Title Analysis of genetic diversity and sites of recombination in human rhinovirus species C.
Author McIntyre, Chloe L; McWilliam Leitch, E Carol; Savolainen-Kopra, Carita; Hovi, Tapani; Simmonds, Peter
Journal J Virol Publication Year/Month 2010-Oct
PMID 20668080 PMCID PMC2937769
Affiliation 1.Centre for Infectious Diseases, University of Edinburgh, Summerhall, Edinburgh EH9 1QH, UK.

Human rhinoviruses (HRVs) are a highly prevalent and diverse group of respiratory viruses. Although HRV-A and HRV-B are traditionally detected by virus isolation, a series of unculturable HRV variants have recently been described and assigned as a new species (HRV-C) within the picornavirus Enterovirus genus. To investigate their genetic diversity and occurrence of recombination, we have performed comprehensive phylogenetic analysis of sequences from the 5\' untranslated region (5\' UTR), VP4/VP2, VP1, and 3Dpol regions amplified from 89 HRV-C-positive respiratory samples and available published sequences. Branching orders of VP4/VP2, VP1, and 3Dpol trees were identical, consistent with the absence of intraspecies recombination in the coding regions. However, numerous tree topology changes were apparent in the 5\' UTR, where >60% of analyzed HRV-C variants showed recombination with species A sequences. Two recombination hot spots in stem-loop 5 and the polypyrimidine tract in the 5\' UTR were mapped using the program GroupingScan. Available HRV-C sequences showed evidence for additional interspecies recombination with HRV-A in the 2A gene, with breakpoints mapping precisely to the boundaries of the C-terminal domain of the encoded proteinase. Pairwise distances between HRV-C variants in VP1 and VP4/VP2 regions fell into two separate distributions, resembling inter- and intraserotype distances of species A and B. These observations suggest that, without serological cross-neutralization data, HRV-C genetic groups may be equivalently classified into types using divergence thresholds derived from distance distributions. The extensive sequence data from multiple genome regions of HRV-C and analyses of recombination in the current study will assist future formulation of consensus criteria for HRV-C type assignment and identification.

StrainID RV Species Serotype Length(nt) Country Year Strain Name
CNC05224 C None 431 United Kingdom 2007 Resp_5867/07
CNC05229 C None 437 United Kingdom 2007 Resp_3832/07
CNC05234 C None 436 United Kingdom 2006 Resp_2964/06
CNC05239 C None 638 United Kingdom 2007 Resp_5404/07
CNC05244 C None 435 United Kingdom 2007 Resp_4097/07
CNC05249 C None 432 United Kingdom 2007 Resp_3727/07
CNC05254 C None 433 United Kingdom 2006 Resp_2744/06
CNC05259 C None 437 United Kingdom 2007 Resp_6145/07
CNC05264 C None 435 United Kingdom 2007 Resp_5571/07
CNC05269 C None 430 United Kingdom 2007 Resp_5279/07
CNC05274 C None 431 United Kingdom 2007 Resp_4857/07
CNC05279 C None 435 United Kingdom 2007 Resp_4122/07
CNC05284 C None 438 United Kingdom 2007 Resp_3972/07
CNC05289 C None 431 United Kingdom 2007 Resp_3607/07
CNC05294 C None 424 United Kingdom 2006 Resp_3266/06
CNC05299 C None 626 United Kingdom 2006 Resp_2789/06
CNC05304 C None 434 United Kingdom 2007 Resp_5570/07
CNC05309 C None 471 United Kingdom 2006 Resp_2921/06
CNC05314 C None 471 United Kingdom 2007 Resp_3451/07
CNC05319 C None 471 United Kingdom 2007 Resp_6142/07
CNC05324 C None 471 United Kingdom 2006 Resp_3053/06
CNC05329 C None 471 United Kingdom 2007 Resp_4101/07
CNC05334 C None 471 United Kingdom 2006 Resp_3028/06
CNC05339 C None 471 United Kingdom 2007 Resp_5279/07
CNC05344 C None 471 United Kingdom 2007 Resp_3787/07
CNC05349 C None 471 United Kingdom 2007 Resp_3837/07
CNC05354 C None 471 United Kingdom 2007 Resp_6131/07
CNC05359 C None 471 United Kingdom 2007 Resp_4742/07
CNC05364 C None 471 United Kingdom 2007 Resp_4122/07
CNC05369 C None 471 United Kingdom 2007 Resp_4098/07
CNC05374 C None 471 United Kingdom 2007 Resp_5568/07
CNC05379 C None 471 United Kingdom 2007 Resp_6082/07
CNC05384 C None 471 United Kingdom 2006 Resp_2744/06
CNC05389 C None 471 United Kingdom 2006 Resp_3105/06
CNC05394 C None 471 United Kingdom 2007 Resp_5096/07
CNC05399 C None 393 United Kingdom 2007 Resp7235/07
CNC05404 C None 391 United Kingdom 2007 Resp6424/07
CNC05409 C None 393 United Kingdom 2008 Resp10888/08
CNC05454 C 31 797 United Kingdom 2007 Resp_4923/07
CNC05459 C 12 831 United Kingdom 2007 Resp_3993/07
CNC05464 C 28 831 United Kingdom 2006 Resp_3105/06
CNC05469 C 29 822 United Kingdom 2007 Resp_5345/07
CNC05474 C 16 822 United Kingdom 2007 Resp_5910/07
CNC05479 C 24 822 United Kingdom 2007 Resp_7147/07
CNC05484 C 33 834 United Kingdom 2007 Resp_4917/07
CNC05489 C 20 822 United Kingdom 2007 Resp_4620/07
CNC05494 C 32 822 United Kingdom 2007 Resp_4037/07
CNC05499 C 1 816 United Kingdom 2007 Resp_3753/07
CNC05504 C 27 818 United Kingdom 2007 Resp_3470/07
CNC05509 C 6 831 United Kingdom 2006 Resp_2957/06
CNC05514 C 26 822 United Kingdom 2006 Resp_2514/06
CNC05519 C 6 825 United Kingdom 2006 Resp_2992/06
CNM05225 C None 435 United Kingdom 2007 Resp_5477/07
CNM05230 C None 436 United Kingdom 2007 Resp_3750/07
CNM05235 C None 438 United Kingdom 2006 Resp_2921/06
CNM05240 C None 625 United Kingdom 2007 Resp_5093/07
CNM05245 C None 425 United Kingdom 2007 Resp_4051/07
CNM05250 C None 436 United Kingdom 2007 Resp_3628/07
CNM05255 C None 437 United Kingdom 2008 Resp_10300/08
CNM05260 C None 432 United Kingdom 2007 Resp_6082/07
CNM05265 C None 437 United Kingdom 2007 Resp_5568/07
CNM05270 C None 436 United Kingdom 2007 Resp_5145/07
CNM05275 C None 629 United Kingdom 2007 Resp_4742/07
CNM05280 C None 424 United Kingdom 2007 Resp_4101/07
CNM05285 C None 435 United Kingdom 2007 Resp_3837/07
CNM05290 C None 430 United Kingdom 2007 Resp_3487/07
CNM05295 C None 425 United Kingdom 2006 Resp_3090/06
CNM05300 C None 424 United Kingdom 2006 Resp_2784/06
CNM05305 C None 432 United Kingdom 2006 Resp_3053/06
CNM05310 C None 471 United Kingdom 2006 Resp_2964/06
CNM05315 C None 471 United Kingdom 2007 Resp_5153/07
CNM05320 C None 471 United Kingdom 2006 Resp_3200/06
CNM05325 C None 471 United Kingdom 2006 Resp_2784/06
CNM05330 C None 471 United Kingdom 2007 Resp_4620/07
CNM05335 C None 471 United Kingdom 2006 Resp_3086/06
CNM05340 C None 471 United Kingdom 2007 Resp_3607/07
CNM05345 C None 471 United Kingdom 2007 Resp_4733/07
CNM05350 C None 471 United Kingdom 2007 Resp_4037/07
CNM05355 C None 471 United Kingdom 2007 Resp_5130/07
CNM05360 C None 471 United Kingdom 2007 Resp_5404/07
CNM05365 C None 471 United Kingdom 2007 Resp_4644/07
CNM05370 C None 471 United Kingdom 2007 Resp_5145/07
CNM05375 C None 466 United Kingdom 2007 Resp_5571/07
CNM05380 C None 471 United Kingdom 2007 Resp_6145/07
CNM05385 C None 471 United Kingdom 2007 Resp_4314/07
CNM05390 C None 471 United Kingdom 2007 Resp_3727/07
CNM05395 C None 471 United Kingdom 2007 Resp_3628/07
CNM05400 C None 394 United Kingdom 2007 Resp7147/07
CNM05405 C None 386 United Kingdom 2007 Resp5998/07
CNM05410 C None 393 United Kingdom 2008 Resp10801/08
CNM05450 C 30 816 United Kingdom 2007 Resp_3898/07
CNM05455 C 15 840 United Kingdom 2007 Resp_4644/07
CNM05460 C 12 834 United Kingdom 2007 Resp_3922/07
CNM05465 C 6 831 United Kingdom 2006 Resp_3089/06
CNM05470 C 18 843 United Kingdom 2007 Resp_3733/07
CNM05475 C 32 822 United Kingdom 2007 Resp_5607/07
CNM05480 C 32 816 United Kingdom 2007 Resp_5554/07
CNM05485 C 6 831 United Kingdom 2007 Resp_4891/07
CNM05490 C 27 813 United Kingdom 2007 Resp_4293/07
CNM05495 C 20 828 United Kingdom 2007 Resp_3995/07
CNM05500 C 18 840 United Kingdom 2007 Resp_3631/07
CNM05505 C 6 825 United Kingdom 2007 Resp_3464/07
CNM05510 C 13 819 United Kingdom 2006 Resp_2951/06
CNM05515 C 21 822 United Kingdom 2007 Resp_5071/07
CNM05520 C 8 825 United Kingdom 2008 Resp_10300/08
CNP05222 C None 435 United Kingdom 2007 Resp_6142/07
CNP05227 C None 430 United Kingdom 2007 Resp_4817/07
CNP05232 C None 434 United Kingdom 2006 Resp_3200/06
CNP05237 C None 435 United Kingdom 2006 Resp_2480/06
CNP05242 C None 606 United Kingdom 2007 Resp_4644/07
CNP05247 C None 434 United Kingdom 2007 Resp_3922/07
CNP05252 C None 429 United Kingdom 2006 Resp_3089/06
CNP05257 C None 424 United Kingdom 2007 Resp_5345/07
CNP05262 C None 437 United Kingdom 2007 Resp_5910/07
CNP05267 C None 430 United Kingdom 2007 Resp_7147/07
CNP05272 C None 424 United Kingdom 2007 Resp_4917/07
CNP05277 C None 426 United Kingdom 2007 Resp_4620/07
CNP05282 C None 438 United Kingdom 2007 Resp_4037/07
CNP05287 C None 429 United Kingdom 2007 Resp_3753/07
CNP05292 C None 423 United Kingdom 2007 Resp_3470/07
CNP05297 C None 430 United Kingdom 2006 Resp_2957/06
CNP05302 C None 436 United Kingdom 2006 Resp_2514/06
CNP05307 C None 430 United Kingdom 2006 Resp_2992/06
CNP05312 C None 471 United Kingdom 2007 Resp_3469/07
CNP05317 C None 470 United Kingdom 2006 Resp_2503/06
CNP05322 C None 471 United Kingdom 2007 Resp_3832/07
CNP05327 C None 471 United Kingdom 2007 Resp_4293/07
CNP05332 C None 471 United Kingdom 2006 Resp_2957/06
CNP05337 C None 471 United Kingdom 2007 Resp_3464/07
CNP05342 C None 471 United Kingdom 2007 Resp_3487/07
CNP05347 C None 471 United Kingdom 2006 Resp_2951/06
CNP05352 C None 471 United Kingdom 2007 Resp_5554/07
CNP05357 C None 471 United Kingdom 2007 Resp_4917/07
CNP05362 C None 471 United Kingdom 2007 Resp_5093/07
CNP05367 C None 471 United Kingdom 2007 Resp_3631/07
CNP05372 C None 471 United Kingdom 2007 Resp_7147/07
CNP05377 C None 471 United Kingdom 2007 Resp_5910/07
CNP05382 C None 471 United Kingdom 2007 Resp_5345/07
CNP05387 C None 471 United Kingdom 2006 Resp_2832/06
CNP05392 C None 471 United Kingdom 2007 Resp_3922/07
CNP05397 C None 471 United Kingdom 2007 Resp_3898/07
CNP05402 C None 391 United Kingdom 2007 Resp6808/07
CNP05407 C None 394 United Kingdom 2007 Resp5910/07
CNP05412 C None 394 United Kingdom 2008 Resp10221/08
CNP05452 C 20 828 United Kingdom 2007 Resp_5093/07
CNP05457 C 9 816 United Kingdom 2007 Resp_4097/07
CNP05462 C 18 843 United Kingdom 2007 Resp_3727/07
CNP05467 C 22 822 United Kingdom 2006 Resp_2744/06
CNP05472 C 23 825 United Kingdom 2007 Resp_6082/07
CNP05477 C 32 822 United Kingdom 2007 Resp_5568/07
CNP05482 C 17 822 United Kingdom 2007 Resp_5145/07
CNP05487 C 19 825 United Kingdom 2007 Resp_4742/07
CNP05492 C 20 822 United Kingdom 2007 Resp_4101/07
CNP05497 C 32 822 United Kingdom 2007 Resp_3837/07
CNP05502 C 1 822 United Kingdom 2007 Resp_3487/07
CNP05507 C 14 822 United Kingdom 2006 Resp_3090/06
CNP05512 C 27 813 United Kingdom 2006 Resp_2784/06
CNP05517 C 23 822 United Kingdom 2006 Resp_3053/06
CNS05223 C None 437 United Kingdom 2007 Resp_5096/07
CNS05228 C None 439 United Kingdom 2007 Resp_4314/07
CNS05233 C None 619 United Kingdom 2006 Resp_3137/06
CNS05238 C None 624 United Kingdom 2007 Resp_5789/07
CNS05243 C None 618 United Kingdom 2007 Resp_4252/07
CNS05248 C None 426 United Kingdom 2007 Resp_3898/07
CNS05253 C None 609 United Kingdom 2006 Resp_2832/06
CNS05258 C None 438 United Kingdom 2007 Resp_3733/07
CNS05263 C None 436 United Kingdom 2007 Resp_5607/07
CNS05268 C None 620 United Kingdom 2007 Resp_5554/07
CNS05273 C None 431 United Kingdom 2007 Resp_4891/07
CNS05278 C None 421 United Kingdom 2007 Resp_4293/07
CNS05283 C None 427 United Kingdom 2007 Resp_3995/07
CNS05288 C None 438 United Kingdom 2007 Resp_3631/07
CNS05293 C None 431 United Kingdom 2007 Resp_3464/07
CNS05298 C None 434 United Kingdom 2006 Resp_2951/06
CNS05303 C None 428 United Kingdom 2007 Resp_5071/07
CNS05308 C None 624 United Kingdom 2007 Resp_3469/07
CNS05313 C None 471 United Kingdom 2006 Resp_3137/06
CNS05318 C None 471 United Kingdom 2007 Resp_3750/07
CNS05323 C None 471 United Kingdom 2008 Resp_10300/08
CNS05328 C None 471 United Kingdom 2007 Resp_3470/07
CNS05333 C None 471 United Kingdom 2006 Resp_2992/06
CNS05338 C None 471 United Kingdom 2007 Resp_4857/07
CNS05343 C None 471 United Kingdom 2007 Resp_3753/07
CNS05348 C None 471 United Kingdom 2007 Resp_4051/07
CNS05353 C None 471 United Kingdom 2007 Resp_5570/07
CNS05358 C None 471 United Kingdom 2006 Resp_3266/06
CNS05363 C None 471 United Kingdom 2007 Resp_4097/07
CNS05368 C None 471 United Kingdom 2007 Resp_5789/07
CNS05373 C None 471 United Kingdom 2007 Resp_5567/07
CNS05378 C None 471 United Kingdom 2007 Resp_5998/07
CNS05383 C None 471 United Kingdom 2007 Resp_3499/07
CNS05388 C None 471 United Kingdom 2006 Resp_3089/06
CNS05393 C None 471 United Kingdom 2007 Resp_3993/07
CNS05398 C None 391 United Kingdom 2008 Resp9449/08
CNS05403 C None 393 United Kingdom 2007 Resp6691/07
CNS05408 C None 391 United Kingdom 2007 Resp5562/07
CNS05413 C None 391 United Kingdom 2008 Resp10061/08
CNS05453 C 33 828 United Kingdom 2007 Resp_5789/07
CNS05458 C 20 822 United Kingdom 2007 Resp_4051/07
CNS05463 C 18 840 United Kingdom 2007 Resp_3628/07
CNS05468 C 18 843 United Kingdom 2007 Resp_3499/07
CNS05473 C 16 822 United Kingdom 2007 Resp_5998/07
CNS05478 C 32 816 United Kingdom 2007 Resp_5567/07
CNS05483 C 9 816 United Kingdom 2007 Resp_5130/07
CNS05488 C 21 816 United Kingdom 2007 Resp_4733/07
CNS05493 C 20 822 United Kingdom 2007 Resp_4098/07
CNS05498 C 7 816 United Kingdom 2007 Resp_3787/07
CNS05503 C 1 816 United Kingdom 2007 Resp_3480/07
CNS05508 C 6 831 United Kingdom 2006 Resp_3028/06
CNS05513 C 22 822 United Kingdom 2006 Resp_2748/06
CNS05518 C 6 825 United Kingdom 2006 Resp_3086/06
CNZ05226 C None 437 United Kingdom 2007 Resp_5153/07
CNZ05231 C None 436 United Kingdom 2007 Resp_3451/07
CNZ05236 C None 436 United Kingdom 2006 Resp_2503/06
CNZ05241 C None 436 United Kingdom 2007 Resp_4923/07
CNZ05246 C None 433 United Kingdom 2007 Resp_3993/07
CNZ05251 C None 437 United Kingdom 2006 Resp_3105/06
CNZ05256 C None 438 United Kingdom 2007 Resp_3499/07
CNZ05261 C None 440 United Kingdom 2007 Resp_5998/07
CNZ05266 C None 436 United Kingdom 2007 Resp_5567/07
CNZ05271 C None 639 United Kingdom 2007 Resp_5130/07
CNZ05276 C None 429 United Kingdom 2007 Resp_4733/07
CNZ05281 C None 426 United Kingdom 2007 Resp_4098/07
CNZ05286 C None 429 United Kingdom 2007 Resp_3787/07
CNZ05291 C None 431 United Kingdom 2007 Resp_3480/07
CNZ05296 C None 431 United Kingdom 2006 Resp_3028/06
CNZ05301 C None 433 United Kingdom 2006 Resp_2748/06
CNZ05306 C None 431 United Kingdom 2006 Resp_3086/06
CNZ05311 C None 471 United Kingdom 2007 Resp_5477/07
CNZ05316 C None 471 United Kingdom 2007 Resp_5867/07
CNZ05321 C None 471 United Kingdom 2006 Resp_2480/06
CNZ05326 C None 471 United Kingdom 2006 Resp_2789/06
CNZ05331 C None 471 United Kingdom 2006 Resp_2748/06
CNZ05336 C None 471 United Kingdom 2007 Resp_4891/07
CNZ05341 C None 471 United Kingdom 2007 Resp_3480/07
CNZ05346 C None 471 United Kingdom 2007 Resp_5071/07
CNZ05351 C None 471 United Kingdom 2007 Resp_3972/07
CNZ05356 C None 471 United Kingdom 2006 Resp_3090/06
CNZ05361 C None 471 United Kingdom 2006 Resp_2514/06
CNZ05366 C None 471 United Kingdom 2007 Resp_4252/07
CNZ05371 C None 471 United Kingdom 2007 Resp_4923/07
CNZ05376 C None 471 United Kingdom 2007 Resp_5607/07
CNZ05381 C None 471 United Kingdom 2007 Resp_3733/07
CNZ05386 C None 471 United Kingdom 2007 Resp_4817/07
CNZ05391 C None 471 United Kingdom 2007 Resp_3856/07
CNZ05396 C None 471 United Kingdom 2007 Resp_3995/07
CNZ05401 C None 394 United Kingdom 2007 Resp6830/07
CNZ05406 C None 394 United Kingdom 2007 Resp5995/07
CNZ05411 C None 391 United Kingdom 2008 Resp10300/08
CNZ05451 C 9 816 United Kingdom 2007 Resp_5404/07
CNZ05456 C 12 834 United Kingdom 2007 Resp_4252/07
CNZ05461 C 26 822 United Kingdom 2007 Resp_3856/07
CNZ05466 C 25 809 United Kingdom 2006 Resp_2832/06
CNZ05471 C 16 798 United Kingdom 2007 Resp_6145/07
CNZ05476 C 32 816 United Kingdom 2007 Resp_5571/07
CNZ05481 C 6 825 United Kingdom 2007 Resp_5279/07
CNZ05486 C 6 831 United Kingdom 2007 Resp_4857/07
CNZ05491 C 9 816 United Kingdom 2007 Resp_4122/07
CNZ05496 C 32 813 United Kingdom 2007 Resp_3972/07
CNZ05501 C 3 831 United Kingdom 2007 Resp_3607/07
CNZ05506 C 19 825 United Kingdom 2006 Resp_3266/06
CNZ05511 C 27 818 United Kingdom 2006 Resp_2789/06
CNZ05516 C 32 822 United Kingdom 2007 Resp_5570/07
CNZ05521 C 32 816 United Kingdom 2007 Resp_6131/07
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