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Topic:Equine Diseases

Equine diseases encompass a wide range of health conditions that can affect horses, including infectious diseases, metabolic disorders, and genetic conditions. These diseases can impact the overall health, performance, and well-being of horses. Common equine diseases include equine influenza, equine herpesvirus, laminitis, and equine metabolic syndrome. Diagnosis and management of these diseases often require a combination of clinical evaluation, laboratory testing, and appropriate treatment strategies. This page gathers peer-reviewed research studies and scholarly articles that explore the etiology, pathophysiology, diagnosis, and treatment options for various equine diseases, providing valuable insights for veterinarians and researchers in the field.
Subpopulations of equine blood lymphocytes expressing regulatory T cell markers.
Veterinary immunology and immunopathology    December 1, 2010   Volume 140, Issue 1-2 90-101 doi: 10.1016/j.vetimm.2010.11.020
Robbin MG, Wagner B, Noronha LE, Antczak DF, de Mestre AM.Several distinct T lymphocyte subpopulations with immunoregulatory activity have been described in a number of mammalian species. This study performed a phenotypic analysis of cells expressing regulatory T cell (Treg) markers in the peripheral blood of a cohort of 18 horses aged 6 months to 23 years, using antibodies to both intracellular and cell surface markers, including Forkhead box P3 (FOXP3), CD4, CD8, CD25, interferon gamma (IFNγ) and interleukin 10 (IL-10). In peripheral blood, a mean of 2.2 ± 0.2% CD4+ and 0.5 ± 0.1% CD8+ lymphocytes expressed FOXP3. The mean percentage of CD4+FOXP...
Equine influenza virus: a jumping virus that races with Thoroughbred horses and greyhounds.
Veterinary journal (London, England : 1997)    November 30, 2010   Volume 189, Issue 1 3-4 doi: 10.1016/j.tvjl.2010.10.023
Murcia PR, Wood JL.No abstract available
Use of in vitro technologies to study phase II conjugation in equine sports drug surveillance.
Bioanalysis    November 30, 2010   Volume 2, Issue 12 1971-1988 doi: 10.4155/bio.10.135
Taylor P, Scarth JP, Hillyer LL.Within equine drug surveillance, there is significant interest in analyzing intact phase II conjugates of drugs in urine, but progress has been limited by a lack of reference material. Methods: In this study, in vitro techniques using equine liver fractions were employed to produce glucuronide and sulfate conjugates of stanozolol, 16β-hydroxystanozolol and nandrolone, the glucuronide conjugate of morphine and the glutathione metabolite of chlordinitrobenzene for the first time in equine sports drug surveillance. Results: The glucuronide conjugate of the synthetic progestagen altrenogest was a...
Prevalence of the mutation in cyclophilin B (PPIB), a causal candidate gene for HERDA, among Quarter Horses in France.
Veterinary dermatology    November 30, 2010   Volume 22, Issue 2 206-208 doi: 10.1111/j.1365-3164.2010.00941.x
White SD, Bourdeau P.Hereditary equine regional dermal asthenia (HERDA) in Quarter Horses is an inherited degenerative skin disease. Initially reported as hyperelastosis cutis, HERDA has a phenotype of hyperextensible, fragile skin, with secondary seromas, haematomas, ulcers and scarring. It primarily affects the dorsal aspect of the body. An autosomal recessive mode of inheritance is considered likely, with affected horses more at risk to produce affected offspring. A mutation in cyclophilin B (PPIB) as a novel, causal candidate gene for HERDA has been described, and verified as segregating with carriers and affe...
Mapping of equine cerebellar abiotrophy to ECA2 and identification of a potential causative mutation affecting expression of MUTYH.
Genomics    November 30, 2010   Volume 97, Issue 2 121-129 doi: 10.1016/j.ygeno.2010.11.006
Brault LS, Cooper CA, Famula TR, Murray JD, Penedo MC.Equine Cerebellar Abiotrophy (CA) is a neurological disease found in Arabian horses. CA is characterized by post-natal degeneration of the Purkinje cells of the cerebellum. Signs of CA include ataxia, head tremors, and a lack of balance equilibrium. We have discovered a linkage of the CA phenotype to a microsatellite marker on ECA2 and identified a region of conserved homozygosity spanning approximately 142 kb. Complete sequencing of the four genes in this region identified one SNP found only in Arabian horses, located in exon 4 of TOE1 and approximately 1200 base pairs upstream of MUTYH, adja...
Evaluation of cytokine mRNA expression in bronchoalveolar lavage cells from horses with inflammatory airway disease.
Veterinary immunology and immunopathology    November 30, 2010   Volume 140, Issue 1-2 82-89 doi: 10.1016/j.vetimm.2010.11.018
Hughes KJ, Nicolson L, Da Costa N, Franklin SH, Allen KJ, Dunham SP.Inflammatory airway disease (IAD) is a common disorder of performance horses and is associated with poor performance and accumulation of mucus and inflammatory cells in lower airway secretions. Horses with IAD frequently have increased relative counts of neutrophils in bronchoalveolar lavage fluid (BALF); less commonly relative counts of eosinophils and/or mast cells may be increased. The aetiopathogenesis of IAD is unknown and may involve innate and/or acquired immune responses to various factors including respirable dust constituents, micro-organisms, noxious gases and unconditioned air. The...
Effects of high mobility group box protein-1, interleukin-1β, and interleukin-6 on cartilage matrix metabolism in three-dimensional equine chondrocyte cultures.
Connective tissue research    November 30, 2010   Volume 52, Issue 4 290-300 doi: 10.3109/03008207.2010.523803
Ley C, Svala E, Nilton A, Lindahl A, Eloranta ML, Ekman S, Skiöldebrand E.The effects of high mobility group box protein (HMGB)-1, interleukin (IL)-1β, and IL-6 on equine articular chondrocytes were investigated, with emphasis on detecting differences between anatomical sites exposed to different loading in vivo, using three-dimensional (3D) cell cultures established with chondrocytes from dorsal radial facet (DRF, highly loaded) and palmar condyle (PC, less loaded) of the third carpal bone (C3). Expression of important genes involved in cartilage metabolism, presence of glycosaminoglycans and cartilage oligomeric matrix protein (COMP) in pellets, and concentration...
RT-qPCR comparison of mast cell populations in whole blood from healthy horses and those with laminitis.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 16-22 doi: 10.1111/j.1365-2052.2010.02093.x
Brooks SA, Bailey E.Inflammatory damage to the digital laminae, a structure responsible for suspension of the distal skeleton within the hoof capsule, results in a painful and often life-threatening disease in horses called laminitis. There can be many diverse causes of laminitis; however, previous work in the horse has suggested that in each case, the inflammation and resulting tissue damage is consistent with the action of mediators released from mast cells (MC), as well as the downstream consequences of their activation. The recent development of molecular genetics tools to characterize cells based on their tr...
Genome-wide SNP association-based localization of a dwarfism gene in Friesian dwarf horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 2-7 doi: 10.1111/j.1365-2052.2010.02091.x
Orr N, Back W, Gu J, Leegwater P, Govindarajan P, Conroy J, Ducro B, Van Arendonk JA, MacHugh DE, Ennis S, Hill EW, Brama PA.The recent completion of the horse genome and commercial availability of an equine SNP genotyping array has facilitated the mapping of disease genes. We report putative localization of the gene responsible for dwarfism, a trait in Friesian horses that is thought to have a recessive mode of inheritance, to a 2-MB region of chromosome 14 using just 10 affected animals and 10 controls. We successfully genotyped 34,429 SNPs that were tested for association with dwarfism using chi-square tests. The most significant SNP in our study, BIEC2-239376 (P(2df)=4.54 × 10(-5), P(rec)=7.74 × 10(-6)), is lo...
Genetics of swayback in American Saddlebred horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 64-71 doi: 10.1111/j.1365-2052.2010.02108.x
Cook D, Gallagher PC, Bailey E.Extreme lordosis, also called swayback, lowback or softback, can occur as a congenital trait or as a degenerative trait associated with ageing. In this study, the hereditary aspect of congenital swayback was investigated using whole genome association studies of 20 affected and 20 unaffected American Saddlebred (ASB) Horses for 48,165 single-nucleotide polymorphisms (SNPs). A statistically significant association was identified on ECA20 (corrected P=0.017) for SNP BIEC2-532523. Of the 20 affected horses, 17 were homozygous for this SNP when compared to seven homozygotes among the unaffected ho...
Fine mapping of a quantitative trait locus for osteochondrosis on horse chromosome 2.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 87-90 doi: 10.1111/j.1365-2052.2010.02113.x
Dierks C, Komm K, Lampe V, Distl O.In this study, we refine a quantitative trait locus for equine osteochondrosis (OC) on horse chromosome (ECA) 2 to a genome-wide significant interval at 20.08-30.94 Mb. The marker set contained 27 newly developed microsatellites equidistantly distributed over ECA2 and 44 nucleotide polymorphisms, located in 16 positional candidate genes for OC. Genotyping was performed in 211 Hanoverian horses from 14 paternal half-sib groups. A NCDN-associated SNP and haplotype were significantly associated with OC in fetlock and/or hock joints. This study is a further step towards the identification of genes...
Genome-wide association analysis of osteochondrosis of the tibiotarsal joint in Norwegian Standardbred trotters.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 111-120 doi: 10.1111/j.1365-2052.2010.02117.x
Lykkjen S, Dolvik NI, McCue ME, Rendahl AK, Mickelson JR, Roed KH.Osteochondrosis (OC), a disturbance in the process of endochondral ossification, is by far the most important equine developmental orthopaedic disease and is also common in other domestic animals and humans. The purpose of this study was to identify quantitative trait loci (QTL) associated with osteochondrosis dissecans (OCD) at the intermediate ridge of the distal tibia in Norwegian Standardbred (SB) using the Illumina Equine SNP50 BeadChip whole-genome single-nucleotide polymorphism (SNP) assay. Radiographic data and blood samples were obtained from 464 SB yearlings. Based on the radiographi...
Comparative human-horse sequence analysis of the CYP3A subfamily gene cluster.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 72-79 doi: 10.1111/j.1365-2052.2010.02111.x
Schmitz A, Demmel S, Peters LM, Leeb T, Mevissen M, Haase B.Cytochrome P450 enzymes (CYP450s) represent a superfamily of haem-thiolate proteins. CYP450s are most abundant in the liver, a major site of drug metabolism, and play key roles in the metabolism of a variety of substrates, including drugs and environmental contaminants. Interaction of two or more different drugs with the same enzyme can account for adverse effects and failure of therapy. Human CYP3A4 metabolizes about 50% of all known drugs, but little is known about the orthologous CYP450s in horses. We report here the genomic organization of the equine CYP3A gene cluster as well as a compara...
Linkage disequilibrium and historical effective population size in the Thoroughbred horse.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 8-15 doi: 10.1111/j.1365-2052.2010.02092.x
Corbin LJ, Blott SC, Swinburne JE, Vaudin M, Bishop SC, Woolliams JA.Many genomic methodologies rely on the presence and extent of linkage disequilibrium (LD) between markers and genetic variants underlying traits of interest, but the extent of LD in the horse has yet to be comprehensively characterized. In this study, we evaluate the extent and decay of LD in a sample of 817 Thoroughbreds. Horses were genotyped for over 50,000 single nucleotide polymorphism (SNP) markers across the genome, with 34,848 autosomal SNPs used in the final analysis. Linkage disequilibrium, as measured by the squared correlation coefficient (r(2)), was found to be relatively high bet...
Refinement of quantitative trait loci on equine chromosome 10 for radiological signs of navicular disease in Hanoverian warmblood horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 36-40 doi: 10.1111/j.1365-2052.2010.02096.x
Lopes MS, Diesterbeck U, Machado Ada C, Distl O.Navicular disease is characterized by a progressive degenerative alteration of the equine podotrochlea. In this study, we refined a previously identified quantitative trait locus (QTL) on horse chromosome 10 for the abnormal development of canales sesamoidales (DCS) of the navicular bone in Hanoverian warmblood horses. Genotyping was done in 192 Hanoverian warmblood horses from 17 paternal half-sib groups. The whole marker set comprised 45 markers including seven newly developed microsatellites and 13 single nucleotide polymorphisms (SNPs) within positional candidate genes. Chromosome-wide sig...
Use of pressor therapy in 34 hypotensive critically ill neonatal foals.
Australian veterinary journal    November 26, 2010   Volume 88, Issue 12 472-477 doi: 10.1111/j.1751-0813.2010.00652.x
Dickey EJ, McKenzie H, Johnson A, Furr MO.Arginine vasopressin (AVP) is used in human medicine in the management of vasodilatory shock and cardiac arrest, but it is not widely used in equine neonatal intensive care because of concerns about potential side effects and suboptimal efficacy. This retrospective study reports the clinical use of AVP and norepinephrine (NE) in foals with refractory hypotension. Objective: To report the cardiovascular responses and fluid balance in critically ill, hypotensive foals receiving either NE or AVP. Methods: The medical records of neonatal foals (<7 days of age) from 2000 to 2007 admitted to the ...
A survey of aged horses in Queensland, Australia. Part 2: Clinical signs and owners’ perceptions of health and welfare.
Australian veterinary journal    November 26, 2010   Volume 88, Issue 12 465-471 doi: 10.1111/j.1751-0813.2010.00638.x
McGowan TW, Pinchbeck G, Phillips CJ, Perkins N, Hodgson DR, McGowan CM.To describe the prevalence and risk factors for clinical signs of disease and owner-reported health or welfare issues of aged horses in Queensland, Australia. Methods: Owners of horses were contacted via an equestrian organisation in Queensland and asked to complete a questionnaire about their horses aged 15 years and older, to gain information about clinical signs observed in the horse and disease history. Owners were asked to identify health or welfare issues they felt were important in aged horses. Results: Owners were able to identify many clinical signs of disease, with 83% of horses havi...
Identification of equine major histocompatibility complex haplotypes using polymorphic microsatellites.
Animal genetics    November 26, 2010   Volume 41 Suppl 2, Issue Suppl 2 150-153 doi: 10.1111/j.1365-2052.2010.02125.x
Tseng CT, Miller D, Cassano J, Bailey E, Antczak DF.A system for identifying equine major histocompatibility complex (MHC) haplotypes was developed based on five polymorphic microsatellites located within the MHC region on ECA 20. Molecular signatures for 50 microsatellite haplotypes were recognized from typing 353 horses. Of these, 23 microsatellite haplotypes were associated with 12 established equine leucocyte antigen (ELA) haplotypes in Thoroughbreds and Standardbreds. Five ELA serotypes were associated with multiple microsatellite subhaplotypes, expanding the estimates of diversity in the equine MHC. The strong correlations between serolog...
Molecular heterogeneity of XY sex reversal in horses.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 41-52 doi: 10.1111/j.1365-2052.2010.02101.x
Raudsepp T, Durkin K, Lear TL, Das PJ, Avila F, Kachroo P, Chowdhary BP.Male-to-female 64,XY sex reversal is a frequently reported chromosome abnormality in horses. Despite this, the molecular causes of the condition are as yet poorly understood. This is partially because only limited molecular information is available for the horse Y chromosome (ECAY). Here, we used the recently developed ECAY map and carried out the first comprehensive study of the Y chromosome in XY mares (n=18). The integrity of the ECAY in XY females was studied by FISH and PCR using markers evenly distributed along the euchromatic region. The results showed that the XY sex reversal condition...
Genomic structure, polymorphism and expression of ACCN1 and ACCN3 genes in the horse.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 138-144 doi: 10.1111/j.1365-2052.2010.02123.x
Mata X, Ducasse A, Vaiman A, Diribarne M, Fraud AS, Guérin G.A category of cation gate proteins was shown to be present in sensory neurons and act as receptors of protons present in tissues such as muscles. The Amiloride-sensitive Cation Channel, Neuronal (ACCN) gene family is known to play a role in the transmission of pain through specialized pH sensitive neurons. Muscles from horses submitted to strenuous exercises produce lactic acid, which may induce variable pain through ACCN differential properties. The sequences of the equine cDNAs were determined to be 2.6 kb in length with an open reading frame of 1539 bp for ACCN1 and 2.1 kb in length with an...
A genome-wide scan for tying-up syndrome in Japanese Thoroughbreds.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 80-86 doi: 10.1111/j.1365-2052.2010.02112.x
Tozaki T, Hirota K, Sugita S, Ishida N, Miyake T, Oki H, Hasegawa T.Tying-up syndrome, also known as recurrent exertional rhabdomyolysis in Thoroughbreds, is a common muscle disorder for racehorses. In this study, we performed a multipoint linkage analysis using LOKI based on the Bayesian Markov chain Monte Carlo method using 5 half-sib families (51 affected and 277 nonaffected horses in total), and a genome-wide association study (GWAS) using microsatellites (144 affected and 144 nonaffected horses) to map candidate regions for tying-up syndrome in Japanese Thoroughbreds. The linkage analysis identified one strong L-score (82.45) between the loci UCDEQ411 and...
Estimated prevalence of the Type 1 Polysaccharide Storage Myopathy mutation in selected North American and European breeds.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 145-149 doi: 10.1111/j.1365-2052.2010.02124.x
McCue ME, Anderson SM, Valberg SJ, Piercy RJ, Barakzai SZ, Binns MM, Distl O, Penedo MC, Wagner ML, Mickelson JR.The GYS1 gene mutation that is causative of Type 1 Polysaccharide Storage Myopathy (PSSM) has been identified in more than 20 breeds of horses. However, the GYS1 mutation frequency or Type 1 PSSM prevalence within any given breed is unknown. The purpose of this study was to determine the frequency of the GYS1 mutation and prevalence of genetic susceptibility to Type 1 PSSM in selected breeds from Europe and North America. The GYS1 mutation was detected in 11 breeds, including, in order of increasing allele frequency, Shires, Morgans, Appaloosas, Quarter Horses, Paints, Exmoor Ponies, Saxon-Thu...
True stress and Poisson’s ratio of tendons during loading.
Journal of biomechanics    November 26, 2010   Volume 44, Issue 4 719-724 doi: 10.1016/j.jbiomech.2010.10.038
Vergari C, Pourcelot P, Holden L, Ravary-Plumioën B, Gerard G, Laugier P, Mitton D, Crevier-Denoix N.Excessive axial tension is very likely involved in the aetiology of tendon lesions, and the most appropriate indicator of tendon stress state is the true stress, the ratio of instantaneous load to instantaneous cross-sectional area (CSA). Difficulties to measure tendon CSA during tension often led to approximate true stress by assuming that CSA is constant during loading (i.e. by the engineering stress) or that tendon is incompressible, implying a Poisson's ratio of 0.5, although these hypotheses have never been tested. The objective of this study was to measure tendon CSA variation during qua...
A conserved segmental duplication within ELA.
Animal genetics    November 26, 2010   Volume 41 Suppl 2 186-195 doi: 10.1111/j.1365-2052.2010.02137.x
Brinkmeyer-Langford CL, Murphy WJ, Childers CP, Skow LC.The assembled genomic sequence of the horse major histocompatibility complex (MHC) (equine lymphocyte antigen, ELA) is very similar to the homologous human HLA, with the notable exception of a large segmental duplication at the boundary of ELA class I and class III that is absent in HLA. The segmental duplication consists of a ∼ 710 kb region of at least 11 repeated blocks: 10 blocks each contain an MHC class I-like sequence and the helicase domain portion of a BAT1-like sequence, and the remaining unit contains the full-length BAT1 gene. Similar genomic features were found in other Perissod...
Clinicopathologic findings following intra-articular injection of autologous and allogeneic placentally derived equine mesenchymal stem cells in horses.
Cytotherapy    November 25, 2010   Volume 13, Issue 4 419-430 doi: 10.3109/14653249.2010.536213
Carrade DD, Owens SD, Galuppo LD, Vidal MA, Ferraro GL, Librach F, Buerchler S, Friedman MS, Walker NJ, Borjesson DL.The development of an allogeneic mesenchymal stem cell (MSC) product to treat equine disorders would be useful; however, there are limited in vivo safety data for horses. We hypothesized that the injection of self (autologous) and non-self (related allogeneic or allogeneic) MSC would not elicit significant alterations in physical examination, gait or synovial fluid parameters when injected into the joints of healthy horses. Methods: Sixteen healthy horses were used in this study. Group 1 consisted of foals (n = 6), group 2 consisted of their dams (n = 5) and group 3 consisted of half-siblings ...
Fatal epizootic equine herpesvirus 1 infections in new and unnatural hosts.
Veterinary microbiology    November 24, 2010   Volume 149, Issue 3-4 456-460 doi: 10.1016/j.vetmic.2010.11.024
Wohlsein P, Lehmbecker A, Spitzbarth I, Algermissen D, Baumgärtner W, Böer M, Kummrow M, Haas L, Grummer B.In a zoological collection, four black bears (Ursus americanus) died from neurological disease within six months. Independently in a geographically different zoo, two Thomson's gazelles (Eudorcas thomsoni) and 18 guinea pigs (Cavia porcellus f. dom.) suffered from neurological disorders. In addition, guinea pigs showed abortions and stillbirths. All affected animals displayed a non suppurative meningoencephalitis with intranuclear inclusion bodies. Immunohistology demonstrated equine herpes virus antigen and ultrastructurally herpes viral particles were detected. Virus isolation and molecular ...
Three recently recognized species of cyathostomes (Nematoda: Strongylidae) in equids in Kentucky.
Parasitology research    November 24, 2010   Volume 108, Issue 5 1179-1184 doi: 10.1007/s00436-010-2160-z
Kuzmina TA, Tolliver SC, Lyons ET.Three species of cyathostomes--Cylicocyclus ashworthi, Cylicostephanus bidentatus, and Cylicostephanus hybridus were identified recently in horses in Kentucky. General characteristics and distinguishing description of these species are presented. Distribution of these species and their role in the horse strongylid community are discussed. The importance of examining the entire contents of the large intestine or alternatively a high number of specimens in order to recover and identify species residing in low numbers is stressed.
Retrospective study of necropsy-associated coagulase-positive staphylococci in horses. Haenni M, Targant H, Forest K, Sévin C, Tapprest J, Laugier C, Madec JY.Coagulase-positive staphylococci (CoPS) are potential causative agents of equine infections, but they are rarely responsible for the death of the animal. In the current study, staphylococci implicated in the death or euthanasia of horses were retrospectively studied in 3,457 necropsies performed over a decade (1995-2006). Morbidity associated with CoPS was 1.7%, representing 60 isolates of CoPS, which were identified as Staphylococcus aureus (59) and Staphylococcus pseudintermedius (1). Coagulase-positive staphylococci (alone or in association with another bacterial species) were associated wi...
Ameloblastic fibro-odontoma associated with a retained molar in an Oldenburg mare. Knowles S, Blas-Machado U, Butler AM, Gomez-Ibañez SE, Lowder MQ, Fayrer-Hosken RA.An 11-year-old Oldenburg mare presented with a 3-month history of weight loss and swelling of the posterior right mandible. Physical examination and radiographs showed a soft-tissue mass in the right mandible with foci of mineralization, periosteal proliferation, and a retained molar. The tumor increased in size over several weeks, and the mare was euthanized. On necropsy, a 24 cm × 15 cm × 15 cm firm, white mass had obliterated the posterior right mandible. The mass was multinodular with discrete clusters of blood-filled cystic spaces. Histologically, the mass was composed of neoplastic ...
A proviral derivative from a reference attenuated EIAV vaccine strain failed to elicit protective immunity.
Virology    November 20, 2010   Volume 410, Issue 1 96-106 doi: 10.1016/j.virol.2010.10.032
Ma J, Shi N, Jiang CG, Lin YZ, Wang XF, Wang S, Lv XL, Zhao LP, Shao YM, Kong XG, Zhou JH, Shen RX.To investigate essential factors that determine the efficacy of vaccines against lentiviruses, an effective attenuated equine infectious anemia virus (EIAV) vaccine strain and a proviral derivative of the vaccine were compared with respect to differences in inducing protective immunity. Although these two strains replicated equally well in vitro and in vivo, the proviral strain induced significantly less protection from disease and infection caused by viral challenge and significantly lower specific neutralizing capability. These findings indicated that the proviral strain had lost the ability...