Jennifer R. A. Taylor

Comparative spring mechanics in mantis shrimp

SUMMARY Elastic mechanisms are fundamental to fast and efficient movements. Mantis shrimp power their fast raptorial appendages using a conserved network of exoskeletal springs, linkages and latches. Their appendages are fantastically diverse, ranging from spears to hammers. We measured the spring mechanics of 12 mantis shrimp species from five different families exhibiting hammer-shaped, spear-shaped and undifferentiated appendages. Across species, spring force and work increase with size of the appendage and spring constant is not correlated with size. Species that hammer their prey exhibit significantly greater spring resilience compared with species that impale evasive prey (‘spearers’); mixed statistical results show that species that hammer prey also produce greater work relative to size during spring loading compared with spearers. Disabling part of the spring mechanism, the ‘saddle’, significantly decreases spring force and work in three smasher species; cross-species analyses show a greater effect of cutting the saddle on the spring force and spring constant in species without hammers compared with species with hammers. Overall, the study shows a more potent spring mechanism in the faster and more powerful hammering species compared with spearing species while also highlighting the challenges of reconciling within-species and cross-species mechanical analyses when different processes may be acting at these two different levels of analysis. The observed mechanical variation in spring mechanics provides insights into the evolutionary history, morphological components and mechanical behavior, which were not discernible in prior single-species studies. The results also suggest that, even with a conserved spring mechanism, spring behavior, potency and component structures can be varied within a clade with implications for the behavioral functions of power-amplified devices.

Structure and mechanical properties of selected protective systems in marine organisms

Marine organisms have developed a wide variety of protective strategies to thrive in their native environments. These biological materials, although formed from simple biopolymer and biomineral constituents, take on many intricate and effective designs. The specific environmental conditions that shape all marine organisms have helped modify these materials into their current forms: complete hydration, and variation in hydrostatic pressure, temperature, salinity, as well as motion from currents and swells. These conditions vary throughout the ocean, being more consistent in the pelagic and deep benthic zones while experiencing more variability in the nearshore and shallows (e.g. intertidal zones, shallow bays and lagoons, salt marshes and mangrove forests). Of note, many marine organisms are capable of migrating between these zones. In this review, the basic building blocks of these structural biological materials and a variety of protective strategies in marine organisms are discussed with a focus on their structure and mechanical properties. Finally, the bioinspired potential of these biological materials is discussed.

Mechanical properties of the rigid and hydrostatic skeletons of molting blue crabs, Callinectes sapidus Rathbun

SUMMARY Molting in crustaceans involves significant changes in the structure and function of the exoskeleton as the old cuticle is shed and a new one is secreted. The flimsy new cuticle takes several days to harden and during this time crabs rely on a hydrostatic skeletal support system for support and movement. This change from a rigid to a hydrostatic skeletal support mechanism implies correlated changes in the function, and thus mechanical properties, of the cuticle. In particular, it must change from primarily resisting compression, bending and torsional forces to resisting tension. This study was designed to explore the changes in the mechanical properties of the crustacean cuticle as the animals switch between two distinct skeletal support mechanisms. Samples of cuticle were removed from blue crabs, Callinectes sapidus, at 1 h (soft-shell stage), 12 h (paper-shell stage), and 7 days (hard-shell stage) following molting. We measured and compared the flexural stiffness, Youngs modulus of elasticity (in tension), and tensile strength for each postmolt stage. We found that the hard-shell cuticle has a flexural stiffness fully four orders of magnitude greater than the soft-shell and paper-shell cuticle. Although the soft-shell cuticle has a Youngs modulus significantly lower than that of the paper-shell and hard-shell cuticle, it has the same tensile strength. Thus, the soft-shell and paper-shell cuticles are unable to resist the significant bending forces associated with a rigid skeletal support system, but can resist the tensile forces that characterize hydrostatic support systems. The mechanical properties of the cuticle thus change dramatically during molting in association with the change in function of the cuticle. These results emphasize the significant role that mechanics plays in the evolution of the molting process in arthropods, and possibly other ecdysozoans.

Ritualized fighting and biological armor: the impact mechanics of the mantis shrimp's telson

SUMMARY Resisting impact and avoiding injury are central to survival in situations ranging from the abiotic forces of crashing waves to biotic collisions with aggressive conspecifics. Although impacts and collisions in biology are ubiquitous, most studies focus on the material properties of biological structures under static loading. Here, we examine the mechanical impact properties of the mantis shrimps telson, a piece of abdominal armor that withstands repeated, intense impacts from the potent hammer-like appendages used by conspecifics during ritualized fighting. We measured the coefficient of restitution, an index of elasticity, of the telson and compared it with that of an adjacent abdominal segment that is not impacted. We found that the telson behaves more like an inelastic punching bag than an elastic trampoline, dissipating 69% of the impact energy. Furthermore, although the abdominal segment provides no mechanical correlates with size, the telsons coefficient of restitution, displacement and impact duration all correlate with body size. The telsons mineralization patterns were determined through micro-CT (Computed Tomography) and correspond to the mechanical behavior of the telson during impact. The mineralized central region of the telson ‘punched’ inward during an impact whereas the surrounding areas provided elasticity owing to their reduced mineralization. Thus, the telson effectively dissipates impact energy while potentially providing the size-related information crucial to its role in conspecific assessment. This study reveals the mechanical infrastructure of impact resistance in biological armor and opens a new window to the biomechanical underpinnings of animal behavior and assessment.

Effects of CO2-induced pH reduction on the exoskeleton structure and biophotonic properties of the shrimp Lysmata californica.

The anticipated effects of CO2-induced ocean acidification on marine calcifiers are generally negative, and include dissolution of calcified elements and reduced calcification rates. Such negative effects are not typical of crustaceans for which comparatively little ocean acidification research has been conducted. Crustaceans, however, depend on their calcified exoskeleton for many critical functions. Here, we conducted a short-term study on a common caridean shrimp, Lysmata californica, to determine the effect of CO2-driven reduction in seawater pH on exoskeleton growth, structure, and mineralization and animal cryptic coloration. Shrimp exposed to ambient (7.99 ± 0.04) and reduced pH (7.53 ± 0.06) for 21 days showed no differences in exoskeleton growth (percent increase in carapace length), but the calcium weight percent of their cuticle increased significantly in reduced pH conditions, resulting in a greater Ca:Mg ratio. Cuticle thickness did not change, indicating an increase in the mineral to matrix ratio, which may have mechanical consequences for exoskeleton function. Furthermore, there was a 5-fold decrease in animal transparency, but no change in overall shrimp coloration (red). These results suggest that even short-term exposure to CO2-induced pH reduction can significantly affect exoskeleton mineralization and shrimp biophotonics, with potential impacts on crypsis, physical defense, and predator avoidance.

A pneumo-hydrostatic skeleton in land crabs

Like their aquatic counterparts, terrestrial crabs repeatedly shed their rigid exoskeleton during moulting. But in the case of land crabs, little water is available to provide a temporary hydrostatic skeleton before the new skeleton hardens, and air does not provide the buoyancy necessary to support the animal. Here we show that whenever its exoskeleton is shed, the blackback land crab Gecarcinus lateralis relies on an unconventional type of hydrostatic skeleton that uses both gas and liquid (a ‘pneumo-hydrostat’). To our knowledge, this is the first experimental evidence for a locomotor skeleton that depends on a gas. It establishes a new category of hydrostatic skeletal support and possibly a critical adaptation to life on land for the Crustacea.

Stress physiology and weapon integrity of intertidal mantis shrimp under future ocean conditions

Calcified marine organisms typically experience increased oxidative stress and changes in mineralization in response to ocean acidification and warming conditions. These effects could hinder the potency of animal weapons, such as the mantis shrimp’s raptorial appendage. The mechanical properties of this calcified weapon enable extremely powerful punches to be delivered to prey and aggressors. We examined oxidative stress and exoskeleton structure, mineral content, and mechanical properties of the raptorial appendage and the carapace under long-term ocean acidification and warming conditions. The predatory appendage had significantly higher % Mg under ocean acidification conditions, while oxidative stress levels as well as the % Ca and mechanical properties of the appendage remained unchanged. Thus, mantis shrimp tolerate expanded ranges of pH and temperature without experiencing oxidative stress or functional changes to their weapons. Our findings suggest that these powerful predators will not be hindered under future ocean conditions.

Biomechanics: a pneumo-hydrostatic skeleton in land crabs.

Like their aquatic counterparts, terrestrial crabs repeatedly shed their rigid exoskeleton during moulting. But in the case of land crabs, little water is available to provide a temporary hydrostatic skeleton before the new skeleton hardens, and air does not provide the buoyancy necessary to support the animal. Here we show that whenever its exoskeleton is shed, the blackback land crab Gecarcinus lateralis relies on an unconventional type of hydrostatic skeleton that uses both gas and liquid (a ‘pneumo-hydrostat’). To our knowledge, this is the first experimental evidence for a locomotor skeleton that depends on a gas. It establishes a new category of hydrostatic skeletal support and possibly a critical adaptation to life on land for the Crustacea.

Assessment of ocean acidification and warming on the growth, calcification, and biophotonics of a California grass shrimp

&NA; Cryptic colouration in crustaceans, important for both camouflage and visual communication, is achieved through physiological and morphological mechanisms that are sensitive to changes in environmental conditions. Consequently, ocean warming and ocean acidification can affect crustaceans’ biophotonic appearance and exoskeleton composition in ways that might disrupt colouration and transparency. In the present study, we measured growth, mineralization, transparency, and spectral reflectance (colouration) of the caridean grass shrimp Hippolyte californiensis in response to pH and temperature stressors. Shrimp were exposed to ambient pH and temperature (pH 8.0, 17 °C), decreased pH (pH 7.5, 17 °C), and decreased pH/increased temperature (pH 7.5, 19 °C) conditions for 7 weeks. There were no differences in either Mg or Ca content in the exoskeleton across treatments nor in the transparency and spectral reflectance. There was a small but significant increase in percent growth in the carapace length of shrimp exposed to decreased pH/increased temperature. Overall, these findings suggest that growth, calcification, and colour of H. californiensis are unaffected by decreases of 0.5 pH units. This tolerance might stem from adaptation to the highly variable pH environment that these grass shrimp inhabit, highlighting the multifarious responses to ocean acidification, within the Crustacea.

A Protocol for Bioinspired Design: A Ground Sampler Based on Sea Urchin Jaws

Bioinspired design is an emerging field that takes inspiration from nature to develop high-performance materials and devices. The sea urchin mouthpiece, known as the Aristotles lantern, is a compelling source of bioinspiration with an intricate network of musculature and calcareous teeth that can scrape, cut, chew food and bore holes into rocky substrates. We describe the bioinspiration process as including animal observation, specimen characterization, device fabrication and mechanism bioexploration. The last step of bioexploration allows for a deeper understanding of the initial biology. The design architecture of the Aristotles lantern is analyzed with micro-computed tomography and individual teeth are examined with scanning electron microscopy to identify the microstructure. Bioinspired designs are fabricated with a 3D printer, assembled and tested to determine the most efficient lantern opening and closing mechanism. Teeth from the bioinspired lantern design are bioexplored via finite element analysis to explain from a mechanical perspective why keeled tooth structures evolved in the modern sea urchins we observed. This circular approach allows for new conclusions to be drawn from biology and nature.