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Energetic trade-offs are particularly pertinent to bio-ballistic systems which impart energy to projectiles exclusively during launch. We investigated such trade-offs in the spring-propelled seeds of Loropetalum chinense , Hamamelis virginiana and Fortunearia sinensis . Using similar seed-shooting mechanisms, fruits of these confamilial plants (Hamamelidaceae) span an order of magnitude in spring and seed mass. We expected that as seed mass increases, launch speed decreases. Instead, launch speed was relatively constant regardless of seed mass. We tested if fruits shoot larger seeds by storing more elastic potential energy (PE). Spring mass and PE increased as seed mass increased (in order of increasing seed mass: L. chinense , H. virginiana , F. sinensis ). As seed mass to spring mass ratio increased (ratios: H. virginiana = 0.50, F. sinensis = 0.65, L. chinense = 0.84), mass-specific PE storage increased. The conversion efficiency of PE to seed kinetic energy (KE) decreased with increasing fruit mass. Therefore, similar launch speeds across scales occurred because (i) larger fruits stored more PE and (ii) smaller fruits had higher mass-specific PE storage and improved PE to KE conversion. By examining integrated spring and projectile mechanics in our focal species, we revealed diverse, energetic scaling strategies relevant to spring-propelled systems navigating energetic trade-offs. 

ABSTRACT Across the tree of life – from fungi to frogs – organisms wield small amounts of energy to generate fast and potent movements. These movements are propelled with elastic structures, and their loading and release are mediated by latch-like opposing forces. They comprise a class of elastic mechanisms termed latch-mediated spring actuation (LaMSA). Energy flow through LaMSA begins when an energy source loads elastic element(s) in the form of elastic potential energy. Opposing forces, often termed latches, prevent movement during loading of elastic potential energy. As the opposing forces are shifted, reduced or removed, elastic potential energy is transformed into kinetic energy of the spring and propelled mass. Removal of the opposing forces can occur instantaneously or throughout the movement, resulting in dramatically different outcomes for consistency and control of the movement. Structures used for storing elastic potential energy are often distinct from mechanisms that propel the mass: elastic potential energy is often distributed across surfaces and then transformed into localized mechanisms for propulsion. Organisms have evolved cascading springs and opposing forces not only to serially reduce the duration of energy release, but often to localize the most energy-dense events outside of the body to sustain use without self-destruction. Principles of energy flow and control in LaMSA biomechanical systems are emerging at a rapid pace. New discoveries are catalyzing remarkable growth of the historic field of elastic mechanisms through experimental biomechanics, synthesis of novel materials and structures, and high-performance robotics systems. 

ABSTRACT Organisms such as jumping froghopper insects and punching mantis shrimp use spring-based propulsion to achieve fast motion. Studies of elastic mechanisms have primarily focused on fully developed and functional mechanisms in adult organisms. However, the ontogeny and development of these mechanisms can provide important insights into the lower size limits of spring-based propulsion, the ecological or behavioral relevance of ultrafast movement, and the scaling of ultrafast movement. Here, we examined the development of the spring-latch mechanism in the bigclaw snapping shrimp, Alpheus heterochaelis (Alpheidae). Adult snapping shrimp use an enlarged claw to produce high-speed strikes that generate cavitation bubbles. However, until now, it was unclear when the elastic mechanism emerges during development and whether juvenile snapping shrimp can generate cavitation at this size. We reared A. heterochaelis from eggs, through their larval and postlarval stages. Starting 1 month after hatching, the snapping shrimp snapping claw gradually developed a spring-actuated mechanism and began snapping. We used high-speed videography (300,000 frames s−1) to measure juvenile snaps. We discovered that juvenile snapping shrimp generate the highest recorded accelerations (5.8×105±3.3×105 m s−2) for repeated-use, underwater motion and are capable of producing cavitation at the millimeter scale. The angular velocity of snaps did not change as juveniles grew; however, juvenile snapping shrimp with larger claws produced faster linear speeds and generated larger, longer-lasting cavitation bubbles. These findings establish the development of the elastic mechanism and cavitation in snapping shrimp and provide insights into early life-history transitions in spring-actuated mechanisms. 

Synopsis Across countless marine invertebrates, coordination of closely spaced swimming appendages is key to producing diverse locomotory behaviors. Using a widespread mechanism termed hybrid metachronal propulsion, mantis shrimp swim by moving five paddle-like pleopods along their abdomen in a posterior to anterior sequence during the power stroke and a near-synchronous motion during the recovery stroke. Despite the ubiquity of this mechanism, it is not clear how hybrid metachronal swimmers coordinate and modify individual appendage movements to achieve a range of swimming capabilities. Using high-speed imaging, we measured pleopod kinematics of mantis shrimp (Neogonodactylus bredini), while they performed two swimming behaviors: burst swimming and taking off from the substrate. By tracking each of the five pleopods, we tested how stroke kinematics vary across swimming speeds and the two swimming behaviors. We found that mantis shrimp achieve faster swimming speeds through a combination of higher beat frequencies, smaller stroke durations, and partially via larger stroke angles. The five pleopods exhibit non-uniform kinematics that contribute to the coordination and forward propulsion of the whole system. Micro-hook structures (retinacula) connect each of the five pleopod pairs and differ in their attachment across pleopods—possibly contributing to passive kinematic control. We compare our findings in N. bredini to previous studies to identify commonalities across hybrid metachronal swimmers at high Reynolds numbers and centimeter scales. Through our large experimental dataset and by tracking each pleopod's movements, our study reveals key parameters by which mantis shrimp adjust and control their swimming, yielding diverse locomotor abilities. 

ABSTRACT Small organisms use propulsive springs rather than muscles to repeatedly actuate high acceleration movements, even when constrained to tiny displacements and limited by inertial forces. Through integration of a large kinematic dataset, measurements of elastic recoil, energetic math modeling and dynamic math modeling, we tested how trap-jaw ants (Odontomachus brunneus) utilize multiple elastic structures to develop ultrafast and precise mandible rotations at small scales. We found that O. brunneus develops torque on each mandible using an intriguing configuration of two springs: their elastic head capsule recoils to push and the recoiling muscle–apodeme unit tugs on each mandible. Mandibles achieved precise, planar, circular trajectories up to 49,100 rad s−1 (470,000 rpm) when powered by spring propulsion. Once spring propulsion ended, the mandibles moved with unconstrained and oscillatory rotation. We term this mechanism a ‘dual spring force couple’, meaning that two springs deliver energy at two locations to develop torque. Dynamic modeling revealed that dual spring force couples reduce the need for joint constraints and thereby reduce dissipative joint losses, which is essential to the repeated use of ultrafast, small systems. Dual spring force couples enable multifunctionality: trap-jaw ants use the same mechanical system to produce ultrafast, planar strikes driven by propulsive springs and for generating slow, multi-degrees of freedom mandible manipulations using muscles, rather than springs, to directly actuate the movement. Dual spring force couples are found in other systems and are likely widespread in biology. These principles can be incorporated into microrobotics to improve multifunctionality, precision and longevity of ultrafast systems. 

Abstract Ultrafast movements propelled by springs and released by latches are thought limited to energetic adjustments prior to movement, and seemingly cannot adjust once movement begins. Even so, across the tree of life, ultrafast organisms navigate dynamic environments and generate a range of movements, suggesting unrecognized capabilities for control. We develop a framework of control pathways leveraging the non-linear dynamics of spring-propelled, latch-released systems. We analytically model spring dynamics and develop reduced-parameter models of latch dynamics to quantify how they can be tuned internally or through changing external environments. Using Lagrangian mechanics, we test feedforward and feedback control implementation via spring and latch dynamics. We establish through empirically-informed modeling that ultrafast movement can be controllably varied during latch release and spring propulsion. A deeper understanding of the interconnection between multiple control pathways, and the tunability of each control pathway, in ultrafast biomechanical systems presented here has the potential to expand the capabilities of synthetic ultra-fast systems and provides a new framework to understand the behaviors of fast organisms subject to perturbations and environmental non-idealities. 

Synopsis Numerous aquatic invertebrates use drag-based metachronal rowing for swimming, in which closely spaced appendages are oscillated starting from the posterior, with each appendage phase-shifted in time relative to its neighbor. Continuously swimming species such as Antarctic krill generally use “pure metachronal rowing” consisting of a metachronal power stroke and a metachronal recovery stroke, while burst swimming species such as many copepods and mantis shrimp typically use “hybrid metachronal rowing” consisting of a metachronal power stroke followed by a synchronous or nearly synchronous recovery stroke. Burst swimming organisms need to rapidly accelerate in order to capture prey and/or escape predation, and it is unknown whether hybrid metachronal rowing can augment acceleration and swimming speed compared to pure metachronal rowing. Simulations of rigid paddles undergoing simple harmonic motion showed that collisions between adjacent paddles restrict the maximum stroke amplitude for pure metachronal rowing. Hybrid metachronal rowing similar to that observed in mantis shrimp (Neogonodactylus bredini) permits oscillation at larger stroke amplitude while avoiding these collisions. We comparatively examined swimming speed, acceleration, and wake structure of pure and hybrid metachronal rowing strategies by using a self-propelling robot. Both swimming speed and peak acceleration of the robot increased with increasing stroke amplitude. Hybrid metachronal rowing permitted operation at larger stroke amplitude without collision of adjacent paddles on the robot, augmenting swimming speed and peak acceleration. Hybrid metachronal rowing generated a dispersed wake unlike narrower, downward-angled jets generated by pure metachronal rowing. Our findings suggest that burst swimming animals with small appendage spacing, such as copepods and mantis shrimp, can use hybrid metachronal rowing to generate large accelerations via increasing stroke amplitude without concern of appendage collision. 

ABSTRACT Latch-mediated spring actuation (LaMSA) is used by small organisms to produce high acceleration movements. Mathematical models predict that acceleration increases as LaMSA systems decrease in size. Adult mantis shrimp use a LaMSA mechanism in their raptorial appendages to produce extremely fast strikes. Until now, however, it was unclear whether mantis shrimp at earlier life-history stages also strike using elastic recoil and latch mediation. We tested whether larval mantis shrimp (Gonodactylaceus falcatus) use LaMSA and, because of their smaller size, achieve higher strike accelerations than adults of other mantis shrimp species. Based on microscopy and kinematic analyses, we discovered that larval G. falcatus possess the components of, and actively use, LaMSA during their fourth larval stage, which is the stage of development when larvae begin feeding. Larvae performed strikes at high acceleration and speed (mean: 4.133×105 rad s−2, 292.7 rad s−1; 12 individuals, 25 strikes), which are of the same order of magnitude as for adults – even though adult appendages are up to two orders of magnitude longer. Larval strike speed (mean: 0.385 m s−1) exceeded the maximum swimming speed of similarly sized organisms from other species by several orders of magnitude. These findings establish the developmental timing and scaling of the mantis shrimp LaMSA mechanism and provide insights into the kinematic consequences of scaling limits in tiny elastic mechanisms.