NN cord-048487-kamvao4b 185 33 times time NNS cord-048487-kamvao4b 185 34 Origin Origin NNP cord-048487-kamvao4b 185 35 and and CC cord-048487-kamvao4b 185 36 evolution evolution NN cord-048487-kamvao4b 185 37 of of IN cord-048487-kamvao4b 185 38 influenza influenza NN cord-048487-kamvao4b 185 39 hemagglutinin hemagglutinin NN cord-048487-kamvao4b 185 40 genes gene NNS cord-048487-kamvao4b 186 1 The the DT cord-048487-kamvao4b 186 2 Clustal Clustal NNP cord-048487-kamvao4b 186 3 X X NNP cord-048487-kamvao4b 186 4 windows windows NNP cord-048487-kamvao4b 186 5 interface interface NN cord-048487-kamvao4b 186 6 : : : cord-048487-kamvao4b 186 7 flexible flexible JJ cord-048487-kamvao4b 186 8 strategies strategy NNS cord-048487-kamvao4b 186 9 for for IN cord-048487-kamvao4b 186 10 multiple multiple JJ cord-048487-kamvao4b 186 11 sequence sequence NN cord-048487-kamvao4b 186 12 alignment alignment NN cord-048487-kamvao4b 186 13 aided aid VBN cord-048487-kamvao4b 186 14 by by IN cord-048487-kamvao4b 186 15 quality quality NN cord-048487-kamvao4b 186 16 analysis analysis NN cord-048487-kamvao4b 186 17 tools tool NNS cord-048487-kamvao4b 186 18 Inferring infer VBG cord-048487-kamvao4b 186 19 the the DT cord-048487-kamvao4b 186 20 rate rate NN cord-048487-kamvao4b 186 21 and and CC cord-048487-kamvao4b 186 22 timescale timescale NN cord-048487-kamvao4b 186 23 of of IN cord-048487-kamvao4b 186 24 Dengue Dengue NNP cord-048487-kamvao4b 186 25 virus virus NN cord-048487-kamvao4b 186 26 evolution evolution NN cord-048487-kamvao4b 186 27 JDO'B JDO'B NNP cord-048487-kamvao4b 186 28 collected collect VBD cord-048487-kamvao4b 186 29 the the DT cord-048487-kamvao4b 186 30 data datum NNS cord-048487-kamvao4b 186 31 , , , cord-048487-kamvao4b 186 32 designed design VBN cord-048487-kamvao4b 186 33 and and CC cord-048487-kamvao4b 186 34 performed perform VBD cord-048487-kamvao4b 186 35 the the DT cord-048487-kamvao4b 186 36 study study NN cord-048487-kamvao4b 186 37 and and CC cord-048487-kamvao4b 186 38 wrote write VBD cord-048487-kamvao4b 186 39 the the DT cord-048487-kamvao4b 186 40 initial initial JJ cord-048487-kamvao4b 186 41 manuscript manuscript NN cord-048487-kamvao4b 186 42 . . . cord-048487-kamvao4b 187 1 ZSS ZSS NNP cord-048487-kamvao4b 187 2 provided provide VBD cord-048487-kamvao4b 187 3 extensive extensive JJ cord-048487-kamvao4b 187 4 review review NN cord-048487-kamvao4b 187 5 of of IN cord-048487-kamvao4b 187 6 the the DT cord-048487-kamvao4b 187 7 study study NN cord-048487-kamvao4b 187 8 design design NN cord-048487-kamvao4b 187 9 and and CC cord-048487-kamvao4b 187 10 provided provide VBN cord-048487-kamvao4b 187 11 assistance assistance NN cord-048487-kamvao4b 187 12 in in IN cord-048487-kamvao4b 187 13 revising revise VBG cord-048487-kamvao4b 187 14 the the DT cord-048487-kamvao4b 187 15 manuscript manuscript NN cord-048487-kamvao4b 187 16 . . . cord-048487-kamvao4b 188 1 MAS MAS NNP cord-048487-kamvao4b 188 2 contributed contribute VBD cord-048487-kamvao4b 188 3 extensive extensive JJ cord-048487-kamvao4b 188 4 work work NN cord-048487-kamvao4b 188 5 in in IN cord-048487-kamvao4b 188 6 reviewing review VBG cord-048487-kamvao4b 188 7 and and CC cord-048487-kamvao4b 188 8 revising revise VBG cord-048487-kamvao4b 188 9 the the DT cord-048487-kamvao4b 188 10 manuscript manuscript NN cord-048487-kamvao4b 188 11 . . . cord-048487-kamvao4b 189 1 Initially initially RB cord-048487-kamvao4b 189 2 , , , cord-048487-kamvao4b 189 3 one one PRP cord-048487-kamvao4b 189 4 might may MD cord-048487-kamvao4b 189 5 define define VB cord-048487-kamvao4b 189 6 an an DT cord-048487-kamvao4b 189 7 estimator estimator NN cord-048487-kamvao4b 189 8 of of IN cord-048487-kamvao4b 189 9 the the DT cord-048487-kamvao4b 189 10 conditional conditional JJ cord-048487-kamvao4b 189 11 pairwise pairwise JJ cord-048487-kamvao4b 189 12 rate rate NN cord-048487-kamvao4b 189 13 to to TO cord-048487-kamvao4b 189 14 be be VB cord-048487-kamvao4b 189 15 that that DT cord-048487-kamvao4b 189 16 has have VBZ cord-048487-kamvao4b 189 17 been be VBN cord-048487-kamvao4b 189 18 previously previously RB cord-048487-kamvao4b 189 19 used use VBN cord-048487-kamvao4b 189 20 in in IN cord-048487-kamvao4b 189 21 the the DT cord-048487-kamvao4b 189 22 paper paper NN cord-048487-kamvao4b 189 23 outlining outline VBG cord-048487-kamvao4b 189 24 the the DT cord-048487-kamvao4b 189 25 TREBLE treble NN cord-048487-kamvao4b 189 26 algorithm algorithm NN cord-048487-kamvao4b 189 27 [ [ -LRB- cord-048487-kamvao4b 189 28 26 26 CD cord-048487-kamvao4b 189 29 ] ] -RRB- cord-048487-kamvao4b 189 30 , , , cord-048487-kamvao4b 189 31 and and CC cord-048487-kamvao4b 189 32 originates originate VBZ cord-048487-kamvao4b 189 33 in in IN cord-048487-kamvao4b 189 34 [ [ -LRB- cord-048487-kamvao4b 189 35 13 13 CD cord-048487-kamvao4b 189 36 ] ] -RRB- cord-048487-kamvao4b 189 37 . . . cord-048487-kamvao4b 190 1 However however RB cord-048487-kamvao4b 190 2 , , , cord-048487-kamvao4b 190 3 this this DT cord-048487-kamvao4b 190 4 apparently apparently RB cord-048487-kamvao4b 190 5 natural natural JJ cord-048487-kamvao4b 190 6 statistic statistic NN cord-048487-kamvao4b 190 7 is be VBZ cord-048487-kamvao4b 190 8 substantially substantially RB cord-048487-kamvao4b 190 9 biased biased JJ cord-048487-kamvao4b 190 10 when when WRB cord-048487-kamvao4b 190 11 the the DT cord-048487-kamvao4b 190 12 sampling sampling NN cord-048487-kamvao4b 190 13 times time NNS cord-048487-kamvao4b 190 14 of of IN cord-048487-kamvao4b 190 15 sequences sequence NNS cord-048487-kamvao4b 190 16 i i PRP cord-048487-kamvao4b 190 17 and and CC cord-048487-kamvao4b 190 18 j j NNP cord-048487-kamvao4b 190 19 are be VBP cord-048487-kamvao4b 190 20 close close JJ cord-048487-kamvao4b 190 21 . . . cord-048487-kamvao4b 191 1 To to TO cord-048487-kamvao4b 191 2 be be VB cord-048487-kamvao4b 191 3 seen see VBN cord-048487-kamvao4b 191 4 in in IN cord-048487-kamvao4b 191 5 the the DT cord-048487-kamvao4b 191 6 following follow VBG cord-048487-kamvao4b 191 7 derivation derivation NN cord-048487-kamvao4b 191 8 , , , cord-048487-kamvao4b 191 9 this this DT cord-048487-kamvao4b 191 10 bias bias NN cord-048487-kamvao4b 191 11 is be VBZ cord-048487-kamvao4b 191 12 the the DT cord-048487-kamvao4b 191 13 result result NN cord-048487-kamvao4b 191 14 of of IN cord-048487-kamvao4b 191 15 the the DT cord-048487-kamvao4b 191 16 time time NN cord-048487-kamvao4b 191 17 sampling sampling NN cord-048487-kamvao4b 191 18 error error NN cord-048487-kamvao4b 191 19 structure structure NN cord-048487-kamvao4b 191 20 . . . cord-048487-kamvao4b 192 1 As as IN cord-048487-kamvao4b 192 2 the the DT cord-048487-kamvao4b 192 3 true true JJ cord-048487-kamvao4b 192 4 value value NN cord-048487-kamvao4b 192 5 of of IN cord-048487-kamvao4b 192 6 the the DT cord-048487-kamvao4b 192 7 rate rate NN cord-048487-kamvao4b 192 8 of of IN cord-048487-kamvao4b 192 9 substitution substitution NN cord-048487-kamvao4b 192 10 is be VBZ cord-048487-kamvao4b 192 11 given give VBN cord-048487-kamvao4b 192 12 by by IN cord-048487-kamvao4b 192 13 we -PRON- PRP cord-048487-kamvao4b 192 14 then then RB cord-048487-kamvao4b 192 15 have have VBP cord-048487-kamvao4b 192 16 an an DT cord-048487-kamvao4b 192 17 expression expression NN cord-048487-kamvao4b 192 18 for for IN cord-048487-kamvao4b 192 19 the the DT cord-048487-kamvao4b 192 20 error error NN cord-048487-kamvao4b 192 21 : : : cord-048487-kamvao4b 192 22 Taking take VBG cord-048487-kamvao4b 192 23 the the DT cord-048487-kamvao4b 192 24 expectation expectation NN cord-048487-kamvao4b 192 25 yields yield VBZ cord-048487-kamvao4b 192 26 the the DT cord-048487-kamvao4b 192 27 bias bias NN cord-048487-kamvao4b 192 28 : : : cord-048487-kamvao4b 192 29 Publish publish VB cord-048487-kamvao4b 192 30 with with IN cord-048487-kamvao4b 192 31 Bio Bio NNP cord-048487-kamvao4b 192 32 Med Med NNP cord-048487-kamvao4b 192 33 Central Central NNP cord-048487-kamvao4b 192 34 and and CC cord-048487-kamvao4b 192 35 every every DT cord-048487-kamvao4b 192 36 scientist scientist NN cord-048487-kamvao4b 192 37 can can MD cord-048487-kamvao4b 192 38 read read VB cord-048487-kamvao4b 192 39 your -PRON- PRP$ cord-048487-kamvao4b 192 40 work work NN cord-048487-kamvao4b 192 41 free free JJ cord-048487-kamvao4b 192 42 of of IN cord-048487-kamvao4b 192 43 charge charge NN cord-048487-kamvao4b 192 44 ) ) -RRB- cord-048487-kamvao4b 192 45 . . . cord-048487-kamvao4b 192 46 _SP