Knee extensor muscle properties

Regarding the knee extensor muscle morphology, we found an increase in the fascicle angle and muscle thickness from the EP to LP. These alterations point towards a radial adaptation of the vastus lateralis muscle indicative of muscle hypertrophy. As there are no comparable studies investigating potential factors leading to muscle growth in pregnant women or gestating animals, in our research article (chapter 5) we discussed several fac-tors afecting muscle architecture.

One of these factors is the increased BM as the increase in muscle thickness of 9 % in the LP was similar to the BM increase of 10 %. However, our statistical analysis demons-trated that the increased BM during pregnancy is unlikely to be the only factor triggering adaptation of the muscle.

Alternatively, we can also assume that conditions during pregnancy such as changes in the endocrine system can foster skeletal muscle hypertrophy since improved regenerative processes in skeletal muscles during pregnancy have been described, at least in animal models (Falick Michaeli et al., 2015). Research on postmenopausal monozygotic twins re-vealed that skeletal muscles do more likely respond to mechanical loading when a hormone therapy with sevenfold levels of estrogen is given (Ronkainen et al., 2009). Thus, pregnan-cy with 40 times higher levels of estrogen compared to non-pregnant women (Martin and Behbehani, 2006) may be a favorable metabolic condition which promotes adaptation to training stimuli and an increased BM.

In our research article, we further suggested that water retention in the intracellular space may have caused a thickening of the muscle fbers resulting in a measurable increase in

muscle thickness. However, again no correlation has been found between the parameters.

In conjunction with the body water assessment, it needs to be mentioned that we were not able to consider the content of the amniotic fuid. This may have afected our results.

Another limitation is that we did not control the fuid intake in our pregnant participants.

However, our results demonstrate that variation in the ICW and ECW of a pregnant wo-man was rather small at each measurement time point. In contrast, diferences between pregnant women were much larger (Figure 10A, B) which is why we would not expect an efect of hydration diferences on the muscle properties.

We speculate that changes in the muscle architecture are attributed to pregnancy-related changes in body posture. It seems conclusive that alterations in body shape during pre-gnancy are accompanied by adaptive mechanisms of the body to maintain balance. As a compensatory efect for the increased anterior load caused by the growing fetus, pre-gnant women in the LP have been found to adapt their body posture by shifting the CoM 4.0 - 6.6 mm in the posterior direction during upright standing (Opala-Berdzik et al., 2010; Bey et al., 2018). During upright standing the muscle fbers of the knee extensors are lengthened beyond the optimum of the force-length relationship (Houtz et al., 1957).

An adaptive increase in the hip angle in pregnant women might further lengthen muscle fbers impairing the unfavorable condition for force production. Thus, the muscle may rely on increased muscular activity to compensate for the impairment in force producti-on. An increased muscle activity could then contribute to radial adaptation in the vastus lateralis muscle. However, if the muscle is continuously active at a diferent part of the force-length relationship, a change in muscle FL is likely (Sharifnezhad et al., 2014; Herzog et al., 1991). As we did not fnd any changes in muscle FL, it appears that this stimulus is not sufciently high to promote a longitudinal adaptation.

It can be summarized that the pregnancy-induced change in muscle architecture is un-likely to be attributed to a single factor. Further research is necessary to clarify which pregnancy-associated combination of changes in the endocrine system, BM, and body composition contributes to this morphological alteration.

Apart from changes in muscle morphology the functional properties of the knee extensors

did not change during pregnancy. The mechanism underlying this efect needs to be fur-ther investigated as our observed increase in muscle thickness may point towards muscle hypertrophy and, thus, to an increased force capability. However, the absolute muscle strength remained constant from the EP to LP and did not difer from the postpartum status or that of the non-pregnant women. This contrasts to fndings that reported the upper extremities becoming weaker during pregnancy (Atay and Basalan Iz, 2015). We argue that the lower extremities are subjected to a diferent adaptation process than the upper extremities.

The constant strength performance of the knee extensors despite increased muscle thick-ness during pregnancy may be attributed to a neural adaptation that is manifested as an increased coactivation of the knee fexors. However, we detected a marginal but not statistically signifcant increase (+ 1.5 %) in the coactivation of the knee fexors from the EP to LP (EP: 8.9 ± 4.2 %, LP: 10.4 ± 5.4 %, p = 0.259, unpublished data, see appendix A.1). Thus, the constant strength performance is unlikely to be the result of an increased force capability of the antagonists.

It is possible that a reduced physical activity level may have sightly reduced the muscle strength performance as we did not control the physical activity level in our participants (Atay and Basalan Iz, 2015). However, even when our pregnant participants were less active in the LP compared to the EP, this did not contribute to a meaningful loss in strength as has been shown for the upper extremities (Atay and Basalan Iz, 2015).

A slight reduction in the muscle strength might also be attributed to a pregnancy-related fast-to-slow fber type shift (Bisch et al., 2006). As primarily fast fbers are involved in force production during the ramp contractions (Gollnick et al., 1974), a reduced number of fast fbers may have diminished the force capabilities of the knee extensors in our par-ticipants.

Psychological factors such as not feeling confdent enough to perform a maximum isome-tric ramp contraction on the dynamometer in the LP might also have afected the force capabilities of our participants. Furthermore, anxiety may also have infuenced the results as increased levels of axiety are a common side efect during pregnancy (Dennis et al., 2017; Nakic Rados et al., 2018).

Finally, muscle strength during pregnancy may have been partly afected by a calcium defciency which is associated with the high prevalence of cramps in the calves during pregnancy (Ireland and Ott, 2000; Hammar et al., 1981). As reported by Herzog et al.

(2015), calcium is essential for force production as it regulates stability and stifness of titin upon activation. If the calcium level was reduced in our pregnant participants, this may have led to a unfavorable stretch state of titin impairing the strength level.

Thus, there is evidence to suggest that pregnancy provides conditions that may be favo-rable for retaining muscle strength in the lower extremities. Both the increased BM and changes in the hormonal milieu are likely to trigger muscle growth. In contrast, changes in the mental state during pregnancy seem most likely to counteract the improved muscle force capabilities during contraction. However, from an evolutionary perspective a reduc-tion in the lower extremities’ strength seems to be unlikely. This may explain why we did not detect any statistical diferences in muscle strength between pregnant fallers (six from 19 women, 32 %) and non-fallers (p > 0.05, unpublished data, see appendix A.2).

Nevertheless, when normalizing muscle strength to BM we detected a decrease in the LP.

Future studies to investigate whether in the event of balance perturbations the legs are actually strong enough to maintain balance are needed. As balance ability has frequently been shown to decrease during pregnancy (Oliveira et al., 2009; Inanir et al., 2014; Bey et al., 2018), while pregnant women have to meet the requirements for a greater physical demand due to the increased BM, a loss in the lower extremities’ muscle strength relative to BM seems likely to be a crucial factor for the increased risk of injury during pregnancy.

Summing up, the lack of absolute muscle strength improvements despite the increased physical demand during pregnancy suggests an increased risk of injury in pregnant wo-men. However, at the same time pregnancy is likely to provide favorable metabolic con-ditions promoting muscle adaptation to training and mechanical loading (Falick Michaeli et al., 2015; Rundgren, 1974; Delmonico et al., 2009). Pregnancy appears to be a suitable period of a woman’s life to be physically active and to continue exercising. Thus, espe-cially in the case of sedentary pregnant women who do not meet the minimum national recommendation of 30 minutes moderate physical activity a day (Mudd et al., 2009; Dob-son et al., 2012; Gaston et al., 2012; Gjestland et al., 2013; Juhl et al., 2012; Zhang and

Savitz, 1996; Sternfeld et al., 1995) the positive efect of pregnancy on muscle adaptation may be a possible key motivator for their participation in a physical activity program.