Understanding how trait diversification alters ecosystem processes is an important goal for ecological and evolutionary studies. Ecological stoichiometry provides a framework for predicting how traits affect ecosystem function. The growth rate hypothesis of ecological stoichiometry links growth and phosphorus (P) body composition in taxa where nucleic acids are a significant pool of body P. In vertebrates, however, most of the P is bound within bone, and organisms with boney structures can vary in terms of the relative contributions of bones to body composition. Threespine stickleback populations have substantial variation in boney armour plating. Shaped by natural selection, this variation provides a model system to study the links between evolution of bone content, elemental body composition, and P excretion. We measure carbon:nitrogen:P body composition from stickleback populations that vary in armour phenotype. We develop a mechanistic mass-balance model to explore factors affecting P excretion, and measure P excretion from two populations with contrasting armour phenotypes. Completely armoured morphs have higher body %P but excrete more P per unit body mass than other morphs. The model suggests that such differences are driven by phenotypic differences in P intake as well as body %P composition. Our results show that while investment in boney traits alters the elemental composition of vertebrate bodies, excretion rates depend on how acquisition and assimilation traits covary with boney trait investment. These results also provide a stoichiometric hypothesis to explain the repeated loss of boney armour in threespine sticklebacks upon colonizing freshwater ecosystems.
As they return to spawn and die in their natal streams, anadromous, semelparous fishes such as Pacific salmon import marine‐derived nutrients to otherwise nutrient‐poor freshwater and riparian ecosystems. Diverse organisms exploit this resource, and previous studies have indicated that riparian tree growth may be enhanced by such marine‐derived nutrients. However, these studies were largely inferential and did not account for all factors affecting tree growth. As an experimental test of the contribution of carcasses to tree growth, for 20 yr, we systematically deposited all sockeye salmon (Oncorhynchus nerka) carcasses (217,055 individual salmon) in the riparian zone on one bank of a 2‐km‐long stream in southwestern Alaska, reducing carcass accumulation on one bank and enhancing it on the other. After accounting for partial consumption and movement of carcasses by brown bears (Ursus arctos) and variation in salmon abundance and body size, we estimated that 267,620 kg of salmon were deposited on the enhanced bank and 45,200 kg on the depleted bank over the 20 yr, for a 5.9‐fold difference in total mass. In 2016, we sampled needles of 84 white spruce trees (Picea glauca) the dominant riparian tree species, for foliar nitrogen (N) content and stable isotope ratios (δ15N), and took core samples for annual growth increments. Stable isotope analysis indicated that marine‐derived N was incorporated into the new growth of the trees on the enhanced bank. Analysis of tree cores indicated that in the two decades prior to our enhancement experiment, trees on the south‐facing (subsequently the depleted) bank grew faster than those on the north‐facing (later enhanced) bank. This difference was reduced significantly during the two decades of fertilization, indicating an effect of the carcass transfer experiment against the background of other factors affecting tree growth.
High-latitude lakes are particularly sensitive to the effects of global climate change, demonstrating earlier ice breakup, longer ice-free seasons, and increased water temperatures. Such physical changes have implications for diverse life-history traits in taxa across entire lake food webs. Here, we use a five-decade time series from an Alaskan lake to explore effects of climate change on growth and reproduction of a widely distributed lacustrine fish, the three-spine stickleback (Gasterosteus aculeatus). We used multivariate autoregressive state-space (MARSS) models to describe trends in the mean length for multiple size classes and to explore the influence of physical (date of ice breakup, surface water temperature) and biological (density of con- and heterospecifics) factors. As predicted, mean size of age 1 and older fish at the end of the growing season increased across years with earlier ice breakup and warmer temperatures. In contrast, mean size of age 0 fish decreased over time. Overall, lower fish density and warmer water temperatures were associated with larger size for all cohorts. Earlier ice breakup was associated with larger size for age 1 and older fish but, paradoxically, with smaller size of age 0 fish. To explore this latter result, we used mixing models on age 0 size distributions, which revealed an additional cohort in years with early ice breakup, lowering the mean size of age 0 fish. Moreover, early ice breakup was associated with earlier breeding, evidenced by earlier capture of age 0 fish. Our results suggest that early ice breakup altered both timing and frequency of breeding; three-spine stickleback spawned earlier and more often in response to earlier ice breakup date. While previous studies have shown the influence of changing conditions in northern lakes on breeding timing and growth, this is the first to document increased breeding frequency, highlighting another pathway by which climate change can alter the ecology of northern lakes.
The Threespine Stickleback Gasterosteus aculeatus is widely distributed across northern hemisphere ecosystems, has ecological influence as an abundant planktivore, and is commonly used as a model organism, but the species lacks a comprehensive model to describe bioenergetic performance in response to varying environmental or ecological conditions. This study parameterized a bioenergetics model for the Threespine Stickleback using laboratory measurements to determine mass-and temperature-dependent functions for maximum consumption and routine respiration costs. Maximum consumption experiments were conducted across a range of temperatures from 7.5 C to 23.0 C and a range of fish weights from 0.5 to 4.5 g. Respiration experiments were conducted across a range of temperatures from 8 C to 28 C. Model sensitivity was consistent with other comparable models in that the mass-dependent parameters for maximum consumption were the most sensitive. Growth estimates based on the Threespine Stickleback bioenergetics model suggested that 22 C is the optimal temperature for growth when food is not limiting. The bioenergetics model performed well when used to predict independent, paired measures of consumption and growth observed from a separate wild population of Threespine Sticklebacks. Predicted values for consumption and growth (expressed as percent body weight per day) only deviated from observed values by 2.0%. Our model should provide insight into the physiological performance of this species across a range of environmental conditions and be useful for quantifying the trophic impact of this species in food webs containing other ecologically or economically important species.
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