Physiological acclimatization in Hawaiian corals following a 22-month shift in baseline seawater temperature and pH.

Climate change poses a major threat to coral reefs. We conducted an outdoor 22-month experiment to investigate if coral could not just survive, but also physiologically cope, with chronic ocean warming and acidification conditions expected later this century under the Paris Climate Agreement. We recorded survivorship and measured eleven phenotypic traits to evaluate the holobiont responses of Hawaiian coral: color, Symbiodiniaceae density, calcification, photosynthesis, respiration, total organic carbon flux, carbon budget, biomass, lipids, protein, and maximum Artemia capture rate. Survivorship was lowest in Montipora capitata and only some survivors were able to meet metabolic demand and physiologically cope with future ocean conditions. Most M. capitata survivors bleached through loss of chlorophyll pigments and simultaneously experienced increased respiration rates and negative carbon budgets due to a 236% increase in total organic carbon losses under combined future ocean conditions. Porites compressa and Porites lobata had the highest survivorship and coped well under future ocean conditions with positive calcification and increased biomass, maintenance of lipids, and the capacity to exceed their metabolic demand through photosynthesis and heterotrophy. Thus, our findings show that significant biological diversity within resilient corals like Porites, and some genotypes of sensitive species, will persist this century provided atmospheric carbon dioxide levels are controlled. Since Porites corals are ubiquitous throughout the world's oceans and often major reef builders, the persistence of this resilient genus provides hope for future reef ecosystem function globally.

Habitat aridity as a determinant of the trade-off between water conservation and evaporative heat loss in bats.

The maintenance of water balance in arid environments might represent a formidable challenge for Chiroptera, since they have high surface-to-volume ratios. In deserts, bats conserve water, for example, using daily torpor, but they also might experience episodic heat bouts, when they may need to increase total evaporative water loss (TEWL) to thermoregulate. We hypothesized that in bats, habitat aridity and its variability determine a trade-off between water conservation and thermoregulation via evaporative means. To test this hypothesis, we collated data from the literature of 22 species of bats on TEWL, body temperature and resting metabolic rate, in torpor and euthermy. We also collected data on ambient temperature (Ta) and precipitation of the locations where bats were captured, calculated an aridity index, and built an index of variability of the environment. After correcting for phylogeny, we found that, as aridity and variability of the environment increased, bats had lower values of TEWL, but the rate at which TEWL increases with Ta was higher, supporting our hypothesis. These results suggest that at high Ta there is a trade-off between water conservation and evaporative heat loss in bats. The evolution of physiological mechanisms that allow water conservation and tolerance to conditions of high Ta without access to free water might thus be crucial to explain the distribution of desert bats.

Lipid composition of the stratum corneum in different regions of the body of Kuhl's pipistrelle from the Negev Desert, Israel.

The most superficial epidermal layer in endotherms is the stratum corneum (SC), which is composed of dead corneocytes embedded in a lipid matrix with free fatty acids, cholesterol, ceramides, and cerebrosides; the lipid composition of the SC determines its permeability to water vapor. Lipids that are more polar, have longer hydrocarbon chains, and are less bulky are often packed in more ordered phase states to slow cutaneous evaporative water loss (CEWL); these lipids also resist transitions to more disordered phases at high ambient temperatures (Ta). In bats, wing and tail membranes (wing patagia and tail uropatagium, respectively) allow powered flight, but increase surface area, and hence CEWL, with implications for survival in arid environments. We captured Pipistrellus kuhlii from an arid habitat and measured the lipid composition of the SC of the plagiopatagium in the wing, the uropatagium, and the non-membranous region (NMR) of the body using thin layer chromatography and reversed phase high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. The patagia contained more cholesterol and shorter-chained ceramides, and fewer cerebrosides than the NMR, indicating that the lipid phase transition temperature in the patagia is lower than in the NMR. Thus, at moderate Ta the lipids in the SC in all body regions will remain in an ordered phase state, allowing water conservation; but as Ta increases, the lipids in the SC of the patagia will more easily transition into a disordered phase, resulting in increased CEWL from the patagia facilitating efficient heat dissipation in hot environments.

Role of the gut microbiome in mediating standard metabolic rate after dietary shifts in the viviparous cockroach, Diploptera punctata.

Diet may be a significant determinant of insect gut microbiome composition. However, the extent to which dietary shifts shape both the composition and relevant functions of insect gut microbiomes, and ultimately impact host energy balance (i.e. metabolic phenotype), is not well understood. We investigated the impacts of diet switching on Diploptera punctata females maintained on a dog food (DF) diet relative to those fed a comparatively sub-optimal cellulose-amended dog food (CADF) diet for 4 weeks. After this period, dietary shift resulted in a significantly higher average mass-specific standard metabolic rate (SMR) in CADF-fed females compared with DF-fed females. We also uncovered significant 13C-enrichment in DF-fed insect samples relative to CADF-fed insect samples and lowered bacterial essential amino acid (EAA) provisioning in CADF-fed samples. Differences in SMR and EAA provisioning were not accompanied by significant differences in overall microbiome composition between the two groups. However, cellulolytic and nitrogen-fixing bacterial families dominant in wild omnivorous cockroaches and wood-feeding termites were significantly enriched in CADF-fed females than in DF-fed females, at the end of the study. We propose that these changes in microbiome composition after dietary shifts are associated with changes in EAA provisioning and possibly SMR. Further studies are needed to comprehensively understand the relative importance of gut microbial functions among the complexity of factors known to underscore SMR responses in insects under varying dietary conditions.

The role of gut microbiota in the regulation of standard metabolic rate in female Periplaneta americana.

Insect gut microbiota contribute significantly to host nutritional ecology. Disrupting insect gut microbial assemblages impacts nutrient provisioning functions, and can potentially affect host standard metabolic rate (SMR), a measure of host energy balance. In this study, we evaluated the effect of disrupting gut microbial assemblages on the SMR of female Periplaneta americana cockroaches fed dog food (DF, high protein/carbohydrate (p/c) ratio), and cellulose-amended dog food (CADF, 30% dog food, 70% cellulose, low p/c ratio) diets, supplemented with none, low, or high antibiotic doses. Bacterial loads decreased significantly between diet types (P = 0.04) and across antibiotic doses (P = 0.04). There was a significant diet type x antibiotic dose interaction on SMR of females on both diets (P = 0.05) by the end of the seven-day experimental period. In CADF-fed females, SMR decreased linearly with decreasing bacterial load. However, SMR of DF-fed females on the low dose was significantly higher than those in the control and high dose groups. This is interpreted as a diet-dependent response by low dose DF-fed females to the loss of nutritional services provided by gut bacteria. Severe reductions in bacterial load at high doses reduced SMR of females on both diet types. This study provides insights into the potential role of gut bacteria as modulators of host energy expenditure under varying dietary conditions.

Compositional differences among female-associated and embryo-associated microbiota of the viviparous Pacific Beetle cockroach, Diploptera punctata.

All cockroach species, except one, harbor the endosymbiont Blattabacterium, transmitted from females to embryos. Adult cockroaches acquire non-Blattabacterium bacteria as part of their gut microbiota over time, but our knowledge of the possible transmission of these non-Blattabacterium bacteria from females to embryos is rudimentary. We characterized the gut microbiota of gravid viviparous Diploptera punctata females and the non-Blattabacterium microbiota of associated developing embryos, as well as the gut microbiota of non-gravid females, and the microbiota of orphan embryos (females not included), following high-throughput sequencing of the 16S rRNA gene to assess bacterial transference. We determined significant differences in community composition between gravid females and associated embryos and overall greater similarity in community composition among embryos than adult females. Results suggest various routes of transference of bacteria from females or the environment to embryos. The bacterial families Halomonadaceae and Shewanellaceae were more abundant in embryos than in gravid females. The functional relevance of these families remains to be elucidated, but provisioning of amino acids deficient in the brood sac secretion is a possibility. Overall, our results highlight the need for further studies investigating the uptake and selective screening of microbes by D. punctata embryos, as well as their functions.

PNPLA1 has a crucial role in skin barrier function by directing acylceramide biosynthesis.

Mutations in patatin-like phospholipase domain-containing 1 (PNPLA1) cause autosomal recessive congenital ichthyosis, but the mechanism involved remains unclear. Here we show that PNPLA1, an enzyme expressed in differentiated keratinocytes, plays a crucial role in the biosynthesis of ω-O-acylceramide, a lipid component essential for skin barrier. Global or keratinocyte-specific Pnpla1-deficient neonates die due to epidermal permeability barrier defects with severe transepidermal water loss, decreased intercellular lipid lamellae in the stratum corneum, and aberrant keratinocyte differentiation. In Pnpla1-/- epidermis, unique linoleate-containing lipids including acylceramides, acylglucosylceramides and (O-acyl)-ω-hydroxy fatty acids are almost absent with reciprocal increases in their putative precursors, indicating that PNPLA1 catalyses the ω-O-esterification with linoleic acid to form acylceramides. Moreover, acylceramide supplementation partially rescues the altered differentiation of Pnpla1-/- keratinocytes. Our findings provide valuable insight into the skin barrier formation and ichthyosis development, and may contribute to novel therapeutic strategies for treatment of epidermal barrier defects.

The cutaneous lipid composition of bat wing and tail membranes: a case of convergent evolution with birds.

The water vapour permeability barrier of mammals and birds resides in the stratum corneum (SC), the outermost layer of the epidermis. The molar ratio and molecular arrangement of lipid classes in the SC determine the integrity of this barrier. Increased chain length and polarity of ceramides, the most abundant lipid class in mammalian SC, contribute to tighter packing and thus to reduced cutaneous evaporative water loss (CEWL). However, tighter lipid packing also causes low SC hydration, making it brittle, whereas high hydration softens the skin at the cost of increasing CEWL. Cerebrosides are not present in the mammalian SC; their pathological accumulation occurs in Gaucher's disease, which leads to a dramatic increase in CEWL. However, cerebrosides occur normally in the SC of birds. We tested the hypothesis that cerebrosides are also present in the SC of bats, because they are probably necessary to confer pliability to the skin, a quality needed for flight. We examined the SC lipid composition of four sympatric bat species and found that, as in birds, their SC has substantial cerebroside contents, not associated with a pathological state, indicating convergent evolution between bats and birds.

Metabolic rate, evaporative water loss and thermoregulatory state in four species of bats in the Negev desert.

Life in deserts is challenging for bats because of their relatively high energy and water requirements; nevertheless bats thrive in desert environments. We postulated that bats from desert environments have lower metabolic rates (MR) and total evaporative water loss (TEWL) than their mesic counterparts. To test this idea, we measured MR and TEWL of four species of bats, which inhabit the Negev desert in Israel, one species mainly restricted to hyper-arid deserts (Otonycteris hemprichii), two species from semi-desert areas (Eptesicus bottae and Plecotus christii), and one widespread species (Pipistrellus kuhlii). We also measured separately, in the same individuals, the two components of TEWL, respiratory water loss (RWL) and cutaneous evaporative water loss (CEWL), using a mask. In all the species, MR and TEWL were significantly reduced during torpor, the latter being a consequence of reductions in both RWL and CEWL. Then, we evaluated whether MR and TEWL in bats differ according to their geographic distributions, and whether those rates change with Ta and the use of torpor. We did not find significant differences in MR among species, but we found that TEWL was lowest in the species restricted to desert habitats, intermediate in the semi-desert dwelling species, and highest in the widespread species, perhaps a consequence of adaptation to life in deserts. Our results were supported by a subsequent analysis of data collected from the literature on rates of TEWL for 35 bat species from desert and mesic habitats.

The importance of the lipoxygenase-hepoxilin pathway in the mammalian epidermal barrier.

This review covers the background to discovery of the two key lipoxygenases (LOX) involved in epidermal barrier function, 12R-LOX and eLOX3, and our current views on their functioning. In the outer epidermis, their consecutive actions oxidize linoleic acid esterified in ω-hydroxy-ceramide to a hepoxilin-related derivative. The relevant background to hepoxilin and trioxilin biochemistry is briefly reviewed. We outline the evidence that linoleate in the ceramide is the natural substrate of the two LOX enzymes and our proposal for its importance in construction of the epidermal water barrier. Our hypothesis is that the oxidation promotes hydrolysis of the oxidized linoleate moiety from the ceramide. The resulting free ω-hydroxyl of the ω-hydroxyceramide is covalently bound to proteins on the surface of the corneocytes to form the corneocyte lipid envelope, a key barrier component. Understanding the role of the LOX enzymes and their hepoxilin products should provide rational approaches to ameliorative therapy for a number of the congenital ichthyoses involving compromised barrier function. This article is part of a Special Issue entitled The Important Role of Lipids in the Epidermis and their Role in the Formation and Maintenance of the Cutaneous Barrier. Guest Editors: Kenneth R. Feingold and Peter Elias.

Waking to drink: rates of evaporative water loss determine arousal frequency in hibernating bats.

Bats hibernate to cope with low ambient temperatures (T(a)) and low food availability during winter. However, hibernation is frequently interrupted by arousals, when bats increase body temperature (T(b)) and metabolic rate (MR) to normothermic levels. Arousals account for more than 85% of a bat's winter energy expenditure. This has been associated with variation in T(b), T(a) or both, leading to a single testable prediction, i.e. that torpor bout length (TBL) is negatively correlated with T(a) and T(b). T(a) and T(b) were both found to be correlated with TBL, but correlations alone cannot establish a causal link between arousal and T(b) or T(a). Because hydration state has also been implicated in arousals from hibernation, we hypothesized that water loss during hibernation creates the need in bats to arouse to drink. We measured TBL of bats (Pipistrellus kuhlii) at the same T(a) but under different conditions of humidity, and found an inverse relationship between TBL and total evaporative water loss, independent of metabolic rate, which directly supports the hypothesis that hydration state is a cue to arousal in bats.

Lipid composition of the stratum corneum and cutaneous water loss in birds along an aridity gradient.

Intercellular and covalently bound lipids within the stratum corneum (SC), the outermost layer of the epidermis, are the primary barrier to cutaneous water loss (CWL) in birds. We compared CWL and intercellular SC lipid composition in 20 species of birds from desert and mesic environments. Furthermore, we compared covalently bound lipids with CWL and intercellular lipids in the lark family (Alaudidae). We found that CWL increases in birds from more mesic environments, and this increase was related to changes in intercellular SC lipid composition. The most consistent pattern that emerged was a decrease in the relative amount of cerebrosides as CWL increased, a pattern that is counterintuitive based on studies of mammals with Gaucher disease. Although covalently bound lipids in larks did not correlate with CWL, we found that covalently bound cerebrosides correlated positively with intercellular cerebrosides and intercellular cholesterol ester, and intercellular cerebrosides correlated positively with covalently bound free fatty acids. Our results led us to propose a new model for the organization of lipids in the avian SC, in which the sugar moieties of cerebrosides lie outside of intercellular lipid layers, where they may interdigitate with adjacent intercellular cerebrosides or with covalently bound cerebrosides.

The relationship between cutaneous water loss and thermoregulatory state in Kuhl's pipistrelle Pipistrellus kuhlii, a Vespertillionid bat.

Total evaporative water loss is the sum of respiratory water loss (RWL) and cutaneous water loss (CWL) and constitutes the main avenue of water loss in bats. Because bats fly and have large surface-to-volume ratios, they potentially have high rates of RWL and CWL. Most species of small insectivorous bats have the ability to reduce their body temperature (T(b)) at rest, which substantially reduces energy expenditure and water loss. We hypothesized that bats reduce evaporative water loss during bouts of deep hypothermia (torpor) by decreasing RWL and CWL. We measured T(b), RWL, CWL, and resting metabolic rate (RMR) in Kuhl's pipistrelle Pipistrellus kuhlii, a small insectivorous bat. In support of our hypothesis, we found that RWL decreased with decreasing RMR. We found that CWL was lower in torpid individuals than in normothermic bats; however, bats in deep torpor had similar or higher CWL than bats in shallow torpor, suggesting that they exert a less effective physiological control over CWL when in deep torpor. Because insectivorous bats spend most of their lives in torpor or hibernation, the regulation of CWL in different heterothermic states has relevant ecological and evolutionary consequences.

A phylogenetic approach to total evaporative water loss in mammals.

Maintaining appropriate water balance is a constant challenge for terrestrial mammals, and this problem can be exacerbated in desiccating environments. It has been proposed that natural selection has provided desert-dwelling mammals physiological mechanisms to reduce rates of total evaporative water loss. In this study, we evaluated the relationship between total evaporative water loss and body mass in mammals by using a recent phylogenetic hypothesis. We compared total evaporative water loss in 80 species of arid-zone mammals to that in 56 species that inhabit mesic regions, ranging in size from 4 g to 3,500 kg, to test the hypothesis that mammals from arid environments have lower rates of total evaporative water loss than mammals from mesic environments once phylogeny is taken into account. We found that arid species had lower rates of total evaporative water loss than mesic species when using a dichotomous variable to describe habitat (arid or mesic). We also found that total evaporative water loss was negatively correlated with the average maximum and minimum environmental temperature as well as the maximum vapor pressure deficit of the environment. Annual precipitation and the variable Q (a measure of habitat aridity) were positively correlated with total evaporative water loss. These results support the hypothesis that desert-dwelling mammals have lower rates of total evaporative water loss than mesic species after controlling for body mass and evolutionary relatedness regardless of whether categorical or continuous variables are used to describe habitat.

Climate change and cutaneous water loss of birds.

There is a crucial need to understand how physiological systems of animals will respond to increases in global air temperature. Water conservation may become more important for some species of birds, especially those living in deserts. Lipids of the stratum corneum (SC), the outer layer of the epidermis, create the barrier to water vapor diffusion, and thus control cutaneous water loss (CWL). An appreciation of the ability of birds to change CWL by altering lipids of the skin will be important to predict responses of birds to global warming. The interactions of these lipids are fundamental to the modulation of water loss through skin. Cerebrosides, with their hexose sugar moiety, are a key component of the SC in birds, but how these lipids interact with other lipids of the SC, or how they form hydrogen bonds with water molecules, to form a barrier to water vapor diffusion remains unknown. An understanding of how cerebrosides interact with other lipids of the SC, and of how the hydroxyl groups of cerebrosides interact with water molecules, may be a key to elucidating the control of CWL by the SC.

Allocation of endogenous and dietary protein in the reconstitution of the gastrointestinal tract in migratory blackcaps at stopover sites.

During migratory flight, the mass of the gastrointestinal tract (GIT) and its associated organs in small birds decreases in size by as much as 40%, compared with the preflight condition because of the catabolism of protein. At stopover sites, birds need 2-3 days to rebuild their GIT so that they can restore body mass and fat reserves to continue migration. The source of protein used to rebuild the GIT may be exogenous (from food ingested) or endogenous (reallocated from other organs) or both. Because the relative contribution of these sources to rebuild the GIT of migratory birds is not yet known, we mimicked in-flight fasting and then re-feeding in two groups of blackcaps (Sylvia atricapilla), a Palearctic migratory passerine. The birds were fed a diet containing either 3% or 20% protein to simulate different refueling scenarios. During re-feeding, birds received known doses of (15)N-(l)-leucine before we measured the isotope concentrations in GIT and associated digestive organs and in locomotory muscles. We then quantified the extent to which blackcaps rebuilt their GIT with endogenous and/or dietary protein while refeeding after a fast. Our results indicate that blackcaps fed the low-protein diet incorporated less exogenous nitrogen into their tissues than birds fed the 20% protein diet. They also allocated relatively more exogenous protein to the GIT than to pectoral muscle than those birds re-fed with the high-protein diet. However, this compensation was not sufficient for birds eating the low-protein diet to rebuild their intestine at the same rate as the birds re-fed the high-protein diet. We concluded that blackcaps must choose stopover sites at which they can maximize protein intake to minimize the time it takes to rebuild their GIT and, thus, resume migration as soon as possible.

Cutaneous water loss and covalently bound lipids of the stratum corneum in nestling house sparrows (Passer domesticus L.) from desert and mesic habitats.

Lipids of the stratum corneum (SC), the outer layer of the epidermis of birds and mammals, provide a barrier to water vapor diffusion through the skin. The SC of birds consists of flat dead cells, called corneocytes, and two lipid compartments: an intercellular matrix and a monolayer of covalently bound lipids (CBLs) attached to the outer surface of the corneocytes. We previously found two classes of sphingolipids, ceramides and cerebrosides, covalently bound to corneocytes in the SC of house sparrows (Passer domesticus L.); these lipids were associated with cutaneous water loss (CWL). In this study, we collected adult and nestling house sparrows from Ohio and nestlings from Saudi Arabia, acclimated them to either high or low humidity, and measured their rates of CWL. We also measured CWL for natural populations of nestlings from Ohio and Saudi Arabia, beginning when chicks were 2 days old until they fledged. We then evaluated the composition of the CBLs of the SC of sparrows using thin layer chromatography. We found that adult house sparrows had a greater diversity of CBLs in their SC than previously described. During ontogeny, nestling sparrows increased the amount of CBLs and developed their CBLs differently, depending on their habitat. Acclimating nestlings to different humidity regimes did not alter the ontogeny of the CBLs, suggesting that these lipids represent a fundamental component of SC organization that does not respond to short-term environmental change.

Cutaneous water loss and lipids of the stratum corneum in two syntopic species of bats.

The lipid matrix of the stratum corneum (SC), the outer layer of the epidermis of mammals and birds, constitutes the barrier to diffusion of water vapor through the skin. The lipids of the SC are structured in the intercellular spaces of the mammalian epidermis in ordered layers, called lamellae, which have been postulated to prevent water loss. Lipids in the mammalian SC are mainly cholesterol, free fatty acids and ceramides, the latter forming the structural support for the lamellae. However, knowledge on how the lipid composition of the SC alters cutaneous water loss (CWL) in mammals is rudimentary, and is largely derived from studies on laboratory animals and humans. We measured CWL of individuals of two species of syntopic bats, Tadarida brasiliensis and Myotis velifer. In the first study of its kind on wild mammals, we correlated CWL with the lipid composition of the SC, measured using thin layer chromatography and high performance liquid chromatography coupled with atmospheric pressure photoionization mass spectrometry. Surface-specific CWL was 20.6% higher in M. velifer than in T. brasiliensis, although differences were not significant. Compared with individuals of M. velifer, individuals of T. brasiliensis had more classes, and a higher proportion, of polar ceramides in the SC, a feature associated with lower CWL. Individuals of T. brasiliensis also had a class of non-polar ceramides that presumably spans the lamellae and gives more cohesiveness to the lipid matrix of the SC. We conclude that qualitative and quantitative modifications of the lipid composition of the SC contribute to regulate CWL of these two species of bats.

Cutaneous water loss and the development of the stratum corneum of nestling house sparrows (Passer domesticus) from desert and mesic environments.

Evaporation through the skin contributes to more than half of the total water loss in birds. Therefore, we expect the regulation of cutaneous water loss (CWL) to be crucial for birds, especially those that live in deserts, to maintain a normal state of hydration. Previous studies in adult birds showed that modifications of the lipid composition of the stratum corneum (SC), the outer layer of the epidermis, were associated with changes in rates of CWL. However, few studies have examined the ontogeny of CWL and the lipids of the SC in nestling birds. In this study, we measured CWL and the lipid composition of the SC during development of nestlings from two populations of house sparrows, one from the deserts of Saudi Arabia and the other from mesic Ohio. We found that desert and mesic nestlings followed different developmental trajectories for CWL. Desert nestlings seemed to make a more frugal use of water than did mesic nestlings. To regulate CWL, nestlings appeared to modify the lipid composition of the SC during ontogeny. Our results also suggest a tighter regulation of CWL in desert nestlings, presumably as a result of the stronger selection pressures to which nestlings are exposed in deserts.

Developmental plasticity of cutaneous water loss and lipid composition in stratum corneum of desert and mesic nestling house sparrows.

Intercellular lipids of the stratum corneum (SC), the outer layer of the epidermis, form a barrier to water vapor diffusion through the skin. Previously, we measured cutaneous water loss (CWL) and lipid composition of the SC of adult house sparrows from two populations, one living in the deserts of Saudi Arabia and another living in mesic Ohio. Adult desert house sparrows had a lower CWL, a lower proportion of free fatty acids, and a higher proportion of ceramides and cerebrosides in the SC compared with mesic sparrows. In this study, we investigated developmental plasticity of CWL and lipid composition of the SC in desert and mesic nestling house sparrows reared in low and high humidity and compared our results with previous work on adults. We measured CWL of nestlings and analyzed the lipid composition of the SC using thin-layer chromatography. We showed that nestling house sparrows from both localities had higher CWL than adults in their natural environment, a result of major modifications of the lipid composition of the SC. The expression of plasticity in CWL seems to be a response to opposed selection pressures, thermoregulation and water conservation, at different life stages, on which regulation of CWL plays a crucial role. Desert nestlings showed a greater degree of plasticity in CWL and lipid composition of the SC than did mesic nestlings, a finding consistent with the idea that organisms exposed to more environmental stress ought to be more plastic than individuals living in more benign environments.