Evaluation of Total Body Electrical Conductivity to Estimate Body Composition of a Small Mammal

ABSTRACT The ability to measure body composition is critical for studying the physiological ecology of animals. This is particularly true for small mammals that have a high metabolic rate. We evaluated a nondestructive method of body composition analysis that would allow accurate assessment of body fat, body water, and lean mass. We used total body electrical conductivity (TOBEC) to estimate body composition in the little brown myotis (Myotis lucifugus). In a sample of 46 individuals, TOBEC was both accurate and precise in estimating total body water and lean mass but was less effective at estimating total body fat. Mass-independent or whole-body compartments (i.e., total body water, total body fat, lean dry mass, and lean body mass) were more accurately estimated than mass-dependent or mass-specific body composition compartments (i.e., water content, fat index, and % lean dry mass). The TOBEC measurements we made using an SA-3000 analyzer were influenced by extremes in body temperature, as well as by aluminum and incoloy wing bands. Our study also presents a new method of restraint especially suited for small mammals and birds that increases precision of TOBEC measurements. This study shows that TOBEC is a potentially valuable tool for studying changes in body composition of small mammals and may provide insight into the physiological impacts of various life history stages such as postnatal growth, reproduction, and hibernation.     

Estimation of human body composition by electrical impedance methods: a comparative study

This study 1) further validated the relationship between total body electrical conductivity (TOBEC) and densitometrically determined lean body mass (LBMd) and 2) compared with existing body composition techniques (densitometry, total body water, total body potassium, and anthropometry) two new electrical methods for the estimation of LBM: TOBEC, a uniform current induction method, and bioelectrical impedance analysis (BIA), a localized current injection method. In a sample of 75 male and female subjects ranging from 4.9 to 54.9% body fat the correlation between LBMd and LBM predicted from TOBEC by use of a previously developed regression equation was extremely strong (r = 0.962), thus confirming the validity of the TOBEC method. LBM predicted from BIA by use of prediction equations provided with the instrument also correlated with LBMd (r = 0.912) but overestimated LBM compared with LBMd in obese subjects. However, no such systematic error was apparent when new prediction equations derived from this heterogeneous sample of subjects were applied. Thus the TOBEC and BIA methods, which are based on the differing electrical properties of lean tissue and fat and which are convenient, rapid, and safe, correlate well with more cumbersome human body composition techniques.     

Non-invasive measure of body composition of snakes using dual-energy X-ray absorptiometry

Non-invasive techniques to measure body composition are critical for longitudinal studies of energetics and life histories and for investigating the link between body condition and physiology. Previous attempts to determine, non-invasively, the body composition of snakes have proven problematic. Therefore, we explored whether dual-energy X-ray absorptiometry (DXA) could be used to determine the body composition of snakes. We analyzed 20 adult diamondback water snakes (Nerodia rhombifer) with a DXA instrument and subsequently quantified their body composition by gravimetric and chemical extraction methods. Body composition components scaled with body mass with mass exponents between 0.88 and 1.53. DXA values for lean tissue mass, fat mass and total-body bone mineral mass were significantly correlated with observed masses of lean tissue, fat and ash from chemical analysis. Using regression models incorporating DXA values we predicted the fat-free tissue mass, lean tissue mass, fat mass, ash mass and total body water content for this sample of water snakes. A cross-validation procedure demonstrated that these models estimated fat-free tissue mass, lean tissue mass, fat mass, ash mass and total-body water content with respective errors of 2.2%, 2.3%, 16.0%, 6.6% and 3.5%. Compared to other non-invasive techniques, include body condition indices, total body electrical conductivity (TOBEC) and cyclopropane absorption, DXA can more easily and accurately be used to determine the body composition of snakes.     

Body composition in adolescents: estimation by total body electrical conductivity

This comparative study, conducted on 28 boys and girls of widely varying fatness, was designed to validate a new whole-body composition method [total body electrical conductivity (TOBEC)], based on bioelectrical properties of the human body. A significant correlation [r = 0.911; standard error of the estimate (SEE) = 5.3 kg] was demonstrated between the transformed TOBEC scores (TOBEC0.5 X Ht) and lean body mass (LBM) determined by hydrodensitometry and corrected for individual variations in hydration (LBMd + W). TOBEC determinations also correlated well with 1) total body water determined by deuterium oxide dilution (r = 0.877; SEE = 4.5 liters), 2) total body potassium determined by means of a 4 pi whole-body counter (r = 0.860; SEE = 430.7 meq), 3) LBM derived from skinfold thicknesses (r = 0.850; SEE = 5.8 kg). The residuals of the regression between LBMd + W and TOBEC scores did not show any significant correlation with either the potassium or the water content of the LBM. The results indicate that TOBEC is a simple, rapid, reliable, and noninvasive technique for delineating changes in body composition that occur in children during growth.     

Estimating lipid and lean body mass in small passerine birds using TOBEC,external morphology and subcutaneous fat‐scoring

To assess regression models for lipid and lean body mass in small birds, we recorded live body mass ±0.1 g, total body electrical conductivity (TOBEC; from “third generation” TOBEC machine EM‐SCAN® SA‐3000) or E‐Value, visual fat score (VisFat), and seven body measurements for 52 migratory passerine birds of 13 species (5–40 g). We determined lipid and lean mass of each bird after petroleum‐ether extraction of lipids. We obtained “net”E‐Value (NEV) for each scanned bird by subtracting the E‐Value of the empty bird‐restraining tube, because these showed an inverse temperature dependence (P<0.005). Leave‐one‐out cross validation was used to assess model selection and construct 95% confidence intervals. Although precision of TOBEC increased with bird size (CV of NEV vs. live mass: r=−0.276, P=0.002) and it explained an increasing proportion of variation in lean mass moving from small‐ to medium‐ to large‐bird classes of our data, it did no better than head length in single‐variable prediction of lean or lipid mass and was included in five of the 14 multivariate models we developed. The best multiple regression to predict lean mass included live weight, VisFat, bill length, tarsus and lnNEV (adjusted R²=99.0%); however, the same model lacking only lnNEV yielded aR²=98.9%. A parallel to the above pair of models, but predicting lipid mass, yielded aR²=90.3% and 90.0%, respectively. Subdividing the data by three size classes and three taxa (American redstart Setophaga ruticilla, ovenbird Seiurus aurocapilla, warblers), best‐subset multiple‐regression models predicted lean mass with aR² from 94.7 to 99.6% and lipid mass with aR² from 85.4 to 98.3%. Best models for the size‐ and species‐groups included VisFat and zero to five body measurements, and most included live weight. lnNEV was included only in the models for ovenbird (lipid), warblers (lipid), all birds (both), and large birds (both). Actual lipid mass of all birds was more highly correlated with multiple‐regression‐predicted lipid mass (r=0.955) than with visual subcutaneous fat‐scoring (r=0.683). These multiple‐regression models predicting lipid content using live‐bird measurements and visual fat score as independent variables represent more accurate and precise estimates of actual lipid content in small passerines than any previously published. They are particularly accurate for placing birds into percentage body‐fat classes.     

Noninvasive estimation of body composition in small mammals: a comparison of conductive and morphometric techniques

Body fat stores may serve as an index of condition in mammals. Thus, techniques that measure fat content accurately are important for assessing the ecological correlates of condition in mammal populations. We compared the ability of two conductive techniques, bioelectrical impedance analysis (BIA) and total body electrical conductivity (TOBEC), to predict body composition with that of morphometric methods in three small mammal species: red squirrels (n=13), snowshoe hares (n=30), and yellow-bellied marmots (n=4). Animals were livetrapped in northern Idaho; BIA (all subjects) and TOBEC (squirrels only) measurements were taken following chemical immobilization in the field, and morphometric measurements were taken postmortem. Information provided by BIA and TOBEC failed to improve upon the predictive power of morphometric equations for total body water (TBW) and lean body mass (LBM) in squirrels and hares, which do not store substantial amounts of fat (<5% body mass comprised of fat). Although the same pattern held with respect to LBM in marmots, which accumulate substantial amounts of body fat (>10% body mass), a BIA-based model proved best at estimating TBW, suggesting that the usefulness of conductive techniques may be a function of fat deposition. However, regardless of the technique used to predict body composition, estimates of body fat furnished by our equations failed to approximate actual fat levels accurately in all three test species, probably because these techniques only provide indirect estimates of fat content. These results highlight the limitations inherent in contemporary methods of animal fat estimation and underscore the need for the development of direct and accurate measures of body fat in mammals.     

Raising young reduces body condition and fat stores in black-legged kittiwakes

We conducted a manipulative experiment to investigate how raising chicks affects the body condition (body mass scaled by body size) and body composition (percent fat vs. lean mass) of black-legged kittiwakes (Rissa tridactyla). For 4 consecutive years (1991–1994) we removed eggs from randomly selected nests and then compared adults raising chicks with adults that had their eggs removed. At the end of the chick-rearing period, adults raising chicks were significantly lighter for their size than adults that had their eggs removed. Adults raising chicks also had a significantly lower percent body fat (by 28%) than adults from manipulated nests. The difference in percent body fat between the two groups was apparent at all levels of condition, suggesting that adults that are raising chicks apportion their reserves differently than adults that are working only to meet their own metabolic needs. End-of-season body condition of adults from manipulated and unmanipulated nests varied significantly among 5 years of study, and appeared to reflect differences in local foraging conditions. In all years, females were in worse condition than males at the end of the breeding season. This sex-specific condition difference did not, however, appear to indicate a greater short-term reproductive cost among females. Females were lighter for their size than males in both the manipulated and unmanipulated groups. Our results suggest that adult kittiwakes compromise their body condition and body composition during chick rearing to increase the likelihood of successfully fledging young, even though such adjustments may decrease their own post-reproductive survival probabilities. Prior to estimating the body composition of the experimental birds, we evaluated the usefulness of several noninvasive techniques for predicting fat mass in kittiwakes. We used cross-validation techniques to compare multiple regression models that included total body electrical conductivity (TOBEC), total body water (TBW), and morphometric measurements as independent variables. The most parsimonious model for predicting fat mass was based on TOBEC and mass measurements. TBW and morphometrics were of little utility in predicting fat mass in kittiwakes. Previous studies that have evaluated the usefulness of TOBEC as a predictor of fat mass have shown mixed results. We suggest that the size of the experimental subject relative to the size of the TOBEC measurement chamber may affect the accuracy of this technique. Received: 30 November 1998 / Accepted: 29 April 1999     

Intraspecific variation in reproductive physiology and egg quality in the European Starling Sturnus vulgaris

Egg mass shows large intraspecific variation in birds and is repeatable within individuals. The mechanisms underlying this variation are unknown. We hypothesized that measures of egg quality (the mass of yolk protein, yolk lipid, and albumen protein) would be positively correlated with the plasma pools of the yolk precursor vitellogenin, and the masses of the oviduct, metabolic machinery (liver, heart, lungs, kidneys, gizzard, small intestine and pancreas), and endogenous stores of protein and lipid. We tested these predictions in European Starlings Sturnus vulgaris collected at the peak of egg production effort. In contrast to our predictions, both yolk protein and yolk lipid were negatively correlated with plasma vitellogenin levels. Albumen protein was positively related to oviduct mass, but other aspects of body composition failed to explain variation in egg quality. Hence, while we observed correlations between egg composition and peripheral systems (circulating precursor pools and the oviduct), we found no evidence that egg quality is determined by more general processes, i.e., the supply and processing of nutrients.     

Relationship between measures of body size and composition and velocity of lactate threshold

The purpose of this investigation was to explore the relationship between velocity of lactate threshold (vLT) and various measures of body mass and composition: mass, lean mass, fat mass, percent body fat (% fat), and body surface area (BSA). We hypothesized that mass would be inversely related to vLT, and that differences in measures of body mass and composition would account for a significant amount of variability in vLT. A total of 21 healthy male runners served as subjects. Body composition was assessed by hydrostatic weighing. A significant negative relationship (r = -0.759, p < or = 0.01) was between body mass and vLT. The coefficient of determination between vLT and body mass indicated that nearly 58% of the variability in vLT was explained by body mass in these subjects. Significant relationships were also between vLT and BSA (r = -0.72, p < or = 0.01), fat mass (r = -0.70, p < or = 0.01), % fat (r = -0.59, p < or = 0.01), and lean mass (r = -0.41, p < or = 0.05). Linear regression yielded the following model: y = 369.48 - 1.7343 (X), where y = predicted vLT (m.min(-1)) and X = body mass (kilograms) (SEE = 15.45). Velocity of lactate threshold was significantly inversely related with body mass in a group of male runners. The calculated coefficient of determination suggests that nearly 58% of the variability in vLT was explained by body mass. The present data suggest consideration of categorizing participants in 'road runs' by body mass to equate competition, as is done in other sports (e.g., weightlifting).     

Measurement of arthropod body composition using quantitative magnetic resonance

Quantitative magnetic resonance (QMR) is a new technology for measuring the body composition (wet lean mass, fat mass, and total body water mass) of unrestrained and unanesthetized animals. We conducted a validation study using two species of crayfish (mass range 5.5–27 g), American lobsters (680–732 g), and Madagascar hissing cockroaches (6.5–14 g) to assess the utility of QMR for quantifying the body composition of crustaceans and other large arthropods. A comparison of crayfish, lobster, and cockroach wet lean, fat, and body water masses calculated by QMR with those obtained from the traditional chemical extraction method demonstrates that QMR is a valid technology for analysis of wet lean mass and body water. Fat mass could not be accurately predicted, although this might be improved with the use of a QMR analyzer designed specifically for animals of low fat content. QMR analysis allows rapid (<4 min) and non‐destructive determination of body composition in field and lab environments, enabling researchers to conduct longitudinal studies and to increase the ethicality and practicality of studying rare or threatened species.     

A replicated single generation test of a restricted selections index in poultry

Summary Predicted and realized responses in a single generation of mass selection for an index and for its component traits were compared. The index included the log transformed traits determining egg mass in chickens to 40 weeks of age (days tested from sexual maturity, egg weight, rate of lay). The index was restricted to allow no increase in log days tested. Other traits measured were egg mass, age at first egg, egg weight, rate of lay, number of eggs and body weight. When averaged over replicates, realized and predicted responses were in close agreement for index values and for the component traits. Significant corresponding correlated responses were obtained for egg mass and weight. The restricted trait, log days tested, and the correlated trait age at first egg did not change. Egg mass was increased solely through change in egg weight.Journal paper no. 6917. Purdue University Agricultural Experiment StationThis investigation was conducted as a part of the cooperative research of the NC-89 Regional Poultry Breeding Project entitled Nature and Utilization of Genetic Variation Influencing Economic Traits in Poultry     

Fatty acid content of eggs determines antipredator performance of fish larvae

Recent work has suggested that provisioning of eggs with certain critical nutrients could be a more meaningful measure of maternal investment and correlate of offspring fitness than traditional measures of egg size. The aim of our study was to assess variability in egg quality and larva quality and to identify connections between them and the implications for larval survival. Egg size, proximate composition, and fatty acid composition were measured for 40 batches of eggs from 8 captive pairs of red drum (Osteichthyes: Sciaenops ocellatus). We reared larvae from these batches of eggs to a common size (10 mm total length, 2-3 weeks posthatching) and assessed routine activity and escape response performance of 671 individuals. Egg fatty acid composition varied more than egg size or proximate composition. Concentrations of certain long chain, highly unsaturated essential fatty acids (e.g., arachidonic acid and docosahexaenoic acid) were the only egg traits that were significantly related to larva quality (measured as escape performance). Reduced escape performance of larvae from eggs with low fatty acid concentrations was not compensated by 3 weeks of feeding on a diet enriched with fatty acids, suggesting irreversible developmental effects. Since fatty acids in eggs originate from the maternal diet, offspring survival may be determined in part by availability of nutrient-rich prey to pre-spawning adults. Migrations, regime shifts, and exploitation of marine communities could operate through this mechanism to influence recruitment in fish populations. Our findings underscore the importance of non-genetic maternal contributions to egg quality and the linkage between environmental conditions experienced by adult females and offspring fitness.     

Intraclutch egg-size variation in the Barnacle Goose Branta leucopsis: an egg-removal experiment

We carried out an egg-removal experiment on Barnacle Geese Branta leucopsis to test the hypothesis that tactile contact between the brood patch and eggs accumulating in the nest during laying was causally related to intraclutch egg-size variation, acting through the development of incubation behaviour. Egg removal had no effect on mean egg mass, relaying interval or clutch size: mean (±s.e.) clutch sizes for first clutches were 4.9 ± 0.9 and 4.7 ± 0.9 eggs for control and experimental females, respectively. We therefore conclude that Barnacle Geese have a determinate pattern of egg laying. Daily removal of all eggs from the first-laid egg did not affect the within-clutch pattern of egg-size variation: in both control and experimental clutches the first egg was relatively small, the second or third egg was largest and there was a linear decline in egg size to the smallest, last-laid egg. There was no significant difference in either the absolute or relative size of the last-laid eggs in control and experimental clutches. Tactile contact with eggs in the nest, therefore, is not required for the expression of intraclutch egg-size variation in Barnacle Geese. In this respect, determinate-laying Barnacle Geese differ from species with indeterminate laying patterns (e.g. gulls) where egg removal does affect the normal pattern of within-clutch egg-size variation. If hormonal changes (e.g. prolactin) associated with onset of incubation are causally related to the intraclutch decrease in egg size, then the stimuli involved appear to differ between determinate and indeterminate layers.     

Diapause duration, survival in relation to desiccation and egg-pod morphology of the Senegalese grasshopper, Oedaleus senegalensis

Abstract. The Senegalese grasshopper, Oedaleus senegalensis (Krauss) (Orthoptera: Acrididae), is periodically a devastating pest of subsistence crops in the sahelian zone of West Africa.Egg diapause has been suggested as an important component of the mechanism acting to generate outbreaks and therefore diapause duration, egg survival in relation to desiccation and egg-pod morphology were investigated in this study.
Three months after oviposition, 50% of initially diapausing egg pods maintained in the laboratory exhibited 'partial emergence' and produced hoppers.No difference in diapause duration was evident between egg pods maintained at a constant 30 ± 1°C and those maintained under alternating temperature conditions of 23 ± 1°C and 35 ± 1°C.Laboratory maintained eggs showed significant variation in diapause duration with diapause lasting between 2 months and 4 years.Diapause duration was shorter in the field with only c. 1.4% of eggs remaining in diapause after a single dry season (6–7 months).Survival of eggs in relation to desiccation was high in the field and low in the laboratory, and field eggs lost only c. 20% of their body weight over the dry season.Survival of diapausing and non-diapausing egg pods in the laboratory was similar, indicating that diapause in this species is primarily a mechanism for preventing emergence at an inappropriate time of year, rather than conferring any additional resistance to desiccation.Egg pods oviposited under diapause-inducing conditions (LD 14:10 h, 25°C) were significantly shorter than those laid under non-diapausing conditions (LD 10:14 h, 40°C).These results are discussed both in relation to possible O.senegalensis 'bet-hedging' strategies and to the probable mechanism operating behind major outbreaks of this species.     

Egg quality and parental ornamentation in the blue tit Parus caeruleus

Components of avian egg quality often show remarkable variation between females and yet the causes of variation in this fitness-related trait remain poorly understood. We investigated the egg size and yolk carotenoid investment of blue tit Parus caeruleus females in relation to multiple parental traits. We show that females produce eggs with more colourful yolks when mated to ultraviolet (UV) attractive males. Since yolk carotenoids may enhance offspring fitness, but may be available to females in limiting amounts, the correlation we found suggests adaptive egg quality adjustment in response to mate attractiveness, but an experimental approach is required to rule out alternative explanations. Yolk colour was also correlated with laying date and yolk mass. Though the effect of laying date could not be explained by ambient temperature during egg formation, it suggests a proximate constraint of general carotenoid availability on yolk composition. Egg size was not affected by male attractiveness; however, females with a brighter crown produced larger eggs, suggesting that crown coloration might indicate individual quality in females too.     

A population-based twin study on sleep duration and body composition

The aim of this study is to investigate the relationship between sleep duration and body composition and to estimate the genetic contribution of sleep duration and body composition in a Chinese twin population. This cross-sectional analysis included 738 men and 511 women aged 21-72 year. Anthropometric and dual-energy X-ray absorptiometry (DXA) measures of body composition were used. Sleep duration was obtained from a standard sleep questionnaire. Multiple regression models were used to examine the association between sleep duration and body composition measures. Structural equation modeling was used to assess the heritability of sleep duration and body composition. Compared with individuals in the 2nd and 3rd age-specific quartiles of sleep duration (reference group), shorter (1st quartile) sleep duration among women but not men was associated with higher z-scores (0.248-0.317) for all adiposity measures--BMI, fat mass index (FMI), percent body fat mass (%BF), and percent trunk fat mass (%TF), P < 0.05 for each--and with 0.306 lower z-scores for percent body lean mass (%LM) and 0.353 lower lean/fat mass ratio (LFR), P < 0.01 for each. The heritability of sleep duration was 0.27 in men and 0.29 in women, while the heritability of body composition was as high as 0.56-0.73 after adjustment for age in both genders. Short sleep duration was associated with increased body fat and decreased lean body mass in women but not in men. Sleep duration was largely influenced by environmental factors while adiposity measures were mainly influenced by genetic factors.     

Reproductive success and the energetic cost of parental care in male smallmouth bass

Direct field measurements of the energetic expenditure on parental care and within-nest reproductive success of individual male smallmouth bass Micropterus dolomieui were determined by measuring the change in total body mass as well as by total body electroconductivity analysis (TOBEC™). With TOBEC, the change in total body lean mass of the same live individual was measured non-destructively at the beginning and end of the parental care period. Lean mass was the primary source of energy utilized during parental care indicating starvation and potential loss of future reproduction. Individual loss in lean mass was related positively to reproductive success suggesting that the energy expended during parental care does affect individual fitness.     

Egg size, egg composition and reproductive success in the Oystercatcher Haematopus ostralegus

We investigated the relationship between egg size and composition and their subsequent effects on hatching and fledging success in Eurasian Oystercatchers Haematopus ostralegus on the island of Schiermonnikoog (53̀30'N, 06̀10'E) in the Dutch Wadden Sea between 1986 and 1990. Egg size variation was considerable. The ratio in egg volume of the largest over the smallest egg was more than 1.5 in each of five years. Differences between females accounted for, on average, 61% of the total variance of egg volume. Individual females produced similarly sized eggs from one year to another. Nevertheless, average egg volume per clutch declined by 0.2 cm³ per year as females aged, but this explains little of the observed variation. Female size and food supplementation had no significant effect on egg volume. In absolute terms, large eggs contain more lean dry matter and lipid than small eggs, but the proportion of both constituents decreases with egg size. Consequently, the combustible energy content of eggs increases less than proportionally with egg size. Hatchability averaged 87% and was not correlated with egg volume. Hatchling weight increased with egg size, averaging 65% of fresh egg weight. Chick survival until fledging did not increase with egg volume. Consequently, within the size range observed, large and small eggs are of comparable quality. Since both the costs and benefits of large eggs compared with small eggs seem small at best, we propose that, within the size range encountered in this population, egg size can be considered an evolutionarily neutral trait.     

山地麻蜥的雌性繁殖和孵出幼体特征

山地麻蜥年产多窝卵。窝卵数、窝卵重和卵重与雌体体长呈正相关,卵重与窝卵数无关。窝卵数、窝卵重和卵重不存在明显的季节变化。卵长径与卵短径呈正相关,卵长径、卵短径与窝卵数无关。雌体通过增加窝卵数和卵大小增加繁殖输出。孵出幼体的体重、体长、尾长、躯干干重、刺余卵黄不存在季节差异,但5月份孵出幼体的脂肪体高于4月份。     

Experimental manipulation of food availability leads to short‐term intra‐clutch adjustment in egg mass but not in yolk androgen or thyroid hormones

In birds, mothers can affect their offspring's phenotype and thereby survival via egg composition. It is not well known to what extent and time‐scales environmental variation in resource availability, either via resource constrains or adaptive adjustment to predicted rearing conditions, influences maternal effects. We experimentally studied whether egg and yolk mass and yolk hormone levels respond to short‐term changes in food availability during laying in wild great tits Parus major. Our treatment groups were: 1) food supplementation (mealworms) from the 1st until the last egg; 2) food supplementation from the 1st until the 5th egg, where the effect of cessation of the supplementary food treatment could also be studied; 3) no food supplementation (controls). We analysed both nutritional resources (egg, yolk and albumen mass), and the important developmental signals, yolk androgens (testosterone and androstenedione), and for the first time in a wild population, yolk thyroid hormones (thyroxine and 3,5,3′‐triiodothyronine). Egg mass is a costly resource for females, androgens most likely non‐costly signals, whereas thyroid hormones may be costly signals, requiring environmental iodine. In the food supplemented group egg, yolk and albumen mass increased rapidly relative to controls and when food supplementation was halted, egg and albumen mass decreased, indicating rapid responses to resource availability. Yolk androgen and thyroid hormone levels were not affected by food supplementation during laying. Thyroxine showed an increase over the laying sequence and its biological meaning needs further study. The rapid changes in egg mass to variation in within‐clutch food availability suggest energetic/protein/nutrient constrains on egg formation. The lack of a response in yolk hormones suggest that perhaps in this species the short‐term changes in resource availability during egg laying do not predict offspring rearing conditions, or (for thyroid hormones) do not cause systemic changes in circulating hormones, and hence do not affect maternal signaling.