Speed skating is an intriguing sport to study, in particular because speed skaters need to adopt the biomechanically favorable crouched position that is essential for speed skating performance (Van Ingen Schenau et al., 1983; Konings et al., 2015). This crouched position combined with a relatively long gliding phase and high intramuscular forces at the same time leads to a physiological disadvantage as it is suggested to increase deoxygenation of the working muscles (Rundell, 1996; Rundell et al., 1997; Konings et al., 2015). The presence and magnitude of deoxygenation during speed skating has been investigated using near infrared spectroscopy, demonstrating a linear relationship with knee-angle in long track speed skating (Foster et al., 1999) and an asymmetry between the legs in short track skating, particularly related to cornering (Hesford et al., 2012, 2013a,b). Most likely, the reduced blood flow to the working muscles reduces the aerobic capacity of the recruited muscle groups (Rundell, 1996), whereas not all the available oxygen can be transported toward the working muscles during speed skating. The reduced blood flow is expected to exacerbate processes related to perceived fatigue and recovery, thereby impacting on elite performance. Due to the technical nature of speed skating that is affecting physiological mechanisms such as oxygenation and blood flow, this sport provides a unique setting allowing us to uncover novel mechanistic insights into these aspects of the physiological response to exercise in elite sports.

Changes in muscle oxygenation can inform on a drop in tissue pO₂ and therefore potentially correlate with fatigue and performance. Global hypoxia decreases arterial oxygenation and can exacerbate peripheral fatigue and fatigability (Katayama et al., 2007; Romer et al., 2007). Local ischemia leading to a 7% or larger reduction in muscle oxygenation can lead to decreased muscle production in the fore-arm muscles (Murthy et al., 2001). Increased intramuscular pressure has been suggested to decrease blood flow and thereby reduce tissue oxygenation at isometric contractions >25–35% of maximal torque capacity in upper leg muscles (de Ruiter et al., 2007). Under these conditions the muscle fibers will largely depend on glycolysis as an energy source, potentially exacerbating fatigue.

Long track and short track speed skating are expected to differ in terms of asymmetry between the legs, associated with the technical nature and characteristics of the different sports. In our communication with coaches and athletes, we became aware that when imposing testing sessions of similar load in short-track (ST) and long-track (LT) skating to elite ST athletes, the athletes felt more fatigued after ST compared to LT skating sessions. As short track athletes compete on a shorter track with shorter straights and more and tighter curves during their race, differences in intra-muscular pressure, aerobic physiology and thereby experienced fatigue and recovery could be expected.

Therefore, the aim of the current study was to measure the influence of skating mode (short-track vs. long-track) on muscle oxygenation, and determine how this affects perceived fatigue and recovery in speed skating. The crouched position of speed skaters leads to a unique form of physiological stress: static occlusion of blood flow that occurs repetitively with every cycle, that might hinder recovery related to oxygenation (Rundell, 1996; Rundell et al., 1997; Foster et al., 1999). In short track speed skating, with tighter corners, longer periods of occlusion are seen for the right leg in particular (Hesford et al., 2012, 2013a,b), and we expect this will impact on recovery and perceived fatigue. It is hypothesized that the working muscles will remain more deoxygenated during short-track speed skating, associated with the relatively long periods of static contraction, and that there is thereby a larger asymmetry between the left and right leg. It is expected that this will lead to higher experienced fatigue, and longer periods of recovery after short-track speed skating compared to long-track speed skating. If a link between muscle oxygenation and fatigue is clear for the same elite athletes in a single sport (speed skating) depending only on the mode of the exercise, this suggests that measures of muscle oxygenation could inform in constructing training and recovery loads in other sports or activities involving local ischemia, even ones that do not have a specific asymmetric component.

Means ± SD of the time-trial performance for the long-track time-trials (LT) and the short-track time-trials (ST) can be found in Table 1. Faster completion times (LT = 63.45 ± 2.20, ST = 60.94 ± 2.90 s; p = 0.005) and a higher mean velocity (LT = 44.36 ± 1.97, ST = 38.26 ± 1.46 km/h; p < 0.001) were found for LT compared to ST. Nevertheless, no differences in heart rate (LT = 178.8 ± 5.8, ST = 179.6 ± 6.6 bpm; p = 0.609), lactate (LT = 10.5 ± 1.5, ST = 11.0 ± 1.1 mmol/l; p = 0.356) or RPE (LT = 8.9 ± 1.2, ST = 9.2 ± 1.1; p = 0.555) were found between the testing conditions, suggesting time-trials could be perceived as of a similar physiological intensity. No differences in knee angle were reported between conditions (LT = 103.4 ± 4.5, ST = 103.8 ± 2.9°; P = 0.843). However, a lower trunk angle was found during ST (LT = 16.6 ± 1.1, ST = 15.3 ± 0.9°; p < 0.001). In addition, deceleration of the participants was higher after ST compared to LT (see Table 1 and Figure 1).

Changes in muscle oxygenation between both legs occurred in both LT as well as ST (see Table 1 and Figures 2–4). Typical NIRS traces are shown for TSI (Figure 2) and HHb, HbO₂, and tHb (Figures 3, 4). They illustrate that at the start of the race compared to baseline there is a fall in muscle oxygen saturation (TSI) caused by rise in HHb and fall in HbO₂. The fall in HbO₂ is larger than the rise in HHb, resulting in a consequent fall in tHb. As the TT progresses, muscle oxygenation (TSI) increases. This is caused entirely by an increase in HbO₂. There is no change in HHb, with the result that tHb increases along with HbO₂.

There is an asymmetry in muscle oxygenation between left and right legs as the TT progresses. Table 1 shows the rate of change in time for the NIRS parameters. Muscle oxygenation significantly increases in the left leg compared to the right in both ST and LT. However, the difference is larger in ST than LT. These oxygenation changes are dominated by HbO₂ and not HHb. HbO₂ increases in both LT and ST with the larger differences being seen in ST. There is no significant change in HHb in either leg. An HbO₂ increase with no change in HHb is reflected in an increase in tHb. Again this increase is larger for the left leg than the right leg in ST compared to LT.

The use of spatially resolved methods enabled an absolute measurement of muscle oxygen saturation (TSI) at the end of the TT. The only difference between legs was seen for the right leg in ST. Muscle oxygenation in left ST, right ST and left ST were equivalent to one another and all higher than right ST.

The temporal changes in TSI, tHb, HbO₂, and HHb reveal significant differences in oxygenation in the course of a single lap. This can be illustrated by higher resolution analysis of a single half lap in LT (Figure 5) and ST (Figure 6). The lap was divided into segments indicated in both figures. An illustration of how these different segments relate to technique can be found in Figure 7 (LT) and Figure 8 (ST) During the straights in LT (Figure 5) the leg desaturates in the gliding phase and push-off (when the leg is on the ice), while the leg resaturates in the repositioning phase (when the leg is off the ice). In ST (Figure 6), the typical way of traveling (hanging) around the corner on only the right leg leads to a resaturation of the left leg and a desaturation of the right leg.

Figure 9 indicates the changes in muscle oxygenation during a 6-s sprint cycling test taken before and after the time trials. No differences were found in the drop of the ΔTSI% during the 6-s sprint between the legs, between the pre and post-test or between skating modes (P > 0.05; Table 2). The recovery halftime in ΔTSI% after the post 6-s sprint test was significantly increased after ST compared to LT. However, there were no differences in recovery rates between the legs for either skating mode (see Table 2). In addition, participants reported they felt more recovered 2 h (P = 0.014) and 4 h (P = 0.026) after LT compared to ST (Table 2).

The aim of this study was to examine the influence of speed skating mode (short-track vs. long-track) on muscle oxygenation in speed skating, and how it consequently affects processes of fatigue and recovery. In order to compare long-track and short-track speed skating, time-trials on both tracks of expected similar physiological intensity were set-up. This was done by asking the participants to perform each time-trial maximally. Although the long-track time-trials were significantly faster and longer in their duration, the lack of difference in heart rate during, and lactate value and rate of perceived exertion directly after the time-trials indicated that both skating modes were of a similar physiological intensity. A comparison of the values for heart rate, lactate, and RPE found in this study with other skating studies indicated that our subjects indeed performed maximally during the time-trials at both tracks (Rundell et al., 1997; Hesford et al., 2013b).

The TT started after a flying lap. The starting values for TSI, tHb, HHb, and HbO₂ therefore need to be compared to those seen after the first lap of our previous studies which used a standing start (Hesford et al., 2012, 2013a,b). The initial changes seen in our new short-track data are entirely consistent with our previous data (Hesford et al., 2012), i.e., an increase in HHb and a larger fall in HbO₂ resulting in a drop in TSI and tHb. The amount of hemoglobin in the optical field of view has decreased and what is seen has less oxygen bound. The simplest explanation of this combination of NIRS data is that there is a decrease in blood volume flow and an increase in muscle oxygen consumption. The same trend is seen in the long-track data, indicating that here as well the combination of increased tissue oxygen demand and physical occlusion has caused a combination of vasoconstriction and increased oxygen consumption.

The two skating modes respond similarly after these initial effects. Consistent with our previous studies (Hesford et al., 2012, 2013a,b) there is an increase in muscle oxygenation with time. This is evidenced by a rise in TSI. However, this TSI increase is a result of an increase in HbO₂ alone. HHb is unaltered with the consequence that tHb increases. The amount of hemoglobin in the optical field of view has now increased and all of this increase is due to hemoglobin that has oxygen bound to it. The rate of oxygen extraction (HHb) does not change with time. The simplest explanation of this data is that the blood volume flow has increased but none of the extra arterial oxygen being delivered is consumed. In effect the blood vessels have expanded but this has not resulted in the muscles consuming more oxygen—the limitation is elsewhere. This does not of course mean that this vasodilation could not be physiologically relevant as it will also increase substrate flow to the muscle and the removal of metabolites.

Where the two skating modes do differ is in their quantitative response to these initial effects, in particular between the asymmetry of the two legs. As shown previously (Hesford et al., 2012), the rate of reoxygenation is faster in the left leg than in the right leg in short-track. However, this is not the case in long-track where no significant differences are seen. At the end if the TT the absolute muscle oxygenation is the same in the right and left legs in long-track and this value is identical to that in the left leg in short-track. However, the right leg in short-track is oxygen deprived compared to these three other states.

The reason for this asymmetry can be seen when exploring the oxygenation changes during a single lap. There are high intramuscular forces on the right leg when traveling around the corner in short-track skating, as the skater hangs on that right leg increasing deoxygenation. Consequently, the left leg that is off the ice, or at least carrying minimal weight, reoxygenates in the corners. In long-track speed skating, the NIRS measurements in combination with video captures showed a cyclic pattern of deoxygenation and reoxygenation during the straights. This pattern is consistent with the repetitive cyclic movement pattern that is executed in long track speed skating, consisting of a gliding phase, a push off phase and a swing phase (De Boer et al., 1987; Allinger and Van den Bogert, 1997; Konings et al., 2015). In the gliding phase and push-off phase, blood flow is occluded and deoxygenation occurs, while in the swing phase reoxygenation occurred. In the corners, a stable and consistent pattern in terms of oxygenation was visible in long-track skating: the difference between the legs is maintained and levels of oxygenation change very little. The movement pattern of leg crossovers in long-track speed skating was rhythmic, cyclic and consistent, and no long on-ice or off-ice phases are seen (Figures 5, 7). Therefore, in long-track deoxygenation largely occurred during the straight not the corners and the changes between the two legs were largely symmetrical. In contrast, in short track speed skating, a very clear asymmetry in reoxygenation occurred during the corners, with the left leg reoxygenating when off the ice and the right leg deoxygenating when on the ice throughout the corner (Figures 6, 8).

Although an asymmetry in muscle oxygenation in long-track speed skating was reported before (Born et al., 2014), the current findings expand on these findings. That study reported that the difference in muscle oxygenation between both legs during the course of a 3000 m TT was only a result of a difference in the initial desaturation between both legs directly after the start (Born et al., 2014). In contrast, the present study shows that the asymmetry between both legs during long-track speed skating is also a result of a slightly enhanced reoxygenation rate in the left leg as the TT progressed (Table 1).

Our results agree with other findings in the literature, relating higher intramuscular pressure to hampered oxygenation and subsequent larger dependency on anaerobic energy supplies, exacerbating fatigue (de Ruiter et al., 2007; Katayama et al., 2007; Romer et al., 2007). In terms of the quantity of differences in muscle oxygenation, we found a reduction of more than 10% in TSI% over the whole test compared to baseline during both skating modes. This is of a similar magnitude to a study in fore-arm muscles, where it was demonstrated that a 7% or larger reduction in muscle oxygenation was found to lead to decreased muscle force production (Murthy et al., 2001).

We hypothesized that the clearly demonstrated differences in deoxygenation and reoxygenation between sports would have an impact on perceived fatigue and recovery, in favor of long-track skating. Indeed, higher experienced fatigue after 2 and 4 h were found after the short-track time-trials compared to the long-track time-trials. This is despite the lack of a difference between lactate values, heart rate data and experienced fatigue when measured directly after the two testing sessions. Some clue to possible mechanisms came from the post-exercise 6-s maximal exercise test. Following short-track skating, there was a clear decrease in the post-exercise reoxygenation rate compared to long-track skating. Intriguingly this difference was seen in both legs, i.e., no differences were found between the right and left legs recovery rates even in short track skating. This might indicate that the differences between both legs are too small to affect processes of recovery after speed skating, however, it might also be a consequence of the relatively long time period (±5 min) between the end of training and the post-test. Nevertheless, whatever the detailed mechanism, the present study provides evidence that short-track speed skating is physiologically more demanding than long track speed skating and that this is associated with a demonstrated difference in hemodynamic patterns both during and post-race and a slower recovery from fatigue.

The application of blood flow restriction in high intensity cycling impacted on lactate accumulation and ratings of perceived exertion (Kim et al., 2015). At low exercise intensities, it was found that RPE was not affected by higher pressures in the range between 40 and 90% of arterial occlusion (Loenneke et al., 2016). A key difference in our skating exercises is that high phases of high intra-muscular pressures in the cycle are alternated with phases of unloading, when the leg is in the swing phase. Alternating pressure patterns and their impact on oxygenation and metabolism in human propulsion have not been studied very often in literature. Our work could inform on the mechanism of Kaatsu training: the application of blood flow restriction during low intensity training that has been shown to produce favorable muscle and vascular adaptations exacerbating training effect. This could be of interest for clinical populations (Loenneke et al., 2016).

In contrast to previous reports that focused on only one skating modality, the current study benefitted from using the same sample of subjects, familiar to both skating modalities, for both long-track and short-track speed skating. In agreement with previous separate studies for short-track (Rundell et al., 1997; Hesford et al., 2012, 2013b) and long-track (Foster et al., 1999), oxygenation changes in the current study are best explained by postulating a reduced blood flow to the working muscles throughout the race for both speed skating disciplines. Most likely, this reduced blood flow caused the relatively high lactate values and low oxygen uptake values in this study. This supports the assumption that a reduced blood flow reduces the aerobic capacity of the recruited muscle groups (Rundell, 1996). The current findings also indicate that the relatively low reported maximal oxygen uptake values in previous long-track speed skating studies (Smith and Roberts, 1991; De Koning et al., 1994) are a consequence of the deoxygenation of the working muscles in speed skating.

In terms of performance, traveling around the corners in short-track skating results in different velocity profiles compared to long-track speed skating. The subjects were slower during short-track speed skating when they traveled around the corner standing at one leg only. To regain their loss in velocity, the subjects accelerated when they came out of the corner and during the straight. Consequently, short-track seems to show more variability in velocity in each lap, while the long-track velocity profile seems more continuous (see Figure 1). Fluctuations in velocity were larger in short-track speed skating, and large fluctuations around the mean velocity are associated with higher aerodynamic energy losses (Van Ingen Schenau et al., 1983; De Koning et al., 1999; Hettinga et al., 2011, 2012). In running, a more variable interval training pattern has been shown to be associated with a slower recovery than demonstrated after continuous training (Townsend et al., 2013). In that sense, this finding also confirms that short-track speed skating is the more demanding sports of the two, and might need a longer recovery after exercise and training.

For trainers and coaches it is important to realize that due to this element of occlusion and reduced blood flow as well as the larger fluctuations in speed, physiological load is higher than expected for their athletes during short-track speed skating compared to long-track speed skating. Moreover, training schedules based on physiological variables achieved during other non-occluded sports, such as commonly used (maximal) cycling tests, might not represent the actual physiological load of the training sessions, and trainers should beware of risks of overtraining. The working muscles of the right leg remained relatively more deoxygenated during short-track compared to long-track speed skating. This indicates that the muscles in short-track speed skating are more anaerobic than in long-track speed skating. However, as oxygen extraction does not change with time even when oxygenation recovers in long-track skating, it is not clear that this has adverse metabolic consequences at least in terms of oxygen metabolism. Nevertheless, the present study shows that a short-track training session leads to higher experienced fatigue in the hours after the training session and seems to need a longer recovery period compared to a similar long-track session.

Our findings emphasize the importance of pacing in speed skating. In particular the ability to maintain the optimal technical skating characteristics (respectively, small knee-, trunk-, and push-off angle) during the whole race, despite processes of fatigue, seemed to be crucial in elite (long-track) speed skating (Hettinga et al., 2011) and lead to different pacing behavior in speed skating compared to for example cycling (Stoter et al., 2016). Less explosive pacing strategies are chosen in speed skating, to make sure not to collapse at the end of the race. The present study showed that the crouched skating position indeed seemed to lead to a physiological disadvantage: the deoxygenation of the working muscles, which may cause an earlier onset of fatigue. As in the short-track time-trials in the present study, a relatively high decrease in velocity toward the end of the race was demonstrated, also in short-track speed skating the ability to maintain optimal technical skating characteristics at the end of the race seems crucial (Konings et al., 2016). In particular in actual competition this will be of importance, as this is performed in heats, competing against opponents (Konings et al., 2016; Noorbergen et al., 2016).

In conclusion, the current study showed that the patterns of reoxygenation and deoxygenation in the working muscles during a race are different for long-track and short-track speed skating. This seems to be a result of the high intramuscular forces on the right leg when traveling around the corner in short-track skating: athletes are usually standing on the right leg throughout the corner, that thereby remains deoxygenated for a long period where the left leg reoxygenates in the same phase when off the ice. In long-track skating, a more symmetric profile was demonstrated, in which over the straights the cyclic pattern of stance phase, push off phase and swing phase was clearly visible in terms of oxygenation. In the corners, the rhythmic leg crossovers resulted in a rather stable value for deoxygenation, that was larger for the right leg compared to the left leg. The results demonstrated that short-track speed skating seems to be the more physiologically demanding sport. This subsequently led to higher experienced fatigue directly after the exercise, as well as to longer periods of recovery needed in the hours after the short-track compared to long-track speed skating. These results can help coaches and athletes in optimizing their training protocols, and provide us with more insights into the mechanistic physiological principles relevant for elite athletes in different sports, and on how technical factors are impacting on those.

The measurements were conducted at InnoSportLab Thialf, with portable equipment available from the laboratory of the University of Essex. FH, MK, and CC all contributed to conception and design of the work. MK collected data. FH, MK, and CC were involved in analysis and interpretation. MK, FH, and CC wrote the paper. All authors have approved the final version of the manuscript, agree to be accountable for all aspects of the work in ensuring that questions related to the accuracy or integrity of any part of the work are appropriately investigated and resolved, and all persons designated as authors qualify for authorship, and all those who qualify for authorship are listed.

The work described here was funded in part by grant EP/F005733/1 from the UK Engineering and Physical Sciences Research Council (EPSRC). There were no additional funding sources for this study. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.