Animal Guts as Nonideal Chemical Reactors: Partial Mixing and Axial Variation in Absorption Kinetics

Animal guts have been idealized as axially uniform plug-flow reactors (PFRs) without significant axial mixing or as combinations in series of such PFRs with other reactor types. To relax these often unrealistic assumptions and to provide a means for relaxing others, I approximated an animal gut as a series of n continuously stirred tank reactors (CSTRs) and examined its performance as a Function of n. For the digestion problem of hydrolysis and absorption in series, I suggest as a first approximation that a tubular gut of length L and diameter D comprises n=L/D tanks in series. For n greater than or equal to 10, there is little difference between performance of the nCSTR model and an ideal PFR in the coupled tasks of hydrolysis and absorption. Relatively thinner and longer guts, characteristic of animals feeding on poorer forage, prove more efficient in both conversion and absorption by restricting axial mixing, in the same total volume, they also give a higher rate of absorption. I then asked how a fixed number of absorptive sites should be distributed among the n compartments. Absorption rate generally is maximized when absorbers are concentrated in the hindmost few compartments, but high food quality or suboptimal ingestion rates decrease the advantage of highly concentrated absorbers. This modeling approach connects gut function and structure at multiple scales and can be extended to include other nonideal reactor behaviors observed in real animals

Animal Guts as Ideal Chemical Reactors: Maximizing Absorption Rates

I solved equations that describe coupled hydrolysis in and absorption from a continuously stirred tank reactor (CSTR), a plug flow reactor (PFR), and a batch reactor (BR) for the rate of ingestion and/or the throughput time that maximizes the rate of absorption (=gross rate of gain from digestion). Predictions are that foods requiring a single hydrolytic step (e.g., disaccharides) yield ingestion rates that vary inversely with the concentration of food substrate ingested, whereas foods that require multiple hydrolytic and absorptive reactions proceeding in parallel (e.g., proteins) yield maximal ingestion rates at intermediate substrate concentrations. Counterintuitively, then, animals acting to maximize their absorption rates should show compensatory ingestion (more rapid feeding on food of lower concentration), except for the lower range of diet quality fur complex diets and except for animals that show purely linear (passive) uptake. At their respective maxima in absorption rates, the PFR and BR yield only modestly higher rates of gain than the CSTR but do so at substantially lower rates of ingestion. All three ideal reactors show milder than linear reduction in rate of absorption when throughput or holding time in the gut is increased (e.g., by scarcity or predation hazard); higher efficiency of hydrolysis and extraction offset lower intake. Hence adding feeding costs and hazards of predation is likely to slow ingestion rates and raise absorption efficiencies substantially over the cost-free optima found here

Deep-sea species diversity: Does it have a characteristic scale?

Dispersion patterns and species diversities of deep-sea macrobenthos were examined for evidence that diversity-controlling processes operate predominantly on any one of several spatial scales. Identification of such scales, if any, would aid in the identification of the diversity-regulating processes themselves. The specific hypothesis that species diversity is independent of scale and location within the deep sea was tested with replicated..

Collaborative Proposal: Form and function of phytoplankton in unsteady, low Reynolds-number flows

Small-scale flow dynamics at low Reynolds numbers (Re) are important to phytoplankton cells in delivery of nutrients, sensory detection by and physical encounter with herbivores, accumulation of bacterial populations in the phycosphere or region immediately surrounding phytoplankton cells and coagulation of cells themselves as a mechanism terminating blooms. In nature most phytoplankton experience unsteady flows, i.e., velocities near the cells that vary with time due to the intermittency of turbulence and to discontinuous, spatially distributed pumping by herbivores. This unsteadiness has not previously been taken into account in models or measurements with plankton. Moreover, there have been decade- and century- long lags in moving relevant models of unsteady flow effects at low Re from applied mathematics and engineering to ecological applications. Engineering models show unsteady effects due to the history of formation of spatially extensive flow perturbations or wakes should be important to unsteady motions of moderately small biota. This project will address these affects. Non-swimming phytoplankton, and in particular diatoms, will be used as the simplest case where important unsteady flow behaviors should arise. This research activity will include a multi-level educational program, aimed at graduate research assistants, undergraduate research interns, undergraduate marine sciences majors and high-school teachers. Low-Re behaviors afford unusual opportunities to experience how mathematics, physics and biology inseparably catalyze understanding of phenomena that run counter to intuition. This activity will also include international collaborations with world experts on organism-flow interaction in Cambridge (T.J. Pedley) and Copenhagen (T. Kiorboe & A.W. Visser). The overall goals of the activity are to accelerate the flow of understanding from modelers to measurers to users of the information and back again. Educational materials that project U.S. national standards will be developed during intensive summer workshops with the high-school teachers and be made available on the web. Unsteady flow effects on phytoplankton will be predicted with explicit models based on singularity solutions (that involve the useful simplification that force is applied to the fluid at a small number of points) and mathematical models that include both the near field at low Re and the far field over a range of Re, both representative of nature. Singularity solutions allow explicit treatment of the role of complex cell shapes. Scaled-up analog models will be placed in a large Couette vessel to better visualize behaviors for both the research and teaching efforts. Natural-scale, but simplified, unsteady flows will be produced in smaller Couettes (nested, counter-rotating cylinders with seawater in the gap between the two cylinders) containing live phytoplankton and will be quantified by magnifying, particle-imaging velocimetry (PIV). Image analysis will be used to measure translation, rotation and flexural deformation of the phytoplankton. These studies will test various hypotheses derived from the general thesis that cell shapes and mechanical properties interact with unsteady flows to produce potentially fitness-enhancing, relative motions of the cell or chain and its surrounding fluids. A basic hypothesis is that unsteady fluid motion will interact with bending of cells to produce relative motion of fluid and phytoplankter. A very exciting prospect is that periodic instabilities known to arise at low Re may allow flexible organisms to act as self-organizing engines - through elasticity to harness energy from decaying turbulence and thereby move relative to the fluid. It is also expected that this study of passively bending structures in unsteady flows will help to understand the use of flexible appendages in swimming. The work is likely to aid significantly in associating functions with the shapes and spines of microplankton that are used in the identification of fossil specimens. By including relevant, unsteady fluid motions at low Re, the study will also provide firmer linkages between form and function in living plankton in the size range from 10 - 1000 mm that many large phytoplankton, invertebrate and fish larvae and other small zooplankton occupy

Developing Tools to Evaluate Spawning & Fertilization Dynamics of the Giant Sea Scallop — Phase II: Field Trials in Experimental Populations

Objective 1 — Sperm advection-diffusion model: Develop a two-dimensional spatial model to predict the concentration o f sperm and effective range of fertilization in a sperm plume at varying distances from a source population of spawning males under scenarios of synchronous and asynchronous spawning. Objective 2 — Fertilization assays in field populations: Conduct a time series of fertilization assays over experimental populations of scallops to (a) further develop the methodology to assess ambient sperm loads in scallop populations over the course of the spawning season, (b) compare model predictions about spatial patterns of sperm concentration and fertilization generated in Objective 1 to field observations on experimental populations, (c) determine the influence of differences in the sperm plume arising from two experimental populations spanning a ten-fold difference in male density, and (d) conduct laboratory flume experiments to evaluate potential biases introduced by Nitex egg baskets to estimates of absolute and relative rates of fertilization

Effects of bivalve siphonal currents on the settlement of inert particles and larvae

Dye studies in a flume revealed that the strong exhalent siphonal current of an individual cockle, Clinocardium nuttallii, behaved much like a cylinder held in the flow, shedding vortices downstream. The inhalent flow was much slower and more diffuse, its effects being limited to the lowermost few centimeters above the bottom. Flume experiments with inert particles having settling velocities similar to those of polychaete larvae revealed that the vortex shedding from the excurrent jet led to variability in deposition of the particles a few centimeters downstream of the jet, and that neither the jet nor the incurrent flow substantially changed the mean number of particles depositing per unit area of bed. Field observations within a few days after settlement of Hobsonia florida, an ampharetid polychaete, using ecologically similar but nonplanktonically recruiting oligochaetes as an internal control, showed similarly enhanced variability in recruitment within a few centimeters of the siphon of resident Mya arenaria (soft-shelled clam). We could find no evidence that isolated clams impede settlement in their immediate surroundings and found, instead, some indication of local settlement being enhanced by the flow convergence toward the incurrent siphon. We thus suggest that any negative influence of suspension-feeding bivalves upon settlement is a larger-scale phenomenon caused by depletion of recruits through the integrated filtering activities of individuals upstream of the settlement site. Hence manipulation of bivalve density in small plots may not be very informative regarding influences upon larval settlement

Hadal community structure: Implications from the Aleutian Trench

A 0.25-m2 box core from the Aleutian Trench (50°58.0\u27N, 171°37.5\u27W) was used to generate hypotheses concerning the regulation of benthic community structure in oceanic trenches. High food supply and the concentrating effects of trench topography are suggested by the unexpectedly high standing crop (1272 individuals of macrofaunal taxa m2) and by the feeding modes of the captured polychaetes...

CMG Collaborative Research: Interactions of Phytoplankton with Dissipative Vortices

The aim of this project is to develop coordinated laboratory experiments and computational models to address a fundamental question in oceanography concerning magnitudes and mechanisms of turbulence effects on phytoplankton and other particles at the spatial scale of individual organisms. The importance of external energy in the form of turbulence in determining relative success of different kinds of phytoplankton dates to the seminal analysis of Munk and Riley (1952) and Margalef (1978). Margalef\u27s mandala asserts that high nutrient concentrations and turbulence intensities favor dominance by diatoms, whereas low values favor non-red-tide dinoflagellates. Subsequent work has revealed a wide spectrum of turbulence effects among species of dinoflagellates, including growth stimulation. The physicochemical mechanisms that govern these effects largely remain to be determined, however.Through iteration between innovative numerical models and experiments, the investigators will close a growing gap between textbook understanding of turbulent flows and understanding of consequences for suspended organisms and particles. Models and experiments have used one-dimensional shear to assess turbulence effects at the level of single cells and chains. Effects of fluid straining on concentration fields and cell rotation have been predicted, and effects on cell growth and motion, documented. Current understanding of turbulence, however, places greater emphasis on vorticity, gradients in vorticity and vortices at dissipation scales experienced by individual phytoplankton cells. We propose to develop a framework for both numerical and analog evaluation of effects that cells experience from being in and near viscous-scale vortices, that capture effects of vorticity as well as fluid deformation, evolution of concentration fields, and fluid-structure interactions. Roles of vorticity and gradients in vorticity in determining cell motions and thereby shaping concentration fields have been underappreciated, partly because a signature feature of turbulence, i.e., vortex stretching, is impossible in the two-dimensional flows that so far have been used as theoretical models and the primary basis of analog devices.Numerical approaches will use two simplified models of small-scale vortex structure and evolution, the Burgers vortex and the Lundgren stretched-spiral vortex, giving particular attention to diffusion of vorticity within and away from both. Both decaying and equilibrium vortices will be explored. Models of cells and chains of cells will be based on shapes and flexural stiffnesses of actual cells and chains. Each will be placed successively at a range of positions within and near a vortex and will be fully coupled mechanically to the fluid. Behaviors of interest are cell and chain translation, rotation and deformation and their feedbacks to local velocity and vorticity fields that could be used by grazers to locate a cell. Also to be modeled is the diffusion of scalars (nutrients with cell as sink or metabolites with cell as source), allowing calculation of diffusive fluxes for nutrient acquisition and prediction of chemical fields used by grazers. The investigators will further take advantage of their existing models of flow around flagella to include motile dinoflagellates in the modeling and measurement scheme.Analog experiments will exploit the fact that flows near Kolmogorov scale are dominated by viscosity, just as in earlier Couette experiments, but will incorporate realistic, 3D time variation. Borrowing from a burgeoning variety of geometries used in microfluidics, the investigators will construct a variety of small devices that utilize shed vortex streets, mild jets and cavity flows to match deformation rates, vorticities and gradients in them that produce interesting effects on phytoplankton in their numerical models of vortices. These analogs will be used to test the model predictions and to pose new questions of the models.Broader impacts: Results for phytoplankton extend easily to other important phenomena such as diffusion of attractants from eggs spawned in a turbulent environment (e.g., by abalone and other benthic invertebrates) and corresponding sperm swimming capabilities. They have implications for other important encounter processes such as particle coagulation and sedimentation, hydrosol filtration, and predator-prey interactions. This new approach provides both a natural bridge from larger-scale, direct numerical simulation (DNS) models of turbulence to these individual-scale effects of turbulence and a logical path to parameterizing these effects in larger-scale fluid dynamic models.Turbulence intensity is one of the parameters most likely to be influenced by climate change, and the investigators will work closely with the Center for Ocean Sciences Education Excellence Ocean Systems (COSEE-OS) that has chosen oceans under climate change as its major focus. They will also build on their history of providing teaching and outreach materials in biomechanics at low Reynolds numbers for graduate students, undergraduates and high-school teachers. They will complement both of these efforts with professionally produced, evocative visual animations of the important phenomena that they identify for incorporation into the COSEE-OS website

New resource axes for deposit feeders?

Recent work on selectivity in deposit feeders has focused on the importance of particle size. In field experiments with exotic sediments of known characteristics (glass beads), we demonstrate that selective ingestion in a multitentaculate, surface deposit feeding ampharetid polychaete depends upon particle specific gravities and surface textures. The degree of selectivity for specific gravity is shown to be dependent upon worm size...