A large-scale validation study of aircraft noise modeling for airport arrivals.

In the U.S., the Federal Aviation Administration's Aviation Environmental Design Tool (AEDT) is approved to predict the impacts of aircraft noise and emissions. AEDT's critical role in regulatory compliance and evaluating the environmental impacts of aviation requires asking how accurate are its noise predictions. Previous studies suggest that AEDT's predictions lack desired accuracy. This paper reports on a large-scale study, using 200 000 flight trajectories paired with measured sound levels for arrivals to Runways 28L/28R at San Francisco International Airport, over 12 months. For each flight, two AEDT studies were run, one using the approved mode for regulatory filing and the other using an advanced non-regulatory mode with exact aircraft trajectories. AEDT's per aircraft noise predictions were compared with curated measured sound levels at two locations. On average, AEDT underestimated LAmax by -3.09 dB and SEL by -2.04 dB, combining the results from both AEDT noise-modeling modes. Discrepancies appear to result from limitations in the physical modeling of flight trajectories and noise generation, combined with input data uncertainties (aircraft weight, airspeed, thrust, and lift configuration) and atmospheric conditions.

The metabolic consequences of repeated anoxic stress in the western painted turtle, Chrysemys picta bellii.

The painted turtle is known for its extreme tolerance to anoxia, but it is unknown whether previous experience with anoxic stress might alter physiological performance during or following a test bout of anoxia. Repeatedly subjecting 25°C-acclimated painted turtles to 2h of anoxic stress every other day for 19days (10 submergence bouts total) caused resting levels of liver glycogen to decrease by 17% and liver citrate synthase (CS) and cytochrome oxidase (COX) activities to increase by 33% and 112%, respectively. When the repeatedly submerged turtles were studied during a subsequent anoxic stress test, liver COX and CS activities decreased during anoxia to the same levels of naïve turtles, which were unchanged, and remained there throughout metabolic recovery. There were no effects of the repeated anoxia treatment on any of the other measured variables, which included lactate dehydrogenase and phosphofructokinase activities in liver, skeletal muscle, and ventricle, blood acid-base status, hemoglobin, hematocrit and plasma ion (Na, K, Ca, Mg, Cl) and metabolite concentrations (lactate, glucose, free-fatty acids), before, during, or after the anoxic stress test. We conclude that although painted turtles can show a physiological reaction to repeated anoxic stress, the changes appear to have no measurable effect on anaerobic physiological performance or ability to recover from anoxia.

Academic genealogy and direct calorimetry: a personal account.

Each of us as a scientist has an academic legacy that consists of our mentors and their mentors continuing back for many generations. Here, I describe two genealogies of my own: one through my PhD advisor, H. T. (Ted) Hammel, and the other through my postdoctoral mentor, Knut Schmidt-Nielsen. Each of these pathways includes distingished scientists who were all major figures in their day. The striking aspect, however, is that of the 14 individuals discussed, including myself, 10 individuals used the technique of direct calorimetry to study metabolic heat production in humans or other animals. Indeed, the patriarchs of my PhD genealogy, Antoine Lavoisier and Pierre Simon Laplace, were the inventors of this technique and the first to use it in animal studies. Brief summaries of the major accomplishments of each my scientific ancestors are given followed by a discussion of the variety of calorimeters and the scientific studies in which they were used. Finally, readers are encouraged to explore their own academic legacies as a way of honoring those who prepared the way for us.

Hibernation and gas exchange.

Hibernation in endotherms and ectotherms is characterized by an energy-conserving metabolic depression due to low body temperatures and poorly understood temperature-independent mechanisms. Rates of gas exchange are correspondly reduced. In hibernating mammals, ventilation falls even more than metabolic rate leading to a relative respiratory acidosis that may contribute to metabolic depression. Breathing in some mammals becomes episodic and in some small mammals significant apneic gas exchange may occur by passive diffusion via airways or skin. In ectothermic vertebrates, extrapulmonary gas exchange predominates and in reptiles and amphibians hibernating underwater accounts for all gas exchange. In aerated water diffusive exchange permits amphibians and many species of turtles to remain fully aerobic, but hypoxic conditions can challenge many of these animals. Oxygen uptake into blood in both endotherms and ectotherms is enhanced by increased affinity of hemoglobin for O2 at low temperature. Regulation of gas exchange in hibernating mammals is predominately linked to CO2/pH, and in episodic breathers, control is principally directed at the duration of the apneic period. Control in submerged hibernating ectotherms is poorly understood, although skin-diffusing capacity may increase under hypoxic conditions. In aerated water blood pH of frogs and turtles either adheres to alphastat regulation (pH ∼8.0) or may even exhibit respiratory alkalosis. Arousal in hibernating mammals leads to restoration of euthermic temperature, metabolic rate, and gas exchange and occurs periodically even as ambient temperatures remain low, whereas body temperature, metabolic rate, and gas exchange of hibernating ectotherms are tightly linked to ambient temperature.

Physiology of hibernation under the ice by turtles and frogs.

Successful overwintering under ice by an air-breathing vertebrate requires either effective aquatic respiration if dissolved O(2) is available or the capacity for prolonged anaerobic metabolism if O(2) supplies are limiting. Frogs can remain aerobic for many weeks when submerged at low temperature, even at water PO(2) as low as 30 mmHg, but are unable to survive even 1 week in anoxic water. Fuel reserves of hibernating frogs limit aerobic submergence, whereas acidosis may limit anoxic submergence. Freshwater turtles can also satisfy all or most of their O(2) needs in well-aerated water at low temperature by aquatic respiration, but certain species, in particular painted and snapping turtles, can also survive for up to 4-5 months without O(2). Key adaptations of the painted turtles, and presumably snapping turtles, include metabolic depression and the exploitation of the shell and other bones to buffer lactic acid. As in frogs, glycogen and glucose are the only fuel sources during anoxia, and stores do not seem to be limiting in the painted turtle. Significant differences in anoxia tolerance exist among chelonian species that can be attributed, at least in part, to the magnitude of metabolic depression, the effectiveness of lactic acid buffering, and the size of glycogen stores.

Fish assemblages in streams subject to anthropogenic disturbances along the natchez trace parkway, Mississippi, USA.

A three-year study (July 2000 - June 2003) of fish assemblages was conducted in four tributaries of the Big Black River: Big Bywy, Little Bywy, Middle Bywy and McCurtain creeks that cross the Natchez Trace Parkway, Choctaw County, Mississippi, USA. Little Bywy and Middle Bywy creeks were within watersheds influenced by the lignite mining. Big Bywy and Middle Bywy creeks were historically impacted by channelisation. McCurtain Creek was chosen as a reference (control) stream. Fish were collected using a portable backpack electrofishing unit (Smith-Root Inc., Washington, USA). Insectivorous fish dominated all of the streams. There were no pronounced differences in relative abundances of fishes among the streams (P > 0.05) but fish assemblages fluctuated seasonally. Although there were some differences among streams with regard to individual species, channelisation and lignite mining had no discernable adverse effects on functional components of fish assemblages suggesting that fishes in these systems are euryceous fluvial generalist species adapted to the variable environments of small stream ecosystems.

Analytical approaches for addressing the variation in back-calculated age-length relationships for fish.

Estimating an age-length relationship is a routine aspect of many fisheries studies and is simplified by the use of commercially available computer programs. These computer programs may be misleading since a result can be produced irrespective of the quality or the extent of the data, and there is some concern that back-calculated age-length relationships are sensitive to the sample size and composition. We investigated this issue by comparing estimates of mean back-calculated lengths at age and growth rates derived from subsets of a large sample of wild channel catfish Ictalurus punctatus (N=788) collected in 2001 and 2002 from 9 rivers in Mississippi, United States. Estimates of growth rate varied among subsets consisting of individual year class (2-6) of channel catfish separated from the overall sample. For nine subsets, comprising randomly-selected and increasing proportions of the overall sample (20%-100% at 10% increments of the overall sample), growth was similar. However, growth differed for a subset representing a random 10% of the overall sample. Lengths at age and growth rates derived from each of the 2001 and 2002 components of the sample both differed. All results were significant at P < 0.05.

Lactate metabolism in anoxic turtles: an integrative review.

Painted turtles can accumulate lactic acid to extremely high concentrations during long-term anoxic submergence, with plasma lactate exceeding 200 mmol l(-1). The aims of this review are twofold: (1) To summarize aspects of lactate metabolism in anoxic turtles that have not been reviewed previously and (2) To identify gaps in our knowledge of turtle lactate metabolism by comparing it with lactate metabolism during and after exercise in other vertebrates. The topics reviewed include analyses of lactate's fate during recovery, the effects of temperature on lactate accumulation and clearance, the interaction of activity and recovery metabolism, fuel utilization during recovery, stress hormone responses during and following anoxia, and cellular lactate transport mechanisms. An analysis of lactate metabolism in anoxic turtles in the context of the 'lactate shuttle' hypothesis is also presented.

Effects of temperature on anoxic submergence: skeletal buffering, lactate distribution, and glycogen utilization in the turtle, Trachemys scripta.

To test the hypothesis that submergence temperature affects the distribution of the lactate load and glycogen utilization during anoxia in turtles, we sampled a variety of tissues after 7 days, 24 h, and 4 h of anoxic submergence at 5, 15, and 25 degrees C, respectively. These anoxic durations were chosen because we found that they produced similar decreases in plasma HCO(3)(-) ( approximately 18-22 meq/l). The sampled tissues included ventricle, liver, small intestine, carapace, and the following muscles: flexor digitorum longus, retrahens capitis, iliofibularis, and pectoralis. Shell and skeleton sequestered 41.9, 34.1, and 26.1% of the estimated lactate load at 5, 15, and 25 degrees C. The changes in plasma Ca(2+) and Mg(2+), relative to the estimated lactate load, decreased with increased temperature, indicating greater buffer release from bone at colder temperatures. Tissue lactate contents, relative to plasma lactate, increased with the temperature of the submergence. Glucose mobilization and tissue glycogen utilization were more pronounced at 15 and 25 degrees C than at 5 degrees C. We conclude that, in slider turtles, the ability of the mineralized tissue to participate in the buffering of lactic acid during anoxia is inversely related to temperature, causing the lactate burden to shift to the tissues at warmer temperatures. Muscles utilize glycogen during anoxia more at warmer temperatures.

Comparative shell buffering properties correlate with anoxia tolerance in freshwater turtles.

Freshwater turtles as a group are more resistant to anoxia than other vertebrates, but some species, such as painted turtles, for reasons not fully understood, can remain anoxic at winter temperatures far longer than others. Because buffering of lactic acid by the shell of the painted turtle is crucial to its long-term anoxic survival, we have tested the hypothesis that previously described differences in anoxia tolerance of five species of North American freshwater turtles may be explained at least in part by differences in their shell composition and buffering capacity. All species tested have large mineralized shells. Shell comparisons included 1) total shell CO2 concentration, 2) volume of titrated acid required to hold incubating shell powder at pH 7.0 for 3 h (an indication of buffer release from shell), and 3) lactate concentration of shell samples incubated to equilibrium in a standard lactate solution. For each measurement, the more anoxia-tolerant species (painted turtle, Chrysemys picta; snapping turtle, Chelydra serpentina) had higher values than the less anoxia-tolerant species (musk turtle, Sternotherus odoratus; map turtle, Graptemys geographica; red-eared slider, Trachemys scripta). We suggest that greater concentrations of accessible CO2 (as carbonate or bicarbonate) in the more tolerant species enable these species, when acidotic, to release more buffer into the extracellular fluid and to take up more lactic acid into their shells. We conclude that the interspecific differences in shell composition and buffering can contribute to, but cannot explain fully, the variations observed in anoxia tolerance among freshwater turtles.

Lactate uptake by skeletal bone in anoxic turtles, Trachemys scripta.

Previous studies have shown that freshwater turtle shells can accumulate lactate during periods of anoxic submergence. Our objective in this study was to determine lactate uptake in other parts of the turtle's skeleton. We measured lactate concentration of 7 skeletal elements and 4 shell samples of red-eared slider turtles, Trachemys scripta, in control animals (N=12) and in animals following submergence for 4-5 days in N(2)-equilibrated water at 10 degrees C (N=8). We also collected blood samples and measured blood pH, PCO(2), and PO(2), and plasma lactate. Contralateral bone samples from 6 control turtles were analyzed for % water and mineral composition; bone from the other 6 were equilibrated with lactate solution in vitro. Anoxic submergence resulted in a combined respiratory/non-respiratory (lactic) acidosis and plasma lactate of 45.6+/-2.5 mmol l(-1). Shell and skeletal lactates all increased significantly in the anoxic animals (30.1-43.9 mmol kg(-1)) with limb bones having the highest levels and skull the least. Skeletal samples equilibrated in lactate solution in vitro for 2 days accumulated lactate in similar fashion with limb bones, except for fibula, higher, and skull significantly less than other bones. We conclude that the entire skeleton of the red-eared slider, like its shell, sequesters lactate and contributes thereby to lactic acid buffering.

Tissue glycogen and extracellular buffering limit the survival of red-eared slider turtles during anoxic submergence at 3 degrees C.

The goal of this study was to identify the factors that limit the survival of the red-eared slider turtle Trachemys scripta during long-term anoxic submergence at 3 degrees C. We measured blood acid-base status and tissue lactate and glycogen contents after 13, 29, and 44 d of submergence from ventricle, liver, carapace (lactate only), and four skeletal muscles. We also measured plasma Ca(2+), Mg(2+), Na(+), K(+), Cl(-), inorganic phosphate (P(i)), lactate, and glucose. After 44 d, one of the six remaining turtles died, while the other turtles were in poor condition and suffered from a severe acidemia (blood pH = 7.09 from 7.77) caused by lactic acidosis (plasma lactate 91.5 mmol L(-1)). An initial respiratory acidosis attenuated after 28 d. Lactate rose to similar concentrations in ventricle and skeletal muscle (39.3-46.1 micromol g(-1)). Liver accumulated the least lactate (21.8 micromol g(-1)), and carapace accumulated the most lactate (68.9 micromol g(-1)). Plasma Ca(2+) and Mg(2+) increased significantly throughout submergence to levels comparable to painted turtles at a similar estimated lactate load. Glycogen depletion was extensive in all tissues tested: by 83% in liver, by 90% in ventricle, and by 62%-88% in muscle. We estimate that the shell buffered 69.1% of the total lactate load, which is comparable to painted turtles. Compared with painted turtles, predive tissue glycogen contents and plasma HCO(-)(3) concentrations were low. We believe these differences contribute to the poorer tolerance to long-term anoxic submergence in red-eared slider turtles compared with painted turtles.

Endothelin-1 causes systemic vasodilatation in anaesthetised turtles (Trachemys scripta) through activation of ETB-receptors.

The effects of endothelin-1 (ET-1) on systemic and pulmonary circulation were investigated in anaesthetised freshwater turtles (Trachemys scripta) instrumented with arterial catheters and blood flow probes. Bolus intra-arterial injections of ET-1 (0.4-400 pmol kg(-1)) caused a dose-dependent systemic vasodilatation that was associated with a decrease in systemic pressure (P(sys)) and a rise in systemic blood flow (Q(sys)), causing systemic conductance (G(sys)) to increase. ET-1 had no significant effects on the pulmonary vasculature, heart rate (fh) or total stroke volume (Vs(tot)). This response differs markedly from mammals, where ET-1 causes an initial vasodilatation that is followed by a pronounced pressor response. In mammals, the initial dilatation is caused by stimulation of ET(B)-receptors, while the subsequent constriction is mediated by ET(A)-receptors. In the turtles, infusion of the ET(B)-receptor agonist BQ-3020 (150 pmol kg(-1)) elicited haemodynamic changes that were similar to those of ET-1, and the effects of ET-1 were not affected by the ET(A)-antagonist BQ-610 (0.15 micromol kg(-1)). Conversely, all effects of ET-1 were virtually abolished after specific ET(B)-receptor blockade with the ET(B)-antagonist BQ-788 (0.15 micromol kg(-1)). The subsequent treatment with the general ET-receptor antagonist tezosentan (15.4 micromol kg(-1)) did not produce effects that differed from the treatment with ET(B)-antagonist, and the blockade of ET-1 responses persisted. This present study indicates, therefore, that ET(B)-receptors are responsible for the majority of the cardiovascular responses to ET-1 in Trachemys.

Effect of graded hypoxic and acidotic stress on contractile force of heart muscle from hypoxia-tolerant and hypoxia-intolerant turtles.

Previous studies have shown that isometric contractile force of in vitro cardiac muscle from the anoxia-tolerant painted turtle, Chrysemys picta bellii, decreases when anoxic and when acidotic. This study sought to define the thresholds for these responses in the isolated ventricular strips of the painted turtle and in the anoxia-intolerant softshell turtles, Apalone spinifera. The ventricular strips were exposed to HCO3- Ringer's solution equilibrated at P(O2) 156, 74, 37, 19, and 0 mmHg (45 min at each grade), at both pH 7.0 and at pH 7.8. Strips were also exposed to graded lactic acidosis with intervals between pH 6.8 and pH 7.8 at P(O2) 156 mmHg (softshell) or 37 mmHg (painted). In painted turtle strips at pH 7.8, force remained at control levels until it decreased by 30% at P(O2) 19 mmHg. No further significant decrease occurred at P(O2) 0. In contrast, softshell turtle muscle force did not fall significantly until P(O2) reached 0. When graded hypoxia was imposed at pH 7.0, strips from both species were more sensitive to hypoxia, but the softshell force decreased at a higher P(O2) than the painted turtle (P(O2) 156 mmHg vs. 37 mmHg), its force fell to a lower level at P(O2) 0 (22 % of control vs. 40 % of control), and unlike painted turtle heart muscle, softshell muscle did not recover fully. In summary, these data indicate that ventricular strips of the painted turtle are no more tolerant of hypoxia alone than strips from the softshell turtle, but that when hypoxia is combined with acidosis, the painted turtle heart muscle functions significantly better during the exposure and recovers more fully after exposure.

The role of mineralized tissue in the buffering of lactic acid during anoxia and exercise in the leopard frog Rana pipiens.

To evaluate the role of mineralized tissues of the leopard frog in buffering acid, we analyzed the composition of femur and auditory capsule, the latter of which encloses a portion of the endolymphatic lime sacs, and investigated the extent to which these tissues are involved in buffering lactic acid after 2.5 h of anoxia and 10-19 min of strenuous exercise at 15 degrees C. We analyzed the following tissues for lactate: plasma, heart, liver, gastrocnemius muscle, femur, auditory capsule and carcass. Plasma [Ca(2+)], [Mg(2+)], [inorganic phosphate (P(i))], [Na(+)] and [K(+)] were also measured. Femur Ca(2+), P(i) and CO(3)(2-) compositions were similar to bone in other vertebrates. Auditory capsule had significantly more CaCO(3) than femur. Lactate was significantly elevated in all tissues after anoxia and exercise, including femur and auditory capsule. Anoxia increased plasma [Ca(2+)], [Mg(2+)], [P(i)] and [K(+)] and had no effect on plasma [Na(+)]. Exercise increased plasma [Mg(2+)], [P(i)] and [K(+)] and had no effect on plasma [Ca(2+)] or [Na(+)]. The skeleton and endolymphatic lime sacs buffered 21% of the total lactate load during anoxia, and 9% after exercise. The exact contribution of the entire endolymphatic sac system to lactate buffering could not be determined in the present study. We conclude that the mineralized tissues function as buffers during anoxia and exercised induced lactic acidosis in amphibians.

Surviving extreme lactic acidosis: the role of calcium lactate formation in the anoxic turtle.

During prolonged anoxia at low temperature, freshwater turtles develop high plasma concentrations of both lactate and calcium. At these concentrations the formation of the complex, calcium lactate, normally of little biological significance because of the low association constant for the reaction, significantly reduces the free concentrations of both lactate and calcium. In addition, lactate is taken up by the shell and skeleton to an extent that strongly indicates that calcium lactate formation participates in these structures as well. The binding of calcium to lactate thus contributes to the efflux of lactic acid from the anoxic cells and to the exploitation of the powerful buffering capacity of the shell and skeleton.

Avenues of extrapulmonary oxygen uptake in western painted turtles (Chrysemys picta belli) at 10 degrees C.

The major avenues of extrapulmonary oxygen uptake were determined on submerged western painted turtles (Chrysemys picta bellii) at 10 degrees C by selectively blocking one or more potential pathways for exchange. Previous work indicated that the skin, the cloaca, and the buccopharyngeal cavity can all contribute significantly in various species of turtles. O(2) uptake was calculated from the rate of fall in water P(O(2)) in a closed chamber. Two series of experiments were conducted: in Series 1, each of the potential avenues was mechanically blocked either singly or in combination; in Series 2, active cloacal and buccal pumping were prevented pharmacologically using the paralytic agent rocuronium. In addition in Series 2, N(2)-breathing preceded submergence in some animals and in one set of Series 2 experiments arterial blood was sampled and analyzed for pH, lactate, P(O(2)), and P(CO(2)). Results in both Series 1 and Series 2 revealed that prevention of cloacal and/or buccopharyngeal exchange did not significantly affect total O(2) uptake. Interfering with skin diffusion in Series 1, however, significantly reduced O(2) uptake by 50%. N(2)-breathing prior to submergence in Series 2 did not affect O(2) uptake in paralyzed turtles but significantly increased uptake in unparalyzed turtles without catheters. Blood analysis revealed that all submerged turtles developed lactic acidosis, but the rate of rise in lactate was significantly lower in paralyzed animals. We conclude that passive diffusion through the integument is the principal avenue of aquatic O(2) uptake in this species.

Lung collapse among aquatic reptiles and amphibians during long-term diving.

Numerous aquatic reptiles and amphibians that typically breathe both air and water can remain fully aerobic in normoxic (aerated) water by taking up oxygen from the water via extrapulmonary avenues. Nevertheless, if air access is available, these animals do breathe air, however infrequently. We suggest that such air breathing does not serve an immediate gas exchange function under these conditions, nor is it necessarily related to buoyancy requirements, but serves to keep lungs inflated that would otherwise collapse during prolonged submergence. We also suggest that lung deflation is routine in hibernating aquatic reptiles and amphibians in the northern portions of their ranges, where ice cover prevents surfacing for extended periods.