广西化石猩猩牙齿釉质厚度研究

华南和东南亚发现大量更新世的猩猩牙齿化石。本研究应用CT扫描三维重建的技术方法研究了广西更新世化石猩猩牙齿釉质厚度,并与现生类人猿、现代人、化石类人猿以及早期人类进行比较分析。结果显示:广西猩猩同类牙齿的釉质厚度与牙齿大小相关性很小;臼齿和前臼齿釉质厚度在上下颌之间不存在显著性差异;来自广西不同地区的猩猩化石牙釉质厚度无显著差异。与早期人科成员相比,广西猩猩的牙釉质相对较薄,平均与相对釉质厚度值都明显小于南方古猿、傍人。与早期人属相比,小于直立人、尼人以及非洲和欧洲的早期人属化石。与现代人和现生灵长类相比,广西化石猩猩釉质厚度明显大于大部分猴类和非洲大猿;平均釉质厚度稍大于现生猩猩,而与现代人更为接近;相对釉质厚度小于现代人,而与现生猩猩差异不大,都属于偏厚型釉质。本文讨论了釉质厚度与系统分类演化、食性适应的相关问题,作者推测釉质厚度可能是物种的特征属性,与牙齿功能适应有密切关联。     

Individual variation in enamel structure of human mandibular first premolars

Nine human mandibular first premolars were examined to assess variation in external morphology and enamel structural organization within a tooth type. The relationship of enamel ultrastructure to gross dental morphology was also studied. The teeth were cut in the mesiodistal direction just lingual to the buccal cusp, and etched. Montages were constructed of the cut enamel surface photographed in the scanning electron microscope at 100 X magnification. Parameters were measured and correlation coefficients were calculated for the comparison of various odontometric features. The mesiodistal and buccolingual dimensions were highly correlated and the occlusal thickness of enamel was significantly correlated to crown height but not crown width. Hunter-Schreger bands were less pronounced in fossa areas than at lateral aspects, cusps, or ridges; these bands were directly related to the geometry of the tooth. It was concluded that within this tooth type, there is a large amount of individual variation not only in gross morphology but also in enamel ultrastructure. This result underscores the fact that interspecific comparisons must be made with care.     

Measurements of tooth wear among Australian Aborigines: I. Serial loss of the enamel crown

The dental casts made from Aboriginal children during the course of a longitudinal growth study in Central Australia provided material for analyzing tooth wear under known environmental conditions. The wear facets produced on the occlusal surfaces were clearly preserved on the dental stone casts and recorded the progress of enamel attrition from ages 6 to 18. These casts were photographed and traced by electronic planimetric methods that automatically recorded the location and size of wear facets on the first and second permanent molars. These areas of worn tooth surface were compared to the total tooth surface. The worn surface was regressed on age to calculate wear rates of each tooth. Discriminant analyses were also performed to determine the significance of dental attrition differences between the sexes at each age group. The total wear on each tooth was highly correlated with age as expected but females wore their teeth at a significantly higher rate than males. The mandibular molars wore more rapidly than maxillary teeth in both sexes. The discriminant analysis successfully grouped 91% of the cases according to age and sex. Pattern of wear, the location, and size of wear facets also differed between age groups and sex. The questions of why there is a difference between male and female wear or why there is greater wear on one arch or arch region have no ready answers. The differing rates and pattern of dental wear do suggest that arch shape and growth rates may be the answer though it has yet to be tested. However, the occlusal surface wear is useful for age estimation in a population and provides a record of shifting masticatory forces during growth.     

Dental pathologies in ten free-ranging chimpanzees from Gombe National Park, Tanzania

The dental remains of ten adult chimpanzees from Gombe National Park, Tanzania, were examined for enamel attrition, caries, abscesses, periodontal disease, and tooth loss. Age was the underlying factor in the development of dental pathology, in that enamel wear was present to some extent in all ten but was uniformly severe only in the three for whom estimated age at death was 39-43 years. In turn, enamel wear appears to have been the direct cause of abscess development, periodontal disease, and tooth loss. Periodontal disease was commonly expressed as alveolar resorption, particularly around the premolars and molars. This involvement was variable in all except the two youngest. Some interesting wear patterns were evident in the form of deep grooves in the upper incisors and dramatic notching of the lower canines. These patterns, and enamel attrition in general, were attributed to normal mastication and to various stripping activities. Only one carious lesion was observed, in a male with an estimated age of 26 years. An accurate assessment of the actual prevalence of caries was obscured by enamel wear and tooth loss in the older individuals.     

Enamel thickness of deciduous and permanent molars in modern Homo sapiens

This study presents data on the enamel thickness of deciduous (dm2) and permanent (M1-M3) molars for a geographically diverse sample of modern humans. Measurements were recorded from sections through the mesial cusps of unworn teeth. Enamel is significantly thinner on deciduous than on permanent molars, and there is a distinct trend for enamel to increase in relative thickness from M1 to M3. The relatively thicker enamel of M2s and especially M3s can be related to the overall reduction in size of more distal molar crowns, which has been attained through a differential loss of the dentine component. Enamel tends to be thicker on the protocone than on the paracone, and thicker on the protoconid than on the metaconid, but its distribution is not wholly concordant with models that predict increased thickness as a means by which to counter heavier attritional loss on these "functional" cusps. Indeed, the thickness of enamel tends to be more variable on cusp tips and occlusal surfaces than over the lateral aspects of cusps. The proportionately thicker enamel over the lateral aspects of the protocone and protoconid more likely serves as a means to prolong functional crown life by preventing cusp fracture, rather than being an adaptation to increase the attritional longevity of wear facets. The present data suggest that the human dentition is not predisposed to develop a helicoidal wear plane through the disposition of molar enamel thickness.     

Dentition of moustached tamarins (Saguinus mystax mystax) from Padre Isla, Peru, part 1: quantitative variation

Analyses of dental variation in geographically restricted, wild populations of primates are extremely rare; however, such data form the best source for models of likely degrees of variation within and between fossil species. Data from dental casts of a geographically restricted population of moustached tamarins (Saguinus mystax mystax) from Padre Isla, Peru, document high levels of dental variability, as measured by coefficients of variation, in a nonsexually dimorphic species, despite its isolation and small population size. Like other primates, moustached tamarins show lower variability in the dimensions of the first molars and increased variability in the dimensions of the final molars in the toothrow. Moustached tamarins from Padre Isla have a distinctive pattern of variability in the remaining teeth, including more stable tooth lengths in the anterior and posterior portions of the toothrow, and more stable tooth widths in the midregion of the toothrow. High variability in incisor width may be due to age effects of a distinctive diet and pattern of dental wear.     

Enamel ultrastructure and masticatory function in molars of the American opossum, Didelphis virginiana

Maxillary and mandibular molars of the American opossum, Didelphis virginiana L., were viewed in the scanning electron microscope (SEM) after acid-etching or after cutting and acid-etching. Observations were made on enamel prism patterns as they relate to functional properties of the tooth at a particular site. Molars at different stages of wear were also observed under a dissecting microscope; worn surfaces were correlated with function and enamel ultrastructure. Pounding surfaces of molar cusps wear more rapidly than near-vertical shearing surfaces or crushing basins (i.e. the trigon and talonid basin). Pounding surfaces are subjected to abrasion by food and arc not normally involved in tooth-tooth contact. Near-vertical shearing surfaces and basins used for crushing do experience tooth-tooth contact, but are surprisingly more resistant to wear. Prisms at pounding sites approach the occlusal surface at a near 90° angle and are surrounded with very thick interprismatic (IP) enamel parallel to the occlusal surface of the tooth. The pounding pattern is present at tips of cusps and at occlusal surfaces of ridges of the tooth. At near-vertical shearing surfaces, the prisms approach the outer surface obliquely and are surrounded with IP crystals which are perpendicular to the vertical surface. The angle between prismatic and IP enamel in these patterns is 60–90° in a cervical to occlusal direction. In basins of the tooth used principally for crushing and some shearing, IP enamel is perpendicular to the changing slope of the basin and the prisms are usually at a 55–65° angle to the IP enamel. When the pounding and shearing-crushing patterns meet at a ridge, a distinct seam is observed. Pounding forces occur parallel to the long axis of the prisms and perpendicular to the thick IP enamel (i.e. perpendicular to the long axis of the IP crystals) lying on either side of the prisms. Shearing and crushing forces occur at an oblique angle to the prism, and interprismatic enamel is more evenly distributed about the prism. A spiral pattern is found at the bottoms of the trigon and talonid basins, but not at the bottom of the trigonid which is a non-occluding basin. It is concluded that the differential rates of wear of the enamel surfaces are necessary in maintaining the sharp cutting edges and effective crushing basins of the tribosphenic molar, and the ultrastructural arrangements of the enamel prisms are of functional significance.     

Tooth cusp sharpness as a dietary correlate in great apes

Mammalian molars have undergone heavy scrutiny to determine correlates between morphology and diet. Here, the relationship between one aspect of occlusal morphology, tooth cusp radius of curvature (RoC), and two broad dietary categories, folivory and frugivory, is analyzed in apes. The author hypothesizes that there is a relationship between tooth cusp RoC and diet, and that folivores have sharper teeth than frugivores, and further test the correlation between tooth cusp RoC and tooth cusp size. Eight measures of tooth cusp RoC (two RoCs per cusp) were taken from 53 M²s from four species and subspecies of frugivorous apes (Pongo pygmaeus, Pan troglodytes troglodytes, Pan troglodytes schweinfurthii, and Gorilla gorilla gorilla) and two subspecies of folivorous apes (Gorilla beringei beringei, and Gorilla beringei graueri). Phylogenetically corrected ANOVAs were run on the full dataset and several subsets of the full dataset, revealing that, when buccolingual RoCs are taken into account, tooth cusp RoCs can successfully differentiate folivores and frugivores. PCAs revealed that folivores consistently had duller teeth than frugivores. In addition, a weak, statistically significant positive correlation exists between tooth cusp size and tooth cusp RoC. The author hypothesizes differences in tooth cusp RoC are correlated with wear rates, where, per vertical unit of wear, duller cusps will have a longer length of exposed enamel ridge than sharper cusps. More data need to be gathered to determine if the correlation between tooth cusp RoC and tooth cusp size holds true when small primates are considered. Am J Phys Anthropol 153:226–235, 2014. © 2013 Wiley Periodicals, Inc.     

Enamel thickness and development in a third permanent molar of Gigantopithecus blacki

A ground section was prepared from a lower right M3 attributed to Gigantopithecus blacki as close as possible to axial plane of the mesial cusps. Daily cross striations were imaged, measured and counted in each cusp using polarised light microscopy. Long-period striae of Retzius were counted in the lateral enamel and their periodicity determined from counts and measurements of daily cross striations between adjacent striae. Cross striation spacings in the cusps were between 3.8 microm at the enamel dentine junction and 6 microm close to the enamel surface. Cuspal enamel formation times were long (800 days in the protoconid and 620 days in the metaconid). Linear enamel thickness was as much as 3.75 mm in the protoconid. There were 63 and 61 long-period striae of Retzius in the mesial aspects of the lateral enamel and the periodicity was 11 days. Lateral enamel formation took 1493 and 1291 days and when summed with cuspal enamel formation times totalled 4 years in the protoconid and 3.5 in the metaconid. Relative enamel thickness was 23, calculated through the mesial cusps. This falls short of that in the so-called 'thick hyper-thick' enamel described in 'robust' australopithecines to which Gigantopithecus blacki has previously been compared in both its dental and mandibular morphology. With respect to enamel thickness, therefore, Gigantopithecus blacki falls squarely among an increasingly large number of Miocene hominoids that can all be described as having 'thick enamel'.     

Intraspecific variation in M1 enamel development in modern humans: implications for human evolution

The timing and sequence of enamel development, as well as enamel thickness, was documented for individual cusps (protoconid, hypoconid, metaconid, entoconid) in 15 unworn permanent lower first molars (M(1)s) from a sample of modern human juveniles. These data were compared with previously published data for modern and fossil species reported in the literature. Crown formation in all teeth was initiated in the protoconid and completed in the hypoconid. These cusps had significantly longer formation times (2.91 and 2.96 yrs, respectively) than the metaconid and entoconid (2.52 and 2.38 yrs, respectively), as well as thicker enamel, and each represented between 92-95% of the total crown formation time. Rates of enamel secretion in all cusps increased significantly from 2.97 microm in the inner enamel to 4.47 microm in the outer enamel. Two cusps of one individual were studied in more detail and did not follow this typical trajectory. Rather, there was a sharp decrease in the middle of enamel formation and then a slow recovery of secretion rates from the mid- to outer enamel. This anomalous trajectory of enamel formation is discussed in the context of other nondental tissue responses to illness. Neither secretion rates nor periodicity differed significantly when compared between the cusps of each molar. Differences in cusp formation times, initiation, and completion suggest a relationship between the rates of enamel formation and enamel thickness. This fits with expectations about the mechanics of the chewing cycle and general lower molar morphology. A comparison with similar data for some nonhuman primates and fossil hominoids suggests this relationship may hold true across several primate taxa. Other aspects of enamel growth differed between this human sample and certain fossil species. The lower molars formed slowly over a longer period of time, which may reflect the extended growth period of modern humans. The methodological approach adopted in this study is discussed in the context of that used in other studies.     

To What Extent is Primate Second Molar Enamel Occlusal Morphology Shaped by the Enamel-Dentine Junction?

The form of two hard tissues of the mammalian tooth, dentine and enamel, is the result of a combination of the phylogenetic inheritance of dental traits and the adaptive selection of these traits during evolution. Recent decades have been significant in unveiling developmental processes controlling tooth morphogenesis, dental variation and the origination of dental novelties. The enamel-dentine junction constitutes a precursor for the morphology of the outer enamel surface through growth of the enamel cap which may go along with the addition of original features. The relative contribution of these two tooth components to morphological variation and their respective response to natural selection is a major issue in paleoanthropology. This study will determine how much enamel morphology relies on the form of the enamel-dentine junction. The outer occlusal enamel surface and the enamel-dentine junction surface of 76 primate second upper molars are represented by polygonal meshes and investigated using tridimensional topometrical analysis. Quantitative criteria (elevation, inclination, orientation, curvature and occlusal patch count) are introduced to show that the enamel-dentine junction significantly constrains the topographical properties of the outer enamel surface. Our results show a significant correlation for elevation, orientation, inclination, curvature and occlusal complexity between the outer enamel surface and the enamel dentine junction for all studied primate taxa with the exception of four modern humans for curvature (p<0.05). Moreover, we show that, for all selected topometrical parameters apart from occlusal patch count, the recorded correlations significantly decrease along with enamel thickening in our sample. While preserving tooth integrity by providing resistance to wear and fractures, the variation of enamel thickness may modify the curvature present at the occlusal enamel surface in relation to enamel-dentine junction, potentially modifying dental functionalities such as blunt versus sharp dental tools. In terms of natural selection, there is a balance between increasing tooth resistance and maintaining efficient dental tools. In this sense the enamel cap acts as a functional buffer for the molar occlusal pattern. In primates, results suggest a primary emergence of dental novelties on the enamel-dentine junction and a secondary transposition of these novelties with no or minor modifications of dental functionalities by the enamel cap. Whereas enamel crenations have been reported by previous studies, our analysis do not support the presence of enamel tubercles without dentine relief nuclei. As is, the enamel cap is, at most, a secondary source of morphological novelty.     

Metamerism,morphogenesis, and the expression of carabelli and other dental traits in humans

The patterning cascade model of tooth morphogenesis has emerged as a useful tool in explaining how tooth shape develops and how tooth evolution may occur. Enamel knots, specialized areas of dental epithelium where cusps initiate, act as signaling centers that direct the growth of surrounding tissues. For a new cusp to form, an enamel knot must form beyond the inhibition fields of other enamel knots. The model predicts that the number and size of cusps depends on the spacing between enamel knots, reflected in the spacing between cusps. Recently, work by our group demonstrated that the model predicted Carabelli trait expression in human first molars. Here we test whether differences in Carabelli trait expression along the molar row can also be predicted by the model. Crown areas and intercusp distances were measured from dental casts of 316 individuals with a digital microscope. Although absolute cusp spacing is similar in first and second molars, the smaller size and more triangular shape of second molars results in larger cusp spacing relative to size and, likely, less opportunity for the Carabelli trait to form. The presence and size of the hypocone (HY) and a range of small accessory cusps in a larger sample of 340 individuals were also found to covary with the Carabelli trait in a complex way. The results of this study lend further support to the view that the dentition develops, varies, and evolves as a single functional complex. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

Enamel structure and microwear: an experimental study of the response of enamel to shearing force.

The anisotropic fracturing and differential wear properties of enamel microstructure represent factors that can obscure the predictive relationship between dental microwear and diet. To assess the impact of enamel structure on microwear, this in vitro experimental study examines the relative contributions to wear of three factors: 1) species differences in microstructure, 2) direction of shearing force relative to enamel prisms and crystallites, and 3) size of abrasive particles. Teeth of Lemur, Ovis, Homo, and Crocodylus, representing, respectively, the structural categories of prismatic patterns 1, 2, and 3 and nonprismatic enamel, were abraded by shearing forces (forces having a component directed parallel to abraded surfaces) and examined by scanning electron microscopy. Striation width increased with particle size for nonprismatic, but not for prismatic, specimens. Direction of shear relative to prism and crystallite orientation had a significant influence on striation width in only some prismatic enamels. The different responses of prismatic and nonprismatic enamels to abrasion reflect the influence of structure, but at the level of organization of crystallites rather than prisms per se. Such interactions explain in part the inability of striation width to discriminate among animals with different dietary habits. Heteroscedasticity and deviations from normality also may confound parametric analyses of microwear variables. Variation in crystallite orientation in prismatic enamels may contribute to optimal dental function through the property of differential wear in functionally distinct regions of teeth.     

Variation in modern human premolar enamel formation times: implications for Neandertals

A recent study demonstrated that variation in enamel cap crown formation in the anterior teeth is greater than that in the molars from two geographically distinct populations: native indigenous southern Africans and northern Europeans. Eighty southern African and 69 northern European premolars (P3 and P4) were analyzed in the present study. Cuspal, lateral, and total enamel formation times were assessed. Although cuspal enamel formation times were not consistently different between the two populations, both lateral and total enamel formation times generally were. Bonferroni-corrected t-tests showed that southern Africans had significantly shorter lateral enamel formation time for five of the six cusps, as well as significantly shorter total enamel formation time for these same cusps. An analysis of covariance performed on the lingual cusps of the upper third and fourth premolars showed that differences in enamel formation times between these populations remained when crown height was statistically controlled. A further goal of this study was to ascertain, based on perikymata counts, what Neandertal periodicities would have to be in order for their teeth to have lateral enamel formation times equivalent to either southern Africans or northern Europeans. To this end, perikymata were counted on 32 Neandertal premolars, and the counts were inserted into regression formulae relating perikymata counts to periodicity for each population and each tooth type. Neandertal enamel formation times could be equivalent to those of southern Africans or northern Europeans only if their hypothetical periodicities fall within the range of periodicities for African apes and modern humans (i.e., 6-12 days). The analysis revealed that both populations could encompass Neandertal timings, with hypothetical periodicities based on the southern African population necessitating a lower range of periodicity (6-8 days) than those based on the northern European population (8-11 days).     

Variation in modern human enamel formation times

Most of what we know about the timing of human enamel formation comes from radiographic studies on children of known age. Here, we present new longitudinal data derived from a histological analysis of tooth enamel. Two samples, one from southern Africa and one from northern Europe, contained all anterior and molar tooth types. Two further samples contained only one tooth type: canines from a medieval Danish sample and third molars from a modern North American sample. Data were collected on 326 molars and 352 anterior teeth. Each tooth was sectioned and prepared for polarized light microscopy. We used daily enamel cross striations to determine cuspal enamel formation time, recorded the periodicity of long-period striae in the lateral enamel, and used this value to calculate enamel formation times for each decile of crown length. We present data that reveal some of the processes whereby differences in enamel formation times arise between our samples. Mean cuspal enamel formation times were similar in southern African and northern European anterior teeth, but differed in certain molar cusps. All the southern African anterior teeth completed enamel formation earlier. The greatest difference in mean chronological age at enamel completion was 5.2 vs. 6.2 years of age in lower canines. However, enamel completion times in the molar teeth showed few differences between the samples, with mean times for the longest forming cusps all falling between 3.0 years and 3.45 years. Our data suggest fewer differences between samples and smaller ranges of variation than in many radiographic studies and present a more realistic picture of worldwide variation in enamel formation times.     

Patterns of dental wear and the role of the canine in Cercopithecinae.

The importance of dental wear patterns in understanding masticatory functions in primates has long been appreciated. However, studies of wear patterns among populations of nonhuman primates are few. The purpose of this investigation is to establish the developmental aspects of dental wear in the Cercopithecinae and to describe certain relevant morphological traits. Studies were made of dental casts from 200 primate specimens of Macaca nemestrina, Macaca mulatta, and Papio cynocephalus. These casts were taken at four-month intervals, beginning at two years of age and continuing over a period of six to seven years. The wear pattern starts with the rounding and eventual flattening of the protoconid and protocone of the erupted first molars. Once this stage is reached, the hypoconid and metaconid of the mandibular, and the hypocone and paracone of the maxillary molars are rounded and eventually flattened. This pattern is maintained until the cusp tips are removed and the dentin exposed, however, the entoconid and metacone are not subjected to significant wear at this stage. Analysis of these dental casts and museum specimens has provided data on the development of dental wear during the maturation of these primates. The distribution of forces acting upon the teeth produce diagnostic patterns of wear, which provide evidence of the force location and magnitude. In examining the data, the hypothesis of canine guidance and its limitation of mandibular motion was evaluated. Specimens whose canines were removed demonstrate that the canines play no significant role in the development or maintenance of dental wear planes.     

Nanoindentation of lemur enamel: an ecological investigation of mechanical property variations within and between sympatric species

The common morphological metrics of size, shape, and enamel thickness of teeth are believed to reflect the functional requirements of a primate's diet. However, the mechanical and material properties of enamel also contribute to tooth function, yet are rarely studied. Substantial wear and tooth loss previously documented in Lemur catta at the Beza Mahafaly Special Reserve suggests that their dental morphology, structure, and possibly their enamel are not adapted for their current fallback food (the mechanically challenging tamarind fruit). In this study, we investigate the nanomechanical properties, mineralization, and microstructure of the enamel of three sympatric lemur species to provide insight into their dietary functional adaptations. Mechanical properties measured by nanoindentation were compared to measurements of mineral content, prism orientation, prism size, and enamel thickness using electron microscopy. Mechanical properties of all species were similar near the enamel dentin junction and variations correlated with changes in microstructure (e.g., prism size) and mineral content. Severe wear and microcracking within L. catta's enamel were associated with up to a 43% reduction in nanomechanical properties in regions of cracking versus intact enamel. The mechanical and material properties of L. catta's enamel are similar to those of sympatric folivores and suggest that they are not uniquely mechanically adapted to consume the physically challenging tamarind fruit. An understanding of the material and mechanical properties of enamel is required to fully elucidate the functional and ecological adaptations of primate teeth.     

Reduced tooth size in 45,X (Turner syndrome) females

Mean values and variances of deciduous and permanent tooth dimensions were compared between 121 45,X (Turner syndrome) females and 171 control subjects to clarify the role of the X chromosome on dental development. Although deciduous molars tended to be smaller than normal in 45,X females, there was no evidence of a reduction in tooth size for deciduous anterior teeth. In the permanent dentition, all mesiodistal dimensions were significantly smaller in 45,X females but only some of the buccolingual dimensions were smaller. The findings for deciduous tooth-size may reflect a sampling effect related to the extremely high frequency of spontaneous abortion in 45,X individuals. Results for permanent teeth are consistent with the concept of a decrease in enamel thickness in 45,X females.