Stratigraphic Variation Within Polar Firn Caused by Differential Accumulation and Ice Flow: Interpretation of a 400 Mhz Short-Pulse Radar Profile from West Antarctica

We investigate causes of the stratigraphic variation revealed in a 177 km, 400 MHz short-pulse radar profile of firn from West Antarctica. The profile covers 56 m depth, and its direction was close to those of the ice flow and mean wind. The average, near-surface accumulation rates calculated from the time delays of one radar horizon consistently show minima on leeward slopes and maxima on windward slopes, confirming an earlier study based on stake observations. The stratigraphic variation includes up to 30 m depth variation in individual horizons over tens of km, fold limbs that become progressively steeper with depth, and fold-hinge loci that change direction or propagate down-ice with depth over distances far less than predicted by the ice speeds. We use an accumulation rate model to show how local rate anomalies and the effect of ice speed upon a periodic variation in accumulation rate cause these phenomena, and we reproduce two key features seen in the stratigraphic variations. We conclude that the model provides an explanation of changes in spatial stratigraphy and local measures of accumulation history given the constraints of surface topography, ice and wind velocities, and a general accumulation rate for an area.

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.

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.