To be able to perform swift and powerful movements, ant mandible closer muscles are composed of two subpopulations of muscle fibres: fast fibres for rapid actions and slow fibres for forceful biting. All these fibres attach to a sturdy and complex apodeme which conveys force into the mandible base. Fast muscle fibres attach directly to the apodeme. Slow fibres may attach directly or insert at individual thin filament processes of the apodeme which vary in length. Comparisons of different ant species suggest two basic principles underlying the design of mandible muscles. (1) Ants specialized for fast mandible movements generally feature long heads which contain long fast muscle fibres that attach to the apodeme at small angles. Their muscles comprise only a few filament-attached fibres and they maximize speed of action at the expense of force output. (2) Ants performing particularly forceful mandible movements, such as seed cracking, rely on many short parallel muscle fibres contained within a broad head capsule. Their slower muscles incorporate a large proportion of filament-attached fibres. Two simple models explain how the attachment angles are optimized with respect to force and velocity output and how filament-attached fibres help to generate the largest power output from the available head capsule volume.

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