ࡱ> C F . / 0 1 2 3 4 5 6 7 8 9 : ; < = > ? @ A B  Tbjbjޤ Afƴƴo'| | {{{8l3D0  "!!!d)z) )CCCCCCCEH\C{,'d),,C!!gDb<b<b<, !{!Cb<,Cb<b<VA@E6A!Ь;5XA pC}D0DdATH8HAH{A) *hb<*T*,)))CCf:)))D,,,,H)))))))))| :   JEPonline  Swimming Physical Training in Rats: Cardiovascular Adaptation to Exercise Training Protocols at Different Intensities L.F. Guerreiro1, A.A. Pereira 1, C.N. Martins 1, C. Wally 1, C.A.N. Gonalves1 1Institute of Biological Sciences / Post - Graduate in Physiological Sciences: Comparative Animal Physiology / Federal University of Rio Grande - FURG, Rio Grande, RS, Brazil  ABSTRACT Guerreiro LF, Pereira AA, Martins CN, Wally C, Gonalves CAN. Swimming Physical Training in Rats: Cardiovascular Adaptation to Exercise Training Protocols at Different Intensities. JEPonline 2015; 18(1):1-12. There have been many studies of physical exercise in laboratory animal models. However, these studies are limited by their experimental protocols. The aim of the present study was to develop physical training protocols of low, moderate, and high intensity and to compare the cardiovascular acclimation of Wistar rats to these protocols. Forty male rats 60 days old at the beginning of the experiment and weighing 310 10 g were divided into four groups (n=10): control, low intensity (LI), moderate intensity (MI), and intermittent high intensity (HI). The training was divided into an acclimation phase and a maintenance phase and took place 5 times a week over 16 wks. After the training, the animals were euthanized. Creatine kinase and heart morphometry analyses were conducted. Lactate values were 4.0 0.5 mmolL-1 for LI, 5.5 0.4 mmolL-1 for MI, and 9.2 1.0 mmolL-1 for HI. Heart rate was significantly reduced in the LI (339 20 beatsmin-1) and MI (344 20 beatsmin-1). Morphometric analyses showed that the thickness of the free wall of the left ventricle was greater in the HI group (2.233 0.29 mm) (Pd"0.05). The thickness of the interventricular septum was greater in the HI group (2.158 0.71 mm) and the area of the left ventricle chamber significantly greater in the MI group (25.657 3.6 mm2) (Pd"0.05). Our results show that these protocols induce different metabolic pathways and promote distinct adaptations in the cardiovascular system to the type of physical training experienced. Key Words: Exercise, Cardiovascular, Swimming Intensity INTRODUCTION Studies on exercise physiology use animal models to simulate conditions of physical stress commonly observed in humans, with the aim of better understanding changes that occur in humans at the systemic, cellular, and molecular levels  ADDIN EN.CITE  ADDIN EN.CITE.DATA (3,5,27). Many studies of physical exercise in laboratory animal models have been conducted  ADDIN EN.CITE  ADDIN EN.CITE.DATA (14,15,20,22,28,30,31). However, these studies are limited by their experimental protocols. Creating physical exercise interventions in experimental physiology is not that simple because some factors, such as training duration, frequency, session duration, exercise intensity, and metabolism used during the physical activity, cannot be controlled  ADDIN EN.CITE Booth201013091309130917Booth, F. W.Laye, M. J.Spangenburg, E. E.Veterinary Medicine, Bldg E102, Univ. of Missouri, Columbia, MO 65211, USA. boothf@missouri.eduGold standards for scientists who are conducting animal-based exercise studiesJ Appl Physiol J Appl Physiol219-2110812009/07/04AnimalsExercise Test/classification/*standards/*veterinaryPhysical Conditioning, Animal/classification/*standardsPhysiology/*standardsReference StandardsScience/*standards2010Jan1522-1601 (Electronic) 0161-7567 (Linking)19574508http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1957450800125.2009 [pii] 10.1152/japplphysiol.00125.2009eng(5). These factors make it difficult to simulate physical activity experienced by humans in experimental models  ADDIN EN.CITE  ADDIN EN.CITE.DATA (12). The transposition of experimental data and physical training protocols to experimental physiology is frequently inadequate, leading to misinterpretations in data before publication  ADDIN EN.CITE Booth201013091309130917Booth, F. W.Laye, M. J.Spangenburg, E. E.Veterinary Medicine, Bldg E102, Univ. of Missouri, Columbia, MO 65211, USA. boothf@missouri.eduGold standards for scientists who are conducting animal-based exercise studiesJ Appl Physiol J Appl Physiol219-2110812009/07/04AnimalsExercise Test/classification/*standards/*veterinaryPhysical Conditioning, Animal/classification/*standardsPhysiology/*standardsReference StandardsScience/*standards2010Jan1522-1601 (Electronic) 0161-7567 (Linking)19574508http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1957450800125.2009 [pii] 10.1152/japplphysiol.00125.2009eng(5). Therefore, it is necessary that protocols for physical exercise in animals properly simulate the situations under investigation  ADDIN EN.CITE Gondim200712681268126817Gondim, F. J.Zoppi, C. C.Pereira-da-Silva, L.de Macedo, D. V.Laboratorio de Imunopatologia Experimental, Universidade Federal de Goias, Goiania, Brasil.Determination of the anaerobic threshold and maximal lactate steady state speed in equines using the lactate minimum speed protocolComp Biochem Physiol A Mol Integr PhysiolComp Biochem Physiol A Mol Integr Physiol375-8014632007/01/20Anaerobic Threshold/*physiologyAnimalsHeart RateHorses/*physiologyLactic Acid/*bloodMalePhysical Conditioning, Animal/*physiologyRunning2007Mar1095-6433 (Print) 1095-6433 (Linking)17234441http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17234441S1095-6433(06)00491-0 [pii] 10.1016/j.cbpa.2006.11.002eng(18). Swimming protocols are frequently used with rats, mainly because rats have an innate swimming ability and acclimate well to training, which can be done for little cost compared to treadmill running  ADDIN EN.CITE Gobatto200112781278127817Gobatto, C. A.de Mello, M. A.Sibuya, C. Y.de Azevedo, J. R.dos Santos, L. A.Kokubun, E.Department of Physical Education, Bioscience Institute, University of Sao Paulo State (UNESP), Avenida 24-A, 1515 Bela Vista, Rio Claro, Sao Paulo, 13506-900 Brazil. cgobatto@uol.com.brMaximal lactate steady state in rats submitted to swimming exerciseComp Biochem Physiol A Mol Integr PhysiolComp Biochem Physiol A Mol Integr Physiol21-713012001/10/24Anaerobic Threshold/*physiologyAnimalsLactic Acid/*bloodMalePhysical Conditioning, Animal/physiologyPhysical Endurance/physiologyRatsRats, WistarSwimming/*physiology2001Aug1095-6433 (Print) 1095-6433 (Linking)11672680http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11672680S1095-6433(01)00362-2 [pii]eng(17). Studies of swimming in rats have shown acclimations similar to those observed in humans  ADDIN EN.CITE  ADDIN EN.CITE.DATA (17,29). Studies using animal models should use physical exercise protocols that simulate situations to which humans are exposed daily, such as continuous aerobic exercise of low and moderate intensity, in addition to intermittent exercise of high intensity as observed in team sports. The intensity of the exercise can differentiate acclimation to training by determining the metabolic pathway, the energetic substrate and the energy source used, as well as the tension overload on the cardiovascular system  ADDIN EN.CITE Hernandez-Torres200929292917Hernandez-Torres, R. P.Ramos-Jimenez, A.Torres-Duran, P. V.Romero-Gonzalez, J.Mascher, D.Posadas-Romero, C.Juarez-Oropeza, M. A.Faculty of Physical Education and Sport Sciences, Autonomous University of Chihuahua (UACH), Mexico. rhernant@uach.mxEffects of single sessions of low-intensity continuous and moderate-intensity intermittent exercise on blood lipids in the same endurance runnersJ Sci Med SportJ Sci Med Sport323-311222008/04/09AdolescentAdultAnaerobic Threshold/*physiologyAthletic Performance/physiologyCholesterol, HDL/*bloodCholesterol, LDL/*bloodExercise/*physiologyHumansMaleRunning/*physiologyTriglycerides/*bloodYoung Adult2009Mar1440-2440 (Print)18396101http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=18396101S1440-2440(07)00292-7 [pii] 10.1016/j.jsams.2007.12.002eng(19). Studies of aerobic and anaerobic metabolism have been developed to improve physical training in rats  ADDIN EN.CITE  ADDIN EN.CITE.DATA (6,10,17,23,25). The intensity of exercise can be controlled by the production, removal, and accumulation of lactate in the bloodstream. The maximal lactate steady state (MLSS) is the highest concentration of lactate and workload that can be maintained over time at the anaerobic threshold  ADDIN EN.CITE  ADDIN EN.CITE.DATA (12,16,23). MLSS intensity can be determined during continuous exercise, considering an exercise intensity equivalent to a concentration of 4 mmolL-1 of lactate in humans  ADDIN EN.CITE Beneke201123012301230117Beneke, R.Leithauser, R. M.Ochentel, O.Centre for Sport and Exercise Science, Department of Biological Sciences, University of Essex, Essex, England.Blood lactate diagnostics in exercise testing and trainingInt J Sports Physiol PerformInt J Sports Physiol Perform8-24612011/04/14Biological Markers/bloodExercise/*physiology*Exercise TestHumansLactic Acid/*blood*Muscle ContractionMuscle, Skeletal/*metabolismPhysical EndurancePredictive Value of TestsTime Factors2011Mar1555-0265 (Print) 1555-0265 (Linking)21487146http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=21487146eng(4). Lactate concentrations at MLSS have been studied in rats and values of 5.5 nmolL-1 observed  ADDIN EN.CITE  ADDIN EN.CITE.DATA (6,17,23). Blood lactate levels higher than MLSS during physical exercise indicate that anaerobic metabolism predominates over aerobic metabolism  ADDIN EN.CITE  ADDIN EN.CITE.DATA (17,23). We hypothesized that the intensity of exercise can promote different adaptations in the cardiovascular system. The aim of the present study was to develop protocols for long-term physical training (swimming) in Wistar rats (Rattus norvegicus) and to compare cardiovascular acclimation to this training below (low intensity), at (moderate intensity), and above (intermittent high intensity) MLSS intensity. METHODS Subjects All experiments were conducted in compliance with the Ethics Commission of Animal Use (Comisso de tica em Uso Animal CEUA-FURG), process number 052/2011 and Guidelines on Animal Ethics and Welfare for Veterinary Journals. Forty male Wistar rats were used. The rats were 60 days old at the beginning of the experiment and weighed 200 to 300 g. The animals were kept in collective cages (5 animals/cage), fed a pelleted diet for rodents (15 g/rat/day) and given fresh water ad libitum, and maintained under a 12-hr light:dark photoperiod (lights on at 07:00) at a temperature of 22 2C. Experimental groups: Below MLSS intensity (Low Intensity [LI]): This low-intensity swimming protocol took place in the individual swimming system (Figure 1) without load, for 60 min, 5 timeswk-1, for 8 wks. The purpose of this methodology was to simulate exercise without controlling intensity by overload, thus allowing the animals to exercise daily at an intensity below MLSS. This protocol aimed to simulate common human physical activity, such as walking or running at low intensity, with controlled frequency and duration.  Figure 1. Swimming Tank. (A) Swimming tank for control, low, and moderate intensity protocols. (B) Swimming tank with pulled system for high intensity protocol. Internal cylinder submerged to intermittent swim, and pulled out to recovery, each movement lasting 15 sec (swim/recovery). MLSS Intensity (Moderate Intensity [MI]) This swimming protocol, which also took place in the individual swimming system (Figure 1), comprised 60 min of swimming, 5 timeswk-1 with a workload equivalent to 5% of the animals body weight. The workload was provided by pieces of lead attached to the back of the animal and was equivalent to a moderate MLSS intensity. Above MLSS Intensity (Intermittent High Intensity [HI]) The rats in this group were submitted to intermittent individual swimming up to 60 min, 5 timeswk-1 for 16 wks. Each 60-min session alternated periods of vigorous swimming for 15 sec with a workload equivalent to 15% of the animals body weight at intensities above MLSS followed by recovery for 15 sec. The entire session comprised 30 min of intermittent vigorous swimming and 30 min of rest. No-Training Control (C) The control animals were placed in a liquid environment to simulate the stress under experimental conditions for 1 min, 5 timeswk-1 for 16 wks. Procedures Training Tanks The physical training period lasted for 16 wks. The animals exercised 5 timeswk-1 between 19:00 and 22:00, respecting the animals natural active period. The continuous swimming system used in the LI and MI conditions consisted of a rectangular tank (90 cm width 100 cm length 80 cm height) filled with water up to 50 cm. Ten polyvinyl chloride (PVC) pipes 20 cm wide and 50 cm long were placed inside the tank. The water temperature was kept close to 32C. The animals were able to exercise individually, each in a different PVC pipe. The intermittent swimming system used in the HI condition consisted of a 60-cm wide cylinder covered at the bottom with mesh-galvanized aluminum and containing 10 PVC pipes. The cylinder, which was on pulleys, was immersed in the tank described previously for 15 sec, forcing the animals to swim vigorously. Then the cylinder was lifted by the pulleys, allowing the animals to recover for 15 sec outside of the water (Figure 1). Training Protocol The 16-wk swimming program to which the animals submitted comprised two phases: progressive acclimation and maintenance (Figure 2A and 2B). In the first 8 wks, the rats acclimated to various intensities of exercise. In Week 1, the animals were progressively acclimated to the liquid environment for 20 min, after which the training load and duration of swimming increased progressively (from 20, 25, 30, and 40 min in Week 1 to 40, 45, 50, 55, and 60 min in Week 2). Starting in Week 2 swimming loads were applied (1% for MI and 6% for HI, increasing progressively to a maximum of 5% [MI] and 15% [HI] in Week 8). This phase lasted for 8 wks.  Figure 2. Training Program Protocol. This Protocol last 16-wk and is divide into two periods: 1. Progressive Acclimation (8 wks) and 2. Load Maintenance (8 wks) (n=10/group). (A) Progressive Acclimation and Maintenance to the Workload (% of body weight BW). (B) Progressive Acclimation and Maintenance to the Total Volume of Exercise (Time in minutes). LI: low-intensity swimming, without workload. MI: moderate-intensity swimming with load equal to 5% BW. HI: high-intensity swimming with load equal to 15% BW.  After this initial acclimation phase, once the animals were able to swim for 60 min, the animals were acclimated to different intensities of swimming (Figure 2A, 2B). This phase was characterized by the maintenance of higher loads and longer duration of exercise (60 min and a workload of 5% [MI] or 15% [HI]) for 8 more weeks. The load was adjusted weekly according to the animals body weight. Determination of Lactate Levels Lactate levels was determined by analysis during exercise. Animals in all groups were removed from the water 10, 20, 30, and 40 min into the exercise session and blood samples (25 l) collected according to Voltarelli et al.  ADDIN EN.CITE <EndNote><Cite><Author>Voltarelli</Author><Year>2002</Year><RecNum>32</RecNum><record><rec-number>32</rec-number><foreign-keys><key app="EN" db-id="pdasws25gf0rp8eaetr5xte79fte9e020p9t">3217Voltarelli, F. A.Gobatto, C. A.de Mello, M. A. R.de Mello, MAR UNESP, Dept Educ Fis 1B, Inst Biol, Ave 24-A 1515, BR-13506900 Rio Claro, SP, Brazil UNESP, Dept Educ Fis 1B, Inst Biol, BR-13506900 Rio Claro, SP, BrazilDetermination of anaerobic threshold in rats using the lactate minimum testBrazilian Journal of Medical and Biological ResearchBrazilian Journal of Medical and Biological Research1389-13943511ratblood lactatelactate minimum testmaximal lactate steady statesteady-stateswimming exercisetest protocolperformanceequilibriumspeed2002Nov0100-879XISI:000179717100018<Go to ISI>://000179717100018English(29). The lactate concentration was determined in mmolL-1 with a lactate analyzer (Accutrend). Determination of Muscle Damage Creatine kinase levels as determined using a Labtest Diagnstica SA specific kit were used as markers of muscle damage. The dosage was defined based on the enzymatic system with final point reaction in serum samples from the animals by spectrophotometry expressed in U/L  ADDIN EN.CITE Chapman200623072307230717Chapman, D.Newton, M.Sacco, P.Nosaka, K.School of Exercise, Biomedical and Health Sciences, Edith Cowan University, Joondalup, Western Australia.Greater muscle damage induced by fast versus slow velocity eccentric exerciseInt J Sports MedInt J Sports Med591-82782006/07/29AdultArm/anatomy & histologyCreatine Kinase/bloodElbow Joint/physiologyExercise TestFemaleHumansMaleMuscle Contraction/*physiologyMuscle Fatigue/*physiologyMuscle, Skeletal/physiologyRange of Motion, Articular/physiologyRecovery of Function/physiologyTorque2006Aug0172-4622 (Print) 0172-4622 (Linking)16874584http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1687458410.1055/s-2005-865920eng(8) . Blood Pressure and Heart Rate Measurements Blood pressure and heart rate were measured using a cuff-tail plethysmograph (Leitica 5001). The rats were contained in a properly heated system at 30C to facilitate caudal artery vasodilation, from which systolic and diastolic arterial blood pressure and heart rate were measured based on arterial pulse pressure. Measurements were taken once every 30 days for 16 wks. Heart Morphometric Analysis The animals were euthanized by decapitation and their hearts removed by the end of the study. Hearts were weighed and processed histologically by hematoxylin and eosin staining. Afterward, the slides were photographed using a digital camera coupled with a stereo microscope. The thickness of the free wall of the left ventricle (mm), the thickness of the interventricular septum (mm), and the total area of the left ventricle chamber (mm2) were measured using the software Image J. Statistical Analyses The data are expressed as means standard deviations (M SD).Tests for the normality of the data distribution (KolmogorovSmirnoff test) and homoscedasticity of variances (Levenes test) were performed to verify the assumptions of the analysis of variance. Parametric data were analyzed using analysis of variance followed by Tukeys post hoc test, and nonparametric data were analyzed using the KruskalWallis test. The statistical significance was set at Pd"0.05. RESULTS The swimming training lasted 16 wks, and in that time, no stress behavior (e.g., cannibalism, aggressiveness, or mortality) was observed. The MI and HI groups at maximum load (5% and 15%, respectively) reached the maximum volume of swimming (60 min) at the end of the progressive acclimation period (Week 8) and sustained that load and volume until the end of the maintenance period (Week 9; Figure 3). The C animals had a lactate concentration of 2.5 0.4 mmolL-1 during the MLSS test. The LI group had a concentration of 4.0 0.5 mmolL-1, which was not significantly different from C. The MI group had a blood lactate concentration of 5.5 mmolL-1, which was significantly higher than C (Pd"0.05). The HI group had a blood lactate concentration of 9.2 1.0 mmolL-1, which was significantly higher than all other groups (Pd"0.05; Figure 3).  Figure 3. Lactate Levels after the Progressive Acclimation Period (Weeks 9) at 10, 20, 30, and 40 min of Exercise. C: control. LI: low-intensity swimming, without workload. MI: moderate-intensity swimming, 5%. HI: high-intensity swimming, 15%. Data are means SD (mgdL-1). Different letters indicate significant differences between groups at the 5% significance level. The results for muscle damage were 320 20 U/L for C, which was not different from LI (340 69 U/L) or HI (380 59 U/L). However, the MI group had a significant reduction in damage of 15% (250 15 U/L) compared with the other groups (Pd"0.05; Figure 4).  Figure 4. Creatine Kinase Values (U/L) at the End of the Load Maintenance Period (16 wks). C: control. LI: low-intensity swimming, without workload. MI: moderate-intensity swimming, 5%. HI: high-intensity swimming, 15%. Data are means SD (mgdL-1). Different letters indicate significant differences between groups at the 5% significance level. The results for cardiac morphometry showed that the thickness of the left ventricle free wall was greater (Pd"0.05) in HI (2.233 0.29 mm) compared with C (1.992 0.29 mm) and LI (2.433 mm) (Figure 5A). The thickness of the interventricular septum was greater in HI (2.158 0.71 mm) than in all other groups (P<0.05) (Figure 5B). Finally, the left ventricle chamber was significantly larger in MI (25.657 3.6 mm2) than in C (19.551 3.8 mm2), and HI (14.858 3.7 mm2) had a significantly lower value than the other groups (Pd"0.05; Figure 5C).    Figure 5. Heart Morphometry. (A) Left ventricle free wall thickness (mm). (B) Interventricular septum thickness (mm). (C) Left ventricle chamber area (mm2). C: control. LI: low-intensity swimming, without workload. MI: moderate-intensity swimming, 5%. HI: high-intensity swimming, 15%. Data are means SD (mgdL-1). Different letters indicate significant differences between groups at the 5% significance level. Heart rate was significantly reduced in the LI (344 20 beatsmin-1) and MI (350 20 beatsmin-1) groups compared with C (420 25 beatsmin-1) and HI (378 23 beatsmin-1; Pd"0.05). Systolic and diastolic blood pressure values were not altered by any training protocol (Table 1). Groups  Heart rate (beatsmin-1) Systolic Pressure (mmHg) Diastolic Pressure (mmHg) InitialFinalInitialFinalInitialFinalControl365 26392 22a136 31158 29107 22 105 20Low Intensity 360 30339 15b155 32165 24118 21115 22Moderate Intensity 368 18344 19b137 23150 23115 16110 24High Intensity358 23378 21a162 26172 31115 13120 23Table 1. Heart Rate, Systolic and Diastolic Blood Pressure. Different letters a,b indicate significant differences between treatment groups at the 5% significance level. DISCUSSION The swimming training protocol investigated here was effective at developing cardiovascular alterations in response to physical exercise. The LI group had lactate levels below MLSS, supporting a low-intensity protocol. However, the rats in the MI group, which bore a workload equivalent to 5% of their body weight, had blood lactate levels (5.5 mmolL-1) similar to MLSS  ADDIN EN.CITE  ADDIN EN.CITE.DATA (17,29). These latter results are similar to those reported by Voltarelli et al.  ADDIN EN.CITE Voltarelli200232323217Voltarelli, F. A.Gobatto, C. A.de Mello, M. A. R.de Mello, MAR UNESP, Dept Educ Fis 1B, Inst Biol, Ave 24-A 1515, BR-13506900 Rio Claro, SP, Brazil UNESP, Dept Educ Fis 1B, Inst Biol, BR-13506900 Rio Claro, SP, BrazilDetermination of anaerobic threshold in rats using the lactate minimum testBrazilian Journal of Medical and Biological ResearchBrazilian Journal of Medical and Biological Research1389-13943511ratblood lactatelactate minimum testmaximal lactate steady statesteady-stateswimming exercisetest protocolperformanceequilibriumspeed2002Nov0100-879XISI:000179717100018<Go to ISI>://000179717100018English(29) using subthreshold workloads (4% of body weight) and threshold equivalent loads (5% of body weight). Swimming training protocols with intensities above MLSS and using loads equivalent to more than 6.5% of body weight resulted in an increase in blood lactate production by means of the anaerobic glycolytic pathway. Such protocols are exhaustive and enable animals to perform only a few minutes of physical exercise. Because of this, experimental protocols of high-intensity intermittent training that biochemically simulates team sports are rare. Rats in our high-intensity (HI) group bore a maximum load of 15% of their body weight, resulting in an increase in blood lactate up to 11 mmolL-1. This was efficient at promoting intense predominantly anaerobic exercise above MLSS during a training volume of 60 min, which is equivalent to the practice of team sports  ADDIN EN.CITE  ADDIN EN.CITE.DATA (4,7,11). The results of the present study show that aerobic exercise at intensities below MLSS (LI) and at MLSS (MI) reduces resting heart rate after 16 wks. Such bradycardia may be due to a possible increase in sympathovagal balance after continuous training, as confirmed by the classical literature on sport science  ADDIN EN.CITE  ADDIN EN.CITE.DATA (24,26,27). According to the data presented here, the training protocols did not have any additional effects on systolic or diastolic pressure (which is understandable, as the rats used in the present study were normotensive), confirming the efficiency of acclimation to progressively greater loads. Cardiac remodeling is a physiological condition that is a morphological consequence of systematic physical training involving an increase in maximum cardiac production, increase in systolic volume, and decrease in resting heart rate  ADDIN EN.CITE  ADDIN EN.CITE.DATA (9). Volume overload and an increase in filling pressures result in an increase in elongation strength on the myocardium  ADDIN EN.CITE Anversa199114981498149817Anversa, P.Capasso, J. M.Cardiac hypertrophy and ventricular remodelingLab InvestLab Invest441-56441991/04/01AnimalsCardiomegaly/*pathologyDisease Models, AnimalHeart Ventricles/pathologyHumansMiceMice, TransgenicMyocardium/*pathology1991Apr0023-6837 (Print) 0023-6837 (Linking)1826750http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=1826750eng(1). The present training protocols resulted in two very distinct types of cardiac remodeling. The continuous aerobic swimming of moderate intensity (MI protocol) increased the left ventricle chamber area, promoting eccentric hypertrophy. Eccentric hypertrophy is an elongation of the cardiac muscle fibers of the left ventricle chamber caused by increased blood flow due to aerobic exercise, which increases blood demand in the left ventricle hemodynamics  ADDIN EN.CITE  ADDIN EN.CITE.DATA (13). Similar to the results of the present study, Evangelista et al.  ADDIN EN.CITE  ADDIN EN.CITE.DATA (15) found an increase in the cardiac chambers using a moderate protocol of two daily sessions over 4 wks and without overload. In contrast, the intermittent high-intensity swimming (HI protocol) resulted in an increase in the thickness of the left ventricle wall and interventricular septum, leading to concentric hypertrophy. Because of the post-load increase, the highest intarventricular pressure during the ejection phase in high-intensity exercise can increase stress on the myocardium wall  ADDIN EN.CITE  ADDIN EN.CITE.DATA (1,13,21), with a consequent increase in wall thickness, characterizing concentric hypertrophy  ADDIN EN.CITE  ADDIN EN.CITE.DATA (2). This cardiovascular adaptation is similar to that observed in our HI group and is determined by the elevated MLSS values observed here. In line with the present results, Kemi et al.  ADDIN EN.CITE  ADDIN EN.CITE.DATA (22) investigated high-intensity training on a treadmill for 2 hrs a day over 8 wks and reported that animals developed concentric cardiac hypertrophy with an increase in the thickness of the posterior wall of the left ventricle and an increase in the thickness of the interventricular septum. Many studies have found concentric hypertrophy in the hearts of trained rats and mice, although their training protocols are controversial. For example, studying low-intensity but high-volume and high-frequency exercise (90 min of swimming for 6 wks), Evangelista et al.  ADDIN EN.CITE Evangelista200315251525152517Evangelista, F. S.Brum, P. C.Krieger, J. E.Instituto do Coracao, Faculdade de Medicina, Universidade de Sao Paulo, Sao Paulo, SP, Brasil.Duration-controlled swimming exercise training induces cardiac hypertrophy in miceBraz J Med Biol ResBraz J Med Biol Res1751-936122003/12/11Adaptation, PhysiologicalAnimalsBlood PressureCardiomegaly/*etiology/pathologyCitrate (si)-Synthase/metabolismHeart RateMaleMiceMice, Inbred C57BLMuscle, Skeletal/enzymology*Physical Conditioning, AnimalPhysical Endurance*SwimmingTime Factors2003Dec0100-879X (Print) 0100-879X (Linking)14666261http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=14666261S0100-879X2003001200018 [pii]eng(14) demonstrated concentric hypertrophy in rats. Lemitsu et al.  ADDIN EN.CITE  ADDIN EN.CITE.DATA (20) observed that rats exercised on a treadmill at moderate intensity for 8 wks increased the mass index and the thickness of the left ventricle wall, indicating that moderate-intensity exercise causes concentric cardiac hypertrophy. Using a protocol of moderate loads of 5% body weight but of long duration, Sun et al.  ADDIN EN.CITE Sun200815831583158317Sun, B.Wang, J. H.Lv, Y. Y.Zhu, S. S.Yang, J.Ma, J. Z.Department of Human Sports Science, Nanjing Institute of Physical Education, Nanjing, 210014, China.Proteomic adaptation to chronic high intensity swimming training in the rat heartComp Biochem Physiol Part D Genomics ProteomicsComp Biochem Physiol Part D Genomics Proteomics108-17312008/03/012008Mar1878-0407 (Electronic) 1744-117X (Linking)20483212http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=20483212S1744-117X(07)00093-7 [pii] 10.1016/j.cbd.2007.11.001eng(28) also found concentric hypertrophy. Authors who observed that moderate- and low-intensity physical training can promote concentric hypertrophy could be misinterpreting their results if they compare these protocols to high-intensity training. Even though low- and moderate-intensity workloads were used, they were used over 8 wks without progressive acclimation, with sessions 120 min in duration, and up to 2 timesd-1, characterizing an exhaustive effort. Indeed, Woodiwiss and Norton  ADDIN EN.CITE Woodiwiss199515861586158617Woodiwiss, A. J.Norton, G. R.Department of Physiology, University of the Witwatersrand Medical School, Johannesburg, South Africa.Exercise-induced cardiac hypertrophy is associated with an increased myocardial complianceJ Appl Physiol (1985)J Appl Physiol (1985)1303-117841995/04/01AnimalsDiastole/physiologyDisease Models, AnimalHemodynamics/physiologyHypertrophy, Left Ventricular/*physiopathology/ultrasonographyMaleMyocardial Contraction/*physiologyOrgan SizePhysical Conditioning, Animal/*physiologyRatsRats, Sprague-DawleySystole/physiology1995Apr8750-7587 (Print) 0161-7567 (Linking)7615437http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=7615437eng(31), analyzing the hearts of rats submitted to volunteer exercise at running wheels in their cages, found physiological hypertrophy associated with an increase in the left ventricle wall, concluding that low-intensity exercise might lead to concentric hypertrophy. However, the authors did not control the intensity of the physical exercise, and the exercise alternated between moments of high and low activity followed by rest, similar to intermittent high-intensity anaerobic exercise. The swimming protocols tested here promoted cardiac development compatible with physiological cardiac hypertrophy, but the manner in which the remodeling happened varied according to the intensity, volume, and frequency of the physical exercise, which was eccentric in the MI training and concentric in the HI training. CONCLUSIONS The protocol of swimming physical training for Wistar rats validated in the present study is a proper method of studying experimental sport science, considering the responses to continuous aerobic exercise of low and moderate intensity as well as high-intensity intermittent anaerobic exercise. The protocol induced different metabolic pathways (according to the MLSS levels observed) and promoted distinct myocardium hypertrophy, bradycardia, and no change in blood pressure in normotensive rats. The success of this protocol is found in its progressive acclimation period, in which animals are acclimated to increasingly greater swimming loads and volumes in such a manner as to guarantee that they can be trained without significant muscle damage, fatigue, or mortality. Therefore, the use of this protocol is suggested because of its safety in terms of training intensity levels. We urge future experimental studies in sports science to properly define all protocols regarding adequate acclimation to loads and exercise volume, therefore avoiding misinterpretations.  ACKNOWLEDGMENTS The authors would like to acknowledge CAPES/CNPq for financial support and a fellowship with L.F. Guerreiro, A.A. Pereira, C. Martins, and C. Wally.  Address for correspondence: L.F. Guerreiro, PhD. Instituto de Cincias Biolgicas, Universidade Federal do Rio Grande - FURG, Av. Itlia km 8 - Campus Carreiros; ZIP CODE: 96.201-900 - Rio Grande - RS Brazil, Email: lfernandoguerreiro@hotmail.com  REFERENCES Anversa P, Capasso JM. Cardiac hypertrophy and ventricular remodeling. Lab Invest. 1991; 64:441-445. Arena R, Myers J, Forman DE, Lavie CJ, Guazzi M. Should high-intensity-aerobic interval training become the clinical standard in heart failure? Heart Fail Rev. 2013;18:95-105. Ascensao A, Ferreira R, Magalhaes J. Exercise-induced cardioprotection--biochemical, morphological and functional evidence in whole tissue and isolated mitochondria. Int J Cardiol. 2007;117:16-30. Beneke R, Leithauser RM, Ochentel O. Blood lactate diagnostics in exercise testing and training. Int J Sports Physiol Perform. 2011;6:8-24. Booth FW, Laye MJ, Spangenburg EE. Gold standards for scientists who are conducting animal-based exercise studies. J Appl Physiol. 2010;108:219-221. 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Swimming training can affect intrinsic calcium current characteristics in rat myocardium. Eur J Appl Physiol. 2008;104:549-555. Woodiwiss AJ, Norton GR. Exercise-induced cardiac hypertrophy is associated with an increased myocardial compliance. J Appl Physiol. 1995;78:1303-1311. Disclaimer The opinions expressed in JEPonline are those of the authors and are not attributable to JEPonline, the editorial staff or the ASEP organization.     PAGE  PAGE 1 Journal of Exercise Physiologyonline February 2015 Volume 18 Number 1  Editor-in-Chief Tommy Boone, PhD, MBA Review Board Todd Astorino, PhD Julien Baker, PhD Steve Brock, PhD Lance Dalleck, PhD Eric Goulet, PhD Robert Gotshall, PhD Alexander Hutchison, PhD M. Knight-Maloney, PhD Len Kravitz, PhD James Laskin, PhD Yit Aun Lim, PhD Lonnie Lowery, PhD Derek Marks, PhD Cristine Mermier, PhD Robert Robergs, PhD Chantal Vella, PhD Dale Wagner, PhD Frank Wyatt, PhD Ben Zhou, PhD Official Research Journal of the American Society of Exercise Physiologists ISSN 1097-9751 Official Research Journal of the American Society of Exercise Physiologists ISSN 1097-9751 )L+L8LLLLLLLLLM.MOMPMQMaMbMdMMMMMMM NNNNQN]N^N_NfNgNnNoNqN|NNN!O"O#O1O2O4OOOOOOOOOOOO[Pɹɹɹɹɹٹّh A^hOJQJ^Jh A^h(OJQJ^Jh A^h+6OJQJ^Jh A^hWu56OJQJ^Jh A^h+56OJQJ^Jh A^h+OJQJ^Jh A^hWuOJQJ^Jh A^hYOJQJ^J9[PmPnPoPPPPPPQQQ Q QQQQQQ'QAQDQJQQQŸŸŸŸqgZCgZ,hhh>4c5B*CJOJQJ^JaJphh>4c5OJQJ\^Jh>4cOJQJ^Jhhh>4c6CJOJQJ^JhY6CJOJQJ^Jhs5B*^JmH phsH h A^hsOJQJ^Jh A^hYOJQJ^Jh A^h+OJQJ^Jh A^hWuOJQJ^Jh A^h+6OJQJ^Jh A^hWu56OJQJ^Jh A^h+56OJQJ^JQQQQQQQQQQQQQQQQQQQQQQQQQQQQȾ~k~\J#h$Lgh(5CJ OJQJ^JaJ h(6CJOJQJ]^J%h2>0JCJOJQJ^JmHnHuh(0JCJOJQJ^J#jh(0JCJOJQJU^Jh( h(0Jjh(0JUhjhUhsOJQJ^Jh>4cOJQJ^J,hhh>4c5B*CJOJQJ^JaJph,hQ5h>4c5B*CJOJQJ^JaJphQQQQQQQQQQQQQQQQQRRRR(R)R*R,R-R$a$ h  hh]hh]h&`#$QRRR(R)R*R+R-R=RHRMRRRSR`RRRRRRR!SESlSSSSSSSSѿ߻}}}}}}}}}n\"h(B*CJOJQJ\^Jphh(6CJOJQJ]^J#hHh(CJOJQJ]^JaJh(CJOJQJ]^JaJ)hHh(5CJOJQJ\]^JaJjN_h(Uh(#h A^h(5CJOJQJ^JaJh A^h(5OJQJ^Jh(OJQJ^J,h$Lgh(5B*CJOJQJ^JaJph-R=RSR`RsRRRRRRRRS"S3SFSWSmSSSSSSSSSST$a$gdWgdu 7$8$H$gduSSTT$T%T&T'TDTGTtTuTTTTTTT»̟°hsOJQJ^Jh >;h$Lgh(CJaJ h$Lgh(CJOJQJ^JaJh( hWh( hgh(h(OJQJ^Jh(CJOJQJ\^J"h(B*CJOJQJ\^Jph(hj.h(B*CJOJQJ\^JphTT%T&T'TtTuTTTTTT$a$gdgdu$a$gdu$a$gdW$a$gdW ?0P1h:p5;+/ =!"#$% DpDShindo200015931593159317Shindo, M.Matsubara, T.Faculty of Sport and Health Science, Fukuoka University.[Physical activity and cardiac function]Nihon RinshoNihon Rinsho330-358 Suppl2000/11/21Cardiac OutputExercise/*physiologyExercise TherapyHeart/*physiologyHeart RateHumansPeptidyl-Dipeptidase A/geneticsPolymorphism, GeneticSports/physiology2000Sep0047-1852 (Print) 0047-1852 (Linking)11085138http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11085138jpnBooth201013091309130917Booth, F. 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A.Kokubun, E.Department of Physical Education, Bioscience Institute, University of Sao Paulo State (UNESP), Avenida 24-A, 1515 Bela Vista, Rio Claro, Sao Paulo, 13506-900 Brazil. cgobatto@uol.com.brMaximal lactate steady state in rats submitted to swimming exerciseComp Biochem Physiol A Mol Integr PhysiolComp Biochem Physiol A Mol Integr Physiol21-713012001/10/24Anaerobic Threshold/*physiologyAnimalsLactic Acid/*bloodMalePhysical Conditioning, Animal/physiologyPhysical Endurance/physiologyRatsRats, WistarSwimming/*physiology2001Aug1095-6433 (Print) 1095-6433 (Linking)11672680http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=11672680S1095-6433(01)00362-2 [pii]engManchado Fde200616581658165817Manchado Fde, B.Gobatto, C. A.Voltarelli, F. A.Rostom de Mello, M. A.Sao Paulo State University-UNESP, Department of Physical Education, Av. 24A, 1515, Bela Vista, 13506-900 - Rio Claro, Sao Paulo, Brazil. fbmanchado@yahoo.com.brNon-exhaustive test for aerobic capacity determination in swimming ratsAppl Physiol Nutr MetabAppl Physiol Nutr Metab731-63162007/01/11AnimalsExercise TestLactic Acid/bloodMale*Physical Conditioning, Animal/physiologyRatsRats, Wistar*Swimming2006Dec1715-5312 (Print)17213888http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17213888h06-079 [pii] 10.1139/h06-079engDManchado Fde200616581658165817Manchado Fde, B.Gobatto, C. A.Voltarelli, F. A.Rostom de Mello, M. A.Sao Paulo State University-UNESP, Department of Physical Education, Av. 24A, 1515, Bela Vista, 13506-900 - Rio Claro, Sao Paulo, Brazil. fbmanchado@yahoo.com.brNon-exhaustive test for aerobic capacity determination in swimming ratsAppl Physiol Nutr MetabAppl Physiol Nutr Metab731-63162007/01/11AnimalsExercise TestLactic Acid/bloodMale*Physical Conditioning, Animal/physiologyRatsRats, Wistar*Swimming2006Dec1715-5312 (Print)17213888http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&dopt=Citation&list_uids=17213888h06-079 [pii] 10.1139/h06-079engGobatto200112781278127817Gobatto, C. A.de Mello, M. A.Sibuya, C. Y.de Azevedo, J. R.dos Santos, L. 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