How reliable is musculoskeletal modelling in producing replicable outputs?
A study aiming to understand how different methods of estimating muscle structure influence our interpretations of limb function
Musculoskeletal models of a human (left) and of AL 288-1, also known as Lucy (right) | Image: Wiseman ALA, Charles J, Hutchinson JR. 2024. Static versus dynamic muscle modelling in extinct species: a biomechanical case study of the Australopithecus afarensis pelvis and lower extremity. PeerJ 12:e16821 http://doi.org/10.7717/peerj.16821
A series of simulations have been carried out to see if the well-known 'Lucy' could stand on one leg and how much weight she would have been able to bear when doing so.
"We conducted a suite of static simulations in which we applied a ground reaction force vector to the foot and increased the amount of force until simulation failure or ‘limb collapse’. In this way, we were able to test if Lucy could stand on a single, erect limb and to see how many multiples of body weight she could do so."
Musculoskeletal models of a human (left) and of AL 288-1, also known as Lucy (right) | Image: Wiseman ALA, Charles J, Hutchinson JR. 2024. Static versus dynamic muscle modelling in extinct species: a biomechanical case study of the Australopithecus afarensis pelvis and lower extremity. PeerJ 12:e16821 http://doi.org/10.7717/peerj.16821
Musculoskeletal models of a human (left) and of AL 288-1, also known as Lucy (right) | Image: Wiseman ALA, Charles J, Hutchinson JR. 2024. Static versus dynamic muscle modelling in extinct species: a biomechanical case study of the Australopithecus afarensis pelvis and lower extremity. PeerJ 12:e16821 http://doi.org/10.7717/peerj.16821
The study delves into the mechanics of muscle function in the hominin Australopithecus afarensis (specimen AL 288-1, affectionately nicknamed Lucy), aiming to understand how different methods of estimating muscle structure influence our interpretations of limb function.
The team digitally reconstructed the muscles and tested their ability to support an upright, single-support limb posture.
The force a muscle generates is dependent on muscle architectural parameters, in which fibre length, pennation angle and tendon elasticity all influence force production and subsequent body movement.
However, muscles are rarely preserved in the fossil record and instead we are left with the bare bones. A scientist cannot simulate the locomotion of an extinct individual without first reconstructing the missing soft tissues of the limbs.
Last summer, Wiseman (2023) published the first 3D muscle reconstruction of Lucy, but this study only reconstructed the muscle paths – that is, the space and path that a muscle occupies within a body.
The model creation progress in which 3D muscles were created for Lucy for all muscles in the pelvis and leg, leading to the musculoskeletal model creation in the biomechanical software OpenSim | Image: Ashleigh Wiseman
To simulate movement, researchers next need to estimate the architectural parameters of each muscle. Yet, there are multiple ways to do this.
The team decided to test the influence of these different methods on subsequent model performance by creating seven identical musculoskeletal models of Lucy and by only changing the input muscle parameters.
Wiseman and colleagues focused on 36 muscles in the pelvis and lower limb. These models used different ways to estimate muscle parameters, such as fibre length, pennation angle, and tendon slack length.
Two types of muscle models were compared: a simpler 'static' model and a more complex 'dynamic' model that accounted for elastic tendons and variable force-length-velocity properties of fibres.
In the simulations, vertical and side-to-side (i.e., mediolateral) ground reaction forces were gradually increased until the limb collapsed (simulation failure).
Interestingly, all Lucy models showed similar muscle activation patterns, but the maximum vertical force the limb could withstand varied amongst the models. Static muscle models struggled to support the weight, whilst dynamic models performed better, suggesting that incorporating more realistic muscle properties improves the strength of the model – although the latter does come at a modelling computational expense.
Comparing our results with a human model, we found that both species required mostly similar muscle activations with just a few exceptions to sustain single-limb support under maximal forces. This study highlights the range of outcomes possible when modelling an extinct individual, emphasizing the importance of considering different factors in estimating muscle function, especially when assessing model strength.
"In this image, we see how the muscles of the pelvis and lower limb are responding to these static simulations – these are simulated muscle activations of Lucy and a human.
We created seven different models of Lucy, each with muscle architectural parameters estimated using a different method.
We conducted static simulations on all models and found that muscle activation patterns were slightly different between the seven different Lucy models (panels A-G), whereby some muscles were maximally recruited (dark blue) and other muscles were inactive (grey).
The more muscles that are activated (and especially with higher activations) will be more metabolically expensive to maintain the erect posture.
We also found that some models were weaker than others. For example, we used the 2023 muscle reconstructions of Lucy (Wiseman 2023) which are here called the ‘3D variant’.
These 3D muscle reconstructions were much stronger and capable of supporting greater amounts of force on a single, erect limb than that of the other models (3.6 x body weight (A) versus the arithmetic variant at 1.8 x body weight (B)).
Most importantly, we found that tendon elasticity must be included or the model(s) was found to be too weak (tendon elasticity included in the dynamic-muscle models, E-G).
If we compare this to a human (H), we find that different muscles in the limb are required to maintain the same, erect-style posture in Lucy and the human. Notably, no hip adductors are required, with fewer muscles required in the foot."
Dr Ashleigh Wiseman
Simulated muscle activations of Lucy and a human | Image: Wiseman ALA, Charles J, Hutchinson JR. 2024. Static versus dynamic muscle modelling in extinct species: a biomechanical case study of the Australopithecus afarensis pelvis and lower extremity. PeerJ 12:e16821 http://doi.org/10.7717/peerj.16821
What does this mean for future studies?
A researcher could create two identical models of an extinct individual. By simply adjusting the input architectural parameters and deciding whether to include or exclude an elastic tendon, they can produce one version of the model that is weak.
On the other hand, they can also create a second version capable of supporting the body with multiples of body weight on a single limb. The latter model could go on to simulate activities like running and jumping, whilst the former model may be unable to simulate a basic walk.
In one scenario, Lucy would be an apt biped, capable of running and jumping. In the other, Lucy would have high metabolic costs for a simple walk, and could lead to the inference that she was not a habitual upright walker.
The researcher must then be aware of the implications when selecting the best method to estimate muscle parameters - the choice of parameter estimation method should be carefully selected alongside research requirements.
The paper is published in PeerJ here
Wiseman ALA, Charles J, Hutchinson JR. 2024. Static versus dynamic muscle modelling in extinct species: a biomechanical case study of the Australopithecus afarensis pelvis and lower extremity. PeerJ 12:e16821 https://doi.org/10.7717/peerj.16821
This research builds upon the paper 'Three-dimensional volumetric muscle reconstruction of the Australopithecus afarensis pelvis and limb, with estimations of limb leverage' published in 2023.
Dr Ashleigh Wiseman collaborated with John R Hutchinson – Royal Veterinary College and James Charles – University of Liverpool on this paper.
The research was supported by a Leverhulme Trust Early Career Fellowship (grant number: ECF-2021–054) and by the Isaac Newton Trust, University of Cambridge.
Dr Ashleigh Wiseman is currently a Leverhulme/Isaac Newton Trust Early Career Fellow at the University of Cambridge investigating hominin evolutionary biomechanics. Her main research interests are evolutionary biomechanics, musculoskeletal modelling, and reconstructing locomotion from fossilised remains.
During her current fellowship based in Cambridge, she is constructing 3D musculoskeletal models of hominins and performing biomechanical simulations to test locomotory capabilities in the hominin lineage.
To keep up to date with her research, see the Walking Hominins website.
Published 31 January 2024
The text in this work is licensed under a Creative Commons Attribution 4.0 International License