Gizzard Mass Research Articles

Antibacterial and anatomical defenses in an oil contaminated, vulnerable seaduck.

Oil spills have killed thousands of birds during the last 100 years, but nonlethal effects of oil spills on birds remain poorly studied. We measured phenotype characters in 819 eiders Somateria mollissima (279 whole birds and 540 wings) of which 13.6% were oiled. We tested the hypotheses that (a) the morphology of eiders does not change due to oil contamination; (b) the anatomy of organs reflects the physiological reaction to contamination, for example, increase in metabolic demand, increase in food intake, and counteracting toxic effects of oil; (c) large locomotion apparatus that facilitates locomotion increases the risk of getting oiled; and (d) individual eiders with a higher production of secretions from the uropygial grand were more likely to have oil on their plumage. We tested whether 19 characters differed between oiled and nonoiled individuals, showing a consistent pattern. The final model retained seven predictor variables showing relationships between eiders contaminated with oil and food consumption, flight, and diving abilities. We tested whether these effects were due to differences in body condition, liver mass, empty gizzard mass, or other characters that could have been affected by impaired flight and diving ability. There was no evidence of such negative impact of oiling on eiders. We found that significant exposure to oil was associated with increased diversity of antibacterial defense. Oiled eiders did not constitute a random sample, and superior diving ability as reflected by large foot area was at a selective disadvantage during oil spills. Thus, specific characteristics predispose eiders to oiling, with an adaptation to swimming, diving, and flying being traded against the costs of oiling. In contrast, individuals with a high degree of physiological plasticity may experience an advantage because their uropygial secretions counteract the effects of oil contamination.

Brain mass explains prey size selection better than beak, gizzard and body size in a benthivorous duck species.

Prey size selection in some bird species is determined by the size of the beak. However, we assumed for bird species swallowing whole prey that a cognitive process may be involved. As cognitive feature, brain mass was used. We hypothesized that the mass of the brain was more strongly positively correlated with prey size than morphological features such as beak volume, gizzard mass and body mass. We tested this hypothesis on eiders Somateria mollissima that swallow the prey whole, by using mean and maximum size of nine prey categories. Eiders were collected at the main wintering grounds in Denmark. As index of brain mass we used head volume, which is positively correlated with brain mass (r2 = 0.73). Head volume of eiders was significantly, positive correlated with mean and maximum size of blue mussels Mytilus edulis, razor clams Ensis directus and all prey sizes combined and the maximum size of draft whelk Hinia reticulata and conch Buccinum undatum. Gizzard mass was also significantly positively correlated with maximum size of draft whelk and conch. Beak volume and body mass was not significantly correlated with the size of any of the nine food items. Analyses of effect size for organs showed that head volume was positively related to prey size, whereas beak volume, gizzard mass and body mass did not show a significant positive relationship. These results indicate that cognitive processes connected to brain mass may be involved in prey size selection by eiders.

Effects of edible environmental enrichments during the rearing and laying periods in a littered aviary—Part 2: physical development of pullets and performance, egg quality, and carcass composition in laying hens

Feather pecking and cannibalism are 2 major problems in laying hens' husbandry. Although additional environmental enrichment material (EM) supply is thought to lessen these problems, consistent evidences are lacking. This study examined the effects of EM supply (pecking stones and alfalfa bales) on biological performance, carcass composition, and animal losses in a littered housing system. 2,000 brown-egg and 2,000 white-egg layer hen pullets of the genetic strains Lohmann Brown classic and Lohmann Selected Leghorn classic, respectively, were reared separately in a 16-compartment aviary system until week 18. 1,320 remaining laying hens were then transferred to a 44-compartment laying stable (weeks 19–48). Both strains were tested under 4 treatment variants (n = 150–180 per strain and per variant): V1—no EM over the entire study period; V2—the rearing period with and the laying period without EM; V3—the rearing period without and the laying period with EM; V4—EM over the entire study period. Data on development, performance, egg quality, feed intake, EM consumption, animal losses, carcass composition, and economic traits were collected. Enrichment material supply during rearing (V2 and V4, both strains) was found to globally increase not only hens' relative gizzard mass (P = 0.036) but also the cracked eggs' percentage (compared with V3; P = 0.008) and to decrease the body mass in weeks 6 (P = 0.023) and 8 (P = 0.023) and the uniformity in week 16 (P = 0.011). Enrichment material provision during the laying period (V3 and V4, both strains) increased egg weights (P = 0.028) and the mean body mass (P = 0.036); however, continuous provision of EM (V4, both strains) increased the floor eggs' percentage (P = 0.019). The EM supply did not affect mortality, loss of production days, losses due to skin and toe cannibalism, or the income over feed costs. However, the income over feed and enrichment costs of V1 hens was higher by 0.55 €/hen than that of V4 hens (P = 0.022). Therefore, EM supply cannot be recommended as a measure to increase laying performance and reduce animal losses, but its positive effects on animal welfare should be considered.

Changes in organ size and nutrient reserves of arctic terns (Sterna paradisaea) breeding near a High Arctic polynya

ABSTRACT The arctic tern (Sterna paradisaea) is a ubiquitous migratory seabird of the High Arctic, currently thought to be in decline in most of the circumpolar world, but surprisingly little is known of its biology at high latitudes. We studied organ size and nutrient reserves of arctic terns breeding beside a High Arctic polynya in Nunavut, Canada, from their arrival at the colony into the chick-rearing period. Both males and females had a decrease in gizzard size through breeding, with gizzard mass during chick-rearing 39 percent lower than on arrival at the breeding grounds. Through the duration of the breeding season, heart, liver, and small intestine showed little change, but females had higher fat and protein stores than males. Terns from this colony likely have increasing fat levels and high body condition due to proximity to a highly productive polynya, where terns appear to gain more energy than they expend during foraging trips. This suggests that though terns at this colony may be near the northern limit of their range, local conditions have a strong impact on organ and nutrient reserve dynamics of these arctic seabirds.

Individual diet differences in a molluscivore shorebird are associated with the size of body instruments for internal processing rather than for feeding

Especially in birds, it is widely found that the size of individual prey items follows the size of the instruments of prey capture, handling and processing, i.e. bill size. In fact, this is the natural history basis of major discoveries on adaptive evolution in the face of changing food resources. In some birds, e.g. the molluscivore shorebirds ingesting hard‐shelled prey, most of the prey processing takes place within the digestive tract. This study of a salvaged sample of actively feeding great knots Calidris tenuirostris accidentally drowned in fishing nets in northern China, is the first documentation of diet selection at the level of the individual in previously well‐studied molluscivore shorebirds. Diet composition was not associated with the length of the bill, but with the mass of the muscular gizzard. Gizzard mass, which unlike bill length is a phenotypically flexible trait, enables great knots to adjust to changing food resources as an individual, i.e. instantly responding to the food on offer. For migratory species like great knots which rely on seasonal sequences of interdistant feeding areas offering prey with a variety of characteristics, the capacity to individually adjust appears a key adaptation.

Within-Individual Canalization Contributes to Age-Related Increases in Trait Repeatability: A Longitudinal Experiment in Red Knots.

Age-related increases in the repeatable expression of labile phenotypic traits are often assumed to arise from an increase in among-individual variance due to differences in developmental plasticity or by means of state-behavior feedbacks. However, age-related increases in repeatability could also arise from a decrease in within-individual variance as a result of stabilizing trait expression, that is, canalization. Here we describe age-related changes in within-individual and among-individual variance components in two correlated traits-gizzard mass and exploration behavior-in a medium-sized shorebird, the red knot (Calidris canutus). Increased repeatability of gizzard mass came about due to an increase in among-individual variance, unrelated to differences in developmental plasticity, together with decreases in within-individual variance consistent with canalization. We also found canalization of exploration but no age-related increase in overall repeatability, which suggests that showing predictable expression of exploration behavior may be advantageous from a very young age onward. Contrasts between juveniles and adults in the first year after their capture provide support for the idea that environmental conditions play a key role in generating among-individual variation in both gizzard mass and exploration behavior. Our study shows that stabilization of traits occurs under constant conditions: with increased exposure to predictable cues, individuals may become more certain in their assessment of the environment allowing traits to become canalized.

Dietary fiber content affects growth, body composition, and feed intake and their associations with a major growth locus in growing male chickens of an advanced intercross population

In this study, we examined the effect of different dietary fiber concentrations on growth, body composition, and feed efficiency of growing male chickens with different genetic background. We tested whether the single nucleotide polymorphisms (SNP) in a previously identified growth locus on chromosome 4 (GGA4) between 69.3 and 77.9 Mb was associated with the traits of biologic and commercial importance in interaction with the diet. The study was conducted with growing male chickens of generation 12 of an advanced intercross population descending from the initial cross between inbred New Hampshire (high-growth) and White Leghorn (low-growth) lines. We performed a 2-way interaction analysis between the dietary fiber content and SNP genotypes. At wk 4, the growing male chickens were randomly divided into 2 diet treatments, which were housed and fed individually with either a low- (LFD, n = 38) or a high-fiber diet (HFD, n = 41) containing 0.8 and 5% lignocellulose with a crude fiber content of 60%, respectively. Body weights were recorded weekly. Body composition was examined at wk 13. Feed intake and excreta were analyzed at wk 6 and 13. The effect of dietary fiber content on body weight gain and feed intake was age-dependent. The feed conversion ratio was not altered. Feeding HFD resulted in 10% greater body weight gain (P = 0.047) and greater feed intake (P = 0.014) than LFD at wk 6. At wk 13, the visceral adipose tissue mass was 38% lower (P = 0.008) and the gizzard mass was 14% greater (P = 0.012). The reduction of total fat content in adipose tissue (P = 0.039) in the HFD was associated with reduced intramuscular fat content in drumsticks-thighs (4.2 vs. 5.1%, P = 0.004). The SNP genotypes at the GGA4 locus interacted with dietary fiber content and influenced body weight gain (P = 0.042), visceral adipose tissue mass (P ≤ 0.029), and feed intake (P ≤ 0.050). The New Hampshire alleles of SNPs on GGA4 led to increased body weight gain and feed intake (P ≤ 0.050), and tended to reduced fat deposition under HFD. Our data indicate that the addition of lignocellulose to the diet would improve productivity and carcass quality through reduced adipose tissue mass without changes in feed efficiency. Furthermore, our findings provide evidence for interaction of the target GGA4 region with dietary fiber usage affecting growth, fat deposition, and feed intake. The particular GGA4 locus may contain a single or several genes associated with growth, body composition, and feed intake.

Evolutionary design of a flexible, seasonally migratory, avian phenotype: why trade gizzard mass against pectoral muscle mass?

Migratory birds undergo impressive body remodelling over the course of an annual cycle. Prior to long-distance flights, red knots ( Calidris canutus islandica) reduce gizzard mass while increasing body mass and pectoral muscle mass. Although body mass and pectoral muscle mass are functionally linked via their joint effects on flight performance, gizzard and pectoral muscle mass are thought to be independently regulated. Current hypotheses for observed negative within-individual covariation between gizzard and pectoral muscle mass in free-living knots are based on a common factor (e.g. migration) simultaneously affecting both traits, and/or protein limitation forcing allocation decisions. We used diet manipulations to generate within-individual variation in gizzard mass and test for independence between gizzard and pectoral muscle mass within individuals outside the period of migration and under conditions of high protein availability. Contrary to our prediction, we observed a negative within-individual covariation between gizzard and pectoral muscle mass. We discuss this result as a potential outcome of an evolved mechanism underlying body remodelling associated with migration. Although our proposed mechanism requires empirical testing, this study echoes earlier calls for greater integration of studies of function and mechanism, and in particular, the need for more explicit consideration of the evolution of mechanisms underlying phenotypic design.

Morphological and digestive adjustments buffer performance: How staging shorebirds cope with severe food declines.

Organisms cope with environmental stressors by behavioral, morphological, and physiological adjustments. Documentation of such adjustments in the wild provides information on the response space in nature and the extent to which behavioral and bodily adjustments lead to appropriate performance effects. Here we studied the morphological and digestive adjustments in a staging population of migrating Great Knots Calidris tenuirostris in response to stark declines in food abundance and quality at the Yalu Jiang estuarine wetland (northern Yellow Sea, China). At Yalu Jiang, from 2011 to 2017 the densities of intertidal mollusks, the food of Great Knots, declined 15‐fold. The staple prey of Great Knots shifted from the relatively soft‐shelled bivalve Potamocorbula laevis in 2011–2012 to harder‐shelled mollusks such as the gastropod Umbonium thomasi in 2016–2017. The crushing of the mollusks in the gizzard would require a threefold to 11‐fold increase in break force. This was partially resolved by a 15% increase in gizzard mass which would yield a 32% increase in shell processing capacity. The consumption of harder‐shelled mollusks was also accompanied by reliance on regurgitates to excrete unbreakable parts of prey, rather than the usual intestinal voidance of shell fragments as feces. Despite the changes in digestive morphology and strategy, there was still an 85% reduction in intake rate in 2016–2017 compared with 2011–2012. With these morphological and digestive adjustments, the Great Knots remaining faithful to the staging site to a certain extent buffered the disadvantageous effects of dramatic food declines. However, compensation was not complete. Locally, birds will have had to extend foraging time and use a greater daily foraging range. This study offers a perspective on how individual animals may mitigate the effects of environmental change by morphological and digestive strategies and the limits to the response space of long‐distance migrating shorebirds in the wild.

Body condition of Eiders at Danish wintering grounds and at pre-breeding grounds in Åland

Breeding Eiders Somateria mollissima in the Baltic/Wadden Sea population have declined in recent decades, and several causes have been suggested. A recent study showed a modest increase in body mass from 2.20 kg by ca. 100 g during winter in Denmark, the main wintering area of the flyway population. This raises the question of whether Eiders could increase their body mass by an additional 200 g during pre-breeding to reach the average body mass of ca. 2.50 kg needed for successful reproduction. We analyzed this question by collecting Eiders at the Danish wintering grounds and pre-breeding grounds at Aland, Finland. Female Eiders increased their body condition (defined as scaled body mass index) from winter to the pre-breeding period, whereas males concurrently decreased in weight. The mean body mass of adult female Eiders was 2.23 kg in late winter, when they initiated spring migration. The mean female body mass at the pre-breeding grounds was 2.44 kg, an increase of 9.4% compared with females at the wintering grounds, while adult males decreased in weight by 3.4%. A decrease in gizzard mass was accompanied by a change in diet from large blue mussels Mytilus edulis at the wintering grounds to the small but more abundant mussels in the gizzard during pre-breeding. We conclude that female Eiders can reach a body mass of ca. 2.50 kg before breeding. Building up body condition for breeding is feasible at Aland; thus, difficulty in acquiring adequate resources for reproduction is unlikely to restrict local population size.

Allocation of body reserves during winter in eider Somateria mollissima as preparation for spring migration and reproduction

Capital breeders, such as the eider duck Somateria mollissima, accumulate resources before the start of breeding. Eiders preferentially feed on blue mussels Mytilus edulis to build up body condition during winter. We explored how body condition and gizzard mass of wintering eiders relate to mussel quality and quantity, winter climate and body condition of females at the breeding grounds. Body condition during winter (defined as scaled body mass index) of eiders increased during winter and the magnitude of the effect depended on age and mussel quality. Gizzard mass of eiders increased during winter with effects of mussel quality, mussel stocks and sex. Body condition in winter of adult females increased from the first half of January to the second half of February on average by 1.5%, equal to c. 96 g. During the same period gizzard mass of adult females increased by 12.2%, i.e., a nearly ten-fold increase compared to that observed in body condition in winter. Body condition of females at the breeding grounds in Finland (defined as body condition at hatching) was significantly positively correlated with gizzard mass in winter, but not significantly correlated with body condition in winter. Thus, eiders allocate body reserves to increase gizzard mass but less so to increase body condition in winter. This can be considered an adaptive migratory strategy of these eiders, whereby large winter (pre-migratory) gizzards increase food processing capacity, making it possible for eiders to arrive at the breeding grounds with superior body condition and a high reproductive potential.

An experimental test of state–behaviour feedbacks: gizzard mass and foraging behaviour in red knots

Summary Animals frequently exhibit consistent among‐individual differences in behavioural and physiological traits that are inherently flexible. Why should individuals differ consistently in their expression of labile traits? Recently, positive feedbacks between state and behaviour have been proposed as a possible explanation for the maintenance of consistent among‐individual differences in both state and behaviour. If state affects behaviour, and behaviour reciprocally affects state, then differences in either state or behaviour that arise among individuals even by chance could be maintained over extended periods of time. We tested for positive feedbacks experimentally using wild‐caught red knots (Calidris canutus islandica). In the wild, knots exhibit consistent among‐individual differences in digestive physiology (the mass of the muscular part of the stomach, the gizzard) and foraging behaviour (diet), two inherently labile traits. Experimentally manipulated diet quality had a large effect on gizzard mass. Experimentally manipulated gizzard mass reciprocally influenced total food eaten during ad libitum trials. The effect of gizzard mass on diet choice, though in the predicted direction, was not statistically significant. Individuals exhibited consistent differences in foraging behaviour of unknown origin independent of current gizzard mass, as well as large residual unexplained variance in foraging behaviour. These two sources of variation in foraging behaviour overruled the gizzard mass‐dependent foraging behaviour and hence eroded the treatment‐related differences in gizzard mass. We conclude that positive feedbacks between diet choice and gizzard mass play at best a limited role in maintaining among‐individual variation in gizzard mass in red knots. Furthermore, we suggest that many models of state–behaviour feedbacks likely overestimate their potential importance in maintaining long‐term among‐individual variation in labile traits because most models of state–behaviour feedbacks fail to account for the effects of additional factors that may act to disrupt the feedback loops. The among‐individual differences in diet choice observed during solitary foraging trials eroded the consistent among‐individual differences in gizzard mass observed following periods of staple diet treatments in which knots foraged in social groups. Social foraging interactions may play an important role determining the expression of foraging behaviours such as intake rate that in turn influence gizzard mass. Further studies are needed to experimentally test the role of social interactions as a mechanism generating consistent among‐individual differences in foraging behaviours and gizzard mass. A lay summary is available for this article.

Erratum for ‘Your tools disappear when you stop eating: phenotypic variation in gizzard mass of eiders’

Erratum for ‘Your tools disappear when you stop eating: phenotypic variation in gizzard mass of eiders’

Diet preferences as the cause of individual differences rather than the consequence.

Behavioural variation within a species is usually explained as the consequence of individual variation in physiology. However, new evidence suggests that the arrow of causality may well be in the reverse direction: behaviours such as diet preferences cause the differences in physiological and morphological traits. Recently, diet preferences were proposed to underlie consistent differences in digestive organ mass and movement patterns (patch residence times) in red knots (Calidris canutus islandica). Red knots are molluscivorous and migrant shorebirds for which the size of the muscular stomach (gizzard) is critical for the food processing rate. In this study, red knots (C.c.canutus, n=46) were caught at Banc d'Arguin, an intertidal flat ecosystem in Mauritania, and released with radio-tags after the measurement of gizzard mass. Using a novel tracking system (time-of-arrival), patch residence times were measured over a period of three weeks. Whether or not gizzard mass determined patch residence times was tested experimentally by offering 12 of the 46 tagged red knots soft diets prior to release; this reduced an individual's gizzard mass by 20-60%. To validate whether the observed range of patch residence times would be expected from individual diet preferences, we simulated patch residence times as a function of diet preferences via a simple departure rule. Consistent with previous empirical studies, patch residence times in the field were positively correlated with gizzard mass. The slope of this correlation, as well as the observed range of patch residence times, was in accordance with the simulated values. The 12 birds with reduced gizzard masses did not decrease patch residence times in response to the reduction in gizzard mass. These findings suggest that diet preferences can indeed cause the observed among-individual variation in gizzard mass and patch residence times. We discuss how early diet experiences can have cascading effects on the individual expression of both behavioural and physiomorphic traits. This emphasizes that to understand the ecological consequences of individual differences, knowledge of the environment during development is required.

Understanding spatial distributions: negative density-dependence in prey causes predators to trade-off prey quantity with quality.

Negative density-dependence is generally studied within a single trophic level, thereby neglecting its effect on higher trophic levels. The 'functional response' couples a predator's intake rate to prey density. Most widespread is a type II functional response, where intake rate increases asymptotically with prey density; this predicts the highest predator densities at the highest prey densities. In one of the most stringent tests of this generality to date, we measured density and quality of bivalve prey (edible cockles Cerastoderma edule) across 50 km² of mudflat, and simultaneously, with a novel time-of-arrival methodology, tracked their avian predators (red knots Calidris canutus). Because of negative density-dependence in the individual quality of cockles, the predicted energy intake rates of red knots declined at high prey densities (a type IV, rather than a type II functional response). Resource-selection modelling revealed that red knots indeed selected areas of intermediate cockle densities where energy intake rates were maximized given their phenotype-specific digestive constraints (as indicated by gizzard mass). Because negative density-dependence is common, we question the current consensus and suggest that predators commonly maximize their energy intake rates at intermediate prey densities. Prey density alone may thus poorly predict intake rates, carrying capacity and spatial distributions of predators.

Your tools disappear when you stop eating: phenotypic variation in gizzard mass of eiders

AbstractAnimals show phenotypic flexibility in their digestive system in response to seasonal changes in diet, activity, metabolic rate and reproduction. Many birds are capital breeders that alternate between periods of extensive feeding used for storage and periods of fasting during reproduction. Here, we analyzed the mass of the gizzard (gizzard mass without content) in 885 male and 348 female adult common eiders Somateria mollissima shot by Danish hunters during winter and spring in relation to alternating periods of foraging and fasting during reproduction. Gizzard mass of adult female eiders varied annually from 31 g (N = 25) during reproduction to 119 g during pre‐breeding (N = 314), or almost a four‐fold difference in mass. During winter, both male and female eiders with large gizzard mass had eaten a larger number of preferred blue mussels Mytilus edulis that constitute the main food. Adult female eiders with large gizzard mass had eaten larger but not more mussels. Individual adult females with large gizzard mass that had larger numbers of mussels in their gizzards were in superior body condition. These findings of phenotypic flexibility in gizzard mass show that individuals with larger gizzards consuming larger food items had an advantage in terms of superior body condition, potentially allowing for elevated reproductive success.

Body and organ mass dynamics during remigial moult in a wing–foot-propelled diving sea duck: the Common Eider (Atlantic) (Somateria mollissima dresseri)

Body and organ dynamics, during remigial moult, have been mainly explored on geese, dabbling ducks, and foot-propelled diving ducks, but weakly on sea ducks. This study investigated the internal changes in a wing–foot-propelled sea duck to determine the adaptive strategies implemented. Forty-five male Common Eiders (Atlantic) (Somateria mollissima dresseri Sharpe, 1871), collected in the Gulf of St. Lawrence, were dissected; their body mass, muscle mass, and organ sizes were measured. We tested three hypotheses: (1) S. m. dresseri use a strategic reduction of body mass to reduce the flightlessness duration; (2) organs will exhibit changes consistent with a trade-off between function and maintenance to save and reallocate energy and proteins to feather growth; (3) S. m. dresseri would show lower flight muscle reduction than foot-propelled diving ducks. Somateria mollissima dresseri did not lose body mass, which does not support the first hypothesis. Atrophy of the heart followed by hypertrophy and opposite changes in leg muscle mass and gizzard mass are consistent with the second hypothesis. Flight muscle mass showed lower variations than in other ducks, validating the third hypothesis. We also suggest that the lipid depletion observed early in the moult could be a strategy to reduce foraging effort and minimize the risk of damaging the growing feathers.

The Effect of Digestive Capacity on the Intake Rate of Toxic and Non-Toxic Prey in an Ecological Context.

Digestive capacity often limits food intake rate in animals. Many species can flexibly adjust digestive organ mass, enabling them to increase intake rate in times of increased energy requirement and/or scarcity of high-quality prey. However, some prey species are defended by secondary compounds, thereby forcing a toxin limitation on the forager’s intake rate, a constraint that potentially cannot be alleviated by enlarging digestive capacity. Hence, physiological flexibility may have a differential effect on intake of different prey types, and consequently on dietary preferences. We tested this effect in red knots (Calidris canutus canutus), medium-sized migratory shorebirds that feed on hard-shelled, usually mollusc, prey. Because they ingest their prey whole and crush the shell in their gizzard, the intake rate of red knots is generally constrained by digestive capacity. However, one of their main prey, the bivalve Loripes lucinalis, imposes a toxin constraint due to its symbiosis with sulphide-oxidizing bacteria. We manipulated gizzard sizes of red knots through prolonged exposure to hard-shelled or soft foods. We then measured maximum intake rates of toxic Loripes versus a non-toxic bivalve, Dosinia isocardia. We found that intake of Dosinia exponentially increased with gizzard mass, confirming earlier results with non-toxic prey, whereas intake of Loripes was independent of gizzard mass. Using linear programming, we show that this leads to markedly different expected diet preferences in red knots that try to maximize energy intake rate with a small versus a large gizzard. Intra- and inter-individual variation in digestive capacity is found in many animal species. Hence, the here proposed functional link with individual differences in foraging decisions may be general. We emphasize the potential relevance of individual variation in physiology when studying trophic interactions.

How salinity and temperature combine to affect physiological state and performance in red knots with contrasting non-breeding environments.

Migratory shorebirds inhabit environments that may yield contrasting salinity-temperature regimes-with widely varying osmoregulatory demands, even within a given species-and the question is: by which physiological means and at which organisational level do they show adjustments with respect to these demands? Red knots Calidris canutus winter in coastal areas over a range of latitudes. The nominal subspecies winters in salty areas in the tropics, whereas the subspecies Calidris canutus islandica winters in north-temperate regions of comparatively lower salinities and temperatures. In this study, both subspecies of red knot were acclimated to different salinity (28/40‰)-temperature (5/35 °C) combinations for 2-week periods. We then measured food/salt intakes, basal metabolic rate (BMR), body mass and temperature, fat and salt gland scores, gizzard mass, heat-shock proteins, heterophils/lymphocytes (H/L) ratio and plasma Na(+) to assess the responses of each taxon to osmoregulatory challenges. High salinity (HS)-warm-acclimated birds reduced food/salt intake, BMR, body mass, fat score and gizzard mass, showing that salt/heat loads constrained energy acquisition rates. Higher salt gland scores in saltier treatments indicated that its size was adjusted to higher osmoregulatory demands. Elevated plasma Na(+) and H/L ratio in high-salinity-warm-acclimated birds indicated that salt/heat loads might have a direct effect on the water-salt balance and stress responses of red knots. Subspecies had little or no effect on most measured parameters, suggesting that most adjustments reflect phenotypic flexibility rather than subspecific adaptations. Our results demonstrate how salinity and temperature affect various phenotypic traits in a migrant shorebird, highlighting the importance of considering these factors jointly when evaluating the environmental tolerances of air-breathing marine taxa.

Personality drives physiological adjustments and is not related to survival

The evolutionary function and maintenance of variation in animal personality is still under debate. Variation in the size of metabolic organs has recently been suggested to cause and maintain variation in personality. Here, we examine two main underlying notions: (i) that organ sizes vary consistently between individuals and cause consistent behavioural patterns, and (ii) that a more exploratory personality is associated with reduced survival. Exploratory behaviour of captive red knots (Calidris canutus, a migrant shorebird) was negatively rather than positively correlated with digestive organ (gizzard) mass, as well as with body mass. In an experiment, we reciprocally reduced and increased individual gizzard masses and found that exploration scores were unaffected. Whether or not these birds were resighted locally over the 19 months after release was negatively correlated with their exploration scores. Moreover, a long-term mark–recapture effort on free-living red knots with known gizzard masses at capture confirmed that local resighting probability (an inverse measure of exploratory behaviour) was correlated with gizzard mass without detrimental effects on survival. We conclude that personality drives physiological adjustments, rather than the other way around, and suggest that physiological adjustments mitigate the survival costs of exploratory behaviour. Our results show that we need to reconsider hypotheses explaining personality variation based on organ sizes and differential survival.