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Uzawa, T., Akiyama, S., Kimura, T., Takahashi, S., Ishimori, K., Morishima, I., et al. (2004). Collapse and search dynamics of apomyoglobin folding revealed by submillisecond observations of alpha-helical content and compactness. Proc. Natl. Acad. Sci. U.S.A., 101(5), 1171–1176.
Abstract: The characterization of protein folding dynamics in terms of secondary and tertiary structures is important in elucidating the features of intraprotein interactions that lead to specific folded structures. Apomyoglobin (apoMb), possessing seven helices termed A-E, G, and H in the native state, has a folding intermediate composed of the A, G, and H helices, whose formation in the submillisecond time domain has not been clearly characterized. In this study, we used a rapid-mixing device combined with circular dichroism and small-angle x-ray scattering to observe the submillisecond folding dynamics of apoMb in terms of helical content (f(H)) and radius of gyration (R(g)), respectively. The folding of apoMb from the acid-unfolded state at pH 2.2 was initiated by a pH jump to 6.0. A significant collapse, corresponding to approximately 50% of the overall change in R(g) from the unfolded to native conformation, was observed within 300 micros after the pH jump. The collapsed intermediate has a f(H) of 33% and a globular shape that involves >80% of all its atoms. Subsequently, a stepwise helix formation was detected, which was interpreted to be associated with a conformational search for the correct tertiary contacts. The characterized folding dynamics of apoMb indicates the importance of the initial collapse event, which is suggested to facilitate the subsequent conformational search and the helix formation leading to the native structure.
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Ballew, R. M., Sabelko, J., & Gruebele, M. (1996). Direct observation of fast protein folding: the initial collapse of apomyoglobin. Proc. Natl. Acad. Sci. U.S.A., 93(12), 5759–5764.
Abstract: The rapid refolding dynamics of apomyoglobin are followed by a new temperature-jump fluorescence technique on a 15-ns to 0.5-ms time scale in vitro. The apparatus measures the protein-folding history in a single sweep in standard aqueous buffers. The earliest steps during folding to a compact state are observed and are complete in under 20 micros. Experiments on mutants and consideration of steady-state CD and fluorescence spectra indicate that the observed microsecond phase monitors assembly of an A x (H x G) helix subunit. Measurements at different viscosities indicate diffusive behavior even at low viscosities, in agreement with motions of a solvent-exposed protein during the initial collapse.
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Ducro, B. J., Koenen, E. P. C., van Tartwijk, J. M. F. M., & Bovenhuis, H. (2007). Genetic relations of movement and free-jumping traits with dressage and show-jumping performance in competition of Dutch Warmblood horses. Livestock Science, 107(2-3), 227–234.
Abstract: Genetic parameters for traits evaluated at the studbook entry inspection and genetic correlations with dressage and show-jumping performance in competition were estimated. Data comprised 36,649 Warmblood horses that entered the studbook between 1992 and 2002. The genetic analyses were performed using univariate and bivariate animal models. Heritabilities of the studbook entry traits were estimated in the range 0.15-0.40. The movement traits showed moderate to strong mutual genetic correlations, whereas the genetic correlations of movement traits with free-jumping traits were weak to moderate. The free-jumping traits showed strong to very strong mutual genetic correlations. Competition results of 33,459 horses with performance in dressage and 30,474 horse with performance in show-jumping were linked to the studbook entry data to estimate the genetic relationship with performance in competition. Heritability estimates for dressage and show jumping were 0.14. Genetic correlations of the movement traits with dressage were moderate to strong, and with show-jumping weak to moderate. Genetic correlations of the free-jumping traits with dressage were weak to moderate and unfavourable. The free-jumping traits were genetically strong to very strong correlated to show-jumping. It was concluded that a selection of the traits evaluated at the studbook entry inspection will favourably contribute to estimation of breeding values for sport performance.
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Ducro, B. J., Koenen, E. P. C., van Tartwijk, J. M. F. M., & van Arendonk, J. A. M. (2007). Genetic relations of First Stallion Inspection traits with dressage and show-jumping performance in competition of Dutch Warmblood horses. Livestock Science, 107(1), 81–85.
Abstract: Genetic parameters for traits evaluated at the First Stallion Inspection (FSI) and genetic correlations with dressage and show-jumping performance in competition were estimated. Data comprised 2361 stallions with FSI-observations from 1994 through 1999. Genetic analyses were performed using univariate and bivariate animal models. Heritability estimates of the FSI-traits ranged from 0.25 to 0.61. FSI-traits related to gaits showed strong genetic correlations (above 0.70) and FSI-traits related to free jumping had correlations close to unity. Competition results of 23,897 horses with performance in dressage and 22,811 horses with performance in show-jumping were linked to the FSI data to estimate the genetic relationship with performance in competition. Heritability estimates for dressage and show-jumping were 0.14. Genetic correlation between FSI-gaits and dressage in competition were positive, ranging from 0.37 to 0.72. Genetic correlation between FSI-jumping traits and show-jumping were all above 0.80. FSI-jumping traits showed negative correlations with dressage (about – 0.48). FSI-gait traits showed negative correlations with show-jumping, except for canter. It is concluded that selection at First Stallion Inspection comprises an important component of the stallion selection program, because FSI-traits have good genetic relationships with performance in competition and, due to the number of animals involved, relative high selection intensities can be achieved.
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Stock, K. F., & Distl, O. (2005). Prediction of breeding values for osseous fragments in fetlock and hock joints, deforming arthropathy in hock joints, and pathologic changes in the navicular bones of Hanoverian Warmblood horses. Livestock Production Science, 92(1), 77–94.
Abstract: The results of a standardised radiological examination of 5928 Hanoverian Warmblood horses selected for sale at auction were used to predict relative breeding values (RBV) in the 23,662 horses included in the last four generations of the probands' pedigrees. The distribution of the RBV was investigated in the whole four-generation pedigree, in the probands and in those stallions that contributed at least three offspring to the probands. The RBV of the probands' sires were further compared with the officially published performance-based relative breeding values, i.e., total indices dressage (TID) and jumping (TIJ). The sires' level of dressage indices was considerably higher (mean TID=110) than of the jumping indices (mean TIJ=98). Total indices radiographic findings (TIR) were calculated for the sires with varying weighting given to the individual RBV. In each case, this resulted in a mean TIR of 99. Finally, total indices were derived from TIR and TID and/or TIJ in order to develop different selection schemes for all-purpose breeding and for breeding focused on dressage and show jumping. All breeding values under consideration increased by between 1% and 19% when radiographic findings were weighted with between 30% and 60% complementary to the corresponding performance parameters, and when only sires were selected with above average total indices. At the same time, the prevalences of the radiographic findings investigated here were lowered by up to 10% each. When only one radiographic finding was considered at a time, the maximum attainable response to selection was a 16-23% increase in the relative breeding values and a relative decrease in prevalences of radiographic findings of between 31% and 52%. The results of this study indicate that it is possible in horse breeding to consider simultaneously health and performance traits. Medical data should be included in the prediction of breeding values in order to improve the radiological status of today's riding horses.
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Koenen, E. P. C., Aldridge, L. I., & Philipsson, J. (2004). An overview of breeding objectives for warmblood sport horses. Livestock Production Science, 88(1-2), 77–84.
Abstract: The aim of this paper is to review the current breeding objectives of organisations that run a selection programme for warmblood riding horses in the light of an increasing trend in trade of semen across countries. In a questionnaire, 19 horse breeding organisations provided information on breeding objective traits. Variation both in length and amount of details used to define individual breeding objectives was large, reflecting that many traits in sport horse breeding are not easy to measure, and therefore, have to be defined in a subjective way. The majority of the breeding objectives included conformation, gaits and performance in show jumping and dressage. Some breeding objectives also included behaviour, soundness, health and fertility. However, several organisations did not specify the sport discipline and the level of competition (amateur, national or international level) in the breeding objective. In general, relative weightings of the traits within the verbally presented breeding objectives were not given, but were assessed by the organisations in response to this study. The relevance of more information on expected future production circumstances and on the genetic parameters of the traits of interest are discussed. A further review of the consistency, completeness and the number of traits of the present breeding objectives for sport horses is recommended to optimise the efficiency of selection decisions.
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Wallin, L., Strandberg, E., & Philipsson, J. (2003). Genetic correlations between field test results of Swedish Warmblood Riding Horses as 4-year-olds and lifetime performance results in dressage and show jumping. Livestock Production Science, 82(1), 61–71.
Abstract: The main objective of this study was to estimate genetic correlations between traits of young sport horses (4 years old) evaluated in the Swedish Riding Horse Quality Test (RHQT) and later competition results in dressage and show jumping. The data comprised 3708 Warmblood horses born between 1968 and 1982 that had participated in the RHQT as 4-year-olds and 25[punctuation space]605 horses born between 1953 and 1995 with competition records. According to the criteria between 1206 and 1879 horses were common to this two files and were available for the estimations of the genetic correlations. Competition performance traits were cumulative points and cumulative placings received during a horse's lifetime, and a log10 transformation was used to achieve a more normal distribution of the data. Genetic correlations between gait traits scored in the RHQT and competition results in dressage were favourable, in the range 0.63-0.75, and between jumping traits scored in the RHQT and results in show jumping 0.83-0.93. Estimated heritabilities for gait and jumping traits scored in the RHQT were in the range 0.09-0.27 and 0.10-0.18, respectively. Estimated heritabilities for the cumulative points and cumulative placings in dressage and show jumping were 0.17/0.16 and 0.23/0.27, respectively. Thus, the results from the RHQT have proved to be useful for early genetic evaluation and selection of both mares and stallions for sport performance traits.
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Dyson, S. (2002). Lameness and poor performance in the sport horse: Dressage, show jumping and horse trials. Journal of Equine Veterinary Science, 22(4), 145–150.
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Wallin, L., Strandberg, E., & Philipsson, J. (2001). Phenotypic relationship between test results of Swedish Warmblood horses as 4-year-olds and longevity. Livestock Production Science, 68(2-3), 97–105.
Abstract: The relationship between longevity and different traits scored in the Swedish Riding Horse Quality Test (RHQT) was studied to evaluate their use as predictors of survival. Data comprised 1815 Warmblood horses born between 1969 and 1982 that had participated in the RHQT as 4-year-olds. Survival information was obtained via a questionnaire sent to owners of horses that had participated in the RHQT between 1973 and 1986. All phenotypic values of traits scored at 4 years of age were adjusted for the effect of place/year (event). Survival analysis was performed taking into account censoring. Traits having significant effects on longevity were: conformation, legs (included in conformation), orthopaedic status, jumping ability, and the horses' combined classification score for dressage and jumping talents, respectively. Orthopaedic health had the greatest influence on longevity, and demonstrated the importance of judging health traits in young sports horses. The results of this study confirmed that there is a significant phenotypic relationship between many of the RHQT traits and longevity, and thus the possibility of using them as predictors of survival.
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Hodson, E. F., Clayton, H. M., & Lanovaz, J. L. (1999). Temporal analysis of walk movements in the Grand Prix dressage test at the 1996 Olympic Games. Appl. Anim. Behav. Sci., 62(2-3), 89–97.
Abstract: Video analysis was used to measure temporal characteristics of the collected walk, extended walk and half pirouette at walk of eleven competitors during the team dressage competition at the 1996 Summer Olympic Games in Atlanta, GA. Forelimb stance durations, hind limb stance durations, lateral step intervals and diagonal step intervals were symmetrical for the right and left sides in the collected and extended walk strides, but there were left-right asymmetries in the forelimb stance duration and in the lateral step interval in the half pirouette strides. For both collected and extended walk strides, hind limb stance duration was significantly longer than forelimb stance duration. The mean values for the group of eleven horses showed that the collected and extended walks had a regular rhythm. The half pirouette strides showed an irregularity in which there was a short interval between footfalls of the outside forelimb and inside hind limb, and along interval between footfalls of the inside hind limb and inside forelimb. This irregularity reflected an early placement of the inside hind limb. The stance times of both hind limbs were prolonged and this finding, in combination with the early placement of the inside hind limb, led to an increase in the period of tripedal support in each stride of the half pirouette. This was interpreted as a means of maintaining the horses' balance in the absence of forward movement.
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