Kathleen S. Paul

A simple rule governs the evolution and development of hominin tooth size

The variation in molar tooth size in humans and our closest relatives (hominins) has strongly influenced our view of human evolution. The reduction in overall size and disproportionate decrease in third molar size have been noted for over a century, and have been attributed to reduced selection for large dentitions owing to changes in diet or the acquisition of cooking. The systematic pattern of size variation along the tooth row has been described as a ‘morphogenetic gradient’ in mammal, and more specifically hominin, teeth since Butler and Dahlberg. However, the underlying controls of tooth size have not been well understood, with hypotheses ranging from morphogenetic fields to the clone theory. In this study we address the following question: are there rules that govern how hominin tooth size evolves? Here we propose that the inhibitory cascade, an activator–inhibitor mechanism that affects relative tooth size in mammals, produces the default pattern of tooth sizes for all lower primary postcanine teeth (deciduous premolars and permanent molars) in hominins. This configuration is also equivalent to a morphogenetic gradient, finally pointing to a mechanism that can generate this gradient. The pattern of tooth size remains constant with absolute size in australopiths (including Ardipithecus, Australopithecus and Paranthropus). However, in species of Homo, including modern humans, there is a tight link between tooth proportions and absolute size such that a single developmental parameter can explain both the relative and absolute sizes of primary postcanine teeth. On the basis of the relationship of inhibitory cascade patterning with size, we can use the size at one tooth position to predict the sizes of the remaining four primary postcanine teeth in the row for hominins. Our study provides a development-based expectation to examine the evolution of the unique proportions of human teeth.

Performance analysis of deciduous morphology for detecting biological siblings

OBJECTIVES Family-centered burial practices influence cemetery structure and can represent social group composition in both modern and ancient contexts. In ancient sites dental phenotypic data are often used as proxies for underlying genotypes to identify potential biological relatives. Here, we test the performance of deciduous dental morphological traits for differentiating sibling pairs from unrelated individuals from the same population. MATERIALS AND METHODS We collected 46 deciduous morphological traits for 69 sibling pairs from the Burlington Growth Centres long term Family Study. Deciduous crown features were recorded following published standards. After variable winnowing, inter-individual Euclidean distances were generated using 20 morphological traits. To determine whether sibling pairs are more phenotypically similar than expected by chance we used bootstrap resampling of distances to generate P values. Multidimensional scaling (MDS) plots were used to evaluate the degree of clustering among sibling pairs. RESULTS Results indicate an average distance between siblings of 0.252, which is significantly less than 9,999 replicated averages of 69 resampled pseudo-distances generated from: 1) a sample of non-relative pairs (P < 0.001), and 2) a sample of relative and non-relative pairs (P < 0.001). MDS plots indicate moderate to strong clustering among siblings; families occupied 3.83% of the multidimensional space on average (versus 63.10% for the total sample). CONCLUSIONS Deciduous crown morphology performed well in identifying related sibling pairs. However, there was considerable variation in the extent to which different families exhibited similarly low levels of phenotypic divergence.

The Patterning Cascade Model and Carabelli's trait expression in metameres of the mixed human dentition: exploring a morphogenetic model

OBJECTIVES The Patterning Cascade Model (PCM) provides an evolutionary developmental framework for exploring diversity in tooth crown form. According to the model, proximity of secondary enamel knots and tooth germ size track underlying developmental processes that dictate ultimate crown morphology (i.e., cusp number, accessory cusp presence/size). Previous research has shown the model to successfully predict variation in Carabellis trait expression between antimeric and metameric pairs of human permanent molars. In this study, we quantify Carabellis trait expression for metameres of the mixed dentition (dm2 and M1) and assess the PCMs potential for explaining differences in expression between the two elements. MATERIALS AND METHODS Crown dimensions, intercusp distances, and Carabellis trait expression were collected from 49 subadults possessing observable dm2/M1 pairs. Wilcoxon signed-rank tests and paired t-tests were performed to assess whether metameres differ significantly in morphometric variables. We explored the relationships between relative intercusp distances (RICDs) and Carabellis trait expression using proportional odds logistic regression. RESULTS Intra-individual dm2/M1 pairs differed significantly in Carabellis trait expression (p = 0.01), with dm2 exhibiting higher grades of expression more commonly despite its smaller crown size. Paired molars differed in only one statistically significant RICD: metacone-hypocone (p < 0.01). Most RICDs shared the predicted negative relationship with Carabellis trait expression, but this relationship was only statistically significant for three RICDs in the dm2 (mean, protocone-paracone, metacone-hypocone). CONCLUSIONS We found mixed support for the PCMs ability to explain differences in Carabellis trait expression between metameres of the mixed molar row. Results suggest that protocone-paracone enamel knot spacing has the greatest influence on Carabellis trait expression. Lack of statistical significance for many of the relationships explored may reflect limitations related to sample composition and sample size.

Heritability and genetic integration of tooth size in the South Carolina Gullah

OBJECTIVES This article provides estimates of narrow-sense heritability and genetic pleiotropy for mesiodistal tooth dimensions for a sample of 20th century African American individuals. Results inform biological distance analysis and offer insights into patterns of integration in the human dentition. MATERIALS AND METHODS Maximum mesiodistal crown dimensions were measured using Hillson-FitzGerald calipers on 469 stone dental casts from the Menegaz-Bock Collection. Narrow-sense heritability estimates and genetic and phenotypic correlations were estimated using SOLAR 8.1.1 with covariate screening for age, sex, age*sex interaction, and birth year. RESULTS Heritability estimates were moderate (∼0.10 - 0.90; h2 mean = 0.51) for most measured variables with sex as the only significant covariate. Patterns of genetic correlation indicate strong integration across tooth classes, except molars. Comparison of these results to previously published work suggests lower overall heritability relative to other human populations and much stronger genetic integration across tooth classes than obtained from nonhuman primate genetic pleiotropy estimates. CONCLUSIONS These results suggest that the high heritabilities previously published may reflect overestimates inherent in previous study designs; as such the standard estimate of 0.55 used in biodistance analyses may not be appropriate. For the Gullah, isolation and endogamy coupled with elevated levels of physiological and economic stress may suppress narrow-sense heritability estimates. Pleiotropy analyses suggest a more highly integrated dentition in humans than in other mammals.

Comparative performance of deciduous and permanent dental morphology in detecting biological relatives

OBJECTIVES Dental morphology plays a key role in reconstructing population history and evolutionary relationships at global, regional, and intracemetery scales. At the inter-individual level, it is assumed that close biological kin exhibit greater phenotypic similarity than non-relatives. Heritability estimates provide one measure of phenotypic resemblance but are not easily incorporated into analyses of archaeological samples. In this study we evaluate the assumption that relatives are more similar phenotypically than non-relatives. We compare results for permanent dental morphology to those obtained using deciduous dental morphology in a matched dataset (Paul & Stojanowski, ). MATERIALS AND METHODS Permanent trait expression was scored from dental casts representing 69 sibling pairs, curated as part of the longitudinal Burlington Growth Study. Simulating a biodistance approach, 22 morphological traits of permanent tooth crowns were used to generate 69 inter-relative and 2,076 non-relative Euclidean distances. Following distance ordination, family-specific dispersion values were calculated from multidimensional scaling coordinates. Output was compared to that of a previous study that focused on deciduous crown variation in the same set of individuals (Paul & Stojanowski, ). Mantel tests were used to evaluate the correlation of a proxy genetic distance matrix to both the permanent and deciduous dental distance matrices. RESULTS On average, inter-relative distances generated from morphological traits of permanent tooth crowns were smaller than expected by chance based on resampling (p < 0.001). While family-specific dispersion varied greatly for both permanent and deciduous datasets, over 75% of the families exhibited greater dispersion with permanent traits. This suggests that morphological traits of the permanent dentition provide a less faithful reflection of biological relatedness than morphological traits of the deciduous dentition. Mantel tests indicate that both the deciduous and permanent distance matrices are significantly correlated with a matrix of genetic relatedness coefficients; however, the magnitude of the correlations was low. DISCUSSION Overall, morphological traits of permanent tooth crowns perform moderately well in distinguishing relatives from non-relatives, but deciduous crown variations may provide a more direct reflection of the underlying genetic structure of intra-site or intra-cemetery samples. These findings have implications for bioarchaeological research and biodistance practices. In particular, morphological traits of the deciduous dentition should be incorporated into standard data collection protocols because of their stronger signal of relatedness.

Heritability and genetic integration of anterior tooth crown variants in the South Carolina Gullah

OBJECTIVES This article presents estimates of narrow-sense heritability and bivariate genetic correlation for a series of morphological crown variants of the anterior dentition. These results provide insight into the value of dental phenotypes as evolutionary proxies, as well as the development of tooth crowns as integrated or modular structures. MATERIALS AND METHODS African American dental casts from the Menegaz-Bock collection were scored for a standard set of dental morphological variables using the Arizona State Dental Anthropology System. Estimates of narrow-sense heritability and genetic correlations were generated using SOLAR v. 8.1.1, controlling for the covariates of age, sex, and birth year. Analyses were run using ordinal/continuous scale variables that were then dichotomized at various breakpoints, consistent with standard practices in dental anthropology. RESULTS Heritability estimates were low to moderate for most traits, and lower in magnitude than those reported for odontometric data from the same study sample. Only winging, canine shoveling, and canine double shoveling returned narrow-sense heritabilities that did not differ significantly from zero. Genetic correlations were high among antimeres and metameres and low for different traits scored on the same tooth crown. These results affirm standard data cleaning practices in dental biodistance. Double shoveling was atypical in returning strong negative correlations with other traits, shoveling in particular. CONCLUSIONS Additive genetic variation contributes to dental morphological variation, although the estimates are uniformly lower than those observed for odontometrics. Patterns of genetic correlation affirm most standard practices in dental biodistance. Patterns of negative pleiotropy involving lingual and labial crown features suggest a genetic architecture and developmental complex that differentially constrain morphological variation of distinct surfaces of the same tooth crown. These patterns warrant greater consideration and cross-population validation.

A morphometric approach to Butler's field concept: The deciduous and permanent molars

Leprosy is one of the few specific infectious diseases that can be studied in bioarchaeology due to its characteristic debilitating and disfiguring skeletal changes. Leprosy has been, and continues to be, one of the most socially stigmatising diseases in history, over-riding all other aspects of social identity for the sufferers and frequently resulting in social exclusion. This study examines the stable isotopic evidence of mobility patterns of children, adolescents, and young adult individuals with the lepromatous form of leprosy in Medieval England (10 th –12 th centuries AD) to assess whether the individuals buried with the disease were non-locals, possibly from further afield. Enamel samples from 19 individuals from the St. Mary Magdalen Leprosy Hospital, Winchester (UK) were selected for strontium ( 87 Sr/ 86 6U DQG R[\JHQ į 18 O) stable isotope analysis based on age at death (<30 years), the presence of bone changes associated with lepromatous leprosy, and the underlying geology of their burial locations. The results from these data indicate that the St. Mary Magdalen Leprosy Hospital received an almost equal mixture of local and non-local individuals from further afield, including early pilgrims. At present, the St. Mary Magdalen Leprosy Hospital is the earliest dedicated leprosaria found within Britain and mobility studies such as these can help elucidate and test some of the broader historical notions and identities associated with the movements of those infected with the disease in Medieval England.