Thermal Effects on Ecological Traits of Salmonids

Here, I review thermal influences on metabolic rates and aerobic scope; growth; adult body size; and reproductive and behavioural traits, such as tendency and timing of the migration of salmonid fishes. A thermal window bounded by the upper and lower incipient lethal temperatures (UILT and LILT) determines where salmonids can survive. For most salmonids, LILT is close to 0 and UILT is between 20 and 30ᵒ C. UILT and LILT are influenced by the acclimation temperature. Thermal tolerance is affected by fish size and ambient oxygen content, which decreases with increasing temperature. Standard metabolic rate (SMR), the energy required to maintain essential functions, increases with temperature, whereas maximum metabolic rate (MMR) increases with temperature until reaching a peak (pejus). Then, it decreases gradually to zero, i.e., the upper critical limit (TCrit). Aerobic scope (AS = MMR-SMR) reaches its maximum at the pejus temperature. Metabolic rates and aerobic scope can be modified by temperatures that the fish experiences during embryogenesis and possibly also as larvae and young fry. At maximum feeding, maximum growth increases to a point at or below the pejus temperature. The optimum temperature for growth decreases with reduced food intake and increased body size. As for metabolic rate, the growth rate is influenced by the temperature during embryonic development. In a warmer climate, adult body size is expected to decrease chiefly because of a younger age at maturity. Parental fish retained at a higher temperature during maturation produce larger eggs, and this change in egg size may also be transferred to next-generation offspring. Furthermore, embryogenesis in warmer water leads to larger gonad and egg sizes at maturity. Water temperature influences locomotion, foraging and migratory activity. In a warmer climate, juveniles migrate to the sea earlier in spring. In addition, higher embryo temperature leads to delayed return of adult salmon from the ocean. Thus, temperature affects life history traits of salmonid fishes, partly as a direct effect on metabolic rates and food consumption and partly induced as a phenotypically plastic effect. The phenotypically plastic response may preadapt offspring to perform better in the expected future thermal environment. adaptive developmental programming; behaviour; climate; life history traits; metabolism; Salmonidae; temperaturepublishedVersio

Thermal conditions during embryogenesis influence metabolic rates of juvenile brown trout Salmo trutta

The projected climate change and increase in thermal conditions in northern latitudes over the next 60 yr has the potential to alter the metabolic scope and potential fitness of aquatic ectotherms. Here, we experimentally tested if elevated egg incubation temperature affected metabolic scope in juvenile brown trout (Salmo trutta) as a phenotypically plastic response. Cohorts of brown trout from anadromous and resident crosses were raised through embryogenesis in either natural river temperatures (cold) or elevated (+3°C, warm) temperatures until they could feed exogenously. The standard metabolic rate (SMR), maximum metabolic rate (MMR), and aerobic scope (AS = MMR − SMR) of juveniles from four anadromous- resident crosses and from both incubation temperatures were tested at 13°C. We found that metabolic measures (SMR, MMR, AS) were lower in warm than cold-incubated fish. There was no difference in the metabolic rates of fish from different anadromous-resident crosses. The results of this experiment are consistent with the countergradient variation hypothesis (CGV) in which phenotypic variation, in this case variation in metabolic rates, is inversely related to thermal conditions, originally proposed in relation to altitudinal or latitudinal gradients. While previous studies have related CGV to genetic differences between populations, our study shows that thermal differences encountered at the embryonic stage can produce a phenotypic pattern consistent with CGV. It is difficult to predict the consequences of these metabolic changes in a future warmer climate, as lower metabolic rates indicate that brown trout will probably expend less energy, but a reduced aerobic scope may counteract this affect, limiting their ability as a top predator and in escaping predators. Our results suggest that there are mechanisms used to adjust to elevated water temperature that can be initiated during embryogenesis. Given that there were no differences among crosses, it is likely that temperature-induced differences are the result of plastic responses. aerobic scope; brown trout; climate change; embryogenesis; incubation temperature; maximal metabolic rate; standard metabolic rate.publishedVersio

Evaluation of the Southern Ocean O2/Ar-based NCP estimates in a model framework

Author Posting. © American Geophysical Union, 2013. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Journal of Geophysical Research: Biogeosciences 118 (2013): 385–399, doi:10.1002/jgrg.20032.The sea-air biological O2 flux assessed from measurements of surface O2 supersaturation in excess of Ar supersaturation (“O2 bioflux”) is increasingly being used to constrain net community production (NCP) in the upper ocean mixed layer. In making these calculations, one generally assumes that NCP is at steady state, mixed layer depth is constant, and there is no O2 exchange across the base of the mixed layer. The object of this paper is to evaluate the magnitude of errors introduced by violations of these assumptions. Therefore, we examine the differences between the sea-air biological O2 flux and NCP in the Southern Ocean mixed layer as calculated using two ocean biogeochemistry general circulation models. In this approach, NCP is considered a known entity in the prognostic model, whereas O2 bioflux is estimated using the model-predicted O2/Ar ratio to compute the mixed layer biological O2 saturation and the gas transfer velocity to calculate flux. We find that the simulated biological O2 flux gives an accurate picture of the regional-scale patterns and trends in model NCP. However, on local scales, violations of the assumptions behind the O2/Ar method lead to significant, non-uniform differences between model NCP and biological O2 flux. These errors arise from two main sources. First, venting of biological O2 to the atmosphere can be misaligned from NCP in both time and space. Second, vertical fluxes of oxygen across the base of the mixed layer complicate the relationship between NCP and the biological O2 flux. Our calculations show that low values of O2 bioflux correctly register that NCP is also low (<10 mmol m−2 day−1), but fractional errors are large when rates are this low. Values between 10 and 40 mmol m−2 day−1 in areas with intermediate mixed layer depths of 30 to 50 m have the smallest absolute and relative errors. Areas with O2 bioflux higher than 30 mmol m−2 day−1 and mixed layers deeper than 40 m tend to underestimate NCP by up to 20 mmol m−2 day−1. Excluding time periods when mixed layer biological O2 is undersaturated, O2 bioflux underestimates time-averaged NCP by 5%–15%. If these time periods are included, O2 bioflux underestimates mixed layer NCP by 20%–35% in the Southern Ocean. The higher error estimate is relevant if one wants to estimate seasonal NCP since a significant amount of biological production takes place when mixed layer biological O2 is undersaturated.This work was supported in part by funding from the National Aeronautic and Space Administration (NASA NNX08AF12G) and National Science Foundation (NSF OPP-0823101)

Evaluating Southern Ocean biological production in two ocean biogeochemical models on daily to seasonal timescales using satellite chlorophyll and O2 / Ar observations

© The Author(s), 2015. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Biogeosciences 12 (2015): 681-695, doi:10.5194/bg-12-681-2015.We assess the ability of ocean biogeochemical models to represent seasonal structures in biomass and net community production (NCP) in the Southern Ocean. Two models are compared to observations on daily to seasonal timescales in four different sections of the region. We use daily satellite fields of chlorophyll (Chl) as a proxy for biomass and in situ observations of O2 and Ar supersaturation (ΔO2 / Ar) to estimate NCP. ΔO2 / Ar is converted to the flux of biologically generated O2 from sea to air (O2 bioflux). All data are aggregated to a climatological year with a daily resolution. To account for potential regional differences within the Southern Ocean, we conduct separate analyses of sections south of South Africa, around the Drake Passage, south of Australia, and south of New Zealand. We find that the models simulate the upper range of Chl concentrations well, underestimate spring levels significantly, and show differences in skill between early and late parts of the growing season. While there is a great deal of scatter in the bioflux observations in general, the four sectors each have distinct patterns that the models pick up. Neither model exhibits a significant distinction between the Australian and New Zealand sectors and between the Drake Passage and African sectors. South of 60° S, the models fail to predict the observed extent of biological O2 undersaturation. We suggest that this shortcoming may be due either to problems with the ecosystem dynamics or problems with the vertical transport of oxygen.This work was supported in part by funding from the National Aeronautic and Space Administration (NASA NNX08AF12G) and the National Science Foundation (NSF OPP-0823101)

Ocean Color Algorithm for the Retrieval of the Particle Size Distribution and Carbon-Based Phytoplankton Size Classes Using a Two-Component Coated-Spheres Backscattering Model

The particle size distribution (PSD) of suspended particles in near-surface seawater is a key property linking biogeochemical and ecosystem characteristics with optical properties that affect ocean color remote sensing. Phytoplankton size affects their physiological characteristics and ecosystem and biogeochemical roles, e.g. in the biological carbon pump, which has an important role in the global carbon cycle and thus climate. It is thus important to develop capabilities for measurement and predictive understanding of the structure and function of oceanic ecosystems, including the PSD, phytoplankton size classes (PSCs) and phytoplankton functional types (PFTs). Here, we present an ocean color satellite algorithm for the retrieval of the parameters of an assumed power-law PSD. The forward optical model considers two distinct particle populations (particle assemblage categories) &mdash; phytoplankton and non-algal particles (NAP). Phytoplankton are modeled as coated spheres following the Equivalent Algal Populations (EAP) framework, and NAP are modeled as homogeneous spheres. The forward model uses Mie and Aden-Kerker scattering computations, for homogeneous and coated spheres (for phytoplankton and NAP, respectively) to model the total particulate spectral backscattering coefficient as the sum of phytoplankton and NAP backscattering. The PSD retrieval is achieved via Spectral Angle Mapping (SAM) which uses backscattering end-members created by the forward model. The PSD is used to retrieve size-partitioned absolute and fractional phytoplankton carbon concentrations (i.e. carbon-based PSCs), as well as particulate organic carbon (POC), using allometric coefficients. The EAP-based formulation allows for the estimation of chlorophyll-a concentration via the retrieved PSD, as well as the estimation of the percent of backscattering due to NAP vs. phytoplankton. The PSD algorithm is operationally applied to the merged Ocean Colour Climate Change Initiative (OC-CCI) v5.0 ocean color data set. Results of an initial validation effort are also presented, using PSD, POC, and pico-phytoplankton carbon in-situ measurements. Validation results indicate the need for an empirical tuning for the absolute phytoplankton carbon concentrations; however these results and comparison with other phytoplankton carbon algorithms are ambiguous as to the need for the tuning. The latter finding illustrates the continued need for high-quality, consistent, large global data sets of phytoplankton carbon and related variables to facilitate future algorithm improvements.</p

Ocean mover’s distance: using optimal transport for analysing oceanographic data

Remote sensing observations from satellites and global biogeochemical models have combined to revolutionize the study of ocean biogeochemical cycling, but comparing the two data streams to each other and across time remains challenging due to the strong spatial-temporal structuring of the ocean. Here, we show that the Wasserstein distance provides a powerful metric for harnessing these structured datasets for better marine ecosystem and climate predictions. The Wasserstein distance complements commonly used point-wise difference methods such as the root-mean-squared error, by quantifying differences in terms of spatial displacement in addition to magnitude. As a test case, we consider chlorophyll (a key indicator of phytoplankton biomass) in the northeast Pacific Ocean, obtained from model simulations, in situ measurements, and satellite observations. We focus on two main applications: (i) comparing model predictions with satellite observations, and (ii) temporal evolution of chlorophyll both seasonally and over longer time frames. The Wasserstein distance successfully isolates temporal and depth variability and quantifies shifts in biogeochemical province boundaries. It also exposes relevant temporal trends in satellite chlorophyll consistent with climate change predictions. Our study shows that optimal transport vectors underlying the Wasserstein distance provide a novel visualization tool for testing models and better understanding temporal dynamics in the ocean

Ocean mover’s distance: using optimal transport for analysing oceanographic data

Remote sensing observations from satellites and global biogeochemical models have combined to revolutionize the study of ocean biogeochemical cycling, but comparing the two data streams to each other and across time remains challenging due to the strong spatial-temporal structuring of the ocean. Here, we show that the Wasserstein distance provides a powerful metric for harnessing these structured datasets for better marine ecosystem and climate predictions. The Wasserstein distance complements commonly used point-wise difference methods such as the root-mean-squared error, by quantifying differences in terms of spatial displacement in addition to magnitude. As a test case, we consider chlorophyll (a key indicator of phytoplankton biomass) in the northeast Pacific Ocean, obtained from model simulations, in situ measurements, and satellite observations. We focus on two main applications: (i) comparing model predictions with satellite observations, and (ii) temporal evolution of chlorophyll both seasonally and over longer time frames. The Wasserstein distance successfully isolates temporal and depth variability and quantifies shifts in biogeochemical province boundaries. It also exposes relevant temporal trends in satellite chlorophyll consistent with climate change predictions. Our study shows that optimal transport vectors underlying the Wasserstein distance provide a novel visualization tool for testing models and better understanding temporal dynamics in the ocean

Fecundity and water flow influence the dynamics of Atlantic salmon

Populations are retained at reduced levels by resource competition and environmental stochasticity. In the Norwegian River Imsa, the relationship between fecundity of Atlantic salmon (Salmo salar) spawners and number of smolts per unit river area was investigated for cohorts spawned from 1976 to 2011. Annual number of smolts produced per unit area was best described by a multiplicative model and increased with the fecundity of the females as proxy for number of eggs deposited and the minimum water flow in August towards the end of the first growth season. Mean monthly water temperature, or water flow in any other month during the first year, had no significant effect on number of smolts produced. At sea, there was an almost linear relationship between number of emigrating smolts and returning adults, possibly because population abundance of Atlantic salmon is low relative to the carrying capacity in the ocean. Thus, both number of eggs spawned and minimum water flow in late summer influenced population abundance in the present river. density dependence, River Imsa, Salmo salar, sea survival, stock–recruitmentacceptedVersio

Differences in growth between offspring of anadromous and freshwater brown trout Salmo trutta

In this study, individual growth of juvenile offspring of anadromous and freshwater resident brown trout Salmo trutta and crosses between the two from the River Imsa, Norway, was estimated. The juveniles were incubated until hatching at two temperatures (±S.D.), either 4.4 ± 1.5 C or 7.1 ± 0.6 C. Growth rate was estimated for 22 days in August–September when the fish on average were c. 8 g in wet mass, and the estimates were standardized to 1 g fish dry mass. Offspring of anadromous S. trutta grew better at both 15 and 18 C than offspring of freshwater resident S. trutta or offspring of crosses between the two S. trutta types. This difference appears not to result from a maternal effect because anadromous S. trutta grew better than the hybrids with anadromous mothers. Instead, this appears to be an inherited difference between the anadromous and the freshwater resident fish lending support to the hypothesis that anadromous and freshwater resident S. trutta in this river differ in genetic expression. Egg incubation temperature of S. trutta appeared not to influence the later growth as reported earlier from the studies of Atlantic salmon Salmo salar.publishedVersio

Maternal inheritance influences homing and growth of hybrid offspring between wild and farmed Atlantic salmon

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