Size Matters: Individual Variation in Ectotherm Growth and Asymptotic Size

Body size, and, by extension, growth has impacts on physiology, survival, attainment of sexual maturity, fecundity, generation time, and population dynamics, especially in ectotherm animals that often exhibit extensive growth following attainment of sexual maturity. Frequently, growth is analyzed at the population level, providing useful population mean growth parameters but ignoring individual variation that is also of ecological and evolutionary significance. Our long-term study of Lake Erie Watersnakes, Nerodia sipedon insularum, provides data sufficient for a detailed analysis of population and individual growth. We describe population mean growth separately for males and females based on size of known age individuals (847 captures of 769 males, 748 captures of 684 females) and annual growth increments of individuals of unknown age (1,152 males, 730 females). We characterize individual variation in asymptotic size based on repeated measurements of 69 males and 71 females that were each captured in five to nine different years. The most striking result of our analyses is that asymptotic size varies dramatically among individuals, ranging from 631–820 mm snout-vent length in males and from 835–1125 mm in females. Because female fecundity increases with increasing body size, we explore the impact of individual variation in asymptotic size on lifetime reproductive success using a range of realistic estimates of annual survival. When all females commence reproduction at the same age, lifetime reproductive success is greatest for females with greater asymptotic size regardless of annual survival. But when reproduction is delayed in females with greater asymptotic size, lifetime reproductive success is greatest for females with lower asymptotic size when annual survival is low. Possible causes of individual variation in asymptotic size, including individual- and cohort-specific variation in size at birth and early growth, warrant further investigation.     

The influence of genetic and environmental factors in estimations of current body size, desired body size, and body dissatisfaction.

The objective was to investigate the genetic epidemiology of figural stimuli. Standard figural stimuli were available from 5,325 complete twin pairs: 1,751 (32.9%) were monozygotic females, 1,068 (20.1%) were dizygotic females, 752 (14.1%) were monozygotic males, 495 (9.3%) were dizygotic males, and 1,259 (23.6%) were dizygotic male-female pairs. Univariate twin analyses were used to examine the influences on the individual variation in current body size and ideal body size. These data were analysed separately for men and women in each of five age groups. A factorial analysis of variance, with polychoric correlations between twin pairs as the dependent variable, and age, sex, zygosity, and the three interaction terms (age x sex, age x zygosity, sex x zygosity) as independent variables, was used to examine trends across the whole data set. Results showed genetic influences had the largest impact on the individual variation in current body size measures, whereas non-shared environmental influences were associated with the majority of individual variation in ideal body size. There was a significant main effect of zygosity (heritability) in predicting polychoric correlations for current body size and body dissatisfaction. There was a significant main effect of gender and zygosity in predicting ideal body size, with a gender x zygosity interaction. In common with BMI, heritability is important in influencing the estimation of current body size. Selection of desired body size for both men and women is more strongly influenced by environmental factors.     

The gender-specific apolipoprotein E genotype influence on the distribution of plasma lipids and apolipoproteins in the population of Rochester, Minnesota. II. Regression relationships with concomitants.

The influence of the apolipoprotein E (Apo E) polymorphism and gender on the regression relationships between each of nine plasma lipid and apolipoprotein traits (total cholesterol; ln triglycerides; high-density-lipoprotein cholesterol; apolipoproteins AI, AII, B, and CII; ln CIII; and ln E) and four concomitants (age, weight, waist-to-hip ratio, and smoking) was studied in 507 unrelated individuals representative of the adult population of Rochester, MN. Analyses are presented separately for females and males. Each lipid and apolipoprotein trait exhibited at least one Apo E genotype-specific regression relationship with the concomitants investigated in this study. In most cases the heterogeneity of regression was associated with differences between the epsilon 32 and epsilon 33 genotype. This study documents that the influence of Apo E genotype on average levels of plasma lipids and apolipoproteins varies among subdivisions of the population defined by age, body size, and smoking status.     

Aging changes in the medullary cavity of the proximal femur in American Blacks and Whites.

This study explores changes with age in the medullary cavity in an area of the femur which is becoming of increasing clinical interest. It is based on measurements of medial-lateral and posterior-anterior radiographs of the proximal one-third of the right femur, below the greater trochanter. The study population consisted of 257 individuals from the Terry collection in the Smithsonian Institution and included Blacks and Whites, males and females, aged 20 to 90 years. Medullary cavity and external diameters were measured at 11 levels, beginning with a line drawn perpendicular to the linea aspera at the level of the inferior border of the greater trochanter. To correct for size differences, the interval between levels was set at one-fourth the individual femoral head diameter. The data were subjected to regression analysis and analysis of variance. Anterior-posterior medullary diameter is more highly correlated with age than medial-lateral diameter. Females show the greatest relative increase with age in both dimensions, especially at subtrochanteric levels, and they tend to have relatively larger medullary cavities than males at all ages. White males have relatively larger medullary diameters than Black males. Younger White females have relatively larger medullary cavities than Black females, but the latter reach parity with White females in middle age and show a greater overall percentage of medullary expansion.     

Genotype-Environment Interaction: Apolipoprotein E (Apoe) Gene Effects and Age as an Index of Time and Spatial Context in the Human

We analyzed the age-dependence of the estimates of the parameters of the genetic architecture of plasma ApoE levels associated with ApoE gene variation. Our study sample included 1988 individuals in multigeneration pedigrees from the Rochester, MN, population. We used a 30-yr sliding window across the age range (5-90 yr) to estimate the age dependency of parameters. Additive ApoE allelic variance of transformed plasma ApoE values for both genders, heritabilities for males and phenotypic and residual variance for females peaked in the 20-40-yr age windows and decreased significantly with age (P < 0.05). Phenotypic and residual variance for males and dominance variance for both genders did not vary significantly with age. All parameter estimates were significantly different from zero across all age windows for both genders. Most studies of ApoE have focused on its functions in the pathophysiology of coronary artery disease (CAD) in middle-aged and older individuals. Our findings suggest the greatest role of this gene is in determining phenotypic differences among younger and middle-aged individuals. These observed genotypic effects on the plasma ApoE levels may contribute to age-dependent differences in physiological health, growth, and risk of disease.     

Analysis of genetic and environmental sources of variation in serum cholesterol in Tecumseh,Michigan. III. Identification of genetic effects using 12 polymorphic genetic blood marker systems.

Four of 12 unlinked polymorphic marker systems were identified as predictors of normal serum cholesterol levels. Consistent effects between males and females and with other studies suggest that these marker loci are themselves involved in cholesterol determination or are closely linked to the involved loci. Two-locus combinations suggest that an 8-9 mg/100 ml difference in nonfasting serum cholesterol may be predicted between phenotypic classes which are not rare in frequency.     

The evolution of genetic architecture. I. Diversification of genetic backgrounds by genetic drift

The genetic architecture of a phenotype plays a critical role in determining phenotypic evolution through its effects on patterns of genetic variation. Genetic architecture is often considered to be constant in evolutionary quantitative genetic models. However, genetic architecture may be variable and itself evolve when there are dominance and epistatic interactions among alleles at the same and different loci, respectively. The evolution of genetic architecture by genetic drift is examined here by testing the breeding value of four standard inbred mouse strains mated across a set of 26 related recombinant quasi-inbred (RqI) lines generated from the intercross of the Large (LG/J) and Small (SM/J) inbred mouse strains. Phenotypes of interest include age-specific body weights, growth, and adult body composition. If the genetic architecture of these traits has differentiated by genetic drift during the production of the RqI strains, we should observe interactions between tester strain and RqI strain. The breeding values of the tester strains will change relative to one another depending on which RqI strain they are crossed to. The study included an average of 15.1 offspring per cross, over a total of 100 different crosses. Multivariate and univariate analyses of variance indicate that there is strongly significant interaction for all traits. Interaction is more pronounced in males than in females and accounted for an average of about 40% of the explained variation in males and 30% in females. These results indicate that the genetic architecture of these traits has differentiated by genetic drift in the RqI strains since their isolation from a common founder population. Further analysis indicates that this differentiation results in changes in the order of tester strain effects so that common patterns of selection in these differentiated populations could result in the fixation of different alleles.     

Otitis media: genetic factors and sex differences.

Although genetic factors are recognised as major contributors to otitis media, the presence of sex differences in heritability needs clarification. The aim of this study was to estimate the relative contribution of genetic and environmental effects in otitis media liability with particular focus on sex differences. Data from a cohort of Norwegian twins born between 1967 and 1979 with repeated measures on recurrent childhood otitis media were analysed. Altogether the sample included 4247 twin pairs. The tetrachoric correlations for monozygotic twins were .71 and .65 for males and females respectively. In dizygotic twins the correlations were .35 and .25 for males and females, respectively, and was.34 in opposite sexed pairs. The contribution of genetic and environmental effects was analyzed using structural equation modeling. The best fitting model showed that additive genetic effects explained 72% and 61% of the variance in males and females, respectively. The remaining variance was attributed to individual environmental effects. A model specifying equal heritability estimates for males and females yielded an almost equivalent fit. We found substantial genetic effects for liability to otitis media. There is no evidence that different sets of genes influence liability in males and females, but there may be sex differences in the relative importance of genetic effects.     

A longitudinal genetic study of vocabulary knowledge in adults.

Vocabulary test scores were obtained from a total of 997 adults, all twins or a sibling of twins in this study. Some (N = 217) individuals were tested twice, around 6 years apart. Heritability varied from 50% at the first test occasion to 63% at the second test occasion. The correlation of scores across time was.74. Structural equation modelling showed that stability in vocabulary knowledge over time can largely (around 76%) be explained by genetic factors. Part of the non-shared environmental variance was stable over time also. Any influence from shared environmental factors could not be detected. Results were similar for the two sexes, except that males generally outperformed females. Results were also similar for two age cohorts, except that the older cohort generally outperformed the younger cohort.     

Conspecific reproductive success affects age of recruitment in a great cormorant, Phalacrocorax carbo sinensis, colony.

Few studies have addressed the proximate factors affecting the age at which individuals of long-lived bird species are recruited into the breeding population. We use capture-recapture analysis of resightings of 16 birth cohorts of colour-ringed great cormorants, Phalacrocorax carbo sinensis, in a Danish colony to assess the evidence for two hypotheses: conspecific attraction (earlier recruitment when the colony is large) and conspecific reproductive success (earlier recruitment following years of high breeding success). For both males and females, conspecific reproductive success was the most important covariate explaining the interannual variation in age of recruitment; colony size was also important for females. These covariates explained nearly 60% of the year-to-year variation for both sexes. The age of recruitment increased for cohorts born after 1990, and this increase was correlated with a decline in breeding success in the colony; we interpret this as an indirect and delayed density-dependent effect. Females were recruited earlier than males (mean age of recruitment for cohorts born before 1990: 2.98 years versus 3.53 years); the most plausible reason for this is a skewed sex ratio in favour of males in the adult population. Recruitment of males may thus, to some extent, be constrained by the availability of females. This study provides the first evidence that conspecific reproductive success can affect the age at which individual birds start to breed.     

The Roles of Sex,Mass and Individual Specialisation in Partitioning Foraging-Depth Niches of a Pursuit-Diving Predator

Intra-specific foraging niche partitioning can arise due to gender differences or individual specialisation in behaviour or prey selection. These may in turn be related to sexual size dimorphism or individual variation in body size through allometry. These variables are often inter-related and challenging to separate statistically. We present a case study in which the effects of sex, body mass and individual specialisation on the dive depths of the South Georgia shag on Bird Island, South Georgia are investigated simultaneously using a linear mixed model. The nested random effects of trip within individual explained a highly significant amount of the variance. The effects of sex and body mass were both significant independently but could not be separated statistically owing to them being strongly interrelated. Variance components analysis revealed that 45.5% of the variation occurred among individuals, 22.6% among trips and 31.8% among Dives, while R² approximations showed gender explained 31.4% and body mass 55.9% of the variation among individuals. Male dive depths were more variable than those of females at the levels of individual, trip and dive. The effect of body mass on individual dive depths was only marginally significant within sexes. The percentage of individual variation in dive depths explained by mass was trivial in males (0.8%) but substantial in females (24.1%), suggesting that differences in dive depths among males was largely due to them adopting different behavioural strategies whereas in females allometry played an additional role. Niche partitioning in the study population therefore appears to be achieved through the interactive effects of individual specialisation and gender upon vertical foraging patch selection, and has the potential to interact in complex ways with other axes of the niche hypervolume such as foraging locations, timing of foraging and diet.     

Genetic determination of plasma apolipoprotein AI in a population-based sample.

Apolipoprotein AI (apo AI) is the major protein of high-density lipoprotein (HDL). Using radioimmunoassay, we measured plasma apo AI levels in 1,880 individuals in 283 pedigrees randomly selected from the population with respect to disease status and risk factors for coronary artery disease. Apo AI levels were first adjusted for date of assay (6.8% of apo AI variation) and then adjusted for variability in age and body mass index (an additional 6.6%, 20.4%, and 23.0% of apo AI variations for males, females not using exogenous hormones, and females using exogenous hormones, respectively). A mixture of two normal distributions fit the adjusted data better than did a single normal distribution. Genetic and environmental models that could explain the mixture of normal distributions were investigated using complex segregation analysis. Heterogeneous etiologies for individual differences in adjusted apo AI levels were suggested by the data in the 283 pedigrees. In a subset of 126 pedigrees, there is evidence for the major effect of a nontransmitted environmental factor that explains the mixture of distributions as well as polygenic loci that influence apo AI levels within each distribution. The environmental factor and polygenic loci account for 32% and 65% of the adjusted variation, respectively. In the other 157 pedigrees there is strong support for a single locus with a major effect that accounts for 27% of the adjusted variation. The effect of the polygenic loci is not different from zero in these 157 pedigrees. This is the first study to present evidence for the segregation of a single unmeasured locus with a major effect on levels of apo AI in a population-based sample of pedigrees.     

Ontogenetic variation in the plumage colour of female European Pied Flycatchers Ficedula hypoleuca

Although variation in the dorsal plumage colour of male European Pied Flycatchers Ficedula hypoleuca has received a great deal of attention, females of the species have been usually considered to be nearly uniformly monochromatic brown. Using reflectance spectrophotometry, we explore the age‐dependent variation of plumage colour in females. We disentangle the within‐ and between‐individual effects of this pattern and show a within‐individual darkening of the mantle colour with age, whereas differences between individuals in structural colour expression may underlie the trend for a more reflective white in the females' breast plumage with advancing age. The darkening of the dorsal plumage as females age reflects the most common pattern of age‐related variation in males in most European populations of the species.     

Lifetime individual plasticity in body condition of a migratory bird

Body condition is an important trait that reflects the capacity of individuals to acquire food or resist disease, eventually allowing successful reproduction and survival. We first quantified the effects of condition on life history and a secondary sexual trait using long‐term lifetime data on condition in a migratory bird species, the barn swallow, Hirundo rustica. Second, we quantified sources of individual variation in condition by separating the within‐individual from the between‐individual components of variation of environmental condition and age on body condition. Therefore, we were able to partition variation as a result of selection and phenotypic plasticity. Within‐individual variation in body condition increased in early life until middle age (i.e. 3–4 years of age) in the two sexes followed by only a slight decrease in body condition during senescence in males but not in females. After accounting for age‐dependent variation, condition could be partitioned into a within‐individual plastic response to environmental conditions during migration and a nonplastic response (i.e. a between‐individual difference) to environmental conditions experienced in the African winter quarters. Specifically, there was a within‐individual increase in body condition as environmental conditions during migration improved in both males and females, independent of age. There was a between‐individual effect of condition found in the winter quarter in body condition of females, but not in males, which was attributed to the disappearance of females in poor body condition from the study population as a result of the higher natal dispersal of low‐quality females compared to high‐quality ones during years with favourable environmental conditions in the African winter quarters. Males and females also tended to be in better body condition during the warmer springs upon arrival at the breeding grounds. There was a temporal decline in female body condition during 1991–2007, whereas no significant trend was detected in males. Therefore, both intrinsic (e.g. age and sex) and extrinsic factors (e.g. climate) affected body condition. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 420–434.     

Morphological variation in adult chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d'Ivoire

Twenty five adult chimpanzee skeletons (Pan troglodytes verus) of known age and sex (15 females, 10 males) from a long‐term study site in Taï National Park, Cote d'Ivoire present new data on variation. These skeletons provide a rare opportunity to measure the cranium and postcranium from the same individuals. We compare measurements and indices of the Taï sample with those of relatively complete Pan troglodytes schweinfurthii skeletons from Gombe National Park, Tanzania. Measurements of Pan paniscus are included as an outside comparison. The Taï and Gombe samples are analyzed by sex; combined sex samples are compared between the two groups, and the two sexes to each other. Taï females and males do not differ in most long bone lengths or in pelvic dimensions, but do differ significantly in cranial capacity, facial measurements, clavicle length, scapular breadth, and femur length. Gombe females and males differ significantly in some facial measurements and in scapular breadth. In combined sex samples, Taï individuals have lower cranial capacity, longer palate and mandible, and greater dimensions in the trunk and limb lengths. Taï females account for most of the variation; males differ from each other only in greater length of humerus and femur. The Taï skeletons provide new data for assessing individual variation and sexual dimorphism within and between populations and species. The combination of cranial and postcranial data provides a clearer picture of chimpanzee intraspecific and interspecific variation than can be gained from either data set alone. Am J Phys Anthropol, 2008. © 2007 Wiley‐Liss, Inc.     

The phenotypic and genetic covariance structure of drosphilid wings

Evolutionary constraint results from the interaction between the distribution of available genetic variation and the position of selective optima. The availability of genetic variance in multitrait systems, as described by the additive genetic variance-covariance matrix (G), has been the subject of recent attempts to assess the prevalence of genetic constraints. However, evolutionary constraints have not yet been considered from the perspective of the phenotypes available to multivariate selection, and whether genetic variance is present in all phenotypes potentially under selection. Determining the rank of the phenotypic variance-covariance matrix (P) to characterize the phenotypes available to selection, and contrasting it with the rank of G, may provide a general approach to determining the prevalence of genetic constraints. In a study of a laboratory population of Drosophila bunnanda from northern Australia we applied factor-analytic modeling to repeated measures of individual wing phenotypes to determine the dimensionality of the phenotypic space described by P. The phenotypic space spanned by the 10 wing traits had 10 statistically supported dimensions. In contrast, factor-analytic modeling of G estimated for the same 10 traits from a paternal half-sibling breeding design suggested G had fewer dimensions than traits. Statistical support was found for only five and two genetic dimensions, describing a total of 99% and 72% of genetic variance in wing morphology in females and males, respectively. The observed mismatch in dimensionality between P and G suggests that although selection might act to shift the intragenerational population mean toward any trait combination, evolution may be restricted to fewer dimensions.     

Primate Innovation: Sex,Age and Social Rank Differences

Analysis of an exhaustive survey of primate behavior collated from the published literature revealed significant variation in rates of innovation among individuals of different sex, age and social rank. We searched approximately 1,000 articles in four primatology journals, together with other relevant databases, for examples of innovation. The reported incidence of innovation is higher in males and adults, and lower in females and nonadults, than would be expected by chance given the estimated relative proportions of these groups. Amongst chimpanzees, the only species for which there are sufficient data to consider alone, there is a similar sex difference in the propensity to innovate, but no effect of age. Chimpanzees of low social rank are reported as innovators more frequently than high-ranking chimpanzees are. Male chimpanzees innovate more often than females in sexual, courtship, mating and display contexts; that is, in contexts likely to increase access to mates. The largest number of recorded observations are in the foraging context, wherein contrary to expectations, there is no evidence for female chimpanzees exhibiting more innovation than males. The study is the first extensive investigation of behavioral innovation in primates and provides evidence that much individual variation in the propensity to innovate can be explained in terms of sex, age, and social rank.     

Effect of mass selection on the within-family genetic variance in finite populations

Summary The adequacy of an expression for the withinfamily genetic variance under pure random drift in an additive infinitesimal model was tested via simulation in populations undergoing mass selection. Two hundred or one thousand unlinked loci with two alleles at initial frequencies of 1/2 were considered. The size of the population was 100 (50 males and 50 females). Full-sib matings were carried out for 15 generations with only one male and one female chosen as parents each generation, either randomly or on an individual phenotypic value. In the unselected population, results obtained from 200 replicates were in agreement with predictions. With mass selection, within-family genetic variance was overpredicted by theory from the 12th and 4th generations for the 1,000 and 200 loci cases, respectively. Taking into account the observed change in gene frequencies in the algorithm led to a much better agreement with observed values. Results for the distribution of gene frequencies and the withinlocus genetic covariance are presented. It is concluded that the expression for the within-family genetic variance derived for pure random drift holds well for mass selection within the limits of an additive infinitesimal model.     

Partitioning of genetic variation across the genome using multimarker methods in a wild bird population

The underlying basis of genetic variation in quantitative traits, in terms of the number of causal variants and the size of their effects, is largely unknown in natural populations. The expectation is that complex quantitative trait variation is attributable to many, possibly interacting, causal variants, whose effects may depend upon the sex, age and the environment in which they are expressed. A recently developed methodology in animal breeding derives a value of relatedness among individuals from high‐density genomic marker data, to estimate additive genetic variance within livestock populations. Here, we adapt and test the effectiveness of these methods to partition genetic variation for complex traits across genomic regions within ecological study populations where individuals have varying degrees of relatedness. We then apply this approach for the first time to a natural population and demonstrate that genetic variation in wing length in the great tit (Parus major) reflects contributions from multiple genomic regions. We show that a polygenic additive mode of gene action best describes the patterns observed, and we find no evidence of dosage compensation for the sex chromosome. Our results suggest that most of the genomic regions that influence wing length have the same effects in both sexes. We found a limited amount of genetic variance in males that is attributed to regions that have no effects in females, which could facilitate the sexual dimorphism observed for this trait. Although this exploratory work focuses on one complex trait, the methodology is generally applicable to any trait for any laboratory or wild population, paving the way for investigating sex‐, age‐ and environment‐specific genetic effects and thus the underlying genetic architecture of phenotype in biological study systems.     

Spectral acoustic structure of barking in roe deer (Capreolus capreolus). Sex-, age- and individual-related variations

In roe deer, barking is a loud call commonly given by males and females during inter- or intraspecific interactions. The analysis of a set of 19 spectral variables computed on 560 calls revealed significant variation between sexes, individuals, and probably age classes. Discriminant analysis predicted the sex of an individual with a 93.5% probability from a small portion of the bark frequency range. Among six males, a linear combination of six variables predicted the identity of the barking individual with a 70% probability. These sexual and individual differences provide the potential for social recognition from vocalizations. These results are consistent with the hypothesis that barking in roe deer may allow remote signalling of presence, location and identity, and play an important role in the territorial system of this species.