Assessing non-metric skeleton characters as a morphological tool

The appearance of non-metric skeletal characters in vertebrates results mainly from basic genetic control, as proved and documented for house mice and humans. Although the heritability of non-metric traits, mainly the presence of foramina and similar structures for blood vessels and nerves or dental occlusal variants, have been evaluated as rather low, the simultaneous consideration of several traits allows estimation of epigenetic variation in time and space as a result of genetic relationship. Thus, the main use of non-metric characters has been aimed at assessing epigenetic variability and divergence among populations. Applications extend from the problem of genetic isolation of populations, the lack of reproductive contact, detection of genetic drift, systematic studies to clarify species taxonomy, to phylogenetic interpretation.

Additionally, non-directional deviations from bilateral symmetry in non-metric characters, e.g. fluctuating asymmetry, could be caused by current environmental conditions in general. Fluctuating asymmetry is regarded as a measure of developmental instability to indicate the presence of genomic changes or the influence of contamination and/or habitat deterioration, and is also used as a further population parameter with integrated information. However, standpoints on the use of fluctuating asymmetry are quite inconsistent, fluctuating themselves between considering it a powerful biomonitoring tool to being merely a curious scientific toy.     

Genetic analysis of bilateral dermatoglyphic asymmetry in twins and their parents

The data for this study were collected on 64 twin pairs (30 MZ and 34 DZ) and their 128 parents. Two following hypotheses were evaluated: 1. Bilateral asymmetry is significantly genetically controlled; 2. The twinning phenomenon would affect the magnitude of bilateral asymmetry. The results revealed no statistically significant differences between mean values of MZ and DZ twins and their parents for the majority of the traits. Significant differences were recorded for only 6 of 96 comparisons (6%). Analysis of variance revealed separated sources of MZ, DZ and singleton variance. F-ratios, contrasting variances between different groups were significant for 26 of 96 comparisons (27%) showing heterogeneity of variance between zygosities and between twins and their parents. In addition, environmental covariance appeared to be larger for MZ than DZ with respect to directional asymmetry (DA) for all 16 traits and fluctuating asymmetry (FA) for 14 traits. These observations showed complex environmental determinism for bilateral asymmetry for the majority of dermatoglyphic traits. Significant genetic variance ratios (GVRs) were observed for four variables (25%) with respect to DA and three variables (18.75%) with respect to FA. All these significant GVRs were rendered insignificant because of evidence of greater environmental covariance for MZ twins, except possibly for DA for URC4.     

Genetic and environmental influences on bilateral dental asymmetry in Northwest Indian twins

The role of genetic and environmental factors on dental asymmetry (in maximum crown dimensions) was examined using 58 pairs of twins (23 MZ and 35 DZ) from Chandigarh, India. The t'-test for equality of means by zygosity showed only one variable significantly different among 56: this is ascribable to Type 1 error. Heterogeneity of MZ-DZ total variance was observed in 42.9% of traits of the two types (fluctuating and directional) of bilateral asymmetry. In general, MZ twins showed higher total variance than DZ pairs. MZ twins also showed stronger environmental covariance for a majority of the traits. Dental asymmetry measures thus yielded consistently low genetic variance ratios and indicated predominantly complex environmental determinism. Since fluctuating asymmetry is widely believed to be an environmental stress indicator, this data set allows confirmation of methods for detecting unequal environmental influences on the zygosities which bias estimates of genetic variance and heritability.     

Fluctuating asymmetry and genetic variability in the roe deer (Capreolus capreolus): a test of the developmental stability hypothesis in mammals using neutral molecular markers

Fluctuating asymmetry (FA), used as an indicator of developmental stability, has long been hypothesized to be negatively correlated with genetic variability as a consequence of more variable organisms being better suited to buffer developmental pathways against environmental stress. However, it is still a matter of debate if this is due to metabolic properties of enzymes encoded by certain key loci or rather to overall genomic heterozygosity. Previous analyses suggest that there might be a general difference between homeo- and poikilotherms in that only the latter tend to exhibit the negative correlation predicted by theory. In the present study, we addressed these questions by analysing roe deer (Capreolus capreolus) from five German populations with regard to FA in metric and non-metric skull and mandible traits as well as variability at eight microsatellite loci. Genetic variability was quantified by heterozygosity and mean d2 parameters, and although the latter did not show any relationship with FA, we found for the first time a statistically significant negative correlation of microsatellite heterozygosity and non-metric FA among populations. Because microsatellites are non-coding markers, this may be interpreted as evidence for the role of overall genomic heterozygosity in determining developmental stability. To test if the threshold character of non-metric traits is responsible for the metric vs non-metric difference we also carried out calculations where we treated our metric traits as threshold values. This, however, did not yield significant correlations between FA and genetic variability either.     

Discrete dental trait asymmetry in Mexican and Belizean groups

Discrete dental trait asymmetry was scored on four Mexican Indian and two Afro-Belizean groups. The Mexican populations show significantly higher mean asymmetry than the two populations from Belize even though the Belizean groups have larger teeth. Point biserial coefficients of correlation between asymmetry of discrete traits and tooth size is low and in most cases not significant. This study indicates that populational and ethnic differences exist in the asymmetry of discrete traits, suggesting that these traits may be used to investigate the etiology of dental asymmetry, and the relative roles of genetics and environment.     

Inbreeding effects on bilateral asymmetry of dermatoglyphic patterns

Bilateral correlations are higher and bilateral variances within individuals smaller in the samples of inbred individuals than in matched control groups of the same sex for pattern intensities on fingers in four series of data and also for pattern intensities on palms, toes, and soles and the palmer main line indices in the data collected from a Muslin population of West Bengal. This trend is not apparent in two series of data from the Yanadi tribe, in which the inbred and noninbred samples are not controlled for random variation of genes and environment. Increased variances between individuals and changes in means and distributions of the traits in the inbred samples of the matched data indicate some influence of homozygosity of genes for the traits on their asymmetry. The reduced variability of asymmetry of the traits in the inbred cannot be explained by homozygosity of genes for either directional or absolute asymmetry. One possible explanation is that heterozygotes for these dermatoglyphic triats are more responsive to environmental stress than homozygotes and/or increased selection in the homozygotes against genetic disorders associated with dermatoglyphic asymmetry may reduce the variability of such asymmetry.     

长阳青铜时代与大同北魏朝代人类牙齿的形态变异

本文记述了湖北长阳县深潭湾青铜时代和山西大同市郊北魏朝代两处墓葬遗址出土的人类牙齿形态变异的状况,提供人们对牙齿形态种种变异的认识,探索非测量性牙齿特征在研究种族和人群之间关系中的意义。     

Directional and Fluctuating Asymmetry in Sexual and Asexual Otiorhynchus alpicola Populations

This study concerns the contribution of directional asymmetry (DA) and fluctuating asymmetry (FA) as a characterization of variation in six sexual (diploid) and two asexual (triploid and tetraploid) populations of the weevil Otiorhynchus alpicola. It is shown that DA in sexual populations is about 1 % of the mean length of each of the seven bilateral traits and the average contribution of DA to trait variation is even lower in asexual populations (about 0.85 in triploids and 0.65 in tetraploids forms). The average contribution of FA to the total phenotypic variance is about 23 %, 12 % and 19 % in diploid, triploid and tetraploid populations, respectively. Since FA is generally regarded as a measure of developmental stability, our data indicate that triploid forms of O. alpicola are developmentally more stable than diploid and tetraploid forms. The relationship between the level of ploidy and FA is discussed.     

Nonmetric tooth crown traits in a Sri Lankan aboriginal Vedda population

This study was conducted to determine the frequencies of non-metric tooth crown traits of Vedda of Sri Lanka and to investigate the affinities of these morphological variations with those of other world populations.Fifty dental plaster casts were observed. The Arizona State University dental anthropology system was adopted for classification of the 16 traits observed. We used 13 traits to compare the Vedda and other world populations. Using the frequencies of 13 traits, Smith Mean Measure of Divergence was calculated to determine inter-population distances. Affinities among the Vedda and other world populations were expressed in two dimensions of the principal coordinate analysis.Cusp number in mandibular second molar and hypocone absence in maxillary second molar had the highest frequency at 95.9% and 93.8%, respectively. Shovelling, double shovelling in the maxillary central incisor and deflecting wrinkle in the mandibular first molar had the lowest frequency at 0%. The principal coordinate analysis showed that Sino American and Western Eurasian populations were separated in negative and positive directions in the first principal coordinate axis. Vedda located with the Western Eurasian population groups. Sahul and Sunda Pacific populations located in the intermediate position between Sino American and Western Eurasian populations.The dental phenotype of Vedda has close affinities with those of early south Asian populations. They are far different from Sino American and Sunda pacific populations. Vedda shows closer affinities to Sahul Pacific and South African (Bantu) populations.     

Bilateral asymmetry in skeletal growth and maturation as an indicator of environmental stress.

This study examined the efficacy of bilateral asymmetry in epiphyseal union as an indicator of environmental stress affecting the skeleton. We compared the extent of asymmetry in the postcranial skeleton between two cemetery samples excavated from Medieval Kulubnarti, Sudanese Nubia. Past studies have strongly suggested that these ancient Nubians experienced environmental stress-the early Christian period (550-750 AD) population to a greater extent than the late Christian period (750-1450 AD) population. We hypothesized that if bilateral asymmetry is a reflection of stress, then it should be present or greater in the more stressed population, the early Christian period population, while absent or found to a lesser extent in the less stressed population, the late Christian period population. We computed two mean values, representative of right-side and left-side epiphyseal union, for each individual in both cemetery samples, and tested for significant differences. Bilateral asymmetry was significant in the combined cemetery sample of 90 individuals (P < 0.019). When cemetery samples were tested separately, bilateral asymmetry was significant for the early Christian period sample (P < 0.001), but not for the late Christian period sample. There were no differences attributable to sex. Finally, we discuss why we conclude that environmental stress was favored over a biomechanic explanation as the cause for asymmetry. To the extent that our results support previous findings that early Christian period individuals were more affected by environmental stress than late Christian period individuals, it is reasonable to consider bilateral asymmetry in skeletal growth and maturation a good indicator of environmental stress.     

The phenetic distances of the living Druze from other human populations suggest a major genetic drift from the Western Eurasian ancestral category

The objectives were to determine the expression frequency and sexual dimorphism of 16 non-metric crown traits on the sample of permanent dentitions of the living Druze population (a Near Eastern genetic isolate) in Jordan, and to assess the biological affinity of this sample to 21 regional groups, and to the living general Jordanian population, based on these traits. Druze schoolchildren (46 males, 40 females; mean age = 16.0, sd = 0.5 years) were studied in 2011. The traits were classified using the Arizona State University dental anthropology system, counted with the individual count method, and dichotomized according to the criteria of Scott and Turner for the purpose of group comparisons. Fisher's exact test for dichotomized scores was used to assess sexual dimorphism in these traits. Smith's mean measure of divergence was used to measure all pairwise distance values among the groups. Sexual dimorphism was found in five traits (i.e., UI2 interruption grooves, 3-cusped UM2, UM1 Carabelli's tubercle/cusp, 4-cusp LM1, and LM2 Y-groove pattern). This study revealed that the dental pattern of living Druze, which is similar to that of the general Jordanian population, is sufficiently distinct from the Western Eurasian pattern and all other known dental patterns to form a distinct dental pattern for the regional group or subcategory to which these two populations belong. Moreover, the relatively large distance values of the living Druze and Jordanians from the other world groups considered, including the Western Eurasian groups, suggest a similar major genetic difference of these two populations from the Western Eurasian ancestry.     

Population history of the middle Euphrates valley: Dental non-metric traits at Tell Ashara,Tell Masaikh and Jebel Mashtale,Syria

Fifty-nine dental non-metric traits were scored using Arizona State University Dental Anthropology System on a sample of teeth from 350 human skeletons excavated at three sites in the lower middle Euphrates valley. The dataset was divided into six chronological subsets: Early Bronze Age, Middle Bronze Age, Early Iron Age with Neo-Assyrian period, Classical/Late Antiquity, Early Islamic (Umayyad and Abbasid) period and Modern period. The matrix of Mean Measure of Divergence values exhibited temporal homogeneity of the sample with only dental non-metric trait scores in the Modern subset differing significantly from most other subsets. Such a result suggests that no major gene flow occurred in the middle Euphrates valley between the 3rd millennium BCE and the early 2nd millennium CE. Only after the Mongolian invasion and large depopulation of northern Mesopotamia in the 13th century CE a major population change occurred when the area was taken over in the 17th century by Bedouin tribes from the Arabian Peninsula.     

Developmental instability in Brassica campestris (Cruciferae): fluctuating asymmetry of foliar and floral traits

The degree of fluctuating asymmetry of bilateral traits provides a measure of developmental instability, which can be influenced by genetic as well as environmental stress. We studied genetic variation between and within two populations of the mustard Brassica campestris for asymmetry of foliar (cotyledon width) and floral (petal length and width) traits as well as for phenological (germination and flowering) and performance (biomass and flowering) traits. The two populations differed in mean expression of most traits, including asymmetry. However, within-population estimates of genetic variability tended to be lower for asymmetry than other traits. Asymmetry was greater in the population that had lower biomass accumulation and flower production, which supports the idea that population-level asymmetry may be indicative of population-level performance. However, within each population, evidence that performance was negatively correlated with asymmetry was equivocal. Within populations there was little or no concordance among estimates of asymmetry based on different structures, i.e., plants that had highly asymmetrical cotyledons did not tend to have highly asymmetrical petals. The lack of a general buffering capacity at the individual level may be explained by developmental processes (e.g., action of different genes or morphogens) as well as evolutionary processes (e.g., selection on asymmetry of different traits).     

Relationships between non-metric skeletal traits and cranial size and shape.

A long-standing controversy exists about the comparative utility of metric and non-metric traits as biological indicators in population studies. We hypothesize that the underlying scale which determines the presence or absence of a cranial non-metric trait is an expression of general and/or local size variation in the cranium. Therefore metric and non-metric traits will share a common developmental determination. The hypothesis implies that the underlying scale of a non-metric trait will be correlated with measures of cranial size and shape. Forty-eight cranial metric and twenty-five cranial non-metric traits were scored on the left side of adult male crania from four North American Indian populations. New threshold traits were generated for each non-metric trait by dichotomizing discriminant scores produced by discriminant function analysis. The discriminant analysis was performed using metric traits to discriminate between groups formed by non-metric trait presence or absence. Every non-metric trait tested was significantly correlated with its threshold trait in at least one population. The correlations were of moderate to high levels depending on the trait and population sample studied. This implies that metric and non-metric traits share a moderate to high degree of developmental determination. The cause of these correlations may lie in the common effects that growth and development of the soft tissue and functional spaces of the cranium exert on both metric and non-metric traits.     

Fluctuating asymmetry in certain morphological traits in laboratory populations of Drosophila ananassae.

Fluctuating asymmetry (FA, subtle random deviations from perfect bilateral symmetry) is often used as a measure of developmental instability (DI), which results from perturbations in developmental pathways caused by genetic or environmental stressors. During the present study, we estimated FA in 5 morphological traits, viz. wing length (WL), wing to thorax ratio (W:T), sternopleural bristle number (SBN), sex-comb tooth number (SCTN), and ovariole number (ON) in 18 laboratory populations of Drosophila ananassae. FA levels of measured traits differed significantly among populations except for SBN (in males and females) and W:T ratio (in females). Positional fluctuating asymmetry (PFA), a sensitive measure of DI, also varied significantly among the populations for SBN in females and SCTN in males. Interestingly, both males and females were similar for nonsexual traits. However, when FA across all traits (sexual and nonsexual) was combined into a single composite index (CFA), significant differences were found for both populations and sexes. Males showed higher CFA values than females, suggesting that males are more prone to developmental perturbations. The magnitude of FA differed significantly among traits, being lowest for nonsexual traits (SBN, WL, W:T ratio) and highest for sexual traits (SCTN and ON). The trait size of sexual traits (SCTN and ON) was positively correlated with their asymmetry. The possible reasons for variation in FA both among traits and among populations, and the usefulness of FA as an indicator of developmental stress and phenotypic quality in D. ananassae are discussed.     

Prenatal stress and increased fluctuating asymmetry in the parietal bones of neonatal rats

Recent studies have reported increased fluctuating dental and long bone asymmetry in neonates as a function of prenatal stress. The present study was designed to assess the effects of prenatal stress on a third calcium-dependent system, membranous bone. Pregnant rats were exposed to cold, heat, or noise from conception through parturition. Bilateral parietal bone lengths were measured in the term neonates. Levels of fluctuating asymmetry were found to be significantly increased (p less than .001) in all three stressed groups compared to unstressed controls. Results support the concept of a generalized stress response, and suggestions are offered for human osteological application.     

Hand dominance and bilateral asymmetry in the structure of the second metacarpal

Bilateral asymmetry in the structure of the second metacarpal was examined in relation to functional hand dominance in a large, clinically nonselected, healthy population sample from the Baltimore Longitudinal Study of Aging. Bilateral bone measurements were made from anteroposterior hand radiographs of a total of 992 individuals, 609 males and 383 females, with an age range of 19–94 years. Hand dominance was determined on the basis of personal impression. Total width and medullary width at the midshaft of the second metacarpal were measured to 0.05 mm using a Helios caliper. These two measurements were used to derive cortical thickness, cortical bone area, periosteal (total) area, medullary area, percent cortical area, and the second moment of area in the mediolateral plane. In both right and left-handed individuals, statistically significant side differences were found in the calculated bone areas and the second moment of area, with the dominant hand being larger. Cortical thickness did not show significant side-related differences for either handedness. These results show that functional handedness leads to periosteal and endosteal expansion of the second metacarpal cortex on the dominant side, increasing bone strength without increasing cortical thickness. This is the first time this pattern of asymmetry has been reported in left-handers as well as right-handers. Our results argue for the primacy of environmental (mechanical) effects in determining bilateral asymmetry of limb bone structural properties. © 1994 Wiley-Liss, Inc.     

Inbreeding and developmental stability in three-spined sticklebacks (Gasterosteus aculeatus L.)

Fluctuating asymmetry, small non-directional departures from perfect symmetry in bilateral traits, results from the inability of individuals to buffer development against genetic and environmental perturbations. Fluctuating asymmetry is a widely used measure of developmental stability, and developmental stability has been hypothesised to be inversely related to heterozygosity. We compared male three-spined sticklebacks (Gasterosteus aculeatus L.) that had been inbred for one generation to outbred control males with respect to the asymmetry of a set of bilateral morphometric traits. Inbred fish developed significantly more asymmetric pectoral fins than their outbred counterparts, whereas neither the magnitude of asymmetry for pelvic spines nor for gill covers significantly responded to the treatment. Our results conform to a pattern of heterogeneity amongst traits in their tendency to develop asymmetrically in response to stress.