An argument for the use of total side frequencies of bilateral nonmetric skeletal traits in population distance analysis: The regression of symmetry on incidence

In population studies based on frequencies of bilateral nonmetric skeletal traits, the choice between sampling by individuals or by sides should depend less on the exigencies imposed by fragmentary remains than on fundamental assumptions about the biological meaning of symmetry/asymmetry. Though the latter has been interpreted in various ways, little attention has focused on the possibility that bilateral correlation is meaningful in quantifying genetic liability for a trait. Analysis of two independent mandibular features, mylohyoid bridge and suppressed third molar, in Indian and Eskimo population samples (total N ? 1,200) reveals a statistically significant pattern of increasing bilateral occurrence with increasing population incidence. This pattern is consistent with the theory that liability for a “quasi-continuous” variant is normally distributed with constant increment between thresholds on an underlying scale. According to theory, phenotypes with more pronounced expression (bilateral occurrence) have greater genetic potential than those with less pronounced expression (unilateral occurrence). Therefore, scoring traits in total left and right sides, by giving greater weight to bilaterally affected individuals, may provide a better estimate of the liability for the trait in the population. Viewed in a theoretical context broader than that of the sampling debate, this pattern of positive regression of symmetry on incidence means that prevalence of unilateral occurrence probably cannot be used to assess the relative strength of genetic versus nongenetic control of threshold variants.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques of Cayo Santiago. III. Relative heritability of skeletal nonmetric and metric traits

This study addresses the long-standing controversy in skeletal biology concerning the relative utility of skeletal metric and nonmetric traits for studies of biological relationship. This controversy centers on the relative heritability of these two trait sets. This paper presents heritabilities for a series of skeletal metric and nonmetric traits measured with the same sample of mother-offspring pairs from the Cayo Santiago skeletal collection of rhesus macaques. Skeletal nonmetric traits display significantly greater heritability estimates than metric traits. This difference is due primarily to the high heritability estimates of hyperostotic nonmetric traits. Foraminal traits are not significantly more heritable than skeletal metric traits. The generality of this pattern of heritability values, in which hyperostotic nonmetric traits are more highly heritable than foraminal nonmetric and metric traits, depends on future empirical study of the correlation of heritability values in populations and theoretical work.     

Quantitative genetics of cranial nonmetric traits in randombred mice: heritability and etiology

Cheverud and Buikstra (1981) demonstrated a tendency for nonmetric traits representing the number of foramina to have lower heritabilities than those representing hyperstotic or hypostotic traits in a sample of rhesus macaques. Based on this observation, Cheverud and Buikstra hypothesize that differences in the heritability of the two sets of traits may be due to differences in trait etiology. This study addresses the proposed relationship between trait heritability and etiology. Heritability values are calculated for 35 cranial nonmetric traits in a sample of 320 randombred mice using analysis of variance. The results are minimally consistent with the etiological hypothesis, but only 4 of the 35 traits showed statistically significant heritability values. These results are discussed with reference to the assumption that nonmetric traits have a strong genetic component. It is concluded that the developmental pathways that genetic variation traverses before being expressed in the form of nonmetric traits must be understood before variation in nonmetric traits can be used to its fullest potential.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. I. Single trait heritabilities

The use of skeletal nonmetric traits in studies of biological relationships often involves the assumption that variation in these traits is genetic. Studies of nonmetric traits in human groups and in inbred strains of mice and rabbits have indicated a genetic component to nonmetric trait variation. Skeletons of animals with known matrilineage membership were obtained from the Cayo Santiago skeletal collection in order to obtain a direct estimate of the heritabilities of several nonmetric traits in the free-ranging population of rhesus macaques on Cayo Santiago. Falconer's (1965) method was used to calculate heritability. Heritability estimates range from zero to one, and half of them are greater than 0.5. This indicates that there is a considerable amount of genetic variation for these traits among the Cayo macaques. There is a significant tendency for traits scoring the number of foramina to have lower heritabilities than those scoring hyperstotic or hypostotic traits.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. II. Phenotypic,genetic, and environmental correlations between traits

The general lack of phenotypic correlation among skeletal nonmetric traits has been interpreted as indicating a lack of genetic correlation among these traits. Nonmetric traits scored on animals in the skeletal collection of rhesus macaques from Cayo Santiago are used to calculate phenotypic, genetic, and environmental correlations between traits. The results show that even when phenotypic correlations are low, there may be large, significant genetic correlations among these traits. The genetic correlation pattern suggests that genes which affect nonmetric trait variation act primarily at a local level in the cranium, even though there are genes with pleiotropic effects on skeletal nonmetric traits throughout the cranium. Environmental and phenotypic correlations do not show this neighborhood pattern of correlation.     

The relationship between cranial metric and nonmetric traits in the rhesus macaques from Cayo Santiago

This study addresses the relationship between cranial metric variables and nonmetric traits using the skeletal sample of rhesus macaques from Cayo Santiago. Discriminant function analysis is used to study the metric differences between macaque crania grouped according to the presence or absence of nine nonmetric traits. The computation of total structure coefficients from the discriminant function analyses provides information regarding how closely each metric variable is related to the discriminant functions derived. Total structure coefficients have not been used previously in the study of the relationship between metric and nonmetric traits. The results of the analysis are interpreted using an explicit approach to cranial morphogenesis-functional cranial analysis. It is concluded that the relationship between cranial metric and nonmetric traits is explicable in terms of a common developmental pathway shared by the two types of traits. Identification of the specific etiology of nonmetric traits depends on future anatomical studies or organisms throughout the period of nonmetric trait development.     

Genetic correlations between sides and heritability of asymmetry for nonmetric traits in rhesus macaques on Cayo Santiago

The use of nonmetric traits for estimation of biological distance is a long-standing practice in biological anthropology. Nonmetric traits can be scored using either the individual or the side of the individual as the unit of measure. If sides of the individual are genetically correlated the use of sides would produce redundant genetic information. For this reason, Korey (Am. J. Phys. Anthropol. 53:19-23, 1980) argues for the use of individuals as the unit of measure for nonmetric traits. Ossenberg (Am. J. Phys, Anthropol. 54:471-479, 1981), however, argues that bilateral occurrence of nonmetric traits indicates greater genetic liability for the trait and that therefore the sides are the more biologically correct unit of measure. Genetic correlations for 13 cranial nonmetric traits are estimated for a sample of rhesus macaque skeletons from Cayo Santiago. In addition, heritability of asymmetry is estimated for these 13 traits as a test of Ossenberg's contention that asymmetry is genetically influenced. Significant genetic correlations between sides support Korey's contention that nonmetric traits should be scored by individual. Only two asymmetry heritabilities were significantly different from zero, providing no significant support for Ossenberg's contention that asymmetry is genetically determined. Our results support the theory that asymmetry represents a measure of the ability of an organism to buffer stresses. Therefore, a measure of the heritability of asymmetry is a measure of the heritability of the ability to buffer stresses. This ability does not appear to be heritable in this sample.     

The use of nonmetric variation in estimating human population admixture: a test case with Brazilian blacks, whites, and mulattos

Measurements in populations which serve as valid indicators of biological relationship should be proportional to genetic distance. In order to test the utility of discrete cranial traits for estimating genetic distances among populations, estimates of admixture are obtained for gene frequency data and nonmetric cranial data in São Paulo mulattos (M). The gene frequency data serve as a control that the three populations are related as stated: estimates of admixture are obtained by using São Paulo whites (W) and blacks (B) as parental populations and by estimating the parameter of admixture, m, in the model p_M = (1 ? m) p_W + mp_B (Elston, 1971) where the p's are either gene frequencies or nonmetric trait frequencies. A test of goodness of fit of the model provides a means of ascertaining whether or not the data fit this linear model. While the gene frequency data indicate distances among the three populations which are highly compatible with the linear model of admixture, the nonmetric data show significant deviations from the model. This implies that the frequencies of the nonmetric traits in the populations used in this analysis are not a linear function of genetic distance. This discourages the use of nonmetric traits in making quantitative conclusions about genetic relationships. It also suggests the need for investigation of the use of other skeletal characters for estimating genetic distance, as well as approaches for such investigations through the study of hybrid individuals.     

An examination of the meaning of cranial discrete traits for human skeletal biological studies

Discrete traits are of increasing interest in comparative skeletal biological research. Characteristics justifying their use have been investigated primarily in mice, however. Using 72 discrete variants, 321 human skulls from the Terry Collection of known race, sex and age have been studied. Significant sex and age differences were detected. Inter-trait correlation was found to be at a low but significant overall level. Multivariate comparison with conventional craniometric analysis was undertaken on subdivisions of the sample, and distance based on metric and nonmetric data were concordant. It is concluded, on the basis of these findings and the discontinuous variant frequency distributions, that discrete traits in isolation are not of paramount value to skeletal genetic studies, but may be vital in comparison and conjunction with other types of data in analyzing the population genetics of extinct groups.     

Population structure and skeletal variation in the Late Woodland of west-central Illinois

This paper analyzes nonmetric trait variation in 11 late Late Woodland (ca. AD 700-1000) and one Mississippian (AD 1000-1300) skeletal samples from west-central Illinois from a population-structure perspective. Most of the sites are of the Bluff phase of Late Woodland in the lower Illinois River valley; others are from a nearby, contemporary archaeological phase. Late Woodland as a whole era (ca. AD 250-1000) was a period of marked population growth and expansion into new regional environments, trends that accompanied horticultural intensification in the area. Overall variation between sites was low, but males, females, and the total sample exhibited a significant geographic component to variation due to interregional morphological differences. The Bluff sites tended to group together relative to the non-Bluff sites. However, there was no significant geographic component to variation among the Bluff sites. The results are only partially consistent with archaeological data suggesting population growth and expansion through fissioning. Previous studies have demonstrated significant heterogeneity for nonmetric trait frequencies among Middle Woodland (ca. 100 BC to AD 250) sites, suggesting a Middle to Late Woodland change in population structure that lowered levels of morphological variation. This supports a model of increased intra- and interregional interaction from Middle to Late Woodland times developed from ceramic data by Braun and by Braun and Plog.     

Dental anthropology of Central-Southern,Iron Age Italy: The evidence of metric versus nonmetric traits

Discrete and metric dental traits are used to assess biological similarities and differences among 13 bioarchaeological populations located on each side of the Apennine mountains in central-southern Italy and dated to the first millennium BC . An initial hypothesis, that the mountain chain might provide a significant geographical barrier for population movement (resulting in greater biological affinities among those groups on the same side), is not supported. Instead, the samples appear to cluster more on the basis of time than geography. Archaeological evidence, however, supports an association between populations on opposite sides of the mountains and thus is in accord with the dental data. As anticipated, discrete dental traits appear to be more useful than metric dental traits in assessing such population affinities. This research represents a beginning to a better comprehension of the complexity of the biological and cultural dynamics of Italian populations during recent millennia. Am J Phys Anthropol 107:371–386, 1998. © 1998 Wiley-Liss, Inc.     

A comparison of sampling methods for bibionid larvae (Diptera: Bibionidae) in grassland

Three extraction methods were compared for the recovery of bibionid larvae from grassland soil samples. Only wet-sieving followed by flotation in a saturated salt solution yielded bibionids. No larvae were recovered either with modified Tullgren funnels or by slow immersion of soil cores into a saturated salt solution. The efficiency with which larval bibionid populations can be estimated is poor. Generally, smaller cores yielded more larvae per volume of soil. Most bibionids were found in the top 4 cm of soil. It is concluded that 10 cm diameter and 6–8 cm deep soil cores are an acceptable compromise between efficiency and sampling effort but sample size will largely be determined by the resources available for processing of samples.     

The statistical treatment of correlated bilateral traits in the analysis of cranial material.

At present there is no standardized method to derive frequencies from bilateral non-metric traits. In this paper the commonly used methods are evaluated in light of statistical considerations and degree of sample preservation. In particular, we explore the question of dependence between sides for the bilateral traits. Several workers have tested for bilateral trait correlation in incorrect ways, confusing tests for differences in side frequencies with tests for independence. An easy method to test for independence, using the chi-squared test, is recommended. This test is used on 16 bilateral traits in a large sample of prehistoric carnia from Central California and significant dependence is found for all 16 of these traits. We suggest the traditional method of deriving frequencies be used. Both sides of the cranium should be considered, dividing the number of times the trait occurs by the number of sides available for observation. This method can be used even in poorly preserved samples. Side to side correlation may be compensated for by modifying the constants subtracted in the mean measure of divergence and in the expression for the approximate variance of the mean measure of divergence when the samples are drawn from identical populations.     

Selection in a cycling population: differential response among skeletal traits

Abstract Population density cycles influence phenotypic evolution through both density‐dependent selection during periods of high density and through enhanced genetic drift during periods of low density. We investigated the response of different phenotypic traits to the same density cycles in a population of the yellow‐necked mouse, Apodemus flavicollis, from Bia?owieza National Park in Poland. We examined nonmetric skull traits, skull and mandible size, skull and mandible shape, and transferrin allele frequencies. We found that all of the traits changed significantly over the seven‐year study period. The greatest changes in nonmetric traits and mandible size occurred during periods of increasing density, and the magnitude of changes in skull and mandible shape was correlated with the magnitude of density changes. Frequencies of transferrin alleles changed the most when population density was in decline. Changes among the five phenotypic traits were generally uncorrelated with one another, except for skull and mandible shape. Nonmetric traits were selectively neutral when assessed with Q_ST/F_ST analysis, whereas mandible size, mandible shape, and skull shape showed evidence of fairly strong selection. Selection on skull size was weak or nonexistent. We discuss how different assumptions about the genetic components of variance affect Q_ST estimates when phenotypic variances are substituted for genetic ones. We also found that change in mandible size, mandible shape, skull size, and skull shape were greater than expected under a neutral model given reasonable assumptions about heritability and effective population size.     

A critical review of the mean measure of divergence and Mahalanobis distances using artificial data and new approaches to the estimation of biodistances employing nonmetric traits

This article reviews the two most common distance measures employed for the calculation of biodistances based on nonmetric traits, the mean measure of divergence (MMD) and the tetrachoric Mahalanobis D² distance (TMD). In addition, two new approaches for the estimation of biodistances from nonmetric traits are proposed and assessed. The first (OMD) is based on the direct application of the Mahalanobis distance to ordinally recorded data before their transformation to binary dichotomies. The second (RMD) approximates the covariances of the Mahalanobis distance by the Pearson correlation coefficients calculated in the binary dataset. The application of all four methods to artificial datasets demonstrates that they overall provide a satisfactory estimation of the biodistance among samples especially when the number of statistically non significant distances is very limited. However, the best performance is observed by the OMD, whereas special attention should be paid to the TMD since its values might come out of an ill‐conditioned system. The influence of the number of traits, the effect of missing values, as well as the validity of the test statistics used to assess biodistance significance are also examined and discussed. Am J Phys Anthropol 157:284–294, 2015. © 2015 Wiley Periodicals, Inc.     

植物功能性状对土壤保持的影响研究述评

植被对土壤保持具有重要的影响,但是从植物功能性状的角度总结评述植被对土壤保持影响的研究并不多见。总结评述了植物地上功能性状、地下功能性状对土壤保持功能的影响以及植物地上、地下功能性状的关系,认为:(1)植被地上部分功能性状对土壤保持的作用主要体现在对溅蚀、面蚀的影响及间接改变土壤理化性质等方面,其功能性状指标主要包括叶面积、叶长、叶宽、枝数、植被高度等;(2)植被地下部分功能性状对土壤保持的作用主要体现在固持土壤、提高土壤抗剪切强度、提高土壤抗侵蚀能力、增强土壤渗透性,植物根系固持土壤与根系抗拉能力密切相关,植物根系土壤的物理和水文性质,与细根比例、根长密度、根表面积等性状密切相关;(3)可以通过植物地上部分功能性状间接反映地下部分功能性状,但是现有研究多为定性认识;(4)在植物功能性状对土壤保持的研究中亟待加强植被地上地下功能性状的长期定位监测,深化植被功能性状尤其是根系特征与土壤保持的作用机理,加强植被地上部分、地下部分功能性状的定量表达,建立植被功能性状与土壤保持功能的定量关系,实现植被功能性状与土壤保持功能特征的动态链接。     

Viscoelastic properties of passive skeletal muscle in compression: stress-relaxation behaviour and constitutive modelling

The compressive properties of skeletal muscle are important in impact biomechanics, rehabilitation engineering and surgical simulation. However, the mechanical behaviour of muscle tissue in compression remains poorly characterised. In this paper, the time-dependent properties of passive skeletal muscle were investigated using a combined experimental and theoretical approach. Uniaxial ramp and hold compression tests were performed in vitro on fresh porcine skeletal muscle at various rates and orientations of the tissue fibres. Results show that above a very small compression rate, the viscoelastic component plays a significant role in muscle mechanical properties; it represents approximately 50% of the total stress reached at a compression rate of 0.5% s⁻¹. A stiffening effect with compression rate is observed especially in directions closer to the muscle fibres. Skeletal muscle viscoelastic behaviour is thus dependent on compression rate and fibre orientation.

A model is proposed to represent the observed experimental behaviour, which is based on the quasi-linear viscoelasticity framework. A previously developed strain-dependent Young's Moduli formulation was extended with Prony series to account for the tissue viscoelastic properties. Parameters of the model were obtained by fitting to stress-relaxation data obtained in the muscle fibre, cross-fibre and 45° directions. The model then successfully predicted stress-relaxation behaviour at 60° from the fibre direction (errors <25%). Simultaneous fitting to data obtained at compression rates of 0.5% s⁻¹, 1% s⁻¹ and 10% s⁻¹ was performed and the model provided a good fit to the data as well as good predictions of muscle behaviour at rates of 0.05% s⁻¹ and 5% s⁻¹ (errors <25%).