Non-Bilaterians as Model Systems for Tissue Mechanics.

In animals, epithelial tissues are barriers against the external environment, providing protection against biological, chemical, and physical damage. Depending on the organism's physiology and behavior, these tissues encounter different types of mechanical forces and need to provide a suitable adaptive response to ensure success. Therefore, understanding tissue mechanics in different contexts is an important research area. Here, we review recent tissue mechanics discoveries in three early divergent non-bilaterian systems-Trichoplax adhaerens, Hydra vulgaris, and Aurelia aurita. We highlight each animal's simple body plan and biology and unique, rapid tissue remodeling phenomena that play a crucial role in its physiology. We also discuss the emergent large-scale mechanics in these systems that arise from small-scale phenomena. Finally, we emphasize the potential of these non-bilaterian animals to be model systems in a bottom-up approach for further investigation in tissue mechanics.

Trends in Stroke Kinematics, Reynolds Number, and Swimming Mode in Shrimp-Like Organisms.

Metachronal propulsion is commonly seen in organisms with the caridoid facies body plan, i.e. shrimp-like organisms, as they beat their pleopods in an adlocomotory sequence. These organisms exist across length scales ranging several orders of Reynolds number magnitude, from 10 to 104, during locomotion. Further, by altering their stroke kinematics, these organisms achieve three distinct swimming modes. To better understand the relationship between Reynolds number, stroke kinematics, and resulting swimming mode, Euphausia pacifica stroke kinematics were quantified using high-speed digital recordings and compared to the results for the larger E. superba. Euphausia pacifica consistently operate with a greater beat frequency and smaller stroke amplitude than E. superba for each swimming mode, suggesting that length scale may affect the kinematics needed to achieve similar swimming modes. To expand on this observation, these euphausiid data are used in combination with previously-published stroke kinematics from mysids and stomatopods to identify broad trends across swimming mode and length scale in metachrony. Principal component analysis (PCA) reveals trends in stroke kinematics and Reynolds number as well as the variation among taxonomic order. Overall, larger beat frequencies, stroke amplitudes, between-cycle phase lags, and Reynolds numbers are more representative of the fast forward swimming mode compared to the slower hovering mode. Additionally, each species has a unique combination of kinematics that result in metachrony, indicating that there are other factors, perhaps morphological, which affect the overall metachronal characteristics of an organism. Finally, uniform phase lag, in which the timing between power strokes of all pleopods is equal, in 5-paddle systems is achieved at different Reynolds numbers for different swimming modes, highlighting the importance of taking into consideration stroke kinematics, length scale, and the resulting swimming mode.

Dual Phase-Shifted Ipsilateral Metachrony in Americamysis bahia.

Previously documented metachrony in euphausiids focused on one, five-paddle metachronal stroke, where contralateral pleopod pairs on the same abdominal segment beat in tandem with each other, propelling the animal forward. In contrast, the mysid shrimp Americamysis bahia's pleopods on the same abdominal segment beat independently of each other, resulting in two, five-paddle metachronal cycles running ipsilaterally along the length of the body, 180° out of phase. The morphology, kinematics, and nondimensional measurements of efficiency are compared primarily with the one-cycle Euphausia superba to determine how the two-cycle approach alters the design and kinematics of metachrony. Pleopodal swimming in A. bahia results in only fast-forward swimming, with speeds greater than 2 BL/s (body lengths per second), and can reach speeds up to 12 BL/s, through a combination of increasing stroke amplitude, increasing beat frequency, and changing their inter-limb phase lag. Trends with Strouhal number and advance ratio suggest that the kinematics of metachrony in A. bahia are favored to achieve large normalized swimming speeds.

Metachronal Motion across Scales: Current Challenges and Future Directions.

Metachronal motion is used across a wide range of organisms for a diverse set of functions. However, despite its ubiquity, analysis of this behavior has been difficult to generalize across systems. Here we provide an overview of known commonalities and differences between systems that use metachrony to generate fluid flow. We also discuss strategies for standardizing terminology and defining future investigative directions that are analogous to other established subfields of biomechanics. Finally, we outline key challenges that are common to many metachronal systems, opportunities that have arisen due to the advent of new technology (both experimental and computational), and next steps for community development and collaboration across the nascent network of metachronal researchers.