Muscle oxygenation during different muscle actions
In respect of a possible objective distinction of the two isometric forms, oxidative variables were compared between HIMA and PIMA. No significant differences were found regarding the maximal deoxygenation, saturation slopes, saturation level at global minimum of rHb and time to leveling off into a saturation steady state. This indicates that these variables cannot discriminate HIMA and PIMA on the basis of the small sample size investigated here. Regarding the saturation slopes, this is in accordance with Booghs et al. (2012) . They utilized the near infrared spectroscopy technique (NIRS), which is comparable with our used white light spectrometry [13, 17]. To our best knowledge, the remaining oxidative variables were considered for the first time regarding the comparison of HIMA and PIMA.
As generally accepted, IMAs should restrict the capillary blood flow due to the high intramuscular pressure already at low intensities [19, 20], but the occlusion threshold might vary between individuals and muscles . In contrast, auxotonic contractions possibly support oxygen and blood supply due to the reduced muscle tension during the lowering phase. Comparisons between HIMA and Twitch did not show any significant differences in the analyzed variables of the present study. Thus, the curve characteristics of SvO2 and rHb seem not to be influenced by twitches. Otherwise, the minimal motion of the limb caused by rapid muscle contractions might not be enough to change the muscle oxygenation and blood filling of microvessels significantly.
Previous studies revealed that increases in capillary blood filling (rHb) [13, 15, 30, 32–34] and homeostatic steady states in the oxygen saturation [11, 13–16] are possible already during IMAs without twitches. These data suggest a probable maintenance of capillary blood flow. Recently, it was discussed that this could be achieved by the anatomical placement of capillaries within muscle tissue and the oscillatory behavior of muscle fibers during IMA .
The behavior of oxygen saturation and blood filling can be differentiated by two types , indicated by the SvO2 level at global minimum of rHb . We have suggested a threshold of ~59% whereby values above this threshold are associated with type I (parallel behavior of SvO2 and rHb) . In contrast, if the saturation decreases below that threshold, rHb starts to increase which is related to type II. The distribution of type I and type II assigned measurements reflects a qualitative behavior of the measured parameters SvO2 and rHb. Because the extents of deoxygenation and SvO2 levels at global minimum of rHb did not differ significantly between HIMA and PIMA as well as not between HIMA and Twitch the amount of type I and type II assigned measurements are, as a consequence, not significantly different (Table 2). Thus, the occurrence of type I and type II seems to be independent of the isometric task.
At last, a high and significant negative correlation (r = –0.88) between the SvO2 decrease and SvO2 slope was found over all measurements. The greater the deoxygenation, the steeper the drop. This is plausible by considering the similar TSSs found over all trials.
Time to task failure during different muscle actions
The TTF as performance variable in the presented study, appeared to be longer during PIMA compared
to HIMA. However, the difference (5.51 ± 18.37 s) was not significantly different (methodological limitations see below). This is in line with the results of other studies which also examined the TTF of the biceps brachii muscle  or elbow flexor muscles  with similar settings for HIMA and PIMA at the same intensity (60 % of the MVIC) and same forearm position (horizontal). However, during lower intensities (≤ 30% of the MVIC) of the elbow flexor muscles, the TTF of HIMA seems to be significantly shorter than the TTF of PIMA [3, 4, 9, 11]. If the IMA is, by contrast, performed in a vertical forearm position or during muscle activities at 45 % and 60 % of the MVIC, the TTF was found to be similar between tasks [3, 4]. This indicates that both, the intensity and forearm position, influences the performance of elbow flexor muscles. Regarding the first dorsal interosseous muscle, Maluf et al. (2005) found differences in the TTF between HIMA and PIMA at 20% of MVIC (TTF HIMA < TTF PIMA) but not at 60% .
However, the order of tasks in the presented study might have influenced the TTFs in favor of HIMA. In case HIMA was performed at first (in 6 of 9 cases), the relation of PIMA/HIMA amounted to ~1.17 ± 0.68; in case PIMA was performed at first (3 of 9 cases) the relation PIMA/HIMA was ~1.61 ± 0.53.
Additionally, it should be mentioned again that holding tasks as performed in the present and the other mentioned studies does not imply a pure IMA. In general, muscles show slight oscillations during isometric muscle actions [2, 10, 36–40]. Thus, minor muscle shortenings and lengthenings are present. In case of weight holding, slight motions around the given joint angle have been accepted. The tolerance in different studies ranged from 2° – 10°. This also includes minor concentric contractions to lift the weight back to the starting angle position. Such muscle actions interrupt a pure HIMA and it was hypothesized that the muscle action could be switched to a PIMA after minor concentric contraction has been executed . In the present study, little concentric contractions were documented during four of nine HIMAs in the HP group. This also might have biased the result and might explain why the trend of a longer TTF during PIMA did not reveal statistical significance. Different experimental procedures were applied by Schaefer and Bittmann (2017, 2021) examining elbow extensors. The methods might repeal the above-mentioned problem for HIMA during weight holding [2, 10]. In the first study, a pneumatically driven measurement system was used to realize HIMA and PIMA  and in the second one, an interaction between two subjects comparable with arm wrestling . The former study controlled for a concentric contraction (failure criterion) and the latter one facilitates the performance of the tasks described above during PIMA (acting part) and HIMA (reacting part), since the holding subject just had to adapt to the input of the pushing partner. In both studies, the forearm was positioned vertically and significant differences regarding the TTF of elbow extensors at 80% of the MVIC (TTF HIMA < TTF PIMA) were found. Thus, not only the intensity of muscle activity and positioning but also the examined muscle and experimental procedure might play a role.
In this regard, the present study revealed that intermittent muscle twitches during holding task extended the TTF significantly (~10 s). It was assumed that the muscle twitches induce a switch of the muscle action from reacting (HIMA) to acting (PIMA) during the isometric phases. Considering PIMA and Twitch together and comparing theses to all HIMA trials, the TTF still differs significantly. As discussed above, the behavior of SvO2 and rHb in conjunction with variables 1. – 5. were similar between tasks. Thus, the reason for a longer TTF during PIMA (incl. Twitch) seems not to be a different oxygen or blood supply. This is further supported by the analyzed correlations. Independently of the isometric task, very low to low, non-significant correlations between the TTF and the SvO2 decrease as well as the TTF and SvO2 slopes (r = –0.13 and 0.31, respectively) were found. This result indicate a reasonable independence of the TTF from the deoxygenation as long as SvO2 levels off into a homeostatic steady state. Booghs et al. (2012) also conclude that the decrease in muscle oxygenation is not a significant predictor of the TTF although they did not rule out its contribution to muscle fatigue . Moreover, an enhancement of muscle oxygenation (oxygen half time recovery) as revealed in rock climbers during fatiguing forearm muscle contractions at 60% of the MVIC by New Zealand blackcurrant extract did not affect the TTF . Consequently, there must be other factors why TTF was found to be extended during Twich. One possibility should be shortly discussed in the following.
It was previously suggested that the neuromuscular control could play a decisive role in the distinction of HIMA and PIMA . This was based on the assumption that PIMA is closer to the motor control processes of concentric contraction, and HIMA might show a proximity to eccentric muscle action . Eccentric muscle actions involve more complex control strategies compared to concentric muscle actions possibly resulting in a greater central fatigue [2, 42–46]. Thus, the more complex control processes suggested for HIMA might be one reason for the often found shorter TTF compared to PIMA [2–4, 9–11]. The significantly longer TTF during Twitch in the present study, could support the assumption of a switch from HIMA to PIMA.
Limitations regarding the used measurement technique, not examined skinfold thickness and different determinations methods of the MVIC have been described previously [13, 17].
Some more limitations need to be emphasized especially regarding the TTF. During HIMA and Twitch, the loading was stopped as soon as the elbow angles exceeded 90° for more than 2 s and if twitches were not visible anymore. No objective instrument (e.g., goniometer) was used to reach highest accuracy during control. The measurement error could have influenced individual trials but on group level the random error ought to be leveled out. However, the amplitude and velocity of twitches might have varied between trials. TTFs could also be influenced by the measurement position, as discussed before. Subjects used a horizontal forearm position in all measurements but changed from standing to sitting position between HIMA and PIMA. Thus, activation of trunk stabilizing and postural muscles were different between tasks with an expected higher activity during HIMA (standing). However, the biceps brachii muscle had to be equally activated with 60% of the MVIC within the muscle chain to maintain a 90° elbow flexion. If the trunk stabilizing muscles were not strong enough, the whole body would be bend forward. This was prevented by the counter bearing between the upper arm and thorax. Furthermore, due to the exclusion of measurements (see statistical analysis) the sequence of tasks of HP-group were not balanced anymore (n = 6 HIMA first; n = 3 PIMA first). Thus, effects of fatigue might have influenced the results as discussed above. In contrast, the order was nearly balanced in HT group (n = 5 HIMA first, n = 4 PIMA first).