The Kinematics and Functional Significance of Chemosensory Tongue-Flicking in Northern Water Snakes (Nerodia sipedon) on Land, in Water, and in between.

As organisms transition between different environments, they must do more than simply move through that transition and those environments. Changes in the environment must be detected via the senses. The types of sensory information and the mechanisms of collecting that information may also change as an individual moves through different environments. We use tongue-flicking in northern water snakes, Nerodia sipedon, to examine the mechanics of sensory behavior as snakes move from terrestrial to aquatic habitats. A combination of high-speed video and mesocosm experiment revealed that water snakes will alter the mechanics of tongue-flicking in the context of their environment. Tongue-flicks on land are distinctive, with multiple oscillations, large protrusion distance, and high velocities. Comparatively, tongue-flicks under water are much shorter events, with reduced protrusion and fewer oscillations. At the surface of the water, in the presence of potential anuran prey, water snakes will tap the tips of the tongue on the surface of the water, without undergoing the full oscillations observed on land or underwater. We attribute the differences in the aerial and underwater tongue-flicks to trade-offs in the physical and chemical properties of the environment. The surface tapping behavior we observed is likely snakes altering their behavior to maximize the encounter and collection of frog-specific chemical cues, which are known to travel on the water's surface. Given the ecological transitions and distinctive biogeographical patterns rooted in water snake ecology, there are likely more examples of changing sensory mechanics to be discovered upon further investigation.

Linking Tooth Shape to Strike Mechanics in the Boa constrictor.

Snakes, with the obvious exception of the fangs, are considered to lack the regional specialization of tooth shape and function which are exemplified by mammals. Recent work in fishes has suggested that the definition of homodont and heterodont are incomplete without a full understanding of the morphology, mechanics, and behavior of feeding. We investigated this idea further by examining changes in tooth shape along the jaw of Boa constrictor and integrating these data with the strike kinematics of boas feeding on rodent prey. We analyzed the shape of every tooth in the skull, from a combination of anesthetized individuals and CT scanned museum specimens. For strike kinematics, we filmed eight adult boas striking at previously killed rats. We determined the regions of the jaws that made first contact with the prey, and extrapolated the relative positions of those teeth at that moment. We further determined the roles of all the teeth throughout the prey capture process, from the initiation of the strike until constriction began. We found that the teeth in the anterior third of the mandible are the most upright, and that teeth become progressively more curved posteriorly. Teeth on the maxilla are more curved than on the mandible, and the anterior teeth are more linear or recurved than the posterior teeth. In a majority of strikes, boas primarily made contact with the anterior third of the mandible first. The momentum from the strike caused the upper jaws and skull to rotate over the rat. The more curved teeth of the upper jaw slid over the rat unimpeded until the snake began to close its jaws. In the remaining strikes, boas made contact with the posterior third of both jaws simultaneously, driving through the prey and quickly retracting, ensnaring the prey on the curved posterior teeth of both jaws. The curved teeth of the palatine and pterygoid bones assist in the process of swallowing.

Buccal pumping mechanics of Xenopus laevis tadpoles: effects of biotic and abiotic factors.

Biotic factors such as body size and shape have long been known to influence kinematics in vertebrates. Movement in aquatic organisms can also be strongly affected by abiotic factors such as the viscosity of the medium. We examined the effects of both biotic factors and abiotic factors on buccal pumping kinematics in Xenopus tadpoles using high-speed imaging of an ontogenetic series of tadpoles combined with experimental manipulation of the medium over a 10-fold range of viscosity. We found influences of both biotic and abiotic factors on tadpole movements; absolute velocities and excursions of the jaws and hyoid were greater in higher viscosity fluid but durations of movements were unaffected. Smaller tadpoles have relatively wider heads and more robust hyoid muscles used in buccal expansion and compression. Lever arm ratios were found to be constant at all sizes; therefore, smaller tadpoles have relatively higher resolved muscle forces and, like tadpoles in more viscous medium, displayed higher absolute velocities of jaw and hyoid movements. Nonetheless, small tadpoles drew in water at lower Reynolds numbers (Re) than predicted by kinematics, due to negative allometry of the buccal pump. Finally, tadpoles transitioned from a flow regime dominated by viscous forces (Re=2) to an intermediate regime (Re=106).

Captivity Affects Head Morphology and Allometry in Headstarted Garter Snakes, Thamnophis sirtalis.

In response to the growing number of amphibian and reptiles species in decline, many conservation managers have implemented captive breeding and headstarting programs in an effort to restore these populations. However, many of these programs suffer from low survival success, and it is often unclear as to why some individuals do not survive after reintroduction. Here I document changes to head morphology in the eastern garter snake, Thamnophis sirtalis, in response to time spent in captivity. Thamnophis raised on three diet treatments all differed in head size from wild individuals, and head size differed between the three treatments. Overall, head size was smaller in all three diet treatments than in wild snakes, potentially limiting the available prey for the captive garter snakes. Allometric patterns of growth in head size were also different for each diet treatment. Several potential implications of these changes in morphology are discussed, and what these changes may mean for other species that are part of headstarting and reintroduction programs.

Ontogeny of strike performance in ball pythons (Python regius): a three-year longitudinal study.

The rapid strike of snakes has long been of interest in terms of mechanical performance. Recently, several nonvenomous taxa have been found to strike with the same incredible strike velocity and acceleration as the high-performing vipers. However, little is known regarding how these patterns change through ontogeny. Here I present ontogenetic strike data on ten ball pythons (Python regius) over a three year time period, from birth to sexual maturity. I found that performance declined rapidly over the first 18 months in nearly all kinematic measures. This puts the adult data out of the currently developing trend of high performance being maintained across the diversity of snakes. The underlying cause of the decline in performance is unclear, but there are several avenues of behavior, morphology, biomechanics, and ecology to be investigated.

Strike kinematics and performance in juvenile ball pythons (Python regius).

The rapid strike of snakes has interested researchers for decades. Although most work has focused on the strike performance of vipers, recent work has shown that other snakes outside of the Viperidae can strike with the same velocities and accelerations. However, to date all of these examples focus on performance in adult snakes. Here, we use high-speed video to measure the strike kinematics and performance of 10 juvenile (<6 months of age) ball pythons, Python regius. We find that juvenile P. regius strike at levels comparable to larger snakes, but with shorter durations and over shorter distances. We conclude that the juvenile P. regius maintain performance likely through manipulation of the axial musculature and accompanying elastic tissues, and that this is a first step to understanding ontogenetic changes in behavior and a potential avenue for understanding how captivity may also impact behavior.

A simple, inexpensive system for digital particle image velocimetry (DPIV) in biomechanics.

Functional morphology and biomechanics seek to reveal the mechanistic bases of organismal functions and the physical principles involved at the phenotype-environment interface. Characterization of fluid flow (air or water) within and around organismal structures is an example of this approach. Digital particle imaging velocimetry (DPIV) has been exploited in a variety of biological systems to visualize fluid flow associated with animal movement. DPIV employs particles suspended in air or water that are illuminated by a laser light sheet and recorded with a high-speed video camera. Software tracks particle movement across a specified number of video frames, generating vector diagrams showing patterns of fluid flow through time. As powerful as DPIV methods are, they are limited in application by the high cost and complexity of the equipment required. In this article, we describe a simple DPIV system that substitutes widely available, inexpensive consumer components for scientific-grade equipment to achieve low cost (<$1,000 total) and high accuracy (total error calculated to be approx. 6%, as compared with 5% in professional systems). We have employed this system successfully in our studies on the fluid dynamics of chemosensory tongue-flicking in snakes. This system can be used for research and teaching in labs that typically cannot afford the expense or commitment of a traditional DPIV apparatus and is particularly suited for obtaining preliminary data required to justify further grant and institutional support.