Field-collected otolith samples of young-of the-year Arctic charr (Salvelinus alpinus) and brook charr (Salvelinus fontinalis) and monitored water temperatures were used to estimate a delta(18)O fractionation equation for the genus Salvelinus. When compared to literature reported equations, the developed fractionation equation had a statistically similar slope but dissimilar intercept. Statistical similarities among fractionation equation slope estimates suggest a common otolith delta(18)O incorporation response among fish species that may be interpreted as widespread equilibrium otolith delta(18)O deposition. Statistical dissimilarities among intercept estimates question broad applicability of any single fractionation equation to all fish species and were interpreted here to have biological meaning as a result of known differences in standard metabolic rates among species. Attempts to statistically cross-validate fractionation equations by prediction of water temperatures used in other fractionation studies indicated significant biases in the range of -7.9 to 6.7 degrees C that preclude the broad use of any single fractionation equation for accurate thermal reconstructions. Differences in equation intercepts and the prevalence of predictive biases do not support the conclusion of previous studies concerning the wide applicability and/or general accuracy of fractionation equations and suggest fractionation equations are best developed at the species- or taxon-specific (e.g., genus) level.
A species-specific fractionation equation for Arctic charr (Salvelinus alpinus (L.)) was developed experimentally for use in ecological studies of temperature-driven phenologies for the species. Juvenile Arctic charr were reared in controlled conditions at different temperatures (2-14°C), with three replicates of each temperature. Otoliths from the fish and water samples from the chambers were analysed for oxygen isotope composition and used to estimate temperature-dependent fractionation equations relating the isotopic ratio to rearing temperature. A linear and a second order polynomial relationship were estimated and validated using comparable Arctic charr data from another study. Temperatures predicted using the polynomial equation were not significantly different from recorded experimental temperatures, whereas with the linear equation there were significant differences between the predicted and measured temperatures. The polynomial equation also showed the least bias as measured by mean predictive error. Statistical comparisons of the polynomial fractionation equation to a similarly estimated equation for brook charr (Salvelinus fontinalis (Mitchill)) indicated significant differences. Results imply the need for species-specific fractionation equations, even for closely related fish. Results further suggest the polynomial form of the fractionation equation will facilitate more accurate characterisation of water temperatures suitable for use in ecological studies of temperature-driven phenologies of Arctic charr.
The effects of early environmental conditions can profoundly affect individual development and adult phenotype. In birds, limiting resources can affect growth as nestlings, but also fitness and survival as adults. Following periods of food restriction, individuals may accelerate development, undergoing a period of rapid "catch-up" growth, in an attempt to reach the appropriate size at adulthood. Previous studies of altricial birds have shown that catch-up growth can have negative consequences in adulthood, although this has not been explored in species with different developmental strategies. Here, we investigated the effects of resource limitation and the subsequent period of catch-up growth, on the morphological and metabolic phenotype of adult Japanese quail (Coturnix japonica), a species with a precocial developmental strategy. Because males and females differ in adult body size, we also test whether food restriction had sex-specific effects. Birds that underwent food restriction early in development had muscles of similar size and functional maturity, but lower adult body mass than controls. There was no evidence of sex-specific sensitivity of food restriction on adult body mass; however, there was evidence for body size. Females fed ad lib were larger than males fed ad lib, while females subjected to food restriction were of similar size to males. Adults that had previously experienced food restriction did not have an elevated metabolic rate, suggesting that in contrast to altricial nestlings, there was no metabolic carry-over effect of catch-up growth into adulthood. While Japanese quail can undergo accelerated growth after re-feeding, timing of food restriction may be important to adult size, particularly in females. However, greater developmental flexibility compared to altricial birds may contribute to the lack of metabolic carryover effects at adulthood.
Stable hydrogen isotopes (δ(2)H) are commonly used in studies of animal movement. Tissue that is metabolically inactive after growth (e.g., feathers) provides spatial or dietary information that reflects only the period of tissue growth, whereas tissues that are metabolically active (e.g., red blood cells) provide a moving window of forensic information. However, using δ(2)H for studies of animal movement relies on the assumption that tissue δ(2)H values reflect dietary δ(2)H values, plus or minus a net diet-tissue discrimination value, and that the turnover rate is known for metabolically active tissue. The metabolic rate of an animal may influence both diet-tissue discrimination values and isotopic tissue turnover rate, but this hypothesis has not been tested experimentally. To examine the metabolic hypothesis, an experimental group of 12 male and 15 female captive Japanese quail (Coturnix japonica) was housed at 8.9°C for 90 d to elevate their metabolic rates (mL CO(2) min(-1)), and a control group of 12 male and 13 female quail was housed at room temperature during the same period. For both experimental and control birds, diet-tissue discrimination values were estimated for red blood cells and feathers. To determine turnover rate, experimental and control birds were switched from a (2)H-enriched diet to a (2)H-depleted diet, with red blood cells sampled before and after diet switch. Metabolic rate did not influence red blood cell hydrogen isotope turnover rate (η(2)(p) = 0.24)) or diet-feather isotope discrimination values (η(2)(p) = 0.86). Diet-feather hydrogen isotopic discrimination had a significant sex plus treatment interaction effect; female feathers were depleted in (2)H relative to food regardless of treatment, whereas male feathers were enriched in (2)H. The effect of sex suggested that experimental studies should examine whether coeval males and females differ in blood δ(2)H levels during certain periods of the annual cycle.
Summary1. The stable-hydrogen and oxygen isotopic composition of animal tissues are important tools for estimating the geographic origin of migrating animals. 2. Estimating the origin of animals using H and O isotopes requires knowledge of how closely and consistently tissues reflect precipitation isotopes at the location in which the tissues were grown. 3. One hypothesis to explain variation in isotope discrimination values in tissues from individuals residing at the same location is metabolic rate. Variation in metabolic rate between individuals could alter the relative proportions of drinking water and food atoms that are integrated into a tissue. Individuals with high metabolic rates ingest greater amounts of water compared with those with lower metabolic rates in order to balance metabolic water losses. Increased water consumption may increase the rate of hydrogen and oxygen atom turnover within the blood plasma, thereby limiting the extent to which tissue precursors isotopically exchange with their surroundings. 4. A first test of this hypothesis was carried out by experimentally elevating the metabolic rate of Japanese quail (Coturnix japonica) over a 64-day period. We spiked drinking water with 99·9% 2 H 2 O and 10% 18 O to trace the incorporation of drinking water to feather tissue while holding the isotopic value of the food constant. 5. We observed a 15% increase in metabolic rate was associated with a 6% decrease in the relative contribution of drinking water hydrogen to feather keratin between treatments. In contrast, no differences in the relative contribution of drinking water oxygen to plasma or feathers were detected because of the small isotopic separation (e.g., only 9&) between initial and final drinking water d 18 O values. Results suggest a difference in metabolic rates between individuals residing in the same geographic location have the potential to partially obscure the relationship between precipitation and feather d 2 H values.
The use of preserved otoliths for stable isotope analysis assumes handling and preservation procedures do not alter the isotopic composition of the otolith. Otoliths from wild and hatcheryreared salmonids (brook charr, Salvelinus fontinalis, and Atlantic salmon, Salmo salar) were used to test for possible d 18O preservation effects in ethanol and formalin preservation experiments at varying temperatures. Analysis of variance (ANOVA) demonstrated a significant interaction effect between species and preservative during preservation. Possible causes for the observed effect are discussed in relation to species-specific differences in otolith chemistry related to growth and environment including: (1) chemical mechanisms of dissolution-recrystallisation involving the precipitation of secondary minerals within and at the otolith surface; (2) adsorption of ions at available binding sites on the otolith surface; and (3) isotopic exchange during otolith surface dissolution and/or reprecipitation processes. Differential occurrence of vaterite and aragonite in otoliths is believed to account for some of the observed effects as a result of otolith density differences. Isotopic exchange is also argued to cause much of the observed variation in speciesspecific preservation effects. Biologically, study findings imply that preserved otoliths should not be used for baseline paleoclimatic or individual fish thermal reconstructions, or the development of d 18 O-fractionation equations, without the prior use of pilot studies to determine preservation effects.
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