The purpose of this study was to identify the situations under which HAM can act as knee extensors. To achieve this purpose, a musculoskeletal model-based computational simulation was designed to quantify the HAM function on the knee joint for the two main contributing factors: the effect of changing the posture and the effect of the foot contact condition. This study revealed three main findings. 1) HAM can act as knee extensors that have contact with the ground (CG) condition but also non-contact with the ground (nCG) condition. 2) The HAM function changes depending on the posture. 3) Finally, the range of the hip joint that HAM can act as knee extensors (RANGEHAM) was expanded to the CG condition from the nCG condition.
The first point to be discussed is the posture dependent HAM function. As mentioned earlier, posture is an important contributing factor. This is because the moment arm ratio and the intersegmental dynamics change depending on the posture. Many researchers have claimed that the biarticular muscle moment arm ratio of the proximal joint to the distal joint is the largest contributor to the counterintuitive function [9, 17, 19, 21]. In addition, some previous studies demonstrated that changing the muscle function was coursed by the posture-induced changes in the effect of the intersegmental dynamics [14, 15, 18, 22]. Despite the situation described above, to the best of our best knowledge, a comprehensive study currently does not exist in terms of how the HAM function changes depending on the posture. This is the first study to show in detail the change in the HAM function on the knee joint depending on the posture.
The next discussion deals with the interaction between the moment arm ratio and the posture-dependent intersegment dynamics. Some researchers believe that the moment arm ratio has the largest impact on the counterintuitive biarticular muscle function [9, 17, 19, 21]. However, the results of the present study denied this notion. A typical example is shown in Fig. 4. Figure 4 indicates that the two different postures have the same moment arm ratio. Even though both postures have the same moment arm, the HAM function is different. This is evidence that the HAM function is determined by the interaction between the moment arm ratio and the intersegmental dynamics. We can conclude that the counterintuitive HAM function arose from the interaction between the moment arm ratio and the intersegmental dynamics, and that the interaction between these factors dynamically changes depending on the posture.
Thirdly, the effect of the foot contact conditions is discussed. In the present study, we confirmed that RANGEHAM was expanded to the CG condition from the nCG condition (Fig. 3). This finding indicates that whether the foot is in contact with the ground has a large impact on determining the HAM function. Previous studies have reported that the ground reaction force has a large effect in terms of determining the muscle function during movement [15, 17, 23]. Frigo et al. [15] investigated the alternation of the HAM function under different CG conditions by using a computational simulation approach. As a result, they clearly showed that the effect of the knee flexion that is induced by HAM is dramatically reduced while in contact with the ground conditions in comparison to the nCG condition. These results are consistent with our results. When considering these observations, we concluded that the exertion of the counterintuitive function of HAM is facilitated by the foot contacts on the ground. This finding suggests that the effect of the ground reaction force must be considered when investigating the muscle function.
We discussed the clinical implications of our findings. This study identified a situation in which HAM was able to act as the knee extensor. Previous studies have reported that the capacity to generate knee extension is one of the most important factors in predicting movement performance, such as gait velocity [1–3]. In addition, it is well known that weakness of the quadriceps muscle causes abnormal knee joint movement during walking, especially in the stance phase of walking [4, 5]. Hence, in the rehabilitation region, therapists often try to improve this matter by enhancing the patients' quadriceps muscle. However, it is impossible for some patients, such as stroke patients with severe motor paralysis on the quadriceps muscles and patients who have undergone partial or total quadriceps resection to enhance the strength of the quadriceps muscle [6–8]. Figure 5 shows the knee angular acceleration that is induced by HAM under the CG condition with the hip and knee joint angles during the stance phase of walking, which was divided from Winter’s data [31]. As demonstrated, the knee angular extension acceleration was exerted by HAM in all stance phases of walking. This indicates that HAM can act as knee extensors during the stance phase of walking. Therefore, our findings suggest that HAM can be used as compensatory movement strategy for patients with a reduced capacity to generate knee extension if the patients have enough HAM strength. We believe that our findings are meaningful in the biomechanics region and the rehabilitation region.
This study successfully provided a theoretical framework for how and when the counterintuitive HAM function is produced. Indeed, the results of the present study can explain the wide range of findings in previous studies [11, 12, 14, 15, 17–20]. However, there are some findings from the present study that we were not able to explain well. Thelen et al. investigated the HAM function during the terminal swing and loading response while walking by using electrical stimulation. As a result, Thelen et al. identified that HAM acted as the knee flexor during both phases [16]. According to the results of the present study, HAM should act as the knee extensor during the loading response while walking. There is a potential explanation why this gap occurred. We ignored the HAM function in the frontal and transverse planes, even though it has a three-dimensional moment arm [39]. The muscle action in the frontal and transverse plane has been demonstrated to contribute to the motion in the sagittal plane [40]. Additional studies are necessary to obtain a better understanding of this matter. However, it was determined that an appreciation of the mapping between the interpretations that are drawn from simple and complex models is needed to advance the understanding of movement mechanics [41]. Therefore, we believe our study provides useful information about determining the HAM function because we can clearly show the relationship between the main contributing factors while exerting the counterintuitive HAM function in the sagittal plane with a simple model.
In the present study, we investigated the independent action of HAM on the knee joint. This study did not consider the co-contraction effect of HAM and the quadriceps. Some studies have suggested that the co-contraction effect of HAM and quadriceps muscles is necessary to determine the HAM function [9, 10, 42]. However, the present study revealed that HAM can act as knee extensors without co-contraction. Frigo et al. investigated the co-contraction effect of HAM and the quadriceps muscle and they showed that it enhances the hip extensor effect of HAM and it reduces the knee flexor effect of HAM [15]. Unfortunately, only a few studies have investigated how much impact the co-contraction of HAM and the quadriceps muscles has in determining the HAM function. Therefore, further studies are necessary to identify the co-contraction between HAM and other muscles.
In conclusion, we have demonstrated that HAM can act as knee extensors. Our findings suggest that HAM might compensate for the capacity to generate knee extension during movement as knee extensors. In addition, our findings contribute to establishing the theoretical framework of how and when the counterintuitive function of HAM is exerted.