Lucas Rivero - Academia.edu (original) (raw)

Papers by Lucas Rivero

Journal of Veterinary Medicine Series A, 2007

The main goal of any conditioning programme in athletic horses is to improve performance by induc... more The main goal of any conditioning programme in athletic horses is to improve performance by inducing physiological changes within the animal's body. Equine skeletal muscles have a considerable potential to adapt during training and these adaptations have important physiological implications that influence stamina, strength and speed. Although there is an extensive specialized literature in this regard, scientific based muscle conditioning methods have not been introduced sufficiently in the equine sport practice. After a brief synopsis of both equine muscle exercise physiology and muscular adaptations to training, including their physiological significance, this review focuses on specific training programmes that induce muscular adaptations in athletic horses. The article addresses the following principal question: what kind of stimuli for what kind of muscular adaptations? The experimental data are discussed separately for racehorses (thoroughbreds, trotters and endurance horses) and sport horses (dressage, show jumpers and carriage). Finally, published results about the influence of relevant training parameters (such as intensity, duration and type of exercise) on muscular responses are discussed, as well as those concerning overtraining and detraining. The article closes with some concluding remarks on importance of their application in practice.

Veterinary Record, 2007

FIG 1: Longitudinal section of a sample of diaphragm from a horse with atypical myopathy (case 1)... more FIG 1: Longitudinal section of a sample of diaphragm from a horse with atypical myopathy (case 1). Multifocal and segmental necrosis and Zenker’s degeneration (arrow) in multiple myofibres are visible. Haematoxylin and eosin. Bar=75 μm FIG 2: Transverse section of the gluteus profundus muscle of a horse with atypical myopathy (case 2). Increased intercellular spaces (oedema) and multiple hypercontracted muscle fibres with pale pink cytoplasm (arrows) and severe degeneration of the myofibrillar apparatus are visible. Haematoxylin and eosin. Bar=80 μm Veterinary Record (2007) 161, 346-348

Pflügers Archiv - European Journal of Physiology, 2002

Fifteen healthy, untrained male subjects (mean age ±SD, 22±5 years) were used to examine the plas... more Fifteen healthy, untrained male subjects (mean age ±SD, 22±5 years) were used to examine the plasticity of myosin heavy chain phenotype, size, oxidative capacity and capillarization of skeletal muscle fibre types with short-term electrical stimulation (ES). Ten subjects were electro-stimulated on both quadriceps muscles with a frequency of 45-60 Hz, with 12 s of stimulation followed by 8 s of recovery for a total of 30 min per day, 3 days per week for 6 weeks. The remaining five subjects served as controls. Two vastus lateralis muscle biopsy samples were removed from each subject before (week 0) and after (week 6) ES training. A standardized exercise test on a cycle ergometer was performed by each subject before and after the experimental period and several indicators of whole-body aerobic capacity were estimated. The so-called electromyographic threshold was also determined during the tests. Muscle biopsy samples were analysed by electrophoresis, immunohistochemistry and quantitative histochemistry. Myosin heavy chain (MHC) composition, muscle fibre type distribution, fibre areas, oxidative capacity and capillaries of each fibre type were estimated. Muscular changes with ES revealed an increase of fibres expressing MHC-IIA, and a decrease of fibres expressing MHC-IIX and MHC-I, as well as an increase of the oxidative capacity and mean number of capillaries of fast-twitch (type II) fibres with minimal muscle fibre hypertrophy. These adaptations seem related to a bi-directional transformation from both MHC isoforms I and IIX towards the MHC-IIA isoform. The aerobic performance and electromyographic variables at the whole-body level were not altered by ES. These results indicate that the particular short-term ES training protocol tested in the present study induces significant adaptations in histochemical and metabolic machineries of human skeletal muscle. The results also offer new perspectives for realistic applications of ES in various clinical situations and sport training. Keywords Anaerobic threshold • Electromyography (EMG) • Exercise physiology • Myosin heavy chain • Muscle fibre types • VO 2 max

Pfl�gers Archiv European Journal of Physiology, 2000

Twenty-four 4-year-old Andalusian (Spanish breed) stallions were used to examine the plasticity o... more Twenty-four 4-year-old Andalusian (Spanish breed) stallions were used to examine the plasticity of myosin heavy chain (MHC) phenotype and the metabolic profile in horse skeletal muscle with long-term endurance-exercise training and detraining. Sixteen horses underwent a training programme based on aerobic exercises for 8 months. Afterwards, they were kept in paddocks for 3 months. The remaining eight horses were used as controls. Three gluteus medius muscle biopsy samples were removed at depths of 20, 40 and 60 mm from each horse before (month 0), during (month 3) and after (month 8) training, and again after 3 months of detraining (month 11). MHC composition was analysed by electrophoresis and immunohistochemistry with anti-MHC monoclonal antibodies. Fibre areas, oxidative capacity and capillaries were studied histochemically. The activities of key muscle enzymes of aerobic (citrate synthase and 3-hydroxy-acyl-CoA-dehydrogenase) and anaerobic (phosphofructokinase and lactic dehydrogenase) metabolism and the intramuscular glycogen and triglyceride contents were also biochemically analysed. Early changes with training (3 months) included hypertrophy of type IIA fibres, a reduction of MHC-IIX with a concomitant increase of MHC-IIA, a rise in the number of high-oxidative fibres and in the activities of aerobic muscle enzymes and glycogen content. Long-term changes with training (8 months) were a further decline in the expression of MHC-IIX, an increase of slow MHC-I, additional increases of high-oxidative fibres, capillary density, activities of aerobic enzymes and endogenous glycogen; intramuscular lipid deposits also increased after 8 months of training whereas the activities of anaerobic enzymes declined. Most of exercise-induced alterations reverted after 3 months of detraining. These results indicate that endurance-exercise training induces a reversible transition of MHC composition in equine muscle in the order IIX-->IIA-->I, which is coordinated with changes in the metabolic properties of the muscle. Furthermore, a dose-response relationship was evident between the duration (in total) of training and the magnitude of muscle adaptations.

Journal of Anatomy, 2001

The distribution of muscle fibres classified on the basis of their content of different myosin he... more The distribution of muscle fibres classified on the basis of their content of different myosin heavy chain (MHC) isoforms was analysed in vastus lateralis muscle biopsies of 15 young men (with an average age of 22 y) by correlating immunohistochemistry with specific anti-MHC monoclonal antibodies, myofibrillar ATPase (mATPase) histochemistry and in situ hybridisation with probes specific for MHC β-slow, MHC-IIA and MHC-IIX. The characterisation of a large number of individual fibres was compared and correlated on a fibre-to-fibre basis. The panel of monoclonal antibodies used in the study allowed classification of human skeletal muscle fibres into 5 categories according to the MHC isoform they express at the protein level, types I, IjIIA, IIA, IIAX and IIX. Hybrid fibres coexpressing two isoforms represented a considerable proportion of the fibre composition (about 14 %) and were clearly underestimated by mATPase histochemistry. For a very high percentage of fibres there was a precise correspondence between the MHC protein isoforms and mRNA transcripts. The integrated methods used demonstrate a high degree of precision of the immunohistochemical procedure used for the identification and quantification of human skeletal muscle fibre types. The monoclonal antibody S5-8H2 is particularly useful for identifying hybrid IIAX fibres. This protocol offers new prospects for muscle fibre classification in human experimental studies.

Equine Veterinary Journal, 2008

When inadequate training stress is applied and recovery time is insufficient, performance reducti... more When inadequate training stress is applied and recovery time is insufficient, performance reduction and chronic maladaptation occurs. Known as overtraining syndrome (OTS), this complex condition afflicts horses in top training. The name of the syndrome implies causation and it is necessary to differentiate it from overreaching , a term used in horses that, after suffering a loss of performance without an obvious clinical reason, recover their performance within 1 or 2 weeks. The term OTS should be used for horses in heavy training losing performance without an obvious clinical reason and which is sustained for >2 weeks. Despite considerable scientific investigations, the underlying mechanisms are still poorly understood and there are no objective biomarkers for OTS. An imbalance between training and recovery seems to be the primary cause of this dysfunction. However, other factors such as transport, feeding, subclinical disease and general management may play a role. There is now sufficient evidence that red cell hypervolaemia is not a mechanism for the OTS in horses. A dysfunction of the hypothalamo-pituitaryadrenocortical axis is momentarily the only mechanism that may be related to the development of this syndrome. In the absence of a scoring system to assess the psychological status of horses, the most effective method for diagnosis is the thorough assessment of the history and presenting complaint (length of unexplained performance deficit, weight loss despite adequate feed intake, concurrence of unspecific subclinical problems, unsuccessful treatments and changes of behaviour). Standardised exercise tests are suggested to provide a way to detect subtle changes in hormonal responses in the individual, which may make an important contribution to the detection of early overtraining. But further longitudinal studies are needed to clarify mechanism(s) underlying OTS and to establish the potential for the use of a physiological test as a predictive tool of this disorder.

Equine Veterinary Journal, 2010

Dressage is one of the Olympic equestrian sports practiced in several countries using different h... more Dressage is one of the Olympic equestrian sports practiced in several countries using different horse breeds. Specific characteristics of the walk, trot and canter are required for dressage. It has been assumed that some of these traits could be selected for genetically and contribute to dressage performance. The purpose of this study was to compare the walk, trot and conformation characteristics in young horses of different breeds used for dressage. A total of 142 horses age 3 years were classified into 3 groups of breeds (German, French and Spanish saddle horses) and tested using the same procedure. The skeletal conformation measurements were made by image analysis. Gait variables of the walk and trot were measured by the accelerometric gait analysis system Equimetrix. Discriminant analysis could explain the variability between the groups by taking into account the walk (P<0.0003), trot (P<0.0001) and conformation variables (P<0.0001). Many gait and conformation variables were significantly different between the breeds. In summary, the German horses had gait characteristics more adapted for dressage competition, and the results of this group could be used as a reference for early evaluation in dressage. Purebred Spanish horses could be considered as a reference for collected gaits used for farm work and old academic dressage. The gait and conformation tests could be applied in a breeding or crossing plan to detect more accurately young horses with good dressage ability.

Equine Veterinary Journal, 2010

The purpose of this study was to determine whether oral L-carnitine supplementation enhances the ... more The purpose of this study was to determine whether oral L-carnitine supplementation enhances the responses of skeletal muscle to training in seven 2-year-old Standardbreds. Four horses were supplemented with 10 g/day L-carnitine for 10 weeks and 3 horses served as controls. All horses were exercised regularly every second day on a treadmill for 5 weeks (training period) and housed in individual boxes for 5 additional weeks (detraining period). The training period consisted of 8 high- and 8 low-speed exercises carried out in alternating sequence. Gluteus medius muscle biopsies were taken at Weeks 0 (pretraining), 5 (post-training) and 10 (detraining). Muscular adaptations to training were observed mainly in the L-carnitine-supplemented horses and included an increase in the percentage of type IIA fibres (delta35%, P<0.05), atrophy of type I fibres (delta24%, P<0.01), a rise in the capillary-to-fibre ratio (delta40%, P<0.01) and an increase in the quantitative reaction of periodic acid Schiff stain (delta11%, P<0.05), used as an indicator of intrafibre glycogen content. After detraining, most of these adaptations reverted towards the pretraining situation. Therefore, exogenous carnitine has an additive effect on muscular responses to training and this should be favourable to improve athletic performance. Nevertheless, further studies are necessary to show whether muscle carnitine content is a limiting factor for fatty acid oxidation.

Equine and Comparative Exercise Physiology, 2007

At the International Conference on Equine Exercise Physiology (ICEEP7), about 70 people attended ... more At the International Conference on Equine Exercise Physiology (ICEEP7), about 70 people attended the workshop on workload and conditioning guided by the authors. Most of the audience were involved in Thoroughbred or Standardbred racing, and only a limited number of people were mainly involved in FEI equestrian disciplines (sport horses). The workshop and this review article address the measurement of workload and conditioning of the Thoroughbred racehorse. It was proposed that workload could be quantified using a few selected parameters commonly recorded in the racing industries, such as velocity and distance, to generate a cumulative workload index. The review of conditioning focuses on the Thoroughbred racehorse and examines what can be modified with training, how training programmes should be designed based upon scientific methods and how training programmes should be routinely designed in current practice. It would appear that, in general, the methods used in practice for traini...

Journal of Veterinary Medicine Series A, 2007

The main goal of any conditioning programme in athletic horses is to improve performance by induc... more The main goal of any conditioning programme in athletic horses is to improve performance by inducing physiological changes within the animal's body. Equine skeletal muscles have a considerable potential to adapt during training and these adaptations have important physiological implications that influence stamina, strength and speed. Although there is an extensive specialized literature in this regard, scientific based muscle conditioning methods have not been introduced sufficiently in the equine sport practice. After a brief synopsis of both equine muscle exercise physiology and muscular adaptations to training, including their physiological significance, this review focuses on specific training programmes that induce muscular adaptations in athletic horses. The article addresses the following principal question: what kind of stimuli for what kind of muscular adaptations? The experimental data are discussed separately for racehorses (thoroughbreds, trotters and endurance horses) and sport horses (dressage, show jumpers and carriage). Finally, published results about the influence of relevant training parameters (such as intensity, duration and type of exercise) on muscular responses are discussed, as well as those concerning overtraining and detraining. The article closes with some concluding remarks on importance of their application in practice.

Veterinary Record, 2007

FIG 1: Longitudinal section of a sample of diaphragm from a horse with atypical myopathy (case 1)... more FIG 1: Longitudinal section of a sample of diaphragm from a horse with atypical myopathy (case 1). Multifocal and segmental necrosis and Zenker’s degeneration (arrow) in multiple myofibres are visible. Haematoxylin and eosin. Bar=75 μm FIG 2: Transverse section of the gluteus profundus muscle of a horse with atypical myopathy (case 2). Increased intercellular spaces (oedema) and multiple hypercontracted muscle fibres with pale pink cytoplasm (arrows) and severe degeneration of the myofibrillar apparatus are visible. Haematoxylin and eosin. Bar=80 μm Veterinary Record (2007) 161, 346-348

Pflügers Archiv - European Journal of Physiology, 2002

Fifteen healthy, untrained male subjects (mean age ±SD, 22±5 years) were used to examine the plas... more Fifteen healthy, untrained male subjects (mean age ±SD, 22±5 years) were used to examine the plasticity of myosin heavy chain phenotype, size, oxidative capacity and capillarization of skeletal muscle fibre types with short-term electrical stimulation (ES). Ten subjects were electro-stimulated on both quadriceps muscles with a frequency of 45-60 Hz, with 12 s of stimulation followed by 8 s of recovery for a total of 30 min per day, 3 days per week for 6 weeks. The remaining five subjects served as controls. Two vastus lateralis muscle biopsy samples were removed from each subject before (week 0) and after (week 6) ES training. A standardized exercise test on a cycle ergometer was performed by each subject before and after the experimental period and several indicators of whole-body aerobic capacity were estimated. The so-called electromyographic threshold was also determined during the tests. Muscle biopsy samples were analysed by electrophoresis, immunohistochemistry and quantitative histochemistry. Myosin heavy chain (MHC) composition, muscle fibre type distribution, fibre areas, oxidative capacity and capillaries of each fibre type were estimated. Muscular changes with ES revealed an increase of fibres expressing MHC-IIA, and a decrease of fibres expressing MHC-IIX and MHC-I, as well as an increase of the oxidative capacity and mean number of capillaries of fast-twitch (type II) fibres with minimal muscle fibre hypertrophy. These adaptations seem related to a bi-directional transformation from both MHC isoforms I and IIX towards the MHC-IIA isoform. The aerobic performance and electromyographic variables at the whole-body level were not altered by ES. These results indicate that the particular short-term ES training protocol tested in the present study induces significant adaptations in histochemical and metabolic machineries of human skeletal muscle. The results also offer new perspectives for realistic applications of ES in various clinical situations and sport training. Keywords Anaerobic threshold • Electromyography (EMG) • Exercise physiology • Myosin heavy chain • Muscle fibre types • VO 2 max

Pfl�gers Archiv European Journal of Physiology, 2000

Twenty-four 4-year-old Andalusian (Spanish breed) stallions were used to examine the plasticity o... more Twenty-four 4-year-old Andalusian (Spanish breed) stallions were used to examine the plasticity of myosin heavy chain (MHC) phenotype and the metabolic profile in horse skeletal muscle with long-term endurance-exercise training and detraining. Sixteen horses underwent a training programme based on aerobic exercises for 8 months. Afterwards, they were kept in paddocks for 3 months. The remaining eight horses were used as controls. Three gluteus medius muscle biopsy samples were removed at depths of 20, 40 and 60 mm from each horse before (month 0), during (month 3) and after (month 8) training, and again after 3 months of detraining (month 11). MHC composition was analysed by electrophoresis and immunohistochemistry with anti-MHC monoclonal antibodies. Fibre areas, oxidative capacity and capillaries were studied histochemically. The activities of key muscle enzymes of aerobic (citrate synthase and 3-hydroxy-acyl-CoA-dehydrogenase) and anaerobic (phosphofructokinase and lactic dehydrogenase) metabolism and the intramuscular glycogen and triglyceride contents were also biochemically analysed. Early changes with training (3 months) included hypertrophy of type IIA fibres, a reduction of MHC-IIX with a concomitant increase of MHC-IIA, a rise in the number of high-oxidative fibres and in the activities of aerobic muscle enzymes and glycogen content. Long-term changes with training (8 months) were a further decline in the expression of MHC-IIX, an increase of slow MHC-I, additional increases of high-oxidative fibres, capillary density, activities of aerobic enzymes and endogenous glycogen; intramuscular lipid deposits also increased after 8 months of training whereas the activities of anaerobic enzymes declined. Most of exercise-induced alterations reverted after 3 months of detraining. These results indicate that endurance-exercise training induces a reversible transition of MHC composition in equine muscle in the order IIX-->IIA-->I, which is coordinated with changes in the metabolic properties of the muscle. Furthermore, a dose-response relationship was evident between the duration (in total) of training and the magnitude of muscle adaptations.

Journal of Anatomy, 2001

The distribution of muscle fibres classified on the basis of their content of different myosin he... more The distribution of muscle fibres classified on the basis of their content of different myosin heavy chain (MHC) isoforms was analysed in vastus lateralis muscle biopsies of 15 young men (with an average age of 22 y) by correlating immunohistochemistry with specific anti-MHC monoclonal antibodies, myofibrillar ATPase (mATPase) histochemistry and in situ hybridisation with probes specific for MHC β-slow, MHC-IIA and MHC-IIX. The characterisation of a large number of individual fibres was compared and correlated on a fibre-to-fibre basis. The panel of monoclonal antibodies used in the study allowed classification of human skeletal muscle fibres into 5 categories according to the MHC isoform they express at the protein level, types I, IjIIA, IIA, IIAX and IIX. Hybrid fibres coexpressing two isoforms represented a considerable proportion of the fibre composition (about 14 %) and were clearly underestimated by mATPase histochemistry. For a very high percentage of fibres there was a precise correspondence between the MHC protein isoforms and mRNA transcripts. The integrated methods used demonstrate a high degree of precision of the immunohistochemical procedure used for the identification and quantification of human skeletal muscle fibre types. The monoclonal antibody S5-8H2 is particularly useful for identifying hybrid IIAX fibres. This protocol offers new prospects for muscle fibre classification in human experimental studies.

Equine Veterinary Journal, 2008

When inadequate training stress is applied and recovery time is insufficient, performance reducti... more When inadequate training stress is applied and recovery time is insufficient, performance reduction and chronic maladaptation occurs. Known as overtraining syndrome (OTS), this complex condition afflicts horses in top training. The name of the syndrome implies causation and it is necessary to differentiate it from overreaching , a term used in horses that, after suffering a loss of performance without an obvious clinical reason, recover their performance within 1 or 2 weeks. The term OTS should be used for horses in heavy training losing performance without an obvious clinical reason and which is sustained for >2 weeks. Despite considerable scientific investigations, the underlying mechanisms are still poorly understood and there are no objective biomarkers for OTS. An imbalance between training and recovery seems to be the primary cause of this dysfunction. However, other factors such as transport, feeding, subclinical disease and general management may play a role. There is now sufficient evidence that red cell hypervolaemia is not a mechanism for the OTS in horses. A dysfunction of the hypothalamo-pituitaryadrenocortical axis is momentarily the only mechanism that may be related to the development of this syndrome. In the absence of a scoring system to assess the psychological status of horses, the most effective method for diagnosis is the thorough assessment of the history and presenting complaint (length of unexplained performance deficit, weight loss despite adequate feed intake, concurrence of unspecific subclinical problems, unsuccessful treatments and changes of behaviour). Standardised exercise tests are suggested to provide a way to detect subtle changes in hormonal responses in the individual, which may make an important contribution to the detection of early overtraining. But further longitudinal studies are needed to clarify mechanism(s) underlying OTS and to establish the potential for the use of a physiological test as a predictive tool of this disorder.

Equine Veterinary Journal, 2010

Dressage is one of the Olympic equestrian sports practiced in several countries using different h... more Dressage is one of the Olympic equestrian sports practiced in several countries using different horse breeds. Specific characteristics of the walk, trot and canter are required for dressage. It has been assumed that some of these traits could be selected for genetically and contribute to dressage performance. The purpose of this study was to compare the walk, trot and conformation characteristics in young horses of different breeds used for dressage. A total of 142 horses age 3 years were classified into 3 groups of breeds (German, French and Spanish saddle horses) and tested using the same procedure. The skeletal conformation measurements were made by image analysis. Gait variables of the walk and trot were measured by the accelerometric gait analysis system Equimetrix. Discriminant analysis could explain the variability between the groups by taking into account the walk (P<0.0003), trot (P<0.0001) and conformation variables (P<0.0001). Many gait and conformation variables were significantly different between the breeds. In summary, the German horses had gait characteristics more adapted for dressage competition, and the results of this group could be used as a reference for early evaluation in dressage. Purebred Spanish horses could be considered as a reference for collected gaits used for farm work and old academic dressage. The gait and conformation tests could be applied in a breeding or crossing plan to detect more accurately young horses with good dressage ability.

Equine Veterinary Journal, 2010

The purpose of this study was to determine whether oral L-carnitine supplementation enhances the ... more The purpose of this study was to determine whether oral L-carnitine supplementation enhances the responses of skeletal muscle to training in seven 2-year-old Standardbreds. Four horses were supplemented with 10 g/day L-carnitine for 10 weeks and 3 horses served as controls. All horses were exercised regularly every second day on a treadmill for 5 weeks (training period) and housed in individual boxes for 5 additional weeks (detraining period). The training period consisted of 8 high- and 8 low-speed exercises carried out in alternating sequence. Gluteus medius muscle biopsies were taken at Weeks 0 (pretraining), 5 (post-training) and 10 (detraining). Muscular adaptations to training were observed mainly in the L-carnitine-supplemented horses and included an increase in the percentage of type IIA fibres (delta35%, P<0.05), atrophy of type I fibres (delta24%, P<0.01), a rise in the capillary-to-fibre ratio (delta40%, P<0.01) and an increase in the quantitative reaction of periodic acid Schiff stain (delta11%, P<0.05), used as an indicator of intrafibre glycogen content. After detraining, most of these adaptations reverted towards the pretraining situation. Therefore, exogenous carnitine has an additive effect on muscular responses to training and this should be favourable to improve athletic performance. Nevertheless, further studies are necessary to show whether muscle carnitine content is a limiting factor for fatty acid oxidation.

Equine and Comparative Exercise Physiology, 2007

At the International Conference on Equine Exercise Physiology (ICEEP7), about 70 people attended ... more At the International Conference on Equine Exercise Physiology (ICEEP7), about 70 people attended the workshop on workload and conditioning guided by the authors. Most of the audience were involved in Thoroughbred or Standardbred racing, and only a limited number of people were mainly involved in FEI equestrian disciplines (sport horses). The workshop and this review article address the measurement of workload and conditioning of the Thoroughbred racehorse. It was proposed that workload could be quantified using a few selected parameters commonly recorded in the racing industries, such as velocity and distance, to generate a cumulative workload index. The review of conditioning focuses on the Thoroughbred racehorse and examines what can be modified with training, how training programmes should be designed based upon scientific methods and how training programmes should be routinely designed in current practice. It would appear that, in general, the methods used in practice for traini...