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The unique structure of the human foot is seen as a crucial adaptation for bipedalism. The foot's arched shape enables stiffening the foot to withstand high loads when pushing off, without compromising foot flexibility. Experimental studies demonstrated that manipulating foot stiffness has considerable effects on gait. In clinical practice, altered foot structure is associated with pathological gait. Yet, experimentally manipulating individual foot properties (e.g. arch height or tendon and ligament stiffness) is hard and therefore our understanding of how foot structure influences gait mechanics is still limited. Predictive simulations are a powerful tool to explore causal relationships between musculoskeletal properties and whole-body gait. However, musculoskeletal models used in three-dimensional predictive simulations assume a rigid foot arch, limiting their use for studying how foot structure influences three-dimensional gait mechanics. Here, we developed a four-segment foot model with a longitudinal arch for use in predictive simulations. We identified three properties of the ankle-foot complex that are important to capture ankle and knee kinematics, soleus activation, and ankle power of healthy adults: (1) compliant Achilles tendon, (2) stiff heel pad, (3) the ability to stiffen the foot. The latter requires sufficient arch height and contributions of plantar fascia, and intrinsic and extrinsic foot muscles. A reduced ability to stiffen the foot results in walking patterns with reduced push-off power. Simulations based on our model also captured the effects of walking with anaesthetised intrinsic foot muscles or an insole limiting arch compression. The ability to reproduce these different experiments indicates that our foot model captures the main mechanical properties of the foot. The presented four-segment foot model is a potentially powerful tool to study the relationship between foot properties and gait mechanics and energetics in health and disease. Author summary: During every step, the foot absorbs the shock as it strikes the ground, supports the body weight, and transfers calf muscle forces to the ground to push off towards the next step. Although experiments have shown that foot structure affects both foot and whole-body movement, a deep understanding of the role of different muscles, ligaments, and other tissues in the foot is still lacking. Examining the relation between alterations in individual foot properties, e.g. muscle strength or arch height, and walking is challenging in experiments, but can be done in model-based simulations. Yet, existing models that are suitable for simulating walking oversimplify the foot. Here, we present a new, more detailed foot model. Simulations with our model captured many features of human walking not captured by simulations with simple models. We found that the stiffness of the foot-ankle complex and especially the ability to stiffen the foot had a large effect on the walking pattern. A reduced ability to stiffen the foot resulted in simulated walking patterns resembling those observed in patients with flexible feet. In the future, we will use our model to investigate how foot deformities alter walking in patients and to design personalized treatments. [ABSTRACT FROM AUTHOR]