Find research datasets worth reusing

Data from: Age-specific patterns of maternal investment in common gull egg yolk

While the general patterns of age-specific changes in reproductive success are quite well established in long-lived animals, we still do not know if allocation patterns of maternally-transmitted compounds are related to maternal age. We measured yolk testosterone, carotenoids and vitamins A and E levels in a population of known-aged common gulls (Larus canus) and found an age-specific pattern in yolk lutein and vitamin A concentrations. Middle-aged mothers allocated more of these substances to yolk compared to young and old mothers. These results can be explained through differences in age-specific foraging, absorption, or deposition patterns of carotenoids and vitamins into yolk. If these molecules play a role in antioxidant defense and immune modulation, our results suggest a possible physiological pathway behind the age-specific changes in reproductive success of long-lived birds in the wild.

opencc-zeroDec 2011View details →

Data from: Strong specificity in the interaction between parasitoids and symbiont-protected hosts

Coevolution between hosts and parasites may promote the maintenance of genetic variation in both antagonists by negative-frequency-dependence if the host-parasite interaction is genotype-specific. Here we tested for specificity in the interaction between parasitoids (Lysiphlebus fabarum) and aphid hosts (Aphis fabae) that are protected by a heritable defensive endosymbiont, the γ-proteobacterium Hamiltonella defensa. Previous studies reported a lack of genotype-specificity between unprotected aphids and parasitoids, but suggested that symbiont-conferred resistance might exhibit a higher degree of specificity. Indeed, in addition to ample variation in host resistance as well as parasitoid infectivity, we found a strong aphid clone-by-parasitoid line interaction on the rates of successful parasitism. This genotype-specificity appears to be mediated by H. defensa, highlighting the important role that endosymbionts can play in host-parasite coevolution.

opencc-zeroDec 2016View details →

Data from: Serum copeptin and neuron specific enolase are markers of neonatal distress and long-term neurodevelopmental outcome

The objective of this study was to evaluate the early changes in serial serum levels of copeptin and neuron-specific enolase (NSE) in neonates diagnosed with birth asphyxia, and to determine whether these biomarkers measured in the first 168 hours after birth are predictive of long-term neurodevelopmental outcome. Copeptin and NSE levels were measured from serum samples collected 6, 12, 24, 48, 72, and 168 hours after birth from 75 term neonates diagnosed with hypoxic-ischemic encephalopathy (HIE) and treated with therapeutic hypothermia for 72 hours. In addition, serum copeptin levels after birth were measured from 10 HIE diagnosed neonates, who were randomized to the normothermic arm of the TOBY cohort. All neonates underwent neurodevelopmental assessment using the Bayley Scales of Infant and Toddler Development-II at two years of age. Copeptin levels were highest at 6 hours after birth and steadily decreased, whereas the highest NSE levels were measured at 24 hours after birth. The biomarker levels correlated with blood-gas parameters (base excess, pH and lactate) at 6 and 12 hours after birth. Copeptin and NSE levels in the early postnatal period were significantly higher in neonates with poor outcome compared to those with favorable outcome at two years of age. Furthermore, in the TOBY cohort, copeptin levels were significantly lower in hypothermic compared to normothermic neonates. To conclude, copeptin and NSE measured in the early postnatal period are potential prognostic biomarkers of long-term neurodevelopmental outcome in term neonates diagnosed with HIE and treated with therapeutic hypothermia.

Data from: High specificity in circulating tumor cell identification is required for accurate evaluation of programmed death-ligand 1

Background: Expression of programmed-death ligand 1 (PD-L1) in non-small cell lung cancer (NSCLC) is typically evaluated through invasive biopsies; however, recent advances in the identification of circulating tumor cells (CTCs) may be a less invasive method to assay tumor cells for these purposes. These liquid biopsies rely on accurate identification of CTCs from the diverse populations in the blood, where some tumor cells share characteristics with normal blood cells. While many blood cells can be excluded by their high expression of CD45, neutrophils and other immature myeloid subsets have low to absent expression of CD45 and also express PD-L1. Furthermore, cytokeratin is typically used to identify CTCs, but neutrophils may stain non-specifically for intracellular antibodies, including cytokeratin, thus preventing accurate evaluation of PD-L1 expression on tumor cells. This holds even greater significance when evaluating PD-L1 in epithelial cell adhesion molecule (EpCAM) positive and EpCAM negative CTCs (as in epithelial-mesenchymal transition (EMT)). Methods: To evaluate the impact of CTC misidentification on PD-L1 evaluation, we utilized CD11b to identify myeloid cells. CTCs were isolated from patients with metastatic NSCLC using EpCAM, MUC1 or Vimentin capture antibodies and exclusion-based sample preparation (ESP) technology. Results: Large populations of CD11b+CD45lo cells were identified in buffy coats and stained non-specifically for intracellular antibodies including cytokeratin. The amount of CD11b+ cells misidentified as CTCs varied among patients; accounting for 33–100% of traditionally identified CTCs. Cells captured with vimentin had a higher frequency of CD11b+ cells at 41%, compared to 20% and 18% with MUC1 or EpCAM, respectively. Cells misidentified as CTCs ultimately skewed PD-L1 expression to varying degrees across patient samples. Conclusions: Interfering myeloid populations can be differentiated from true CTCs with additional staining criteria, thus improving the specificity of CTC identification and the accuracy of biomarker evaluation.

opencc-zeroDec 2014View details →

Data from: Evidence for specificity in symbiont-conferred protection against parasitoids

Many insects harbour facultative symbiotic bacteria, some of which have been shown to provide resistance against natural enemies. One of the best-known protective symbionts is Hamiltonella defensa, which in pea aphid (Acyrthosiphon pisum) confers resistance against attack by parasitoid wasps in the genus Aphidius (Braconidae). We asked (i) whether this symbiont also confers protection against a phylogenetically distant group of parasitoids (Aphelinidae) and (ii) whether there are consistent differences in the effects of bacteria found in pea aphid biotypes adapted to different host plants. We found that some H. defensa strains do provide protection against an aphelinid parasitoid Aphelinus abdominalis. Hamiltonella defensa from the Lotus biotype provided high resistance to A. abdominalis and moderate to low resistance to Aphidius ervi, while the reverse was seen from Medicago biotype isolates. Aphids from Ononis showed no evidence of symbiont-mediated protection against either wasp species and were relatively vulnerable to both. Our results may reflect the different selection pressures exerted by the parasitoid community on aphids feeding on different host plants, and could help explain the maintenance of genetic diversity in bacterial symbionts.

Data from: Global assessment of the impact of type 2 diabetes on sleep through specific questionnaires. A case-control study

Type 2 diabetes (T2D) is an independent risk factor for sleep breathing disorders. However, it is unknown whether T2D affects daily somnolence and quality of sleep independently of the impairment of polysomnographic parameters. Material and Methods: A case-control study including 413 patients with T2D and 413 non-diabetic subjects, matched by age, gender, BMI, and waist and neck circumferences. A polysomnography was performed and daytime sleepiness was evaluated using the Epworth Sleepiness Scale (ESS). In addition, 135 subjects with T2D and 45 controls matched by the same previous parameters were also evaluated through the Pittsburgh Sleep Quality Index (PSQI) to calculate sleep quality. Results: Daytime sleepiness was higher in T2D than in control subjects (p=0.003), with 23.9% of subjects presenting an excessive daytime sleepiness (ESS>10). Patients with fasting plasma glucose (FPG ?13.1 mmol/l) were identified as the group with a higher risk associated with an ESS>10 (OR 3.9, 95% CI 1.8-7.9, p=0.0003). A stepwise regression analyses showed that the presence of T2D, baseline glucose levels and gender but not polysomnographic parameters (i.e apnea-hyoapnea index or sleeping time spent with oxigen saturation lower than 90%) independently predicted the ESS score. In addition, subjects with T2D showed higher sleep disturbances [PSQI: 7.0 (1.0-18.0) vs. 4 (0.0-12.0), p<0.001]. Conclusion: The presence of T2D and high levels of FPG are independent risk factors for daytime sleepiness and adversely affect sleep quality. Prospective studies addressed to demonstrate whether glycemia optimization could improve the sleep quality in T2D patients seem warranted.

Data from: Evolution of the exclusively human-pathogen Neisseria gonorrhoeae: human-specific engagement of immunoregulatory Siglecs

Neisseria gonorrhoeae causes the sexually transmitted disease gonorrhea exclusively in humans and uses multiple strategies to infect, including acquisition of host sialic acids that cap and mask lipooligosaccharide termini, while restricting complement activation. We hypothesized that gonococci selectively target human anti-inflammatory sialic acid-recognizing Siglec receptors on innate immune cells to blunt host responses, and that pro-inflammatory Siglecs and SIGLEC pseudogene polymorphisms represent host evolutionary adaptions to counteract this interaction. N. gonorrhoeae can indeed engage multiple human but not chimpanzee CD33rSiglecs expressed on innate immune cells and in the genitourinary tract––including Siglec-11 (inhibitory) and Siglec-16 (activating), which we detected for the first time on human cervical epithelium. Surprisingly, in addition to LOS-sialic acid, we found that gonococcal porin (PorB) mediated binding to multiple Siglecs. PorB also bound preferentially to human Siglecs and not chimpanzee orthologs, modulating host immune reactions in a human-specific manner. Lastly, we studied the distribution of null SIGLEC polymorphisms in a Namibian cohort with a high prevalence of gonorrhea, and found that uninfected women preferentially harbor functional SIGLEC16 alleles encoding an activating immune receptor. These results contribute to the understanding of the human specificity of N. gonorrhoeae and how it evolved to evade the human immune defense.

Data from: Plant odour and sex pheromone are integral elements of specific mate recognition in an insect herbivore

Specific mate recognition relies on the chemical senses in most animals, and especially in nocturnal insects. Two signal types mediate premating olfactory communication in terrestrial habitats: sex pheromones, which blend into an atmosphere of plant odorants. We show that host plant volatiles affect the perception of sex pheromone in males of the African cotton leafworm Spodoptera littoralis and that pheromone and plant volatiles are not perceived as independent messages. In clean air, S. littoralis males are attracted to single synthetic pheromone components or even the pheromone of a sibling species, Oriental cotton leafworm S. litura. Presence of host plant volatiles, however, reduces the male response to deficient or heterospecific pheromone signals. That plant cues enhance discrimination of sex pheromone quality confirms the idea that specific mate recognition in noctuid moths has evolved in concert with adaptation to host plants. Shifts in either female host preference or sex pheromone biosynthesis give rise to new communication channels that have the potential to initiate or contribute to reproductive isolation.

opencc-zeroDec 2013View details →

Data from: Fascicles from energy-storing tendons show an age-specific response to cyclic fatigue loading

Some tendons, such as the human Achilles and equine superficial digital flexor tendon (SDFT), act as energy stores, stretching and recoiling to increase efficiency during locomotion. Our previous observations of rotation in response to applied strain in SDFT fascicles suggest a helical structure, which may provide energy-storing tendons with a greater ability to extend and recoil efficiently. Despite this specialization, energy-storing tendons are prone to age-related tendinopathy. The aim of this study was to assess the effect of cyclic fatigue loading (FL) on the microstructural strain response of SDFT fascicles from young and old horses. The data demonstrate two independent age-related mechanisms of fatigue failure; in young horses, FL caused low levels of matrix damage and decreased rotation. This suggests that loading causes alterations to the helix substructure, which may reduce their ability to recoil and recover. By contrast, fascicles from old horses, in which the helix is already compromised, showed greater evidence of matrix damage and suffer increased fibre sliding after FL, which may partially explain the age-related increase in tendinopathy. Elucidation of helix structure and the precise alterations occurring owing to both ageing and FL will help to develop appropriate preventative and repair strategies for tendinopathy.

Data from: Genetic diversity analyses reveal first insights into breed-specific selection signatures within Swiss goat breeds

We used genotype data from the caprine 50k Illumina BeadChip for the assessment of genetic diversity within and between 10 local Swiss goat breeds. Three different cluster methods allowed the goat samples to be assigned to the respective breed groups, whilst the samples of Nera Verzasca and Tessin Grey goats could not be differentiated from each other. The results of the different genetic diversity measures show that Appenzell, Toggenburg, Valais and Booted goats should be prioritized in future conservation activities. Furthermore, we examined runs of homozygosity (ROH) and compared genomic inbreeding coefficients based on ROH (FROH) with pedigree-based inbreeding coefficients (FPED). The linear relationship between FROH and FPED was confirmed for goats by including samples from the three main breeds (Saanen, Chamois and Toggenburg goats). FROH appears to be a suitable measure for describing levels of inbreeding in goat breeds with missing pedigree information. Finally, we derived selection signatures between the breeds. We report a total of 384 putative selection signals. The 25 most significant windows contained genes known for traits such as: coat color variation (MITF, KIT, ASIP), growth (IGF2, IGF2R, HRAS, FGFR3) and milk composition (PITX2). Several other putative genes involved in the formation of populations, which might have been selected for adaptation to the alpine environment, are highlighted. The results provide a contemporary background for the management of genetic diversity in local Swiss goat breeds.

opencc-zeroDec 2011View details →

Data from: Inbreeding depression in self-incompatible North American Arabidopsis lyrata: disentangling genomic and S-locus specific genetic load

Newly formed selfing lineages may express recessive genetic load and suffer inbreeding depression. This can have a genome-wide genetic basis, or be due to loci linked to genes under balancing selection. Understanding the genetic architecture of inbreeding depression is important in the context of the maintenance of self-incompatibility and understanding the evolutionary dynamics of S-alleles. We addressed this using North-American subspecies of Arabidopsis lyrata. This species is normally self-incompatible and outcrossing, but some populations have undergone a transition to selfing. The goals of this study were to: (1) quantify the strength of inbreeding depression in North-American populations of A. lyrata; and (2) disentangle the relative contribution of S-linked genetic load compared with overall inbreeding depression. We enforced selfing in self-incompatible plants with known S-locus genotype by treatment with CO2, and compared the performance of selfed vs outcrossed progeny. We found significant inbreeding depression for germination rate (δ=0.33), survival rate to 4 weeks (δ=0.45) and early growth (δ=0.07), but not for flowering rate. For two out of four S-alleles in our design, we detected significant S-linked load reflected by an under-representation of S-locus homozygotes in selfed progeny. The presence or absence of S-linked load could not be explained by the dominance level of S-alleles. Instead, the random nature of the mutation process may explain differences in the recessive deleterious load among lineages.

Data from: The evolution of fungal substrate specificity in a widespread group of crustose lichens

Lichens exhibit varying degrees of specialization with regard to the surfaces they colonize, ranging from substrate generalists to strict substrate specialists. Though long recognized, the causes and consequences of substrate specialization are poorly known. Using a phylogeny of a 150-200 MYA clade of lichen fungi, we asked whether substrate niche is phylogenetically conserved, which substrates are ancestral, whether specialists arise from generalists or vice versa, and how specialization affects speciation/extinction processes. We found strong phylogenetic signal for niche conservatism. Specialists evolved into generalists and back again, but transitions from generalism to specialism were more common than the reverse. Our models suggest that for this group of fungi, "escape" from specialization for soil, rock and bark occurred, but specialization for wood foreclosed evolution away from that substrate type. In parallel, speciation models showed positive diversification rates for soil and rock dwellers but not other specialists. Patterns in the studied group suggest that fungal substrate specificity is a key determinant of evolutionary trajectory for the entire lichen symbiosis.

opencc-zeroDec 2013View details →

Data from: Metabolic erosion primarily through mutation accumulation, and not tradeoffs, drives limited evolution of substrate specificity in Escherichia coli

Evolutionary adaptation to a constant environment is often accompanied by specialization and a reduction of fitness in other environments. We assayed the ability of the Lenski Escherichia coli populations to grow on a range of carbon sources after 50,000 generations of adaptation on glucose. Using direct measurements of growth rates, we demonstrated that declines in performance were much less widespread than suggested by previous results from Biolog assays of cellular respiration. Surprisingly, there were many performance increases on a variety of substrates. In addition to the now famous example of citrate, we observed several other novel gains of function for organic acids that the ancestral strain only marginally utilized. Quantitative growth data also showed that strains with higher mutation rate exhibited significantly more declines, suggesting that most metabolic erosion was driven by mutation accumulation and not by physiological tradeoffs. These reductions in growth by mutator strains were ameliorated by growth at lower temperature, consistent with the hypothesis that this metabolic erosion is largely caused by destabilizing mutations to the associated enzymes. We further hypothesized that reductions in growth rate would be greatest for substrates used most differently from glucose, and we used flux balance analysis to formulate this question quantitatively. To our surprise, we found no significant relationship between decreases in growth and dissimilarity to glucose metabolism. Taken as a whole, these data suggest that in a single resource environment, specialization does not mainly result as an inevitable consequence of adaptive tradeoffs, but rather due to the gradual accumulation of disabling mutations in unused portions of the genome.

Data from: Venom of prey-specialised spiders is more toxic to their preferred prey: a result of prey-specific toxins

1. In specialised predators a variety of adaptations have evolved to such a level of specificity that they allow very effective exploitation of focal prey. Venom is an essential adaptive trait of predatory venomous species, such as spiders, yet our knowledge of spider venom is incomplete. 2. In agreement with the prey preference hypothesis, we expected that the venom of spider specialists should be more toxic to focal than to alternative prey, because it is composed of prey-specific toxins. 3. Here we used spiders with three types of trophic specialisations: specialists that were ant-eating, termite-eating, and spider-eating. We compared the efficacy of prey capture of preferred and alternative prey (measured as paralysis latency) with that of related generalists and profiled the venom of the studied species using proteomic methods. We used 22 spider species: six myrmecophagous, two termitophagous, three araneophagous, and 11 euryphagous generalist species belonging to different families. 4. We found that ten of the eleven specialist species induced significantly shorter paralysis latency in preferred prey than in alternative prey. Generalists exhibited either similar efficiency on both prey types or slightly higher efficiency on preferred prey. 5. Multivariate analysis of proteomic profiles (peptides and proteins) revealed significant differences between trophic specialisations, particularly in peptides. Specialists appear to have venom composed of unique specific compounds as revealed by the multivariate ordination and indicator analysis. These components are likely prey-specific toxins.

opencc-zeroJul 2019View details →

Data from: Modality-specific impairment of learning by a neonicotinoid pesticide

Neonicotinoid pesticides can impair bees' ability to learn and remember information about flowers, skills that are critical for effective foraging. Although these cognitive impacts may contribute to broader effects on health and performance, to date they have largely been assayed in simplified protocols that consider learning in a single sensory modality, usually olfaction. Given that real flowers display a variety of potentially useful signals, we assessed the effects of acute neonicotinoid exposure on multimodal learning in free-flying bumblebees. We found that neonicotinoid consumption differentially impacted learning of floral stimuli, impairing scent, but not colour, learning. These findings raise questions about the mechanisms by which pesticides might differentially impair sensory systems, with implications for how neonicotinoids affect multiple aspects of bee ecology.

Data from: Visual landmarks sharpen grid cell metric and confer context specificity to neurons of the medial entorhinal cortex

Neurons of the medial entorhinal cortex (MEC) provide spatial representations critical for navigation. In this network, the periodic firing fields of grid cells act as a metric element for position. The location of the grid firing fields depends on interactions between self-motion information, geometrical properties of the environment and nonmetric contextual cues. Here, we test whether visual information, including nonmetric contextual cues, also regulates the firing rate of MEC neurons. Removal of visual landmarks caused a profound impairment in grid cell periodicity. Moreover, the speed code of MEC neurons changed in darkness and the activity of border cells became less confined to environmental boundaries. Half of the MEC neurons changed their firing rate in darkness. Manipulations of nonmetric visual cues that left the boundaries of a 1D environment in place caused rate changes in grid cells. These findings reveal context specificity in the rate code of MEC neurons.

opencc-zeroDec 2016View details →

Data from: Optimal nutrient exchange and immune responses operate in partner specificity in the cnidarian-dinoflagellate symbiosis

The relationship between reef-building corals and phototrophic dinoflagellates of the genus Symbiodinium is fundamental to the functioning of coral reef ecosystems. It has been suggested that reef corals may adapt to climate change by changing their dominant symbiont type to a more thermally tolerant one, although the capacity for such a community shift is potentially hindered by the compatibility of different host-symbiont pairings. Here we combined transcriptomic and metabolomic analyses to characterize the molecular, cellular, and physiological processes that underlie this compatibility, with a particular focus on Symbiodinium trenchii, an opportunistic, thermally tolerant symbiont that flourishes in coral tissues after bleaching events. Symbiont-free individuals of the sea anemone Exaiptasia pallida (commonly referred to as Aiptasia), an established model system for the study of the cnidarian-dinoflagellate symbiosis, were colonized with the "normal" (homologous) symbiont Symbiodinium minutum and the heterologous S. trenchii. Analysis of the host gene and metabolite expression profiles revealed that heterologous symbionts induced an expression pattern intermediate between the typical symbiotic state and the aposymbiotic state. Furthermore, integrated pathway analysis revealed that increased catabolism of fixed carbon stores, metabolic signaling, and immune processes occurred in response to the heterologous symbiont type. Our data suggest that both nutritional provisioning and the immune response induced by the foreign "invader" are important factors in determining the capacity of corals to adapt to climate change through the establishment of novel symbioses.

opencc-zeroDec 2016View details →

Data from: Linking intra-specific trait variation to community abundance dynamics improves ecological predictability by revealing a growth-defence trade-off

1.Intraspecific trait change, including altered behaviour or morphology, can drive temporal variation in inter-specific interactions and population dynamics. In turn, variation in species' interactions and densities can alter the strength and direction of trait change. The resulting feedback between species′ traits and abundance permits a wide range of community dynamics that would not be expected from ecological theories purely based on species abundances. Despite the theoretical importance of these interrelated processes, unambiguous experimental evidence of how intraspecific trait variation modifies species interactions and population dynamics and how this feeds back to influence trait variation is currently required. 2.We investigate the role of trait-mediated demography in determining community dynamics and examine how ecological interactions influence trait change. We concurrently monitored the dynamics of community abundances and individual traits in an experimental microbial predator-prey-resource system. Using this data, we parameterized a trait dependent community model to identify key ecologically relevant traits and to link trait dynamics with those of species abundances. 3.Our results provide clear evidence of a feedback between trait change, demographic rates and species dynamics. The inclusion of trait-abundance feedbacks into our population model improved the predictability of ecological dynamics from r-squared of 34% to 57% and confirmed theoretical expectations of density dependent population growth and species interactions in the system. 4.Additionally, our model revealed that the feedbacks were underpinned by a trade-off between population growth and anti-predatory defence. High predator abundance was linked to a reduction in prey body size. This prey size decrease was associated with a reduction in its rate of consumption by predators and a decrease in its resource consumption. 5.Modelling trait-abundance feedbacks allowed us to pinpoint the underlying life history trade-off which links trait and abundance dynamics. These results show that accounting for trait-abundance feedbacks has the potential to improve understanding and predictability of ecological dynamics.

Data from: Nutrient-specific compensation for seasonal cold stress in a free-ranging temperate colobine monkey

1. Homeostatic responses of animals to environmentally-induced changes in nutrient requirements provide a powerful basis for predictive ecological models, and yet such responses are virtually unstudied in the wild. 2. We tested for macronutrient-specific compensatory feeding responses by free-ranging golden snub-nosed monkeys (Rhinopithecus roxellana) inhabiting high altitude temperate forests where they experience a substantial difference in ambient temperature in cold winters vs. warmer springs. The monkeys had free access to natural foods throughout the year, and to ensure that any seasonal differences in nutrient intake were due to homeostatic compensation and not constraints on food availability, we studied the monkeys during periods in which they were provisioned with the same amount of supplementary foods in winter and spring. 3. Thermoregulatory energy costs in winter and spring were calculated using partitional calorimetric estimations of convective and radiative heat loss obtained from thermal imaging of free-ranging monkeys in situ. Daily nutrient intakes were measured using continuous focal follows (average 6.9 h/day) of free-ranging individuals (27 in spring and 28 in winter). 4. We used a nutritional geometry framework to integrate these data and test three predictions: i. In order to remain thermoneutral (balance heat loss with heat expenditure), golden snub-nosed monkeys decrease daily energy consumption during the spring compared to winter, ii. Decreased energy intake is accomplished specifically by reducing intake of the primary energetic nutrients, carbohydrate and lipid, relative to protein, and iii. The seasonal reduction in ingested fat and carbohydrate calories will quantitatively match the reduction in thermoregulatory costs in spring compared with winter. 5. Our results showed that energy intake in spring was reduced to almost half (55%) of that in winter. As predicted, this was achieved by specifically reducing fat and carbohydrate consumption with protein intake unchanged, by a quantity (326 kJ/mbm) that almost exactly matched the seasonal difference in the daily energetic costs of thermoregulation (329 kJ/mbm). 6. This is the first study to test for a match between nutrient-specific homeostatic compensation and environmentally-induced perturbations in nutrient requirements in free-ranging animals, and underpins the potential for the homeostasis framework to provide predictive power to ecological models.