Genomic insights into the Ixodes scapularis tick vector of Lyme disease

Nature Communications

Volumn 7, Issue , 2016, Pages

  Gulia Nuss, Monika ^a,as   Nuss, Andrew B ^a,as   Meyer, Jason M ^a,at   Sonenshine, Daniel E ^b   Roe, R Michael ^c   Waterhouse, Robert M ^d,e,f,g   Sattelle, David B ^h   De La Fuente, José ^i,j   Ribeiro, Jose M ^k   Megy, Karine ^l,af   Thimmapuram, Jyothi ^a   Miller, Jason R ^m   Walenz, Brian P ^m,au   Koren, Sergey ^m,au   Hostetler, Jessica B ^k,m   Thiagarajan, Mathangi ^m,av   Joardar, Vinita S ^m,aw   Hannick, Linda I ^m,av   Bidwell, Shelby ^m,aw   Hammond, Martin P ^l   Young, Sarah ^g   Zeng, Qiandong ^g   Abrudan, Jenica L ^n,ax   Almeida, Francisca C ^o   Ayllón, Nieves ⁱ   Bhide, Ketaki ^a   Bissinger, Brooke W ^c,ay   Bonzon Kulichenko, Elena ^p   Buckingham, Steven D ^h   Caffrey, Daniel R ^q   Caimano, Melissa J ^r   Croset, Vincent ^s,az   Driscoll, Timothy ^t,ba   Gilbert, Don ^u   Gillespie, Joseph J ^t,ar   Giraldo Calderón, Gloria I ^a,n   Grabowski, Jeffrey M ^a,v,bb   Jiang, David ^w   Khalil, Sayed M S ^x   Kim, Donghun ^y,z   Kocan, Katherine M ^j   Koči, Juraj ^z,ar   Kuhn, Richard J ^v   Kurtti, Timothy J ^aa   Lees, Kristin ^ab,bc   Lang, Emma G ^a   Kennedy, Ryan C ^ac   Kwon, Hyeogsun ^y,bd   Perera, Rushika ^v,be   Qi, Yumin ^w   Radolf, Justin D ^r   Sakamoto, Joyce M ^ad   Sánchez Gracia, Alejandro ^o   Severo, Maiara S ^ae,bf   Silverman, Neal ^q   Šimo, Ladislav ^z,bg   Tojo, Marta ^af,ag   Tornador, Cristian ^ah   Van Zee, Janice P ^a   Vázquez, Jesús ^p   Vieira, Filipe G ^o   Villar, Margarita ⁱ   Wespiser, Adam R ^q   Yang, Yunlong ^y   Zhu, Jiwei ^c   Arensburger, Peter ^ai   Pietrantonio, Patricia V ^y   Barker, Stephen C ^aj   Shao, Renfu ^ak   Zdobnov, Evgeny M ^d,e   Hauser, Frank ^al   Grimmelikhuijzen, Cornelis J P ^al   Park, Yoonseong ^z   Rozas, Julio ^o   Benton, Richard ^s   Pedra, Joao H F ^ae,ar   Nelson, David R ^am   Unger, Maria F ⁿ   Tubio, Jose M C ^an,ao   Tu, Zhijian ^w   Robertson, Hugh M ^ap   Shumway, Martin ^m,bf   Sutton, Granger ^m   Wortman, Jennifer R ^m,bh   Lawson, Daniel ^l   Wikel, Stephen K ^aq   Nene, Vishvanath M ^m,bi   Fraser, Claire M ^ar   Collins, Frank H ⁿ   Birren, Bruce ^g   Nelson, Karen E ^m   Caler, Elisabet ^m,bj   Hill, Catherine A ^a   more..

^a PURDUE UNIVERSITY   (United States)

^b OLD DOMINION UNIVERSITY   (United States)

^c NORTH CAROLINA STATE UNIVERSITY   (United States)

^d UNIVERSITY OF GENEVA MEDICAL SCHOOL   (Switzerland)

^e SWISS INSTITUTE OF BIOINFORMATICS   (Switzerland)

^f MASSACHUSETTS INSTITUTE OF TECHNOLOGY   (United States)

^g BROAD INSTITUTE OF MIT AND HARVARD   (United States)

^h UNIVERSITY COLLEGE LONDON   (United Kingdom)

ⁱ INSTITUTO DE INVESTIGACIÓN EN RECURSOS CINEGÉTICOS IREC CSIC UCLM JCCM   (Spain)

^j OKLAHOMA STATE UNIVERSITY   (United States)

^k NATIONAL INSTITUTE OF ALLERGY AND INFECTIOUS DISEASES   (United States)

^l WELLCOME TRUST SANGER INSTITUTE   (United Kingdom)

^m J CRAIG VENTER INSTITUTE   (United States)

ⁿ UNIVERSITY OF NOTRE DAME   (United States)

^o UNIVERSITY OF BARCELONA   (Spain)

^p CENTRO NACIONAL DE INVESTIGACIONES CARDIOVASCULARES CNIC   (Spain)

^q UNIVERSITY OF MASSACHUSETTS MEDICAL SCHOOL   (United States)

^r UNIVERSITY OF CONNECTICUT HEALTH CENTER   (United States)

^s UNIVERSITY OF LAUSANNE   (Switzerland)

^t VIRGINIA BIOINFORMATICS INSTITUTE   (United States)

^u INDIANA UNIVERSITY   (United States)

^v PURDUE UNIVERSITY   (United States)

^w VIRGINIA POLYTECHNIC INSTITUTE AND STATE UNIVERSITY   (United States)

^x Agricultural Genetic Engineering Research Institute   (Egypt)

^y TEXAS A AND M UNIVERSITY   (United States)

^z KANSAS STATE UNIVERSITY   (United States)

^aa UNIVERSITY OF MINNESOTA   (United States)

^ab UNIVERSITY OF MANCHESTER   (United Kingdom)

^ac UNIVERSITY OF CALIFORNIA   (United States)

^ad PENNSYLVANIA STATE UNIVERSITY   (United States)

^ae UNIVERSITY OF CALIFORNIA   (United States)

^af UNIVERSITY OF CAMBRIDGE   (United Kingdom)

^ag UNIVERSITY OF SANTIAGO DE COMPOSTELA   (Spain)

^ah UNIVERSITAT POMPEU FABRA   (Spain)

^ai CALIFORNIA STATE POLYTECHNIC UNIVERSITY   (United States)

^aj UNIVERSITY OF QUEENSLAND   (Australia)

^ak UNIVERSITY OF THE SUNSHINE COAST   (Australia)

^al UNIVERSITY OF COPENHAGEN   (Denmark)

^am UNIVERSITY OF TENNESSEE HEALTH SCIENCE CENTER   (United States)

^an WELLCOME TRUST SANGER INSTITUTE   (United Kingdom)

^ao UNIVERSITY OF VIGO   (Spain)

^ap UNIVERSITY OF ILLINOIS AT URBANA CHAMPAIGN   (United States)

^aq Quinnipiac University   (United States)

^ar UNIVERSITY OF MARYLAND SCHOOL OF MEDICINE   (United States)

^as UNIVERSITY OF NEVADA   (United States)

^at MONSANTO COMPANY   (United States)

^au NATIONAL HUMAN GENOME RESEARCH INSTITUTE   (United States)

^av FREDERICK NATIONAL LABORATORY FOR CANCER RESEARCH   (United States)

^aw NATIONAL INSTITUTES OF HEALTH   (United States)

^ax UNIVERSITY OF NEVADA   (United States)

^ay AGBIOME   (United States)

^az UNIVERSITY OF OXFORD   (United Kingdom)

^ba WEST VIRGINIA UNIVERSITY   (United States)

^bb NATIONAL INSTITUTES OF HEALTH   (United States)

^bc Dept Medicine Endocrinol Diabet   (Singapore)

^bd IOWA STATE UNIVERSITY   (United States)

^be Department of Environmental and Radiological Health Sciences   (United States)

^bf MAX PLANCK INSTITUTE FOR INFECTION BIOLOGY   (Germany)

^bg ECOLE NATIONALE VÉTÉRINAIRE D ALFORT   (France)

^bh Seres Therapeutics   (United States)

^bi INTERNATIONAL LIVESTOCK RESEARCH INSTITUTE   (Kenya)

^bj NATIONAL HEART LUNG AND BLOOD INSTITUTE   (United States)

more..

Author keywords

[No Author keywords available]

Indexed keywords

REPETITIVE DNA; LIGAND GATED ION CHANNEL;

ARTHROPOD; BLOOD; CELLS AND CELL COMPONENTS; DISEASE INCIDENCE; DISEASE TRANSMISSION; DISEASE VECTOR; GENE EXPRESSION; GENOME; HOST-PARASITE INTERACTION; LIFE HISTORY TRAIT; LYME DISEASE; METAZOAN; PATHOGEN; POPULATION STRUCTURE; PROTEIN; TICK; VIRUS;

ARTICLE; CONTROLLED STUDY; DETOXIFICATION; ENCEPHALITIS; FEMALE; GENOME; HUMAN GRANULOCYTIC ANAPLASMOSIS; IXODES SCAPULARIS; LANGAT VIRUS; LIFE HISTORY TRAIT; LYME DISEASE; MULTIGENE FAMILY; NONHUMAN; PIROPLASMOSIS; POPULATION STRUCTURE; RETROPOSON; VITELLOGENESIS; ANAPLASMA PHAGOCYTOPHILUM; ANIMAL; ARACHNID VECTOR; GENE EXPRESSION PROFILING; GENETICS; GENOMICS; IXODES; OOCYTE; TRANSMISSION; XENOPUS LAEVIS;

ANIMALIA; ARTHROPODA; IXODES SCAPULARIS; IXODIDA; LANGAT VIRUS; METAZOA; PANCRUSTACEA;

ANAPLASMA PHAGOCYTOPHILUM; ANIMALS; ARACHNID VECTORS; GENE EXPRESSION PROFILING; GENOME; GENOMICS; IXODES; LIGAND-GATED ION CHANNELS; LYME DISEASE; OOCYTES; XENOPUS LAEVIS;

EID: 84957895144     PISSN: None     EISSN: 20411723     Source Type: Journal    
DOI: 10.1038/ncomms10507     Document Type: Article

References (69)

1. Lindgren, E. & Jaenson, T. G. T. Lyme borreliosis in Europe: influences of climate and climate change, epidemiology, ecology and adaptation measures(World Health Organization, 2006). http://www.euro.who.int/--data/assets/pdf-file/0006/96819/E89522.pdf.
2. MMWR. Morbidity and Mortality Weekly Report. Report No. 61, 1-124 (Centers for Disease Control and Prevention (CDC), Atlanta, GA, USA, 2014) http://www.cdc.gov/mmwr/preview/mmwrhtml/mm6153a1.htm.
3. Walker, D. Tick-transmitted infectious diseases in the United States. Annu. Rev. Public Health 19, 237-269 (1998).
4. Rizzoli, A. et al. Lyme borreliosis in Europe. Eurosurveillance 16: pii 19906 (2011).
5. Wormser, G. P. et al. The clinical assessment, treatment, and prevention of Lyme disease, human granulocytic anaplasmosis, and babesiosis: clinical practice guidelines by the Infectious Diseases Society of America. Clin. Infect. Dis. 43, 1089-1134 (2006).
6. Rehm, P., Meusemann, K., Borner, J., Misof, B. & Burmester, T. Phylogenetic position of Myriapoda revealed by 454 transcriptome sequencing. Mol. Phylogenet. Evol. 77, 25-33 (2014) Erratum 80, 340 (2014).
7. Kaufman, W. R. in Integument and ecdysis. Ch. 5 Volume I (eds Sonenshine, D. E. & Roe, R. M.) pp 416-448 (Oxford University Press, 2014).
8. Geraci, N. S., Johnston, J. S., Robinson, J. P., Wikel, S. K. & Hill, C. A. Variation in genome size of argasid and ixodid ticks. Insect Biochem. Mol. Biol. 37, 399-408 (2007).
9. Grbić, M. et al. The genome of Tetranychus urticae reveals herbivorous pest adaptations. Nature 479, 487-492 (2011).
10. Ullmann, A. J., Lima, C. M., Guerrero, F. D., Piesman, J. & Black, IV W. C. Genome size and organization in the blacklegged tick, Ixodes scapularis and the Southern cattle tick, Boophilus microplus. Insect Mol. Biol. 14, 217-222 (2005).
11. Oliver, Jr J. H. Cytogenetics of mites and ticks. Annu. Rev. Entomol. 22, 407-429 (1977).
12. Meyer, J. M., Kurtti, T. J., Van Zee, J. P. & Hill, C. A. Genome organization of major tandem repeats in the hard tick, Ixodes scapularis. Chrom. Res. 18, 357-370 (2010).
13. Ugarković, D., Podnar, M. & Plohl, M. Satellite DNA of the red flour beetle Tribolium castaneum-comparative study of satellites from the genus Tribolium. Mol. Biol. Evol. 13, 1059-1066 (1996).
14. Tubio, J. M., Naveira, H. & Costas, J. Structural and evolutionary analyses of the ty3/gypsy group of LTR retrotransposons in the genome of Anopheles gambiae. Mol. Biol. Evol. 22, 29-39 (2005).
15. Li, W.-H., Gu, Z., Wang, H. & Nekrutenko, A. Evolutionary analyses of the human genome. Nature 409, 847-849 (2001).
16. Lynch, M. & Conery, J. The evolutionary fate and consequences of duplicate genes. Science 290, 1151-1155 (2000).
17. Van Zee, J. P. et al. Paralog analyses reveal gene duplication events and genes under positive selection in Ixodes scapularis and other ixodid ticks BMC Genomics. doi:10.1186/s12864-015-2314-6.
18. Shao, R. & Barker, S. C. Mitochondrial genomes of parasitic arthropods: implications for studies of population genetics and evolution. Parasitology 134, 153-167 (2007).
19. Waterhouse, R. M., Zdobnov, E. M., Tegenfeldt, F., Li, J. & Kriventseva, E. V. OrthoDB: the hierarchical catalog of eukaryotic orthologs in 2011. Nucleic Acids Res. 39, D283-D288 (2011).
20. Csurös, M., Rogozin, I. B. & Koonin, E. V. A detailed history of intron-rich eukaryotic ancestors inferred from a global survey of 100 complete genomes. PLoS Comput. Biol. 7, e1002150 (2011).
21. Pichu, S., Ribeiro, J. M. C. & Mather, T. N. Purification and characterization of a novel salivary antimicrobial peptide from the tick, Ixodes scapularis. Biochem. Biophys. Res. Commun. 390, 511-515 (2009).
22. Chmelar, J., Calvo, E., Pedra, J. H., Francischetti, I. M. & Kotsyfakis, M. Tick salivary secretion as a source of antihemostatics. J. Proteomics 75, 3842-3854 (2012).
23. Schultz, J., Copley, R. R., Doerks, T., Ponting, C. P. & Bork, P. SMART: a web-based tool for the study of genetically mobile domains. Nucleic Acids Res. 28, 231-234 (2000).
24. Sangamnatdej, S., Paesen, G. C., Slovak, M. & Nuttall, P. A. A high affinity serotonin- and histamine-binding lipocalin from tick saliva. Insect Mol. Biol. 11, 79-86 (2002).
25. Francischetti, I. M., Mather, T. N. & Ribeiro, J. M. Cloning of a salivary gland metalloprotease and characterization of gelatinase and fibrin(ogen)lytic activities in the saliva of the Lyme disease tick vector Ixodes scapularis. Biochem. Biophys. Res. Commun. 305, 869-875 (2003).
26. Horn, M. et al. Hemoglobin digestion in blood-feeding ticks: mapping a multipeptidase pathway by functional proteomics. Chem. Biol. 16, 1053-1063 (2009).
27. Yamaji, K. et al. Hemoglobinase activity of a cysteine protease from the ixodid tick Haemaphysalis longicornis. Parasitol. Int. 58, 232-237 (2009).
28. Estrela, A. B., Seixas, A., Teixeira Vde, O., Pinto, A. F. & Termignoni, C. Vitellin- and hemoglobin-digesting enzymes in Rhipicephalus (Boophilus) microplus larvae and females. Comp. Biochem. Physiol. B Biochem. Mol. Biol. 157, 326-335 (2010).
29. Lara, F. A., Lins, U., Bechara, G. H. & Oliveira, P. L. Tracing heme in a living cell: hemoglobin degradation and heme traffic in digest cells of the cattle tick Boophilus microplus. J. Exp. Biol. 208, 3093-3101 (2005).
30. Cavallaro, G., Decaria, L. & Rosato, A. Genome-based analysis of heme biosynthesis and uptake in prokaryotic systems. J. Proteome Res. 7, 4946-4954 (2008).
31. Braz, G. R. C., Coelho, H. S. L., Masuda, H. & Oliveira, P. L. A missing metabolic pathway in the cattle tick Boophilus microplus. Curr. Biol. 9, 703-706 (1999).
32. Graca-Souza, A. V. et al. Adaptations against heme toxicity in blood-feeding arthropods. Insect Biochem. Mol. Biol. 36, 322-335 (2006).
33. Roe, R. M. et al. in Hormonal regulation of metamorphosis and reproduction in ticks (eds Sonenshine, D. E. & Roe, R. M.) Volume I pp 416-448 (Oxford University Press, 2014).
34. Roller, L. et al. Ecdysis triggering hormone signaling in arthropods. Peptides 31, 429-441 (2010).
35. Donohue, K. V. et al. Neuropeptide signaling sequences identified by pyrosequencing of the American dog tick synganglion transcriptome during blood feeding and reproduction. Insect Biochem. Mol. Biol. 40, 79-90 (2010).
36. Egekwu, N. I. et al. Comparing synganglion neuropeptides, neuropeptide receptors and neurotransmitter receptors and their gene expression in response to feeding in Ixodes scapularis (Ixodidae) versus Ornithodoros turicata (Argasidae) Insect Mol. Biol. doi:10:1111/imb.12202.
37. Hauser, F. et al. A genome-wide inventory of neurohormone GPCRs in the red flour beetle Tribolium castaneum. Front. Neuroendocrinol. 29, 142-165 (2008).
38. Nässel, D. R. & Winther, A. M. Drosophila neuropeptides in regulation of physiology and behavior. Prog. Neurobiol. 92, 42-104 (2010).
39. Zhu, J. et al. Mevalonate-farnesal biosynthesis in ticks: comparative synganglion transcriptomics and a new perspective PLoS ONE. doi:10.1371/journal.pone.0141084.
40. Kirkness, E. F. et al. Genome sequences of the human body louse and its primary endosymbiont provide insights into the permanent parasitic lifestyle. Proc. Natl Acad. Sci. USA 107, 12168-12173 (2010).
41. Touhara, K. & Vosshall, L. B. Sensing odorants and pheromones with chemosensory receptors. Annu. Rev. Physiol. 71, 307-332 (2009).
42. Peñalva-Arana, D. C., Lynch, M. & Robertson, H. M. The chemoreceptor genes of the waterflea, Daphnia pulex: many Grs but no Ors. BMC Evol. Biol. 9, 79 (2009).
43. Starostina, E., Xu, A., Lin, H. & Pikielny, C. W. A Drosophila protein family implicated in pheromone perception is related to Tay-Sachs GM2-activator protein. J. Biol. Chem. 284, 585-594 (2009).
44. Vieira, F. G. & Rozas, J. Comparative genomics of the odorant-binding and chemosensory protein gene families across the Arthropoda: origin and evolutionary history of the chemosensory system. Genome Biol. Evol. 3, 476-490 (2011).
45. Croset, V. et al. Ancient protostome origin of chemosensory ionotropic glutamate receptors and the evolution of insect taste and olfaction. PLoS. Genet. 6, e1001064 (2010).
46. Jones, W. D., Cayirlioglu, P., Kadow, I. G. & Vosshall, L. B. Two chemosensory receptors together mediate carbon dioxide detection in Drosophila. Nature 445, 86-90 (2007).
47. Carr, A. L. & Roe, R. M. Acarine attractants: chemoreception, bioassay, chemistry and control Pest. Biochem. Physiol. doi:10.1016/j.pestbp.2015.12.009.
48. Carr, A. L., Sonenshine, D. E., Strider, Jr J. B. & Roe, R. M. Evidence of female sex pheromones and characterization of the cuticular lipids of unfed, adult male versus female blacklegged ticks, Ixodes scapularis Exp. Appl. Acarol. doi:10.1007/s10493-015-0009-y.
49. Velarde, R. A., Sauer, C. D., Walden, K. K., Fahrbach, S. E. & Robertson, H. M. Pteropsin: a vertebrate-like non-visual opsin expressed in the honey bee brain. Insect Biochem. Mol. Biol. 35, 1367-1377 (2005).
50. Brody, T. & Cravchik, A. Drosophila melanogaster G protein-coupled receptors. J. Cell Biol. 150, F83-F88 (2000).
51. Hajdušek, O. et al. Interactions of the tick immune system with transmitted pathogens. Front. Cell. Infect. Microbiol. 3, 1-15 (2013).
52. Rikihisa, Y. Mechanisms of obligatory intracellular infection with Anaplasma phagocytophilum. Clin. Microbiol. Rev. 24, 469-489 (2011).
53. Severo, M. S., Pedra, J. H. F., Ayllón, N., Kocan, K. M. & de la Fuente, J. in Molecular Medical Microbiology 2nd edn, Volume 3, Ch. 110 (eds Tang, Y.-W., Sussman, M., Liu, D., Poxton, I. & Schwartzman, J.) pp 2033-2042 (Academic Press, Elsevier, 2015).
54. Sultana, H. et al. Anaplasma phagocytophilum induces actin phosphorylation to selectively regulate gene transcription in Ixodes scapularis ticks. J. Exp. Med. 207, 1727-1743 (2010).
55. Ayllón, N. et al. Anaplasma phagocytophilum inhibits apoptosis and promotes cytoskeleton rearrangement for infection of tick cells. Infect. Immun. 81, 2415-2425 (2013).
56. Ayllón, N. et al. Systems biology of tissue-specific response to Anaplasma phagocytophilum reveals differentiated apoptosis in the tick vector Ixodes scapularis. PLoS. Genet. 11, e1005120 (2015).
57. Villar, M. et al. Integrated metabolomics, transcriptomics and proteomics identifies metabolic pathways affected by Anaplasma phagocytophilum infection in tick cells. Mol. Cell. Proteomics 14, 3154-3172 (2015).
58. Huang, H., Wang, X., Kikuchi, T., Kumagai, Y. & Rikihisa, Y. Porin activity of Anaplasma phagocytophilum outer membrane fraction and purified P44. J. Bacteriol. 189, 1998-2006 (2007).
59. Neelakanta, G., Sultana, H., Fish, D., Anderson, J. F. & Fikrig, E. Anaplasma phagocytophilum induces Ixodes scapularis ticks to express an antifreeze glycoprotein gene that enhances their survival in the cold. J. Clin. Invest. 120, 3179-3190 (2010).
60. Busby, A. T. et al. Expression of heat-shock proteins and subolesin affects stress responses, Anaplasma phagocytophilum infection and questing behavior in the tick, Ixodes scapularis. Med. Vet. Entomol. 26, 92-102 (2012).
61. Gillespie, J. J. et al. A Rickettsia genome overrun by mobile genetic elements provides insight into the acquisition of genes characteristic of an obligate intracellular lifestyle. J. Bacteriol. 194, 376-394 (2012).
62. Schnettler, E. et al. Induction and suppression of tick cell antiviral RNAi responses by tick-borne flaviviruses. Nucleic Acids Res. 42, 9436-9446 (2014).
63. Grabowski, J. M. et al. Changes in the proteome of Langat-infected Ixodes scapularis ISE6 cells: metabolic pathways associated with flavivirus infection PLoS Negl. Trop. Dis. doi:10.1371/journal.pntd.0004180.
64. Oliver, Jr J. H. et al. Conspecificity of the ticks Ixodes scapularis and I. dammini (Acari: Ixodidae). J. Med. Entomol. 30, 54-63 (1993).
65. Diuk-Wasser, M. A. et al. Spatiotemporal patterns of host-seeking Ixodes scapularis nymphs (Acari: Ixodidae) in the United States. J. Med. Entomol. 43, 166-176 (2006).
66. Nuss, A. B. et al. Dopamine receptor antagonists as new mode-of-action insecticide leads for control of Aedes and Culex mosquito vectors. PLoS Negl. Trop. Dis. 9, e0003515 (2015).
67. Lees, K. & Bowman, A. S. Tick neurobiology: recent advances and the post-genomic era. Invert. Neurosci 7, 183-198 (2007).
68. Hill, C. A. Genome analysis of the major tick and mite vectors of human pathogens. NIH-NIAID-NHGRI, Preprint at http://www.genome.gov/Pages/Research/DER/PathogensandVectors/Tick-and-Mite-Genomes-Cluster-White-Paper-12Jan2011.pdf (2010).
69. King, L. J. et al. Executive summary of the AVMA One Health Initiative Task Force report. J. Am. Vet. Med. Assoc. 233, 259-261 (2008).