Complex Systems Are More than the Sum of Their Parts: Using Integration to Understand Performance, Biomechanics, and Diversity.

Organisms are comprised of many interacting parts, and an increased number or specialization of those parts leads to greater complexity and the necessity for increased integration (the ability of those parts to perform together and maintain a functioning organism). Although this idea is widely recognized among biologists, organisms are more tangibly studied when those parts are considered independently. This reductionist approach has successfully advanced our understanding of organisms' performance. However, performance of one system might (or might not) be dependent on performance of another system to achieve a relevant outcome, and the mechanism of this dependence is poorly understood. We synthesize the concepts of complexity and integration and discuss their application in a biomechanical context. Capture of prey by predatory fishes is used as an example to highlight the application of these ideas. We provide a theoretical framework for future hypotheses of integration and predict an "integration space" for fishes that is then populated with data extracted from the literature. Additionally, using the kinematics of prey-capture in two species of sculpin (Scorpaeniformes: Cottidae), we show that species exhibit multivariate integration in distinct ways, and that these differences add additional insight into ecological divergence that would not be apparent by considering systems independently. Finally, we discuss new insights into organismal performance gained through the study of integration as an emergent property of kinematic systems working together during a common task. Integration is rarely the trait of interest, but we show that future work should adopt a more holistic approach to understand why and how animals perform complex behaviors.