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Many marine invertebrates have larval stages covered in linear arrays of beating cilia, which propel the animal while simultaneously entraining planktonic prey. 1 These bands are strongly conserved across taxa spanning four major superphyla, 2, 3 and they are responsible for the unusual morphologies of many invertebrate larvae. 4, 5 However, few studies have investigated their underlying hydrodynamics. 6,7 Here, we study the ciliary bands of starfish larvae, and discover a beautiful pattern of slowly-evolving vortices that surrounds the swimming animals. Closer inspection of the bands reveals unusual ciliary "tangles" analogous to topological defects that break-up and re-form as the animal adjusts its swimming stroke. Quantitative experiments and modeling demonstrate that these vortices create a physical tradeoff between feeding and swimming in heterogenous environments, which manifests as distinct flow patterns or "eigenstrokes" representing each behavior-potentially implicating neuronal control of cilia. This quantitative interplay between larval form and hydrodynamic function may generalize to other invertebrates with ciliary bands, and illustrates the potential effects of active boundary conditions in other biological and synthetic systems.

The mechanical properties of cells and tissues play a well-known role in physiology and disease. The model organism Caenorhabditis elegans exhibits mechanical properties that are still poorly understood, but are thought to be dominated by its collagen-rich outer cuticle. To our knowledge, we use a novel microfluidic technique to reveal that the worm responds linearly to low applied hydrostatic stress, exhibiting a volumetric compression with a bulk modulus, k ¼ 140 5 20 kPa; applying negative pressures leads to volumetric expansion of the worm, with a similar bulk modulus. Surprisingly, however, we find that a variety of collagen mutants and pharmacological perturbations targeting the cuticle do not impact the bulk modulus. Moreover, the worm exhibits dramatic stiffening at higher stresses-behavior that is also independent of the cuticle. The stress-strain curves for all conditions can be scaled onto a master equation, suggesting that C. elegans exhibits a universal elastic response dominated by the mechanics of pressurized internal organs.

Archaeologists argue that the replacement of Neanderthals by modern humans was driven by interspecific competition due to a difference in culture level. To assess the cogency of this argument, we construct and analyze an interspecific cultural competition model based on the Lotka−Volterra model, which is widely used in ecology, but which incorporates the culture level of a species as a variable interacting with population size. We investigate the conditions under which a difference in culture level between cognitively equivalent species, or alternatively a difference in underlying learning ability, may produce competitive exclusion of a comparatively (although not absolutely) large local Neanderthal population by an initially smaller modern human population. We find, in particular, that this competitive exclusion is more likely to occur when population growth occurs on a shorter timescale than cultural change, or when the competition coefficients of the Lotka−Volterra model depend on the difference in the culture levels of the interacting species.Lotka−Volterra | cultural evolution | Paleolithic | feedback | replacement N eanderthals are a human species (or subspecies) that went extinct, after making a small contribution to the modern human genome (1, 2). Hypotheses for the Neanderthal extinction and their replacement by modern humans, in particular as recorded in Europe, can be classified into those emphasizing competition with modern humans and those arguing that interspecific competition was of minor relevance. Among the latter are the climate change (3) and epidemic/endemic (4) hypotheses. However, an ecocultural niche modeling study has shown that Neanderthals and modern humans exploited similar niches in Europe (5), which, together with a recent reassessment of European Paleolithic chronology showing significant spatiotemporal overlap of the two species (6), suggests a major role for interspecific competition in the demise of the Neanderthals.Replacement of one species (or population) by another is ultimately a matter of numbers. One competing species survives while the other is reduced to, or approaches, zero in size. In the classical Lotka−Volterra model of interspecific competition, this process is called competitive exclusion (7). If Neanderthals were indeed outcompeted by modern humans, the question arises: Wherein lay the advantage to the latter species? Many suggestions have been made, including better tools (8), better clothing (9, 10), and better economic organization (11). These hypotheses share the premise that modern humans were culturally more advanced than the coeval Neanderthals.The purpose of our paper is threefold. First, we extend the Lotka−Volterra-type model of interspecific competition by incorporating the "culture level" of a species as a variable that interacts with population size (12, 13). Here, culture level may be interpreted as the number of cultural traits, toolkit size, toolkit sophistication, etc. Although, as noted above, many anthropological and archaeological discussions i...