Associations between Carabelli trait and cusp areas in human permanent maxillary first molars

Few dental anthropological studies have investigated the associations between tooth crown size and crown traits in humans using quantitative methods. We tested several hypotheses about overall crown size, individual cusp areas, and expression of Carabelli cusps in human permanent first molars by obtaining data from standardized occlusal photographs of 308 Australians of European descent (171 males and 137 females). Specifically, we aimed to calculate the areas of the four main molar cusps, and also Carabelli cusp, and to compare the relative variability of cusp areas in relation to timing of development. We also aimed to compare cusp areas between males and females and to describe how Carabelli cusp interacted with other molar cusps. Measurements included maximum crown diameters (mesiodistal and buccolingual crown diameters), the areas of the four main cusps, and the area of Carabelli cusp. The pattern of relative variability in absolute areas of molar cusps corresponded with their order of formation, the first-forming paracone displaying the least variation, and the last-forming Carabelli cusp showing the greatest. Overall crown size and areas of individual cusps all showed sexual dimorphism, with values in males exceeding those in females. Sexual dimorphism was smallest for paracone area and greatest for Carabelli cusp area. Overall crown size and cusp areas were larger in individuals displaying a Carabelli cusp, especially the hypocone area. Although the combined area of the protocone and a Carabelli cusp was greater in cuspal forms than noncuspal forms, protocone area alone was significantly smaller in the former. Our findings lead us to propose that, in individuals with the genotype for Carabelli trait expression, larger molar crowns are more likely to display Carabelli cusps, whereas molars with smaller crowns are more likely to display reduced forms of expression of the trait. We suggest that the pattern of folding of the internal enamel epithelium in developing molar crowns, particularly in the protocone region, can be modified by a developing Carabelli cusp.     

Study of the cuspal ridges of the upper first molars in a modern Japanese population

Materials used were dental casts of the upper first molars of modern Japanese subjects, comprising 29 males and 25 females. Their molar occlusal surfaces were photographed by moiré contourography using the standard trigonal plane. The ridges of a cusp, comprising a central ridge and mesial and distal accessory ridges, were identified from the patterns of the moiré fringes. The central ridge was observed in all cusps except for the hypocone in both sexes. Frequencies of the mesial and distal accessory ridges of trigonal cusps were over 90% except for the distal accessory ridge of the metacone, and those of the hypocone were under 25% in both sexes. These values were generally higher in males than in females, especially for the distal accessory ridge of the metacone. The running pattern of the cuspal ridges showed little difference between sexes. The oblique ridge which was higher than the central groove formed a saddle-like structure. This ridge was observed in all materials, but its heights and structural components varied remarkably. In this study, the distal accessory ridge of the metacone was found to be incorporated into the oblique ridge in about 13% of cases. Variability in the running pattern of the ridges within a single cusp was highest in the hypocone and lowest in both the paracone and protocone. The results obtained are considered to represent the stability or reductive tendency of cusps in the upper first molars.     

Genetic integration of molar cusp size variation in baboons

Many studies of primate diversity and evolution rely on dental morphology for insight into diet, behavior, and phylogenetic relationships. Consequently, variation in molar cusp size has increasingly become a phenotype of interest. In 2007 we published a quantitative genetic analysis of mandibular molar cusp size variation in baboons. Those results provided more questions than answers, as the pattern of genetic integration did not fit predictions from odontogenesis. To follow up, we expanded our study to include data from the maxillary molar cusps. Here we report on these later analyses, as well as inter‐arch comparisons with the mandibular data. We analyzed variation in two‐dimensional maxillary molar cusp size using data collected from a captive pedigreed breeding colony of baboons, Papio hamadryas, housed at the Southwest National Primate Research Center. These analyses show that variation in maxillary molar cusp size is heritable and sexually dimorphic. We also estimated additive genetic correlations between cusps on the same crown, homologous cusps along the tooth row, and maxillary and mandibular cusps. The pattern for maxillary molars yields genetic correlations of one between the paracone–metacone and protocone–hypocone. Bivariate analyses of cuspal homologues on adjacent teeth yield correlations that are high or not significantly different from one. Between dental arcades, the nonoccluding cusps consistently yield high genetic correlations, especially the metaconid–paracone and metaconid–metacone. This pattern of genetic correlation does not immediately accord with the pattern of development and/or calcification, however these results do follow predictions that can be made from the evolutionary history of the tribosphenic molar. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

现代中国人群形成与分化的形态证据——亚洲与非洲现代人群上颌第一臼齿齿尖相对面积的对比分析

现代人群形成与分化造成的具有地域差别的人群标志性体质特征是研究人群之间关系的重要信息。作为臼齿形态测量的组成部分,上颌第一臼齿齿尖相对面积在人类演化以及现代人群关系上的研究价值已经引起了人类学界的关注。然而,迄今对上颌第一臼齿齿尖相对面积在世界各地的现代人群之间是否具有地域性差异还不是很清楚。本文对代表亚洲和非洲现代人群的208枚上颌第一臼齿齿尖相对面积的研究发现,上颌第一臼齿四个主要齿尖相对面积在亚洲和非洲现代人群之间都存在明显差异。亚洲现代人上颌第一臼齿近中齿尖(原尖和前尖)相对面积较大,而远中齿尖(后尖和次尖)相对面积较小。非洲现代人上颌第一臼齿各齿尖相对面积均较亚洲人群具有更大的变异,尤其在原尖和次尖相对面积上。采用判别分析可以将69.2%的标本正确地判别归入其原来所属的组群。通过与化石人类相关数据的对比发现,非洲现代人比亚洲现代人在原尖和次尖相对面积以及前尖/后尖相对面积比例上更加接近于人类演化的早期形式。现有的证据显示,亚洲和非洲现代人群在齿尖相对面积的分化至少可以追溯到全新世早期,更加精确的分化时间需要结合更新世晚期甚至中期化石人类数据去获得。本研究揭示出的非洲现代人上颌第一臼齿齿尖相对面积的高异质性(heterogeneity)和相对原始性在现代人形成与分化方面的作用尚需进一步的研究。     

Intercusp relationships of the permanent maxillary first and second molars in American whites

Much of a human molar's morphology is concentrated on its occlusal surface. In view of embryologists' recent attention on the determination of crown morphology by enamel knots that initiate cusp formation, we were interested in the arrangement of cusp apices in the definitive tooth. Computer-assisted image analysis was used to measure intercusp distances and angles on permanent maxillary M1 and M2 in a sample of 160 contemporary North American whites. The intent was to generate normative data and to compare the size and variability gradients from M1 to M2. There is little sexual dimorphism in intercusp distances or angles, even though the conventional mesiodistal (MD) and buccolingual (BL) crown size is 2.0% and 4.0% larger in males, respectively, in these same teeth. Dimensions decreased in size and increased in variability from M1 to M2, but differentially. Cusps of the trigon were more stable between teeth, especially the paracone-protocone relationship. Principal components analysis on the six M1 distances disclosed only one eigenvalue above 1.0, indicating that overall crown size itself is the paramount controlling factor in this tooth that almost invariably exhibits a hypocone. In contrast, four components were extracted from among the 12 angular cusp relationships in M1. These axes of variation may prove useful in studies of intergroup differences. A shape difference occurs in M2, depending on whether the hypocone is present; when absent, the metacone is moved lingually, creating more of an isosceles arrangement for the cusps of the trigon. Statistically, correlations are low between occlusal intercusp relationships and conventional crown diameters measured at the margins of the crowns that form later. Weak statistical dependence between cusp relationships and traditional MD and BL diameters suggest that separate stage- and location-specific molecular signals control these different parts (and different stages) of crown formation.     

A new molar from the Middle Pleistocene hominid assemblage of Yanhuidong,Tongzi, South China

1988年,贵州省博物馆对桐梓岩灰洞的支洞进行了最后一次发掘。2014年,在洞内第四层堆积物中鉴定出逾2000件的动物牙齿化石,以及一枚古人类上颊齿(编号:TZ-1)。1991年,铀系法测定这些次生堆积物的沉积年代约为距今24万年。本文运用高精度CT(巴黎自然历史博物馆)对TZ-1的釉质齿质界面(EDJ)和牙髓腔几何形态进行了分析。TZ-1的冠面形态有如下特征:次尖小且在远中舌侧不发育,咀嚼面轮廓呈四边形,颊舌径稍大过近中远中径,原尖舌侧齿带发育,齿尖从大到小依次为原尖、后尖、前尖和次尖。TZ-1牙髓腔的髓角与其釉质齿质界面以及釉质表面的形态都具有相关性。TZ-1的形态与M1虽有相似之处,但完全不同于1983年出土于同一层位的另两颗M1;其应被鉴定为dm2,并可被归入中中更新世的中国直立人支系。岩灰洞上臼齿PA 875的形态与建始龙骨洞PA 1279和周口店直立人等古老型直立人相似。岩灰洞另一枚刚萌发的臼齿PA 874具有凸出的次尖和长菱形的外廓,接近于爪哇型直立人Sangiran NG 91-G10;这也是智人和尼安德特人的共有衍征。但PA 874的冠面仍保留了亚洲型的齿带,因而被归入人属未定种。因出土自次生堆积,岩灰洞的三种古人类类型未必曾同时并存,但却揭示了华南地区人类演化进程中的多样面貌。     

记江苏泗洪首次发现森林古猿类化石

本文记述了在江苏泗洪松林庄发现的一种古猿类化石,它以个体小、颊齿宽、有发达的齿带等特征有别于我国云南的腊玛古猿、西瓦古猿;它也有别于在同一地点、同一层位发现的双沟醉猿。其形态与非洲的Proconsul属接近,根据这些形态特点和它的地史分布,我们订立了一新属一新种:Platodontopithecus jianghuaiensis,地质时代为中新世。     

全新世中国北方人群第一臼齿齿冠面积和齿尖相对面积的测量分析

通过对532枚中国北方地区新石器时代到青铜时代和铁器时代人类上、下颌骨第一臼齿(M1和M1)齿冠面积和齿尖面积的测量和分析, 得到以下关于中国北方近代人群牙齿尺寸的准确数据和变异情况的结果: 两性M1和M1齿冠面积存在显著性差异, 男性齿冠面积大于女性齿冠面积, 齿尖相对面积不存在性别差异; 从新石器时代到青铜时代男性M1和M1分别缩小了1.16%和4.96%, 女性分别缩小了5.68%和6.11%,青铜时代到铁器时代齿冠面积无明显变化; M1颊侧尖相对基底面积大于舌侧尖, 形成齿尖大小差异的原因很可能是在人类演化过程中, 齿冠面积整体缩小的趋势下, 为使咬合面最大化使用, 各齿尖朝着更利于扩大咬合面基底面积的方式进化的结果; 世界各地区近代人群M1和M1各齿尖大小顺序基本一致, 其中M1为原尖>前尖>后尖>次尖, M1下原尖最大, 下次小尖最小, 其他三个齿尖面积非常接近, 不存在明显大小变化。     

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.     

Reduction of maxillary molars in Homo sapiens sapiens: a different perspective.

Crown and cusp areas, and buccolingual and mesiodistal diameters of maxillary molars of complete upper tooth rows (30 males, 30 females) were analysed in order to quantify changes in size and shape from the first to the third molar. Uni- and multivariate analyses revealed the mesial cusps, in particular the protocone (mesiolingual cusp), to be more stable than the other cusps. Although there is a gradient in size from the first to third molar, shape changes were found to be marked. Overall, the findings are in keeping with the field theory and the hypotheses of environmental constraints on later developing teeth. However, not all of the results could be entirely explained by these concepts. Functional aspects seem to account for the relative stability of the protocone and the buccolingual crown diameter. It appears that this functional complex is relatively stable despite the overall reduction of tooth size, which is probably secondary to processes occurring in the jaws and the cranium. This finding may have implications for studies on tooth reduction between populations of different time periods.     

Cusp size, sexual dimorphism, and heritability of cusp size in twins.

Overall measures of mandibular molars reflect the combined size contributions of the component cusps and ridges. Until now, the size hierarchy of primary and permanent mandibular molar cusps remained unclear. This paper utilizes the relative plane surface areas (basal area dimensions) of the individual molar cusps, as assays of cusp size to demonstrate cusp size variations within populations, antimere cuspal variations, sexual dimorphism, and, the heritability of cusp size. Duplicate dental casts from 199 pairs of like-sexed twins provide the raw dats. Defined anatomic landmarks on the occlusal surfaces were reduced to X-Y rectangular coordinates prior to the computation of the basal areas dimensions. The results establish a cusp size hierarchy specific for molar type, i.e., five-cusped molars with a distal fovea and distal marginal ridge (5fd), five-cusped molars without a distal fovea and without a distal marginal ridge (5o), and four-cusped molars (4c). Sexual dimorphism in cusp size is apparent in 5fd molar cusped but not in 5o molar cusps. However, males have a significantly higher frequency of 5fd molars. Females have a higher frequency of smaller 5o and 4c molars which have fewer crown components. Moreover, female 5o molars have cusps as large as or larger than 5o male molor cusps. Right-side-left-side differences exist between antimere cusps based on relatively low correlations. The mirroring of molor types occurs infrequently. When observed, most intrapair differences for cusp size, using F-ratios, indicate a low component of hereditary variability.     

Carabelli's trait and tooth size of human maxillary first molars

Carabelli's trait is a morphological feature that can occur on the protocone of human maxillary molars. This study tests the hypothesis that Carabelli's trait is correlated statistically with the dimensions of the crown's four principal cusps or whether, as a cingular feature, the trait truly accretes onto an otherwise unaffected crown. Computer-assisted image analysis was used to measure the 6 intercusp distances and 12 angular relationships among cusp tips on the permanent first molar of 300 young adult American whites. Carabelli's complex was scored using an 8-grade ordinal scheme. Crown size was quantified in three ways, namely as 1) maximum mesiodistal and buccolingual diameters, 2) the 6 intercusp distances, and 3) the 12 angular cusp arrangements. There was no sex difference in the morphological expression of Carabelli's trait in this sample. Overall crown size and intercusp distances were significantly and progressively larger in molars with larger Carabelli's trait expressions. There are graded size responses between crown size (mesiodistal and buccolingual diameters), sizes of the four principal cusps, and morphological stage of Carabelli's complex, though the statistical relationships are appreciably stronger in males than females. Carabelli's trait occurs preferentially in larger molars. In contrast, angular (shape) relationships among cusp tips are not discernibly affected by trait size in either sex. There is the situation, then, that Carabelli's trait is developmentally correlated with crown size, but with no apparent alteration of cusp arrangements, suggesting that the increases are isometric across the occlusal table. Why the association is much weaker in females remains speculative, but these data provide yet another line of evidence that, within a population, tooth size is associated in a positive fashion with crown complexity.     

豫鼠化石在我国阿尔金地区的发现

记述了在新疆巴音郭楞蒙古族自治州若羌县阿尔金山地区彩虹沟首次发现的豫鼠一新种:阿尔金豫鼠(Yuomys altunensis sp.nov.).其主要特征是臼齿的尺寸较大,比例上较宽,齿冠较高,臼齿的后小尖与后尖明显分开,后脊相对较长,但不完全,次尖明显小于原尖,舌侧内凹伸达臼齿齿冠基部,后齿带与后尖舌侧连;M3后尖为新月形,后齿带较短等.根据豫鼠臼齿的进化趋势和新种臼齿的尺寸较大、齿冠较高和舌侧内凹伸达齿冠基部的特征与Y.cavioides,Y.eleganes和Y.huangzhuangensis相近,但颊齿比例较宽判断,Y.altunensis可能与该3种处于同样的进化阶段或稍进步.其产出的地层时代很可能与它们相近或稍晚,即为晚中始新世或稍晚.     

Geometric morphometric 3D shape analysis and covariation of human mandibular and maxillary first molars

Dental casts of 160 Greek subjects (80 males, 80 females) were scanned by a structured‐light scanner. The upper and lower right first molar occlusal surface 3D meshes were processed using geometric morphometric methods. A total of 265 and 274 curve and surface sliding semilandmarks were placed on the upper and lower molar surfaces, respectively. Principal component analysis and partial least square analysis were performed to assess shape parameters. Molars tended to vary between an elongated and a more square form. The first two principal components (PCs), comprising almost 1/3 of molar shape variation, were related to mesiodistal–buccolingual ratios and relative cusp position. Distal cusps displayed the greatest shape variability. Molars of males were larger than those of females (2.8 and 3.2% for upper and lower molars respectively), but no shape dimorphism was observed. Upper and lower molar sizes were significantly correlated (r² = 0.689). Allometry was observed for both teeth. Larger lower molars were associated with shorter cusps, expansion of the distal cusp, and constriction of the mesial cusps (predicted variance 3.25%). Upper molars displayed weaker allometry (predicted variance 1.59%). Upper and lower molar shape covariation proved significant (RV = 17.26%, P < 0.0001). The main parameter of molar covariation in partial least square axis 1, contributing to 30% of total covariation, was cusp height, in contrast to the primary variability traits exhibited by PC1 and PC2. The aim of this study was to evaluate shape variation and covariation, including allometry and sexual dimorphism, of maxillary and mandibular first permanent molar occlusal surfaces. Am J Phys Anthropol 152:186–196, 2013. © 2013 Wiley Periodicals, Inc.     

A morphometric analysis of maxillary molar crowns of Middle-Late Pleistocene hominins

This study explores the significance of shape differences in the maxillary first molar crowns of Neandertals and anatomically modern humans. It uses morphometric analysis to quantify these differences and to investigate how the orientation of major cusps, relative cusp base areas and occlusal polygon area influence crown shape. The aims of this study were to 1) quantify these data to test whether the tooth shapes of Neandertals and anatomically modern humans differ significantly and 2) to explore if either of the shapes is derived relative to earlier fossil hominins. Data were collected from digital occlusal photographs using image-processing software. Cusp angles, relative cusp base areas and occlusal polygon areas were measured on Neandertals (n=15), contemporary modern humans (n=62), Upper Paleolithic humans (n=6), early anatomically modern humans (n=3) and Homo erectus (n=3). Univariate and multivariate statistical tests were used to evaluate the differences between contemporary modern humans and Neandertals, while the much sparser data sets from the other fossil samples were included primarily for comparison. Statistically significant differences reflecting overall crown shape and internal placement of the crown apices were found. Neandertals are distinguished from contemporary humans by possessing maxillary first molars that 1) are markedly skewed; 2) possess a narrower distal segment of the occlusal polygon compared to the mesial segment; 3) possess a significantly smaller metacone and a significantly larger hypocone; and 4) possess a significantly smaller relative occlusal polygon area reflecting internally placed cusps. Differences in relative cusp base areas of the hypocone and metacone may contribute to the shape differences observed in Neandertals. However, early anatomically modern humans possessing a pattern of relative cusp base areas similar to Neandertals lack their unusual shape. That the morphology observed in non-Neandertal fossil hominins is more anatomically modern human-like than Neandertal-like, suggests that this distinctive morphology may be derived in Neandertals.     

Ursavus在中国的首次发现——并记禄丰古猿化石产地的其他熊类标本

本文记述的是1981年在禄丰古猿化石产地发掘中发现的熊类标本,共有二属二种和一种暂不能确定属、种的类型。其中祖熊(Ursavus)过去只见于欧洲、北美和西亚的中新世地层,在我国还是首次发现。禄丰的标本,就其大小和特征可归于Ursavus depereti;印度熊(Indarctos)过去在我国发现过两个种:中国印度熊(I.sinensis)和拉氏印度熊(I.lagrelli),禄丰的标本为前一个种;第三类型与Protursus和Ursus都有相似之处,但又有区别,鉴于材料不足,暂不订属和种。     

Hypocone and metaconule: identification and variability on human molars

Hypocone and metaconule variability was studied in a large sample of upper molars belonging to several human populations. The identification of these cusps does not generally show problems. However, on second and especially on third molars, it is occasionally difficult to decide if a little cup located on the crown distal border is a hypocone or a metaconule. The identification of supernumerary cusps located between the metacone and the hypocone seldom is clarified. We suggest the utility of differentiating if such cusps proceed the hypocone or metacone division and of using metacone variability in dental anthropology studies. An objective classification of the metacone size is proposed to standardize observations. Paper presented at the 4^th congress of the European Anthropology Association (Florence, Sept. 1984).     

山西垣曲先炭兽类一新种

本文主要报道1983年9月,在山西省垣曲县寨里发现的一件较完整的先炭兽类老年个体上颌骨.带有颇为完整的左、右齿列. I~1-P~2 各齿前、后均有较长的齿隙. P~1、P~2、P~3 前的齿隙颌骨上,保留有乳前臼齿的齿槽.这块标本在大小及形态特征上,与目前所知先炭兽属 Anthracokeryx 中任何种都有相当大的差异.它代表了该属中的一个新种.     

Dentition and Jaw of Kokopellia juddi, a Primitive Marsupial or Near-Marsupial from the Medial Cretaceous of Utah

We add to the knowledge of the dentition and lower jaw of the primitive marsupial or near marsupial, Kokopellia juddi, based on newly collected materials from the medial Cretaceous (Albian–Cenomanian) of central Utah. The dental formula, i4 c1 p3 m4, is primitive for (or with respect to) Marsupialia, as are a number of features of the dentary and dentition: presence of a labial mandibular foramen, ?an inflected angle, ?and a trace of the meckelian groove; lack of “staggering” of the lower incisor series; lack of “twinning” between entoconid and hypoconulid on lower molars; incompletely lingual position of lower molar paraconid; upper molar protocone relatively small and mesiodistally narrow; and conules placed about halfway between the protocone and the paracone–metacone. Other than the stylocone, cusps are lacking from the stylar shelf; we argue that this represents the primitive marsupial condition based on the economy of character change and the stratigraphic record of marsupials in the Cretaceous of North America. Recent discoveries of early marsupials, eutherians, and therians of metatherian–eutherian grade provide data indicating that some derived features of the dentary and dentition (e.g., loss of coronoid, meckelian groove, and labial mandibular foramen; acquisition of strong, “winged” conules, double rank postvallum/prevallid shearing, and stylar cusp D) probably arose independently, in some cases more than once, among the major groups of tribosphenic mammals. In turn, this suggests that a common ancestor for marsupials and placentals was more primitive than has generally been appreciated.     

Evolution of the Tribosphenic Molar Pattern in Early Mammals,with Comments on the “Dual-Origin” Hypothesis

Development of the tribosphenic molar was a fundamental event that likely influenced the rise of modern mammals. This multi-functional complex combined shearing and grinding in a single chewing stroke, and provided the base morphology for the later evolution of the myriad dental morphologies employed by mammals today. Here a series of morphotypes are presented that represent stepwise acquisition of characters of the molar crown, in an effort to clarify homologies and functional analogies among molars of tribosphenic and tribosphenic-like mammals, as well as their putative sister groups. This is accomplished by evaluation of wear features, which provide direct evidence of occlusal function, and mapping these features on molars of the various morphotypes demonstrates their utility in determining homology. The original singular lower molar talonid cusp is homologous with the hypoconid, and upper molar cusp C in early mammals is homologous with the metacone (cusp “C” is a neomorph with variable occurrence). The lingual translation of the metacone to a position more directly distal to the paracone (as in Peramus) creates an embrasure for the lower molar hypoconid, and is accompanied by the development of the hypoconulid and a new shearing surface. Lastly, the Gondwanan radiation of tribosphenic-like mammals, the Australosphenida (including monotremes), is determined to be functionally non-tribosphenic. The Tribosphenida are restricted to Laurasian taxa, with an origin at or just prior to the Jurassic-Cretaceous boundary.