Spatial accuracy of a rapid defense behavior in caterpillars.

Aimed movements require that an animal accurately locates the target and correctly reaches that location. One such behavior is the defensive strike seen in Manduca sexta larva. These caterpillars respond to noxious mechanical stimuli applied to their abdomen with a strike of the mandibles towards the location of the stimulus. The accuracy with which the first strike movement reaches the stimulus site depends on the location of the stimulus. Reponses to dorsal stimuli are less accurate than those to ventral stimuli and the mandibles generally land ventral to the stimulus site. Responses to stimuli applied to anterior abdominal segments are less accurate than responses to stimuli applied to more posterior segments and the mandibles generally land posterior to the stimulus site. A trade-off between duration of the strike and radial accuracy is only seen in the anterior stimulus location (body segment A4). The lower accuracy of the responses to anterior and dorsal stimuli can be explained by the morphology of the animal; to reach these areas the caterpillar needs to move its body into a tight curve. Nevertheless, the accuracy is not exact in locations that the animal has shown it can reach, which suggests that consistently aiming more ventral and posterior of the stimulation site might be a defense strategy.

Visceral-locomotory pistoning in crawling caterpillars.

Animals with an open coelom do not fully constrain internal tissues, and changes in tissue or organ position during body movements cannot be readily discerned from outside of the body. This complicates modeling of soft-bodied locomotion, because it obscures potentially important changes in the center of mass as a result of internal tissue movements. We used phase-contrast synchrotron X-ray imaging and transmission light microscopy to directly visualize internal soft-tissue movements in freely crawling caterpillars. Here we report a novel visceral-locomotory piston in crawling Manduca sexta larvae, in which the gut slides forward in advance of surrounding tissues. The initiation of gut sliding is synchronous with the start of the terminal prolegs' swing phase, suggesting that the animal's center of mass advances forward during the midabdominal prolegs' stance phase and is therefore decoupled from visible translations of the body. Based on synchrotron X-ray data and transmission light microscopy results, we present evidence for a two-body mechanical system with a nonlinear elastic gut that changes size and translates between the anterior and posterior of the animal. The proposed two-body system--the container and the contained--is unlike any form of legged locomotion previously reported and represents a new feature in our emerging understanding of crawling.

Caterpillar crawling over irregular terrain: anticipation and local sensing.

Animal locomotion is produced by co-coordinated patterns of motor activity that are generally organized by central pattern generators and modified by sensory feedback. Animals with remote sensing can anticipate obstacles and make adjustments in their gait to accommodate them. It is largely unknown how animals that rely on touch might use such information to adjust their gait. One possibility is immediate (reflexive) change in motor activity. Elongated animals, however, might modulate movements by passing information from anterior to posterior segments. Using the caterpillar Manduca sexta we examined the movements of the most anterior abdominal prolegs as they approached an obstacle. The first pair of prolegs anticipated the obstacle by lifting more quickly in the earliest part of the swing phase: the caterpillar had information about the obstacle at proleg lift-off. Sometimes the prolegs corrected their trajectory mid-step. Removal of sensory hairs on the stepping leg did not affect the early anticipatory movements, but did change the distance at which the mid-step corrections occurred. We conclude that anterior sensory information can be passed backwards and used to modulate an ongoing crawl. The local sensory hairs on each body segment can then fine-tune movements of the prolegs as they approach an obstacle.

Kinematics of horizontal and vertical caterpillar crawling.

Unlike horizontal crawling, vertical crawling involves two counteracting forces: torque rotating the body around its center of mass and gravity resisting forward movement. The influence of these forces on kinematics has been examined in the soft-bodied larval stage of Manduca sexta. We found that crawling and climbing are accomplished using the same movements, with both segment timing and proleg lift indistinguishable in horizontal and vertical locomotion. Minor differences were detected in stride length and in the delay between crawls, which led to a lower crawling speed in the vertical orientation. Although these differences were statistically significant, they were much smaller than the variation in kinematic parameters between animals. The ability of Manduca to crawl and climb using the same movements is best explained by Manduca's relatively small size, slow speed and strong, controlled, passive grip made possible by its proleg/crochets.

Exogenous and endogenous corticosterone alter feather quality.

We investigated how exogenous and endogenous glucocorticoids affect feather replacement in European starlings (Sturnus vulgaris) after approximately 56% of flight feathers were removed. We hypothesized that corticosterone would retard feather regrowth and decrease feather quality. After feather regrowth began, birds were treated with exogenous corticosterone or sham implants, or endogenous corticosterone by applying psychological or physical (food restriction) stressors. Exogenous corticosterone had no impact on feather length and vane area, but rectrices were lighter than controls. Exogenous corticosterone also decreased inter-barb distance for all feathers and increased barbule number for secondaries and rectrices. Although exogenous corticosterone had no affect on rachis tensile strength and stiffness, barbicel hooking strength was reduced. Finally, exogenous corticosterone did not alter the ability of Bacillus licheniformis to degrade feathers or affect the number of feathers that failed to regrow. In contrast, endogenous corticosterone via food restriction resulted in greater inter-barb distances in primaries and secondaries, and acute and chronic stress resulted in greater inter-barb distances in rectrices. Food-restricted birds had significantly fewer barbules in primaries than chronic stress birds and weaker feathers compared to controls. We conclude that, although exogenous and endogenous corticosterone had slightly different effects, some flight feathers grown in the presence of high circulating corticosterone are lighter, potentially weaker, and with altered feather micro-structure.