Technical note: A new three‐dimensional technique for high resolution quantitative recording of perikymata

The number and spacing of incremental markings at the enamel surface, known as perikymata, are considered important indicators of dental growth patterns, as they provide information on crown formation times and the underlying developmental processes. This study explores the potential of a new three‐dimensional technique for the reconstruction of dental growth profiles, using teeth from a medieval child from Abingdon, Oxfordshire. The crowns of three anterior teeth were imaged and analyzed using the Alicona 3D InfiniteFocus imaging microscope. Individual perikyma grooves can be unambiguously identified on a profile of the reconstructed enamel surface and direct distances between successive pairs of perikyma grooves can be calculated from coordinate data. This quantitative approach constitutes a more objective way to record perikymata spacing than current methods. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Brief communication: The distribution of perikymata on Qafzeh anterior teeth

Recent studies have suggested that Neandertals and modern humans differ in the distribution of perikymata (enamel growth increments) over their permanent anterior tooth crowns. In modern humans, perikymata become increasingly more compact toward the cervix than they do in Neandertals. Previous studies have suggested that a more homogeneous distribution of perikymata, like that of Neandertals, characterizes the anterior teeth of Homo heidelbergensis and Homo erectus as well. Here, we investigated whether Qafzeh anterior teeth (N = 14) differ from those of modern southern Africans, northern Europeans, and Alaskans (N = 47–74 depending on tooth type) in the percentage of perikymata present in their cervical halves. Using the normally distributed modern human values for each tooth type, we calculated Z‐scores for the 14 Qafzeh teeth. All but two of the 14 Qafzeh teeth had negative Z‐scores, meaning that values equal to these would be found in the bottom 50% of the modern human samples. Seven of the 14 would be found in the lowest 5% of the modern human distribution. Qafzeh teeth therefore appear to differ from those of modern humans in the same direction that Neandertals do: with generally lower percentages of perikymata in their cervical regions. The similarity between them appears to represent the retention of a perikymata distribution pattern present in earlier members of the genus Homo, but not generally characteristic of modern humans from diverse regions of the world. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Linear enamel hypoplasia as an indicator of physiological stress in great apes: reviewing the evidence in light of enamel growth variation

Physiological stress, such as malnutrition or illness, can disrupt normal enamel growth, resulting in linear enamel hypoplasias (LEHs). Although ecological factors may contribute to LEH expression, other factors, such as surface abrasion and enamel growth variables, are also likely to be involved. Attention to these other factors is necessary before we can begin to understand what LEH might signify in terms of ecological sources of physiological stress in non-human primates. This study focuses on assessing the contribution of these other factors to variation in LEH expression within and across great ape taxa. Here, we present LEH data from unabraded crown regions in samples of seven great ape species. We analyze these data with respect to lateral enamel formation time and the angles that striae of Retzius make with the enamel surface, as these variables are expected to affect variation in LEH expression. We find that although the duration of enamel formation is associated with sex differences in LEH expression, it is not clearly related to taxonomic variation in LEH expression, and does not explain the low frequency of LEH in mountain gorillas found in this and a previous study. Our data on striae of Retzius angles suggest that these influence LEH expression along the tooth crown and may contribute to the consistently high frequencies of LEH seen in Pongo in this and previous studies. We suggest that future work aimed at understanding species variation in these angles is crucial to evaluating taxonomic patterns of LEH expression in great apes.     

Retrieving chronological age from dental remains of early fossil hominins to reconstruct human growth in the past

A chronology of dental development in Pan troglodytes is arguably the best available model with which to compare and contrast reconstructed dental chronologies of the earliest fossil hominins. Establishing a time scale for growth is a requirement for being able to make further comparative observations about timing and rate during both dento-skeletal growth and brain growth. The absolute timing of anterior tooth crown and root formation appears not to reflect the period of somatic growth. In contrast, the molar dentition best reflects changes to the total growth period. Earlier initiation of molar mineralization, shorter crown formation times, less root length formed at gingival emergence into functional occlusion are cumulatively expressed as earlier ages at molar eruption. Things that are similar in modern humans and Pan, such as the total length of time taken to form individual teeth, raise expectations that these would also have been the same in fossil hominins. The best evidence there is from the youngest fossil hominin specimens suggests a close resemblance to the model for Pan but also hints that Gorilla may be a better developmental model for some. A mosaic of great ape-like features currently best describes the timing of early hominin dental development.     

Meaningful measures of enamel hypoplasia: Prevalence and comparative intensity of developmental stress

Objectives

Developmental stress causing enamel thinning is an important topic in primate biology. Because taxa differ in growth rates and enamel thickness, the goal is to provide a new method allowing direct comparison of prevalence and salience of enamel defects among samples.

Materials and Methods

Casts of ape teeth spanning the Late Pleistocene to Late Miocene from three site areas of increasing seasonality, equator (Sumatra) to 20° (Vietnam) and 25°N latitude (China), were examined for enamel defects among paleo-orangutans (n = 571, 222, respectively) and Lufengpithecus lufengensis (n = 198). Frequency of affected teeth and number of linear enamel hypoplasia were recorded. Defect dimensions were measured with a confocal microscope. Simple prevalence is compared to weighted prevalence (%), calculated by dividing “number of LEH from specific tooth groups” by “specific tooth sample size”; this quantity divided by “tooth-specific years of imbricational enamel formation.” Defect dimensions are reduced to a dimensionless index termed “enamel deficit ratio” through dividing “daily enamel deficit” by “daily secretion rate.”

Results

Weighted prevalence increases to the North, highlighting latitudinal similarities. In contrast, “enamel deficit ratio,” designed to express comparative severity of developmental stress among samples, was least in the high latitude sample and differed little between paleo-orangutan samples.

Discussion

The actual numbers generated are not as important as efficacy of the proposed methods for other taxa. Developmental stress appears least severe in the high latitude (Lufengpithecus) sample but affects a greater proportion, compared to paleo-orangutans. Regardless of findings, the proposed solutions to improve comparability of disparate samples, yield reasonable results.     

Macroscopic and microscopic analyses of linear enamel hypoplasia in Plio-Pleistocene South African hominins with respect to aspects of enamel development and morphology

This study uses macroscopic and microscopic methods to analyze the expression of linear enamel hypoplasia (LEH) in Plio-Pleistocene South African hominins. LEH is a developmental defect of enamel that is used in many anthropological contexts as a physiological stress indicator. Previous research has not settled the question as to whether differences in LEH expression exist between Paranthropus and Australopithecus and if they exist, to what extent these differences might be explained simply by taxonomic differences in enamel development and morphology rather than by differential stress experience. In this study, the analysis of LEH is conducted with respect to differences between Paranthropus and Australopithecus in aspects of enamel development and morphology that are thought to influence LEH expression. Two factors impacting LEH expression are considered: the duration of enamel formation, and the spacing of perikymata. It is predicted that if the first factor strongly influences the expression of LEH, then there should be fewer defects per tooth in Paranthropus because of its abbreviated crown formation spans (and fast extension rates) relative to Australopithecus. It is also predicted that because Australopithecus has more densely packed perikymata in comparable regions of the crown than Paranthropus, this taxon should, on average, have narrower defects than Paranthropus. To address these questions, 200 Australopithecus and 137 Paranthropus teeth were examined for LEH, and the analysis of defect width with respect to perikymata spacing was conducted on tooth impressions examined under a scanning electron microscope using INCA (Oxford Instruments) measurement software. Data support the first prediction: Australopithecus does have significantly more defects per canine tooth than Paranthropus. Data do not support the second prediction in large part because several Australopithecus specimens have wide groove defects in which perikymata are not visible and enamel is irregular. Such wide grooves are not predicted by perikymata spacing such that alternative explanations, including taxonomic differences in ameloblast sensitivity and the duration/severity of disruptions to enamel growth, must be considered.     

Relationship of enamel hypoplasia to the pattern of tooth crown growth: A discussion

The defects of enamel hypoplasia can be related to the layered structure of enamel which represents the sequence of development in tooth crowns. From such studies, it is possible to see that furrow-type enamel defects (the most common form of hypoplasia seen with the naked eye) are just the most prominent expression of a continuum which extends ever smaller, down to a microscopic disturbance to a single layer in the crown formation sequence. Furthermore, the progressive decrease in spacing between development layers which occurs down the crown sides, from occlusal to cervical, affects both the prominence and apparent width of the defects. This makes it difficult to use measurements as a means of estimating the duration of the disturbance causing a particular defect. The difficulty is even greater for the less common pitted or exposed-plane-type defects, for which the apparent width bears very little relationship with the duration of the growth disturbance. The defects of enamel hypoplasia can therefore be understood clearly only when examined under the microscope in relation to the structures which mark the development sequence of the tooth crown. Am J Phys Anthropol 104:89–103, 1997. © 1997 Wiley-Liss, Inc.     

Life history, enamel formation, and linear enamel hypoplasia in the Ceboidea

Linear enamel hypoplasia (LEH), a developmental defect of enamel, increases in frequency from prosimian to monkey to lesser ape to great ape grades (Guatelli-Steinberg 2000 Am. J. Phys. Anthropol. 112:395-410, [2001] Evol. Anthropol. 10:138-151; Newell 1998 Ph.D. dissertation, Temple University). This taxonomic pattern in the distribution of LEH is closely related to maturation length across the primate order (Newell 1998 Ph.D. dissertation, Temple University, 2000 Am. J. Phys. Anthropol. [Suppl.] 30:236). Longer maturation periods are associated with higher LEH frequencies; they appear to provide greater opportunity for defects to form. The present study explores the relationship between maturation length and LEH frequency within the Ceboidea. Because of its prolonged period of growth, Cebus is predicted to manifest LEH at a higher frequency than the more rapidly maturing ceboid genera. To test this hypothesis, two separate researchers (E.A.N. and D.G.-S.) examined LEH in nonoverlapping museum series of ceboids. The results support the hypothesis: in 13 genera (n = 1,276), E.A.N. found that LEH frequencies ranged from 0% in Callicebus, Cebuella, and Saimiri to 20% in Cebus. D.G.-S. found similar frequencies among five genera (n = 107), from 0% in Saimiri to 32% in Cebus. Thus, the broad pattern of LEH distribution evident across major taxonomic groups of primates is repeated within the Ceboidea. We also examined a related hypothesis linking the spacing of perikymata, which is influenced by enamel extension rates (Shellis 1998 J. Hum. Evol. 35:387-400), to LEH. The most likely areas of tooth crowns to exhibit LEH in human teeth are those in which perikymata are most closely spaced (Hillson and Bond 1997 Am. J. Phys. Anthropol. 104:89-103). We hypothesized that the longer-maturing Cebus, with its elevated LEH frequency, will also exhibit more closely spaced perikymata than other ceboids. Analysis of a small microscopic subsample (n = 8) lends limited support to this second hypothesis.     

Preferential expression of linear enamel hypoplasia on the sectorial premolars of rhesus monkeys (Macaca mulatta)

Three hundred and sixty rhesus macaque specimens at the Caribbean Primate Research Center were examined for evidence of linear enamel hypoplasia (LEH). A previously unreported intertooth pattern in LEH was observed. Defects occur preferentially on the sectorial premolar of both males and females. Relative to other teeth, the sectorial premolar exhibits more prominent defects and is more likely to exhibit multiple defects. This pattern is unlike the human intertooth LEH pattern and unlike patterns previously reported for monkeys and apes. These observations are discussed in the context of factors thought to influence the intertooth distribution of LEH in humans and in nonhuman primates. The authors reject crown height, the timing of crown development, and the duration of crown formation as factors contributing to the observed pattern and favor an explanation involving enamel thickness, perikymata spacing, and/or prism orientation. Am J Phys Anthropol 107:179–186, 1998. © 1998 Wiley-Liss, Inc.     

Analysis and significance of linear enamel hypoplasia in Plio-Pleistocene hominins

This study of linear enamel hypoplasia (LEH) in Plio-Pleistocene hominins builds on a previous study (Guatelli-Steinberg [2003] Am. J. Phys. Anthropol. 120:309-322) that focused on LEH in early South African hominins. The present study is more comprehensive, encompassing dental specimens of hominins from East Africa as well, including early Homo. As a developmental defect of enamel, LEH is used in anthropological contexts to reveal information about physiological stress. However, intrinsic aspects of enamel development and morphology can affect the expression of LEH, complicating efforts to understand the significance of these defects. In this study, the analysis of LEH is conducted with respect to enamel development and morphology. It is predicted that Paranthropus should have fewer defects on its canine teeth than Australopithecus and Homo, owing to its abbreviated period of enamel formation. This prediction is supported: Paranthropus has statistically significantly fewer defects per canine than Australopithecus and Homo. The previous study demonstrated that despite the wider spacing of perikymata on the teeth of South African Paranthropus, defects on the canine teeth of this genus were not wider than those of Australopithecus. A multiple linear regression analysis in that study, as well as a separate analysis in the present study, indicate that the number of perikymata within defects is a better predictor of defect width than perikymata spacing. In this study, it was additionally found that the average number of perikymata within Australopithecus defects is statistically significantly greater than it is in Paranthropus, thus explaining why Paranthropus defects are not wider than those of Australopithecus. The biological significance of this difference in the number of perikymata within the defects of Australopithecus and Paranthropus is considered in light of several factors, including: 1) the possibility that other intrinsic attributes of enamel morphology may be involved (specifically the faster extension rates of Paranthropus that result in shallower defects), 2) generic differences in the canalization of enamel development, and 3) generic differences in the duration of disruptions to enamel growth.     

Incremental enamel development in modern human deciduous anterior teeth

This study reconstructs incremental enamel development for a sample of modern human deciduous mandibular (n = 42) and maxillary (n = 42) anterior (incisors and canines) teeth. Results are compared between anterior teeth, and with previous research for deciduous molars (Mahoney: Am J Phys Anthropol 144 (2011) 204-214) to identify developmental differences along the tooth row. Two hypotheses are tested: Retzius line periodicity will remain constant in teeth from the same jaw and range from 6 to 12 days among individuals, as in human permanent teeth; daily enamel secretion rates (DSRs) will not vary between deciduous teeth, as in some human permanent tooth types. A further aim is to search for links between deciduous incremental enamel development and the previously reported eruptionsequence. Retzius line periodicity in anterior teeth ranged between 5 and 6 days, but did not differ between an incisor and molar of one individual. Intradian line periodicity was 12 h. Mean cuspal DSRs varied slightly between equivalent regions along the tooth row. Mandibular incisors initiated enamel formation first, had the fastest mean DSRs, the greatest prenatal formation time, and based upon prior studies are the first deciduous tooth to erupt. Relatively rapid development in mandibular incisors in advance of early eruption may explain some of the variation in DSRs along the tooth row that cannot be explained by birth. Links between DSRs, enamel initiation times, and the deciduous eruption sequence are proposed. Anterior crown formation times presented here can contribute toward human infant age-at-death estimates. Regression equations for reconstructing formation time in worn incisors are given.     

Hominid teeth and their relationship to hominid phylogeny

Hominid fossil teeth are analyzed in terms of size and shape and contrasted with teeth from a modern population to determine boundaries that can be placed on a reconstruction of the hominid phylogeny. Teeth alone are considered as they are the only material preserved in large enough quantities to measure population variability accurately. Problems with the use of indices such as size and shape as species markers are discussed. Conclusions that are made about possible phylogenies are based on the analysis of size and shape.     

Dental fast track: Prenatal enamel growth,incisor eruption,and weaning in human infants

Correlation between the timing of permanent first molar eruption and weaning age in extant primates has provided a way to infer a life history event in fossil species, but recent debate has questioned whether the same link is present in human infants. Deciduous incisors erupt at an age when breast milk can be supplemented with additional foods (mixed feeding), and weaning is typically complete before permanent first molars erupt. Here, I use histological methods to calculate the prenatal rate by which enamel increases in thickness and height on human deciduous incisors, canines, and molars (n = 125). Growth trajectories for each tooth type are related to the trimesters and assessed against the eruption sequence and final crown height. Analyses show that central incisors initiate early in the second trimester with significantly faster secretion rates relative to canines and second molars, which initiate closer to birth. Even though initial extension rates were correlated with crown height and scaled with positive allometry within each tooth class, the relatively short incisors still increased in height at a significantly faster rate than the taller canines and molars. The incisor prenatal “fast track” produces a greater proportion of the crown before birth than all other tooth types. This growth mechanism likely facilitates early incisor eruption at a time when the mixed feeding of infants can be initiated as part of the weaning process. Findings provide a basis from which to explore new links between developmental trends along the tooth row and mixed feeding age in other primates. Am J Phys Anthropol 156:407–421, 2015. © 2014 Wiley Periodicals, Inc.     

Enamel microstructure of the hominid KB 5223 from Kromdraai, South Africa

The Plio-Pleistocene site of Kromdraai, South Africa, is well known for the recovery of the holotype of Paranthropus robustus, one of nine individual hominids recovered from this site to date. Among the Kromdraai sample, the specimen KB 5223 comprises several isolated deciduous and permanent lower teeth assigned to Paranthropus, the only recognized genus at this site. However, a more recent analysis of this specimen suggested that it should be classified as Homo. The lower right first permanent molar of KB 5223 had been previously sectioned along the tips of the mesial cusps, exposing its enamel microstructure. Previous studies had indicated differences between Homo and Paranthropus at the microstructural level. A portable confocal scanning microscope was used to describe details of the enamel microstructure of the M1 and I1 of this specimen. Angles formed between the striae of Retzius and the enamel dentine junction (EDJ), daily secretion rates in cuspal enamel of the protoconid and metaconid and crown formation time of the RM1 are provided. The number of perikymata on the right I1 was counted. Results indicate that some features recorded in the KB 5223 molar differ from those of Paranthropus. However, the number of perikymata on the I1 is lower than values so far reported for early Homo but similar to Paranthropus. Crown formation time of KB 5223 M1 was markedly lower than mean values of M1 in H. sapiens, but similar to other early hominids. Daily secretion rates in the cuspal enamel of KB 5223 M1 were higher than in modern humans.