Enamel thickness per masticatory phases (ETMP): A new approach to assess the relationship between macrowear and enamel thickness in the human lower first molar

Maxillary molar enamel thickness of Plio-Pleistocene hominins

Enamel thickness variation of deciduous first and second upper molars in modern humans and Neanderthals

The significance of a gradient in enamel thickness along the human permanent molar row has been debated in the literature. Some attribute increased enamel thickness from first to third molars to greater bite force during chewing. Others argue that thicker third molar enamel relates to a smaller crown size facilitated by a reduced dentin component. Thus, differences in morphology, not function, explains enamel thickness. This study draws on these different interpretive models to assess enamel thickness along the entire human deciduous tooth row. Average enamel thickness (AET), the area and proportion of crown enamel and dentin, and a crown size proxy are calculated for incisors, canines, and molars. Allometric scaling relationships are assessed within each tooth class, and then comparisons are undertaken along the row. Generally, AET was correlated with crown size and scaled with isometry, except for second molars which scaled with positive allometry. Mean AET increased along the row and was greater on molars, where bite forces are reported to be higher. Second molars combined the largest crown size with the thickest enamel and the smallest proportion of dentin, which is consistent with a reduction in the potential for cusp fracture under high bite forces. Resistance to wear may also account for some enamel thickness variation between tooth classes. Dental reduction did not explain the trend in AET from central to lateral incisors, or from first to second molars. The gradient in AET along the deciduous tooth row is partly consistent with a functional interpretation of enamel thickness.

Dental enamel thickness continues to feature prominently in anthropological studies of ape and human evolution, as well as studies of preventative oral care and treatment. Traditional studies of enamel thickness require physical sectioning of teeth for linear and scaled measurements. Recent applications of microtomographic imaging allow scientists to employ larger and more diverse samples, including global samples of recent humans as well as fossil hominin teeth. Unfortunately, little is known about the degree of enamel thickness variation among human populations, particularly across the dentition. This study employed microtomography to virtually image, section, and quantify the average enamel thickness of a sample of clinically extracted Indonesian canine and premolar teeth. This virtual sample was compared to physically sectioned African and European teeth. The results demonstrate that average enamel thickness is similar among human dentitions; no significant differences were detected within tooth positions, which is surprising given developmental differences between European and African canines and premolars. When populations were combined, differences were found in average enamel thickness between maxillary and mandibular premolars, and between canines and premolars within both dental arcades. This finding is potentially due to differences in premolar morphology and a trend of increasing enamel thickness distally throughout the dentition. The finding of limited population variation within tooth positions and significant variation between tooth positions is consistent with previous two-dimensional and three-dimensional studies of human molar enamel thickness. Average enamel thickness in canines and premolars does not differ between the sexes in our sample, although male teeth tend to have larger enamel and dentine cross-sectional areas, enamel–dentine junction lengths, and bi-cervical diameters. Males have significantly greater dentine area and enamel–dentine junction length than females for maxillary canines and premolars. The results of this study suggest that enamel thickness values in mixed-populations of humans are appropriate for comparisons with fossil hominins.

Enamel thickness has been linked to functional aspects of masticatory biomechanics and has been demonstrated to be an evolutionary plastic trait, selectively responsive to dietary changes, wear and tooth fracture. European Late Paleolithic and Mesolithic hunter-gatherers mainly show a flat wear pattern, while oblique molar wear has been reported as characteristic of Neolithic agriculturalists. We investigate the relationships between enamel thickness distribution and molar wear pattern in two Neolithic and medieval populations. Under the assumption that dietary and/or non-dietary constraints result in directional selective pressure leading to variations in enamel thickness, we test the hypothesis that these two populations will exhibit significant differences in wear and enamel thickness patterns. Occlusal wear patterns were scored in upper permanent second molars (UM2) of 64 Neolithic and 311 medieval subadult and adult individuals. Enamel thickness was evaluated by microtomography in subsamples of 17 Neolithic and 25 medieval individuals. Eight variables describing enamel thickness were assessed. The results show that oblique molar wear is dominant in the Neolithic sample (87%), while oblique wear affects only a minority (42%) of the medieval sample. Moreover, in the Neolithic molars, where buccolingually directed oblique wear is dominant and greatest enamel lost occurs in the distolingual quadrant, thickest enamel is found where occlusal stresses are the most important-on the distolingual cusp. These results reveal a correlation between molar wear pattern and enamel thickness that has been associated to dietary changes. In particular, relatively thicker molar enamel may have evolved as a plastic response to resist wear.

3D enamel thickness in Neandertal and modern human permanent canines

Dental morphology and wear pattern of bamboo rats, genus Rhizomys (Rodentia: Spalacidae)

Tachyoryctes, the most derived genus within the tribe Tachyoryctini (Rhizomyinae, Spalacidae, Rodentia), is known from Pliocene to Recent. It comprises four species: the fossil T. pliocaenicus and T. konjiti and the extant T. splendens and T. macrocephalus. A detailed description of the morphology and dental wear pattern of the cheek teeth of T. splendens is provided based on the study of 297 skulls. For the first and second lower and upper molars, five stages of wear are recognized (0–4). The third lower and upper molars show four stages of wear (0–3). The dental wear pattern in T. splendens is comparable to that of the Pleistocene T. konjiti.

Enamel thickness trends in Plio-Pleistocene hominin mandibular molars

Modern human molar enamel thickness and enamel–dentine junction shape

Dental enamel thickness has received considerable attention in ecological models of the adaptive significance of primate morphology. Several authors have theorized that the degree of enamel thickness may reflect selective pressures related to the consumption of fallback foods (dietary items that may require complex processing and/or have low nutritional value) during times of preferred food scarcity. Others have speculated that enamel thickness reflects selection during mastication of foods with particular material properties (i.e., toughness and hardness). Orangutans prefer ripe fruit when available, but show interspecific and sex differences in the consumption of fallback foods (bark, leaves, and figs) and other preferred foods (certain seeds). Bornean orangutans (Pongo pygmaeus) have also been reported to masticate more mechanically demanding foods than Sumatran orangutans (Pongo abelii). To test these ecological models, we assessed two-dimensional enamel thickness in orangutan full dentitions using established histological and virtual quantification methods. No significant differences in average enamel thickness (AET) were found between species. We found significant differences in the components of enamel thickness indices between sexes, with males showing greater enamel-dentine junction lengths and dentine core areas, and thus relatively thinner enamel than females. Comparisons of individuals of known sex and species revealed a dentition-wide trend for Bornean females to show greater AET than Sumatran females. Differences between small samples of males were less evident. These data provide only limited support for ecological explanations of enamel thickness patterns within great ape genera. Future studies of dietary ecology and enamel thickness should consider sex differences more systematically.

Molar enamel thickness is a key feature in the study of hominid evolution. Our understanding of enamel thickness and distribution patterns, however, has so far been based mostly on the limited information available from physical cross sections of the crown. In this study, the 3-dimensional (3D) whole crown enamel distribution pattern was explored in 74 extant great ape and modern human molars. Serial cross sections obtained from microfocal X-ray computed tomography were used to generate digital molar reconstructions at 50 to 80 micron voxel resolution, each crown represented by two to five million voxels. Surface data of both enamel dentine junction (EDJ) and outer enamel were extracted to derive volumetric measures, surface areas, curvilinear distances, and whole crown radial thickness maps. Three-dimensional average enamel thickness (AET) was defined as enamel volume divided by EDJ surface area. In 3D AET relative to tooth size, Homo exhibited the thickest, Gorilla the thinnest, and Pan and Pongo intermediately thick enamel. This result differs from previous claims that molar enamel of Pongo is relatively thicker than that of Pan. The discrepancy between three and two-dimensional (2D) values of AET stems from a combination of local differences in within tooth enamel distribution pattern and EDJ topography between Pan and Pongo molars. It demonstrates that 2D AET is not an appropriate substitute or estimator of whole crown AET. Examination of whole crown 3D distributions of molar enamel revealed a pattern common to all four examined species, the “functional” side of the molar having thicker enamel than the opposite side. However, some unique aspects of each species were also apparent. While the Gorilla molar has relatively thin enamel throughout its crown, Pan molars are characterized by particularly thin enamel in the occlusal fovea, and Pongo molars by an accentuation of relatively thin basal and thick occlusal enamel. Human molars are characterized by relatively thick enamel throughout the crown, with relatively large contrasts between buccal and lingual, and between mesial and distal crown portions. The ancestral condition common to the four extant species can be estimated by interpreting molar enamel distribution patterns unique to each genus as likely to be derived. We hypothesize that the last common ancestor likely had intermediately thick enamel, without particular thickening (or thinning) of enamel either occlusally or basally.

Enamel thickness is highly susceptible to natural selection because thick enamel may prevent tooth failure. Consequently, it has been suggested that primates consuming stress-limited food on a regular basis would have thick-enameled molars in comparison to primates consuming soft food. Furthermore, the spatial distribution of enamel over a single tooth crown is not homogeneous, and thick enamel is expected to be more unevenly distributed in durophagous primates. Still, a proper methodology to quantitatively characterize enamel 3D distribution and test this hypothesis is yet to be developed. Unworn to slightly worn upper second molars belonging to 32 species of anthropoid primates and corresponding to a wide range of diets were digitized using high resolution microcomputed tomography. In addition, their durophagous ability was scored from existing literature. 3D average and relative enamel thickness were computed based on the volumetric reconstruction of the enamel cap. Geometric estimates of their average and relative enamel-dentine distance were also computed using 3D dental topography. Both methods gave different estimations of average and relative enamel thickness. This study also introduces pachymetric profiles, a method inspired from traditional topography to graphically characterize thick enamel distribution. Pachymetric profiles and topographic maps of enamel-dentine distance are combined to assess the evenness of thick enamel distribution. Both pachymetric profiles and topographic maps indicate that thick enamel is not significantly more unevenly distributed in durophagous species, except in Cercopithecidae. In this family, durophagous species such as mangabeys are characterized by an uneven thick enamel and high pachymetric profile slopes at the average enamel thickness, whereas non-durophagous species such as colobine monkeys are not. These results indicate that the distribution of thick enamel follows different patterns across anthropoids. Primates might have developed different durophagous strategies to answer the selective pressure exerted by stress-limited food.

Dental tissue proportions and enamel thickness in Neandertal and modern human molars

Orangutans (Pongo) are the only great ape genus with a substantial Pleistocene and Holocene fossil record, demonstrating a much larger geographic range than extant populations. In addition to having an extensive fossil record, Pongo shows several convergent morphological similarities with Homo, including a trend of dental reduction during the past million years. While studies have documented variation in dental tissue proportions among species of Homo, little is known about variation in enamel thickness within fossil orangutans. Here we assess dental tissue proportions, including conventional enamel thickness indices, in a large sample of fossil orangutan postcanine teeth from mainland Asia and Indonesia. We find few differences between regions, except for significantly lower average enamel thickness (AET) values in Indonesian mandibular first molars. Differences between fossil and extant orangutans are more marked, with fossil Pongo showing higher AET in most postcanine teeth. These differences are significant for maxillary and mandibular first molars. Fossil orangutans show higher AET than extant Pongo due to greater enamel cap areas, which exceed increases in enamel-dentine junction length (due to geometric scaling of areas and lengths for the AET index calculation). We also find greater dentine areas in fossil orangutans, but relative enamel thickness indices do not differ between fossil and extant taxa. When changes in dental tissue proportions between fossil and extant orangutans are compared with fossil and recent Homo sapiens, Pongo appears to show isometric reduction in enamel and dentine, while crown reduction in H. sapiens appears to be due to preferential loss of dentine. Disparate selective pressures or developmental constraints may underlie these patterns. Finally, the finding of moderately thick molar enamel in fossil orangutans may represent an additional convergent dental similarity with Homo erectus, complicating attempts to distinguish these taxa in mixed Asian faunas.