Characteristics of elite open-water swimmers

Open-water swimming (5, 10, and 25 km) has many unique challenges that separate it from other endurance sports, like marathon running and cycling. The characteristics of a successful open-water swimmer are unclear. The purpose of this study was to determine the physical and metabolic characteristics of a group of elite-level open-water swimmers. The open-water swimmers were participating in a 1-week training camp. Anthropometric, metabolic, and blood chemistry assessments were performed on the athletes. The swimmers had a VO(2)peak of 5.51 +/- 0.96 and 5.06 +/- 0.57 ml.kg(-1).min(-1) for males and females, respectively. Their lactate threshold (LT) occurred at a pace equal to 88.75% of peak pace for males and 93.75% for females. These elite open-water swimmers were smaller and lighter than competitive pool swimmers. They possess aerobic metabolic alterations that resulted in enhanced performance in distance swimming. Trainers and coaches should develop dry-land programs that will improve the athlete's muscular endurance. Furthermore, programs should be designed to increase the LT velocity as a percentage of peak swimming velocity.     

Modeling and relationship of respiratory exchange ratio to athletic performance

Previous research has related the results of tests of maximum aerobic capacity to performance for endurance athletes. These results are often only able to predict the running velocity of races such as the marathon. This investigation sought to determine the absolute V[Combining Dot Above]O2 at various respiratory exchange ratio (RER) values (0.85, 0.90, 0.95, 1.0, 1.05, and 1.10) by using a third-order polynomial regression to model the physiological responses for V[Combining Dot Above]O2 and RER obtained from an assessment of maximum aerobic capacity. The V[Combining Dot Above]O2 determined was subsequently correlated to race performance. The participants in the study were selected from a population of National Collegiate Athletic Association Division 1 crosscountry runners (male n = 7, female n = 7, age 20.5 ± 0.9 years; height 170.3 ± 8.2 cm; weight 59.7 ± 8.7 kg; V[Combining Dot Above]O2max 57.0 ± 7.8 ml O2·kg·min). Third-order regression analysis resulted in strong curve fitting between the variables (r = 0.949 ± 0.03). Partial correlations (controlled for weight) were used to assess the relationship between oxygen consumption at the desired points of RER and race performance. The partial correlations revealed that the absolute oxygen consumptions at all RER points of interest were significantly correlated to race performance (r > 0.740, p < 0.01). There was a significant difference in the strength of the correlations for the points RER 0.95 (t = 2.68957, p = 0.01), 1.0 (t = 2.18516, p = 0.03), and 1.05 (t = 1.85668, p = 0.04) and the correlations found for RER 0.85. After converting the oxygen consumption at the RER points to estimated horizontal running speeds, only the estimate at RER 1.05 was not statistically different from the actual speed achieved in the culminating XC race. It can be suggested based upon these results that coaches of collegiate crosscountry runners who engage in metabolic testing of athletes examine the estimated running pace at RER 1.05 to gain an insight into a runner's potential.     

Running Pace Decrease during a Marathon Is Positively Related to Blood Markers of Muscle Damage

Background

Completing a marathon is one of the most challenging sports activities, yet the source of running fatigue during this event is not completely understood. The aim of this investigation was to determine the cause(s) of running fatigue during a marathon in warm weather.

Methodology/Principal Findings

We recruited 40 amateur runners (34 men and 6 women) for the study. Before the race, body core temperature, body mass, leg muscle power output during a countermovement jump, and blood samples were obtained. During the marathon (27 °C; 27% relative humidity) running fatigue was measured as the pace reduction from the first 5-km to the end of the race. Within 3 min after the marathon, the same pre-exercise variables were obtained.

Results

Marathoners reduced their running pace from 3.5 ± 0.4 m/s after 5-km to 2.9 ± 0.6 m/s at the end of the race (P<0.05), although the running fatigue experienced by the marathoners was uneven. Marathoners with greater running fatigue (> 15% pace reduction) had elevated post-race myoglobin (1318 ± 1411 v 623 ± 391 µg L⁻¹; P<0.05), lactate dehydrogenase (687 ± 151 v 583 ± 117 U L⁻¹; P<0.05), and creatine kinase (564 ± 469 v 363 ± 158 U L⁻¹; P = 0.07) in comparison with marathoners that preserved their running pace reasonably well throughout the race. However, they did not differ in their body mass change (−3.1 ± 1.0 v −3.0 ± 1.0%; P = 0.60) or post-race body temperature (38.7 ± 0.7 v 38.9 ± 0.9 °C; P = 0.35).

Conclusions/Significance

Running pace decline during a marathon was positively related with muscle breakdown blood markers. To elucidate if muscle damage during a marathon is related to mechanistic or metabolic factors requires further investigation.     

Are mitochondrial haplogroups associated with elite athletic status? A study on a Spanish cohort

There is increasing evidence regarding the association between mitochondrial DNA (mtDNA) and aerobic capacity; however, whether mtDNA haplogroups are associated with the status of being an elite endurance athlete is more controversial. We compared the frequency distribution of mtDNA haplogroups among the following groups of Spanish (Caucasian) men: 102 elite endurance athletes (professional road cyclists, endurance runners), 51 elite power athletes (jumpers, throwers and sprinters), and 478 non-athletic controls. We observed a significant difference between endurance athletes and controls (Fisher exact test = 17.89, P = 0.015; Bonferroni's significant threshold = 0.017), yet not between power athletes and controls (Fisher exact test = 47.99, P = 0.381) or between endurance and power athletes (Fisher exact test = 5.53, P = 0.597). We observed that the V haplogroup was overrepresented in endurance athletes (15.7%) compared with controls (7.5%) (odds ratio: 2.284; 95% confidence interval: 1.237, 4.322). In conclusion, our findings overall support the idea that mtDNA variations could be among the numerous contributors to the status of being an elite endurance athlete, whereas no association was found with elite power athletic status.     

Physiological analysis of running performance: revision of the hyperbolic model

1 This paper presents a physiological model for the analysis of the "running curve" (i.e. the relationship between running time and power output) (Table I and Fig. 1). This model is derived from the basic hyperbolic model (Eq. 2: Fig. 2 a) modified in order to take into account the slow adjustment of oxygen utilization and of glycolysis at the onset of exercise (Eq. 7) and the progressive reduction of average aerobic power sustained with increasing running time for races lasting longer than 7 min (Eq. 10; Fig. 2 b, 2 d). 2 This model provides a satisfactory smoothing of the running curve from 200 m to marathon. Average errors (in absolute value) between actual and estimated power output are 0.9 +/- 0.2% (+/- SE) and 1.3 +/- 0.4% for male and female world records respectively (Table II). The basic hyperbolic model (Ptc = A/tc + B) provides a satisfactory smoothing of the running curve on comparatively narrow ranges of events (Table III). 3 Estimations of anaerobic energy stores and of VO2 max made for male and female from world records (1,219 J.kg-1 and 76.4 ml.kg-1.min-1; 1,118 J.kg-1 and 67.8 ml.kg-1.min-1 respectively) are in reasonable agreement with expected values in elite runners (Table IV). 4 The model allows the computation of an objective measure of endurance, i.e. the so-called "ability to sustain a high percentage of VO2 max for a long period of time".(ABSTRACT TRUNCATED AT 250 WORDS)     

Advantages of a smaller bodymass in humans when distance-running in warm, humid conditions

Using a 65-kg athlete running a 2 h 10 min marathon as an example, we estimated that imbalances between approximately 1400 W of heat production and dissipation would occur in ambient temperatures of 17 degrees C at 90% relative humidity (rh) to 37 degrees C at 50% rh. Because heat production during running depends on body mass and heat loss depends on surface area, intercepts between predicted heat production and maximal heat loss with increasing speeds depend on an athlete's body mass. At 35 degrees C and 60% rh, a 45-kg athlete could maintain thermal balance by running a 2 h 13 min marathon at 19.1 km x h(-1) but a 75-kg athlete would only be able run a 3 h 28 min marathon at 12.2 km x h(-1). In both cases, the production of 970-1020 W of heat would necessitate the evaporation of at least 1.5-1.6 l of sweat per hour. A lower metabolic heat production in lighter runners at any given speed may be one reason why smallness of stature is an asset in distance running.     

The impact of different pacing strategies on five-kilometer running time trial performance

The purpose of this study was to determine the optimal 1.63-km (1-mile) pacing strategy for 5-km running performance in moderately trained women distance runners. Eleven women distance runners (20.7 +/- 0.8 years, 163.8 +/- 2.0 cm, 57.0 +/- 2.2 kg, 51.7 +/- 1.0 ml.kg(-1).min(-1), 18.9 +/- 0.8% fat, 78.1 +/- 1.4% VO(2)max at lactate threshold) performed 2 preliminary 5-km time trials on a treadmill to establish baseline 5-km times. The average 1.63-km split pace of the fastest preliminary trial was manipulated for the first 1.63 km of the experimental trials and run either equal to (EVEN), 3% faster than (3%), or 6% faster than (6%) the current baseline average 1.63-km pace for each subject. Ventilation (V(E)), oxygen consumption VO(2)max )), respiratory exchange ratio, and heart rate were measured continuously. Overall 5-km times were not different (p > 0.05) for the EVEN, 3% and 6% trials finishing in 21:11 (minutes/seconds) +/- 29 seconds, 20:52 +/- 36 seconds and 20:39 +/- 29 seconds, respectively. The fastest time for 8 subjects resulted from the 6% trial and the other 3 subjects' fastest times resulted from the 3% trial. The overall exercise intensity (%VO(2)max , %VO(2)max above lactate threshold, V(E), and respiratory exchange ratio) of the first 1.63-km split was not different between the 3 and 6% trials, despite the 6% trial being 13 seconds faster than the 3% trial. Based on these findings, initial 1.63-km starting paces of a 5-km race can be 3 to 6% greater than current average race pace without negatively impacting performance. In order to optimize 5-km performance, runners should start the initial 1.63 km of a 5-km race at paces 3-6% greater than their current average race pace.     

Lactate-related factors as a critical determinant of endurance

Many interrelated physiological and/or morphological factors have been demonstrated to influence endurance exercise performance. Some of these factors include skeletal musculature, running economy, maximal oxygen uptake (VO2max), maximal steady state (MSS), onset of blood lactate accumulation (OBLA), onset of plasma lactate accumulation (OPLA), and anaerobic (or lactate) threshold (AT or LT). The present paper focuses mainly on VO2max, MSS, OBLA, OPLA and LT, all of which have been postulated as a prerequisite in endurance exercise success. This paper consists of: (1) significance of La-related variables, (2) longitudinal studies, (3) comments, and (4) conclusion. Briefly, it is suggested that estimation of endurance exercise potential could be obtained with relatively high precision using laboratoriously measured La-related variables. The most critical determinant of endurance exercise performance such as marathon time is considered running velocity (V) at which LT is detected (V / LT), VO2 / LT, or V / MSS, while V / OBLA appears to be the best predictor of performance in endurance events of 16 km or shorter distances.     

Isotonic and isoelectric endurance tests for the upper trapezius muscle

As an alternative to a conventional endurance electromyogram (EMG) test for assessment of muscle capacity in the upper trapezius muscle (isotonic test, IT), an isoelectric test (IE) has been investigated. Nine subjects performed the two endurance tests. The tests were performed with a straight horizontal arm in the plane of the scapula. In IE, EMG amplitude was fed back to the subject and the subject was instructed to maintain a constant EMG activity equal to the level with the arm unsupported. Subjective ratings of local fatigue were obtained during the experiments. The EMG recordings from both tests were analysed for the root mean square value as well as according to the mean power frequency (MPF) technique. All the subjects endured maximal 15 min IE while in IT the median endurance was 11.9 min. Average subjective ratings of perceived fatigue increased more in IT compared to IE. The average normalised MPF curve from IT increased while that from IE was unaffected. There was a significant difference between the MPF results at 6 and 8 min. It is noteworthy that the MPF was higher in IT in spite of a higher accumulated biomechanical load. It was concluded from these studies that the interpretation of decreased MPF as an indicator of increased local muscle fatigue is doubtful at low contraction levels. It is suggested that an IE is a more appropriate method for the functional evaluation of low threshold motor units of the upper trapezius muscle in research into occupational disorders. Accepted: 15 October 1996     

No association between Angiotensin Converting Enzyme (ACE) gene variation and endurance athlete status in Kenyans

East African runners are continually successful in international distance running. The extent to which genetic factors influence this phenomenon is unknown. The insertion (I) rather than deletion (D) of a 287 bp fragment in the human angiotensin converting enzyme (ACE) gene is associated with lower circulating and tissue ACE activity and with endurance performance amongst Caucasians. To assess the association between ACE gene variation and elite endurance athlete status in an African population successful in distance running, DNA samples were obtained from 221 national Kenyan athletes (N), 70 international Kenyan athletes (I), and 85 members of the general Kenyan population (C). Blood samples were obtained from C and assayed for circulating ACE activity. ACE I/D (rs????--from NCBI SNPdb first time poly mentioned) genotype was determined, as was genotype at A22982GD (rs????--from NCBI SNPdb first time poly mentioned) which has been shown to associate more closely with ACE levels in African subjects than the I/D polymorphism. ACE I/D and A22982G genotypes explained 13 and 24% of variation in circulating ACE activity levels (P = 0.034 and <0.001 respectively). I/D genotype was not associated with elite endurance athlete status (df = 4, chi(2) = 4.1, P=0.39). In addition, genotype at 22982 was not associated with elite endurance athlete status (df = 4, chi(2) = 5.7, P = 0.23). Nor was the A allele at 22982, which is associated with lower ACE activity, more prevalent in N (0.52) or I (0.41) relative to C (0.53). We conclude that ACE I/D and A22982G polymorphisms are not strongly associated with elite endurance athlete status amongst Kenyans.     

Modeling: optimal marathon performance on the basis of physiological factors

This paper examines current concepts concerning "limiting" factors in human endurance performance by modeling marathon running times on the basis of various combinations of previously reported values of maximal O2 uptake (VO2max), lactate threshold, and running economy in elite distance runners. The current concept is that VO2max sets the upper limit for aerobic metabolism while the blood lactate threshold is related to the fraction of VO2max that can be sustained in competitive events greater than approximately 3,000 m. Running economy then appears to interact with VO2max and blood lactate threshold to determine the actual running speed at lactate threshold, which is generally a speed similar to (or slightly slower than) that sustained by individual runners in the marathon. A variety of combinations of these variables from elite runners results in estimated running times that are significantly faster than the current world record (2:06:50). The fastest time for the marathon predicted by this model is 1:57:58 in a hypothetical subject with a VO2max of 84 ml.kg-1.min-1, a lactate threshold of 85% of VO2max, and exceptional running economy. This analysis suggests that substantial improvements in marathon performance are "physiologically" possible or that current concepts regarding limiting factors in endurance running need additional refinement and empirical testing.     

Effects of three types of resisted sprint training devices on the kinematics of sprinting at maximum velocity

Resisted sprint running is a common training method for improving sprint-specific strength. For maximum specificity of training, the athlete's movement patterns during the training exercise should closely resemble those used when performing the sport. The purpose of this study was to compare the kinematics of sprinting at maximum velocity to the kinematics of sprinting when using three of types of resisted sprint training devices (sled, parachute, and weight belt). Eleven men and 7 women participated in the study. Flying sprints greater than 30 m were recorded by video and digitized with the use of biomechanical analysis software. The test conditions were compared using a 2-way analysis of variance with a post-hoc Tukey test of honestly significant differences. We found that the 3 types of resisted sprint training devices are appropriate devices for training the maximum velocity phase in sprinting. These devices exerted a substantial overload on the athlete, as indicated by reductions in stride length and running velocity, but induced only minor changes in the athlete's running technique. When training with resisted sprint training devices, the coach should use a high resistance so that the athlete experiences a large training stimulus, but not so high that the device induces substantial changes in sprinting technique. We recommend using a video overlay system to visually compare the movement patterns of the athlete in unloaded sprinting to sprinting with the training device. In particular, the coach should look for changes in the athlete's forward lean and changes in the angles of the support leg during the ground contact phase of the stride.     

The effects of training on the metabolic and respiratory profile of high-intensity cycle ergometer exercise

The tolerable work duration (t) for high-intensity cycling is well described as a hyperbolic function of power (W): W = (W'.t-1) + Wa, where Wa is the upper limit for sustainable power (lying between maximum W and the threshold for sustained blood [lactate] increase, theta lac), and W' is a constant which defines the amount of work which can be performed greater than Wa. As training increases the tolerable duration of high-intensity cycling, we explored whether this reflected an alteration of Wa, W' or both. Before and after a 7-week regimen of intense interval cycle-training by healthy males, we estimated ( ) theta lac and determined maximum O2 uptake (mu VO2); Wa; W'; and the temporal profiles of pulmonary gas exchange, blood gas, acid-base and metabolic response to constant-load cycling at and above Wa. Although training increased theta lac (24%), mu VO2 (15%) and Wa (15%), W' was unaffected. For exercise at Wa, a steady state was attained for VO2, [lactate] and pH both pre- and post-training, despite blood [norepinephrine] and [epinephrine] ([NE], [E]) and rectal temperature continuing to rise. For exercise greater than Wa, there was a progressive increase in VO2 (resulting in mu VO2 at fatigue), [lactate], [NE], [E] and rectal temperature, and a progressive decrease for pH. We conclude that the increased endurance capacity for high-intensity exercise following training reflects an increased W asymptote of the W-t relationship with no effect on its curvature; consequently, there is no appreciable change in the amount of work which can be performed above Wa.(ABSTRACT TRUNCATED AT 250 WORDS)     

Some biochemical threshold mechanisms

Certain arrangements of enzymatic (bimolecular) subsystems lead to characteristic threshold-type response. Two simple cases of such systems are studied here in terms of steady state behavior and explicit relationships between system and curve parameters. It is found that the curvature of the threshold curve is directly related to the equivalent Michaelis constant and, in the case of saturated threshold curve, the slope of the curve at the idealized threshold is limited by the ratio of saturation to threshold. This slope may be appreciably increased up to a stepwise response at the threshold if a multisubstrate complex of the enzyme is the only species which affects the enzyme mediated transport.     

Changes in H reflex and V wave following short-term endurance and strength training

This study examined the effects of 3 wk of either endurance or strength training on plasticity of the neural mechanisms involved in the soleus H reflex and V wave. Twenty-five sedentary healthy subjects were randomized into an endurance group (n = 13) or strength group (n = 12). Evoked V-wave, H-reflex, and M-wave recruitment curves, maximal voluntary contraction (MVC), and time-to-task-failure (isometric contraction at 40% MVC) of the plantar flexors were recorded before and after training. Following strength training, MVC of the plantar flexors increased by 14.4 ± 5.2% in the strength group (P < 0.001), whereas time-to-task-failure was prolonged in the endurance group (22.7 ± 17.1%; P < 0.05). The V wave-to-maximal M wave (V/M(max)) ratio increased significantly (55.1 ± 28.3%; P < 0.001) following strength training, but the maximal H wave-to-maximal M wave (H(max)/M(max)) ratio remained unchanged. Conversely, in the endurance group the V/M(max) ratio was not altered, whereas the H(max)/M(max) ratio increased by 30.8 ± 21.7% (P < 0.05). The endurance training group also displayed a reduction in the H-reflex excitability threshold while the H-reflex amplitude on the ascending limb of the recruitment curve increased. Strength training only elicited a significant decrease in H-reflex excitability threshold, while H-reflex amplitudes over the ascending limb remained unchanged. These observations indicate that the H-reflex pathway is strongly involved in the enhanced endurance resistance that occurs following endurance training. On the contrary, the improvements in MVC following strength training are likely attributed to increased descending drive and/or modulation in afferents other than Ia afferents.     

Validation of two running tests as estimates of maximal aerobic power in children

In order to validate the "Maximal Multistage 20 Meter Shuttle Run Test" by Leger and Lambert (1982) (20-MST) as an estimate of maximal aerobic power (VO2max) and to compare the results of this test with the results of a 6 min endurance run, 82 subjects (41 boys and 41 girls) aged 12-14 performed the 20-MST and the 6 min endurance run, and had their VO2max directly measured during maximal treadmill running. The 20-MST is a maximal running test starting at a running speed of 8.0 km X h-1, which is increased every minute and in which the pace is set by an audio signal. Performing the test, one runs a 20-meter course back and forth. The test result is expressed as "palier" (one palier is approximately one minute). The mean results of the 20-MST were, for boys, 8.0 palier (+/- 1.7) and for girls, 6.4 palier (+/- 1.5). The mean results of the 6 min endurance run were for boys, 1264.4 meters (+/- 160.8), and for girls, 1103.9 meters (+/- 144.7). The mean VO2max for boys was 53.2 ml X kg-1 X min-1 (+/- 5.4) and for girls, 44.1 (+/- 4.8) ml X kg-1 X min-1. The correlation coefficient between VO2max and the 20-MST was found to be 0.68 (+/- 3.9) for boys, 0.69 (+/- 3.4) for girls and 0.76 (+/- 4.4) for both sexes, and that of VO2max with the 6 min endurance run was 0.51 (+/- 4.6) for boys, 0.45 (+/- 4.3) for girls and 0.63 (+/- 5.3) for both sexes. The conclusion is that the 20-MST is a suitable tool for the evaluation of maximal aerobic power. Although the differences in validity between the 20-MST and the 6 minutes endurance run were statistically not significant (p greater than 0.05), for reasons of practicability the 20-MST should be preferred to the 6 minutes endurance run when used in physical education classes.     

Effects of gradual-elastic compression stockings on running economy, kinematics, and performance in runners

We investigated the effect of gradual-elastic compression stockings (GCSs) on running economy (RE), kinematics, and performance in endurance runners. Sixteen endurance trained athletes (age: 34.73 ± 6.27 years; VO2max: 62.83 ± 9.03 ml·kg(-1)·min(-1); 38 minutes in 10 km; 1 hour 24 minutes in half marathon) performed in random order 4 bouts of 6 minutes at a recent half-marathon pace on a treadmill to evaluate RE with or without GCSs. Subsequently, 12 athletes were divided into 2 equal groups matched by their VO2max, and they performed a time limit test (T(lim)) on a treadmill at 105% of a recent 10-km pace with or without GCSs for evaluation of physiological responses and running kinematics. There were no significant differences in the RE test in all of the variables analyzed for the conditions, but a moderate reproducibility for some physiological responses was detected in the condition with GCSs. In the T(lim), the group that wore GCSs reached a lower % of maximum heart rate (HRmax) compared with the control group (96.00 ± 2.94 vs. 99.83 ± 0.40) (p = 0.01). Kinematics did not differ between conditions during the T(lim) (p > 0.05). There were improvement trends for time to fatigue (337 vs. 387 seconds; d = 0.32) and a lower VO2peak (≈53 vs. 62 ml·kg(-1)·min(-1); d = 1.19) that were detected with GCSs during the T(lim). These results indicate that GCSs reduce the % of HRmax reached during a test at competition pace. The lower reproducibility of the condition with GCSs perhaps suggests that athletes may possibly need an accommodation period for systematically experiencing the benefits of this garment, but this hypothesis should be further investigated.     

The significance of the aerobic-anaerobic transition for the determination of work load intensities during endurance training.

Anaerobic and aerobic-anaerobic threshold (4 mmol/l lactate), as well as maximal capacity, were determined in seven cross country skiers of national level. All of them ran in a treadmill exercise for at least 30 min at constant heart rates as well as at constant running speed, both as previously determined for the aerobic-anaerobic threshold. During the exercise performed with a constant speed, lactate concentration initially rose to values of nearly 4 mmol/l and then remained essentially constant during the rest of the exercise. Heart rate displayed a slight but permanent increase and was on the average above 170 beats/min. A new arrangement of concepts for the anaerobic and aerobic-anaerobic threshold (as derived from energy metabolism) is suggested, that will make possible the determination of optimal work load intensities during endurance training by regulating heart rate.     

Holding potential affects the apparent voltage-sensitivity of sodium channel activation in crayfish giant axons.

Sodium channel activations, measured as the fraction of channels open to peak conductance for different test potentials (F[V]), shows two statistically different slopes from holding potential more positive than -90 mV. A high valence of 4-6e is indicated a test potentials within 35 mV of the apparent threshold potential (circa -65 mV at -85 mV holding potential). However, for test potentials positive to -30 mV, the F(V) curve shows a 2e valence. The F(V) curve for crayfish axon sodium channels at these "depolarized" holding potentials thus closely resembles classic data obtained from other preparations at holding potentials between -80 and -60 mV. In contrast, at holding potentials more negative than -100 mV, the high slope essentially disappears and the F(V) curve follows a single Boltzmann distribution with a valence of approximately 2e at all potentials. Neither the slope of this simple distribution nor its midpoint (-20 mV) was significantly affected by removal of fast inactivation with pronase. The change in F(V) slope, when holding potential is increased from -85 to -120 mV, does not appear to be caused by the contribution of a second channel type. The simple voltage dependence of sodium current found at Vh -120 mV be used by to discriminate between models of sodium channel activation, and rules out models with three particles of equal valence.