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Renaud, Limoilou-Amelie; Blanchet, F. Guillaume; Cohen, Alan A.; Pelletier, Fanie 2019-03-22 1.Ecologists seek to understand the fitness consequences of variation in physiological markers, under the hypothesis that physiological state is linked to variability in individual condition and life history. 2.Thus, ecologists are often interested in estimating correlations between entire suites of correlated traits, or biomarkers, but sample size limitations often do not allow us to do this properly when large numbers of traits or biomarkers are considered. 3.Latent variables are a powerful tool to overcome this complexity. Recent statistical advances have enabled a new class of multivariate models – Multivariate Hierarchical Modeling (MHM) with latent variables − which allow to statistically estimate unstructured covariances/correlations among traits with reduced constraints on the number of degrees of freedom to account in the model. It is thus possible to highlight correlated structures in potentially very large numbers of traits. 4.Here, we apply MHM to evaluate the relative importance of individual differences and environmental effects on milk composition and identify the drivers of this variation. We ask whether variation in bighorn sheep milk affects offspring fitness. 5.We evaluate whether mothers show repeatable individual differences in the concentrations of 11 markers of milk composition and we investigate the relative importance of annual variability, maternal identity and morphological traits in structuring milk composition. We then use variance estimates to investigate how a subset of repeatable milk markers influence lamb summer survival. 6.Repeatability of milk markers ranged from 0.05 to 0.64 after accounting for year‐to‐year variations. Milk composition was weakly but significantly associated with maternal mass in June and September, summer mass gain and winter mass loss. Variation explained by year‐to year fluctuations ranged from 0.07 to 0.91 suggesting a strong influence of environmental variability on milk composition. Milk composition did not affect lamb survival to weaning. 7.Using joint models in ecological, physiological or behavioural contexts has the major advantage of decomposing a (co)variance/correlation matrix while being estimated with fewer parameters than in a ‘traditional’ mixed‐effects model. The joint models presented here complement a growing list of tools to analyse correlations at different hierarchical levels separately and may thus represent a partial solution to the conundrum of physiological complexity.
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Fowler, Melinda A.; Paquet, Mélissa; Legault, Véronique; Cohen, Alan A.; Williams, Tony D. 2018-11-26 Background: It is widely assumed that variation in fitness components has a physiological basis that might underlie selection on trade-offs, but the mechanisms driving decreased survival and future fecundity remain elusive. Here, we assessed whether physiological variables are related to workload ability or immediate fitness consequences and if they mediate future survival or reproductive success. We used data on 13 physiological variables measured in 93 female European starlings (Sturnus vulgaris) at two breeding stages (incubation, chick-rearing), for first-and second-broods over two years (152 observations). Results: There was little co-variation among the physiological variables, either in incubating or chick-rearing birds, but some systematic physiological differences between the two stages. Chick-rearing birds had lower hematocrit and plasma creatine kinase but higher hemoglobin, triglyceride and uric acid levels. Only plasma corticosterone was repeatable between incubation and chick-rearing. We assessed relationships between incubation or chick-rearing physiology and measures of workload, current productivity, future fecundity or survival in a univariate manner, and found very few significant relationships. Thus, we next explored the utility of multivariate analysis (principal components analysis, Mahalanobis distance) to account for potentially complex physiological integration, but still found no clear associations. Conclusions: This implies either that a) birds maintained physiological variables within a homeostatic range that did not affect their performance, b) there are relatively few links between physiology and performance, or, more likely, c) that the complexity of these relationships exceeds our ability to measure it. Variability in ecological context may complicate the relationship between physiology and behavior. We thus urge caution regarding the over-interpretation of isolated significant findings, based on single traits in single years, in the literature.
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Milot, Emmanuel; Cohen, Alan A.; Vézina, François; Buehler, Deborah M.; Matson, Kevin D.; Piersman, Theunis; Piersma, Theunis 2014-12-06 1. The body condition of free-ranging animals affects their response to stress, decisions, ability to fulfil vital needs and, ultimately, fitness. However, this key attribute in ecology remains difficult to assess, and there is a clear need for more integrative measures than the common univariate proxies. 2. We propose a systems biology approach that positions individuals along a gradient from a ‘normal/optimal’ to ‘abnormal/suboptimal’ physiological state based on Mahalanobis distance computed from physiological biomarkers. We previously demonstrated the validity of this approach for studying ageing in humans; here, we illustrate its broad potential for ecological studies. 3. As an example, we used biomarker data on shorebirds and found that birds with an abnormal condition had a lower maximal thermogenic capacity and higher scores of inflammation, with important implications for their ecology and health. Moreover, Mahalanobis distance captured a signal of condition not detected by the individual biomarkers. 4. Overall, our results on birds and humans show that individuals with abnormal physiologies are indeed in worse condition. Moreover, our approach appears not to be particularly sensitive to which set of biomarkers is used to assess condition. Consequently, it could be applied easily to existing ecological data sets. 5. Our approach provides a general, powerful way to measure condition that helps resolve confusion as to how to deal with complex interactions and interdependence among multiple physiological and condition measures. It can be applied directly to topics such as the effect of environmental quality on body condition, risks of health outcomes, mechanisms of adaptive phenotypic plasticity, and mechanisms behind long-term processes such as senescence. https://creativecommons.org/publicdomain/zero/1.0/

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