[Show abstract][Hide abstract] ABSTRACT: The domestication of wolves is currently under debate. Where, when and from which wolf sub-species dogs originated are being investigated both by osteoarchaeologists and geneticists. While DNA research is rapidly becoming more active and popular, morphological methods have been the gold standard in the past. But even today morphological details are routinely employed to discern archaeological wolves from dogs. One such morphological similarity between Canis lupus chanco and dogs was published in 1977 by Olsen and Olsen. This concerns the “turned back” anatomy of the dorsal part of the vertical ramus of the mandible that was claimed to be specific to domestic dogs and Chinese wolves C. lupus chanco, and “absent from other canids”. Based on this characteristic, C. lupus chanco was said to be the progenitor of Asian and American dogs, and this specific morphology has been continuously used as an argument to assign archaeological specimens, including non-Asian and non-American, to the dog clade. We challenged this statement by examining 384 dog skulls of 72 breeds and 60 skulls of four wolf sub-species. Only 20 % of dog mandibles and 80 % of C. lupus chanco showed the specific anatomy. In addition, 12 % of Canis lupus pallipes mandibles showed the “turned back” morphology. It can be concluded that the shape of the coronoid process of the mandible cannot be used as a morphological trait to determine whether a specimen belongs to a dog or as an argument in favour of chanco as the progenitor to dogs.
[Show abstract][Hide abstract] ABSTRACT: For more than a century, the orbital angle has been studied by many authors to distinguish dog skulls from their progenitor, the wolf. In early studies, the angle was reported to be different between dogs (49°–55°) and wolves (39°–46°). This clear difference was, however, questioned in a more recent Scandinavian study that shows some overlap. It is clear that in all studies several methodological issues were unexplored or unclear and that group sizes and the variety of breeds and wolf subspecies were small. Archaeological dog skulls had also not been studied. Our goal was to test larger and more varied groups and add archaeological samples as they are an evolutionary stage between wolves and modern dogs. We also tested the influence of measuring methods, intra- and inter-reliability, angle symmetry, the influence of variations in skull position and the possibility of measuring and comparing this angle on 3D CT scan images. Our results indicate that there is about 50 % overlap between the angle range in wolves and modern dogs. However, skulls with a very narrow orbital angle were only found in wolves and those with a very wide angle only in dogs. Archaeological dogs have a mean angle very close to the one of the wolves. Symmetry is highest in wolves and lowest in archaeological dogs. The measuring method is very reliable, for both inter- and intra-reliability (0.99–0.97), and most skull position changes have no statistical influence on the angle measured. Three-dimensional CT scan images can be used to measure OA, but the angles differ from direct measuring and cannot be used for comparison. Evolutionary changes in dog skulls responsible for the wider OA compared to wolf skulls are mainly the lateralisation of the zygomatic process of the frontal bone. Our conclusion is that the orbital angle can be used as an additional morphological measuring method to discern wolves from recent and archaeological dogs. Angles above 60° are certainly from recent dogs. Angles under 35° are certainly of wolves.
[Show abstract][Hide abstract] ABSTRACT: Digit length ratio (primarily 2D:4D) has become increasingly popular as a possible biomarker of intrauterine steroid exposure in the human medical, social and psychological literature. Human males tend to have lower digit ratios than females, and individuals with low ratios tend to excel in physical performance, especially in endurance-related sports. Because early limb development is evolutionarily conservative, it has been speculated that these trends should also be visible in other tetrapod vertebrates. However, studies on non-human vertebrates are scant, and their results suggest that sexual dimorphism in digit ratios and the associations with physical performance are much more intricate and taxon-specific than presumed. In this study, we compared digit ratios of two Podarcis lizards among sexes, colour morphs and species. We also tested for associations with three performance characteristics that are of ecological relevance. Both species examined exhibit male-larger sexual dimorphism in digit ratio. 2D:4D, 3D:4D and 2D:3D ratios are tightly correlated within the manus and the pes, but less so between manus and pes. In the colour polymorphic species P. melisellensis, the yellow morph exhibits higher dimorphism than the orange and white morphs. Digit ratios did not correlate with individual performance for sprint speed or endurance, but within males of P. melisellensis, individuals with higher digit ratios correlated positively with head size and bite force. We conclude that digit ratios in lizards deserve attention, because they exhibit sexual dimorphism and correlate with ecologically relevant morphological and performance variables. As lizard species differ widely in mating systems, reproductive mode, habitat use and locomotor behaviour, they seem excellent model animals for studying patterns in digit length ratios.
[Show abstract][Hide abstract] ABSTRACT: The importance of measurement error in studies of asymmetry has been acknowledged for a long time. It is now common practice to acquire independent repeated measurements of trait values and to estimate the degree of measurement error relative to the amount of asymmetry. Methods also allow obtaining unbiased estimates of asymmetry, both at the population and individual level. One aspect that has been ignored is potential between-individual variation in measurement error. In this paper, I develop a new method to investigate this variation in measurement error and to generate unbiased estimates of individual asymmetries. Simulations show that variation in measurement error can indeed result in biased estimates of individual asymmetry and that the proposed method adequately provides unbiased estimates. The method is applied to two empirical datasets and shows that, at least in some traits, substantial variations in measurement occur. The limitations of the model are discussed.
[Show abstract][Hide abstract] ABSTRACT: As both degree of masculinity/femininity in sexually dimorphic organisms and developmental instability are put forward as indicators of individual quality, they are expected to correlate positively. However, the results in the literature are equivocal. One reason may be that most research has been performed in populations exposed to relatively low levels of parasitism. This article studies associations between the degree of masculinity/femininity and the fluctuating asymmetry (a measure of developmental instability) of skulls of three primate species [baboon (Papio anubis), eastern lowland gorilla (Gorilla beringei graueri), and chimpanzee (Pan troglodytes)] collected in the wild. The exposure to infection and parasites is likely to be higher in these individuals compared to captive primates and humans, and therefore likely to increase the levels of association. While the degree of individual masculinity/femininity and levels of fluctuating asymmetry were quantified accurately, I found no evidence of an association between them. My results thus are in line with the conclusion of a recent meta-analysis, that there is little evidence for associations between masculinity/femininity and developmental instability.
No preview · Article · Feb 2015 · International Journal of Primatology
[Show abstract][Hide abstract] ABSTRACT: The often observed directional asymmetry (DA) in human limb bones may have a genetic/developmental basis and/or could emerge from different mechanical loadings across sides due to handedness. Because behavioural lateralization in itself has a genetic basis, it has been suggested that DA in limbs could develop prenatally as a pre-adaptation to adult life. However, the presence of consistent differences in the size of left and right limb bones in early development is understudied. We study asymmetry in limb bones during early development (10-20 weeks of gestation) in a sample of 178 aborted foetuses. Statistically significant DA was found in several upper and lower limb bones, where the right-hand side was consistently larger than the left. We argue that this pattern is probably the consequence of developmental processes related to internal asymmetric positioning of organs.
[Show abstract][Hide abstract] ABSTRACT: Abstract Background: Studies of the process of human mate selection and attractiveness have assumed that selection favours morphological features that correlate with (genetic) quality. Degree of masculinity/femininity and fluctuating asymmetry (FA) may signal (genetic) quality, but what information they harboured and how they relate to fitness is still debated. Aim: To study strength of associations between facial masculinity/femininity, facial FA, attractiveness and physical strength in humans. Subjects and methods: Two-hundred young males and females were studied by measuring facial asymmetry and masculinity on the basis of frontal photographs. Attractiveness was determined on the basis of scores given by an anonymous panel, and physical strength using hand grip strength. Results: Patterns differed markedly between males and females and analysis method used (univariate vs multivariate). Overall, no associations between FA and attractiveness, masculinity and physical strength were found. In females, but not males, masculinity and attractiveness correlated negatively and masculinity and physical strength correlated positively. Conclusion: Further research into the differences between males and females in associations between facial morphology, attractiveness and physical strength is clearly needed. The use of a multivariate approach can increase our understanding of which regions of the face harbour specific information of hormone levels and perhaps behavioural traits.
No preview · Article · Feb 2014 · Annals of Human Biology
[Show abstract][Hide abstract] ABSTRACT: Ratios of digit lengths are studied intensively as markers of prenatal sex hormone levels.
Study sexual dimorphism in ratios of metacarpals, which received less attention.
We studied six metacarpal ratios in deceased human fetuses of ages 10 to 42weeks.
We found no indication of a sexual dimorphism at this early stage of development.
Full-text · Article · Jan 2014 · Early human development
[Show abstract][Hide abstract] ABSTRACT: Most gait-scoring scales for pigs have a limited number of categories, supposedly to improve repeatability. However, reducing the number of categories could lead to loss of information if the observers' discriminative capacities are underused. With a recently estimated within-herd prevalence of sow lameness of 8.8% to 16.9% in the European Union and the associated losses, the availability of reliable tools for the timely detection of initial cases warrants attention. This study investigated the intra-and inter-observer repeatability (intra-OR and inter-OR) of three gait-scoring scales for sows: a continuous 'tagged' visual analogue scale (tVAS, measured in mm), a 5-point and a 2-point ordinal scale (5P and 2P), all with the same descriptors. Veterinary medicine students (n = 108) were trained to use the scales and then asked to score 90 videos (30 per scale) of sows with normal and abnormal gait. Thirty-six videos were shown once and 18 were randomly shown three times, of which one mirrored horizontally. The students' opinions on the scales were also collected. Intra-and inter-OR were higher with the tVAS than the 2P scale (inter-OR: 0.73 v. 0.60; P
[Show abstract][Hide abstract] ABSTRACT: Aneuploidies cause gene-dosage imbalances that presumably result in a generalized decreased developmental homeostasis, which is expected to be detectable through an increase in fluctuating asymmetry (FA) of bilateral symmetric traits. However, support for the link between aneuploidy and FA is currently limited and no comparisons among different aneuploidies have been made. Here, we study FA in deceased human fetuses and infants from a 20-year hospital collection. Mean FA of limb bones was compared among groups of aneuploidies with different prenatal and postnatal survival chances and two reference groups (normal karyogram or no congenital anomalies). Limb asymmetry was 1.5 times higher for aneuploid cases with generally very short life expectancies (trisomy 13, trisomy 18, monosomy X, triploidy) than for trisomy 21 patients and both reference groups with higher life expectancies. Thus, FA levels are highest in groups for which developmental disturbances have been highest. Our results show a significant relationship between fluctuating asymmetry, human genetic disorders and severity of the associated abnormalities.
Full-text · Article · Jan 2014 · Scientific Reports
[Show abstract][Hide abstract] ABSTRACT: Fluctuating asymmetry (FA), as an indirect measure of developmental instability (DI), has been intensively studied for associations with stress and fitness. Patterns, however, appear heterogeneous and the underlying causes remain largely unknown. One aspect that has received relatively little attention in the literature is the consequence of direct mechanical effects on asymmetries. The crucial prerequisite for FA to reflect DI is that environmental conditions on both sides should be identical. This condition may be violated during early human development if amniotic fluid volume is deficient, as the resulting mechanical pressures may increase asymmetries. Indeed, we showed that limb bones of deceased human fetuses exhibited increased asymmetry, when there was not sufficient amniotic fluid (and, thus, space) in the uterine cavity. As amniotic fluid deficiency is known to cause substantial asymmetries and abnormal limb development, these subtle asymmetries are probably at least in part caused by the mechanical pressures. On the other hand, deficiencies in amniotic fluid volume are known to be associated with other congenital abnormalities that may disturb DI. More specifically, urogenital abnormalities can directly affect/reduce amniotic fluid volume. We disentangled the direct mechanical effects on FA from the indirect effects of urogenital abnormalities, the latter presumably representing DI. We discovered that both factors contributed significantly to the increase in FA. However, the direct mechanical effect of uterine pressure, albeit statistically significant, appeared less important than the effects of urogenital abnormalities, with an effect size only two-third as large. We, thus, conclude that correcting for the relevant direct factors allowed for a representative test of the association between DI and stress, and confirmed that fetuses form a suitable model system to increase our understanding in patterns of FA and symmetry development.
[Show abstract][Hide abstract] ABSTRACT: Directional asymmetry (DA), where at the population level symmetry differs from zero, has been reported in a wide range of traits and taxa, even for traits in which symmetry is expected to be the target of selection such as limbs or wings. In invertebrates, DA has been suggested to be non-adaptive. In vertebrates, there has been a wealth of research linking morphological asymmetry to behavioural lateralisation. On the other hand, the prenatal expression of DA and evidences for quantitative genetic variation for asymmetry may suggest it is not solely induced by differences in mechanic loading between sides. We estimate quantitative genetic variation of fetal limb asymmetry in a large dataset of rabbits. Our results showed a low but highly significant level of DA that is partially under genetic control for all traits, with forelimbs displaying higher levels of asymmetry. Genetic correlations were positive within limbs, but negative across bones of fore and hind limbs. Environmental correlations were positive for all, but smaller across fore and hind limbs. We discuss our results in light of the existence and maintenance of DA in locomotory traits.
[Show abstract][Hide abstract] ABSTRACT: Fluctuating asymmetry (the directionally random asymmetry of bilateral structures, FA) is commonly used as a measure of developmental instability, and may increase with stress. As several studies reported a relation between FA and developmental abnormalities, we investigate whether FA could be an additional perhaps more sensitive marker of developmental toxicity. The aim of this work is analyzing patterns of FA in multiple traits in a large dataset of rabbit fetuses, which were prenatally exposed to a toxic compound and sacrificed just before natural delivery. Gravid females were exposed to three doses of this compound, inducing abnormalities in the fetuses at the high dose only. The average FA, however, was already higher than control in rabbit fetuses of the low-dose group but did not further increase with higher concentrations. Moreover, the increase in FA differed between traits, with the hindlimbs showing the strongest response. In addition, we did not find any association between FA and the presence of fetal abnormalities at the individual level. Although these results suggest that FA may act as "an early warning system," we did not find a dose-response relationship with increasing stress and effects were trait-specific. Further testing is needed before FA may be considered as a sensitive marker in developmental toxicity studies.
No preview · Article · Aug 2013 · Birth Defects Research Part B Developmental and Reproductive Toxicology
[Show abstract][Hide abstract] ABSTRACT: The mechanisms of developmental buffering and its relevance to the evolutionary process have recently attracted a lot of attention in both developmental and evolutionary biology. Among other things, whether the two components of developmental buffering [i.e. canalization and developmental stability (DS)] have a common basis has long been the subject of debate. In the present study, we examine the association between fluctuating asymmetry (i.e. the directionally random asymmetry of bilateral structures), a measure of DS, and between-individual variation of long bones in over 1000 rabbit foetuses. The lack of correlations between fluctuating asymmetry and between-individual variation at the individual, litter and treatment level, in combination with the absence of correspondence among covariance matrices, supports distinct developmental mechanisms for DS and canalization. We discuss our results in the context of recent insights into the mechanisms of developmental buffering.
No preview · Article · Jun 2013 · Biological Journal of the Linnean Society
[Show abstract][Hide abstract] ABSTRACT: The genetic basis of developmental instability (DI) remains largely unknown as a result of its morphological expression, fluctuating asymmetry (FA), poorly reflecting DI, especially if few traits are studied. The typically low values of heritability of FA (h2FA) can be translated into higher values of DI (h2DI) by the hypothetical repeatability, yet leading to wide confidence intervals. Thus, high sample sizes and/or several traits are indispensible for reaching meaningful conclusions. To obtain more insights into quantitative genetic variation of DI, we investigated between-family variance in DI in six long bones of 1126 foetuses of the New Zealand white rabbit from a full-sib experiment. We applied different approaches to obtain genetic parameters for DI. Heritabilities and the coefficients of between-family variation (CVB) were calculated for six individual traits and composite indices. The results obtained, despite a likely upward bias as a result of maternal and non-additive effects, lend support to the presence of moderate additive genetic variance for DI. It is suggested that, in foetal traits, the environmental variance was minimal, leading to a high likelihood of detecting genetic variation in DI, thus creating an ideal model system for studying the genetic basis of DI.
No preview · Article · May 2013 · Biological Journal of the Linnean Society
[Show abstract][Hide abstract] ABSTRACT: Although bird-tick systems affect the human risk of tick-borne diseases, very little is known about the ecological factors that shape the spatio-temporal variation of tick infestations in terrestrial songbirds. We present a risk model that explains the levels of infestation of Ixodes ricinus, the main vector of Borrelia burgdorferi s.l., during the breeding season of the great tit (Parus major), one of Europe's most abundant avian reservoir hosts of Borrelia burgdorferi s.l.. Tit tick burden were modeled as a function of variables summarizing vegetation, climate, proxies for mammal abundance and characteristics of individual birds and their nests. Tick loads were positively associated with the relative humidity prior to capture of the bird and the cover of bracken inside its territory. The number of cold winter days prior to the bird's breeding season showed a negative association with tick loads. None of the proxies for mammal abundance correlated with tick loads. Tick loads decreased with age in female tits, whereas they increased with age in male tits. Tick burdens in the parental tits were positively associated with their brood size and negatively correlated with the average nestling body weight. Possible mechanisms include: how tit foraging influences tick encounter rates, host tick resistance mechanisms and the environmental conditions that simultaneously affect tick exposure risk and brood characteristics. We believe this study provides the first detailed insights into the ecological factors that shape tick burden in a terrestrial songbird.
Full-text · Article · Apr 2013 · International journal for parasitology
[Show abstract][Hide abstract] ABSTRACT: The study of asymmetry can provide insights into genetic and environmental influences on organismal development. Directional asymmetry (DA) can be either adaptive or non-adaptive, whereas fluctuating asymmetry (FA) – defined as small non-directional departures from symmetry in bilateral traits – is thought to be an indicator of genetic or environmental stress experienced during development. Using data from 28 European populations, we assessed the degree of DA and FA in the lateral plates of threespine sticklebacks Gasterosteus aculeatus and surveyed the direction of DA and differences in levels of DA and FA in different habitat types (viz. marine, lake and river populations). DA differed between habitats, with right-biased DA found in the marine populations and no directional bias found in lake and river populations. Differences in DA among habitats may be a by-product of habitat-specific developmental instability resulting in asymmetry, or it may indicate habitat-specific differences in selection against/for symmetry, as has been proposed in previous research of sticklebacks. Also, the presence of FA varied depending upon habitat type, but it also depended on plate morph – a variable confounded with the habitat effect. While we cannot rule out factors such as stress as a cause of population differences in FA, it may also simply be a by-product of other evolutionary processes (e.g. lateral plate number reduction) without functional basis.
Full-text · Article · Apr 2013 · Journal of Zoology