How foresight might support the behavioral flexibility of arthropods

The small brains of insects and other invertebrates are often thought to constrain these animals to live entirely in the moment'. In this view, each one of their many seemingly hard-wired behavioral routines is triggered by a precisely defined environmental stimulus configuration, but there is no mental appreciation of the possible outcomes of one's actions, and therefore little flexibility. However, many studies show problem-solving behavior in various arthropod species that falls outside the range of fixed behavior routines. We propose that a basic form of foresight, the ability to predict the outcomes of one's own actions, is at the heart of such behavioral flexibility, and that the evolutionary roots of such outcome expectation are found in the need to disentangle sensory input that is predictable from self-generated motion versus input generated by changes in the outside world. Based on this, locusts, grasshoppers, dragonflies and flies seem to use internal models of the surrounding world to tailor their actions adaptively to predict the imminent future. Honeybees and orb-weaving spiders appear to act towards a desired outcome of their respective constructions, and the genetically pre-programmed routines that govern these constructions are subordinate to achieving the desired goal. Jumping spiders seem to preplan their route to prey suggesting they recognize the spatial challenge and actions necessary to obtain prey. Bumblebees and ants utilize objects not encountered in the wild as types of tools to solve problems in a manner that suggests an awareness of the desired outcome. Here we speculate that it may be simpler, in terms of the required evolutionary changes, computation and neural architecture, for arthropods to recognize their goal and predict the outcomes of their actions towards that goal, rather than having a large number of pre-programmed behaviors necessary to account for their observed behavioral flexibility.