Influence of two different green algal diets on specific dynamic action and incorporation of carbon into biochemical fractions in the copepod Acartia tonsa

Previous studies have shown that the two green algae Tetraselmissp. (Prasinophyceae) and Dunaliella tertiolecta (Chlorophyceae)induce high and low egg production rates in Acartia tonsa. Theprimary goal of the present study was to investigate if thisis attributable to differences in the specific dynamic action(SDA) of the two diets. Secondly, we wanted to investigate ifany qualitative differences in the incorporation of nutritionalconstituents from the two diets are influencing SDA. The functionalresponse of ingestion was very different with the two diets.Ingestion of T. impellucida was relatively high even at lowfood concentrations with a maximum of 19 µg C ind^-1 day^-1.The functional response was more clearly sigmoidal on D. tertiolectawith a maximum of 7.3 µg C ind^-1 day^-1. The higher ingestionrate of T. impellucida also induced higher respiration rates.Maximum respiration rates were 3.0 nl O₂ ind^-1 min^-1 on T. impellucidaand 1.5 nl O₂ ind^-1 min^-1 onD. tertiolecta. This created significantlydifferent SDA coefficients: 0.19 on T. impellucida and 0.06on D. tertiolecta, which implies that the magnitude of SDA isstrongly influenced by the composition of the diet. The incorporationof carbon into lipids was significantly higher on D. tertiolecta.However, because of lack of longer chain fatty acids in D. tertiolectathe copepods did not benefit from this. Thus, the proportionof carbon allocated to egg lipids was much lower than when feedingon T. impellucida. Acartia tonsa incorporated relatively morecarbon into proteins when feeding on T. impellucida than onD. tertiolecta. Since protein synthesis is energetically verydemanding this is probably the reason for the higher SDA coefficientin those feeding on T. impellucida.     

Protein synthesis and specific dynamic action in crustaceans: effects of temperature

Temperature influences the specific dynamic action (SDA), or rise in oxygen uptake rate after feeding, in eurythermal and stenothermal crustaceans by changing the timing and the magnitude of the response. Intra-specific studies on the eurythermal crab, Carcinus maenas, show that a reduction in acclimation temperature is associated with a decrease in SDA magnitude, resulting from an increase in SDA duration but a decrease in peak factorial scope (the factorial rise in peak SDA over prefeeding values). Inter-specific feeding studies on stenothermal polar isopods revealed marked differences in SDA response between the Antarctic species, Glyptonotus antarcticus and the Arctic species, Saduria entomon. Compared to S. entomon held at 4 and 13 degrees C, the SDA response in G. antarcticus held at 1 degrees C was characterised by a lower absolute oxygen uptake rate at peak SDA and an extended SDA duration. At peak SDA, whole animal rates of protein synthesis increased in proportion to the postprandial increase in oxygen uptake rate in the Antarctic and the Arctic species. Rates of oxygen uptake plotted against whole animal rates of protein synthesis gave similar relationships in both isopod species, indicating similar costs of protein synthesis after a meal, despite their differences in SDA response and thermal habitat.     

Specific dynamic action and carbon incorporation in Calanus finmarchicus copepodites and females

Specific dynamic action (SDA) and incorporation of carbon into protein, lipids, and polysaccharides were measured in copepodite CV and adult female Calanus finmarchicus during the spring/summer growth season in Raunefjorden, Norway. Organic carbon from the food (Rhodomonas baltica) was allocated differently in the two developmental stages. Copepodites incorporated 50-80% of the carbon into lipids and only 7-22% into protein. Carbon incorporation into protein was higher in females, constituting 23% in May at 7 °C and 34% in June at 11 °C. This resulted in significant differences in the magnitude of SDA, measured as the increase in oxygen consumption during and after an 8-h feeding event. On average, the rate of carbon incorporation into protein was 2 times higher and the magnitude of SDA was 2.5 times higher in females than in copepodites. There was a significant correlation between the magnitude of SDA and carbon incorporation into protein suggesting that SDA is linked to protein synthesis. When comparing ATP equivalents of the magnitude of SDA with ATP equivalents of the total amount of carbon incorporated, more energy was produced than consumed. This supports speculations of an energy demand associated with a rapid turnover of newly synthesised protein.     

Comparison of the metabolism of Acartia clausi and A. tonsa: influence of temperature and salinity

In the Marseilles region (French Mediterranean coast), A. clausi is one of the most abundant copepod species of the Gulf of Fos while A. tonsa constitutes the almost exclusive copepod species of the Berre lagoon, a neighbouring semi-closed brackish area communicating with the gulf. As different ecophysiological capabilities to stand the various temperature, salinity and food conditions could explain why these two species do not coexist in the same environment, comparative experiments were performed on metabolism and feeding. The respiration and ammonia excretion of the two species were measured in different combinations of temperature (10, 15 and 20 degrees C) and salinity (15, 25 and 35 per thousand). For each temperature, at the salinity of 35 per thousand, respiration rates were less in A. clausi than in A. tonsa, the contrary being observed at the lowest salinity. At any temperature ammonia excretion was greater at the intermediate salinity in A. tonsa and least in A. clausi. In Acartia tonsa, Q(10) of respiration and excretion were minimum at the lowest salinity, while in A. clausi they were unaffected by salinity variation. The O:N atomic ratio (from respiration and ammonia excretion rates) was significantly more elevated in A. clausi (mean 21.2; range 13.6-28.7) than in A. tonsa (mean 11.3; range 4.2-25) suggesting a more proteinic oriented metabolism in the later. Feeding experiments where Dunaliella tertiolecta30 per thousand) or lagoon (<16 per thousand) salinity. The relationships between ingestion and food concentration in the two species were not significantly different. These different results are compared to other ecophysiological information concerning these Acartia species (survival tolerances, osmotic regulation, feeding behaviour) and are discussed in relation with the characteristics of their niches in the studied region.     

Effect of protein synthesis inhibitor cycloheximide on starvation,fasting and feeding oxygen consumption in juvenile spiny lobster Sagmariasus verreauxi

Metabolism in aquatic ectotherms evaluated by oxygen consumption rates reflects energetic costs including those associated with protein synthesis. Metabolism is influenced by nutritional status governed by feeding, nutrient intake and quality, and time without food. However, little is understood about contribution of protein synthesis to crustacean energy metabolism. This study is the first using a protein synthesis inhibitor cycloheximide to research contribution of cycloheximide-sensitive protein synthesis to decapod crustacean metabolism. Juvenile Sagmariasus verreauxi were subject to five treatments: 2-day fasted lobsters sham injected with saline; 2-day fasted lobsters injected with cycloheximide; 10-day starved lobsters injected with cycloheximide; post-prandial lobsters fed with squid Nototodarus sloanii with no further treatment; and post-prandial lobsters injected with cycloheximide. Standard and routine metabolic rates in starved lobsters were reduced by 32% and 41%, respectively, compared to fasted lobsters, demonstrating metabolic downregulation with starvation. Oxygen consumption rates of fasted and starved lobsters following cycloheximide injection were reduced by 29% and 13%, respectively, demonstrating protein synthesis represents only a minor component of energy metabolism in unfed lobsters. Oxygen consumption rate of fed lobsters was reduced by 96% following cycloheximide injection, demonstrating protein synthesis in decapods contributes a major proportion of specific dynamic action (SDA). SDA in decapods is predominantly a post-absorptive process likely related to somatic growth. This work extends previously limited knowledge on contribution of protein synthesis to crustacean metabolism, which is crucial to explore the relationship between nutritional status and diet quality and how this will affect growth potential in aquaculture species.

     

The relationship between specific dynamic action and growth in grass carp, Ctenophavyngodon idella (Val.)

Individual grass carp, Ctenopharyngodon idellu , were maintained in a respirometer for a month and fed pelleted diets containing various proportions of carbohydrate, fat and protein at different ration levels. Oxygen consumption was measured continuously, allowing the effects of consecutive daily feeding on respiration to be studied. The relationships established between daily food intake and oxygen consumption showed that, on average, 23.3% (high protein diet), 15.3% (high carbohydrate diet), 20.7% (high lipid diet) and 7.0% ( Lemnu diet ) of the absorbed energy was partitioned into specific dynamic action (SDA). (Here the term SDA is used to describe the oxygen consumption of a feeding fish in excess of the routine metabolic rate.) In terms of the overall energy budgets of growing fish, SDA represented between 12 and 58% of the total heat lost over the experimental period and was equivalent to between 14 and 33% of the consumed energy. Ration was positively correlated with heat loss due to total respiration ( r = 0.881) and with heat loss due to SDA ( r = 0.762). As ration increased, the size of SDA relative to total respiration increased. Significant positive correlations were found between oxygen consumption (total or due to SDA) and specific growth rate, and between oxygen consumption and the deposition of protein and energy. However, growth rate had a minimal influence on daily oxygen consumption when compared with food intake.     

The effect of meal composition on specific dynamic action in burmese pythons (Python molurus)

We quantified the specific dynamic action (SDA) resulting from the ingestion of various meal types in Burmese pythons (Python molurus) at 30 degrees C. Each snake was fed a series of experimental meals consisting of amino acid mixtures, simple proteins, simple or complex carbohydrates, or lipids as well as meals of whole animal tissue (chicken breast, beef suet, and mouse). Rates of oxygen consumption were measured for approximately 4 d after feeding, and the increment above standard metabolic rate was determined and compared to energy content of the meals. While food type (protein, carbohydrate, and lipid) had a general influence, SDA was highly dependent on meal composition (i.e., amino acid composition and carbohydrate structure). For chicken breast and simple carbohydrates, the SDA coefficient was approximately one-third the energetic content of the meal. Lard, suet, cellulose, and starch were not digested and did not produce measurable SDA. We conclude that the cost of de novo protein synthesis is an important component of SDA after ingestion of protein meals because (1) simple proteins, such as gelatin and collagen, did not stimulate levels of SDA attained after consumption of complete protein, (2) incomplete mixtures of amino acids failed to elicit the SDA of a complete mixture, and (3) the inhibition of de novo protein synthesis with the drug cycloheximide caused a more than 70% decrease in SDA. Stomach distension and mechanical digestion of intact prey did not cause measurable SDA.     

Effects of feeding on metabolic rate, and the Specific Dynamic Action in plaice, Pleuronectes platessa L.

The rate of oxygen consumption of plaice increases after feeding and declines to a resting level after 24–72 h. The maximum increase corresponds to a level which is approximately twice the resting rate of oxygen consumption. This increase corresponds to the Specific Dynamic Action (SDA) and increases in magnitude with increase in food intake. The magnitude is greatest with high protein content diets. The duration of the SDA effect is reduced with increase in temperature and increases with the percentage of protein in the diet.     

Naupliar development times and survival of the copepods Calanus helgolandicus and Calanus finmarchicus in relation to food and temperature

We measured egg hatching times and stage specific naupliar developmenttimes of the key calanoid copepods Calanus helgolandicus andC. finmarchicus in response to temperature, food quantity andfood type. Calanus helgolandicus development times decreasedwith increasing temperature and when fed Isochrysis galbana(4.4 µm ESD) rather than Prorocentrum micans (29.5 µmESD). Nauplii needed higher food carbon concentration to developpast the first feeding stage (N3) when fed I. galbana comparedwith P. micans. At low food carbon concentrations nauplii developedmore slowly past N3 than at more saturated levels. The survivalof nauplii fed P. micans increased with temperature, but starvednauplii survival decreased at higher temperatures. We parameterizeda temperature-dependent model of development for both specieswhich fitted the observed stage durations under non-limitingfood extremely well and demonstrated that C. finmarchicus developsfaster than C. helgolandicus. Further data are needed to clarifythe effect of food-temperature interactions on development rates.     

Production of Bacterial Substrate by Marine Copepods: Effect of Phytoplankton Biomass and Cell Size

Laboratory experiments were performed to evaluate the importanceof grazing activity of the marine copepod Acartia tonsa forproduction of substrate for bacteria. Acartia tonsa were feda range of concentrations of the nanoflagellate Rhodomonas baltica,the diatom Ditylum brightwelli and the dinoflagellate Ceratiumlineatum. Regardless of the concentration of R. baltica, nodetectable response in bacterial biomass was observed due tograzing. However, when A. tonsa grazed the larger phytoplanktoncells of D. brightwelli and C. lineatum, responses in bacterioplanktonwere observed. It was estimated that approximately 54–69%of the grazed carbon was lost to the surroundings when A. tonsawas feeding on these large phytoplankton species. The laboratoryresults were applied to a dataset from a coastal temperate ecosystem.This analysis showed that the copepod contribution to the DOCpool was as important as the leakage from the primary producers.It is concluded, that the DOM contribution from copepods willbe largest when grazing plankton communities are composed oflarge species.     

Specific dynamic action and feeding metabolism in common carp,Cyprinus carpio L.

• 1.1. Oxygen uptake attributable to Specific Dynamic Action (SDA) was measured in common carp, Cyprinus carpio L. (63.6–84.0 g) fed on 20, 35 and 50% dietary protein at 0.40 to 1.00% ration levels at 28°C.
• 2.2. After feeding both SDA magnitude and mean peak oxygen consumption increased directly with dietary protein and ration levels. SDA duration was not significantly related to dietary protein but significantly increased with ration levels.
• 3.3. SDA coefficients were 8.99, 13.51 and 15.94% with 20, 35 and 50% dietary protein showing a direction relationship to the protein content. The SDA coefficient did not change with ration size.
• 4.4. SDA models resulting from this work are of great interest for the aquaculturist, as post-feeding oxygen requirements in an intensive fish culture can be predicted where dietary protein and ration levels are known.

     

Energy metabolism and the postprandial response of the Chilean tarantulas, Euathlus truculentus (Araneae: Theraphosidae)

One of the most ubiquitous consequences of feeding in animals is specific dynamic action (SDA), a drastic increment in metabolic rate after a meal, which lasts from a few hours to several days. According to a recent exhaustive review by Secor (2009), studies in SDA are abundant, encompassing all kinds of vertebrates and invertebrates. However, important exceptions are arachnids, as few studies have characterized SDA in this group. Here, we measured the standard metabolic rate (SMR) of the Chilean tarantulas Euathlus truculentus (body mass=7.32±0.7 g; N=32; T(A)=25°C), its inter-individual variation (i.e., repeatability) and its SDA. We measured SMR three or four times in each individual, and we also conducted predation experiments where a prey was consumed by each spider, during a respirometry trial. The SMR of E. truculentus was 0.00049±0.000079 mlCO(2) g(-1) min(-1) which corresponds to 1524 μW (assuming a protein-based diet), 108.4% of the predicted value for arachnids. According to the standard nomenclature for SDA studies, the scope of the SDA for a meal size of 1.26±0.04 g (18% of the spider size) was 6.55±1.1 times the baseline, the time to peak was 45 min, and the magnitude of the SDA was 0.28±0.03 kj, which is 85% of the expected value for invertebrates. Our SMR data are in concordance with previous findings suggesting remarkably low energy metabolism in arachnids, compared with other arthropods. On the other hand, the exceedingly high scope of the postprandial response contrasts with the comparatively low SDA. This fact suggests that spiders spend most of the energy for digestion in a short period after prey capture, which could be a consequence of their external digestion.     

Postprandial responses in the African rhombig egg eater (Dasypeltis scabra)

The African rhombic egg eater (Dasypeltis scabra) is a colubrid snake feeding exclusively on bird eggs. Frequency of feeding is governed by the seasonal availability of bird eggs; i.e., long fasting intervals change with relatively short periods when plenty of food is available. Intermittent feeding snakes show a remarkable postprandial increase of metabolic rate and digestive organ size. The postprandial increase in metabolic rate (specific dynamic action, SDA) in snakes is affected by meal size, temperature, and meal composition. A major portion of SDA in snakes is allocated to gastric function and the breakdown of the meal. We hypothesize that SDA in egg eaters is lower than in other snake species, because egg eaters feed on “liquid” food that does not require enzymatic breakdown in the stomach. We also hypothesized that other components of the postprandial response of egg eaters (e.g., size changes of the intestine and the liver) do not differ from other snakes. The standard metabolic rate and metabolic response to feeding were measured using closed-chamber respirometry. Size changes of small intestine and liver were measured using high-resolution transcutaneous ultrasonography. Standard metabolic rates of fasting egg eaters were in the same range of mass specific values as known from other snakes. Within 24 h after feeding, oxygen consumption doubled and peaked at 2 days after feeding. At the same time, the size of the small intestine and the cross-sectional diameter of the liver increased. Within 2 days after feeding, the size of the mucosal epithelium doubled its thickness. Liver size increased significantly within 24 h reaching maximum size 2–4 days after feeding. The size of both organs returned to fasting values within 7–10 days after feeding. The postprandial response of African rhombic egg eaters shows the same pattern and dynamics as known from other snake species. However, the factorial increase of metabolic rate during SDA is the lowest reported for any snake. A comparison with literature data supports the idea that SDA is mainly determined by gastric function and that it is low in egg eaters because they do not have to break down solid meals in the stomach as other snake species do.     

Nuclear protein synthesis in a temperature-sensitive Chinese hamster ovary cell line at 40 degrees C

We examined the incorporation of radioactive amino acids into nuclear proteins occurring at nonpermissive conditions in tsH1 Chinese hamster ovary cells with a temperature-sensitive defect in cytosol nonmitochondrial protein synthesis. In leucine-free medium at 40 degrees C, total cellular protein synthesis declined by 1-1.5%/min. As reported by others, preincubating these cells at 42 degrees C for 5-10 min sharply increased the rate of decline. The synthesis of acidic nuclear proteins at nonpermissive conditions (40 degrees C + 300 micrograms/ml cycloheximide) was demonstrated by the nuclear incorporation of 3H-tryptophan. Radioactivity, seen by autoradiography to be associated with these isolated Triton-X-100-washed nuclei, was released after incubating labelled nuclei with proteolytic enzymes. During incubation of tsH1 cells at nonpermissive conditions, pulse/chase experiments were consistent with the loss of some nuclear radioactivity into the cytoplasm. The distribution of cytosol and nuclear proteins, labelled at permissive or nonpermissive conditions and separated by isoelectric focusing, differed quantitatively and probably qualitatively, confirming the residual synthesis of acidic nuclear proteins at 40 degrees C in the presence of cycloheximide. Most newly synthesized acidic proteins retained by nuclei from cells labeled at nonpermissive conditions were present in a transciptionally active chromatin fraction. Although under these conditions the apparent rate of cellular RNA synthesis was unchanged, inhibiting residual cycloheximide-resistant nuclear protein synthesis with puromycin proportionately reduced RNA synthesis. Preincubating cells with 20 micrograms/ml of actinomycin D did not inhibit residual labelling of nuclear proteins; effects on residual nuclear labelling of impaired mitochondrial respiration were ambiguous. Nuclear proteins labelled under nonpermissive conditions probably included some of the 'prompt' heat shock proteins recently described. Provided certain assumptions are correct, our results are consistent with very limited protein synthesis associated with and even intrinsic to cell nuclei. They also suggest that this residual cycloheximide-resistant protein synthesis could be concerned with optimum synthesis or processing of certain nuclear RNA species.     

Gastrointestinal blood flow and postprandial metabolism in swimming sea bass Dicentrarchus labrax

In trout and salmon, the metabolic costs of exercise and feeding are additive, which would suggest that gastrointestinal blood flow during exercise is maintained to preserve digestive and absorptive processes related to the specific dynamic action (SDA) of food. However, in most published studies, gastrointestinal blood flow drops during swimming, hypoxia, and general stress. To test whether gastrointestinal blood flow is spared during exercise after feeding, sea bass were instrumented with flow probes to measure cardiac output and celiacomesenteric blood flow while swimming in a respirometer before and after feeding. Swimming at 2 body lengths per second (bl s(-1)) increased metabolic rate considerably more than did feeding (208% vs. 32% increase, respectively, relative to resting), and a similar pattern was observed for cardiac output. In unfed fish, resting gastrointestinal blood flow was 13.8+/-0.5 mL min(-1) kg(-1). After feeding, resting gastrointestinal blood flow increased by 82% but then decreased progressively with increasing swimming speeds. At 2 bl s(-1), gastrointestinal blood flow in fed fish was not significantly different compared with that in unfed swimming fish, and, therefore, the data do not support the gastrointestinal sparing hypothesis. The magnitude of the SDA was maintained despite the decrease in gastrointestinal blood flow and the consequent reduction in oxygen supply to the gut. An estimate of maximal oxygen flow to the gastrointestinal tract after feeding yielded 2.6 mmol O(2) h(-1) kg(-1), but this amount is not able to cover the oxygen demand of 3.16 mmol O(2) h(-1) kg(-1). Therefore, the SDA must reflect metabolic processes in tissues other than those directly perfused by the celiacomesenteric artery.     

Algal toxins alter copepod feeding behavior

Using digital holographic cinematography, we quantify and compare the feeding behavior of free-swimming copepods, Acartia tonsa, on nutritional prey (Storeatula major) to that occurring during exposure to toxic and non-toxic strains of Karenia brevis and Karlodinium veneficum. These two harmful algal species produce polyketide toxins with different modes of action and potency. We distinguish between two different beating modes of the copepod's feeding appendages-a "sampling beating" that has short durations (<100 ms) and involves little fluid entrainment and a longer duration "grazing beating" that persists up to 1200 ms and generates feeding currents. The durations of both beating modes have log-normal distributions. Without prey, A. tonsa only samples the environment at low frequency. Upon introduction of non-toxic food, it increases its sampling time moderately and the grazing period substantially. On mono algal diets for either of the toxic dinoflagellates, sampling time fraction is high but the grazing is very limited. A. tonsa demonstrates aversion to both toxic algal species. In mixtures of S. major and the neurotoxin producing K. brevis, sampling and grazing diminish rapidly, presumably due to neurological effects of consuming brevetoxins while trying to feed on S. major. In contrast, on mixtures of cytotoxin producing K. veneficum, both behavioral modes persist, indicating that intake of karlotoxins does not immediately inhibit the copepod's grazing behavior. These findings add critical insight into how these algal toxins may influence the copepod's feeding behavior, and suggest how some harmful algal species may alter top-down control exerted by grazers like copepods.     

Cycloheximide and heat shock induce new polypeptide synthesis in Neurospora crassa.

Treatment of Neurospora crassa with 0.1 microgram of cycloheximide per ml, a concentration which inhibited protein synthesis by about 70%, resulted in the greatly enhanced synthesis of at least three polypeptide bands with estimated molecular weights of 88,000, 30,000, and 28,000. A temperature shift from 25 to 37 degrees C resulted in the appearance of a single new polypeptide band of 70,000 daltons, the same size as the major heat shock-induced proteins observed in species of Drosophila and Dictyostelium. Synthesis of the cycloheximide-stimulated polypeptide bands was on cytoplasmic ribosomes rather than on mitochondrial ribosomes, as incorporation of isotope into the polypeptide bands was inhibited by 1.0 microgram of cycloheximide per ml but not by 1 mg of chloramphenicol per ml. In a mutant with cycloheximide-resistant ribosomes, 0.1 microgram of cycloheximide per ml failed to alter the pattern of protein synthesis from that of the controls. It is suggested that the new synthesis of the polypeptide bands reflects specific mechanisms of adaptation to different kinds of environmental stress, including inhibition of protein synthesis and temperature increases.     

Temperature compensation in the escape response of a marine copepod, Calanus finmarchicus (Crustacea)

Calanus finmarchicus, the dominant mesozooplankter of the North Atlantic, is an important food source for many fishes and other planktivores. This species, which has limited diel vertical migration, depends on its fast-start escape response to evade predators. It has myelinated neuronal axons, which contribute to its rapid and powerful escape response. The thermal environment that C. finmarchicus inhabits ranges from below 0 degrees C to 16 degrees C. Previous studies have shown that respiration, growth, and reproductive rates are strongly dependent on temperature, with Q10 > 2.5. A comparable dependence of the escape response could place the animal at higher risk for cold-compensated predators. Our work focused on the temperature dependence of the behavioral response to stimuli that mimic predatory attacks. We found that in contrast to other biological processes, all aspects of the escape response showed a low dependence on temperature, with Q10 values below 2. This low temperature dependence was consistent for escape parameters that involved neural as well as muscle components of the behavioral response. These findings are discussed in the contexts of the predator-prey relations of copepods and the thermal dependence of behavior in other taxa.     

The respiratory consequences of feeding in amphibians and reptiles

Many ectothermic vertebrates ingest very large meals at infrequent intervals. The digestive processes associated with these meals, often coupled with an extensive hypertrophy of the gastrointestinal organs, are energetically expensive and metabolic rate, therefore, increases substantially after feeding (specific dynamic action, SDA). Here, we review the cardio-respiratory consequences of SDA in amphibians and reptiles. For some snakes, the increased oxygen uptake during SDA is of similar magnitude to that of muscular exercise, and the two physiological states, therefore, exert similar and profound demands on oxygen transport by the cardiorespiratory systems. In several species, SDA is attended by increases in heart rate and overall systemic blood flows, but changes in blood flow distribution remain to be investigated. In snakes, the regulation of heart rate appears to involve a non-adrenergic-non-cholinergic mechanism, which may be a regulatory peptide released from the gastrointestinal system during digestion. Digestion is also associated with a net acid secretion to the stomach that causes an increase in plasma HCO3- concentration (the 'alkaline tide'). Experiments on chronically cannulated amphibians and reptiles, show that this metabolic alkalosis is countered by an increased P(CO2), so that the change in arterial pH is reduced. This respiratory compensation of arterial pH is accomplished through a reduction in ventilation relative to metabolism, but the estimated reductions in lung P(O2) are relatively small. The SDA response is also associated with haematological changes, but large interspecific differences exist. The studies on cardiorespiratory responses to digestion may allow for a further understanding of the physiological and structural constraints that limits the ability of reptiles and amphibians to sustain high metabolic rates.     

Effects of body mass, meal size, fast length, and temperature on specific dynamic action in the timber rattlesnake (Crotalus horridus)

Detailed analysis of animal energy budgets requires information on the cost of digestion (specific dynamic action [SDA]), which can represent a significant proportion of ingested energy (up to 30% in infrequent feeders). We studied the effects of snake mass, temperature (25 degrees and 30 degrees C), fasting time (1 and 5 mo), and prey size (10%-50% of snake mass) on SDA in 26 timber rattlesnakes (Crotalus horridus). We used flow-through respirometry to measure hourly CO(2) production rates (VCO2) for 1 d before and up to 17 d after feeding. Crotalus horridus, like previously studied viperids and boids, show large and ecologically relevant increases in metabolism due to feeding. Depending on treatment and individual, VCO2 increased to 2.8-11.8 times the resting metabolic rate within 12-45 h postfeeding and decreased to baseline within 4.3-15.4 d. Significant effects of snake mass, meal mass, and fast length were detected. Increased temperature decreased the time required to complete the process but had little effect on total energy expended on SDA. Energy expended on SDA increased with increasing fast length, snake mass, and prey mass. Considering all of our data, we found that a simple allometric relationship explained 96.7% of the variation in total CO(2) production during SDA. Calculations suggest that energy devoted to SDA may approach 20% of the total annual energy budget of snakes in nature. Discrepancies between our data and some previous studies draw attention to the fact that the measurement, expression, and analysis of SDA may be sensitive to several methodological and statistical assumptions.