Brief communication: dental development and enamel thickness in the Lakonis Neanderthal molar

Developmental and structural affinities between modern human and Neanderthal dental remains continue to be a subject of debate as well as their utility for informing assessments of life history and taxonomy. Excavation of the Middle Paleolithic cave site Lakonis in southern Greece has yielded a lower third molar (LKH 1). Here, we detail the crown development and enamel thickness of the distal cusps of the LKH 1 specimen, which has been classified as a Neanderthal based on the presence of an anterior fovea and mid-trigonid crest. Crown formation was determined using standard histological techniques, and enamel thickness was measured from a virtual plane of section. Developmental differences include thinner cuspal enamel and a lower periodicity than modern humans. Crown formation in the LKH 1 hypoconid is estimated to be 2.6-2.7 years, which is shorter than modern human times. The LKH 1 hypoconid also shows a more rapid overall crown extension rate than modern humans. Relative enamel thickness was approximately half that of a modern human sample mean; enamel on the distal cusps of modern human third molars is extremely thick in absolute and relative terms. These findings are consistent with recent studies that demonstrate differences in crown development, tissue proportions, and enamel thickness between Neanderthals and modern humans. Although overlap in some developmental variables may be found, the results of this and other studies suggest that Neanderthal molars formed in shorter periods of time than modern humans, due in part to thinner enamel and faster crown extension rates.     

人类牙齿齿冠和齿根分离两种技术方法对牙釉质厚度测量的影响

在基于计算机断层扫描技术（CT）和虚拟图像处理技术的灵长类牙齿测量学研究中,经常需要分离三维虚拟模型的齿冠和齿根,再进行后续测量工作,如计算机辅助的生物力学分析、釉质厚度测量等。而分离齿冠和齿根这一步骤,目前有多种方法,如,1)根据齿颈线切分齿冠,或2)人工建立基底平面切分齿冠。为了评估这两种不同的处理方式对后续的牙齿测量学上的影响,本文使用三维方法测量了82例化石和现代人类下颌后部牙齿的釉质厚度,包括南方古猿、早期人属、尼安德特人和现代人。使用配对t检验对比发现,两种方法得到的釉质厚度数值上没有显著差别,但随后进行的种间比较发现,使用基底平面切分齿冠的方法比较费时,更依赖于测量者的人工操作,并且可能弱化了物种间前臼齿绝对釉质厚度的差异,造成系统误差。其原因是对于前臼齿和前部牙齿等齿颈线形状不规则的标本,基底平面难以建立或误差较大。在未来对釉质厚度的种间差异的研究中,特别对齿颈线形状不规则的标本（如人类前部牙齿及猩猩、黑猩猩的牙齿等）,本文推荐使用齿颈线分离齿冠和齿根,测量和计算齿颈线之上的釉质厚度。釉质厚度有一定的分类学、功能形态学和系统发育学意义。本文积累了一批可供未来对比研究的原始数据,并且发现尼安德特人前臼齿的相对釉质厚度显著小于现代人,这与前人利用臼齿、犬齿所做的对比研究结果相同,支持了尼安德特人拥有较薄的相对釉质厚度这一观点。     

Tissue proportions and enamel thickness distribution in the early Middle Pleistocene human deciduous molars from Tighenif,Algeria

The present study of three human upper deciduous molars from the early Middle Pleistocene site of Tighenif, Algeria, constitutes the first microtomographic-based endostructural exploration of African fossil teeth likely representative of the Homo heidelbergensis morph. Comparative morphological observations and 2-3D measurements describing subtle tooth organization (crown tissue proportions) and enamel thickness topography (site-specific distribution and global patterning) indicate that their virtual extracted structural signature better fits the modern human, rather than the Neanderthal condition. Accordingly, we predict that the inner structural morphology of the deciduous molars from the Middle Pleistocene western European series better fits the primitive, and not the derived Neanderthal figures.     

Enamel and dentine dimensions of the Pleistocene hominins from Atapuerca (Burgos,Spain): A comparative study of canine teeth

Enamel and dentin patterns have awakened a considerable interest in phylogenetic studies. However, almost nothing is known about the dental tissue proportions of European Pleistocene hominins, apart from Neanderthal populations. This study aims to assess the three-dimensional dental tissue proportions of permanent canines belonging to the extensive sample of hominin teeth at Sierra de Atapuerca (Spain) through the use of microtomographic techniques. Our results show that early and middle Pleistocene populations from Atapuerca exhibit large coronal and root dentine dimensions, as well as a thinly enamelled pattern, which has been traditionally considered an autapomorphic Neanderthal trait. Therefore, these results might support an early enamel thickness decrease which is already observed 800 kyr ago in Homo antecessor and maintained in later groups such as Sima de los Huesos and Neanderthal populations during the middle Pleistocene.     

Cervical and crown outline analysis of worn Neanderthal and modern human lower second deciduous molars

Despite the general increase in digital techniques for dental morphometric analyses, only a few methods are available to study worn teeth. Moreover, permanent dentitions are studied much more frequently than deciduous teeth. In this study, we address both issues by providing a taxonomic classification of Neanderthal and modern human (MH) lower second deciduous molars (dm₂s) through the analysis of crown and cervical outlines. Crown and cervical outlines were obtained from a three‐dimensional (3D) digital sample of uniformly oriented dm₂s. Both outlines were centered on the centroid of their area and represented by 16 pseudolandmarks obtained by equiangularly spaced radial vectors out of the centroid. We removed size information from the oriented and centered outlines with a uniform scaling of the pseudolandmark configurations to unit Centroid Size. Group shape variation was evaluated separately for the dm₂ crown and cervical outlines through a shape–space principal component (PC) analysis. Finally, quadratic discriminant analysis of a subset of PCs was used to classify the specimens. Our results demonstrate that both outlines successfully separate the two groups. Neanderthals showed a buccodistal expansion and convex lingual outline shape, whilst MHs have buccodistal reduction and straight lingual outline shape. Therefore, we confirmed that the cervical outline represents an effective parameter for distinguishing between the two taxa when dealing with worn or damaged dm₂s. Am J Phys Anthropol 2012. © 2012 Wiley Periodicals, Inc.     

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

The thickness of dental enamel is often discussed in paleoanthropological literature, particularly with regard to differences in growth, health, and diet between Neandertals and modern humans. Paleoanthropologists employ enamel thickness in paleodietary and taxonomic studies regarding earlier hominins, but variation in enamel thickness within the genus Homo has not been thoroughly explored despite its potential to discriminate species and its relevance to studies of growth and development. Radiographic two-dimensional studies indicate that Neandertal molar enamel is thin relative to the thick enamel of modern humans, although such methods have limited accuracy. Here we show that, measured via accurate high-resolution microtomographic imaging, Neandertal molar enamel is absolutely and relatively thinner than modern human enamel at most molar positions. However, this difference relates to the ratio of coronal dentine volume to total crown volume, rather than the quantity of enamel per se. The absolute volume of Neandertal molar enamel is similar to that of modern humans, but Neandertal enamel is deposited over a larger volume of coronal dentine, resulting in lower average (and relative) enamel thickness values. Sample sizes do not permit rigorous intragroup comparisons, but Neandertal molar tissue proportions evince less variation than the modern human sample. Differences in three- and two-dimensional enamel thickness data describing Neandertal molars may be explained by dimensional reduction. Although molar tissue proportions distinguish Neanderthals from recent Homo sapiens, additional study is necessary to assess trends in tissue proportions in the genus Homo throughout the Pleistocene.     

Hunter-Schreger band orientation in interproximal enamel of modern human molars with complex morphological traits

A recent paper concerning characteristic groove-shaped wear in interproximal attritional facets of Neandertal posterior teeth correlated groove formation with deep enamel structural traits (Villa & Giacobini, 1995). Further studies were carried out on modern human teeth in order to evaluate a possible correlation between deep enamel structure and some crown complex morphological features. Seventy-five isolated molars from medieval and recent ossuaries as well as five molars recently extracted for therapeutic purpose were examined using scanning electron and optic microscope ground sections of interproximal enamel. The results showed possible correlations between deep enamel structural characteristics (i.e., Hunter-Schreger band verticalization) and some crown complex traits (i.e., occlusal groove pattern, mesial and distalfovea, mesial and distal accessory tubercules, interproximal marginal ridges). Observations reported in this study showed a significantly higher frequency of Hunter-Schreger band verticalization in lower molars with an occlusal complex pattern (i.e. primitive fissure pattern “Y” or “+”). This observation suggests a positive correlation among crown complex traits (which occur frequently in Neandertal teeth) with deep enamel structural characteristics. In addition, analysis of teeth grouped according to individual showed crown morphological traits and deep enamel structural characteristics corresponding to the same individual. This observation supports a genetic basis in occlusal shape and microstructural dental traits.     

Variation in enamel thickness within the genus Homo

Recent humans and their fossil relatives are classified as having thick molar enamel, one of very few dental traits that distinguish hominins from living African apes. However, little is known about enamel thickness in the earliest members of the genus Homo, and recent studies of later Homo report considerable intra- and inter-specific variation. In order to assess taxonomic, geographic, and temporal trends in enamel thickness, we applied micro-computed tomographic imaging to 150 fossil Homo teeth spanning two million years. Early Homo postcanine teeth from Africa and Asia show highly variable average and relative enamel thickness (AET and RET) values. Three molars from South Africa exceed Homo AET and RET ranges, resembling the hyper thick Paranthropus condition. Most later Homo groups (archaic European and north African Homo, and fossil and recent Homo sapiens) possess absolutely and relatively thick enamel across the entire dentition. In contrast, Neanderthals show relatively thin enamel in their incisors, canines, premolars, and molars, although incisor AET values are similar to H. sapiens. Comparisons of recent and fossil H. sapiens reveal that dental size reduction has led to a disproportionate decrease in coronal dentine compared with enamel (although both are reduced), leading to relatively thicker enamel in recent humans. General characterizations of hominins as having ‘thick enamel’ thus oversimplify a surprisingly variable craniodental trait with limited taxonomic utility within a genus. Moreover, estimates of dental attrition rates employed in paleodemographic reconstruction may be biased when this variation is not considered. Additional research is necessary to reconstruct hominin dietary ecology since thick enamel is not a prerequisite for hard-object feeding, and it is present in most later Homo species despite advances in technology and food processing.     

The Neanderthalian molar from Hunas, Germany.

In this paper, we present a well-preserved isolated human molar found in 1986 in the Hunas cave ruin, south-east Bavaria. The tooth was located at the bottom of layer F2, which belongs to a long stratigraphic sequence comprising faunal remains as well as archaeological levels (Mousterian). A stalagmite from layer P at the base of the stratigraphic sequence was recently dated to 79.373+/-8.237 ka (base) and 76.872+/-9.686 ka (tip) by TIMS-U/Th (Stanford University). We identified the tooth as a right (possibly third) mandibular molar. Characteristic parameters such as crown and root morphology, fissure pattern, enamel thickness, occlusal and interproximal wear, dental dimensions and indices, and radiological features indicate that the Hunas molar represents the tooth of a Neanderthal. This is corroborated by both the palaeontological and archaeological findings (Mousterian) of layer F2.     

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。     

An examination of dental development in Graecopithecus freybergi (=Ouranopithecus macedoniensis)

This study examined enamel thickness and dental development in Graecopithecus freybergi (=Ouranopithecus macedoniensis), a late Miocene hominoid from Greece. Comparative emphasis was placed on Proconsul, Afropithecus, Dryopithecus, Lufengpithecus, and Gigantopithecus, fossil apes that vary in enamel thickness and patterns of development. In addition, comparisons were made with Paranthropus to investigate reported similarities in enamel thickness. Several sections of a right lower third molar were generated, from which enamel thickness and aspects of the enamel and dentine microstructure were determined. Data from parallel sections shed light on the effects of section obliquity, which may influence determination of both enamel thickness and crown formation time. Graecopithecus has relatively thick enamel, greater than any fossil ape but less than Paranthropus, with which it does show similarity in prism path and Hunter-Schreger band morphology. Aspects of enamel microstructure, including the periodicity and daily secretion rate, are similar to most extant and fossil apes, especially Afropithecus. Total crown formation time was estimated to be 3.5 years, which is greater than published values for modern Homo, similar to Pan, and less than Gigantopithecus. Data on dentine secretion and extension rates suggest that coronal dentine formation was relatively slow, but comparative data are very limited. Graecopithecus shares a crown formation pattern with several thick-enamelled hominoids, in which cuspal enamel makes up a very large portion of crown area, is formed by a large cell cohort, and is formed in less than half of the total time of formation. In Paranthropus, this pattern appears to be even more extreme, which may result in thicker enamel formed in an even shorter time. Developmental similarities between Paranthropus and Graecopithecus are interpreted to be parallelisms due to similarities in the mechanical demands of their diets.     

Talonid crests expression at the enamel–dentine junction of hominin lower permanent and deciduous molars

The application of microtomography (mCT) to dental morphological studies has unveiled a new source of palaeobiological information, particularly in the analysis of the internal structures of teeth. In this study, we assess the expression of talonid crests at the enamel and dentine surfaces in lower permanent and second deciduous molars (M₂ and dm₂) of H. sapiens, H. neanderthalensis and Atapuerca-Sima de los Huesos (SH) hominins. In modern humans, talonid crests are described exclusively in the deciduous teeth (Korenhof, 1982) and interpreted as a primitive mammalian remnant of the talonid attachment to the trigonid. Here we report for the first time the expression of talonid crests of deciduous and permanent molars in H. sapiens, H. neanderthalensis and Middle Pleistocene hominins. We discuss possible evolutionary interpretations and suggest the importance of recording this feature in future studies.     

«Relative growth» from the dentino-enamel junction in primate maxillary molars

The dentino-enamel junction is critical throughout growth to mature crown configurations, being the interface between the papilla and the dental cap. Enamel deposition occurs relatively late and often causes changes from the pattern residing in the dentino-enamel junction. Primate teeth (mostly M¹) have been stripped of enamel after measurement and mapping of the original crown. Relative growth, a variant of static adult (allomorphic) allometries, is assessed by displacement of enamel basal crown component landmarks from dentine homologues relative to tooth size. The hypothesis that differential enamel growth reflects evolutionary history is supported by the positive allometry and shape differences in enamel versus dentine landmarks among phyletically enlarged and dentally-reduced primates.     

The Early Aurignacian human remains from La Quina-Aval (France)

There is a dearth of diagnostic human remains securely associated with the Early Aurignacian of western Europe, despite the presence of similarly aged early modern human remains from further east. One small and fragmentary sample of such remains consists of the two partial immature mandibles plus teeth from the Early Aurignacian of La Quina-Aval, Charente, France. The La Quina-Aval 4 mandible exhibits a prominent anterior symphyseal tuber symphyseos on a vertical symphysis and a narrow anterior dental arcade, both features of early modern humans. The dental remains from La Quina-Aval 1 to 4 (a dm₁, 2 dm₂, a P⁴ and a P₄) are unexceptional in size and present occlusal configurations that combine early modern human features with a few retained ancestral ones. Securely dated to ∼33 ka ¹⁴C BP (∼38 ka cal BP), these remains serve to confirm the association of early modern humans with the Early Aurignacian in western Europe.     

Technical note: Estimating original crown height in worn mandibular canines using aspects of dentin morphology

We present a novel method to estimate original crown height (OCH) for worn human mandibular canines using a cubic regression equation based on ratios of worn crown height and exposed dentin. This method may help alleviate issues frequently presented by worn teeth in dental analyses, including those in bioarchaeology. Mandibular canines (n = 28) from modern day New Zealand and English populations were selected. Crown height and dentin thickness were measured on dental thin sections (n = 19) and the resulting (log10) ratios were fitted to a cubic regression curve allowing OCH in worn crowns to be predicted. Variation in the dentin apex position was recorded and effects of angled wear slopes investigated allowing adjusted values to be generated. Our method is trialed for use on intact and sectioned teeth (n = 17). A cubic regression curve best describes the relationship between (log10) ratios and crown height deciles (R² = 0.996, df₁ = 3, df₂ = 336, p < 0.001). No significant differences were detected between OCH estimates using our method and digitally recreated cusp outlines of the same crowns (t = 1.024, df = 16, p > 0.05), with a mean absolute error of 0.171 mm and an adjusted coefficient of determination of 0.923. Our approach offers a quantitative method to estimate the percentage of OCH remaining on worn mandibular canines, and by extension, the OCH. Our estimates are comparable to digitally recreated cusps but less subjective and not limited to crowns with minimal wear.     

Postcanine microstructure in Cricodon metabolus,a Middle Triassic gomphodont cynodont from south‐eastern Africa

Cricodon metabolus is a trirachodontid cynodont from the Anisian (Middle Triassic) of eastern and southern Africa. It has labiolingually expanded (gomphodont) postcanines but also a sectorial tooth in the last postcanine locus. In this paper, we examine the crown microstructure of isolated sectorial and gomphodont postcanines belonging to the holotype specimen of this taxon using scanning electron microscopy. The enamel of both teeth is prismless and composed of discontinuous columnar divergence units, supporting the consistent presence of synapsid columnar enamel in cynognathians. Abundant tubules and numerous irregularly spaced incremental lines are also visible in the enamel and dentine layers in each tooth. This study reveals that the enamel thickness varies along the tooth row in Cricodon as the enamel layer of the gomphodont postcanines is 11.5 times thicker than that of the sectorial crown. It is likely that this difference reflects occlusal stresses and fewer replacements in gomphodont postcanines relative to sectorial teeth. Approximately 100 incremental growth lines of von Ebner are present in the dentine layer, indicating that the deposition of the dentine by odontoblasts occurred for three months before the animal's death.     

Enamel thickness and the helicoidal occlusal plane

In the present study 38 unworn maxillary molars (M¹ = 16, M²= 12, M³ = 10) of modern humans from a Slavic necropolis were sectioned through the mesial cusps in a plane perpendicular to the cervical margin of the crown. Five slightly worn M¹s and one slightly worn M³ were also used thus increasing the total sample to 44, but measurements made on the worn areas were coded as missing values. Seven measurements of enamel thickness as well as the heights of the protocone and the paracone dentine horns were recorded in order to analyze whether changes in these dimensions in anteroposterior direction can be related to the helicoidal occlusal plane. Uni- and multivariate analyses revealed that the distribution of enamel thickness within and between maxillary molars corresponds to a helicoidal occlusal wear pattern. Enamel thickness along the occlusal basin increases from anterior to posterior, which may lead to rapid development of a reverse curve of Monson in first molars when compared to posterior teeth. However, although these overall differences together with the serial, especially delayed eruption pattern of human molars, contribute to the marked expression of the helicoidal occlusal plane in Homo, differences in enamel patterning between molars indicate that a helicoidal plane is a structural feature of the orofacial skeleton. In contrast to first upper molars, second and third molars show absolutely and relatively thicker enamel under the Phase I wear facet of the paracone, i. e., the lingual slope of the paracone, than under the Phase II facet of the protocone, i. e., the buccal slope of that cusp. These proportional differences are most pronounced in M³, as evidenced by uni- and multivariate statistics. It thus appears that the pattern of enamel thickness distribution from M¹ to M³ follows a trend towards providing additional tooth material in areas that are under greater functional demands, that is, corresponding to a lingual slope of wear anteriorly and to a flat or even buccal one posteriorly. In addition, the heights of the dentine horns in anteroposterior direction change in a way that lends support to the hypothesis that the axial inclination of teeth could be one of the most important factors for the development of the helicoidal occlusal plane. Finally, the changes in morphology and enamel thickness distribution from first to third upper molars found in this study suggest that molars could be “specialized” in their function, i. e., from performing proportionally more shearing anteriorly to increased crushing and grinding activities posteriorly. © 1994 Wiley-Liss, Inc.     

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.