Jeannette Yen

Advertisement and concealment in the plankton: what makes a copepod hydrodynamically conspicuous?

Euchaeta rimana, a pelagic marine copepod, roams a 3-dimensional environment and its antennular setal sensors are oriented to detect water-borne signals in 3 dimensions. When the copepod moves through water or moves water around itself, it creates a fluid dis- turbance distinct from the ambient fluid motion. As it swims and hovers, the copepods laminar feeding current takes the unstable nature of small-scale turbulence, organizes it, and makes the local domain a familiar territory within which signals can be specified in time and space. The streamlines betray both the 3-dimensional spatial location (x, y, z) as well as the time (t) separating a signal caught in the feeding current and the copepod receptor-giving the copepod early warning of the approach of a prey, predator, or mate. The copepod reduces the complexity of its environment by fixing information from a turbulent field into a simpler, more defined laminar field. We quantitatively analysed small-scale fluid deformations created by copepods to document the strength of the signal. Physiological and behavioral tests confirm (a) that these disturbances are relevant signals transmitting information between zooplankters and (b) that hydrodynami- cally conspicuous structures, such as feeding currents, wakes, and vibrations, elicit specific responses from copepods. Since fluid mechanical signals do elicit responses, copepods shape their fluid motion to advertise or to conceal their hydrodynamic presence. When swimming, a copepod barely leaves a trace in the water; the animal generates its flow and advances into the area from which the water is taken, covering up its tracks with the velocity gradient it creates around itself. When escaping, it sheds conspicuous vortices. Prey caught in a flow field expose their location by hopping. These escape hops shed jet-like wakes detected by copepod mech- anoreceptors. Copepods recognize the wakes and respond adaptively.

DANE: Fostering Creativity in and through Biologically Inspired Design

In this paper, we present an initial attempt at systemizing knowledge of biological systems from an engineering perspective. In particular, we describe an interactive knowledge-based design environment called DANE that uses the Structure-Behavior-Function (SBF) schema for capturing the functioning of biological systems. We present preliminary results from deploying DANE in an interdisciplinary class on biologically inspired design, indicating that designers found the SBF schema useful for conceptualizing complex systems.

Quantitative analysis of tethered and free-swimming copepodid flow fields

SUMMARY We quantified the flow field generated by tethered and free-swimming Euchaeta antarctica using the particle image velocimetry (PIV) technique. The streamlines around the free-swimming specimens were generally parallel to the body axis, whereas the streamlines around all of the tethered copepodids demonstrated increased curvature. Differences noted in the streamline pattern, and hence the vorticity, dissipation rate and strain rate fields, are explained by considering the forces on the free-swimming specimen compared to the tethered specimen. Viscous flow theory demonstrates that the force on the fluid due to the presence of the tether irrevocably modifies the flow field in a manner that is consistent with the measurements. Hence, analysis of the flow field and all associated calculations differ for tethered versus free-swimming conditions. Consideration of the flow field of the free-swimming predatory copepodid shows the intensity of the biologically generated flow and the extent of the mechanoreceptive signal quantified in terms of shear strain rate. The area in the dorso-ventral view surrounded by the 0.5 s-1 contour of exy, which is a likely threshold to induce an escape response, is 11 times the area of the exoskeletal form for the free-swimming case. Thus, mechanoreceptive predators will perceive a more spatially extended signal than the body size.

Analysis of the flow field of the krill, Euphausia pacifica

Velocity measurements were performed for the flow field generated by tethered krill Euphausia pacifica. The particle image velocimetry (PIV) technique was used to measure the velocity field in vertical planes aligned with the krill body axis. The krill generates a narrow jet-like flow behind and below the pleopods (roughly 25° below horizontal). The volume of fluid moving at greater than 10% of the maximum velocity near the pleopods is roughly 18 times larger than the volume of the krill. Thus, the hydrodynamic disturbance occupies a significantly larger region than the animal body. Other krill, sensing the flow disturbance, may take advantage of the flow induced by a neighbor to locate a mate or to draft for efficient propulsion.

The prevalence and implications of copepod behavioral responses to oceanographic gradients and biological patchiness

Several species and developmental stages of calanoid copepods were tested for responses to environmental cues in a laboratory apparatus that mimicked conditions commonly associated with patches of food in the ocean. All species responded to the presence of phytoplankton by feeding. All species responded by increasing proportional residence time in one, but not both, of the treatments defined by gradients of velocity or density. Most species increased swimming speed and frequency of turning in response to the presence of chemical exudates or gradients of velocity. Only one species, Eurytemora affinis, increased proportional time of residence in response to gradients in density of the water. Responses of E. affinis to combined cues did not definitively demonstrate a hierarchical use of different cues as previously observed for Temora longicornis and Acartia tonsa. A simple foraging simulation was developed to assess the applicability in the field of the behavioral results observed in the laboratory. These simulations suggest that observed fine-scale behaviors could lead to copepod aggregations observed in situ. The present study demonstrates that behavioral response to cues associated with fine-scale oceanographic gradients and biological patchiness is functionally important and prevalent among copepods and likely has significant impacts on larger-scale distributional patterns.

The hydrodynamic disturbances of two species of krill: implications for aggregation structure

SUMMARY Krill aggregations vary in size, krill density and uniformity depending on the species of krill. These aggregations may be structured to allow individuals to sense the hydrodynamic cues of neighboring krill or to avoid the flow fields of neighboring krill, which may increase drag forces on an individual krill. To determine the strength and location of the flow disturbance generated by krill, we used infrared particle image velocimetry measurements to analyze the flow field of free-swimming solitary specimens (Euphausia superba and Euphausia pacifica) and small, coordinated groups of three to six E. superba. Euphausia pacifica individuals possessed shorter body lengths, steeper body orientations relative to horizontal, slower swimming speeds and faster pleopod beat frequencies compared with E. superba. The downward-directed flow produced by E. pacifica has a smaller maximum velocity and smaller horizontal extent of the flow pattern compared with the flow produced by E. superba, which suggests that the flow disturbance is less persistent as a potential hydrodynamic cue for E. pacifica. Time record analysis reveals that the hydrodynamic disturbance is very weak beyond two body lengths for E. pacifica, whereas the hydrodynamic disturbance is observable above background level at four body lengths for E. superba. Because the nearest neighbor separation distance of E. superba within a school is less than two body lengths, hydrodynamic disturbances are a viable cue for intraspecies communication. The orientation of the position of the nearest neighbor is not coincident with the orientation of the flow disturbance, however, which indicates that E. superba are avoiding the region of strongest flow.

Coordination of multiple appendages in drag-based swimming

Krill are aquatic crustaceans that engage in long distance migrations, either vertically in the water column or horizontally for 10 km (over 200 000 body lengths) per day. Hence efficient locomotory performance is crucial for their survival. We study the swimming kinematics of krill using a combination of experiment and analysis. We quantify the propulsor kinematics for tethered and freely swimming krill in experiments, and find kinematics that are very nearly metachronal. We then formulate a drag coefficient model which compares metachronal, synchronous and intermediate motions for a freely swimming body with two legs. With fixed leg velocity amplitude, metachronal kinematics give the highest average body speed for both linear and quadratic drag laws. The same result holds for five legs with the quadratic drag law. When metachronal kinematics is perturbed towards synchronous kinematics, an analysis shows that the velocity increase on the power stroke is outweighed by the velocity decrease on the recovery stroke. With fixed time-averaged work done by the legs, metachronal kinematics again gives the highest average body speed, although the advantage over synchronous kinematics is reduced.

Underwater flight by the planktonic sea butterfly.

ABSTRACT In a remarkable example of convergent evolution, we show that the zooplanktonic sea butterfly Limacina helicina ‘flies’ underwater in the same way that very small insects fly in the air. Both sea butterflies and flying insects stroke their wings in a characteristic figure-of-eight pattern to produce lift, and both generate extra lift by peeling their wings apart at the beginning of the power stroke (the well-known Weis-Fogh ‘clap-and-fling’ mechanism). It is highly surprising to find a zooplankter ‘mimicking’ insect flight as almost all zooplankton swim in this intermediate Reynolds number range (Re=10–100) by using their appendages as paddles rather than wings. The sea butterfly is also unique in that it accomplishes its insect-like figure-of-eight wing stroke by extreme rotation of its body (what we call ‘hyper-pitching’), a paradigm that has implications for micro aerial vehicle (MAV) design. No other animal, to our knowledge, pitches to this extent under normal locomotion. Highlighted Article: The zooplanktonic sea butterfly Limacina helicina ‘flies’ underwater using many of the same fluid dynamic ‘tricks’ that very small insects use to fly in air.

A study of biologically-inspired design as a context for enhancing student innovation

This article describes an investigation of the use of biologically-inspired design as a context from which to teach innovative design. The research compared ideation behavior among mechanical engineering students from a capstone design class to mechanical engineering students who had taken a semester-long course specifically focused on biologicallyinspired design. Both groups of students were presented with the same design challenge, and pre-established metrics were used to characterize the novelty and variety of the resultant designs generated by the students. The designs from the biologically-inspired design students had an average novelty score 80% higher than those from the control group of capstone students, and the result was statistically-significant. The biologically-inspired design students also had a 37% higher average variety score, although a small sample size led to a high variance and prevented statistical significance. The increased scores for novelty and variety imply a greater tendency toward innovative design among the biologically-inspired design students. The source of greater innovation is unclear but may be due to improved analogical reasoning capabilities among the biologically-inspired design students.

Particle and prey detection by mechanoreceptive copepods: a mathematical analysis

When particles move through fluids, they produce far-field pressure differences and near-field fluid deformations. Here we evaluate if a copepod, relying on mechanoreceptive antennulary setal hairs, can detect pressure changes caused by a variety of signal sources. We first provide a correction of the copepod mechanoreception model of Legier-Visser et al. (1986), showing how an object above a minimum size should be detectable. The pressure change ΔP created by an object of this minimum size was 385 dynes/cm2, based on biomechanical relationships for a rigid seta bending with respect to the exoskeletal body and using the neurophysiological detection threshold of a 10 nm bend of the sensory seta (Yen et al., 1992). The ΔP for: a 3 μm particle = 0.01 dynes/cm2, a 50 μm particle = 0.16 dynes/cm2, an escaping nauplius = 78 dynes/cm2, a revolving prey = 10−5 dynes/cm2, a 1 mm copepod escaping at 1 m/s at a distance of 1 mm from the mechanoreceptive sensory hairs of its captor = 312 dynes/cm2. Only the copepod escaping at high-speed close to the captor would create a pressure difference that could elicit a response. At this point, we conclude that pressure differences are rarely of a magnitude that is perceptible and that additional information must be derived for a copepod to detect prey. Other signals include fluid deformations as well as other types of stimuli (odor, shadows). Like most organisms, a copepod will rely on all sensory modalities to find food, avoid predators, and track mates, assuring their survival in the aquatic environment. It also is possible that the biomechanical model is insufficient for estimating pressure differences causing the cuticular deformation or that further analysis is necessary to improve our certainty of the sensitivity of the copepod seta.