Dense and pleiotropic regulatory information in a developmental enhancer.

Changes in gene regulation underlie much of phenotypic evolution1. However, our understanding of the potential for regulatory evolution is biased, because most evidence comes from either natural variation or limited experimental perturbations2. Using an automated robotics pipeline, we surveyed an unbiased mutation library for a developmental enhancer in Drosophila melanogaster. We found that almost all mutations altered gene expression and that parameters of gene expression-levels, location, and state-were convolved. The widespread pleiotropic effects of most mutations may constrain the evolvability of developmental enhancers. Consistent with these observations, comparisons of diverse Drosophila larvae revealed apparent biases in the phenotypes influenced by the enhancer. Developmental enhancers may encode a higher density of regulatory information than has been appreciated previously, imposing constraints on regulatory evolution.

A simple strategy for detecting moving objects during locomotion revealed by animal-robot interactions.

An important role of visual systems is to detect nearby predators, prey, and potential mates, which may be distinguished in part by their motion. When an animal is at rest, an object moving in any direction may easily be detected by motion-sensitive visual circuits. During locomotion, however, this strategy is compromised because the observer must detect a moving object within the pattern of optic flow created by its own motion through the stationary background. However, objects that move creating back-to-front (regressive) motion may be unambiguously distinguished from stationary objects because forward locomotion creates only front-to-back (progressive) optic flow. Thus, moving animals should exhibit an enhanced sensitivity to regressively moving objects. We explicitly tested this hypothesis by constructing a simple fly-sized robot that was programmed to interact with a real fly. Our measurements indicate that whereas walking female flies freeze in response to a regressively moving object, they ignore a progressively moving one. Regressive motion salience also explains observations of behaviors exhibited by pairs of walking flies. Because the assumptions underlying the regressive motion salience hypothesis are general, we suspect that the behavior we have observed in Drosophila may be widespread among eyed, motile organisms.

A linear systems analysis of the yaw dynamics of a dynamically scaled insect model.

Recent studies suggest that fruit flies use subtle changes to their wing motion to actively generate forces during aerial maneuvers. In addition, it has been estimated that the passive rotational damping caused by the flapping wings of an insect is around two orders of magnitude greater than that for the body alone. At present, however, the relationships between the active regulation of wing kinematics, passive damping produced by the flapping wings and the overall trajectory of the animal are still poorly understood. In this study, we use a dynamically scaled robotic model equipped with a torque feedback mechanism to study the dynamics of yaw turns in the fruit fly Drosophila melanogaster. Four plausible mechanisms for the active generation of yaw torque are examined. The mechanisms deform the wing kinematics of hovering in order to introduce asymmetry that results in the active production of yaw torque by the flapping wings. The results demonstrate that the stroke-averaged yaw torque is well approximated by a model that is linear with respect to both the yaw velocity and the magnitude of the kinematic deformations. Dynamic measurements, in which the yaw torque produced by the flapping wings was used in real-time to determine the rotation of the robot, suggest that a first-order linear model with stroke-average coefficients accurately captures the yaw dynamics of the system. Finally, an analysis of the stroke-average dynamics suggests that both damping and inertia will be important factors during rapid body saccades of a fruit fly.