Enamel extension rate patterns in modern human teeth: Two approaches designed to establish an integrated comparative context for fossil primates

Enamel extension rates (EERs), the rates at which ameloblasts differentiate, determine how fast tooth crowns grow in height. Studies of fossil primate (including hominin) enamel microstructure usually focus on species differences in enamel formation time, but they have also begun to address species-level variation in enamel extension rates. To improve our ability to compare EERs among primate species, a better understanding how EERs vary within species is necessary. Using a large and diverse modern human histological sample, we find that initial EERs and patterns of EER change along the enamel-dentine junction (EDJ) vary in relation to EDJ length. We also find that enamel formation time varies in relation to EDJ length, but that it does so independently of initial EERs. These results suggest that EDJ length variation within a species sample can affect interspecific comparisons not only of EERs but also of enamel formation times. Additionally, these results lend within-species support to the hypothesis, based on comparisons among hominin species, that EERs and crown formation times can vary independently (Dean, 2009). In a second approach, we analyzed EER changes specifically in the lateral enamel of two modern human population samples as these changes relate to the distribution of perikymata. As surface manifestations of internal enamel growth increments, perikymata provide a valuable source of information about enamel growth in fossils. We find that EER declines in the lateral enamel are associated with an increase in perikymata density from first to last-formed lateral enamel. Moreover, variation in the extent of EER decline among individuals is associated with variation in the distribution of perikymata along their enamel surfaces. These latter findings suggest that the distribution of perikymata on the enamel surface provides information about rates of EER decline in lateral enamel, at least in modern humans.     

Perikymata spacing and distribution on hominid anterior teeth

We documented the spacing and distribution of perikymata on the buccal enamel surface of fossil hominin anterior teeth with reference to a sample of modern human and modern great ape teeth. A sample of 27 anterior teeth attributed to Australopithecus (5 to A. afarensis, 22 to A. africanus) and of 33 attributed to Paranthropus (6 to P. boisei, and 27 to P. robustus) were replicated and sputter-coated with gold to enable reflected light microscopy of their surface topography. Anterior teeth were then divided into 10 equal divisions of buccal crown height. The total perikymata count in each division of crown height was recorded using a binocular microscope fitted with a vernier micrometer eyepiece. Then the mean number of perikymata per millimeter was calculated for each division. Similar comparative data for a modern sample of 115 unworn human anterior teeth and 30 African great ape anterior teeth were collected from ground sections. Perikymata counts in each taxon (together with either known or presumed periodicities of perikymata) were then used to estimate enamel formation times in each division of crown height, for all anterior tooth types combined. The distributions of these estimates of time taken to form each division of crown height follow the same trends as the actual perikymata counts and differ between taxa in the same basic way. The distinction between modern African great apes and fossil hominins is particularly clear. Finally, we calculated crown formation times for each anterior tooth type by summing cuspal and lateral enamel formation times. Estimates of average crown formation times in australopiths are shorter than those calculated for both modern human and African great ape anterior teeth. The data presented here provide a better basis for exploring differences in perikymata spacing and distribution among fossil hominins, and provide the first opportunity to describe four specimens attributed to Homo in this context. Preliminary data indicate that differences may exist among the species attributed to early Homo, especially between Homo ergaster and Homo rudolfensis on the one hand, and Homo habilis sensu strico on the other.     

“Missing perikymata”—fact or fiction? A study on chimpanzee (Pan troglodytes verus) canines

Recently, a lower than expected number of perikymata between repetitive furrow‐type hypoplastic defects has been reported in chimpanzee canines from the Fongoli site, Senegal (Skinner and Pruetz: Am J Phys Anthropol 149 (2012) 468–482). Based on an observation in a localized enamel fracture surface of a canine of a chimpanzee from the Taï Forest (Ivory Coast), these authors inferred that a nonemergence of striae of Retzius could be the cause for the “missing perikymata” phenomenon in the Fongoli chimpanzees. To check this inference, we analyzed the structure of outer enamel in three chimpanzee canines. The teeth were studied using light‐microscopic and scanning‐electron microscopic techniques. Our analysis of the specimen upon which Skinner and Pruetz (Am J Phys Anthropol 149 (2012) 468–482) had made their original observation does not support their hypothesis. We demonstrate that the enamel morphology described by them is not caused by a nonemergence of striae of Retzius but can be attributed to structural variations in outer enamel that result in a differential fracture behavior. Although rejecting the presumed existence of nonemergent striae of Retzius, our study provided evidence that, in furrow‐type hypoplastic defects, a pronounced tapering of Retzius increments can occur, with the striae of Retzius forming acute angles with the outer enamel surface. We suggest that in such cases the outcrop of some striae of Retzius is essentially unobservable at the enamel surface, causing too low perikymata counts. The pronounced tapering of Retzius increments in outer enamel presumably reflects a mild to moderate disturbance of the function of late secretory ameloblasts. Am J Phys Anthropol 157:276–283, 2015. © 2015 Wiley Periodicals, Inc.     

Accessing Developmental Information of Fossil Hominin Teeth Using New Synchrotron Microtomography-Based Visualization Techniques of Dental Surfaces and Interfaces

Quantification of dental long-period growth lines (Retzius lines in enamel and Andresen lines in dentine) and matching of stress patterns (internal accentuated lines and hypoplasias) are used in determining crown formation time and age at death in juvenile fossil hominins. They yield the chronology employed for inferences of life history. Synchrotron virtual histology has been demonstrated as a non-destructive alternative to conventional invasive approaches. Nevertheless, fossil teeth are sometimes poorly preserved or physically inaccessible, preventing observation of the external expression of incremental lines (perikymata and periradicular bands). Here we present a new approach combining synchrotron virtual histology and high quality three-dimensional rendering of dental surfaces and internal interfaces. We illustrate this approach with seventeen permanent fossil hominin teeth. The outer enamel surface and enamel-dentine junction (EDJ) were segmented by capturing the phase contrast fringes at the structural interfaces. Three-dimensional models were rendered with Phong’s algorithm, and a combination of directional colored lights to enhance surface topography and the pattern of subtle variations in tissue density. The process reveals perikymata and linear enamel hypoplasias on the entire crown surface, including unerupted teeth. Using this method, highly detailed stress patterns at the EDJ allow precise matching of teeth within an individual’s dentition when virtual histology is not sufficient. We highlight that taphonomical altered enamel can in particular cases yield artificial subdivisions of perikymata when imaged using X-ray microtomography with insufficient resolution. This may complicate assessments of developmental time, although this can be circumvented by a careful analysis of external and internal structures in parallel. We further present new crown formation times for two unerupted canines from South African Australopiths, which were found to form over a rather surprisingly long time (> 4.5 years). This approach provides tools for maximizing the recovery of developmental information in teeth, especially in the most difficult cases.     

What molars contribute to an emerging understanding of lateral enamel formation in Neandertals vs. modern humans

Two hypotheses, based on previous work on Neandertal anterior and premolar teeth, are investigated here: (1) that estimated molar lateral enamel formation times in Neandertals are likely to fall within the range of modern human population variation, and (2) that perikymata (lateral enamel growth increments) are distributed across cervical and occlusal halves of the crown differently in Neandertals than they are in modern humans. To investigate these hypotheses, total perikymata numbers and the distribution of perikymata across deciles of crown height were compared for Neandertal, northern European, and southern African upper molar mesiobuccal (mb) cusps, lower molar mesiobuccal cusps, and the lower first molar distobuccal (db) cusp. Sample sizes range from five (Neandertal M(1)db) to 29 (southern African M(1)mb). Neandertal mean perikymata numbers were found to differ significantly from those of both modern human samples (with the Neandertal mean higher) only for the M(2)mb. Regression analysis suggests that, with the exception of the M(2)mb, the hypothesis of equivalence between Neandertal and modern human lateral enamel formation time cannot be rejected. For the M(2)mb, regression analysis strongly suggests that this cusp took longer to form in the Neandertal sample than it did in the southern African sample. Plots of perikymata numbers across deciles of crown height demonstrate that Neandertal perikymata are distributed more evenly across the cervical and occlusal halves of molar crowns than they are in the modern human samples. These results are integrated into a discussion of Neandertal and modern human lateral enamel formation across the dentition, with reference to issues of life history and enamel growth processes.     

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.     

Under restrictive conditions,can the widths of linear enamel hypoplasias be used as relative indicators of stress episode duration?

Linear enamel hypoplasia (LEH), a type of enamel defect reflecting nonspecific physiological stress, has traditionally been used by bioarchaeologists to assess human health. Initially, measurements of defect width were used to estimate the duration of stress episodes. More recently, methods of counting within-defect perikymata (enamel growth increments) were developed to more accurately assess duration. Because perikymata are often not continuously visible within defects, while widths can usually be measured, the primary purpose of this article was to determine if, under restrictive conditions, the widths of LEH defects might be used as relative indicators of stress episode duration. Using a set of dental replicas from the prehistoric Irene Mound (1150-1400 A.D.), this study also investigated potential sources of variation in defect widths and how often defect widths could be measured and within-defect perikymata counted. Of 120 defects, only 47 contained both measurable defect widths and total within-defect perikymata, while 79 had measurable defect widths. Regression analysis revealed that, for these 47 defects, defect widths were more strongly related to the total number of within-defect perikymata than they were to crown region or tooth type. Although wide prediction intervals indicated that a defect's width could not be used to predict the number of within-defect perikymata for an individual, narrower confidence intervals associated with hypothetical mean population widths suggested that mean defect widths might be used to rank populations in terms of relative average stress episode duration.     

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.     

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.     

Did the lateral enamel of Neandertal anterior teeth grow differently from that of modern humans?

The formation of lateral enamel in Neandertal anterior teeth has been the subject of recent studies. When compared to the anterior teeth of modern humans from diverse regions (Point Hope, Alaska; Newcastle upon Tyne, England; southern Africa), Neandertal anterior teeth appear to fall within the modern human range of variation for lateral enamel formation time. However, the lateral enamel growth curves of Neandertals are more linear than those of these modern human samples. Other researchers have found that the lateral enamel growth curves of Neandertals are more linear than those of Upper Paleolithic and Mesolithic modern humans as well. The statistical significance of this apparent difference between Neandertal and modern human lateral enamel growth curves is analyzed here. The more linear Neandertal enamel growth curves result from the smaller percentage of total perikymata located in the cervical halves of their teeth. The percentage of total perikymata in the cervical halves of teeth is therefore compared between the Neandertal sample (n=56 teeth) and each modern human population sample: Inuit (n=65 teeth), southern African (n=114 teeth), and northern European (n=115 teeth). There are 18 such comparisons (6 tooth types, Neandertals vs. each of the three modern human populations). Eighteen additional comparisons are made among the modern human population samples. Statistically significant differences are found for 16 of the 18 Neandertal vs. modern human comparisons but for only two of the 18 modern human comparisons. Statistical analyses repeated for subsamples of less worn teeth show a similar pattern. Because surface curvature is thought to affect perikymata spacing, we also conducted measurements to assess surface curvature in thirty teeth. Our analysis shows that surface curvature is not a factor in this lateral enamel growth difference between Neandertals and modern humans.     

Estimating striae of Retzius periodicity nondestructively using partial counts of perikymata

Accurate age estimations for enamel formation and the timing of enamel hypoplasia have traditionally only been available through histological analyses of dental thin sections, which is a difficult and destructive process. However, an association between striae of Retzius periodicity, crucial for accurate aging, and the total number of striae in imbricational enamel has been reported in the literature. This means periodicity can be estimated nondestructively but is reliant on all perikymata being visible along the crown surface. Therefore, crowns with worn or damaged surfaces may not be able to be assessed, potentially limiting sample sizes. We tested this relationship in a modern New Zealand sample and investigated whether reliable associations might be identified using only partial perikymata counts from the cervical half of the crown. Using mandibular canines (n = 11), the distribution of perikymata per decile was recorded using high definition replica surfaces. Thin sections of the same crowns were used to assess periodicity histologically along with striae of Retzius distributions. A strong correlation between total striae numbers and periodicity was also identified in our sample. Furthermore, we report strong correlations that allow periodicity to be estimated from perikymata counts using only 10% of crown height when certain deciles are used. Based on these findings, we propose a simple matrix that can be developed for nondestructively estimating periodicity based on the range of perikymata counts in the sixth to ninth deciles. Am J Phys Anthropol 154:251–258, 2014. © 2014 Wiley Periodicals, Inc.     

Prevalence and the duration of linear enamel hypoplasia: a comparative study of Neandertals and Inuit foragers

As a dental indicator of generalized physiological stress, enamel hypoplasia has been the subject of several Neandertal studies. While previous studies generally have found high frequencies of enamel hypoplasia in Neandertals, the significance of this finding varies with frequencies of enamel hypoplasia in comparative samples. The present investigation was undertaken to ascertain if the enamel hypoplasia evidence in Neandertals suggests a high level of physiological stress relative to a modern human foraging group, represented here by an archaeological sample of Inuit from Point Hope, Alaska. Unlike previous studies, this study focused specifically on linear enamel hypoplasia (LEH), emphasizing systemic over localized causes of this defect by considering LEH to be present in an individual only if LEH defects occur on two anterior teeth with overlapping crown formation periods. Moreover, this study is the first to evaluate the average growth disruption duration represented by these defects in Neandertals and a comparative foraging group. In the prevalence analysis, 7/18 Neandertal individuals (from Krapina and southern France) and 21/56 Neandertal anterior teeth were affected by LEH, or 38.9% and 37.5% respectively. These values do not differ significantly from those of the Inuit sample in which 8/21, or 38.1% of individuals, and 32/111, or 28.8% of anterior teeth were affected. For the growth disruption duration analysis, 22 defects representing separate episodes of growth disruption in Neandertals were compared with 22 defects in the Inuit group using three indicators of duration: the number of perikymata (growth increments) in the occlusal walls of LEH defects, the total number of perikymata within them, and defect width. Only one indicator, the total number of perikymata within defects, differed significantly between the Inuit and Neandertal groups (an average of 13.4 vs. 7.3 perikymata), suggesting that if there is any difference between them, the Inuit defects may actually represent longer growth disruptions than the Neandertal defects. Thus, while stress indicators other than linear enamel hypoplasia may eventually show that Neandertal populations were more stressed than those of modern foragers, the evidence from linear enamel hypoplasia does not lend support to this idea.     

Growth layers and incremental markings in hard tissues; a review of the literature and some preliminary observations about enamel structure in Paranthropus boisei

The general factors underlying the formation of growth layers and incremental markings in hard tissues are reviewed with particular reference to fossil hominid tooth enamel. The experimental and circumstantial evidence that point to a slowing of enamel matrix secretion in a daily (circadian) and near weekly (circaseptan) mode during tooth formation is also reviewed. Data from previous studies in which the number of daily increments between adjacent striae of Retzius have been recorded in primates are reviewed and new data are presented for this repeat interval in fossil hominids. The factors likely to influence the number of striae of Retzius beneath the cuspal regions of anterior teeth are outlined and the limitations of employing surface incremental features to obtain estimates for age at death of an individual are also discussed. It is concluded that there is good evidence to support the hypothesis that perikymata are near weekly incremental phenomena with a likely periodicity of 7,8 or 9 days in fossil hominids. It can also be concluded that at present, better estimates for the age at death of an individual during early phases of the growth period can be obtained from studies of perikymata than by any other non-destructive technique.     

Scanning electron microscope study of cat and dog enamel structure

Scanning electron microscopy revealed several similarities as well as significant differences in the enamel structure between cat and dog teeth. Three enamel layers were present in both species; a surface rodless (aprismatic) layer, an outer layer of parallel rods (only at some sites), and an inner layer with prominent Hunter-Schreger bands. In the inner layer of both carnivores, the diameter of individual rods varied significantly and frequently their course changed abruptly with respect to neighboring rods. In dog teeth the cross-sectional shape of inner enamel rods was pleomorphic, but hexagonal in outer enamel. In contrast, cat enamel rods were rounded in both inner and outer enamel layers. Hunter-Schreger bands of cats circumscribed the teeth in relatively straight segments, but these bands showed pronounced waviness in dog teeth. In cats and dogs the surface rodless layer was structurally continuous with subjacent interrod enamel and covered all tooth surfaces with the exception of the cervical areas. The data show that the structure of inner and outer enamel layers differ between these two carnivore species and that the enamel structure of the cat was most similar to that described in humans. One principal difference between carnivore and human teeth is that the growth lines of carnivores do not terminate at perikymata on the tooth surface.     

Plasticity and constraint in response to early‐life stressors among late/final jomon period foragers from Japan: Evidence for life history trade‐offs from incremental microstructures of enamel

This study evaluates two hypotheses that address how Late/Final Jomon period people responded to early‐life stress using linear enamel hypoplasia (LEH) and incremental microstructures of enamel. The first hypothesis predicts that Jomon people who experienced early‐life stressors had greater physiological competence in responding to future stress events (predictive adaptive response). The second hypothesis predicts that Jomon people traded‐off in future growth and maintenance when early investment in growth and survival was required (plasticity/constraint). High resolution tooth impressions were collected from intact, anterior teeth and studied under an engineer's measuring microscope. LEH were identified based on accentuated perikymata and depressions in the enamel surface profile. Age of formation for each LEH was estimated by summing counts of perikymata and constants associated with crown initiation and cuspal enamel formation times. The relationship between age‐at‐first‐defect formation, number of LEH, periodicity between LEH, and mortality was evaluated using multiple regression and hazards analysis. A significant, positive relationship was found between age‐at‐death relative to age‐at‐first‐defect formation and a significant, negative relationship was found between number of LEH relative to age‐at‐first‐defect formation. Individuals with earlier forming defects were at a significantly greater risk of forming defects at later stages of development and dying at younger ages. These results suggest that Late/Final Jomon period foragers responded to early‐life stressors in a manner consistent with the plasticity/constraint hypothesis of human life history. Late/Final Jomon period individuals were able to survive early‐life stressors, but this investment weakened responses to future stress events and exacerbated mortality schedules. Am J Phys Anthropol 155:537–545, 2014. © 2014 Wiley Periodicals, Inc.     

Enamel incremental periodicity in the pig-tailed macaque: A polychrome fluorescent labeling study of dental hard tissues

Bromage and Dean originally outlined a nondestructive method for the study of enamel formation and concluded that early hominids resembled the extant apes more closely than they did modern humans in their rates of growth and maturation. The method used assumed that an enamel circadian rhythmicity was referable to a longer near-weekly period represented by perikymata (periodic surface growth features). This assumption became a matter of debate and discussion. In this study, developing teeth in Macaca nemestrina were labeled with polychrome fluorescent dyes. Examination of the distribution of these dyes in two sectioned teeth provides experimental confirmation of enamel circadian periodicity.     

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).     

禄丰古猿(Lufengpithecus lufengensis)牙齿釉质生长线与个体发育问题研究

运用扫描电子显微镜,对4枚禄丰古猿牙齿（恒齿）的釉质结构进行了观察研究。发现：禄丰古猿牙齿釉质表面有明显的釉面横纹结构;釉面横纹的密度向牙颈方向逐渐增大;观察记数了4枚牙齿的釉面横纹数,进而推算出牙冠的形成时间和年龄。与化石人科成员、现代人及现生大猿比较,禄丰古猿牙冠发育模式及时间,与南方古猿纤细种比较接近或相似,明显长于南方古猿粗壮种,有别于现生大猿。     

Pig enamel revisited – Incremental markings in enamel of wild boars and domestic pigs

The nature and periodicity of incremental markings in pig enamel is currently debated. To broaden the basis for a correct interpretation of growth marks in pig enamel, we analyzed their periodicity in teeth of wild boars and domestic pigs. For that, the numbers of enamel incremental markings were recorded in ground sections and compared with crown formation times for the respective teeth derived from literature data on tooth development and eruption in Sus scrofa. Our study revealed that laminations with a daily periodicity are the dominant incremental feature of pig enamel. In wild boar M₃s, daily enamel secretion (apposition) rates ranged between a minimum of 6.1?µm in the inner and a maximum of 30.6?µm in the outer enamel.Long-period (supra-daily) incremental markings were present as perikymata at the outer enamel surface (OES). Contrary to the situation in primate enamel, in pig enamel the long-period incremental lines terminating in perikyma grooves were mostly structurally indistinguishable from the daily laminations. Typically, five sub-daily increments were present between successive laminations. The incremental pattern in pig enamel can be misinterpreted if the laminations are mistaken for long-period markings (striae of Retzius) and the sub-daily growth marks for daily prism cross-striations. The findings of the present study demonstrate the critical importance of correctly characterizing the incremental markings and their periodicity in enamel, and caution against an uncritical transfer of the interpretation of the nature of incremental markings in primate enamel to other mammalian taxa.     

Reconstruction of periodicity of repetitive linear enamel hypoplasia from perikymata counts on imbricational enamel among dry‐adapted chimpanzees (Pan troglodytes verus) from Fongoli,Senegal

Periodicity of repetitive linear enamel hypoplasia (rLEH) in apes from high latitudes with single wet and dry seasons annually has not been described. We reconstruct periodicity and duration of rLEH in canine teeth from three recently deceased chimpanzees from Fongoli, Senegal with a marked seven‐month dry season. High‐resolution dental molds were taken in the field for magnified imaging with digital microscopy. Photomontages allowed counting of perikymata between episodes of rLEH for reconstruction of periodicity and duration of physiological stress. Where rLEH spans the imbricational enamel, the number of events is consistent with years required to form canine imbricational enamel; i.e., periodicity of rLEH seems circannual. We predicted perikymata counts between rLEH events ranging from 52 to 61 based on reported “long counts” of 7–6 days. Counts ranged from 29.5 to 44, individual mean of 36.7. This discrepancy could be explained by recurrent stress with a periodicity of 7.2–8.4 months, or by long counts of 10 days per stria. Neither is supported in the literature. Since we find evidence of rLEH with circannual periodicity, we postulate the existence of non‐emergent imbricational striae. Based on evidence that stress at Fongoli recurs annually, we reconstruct stress duration of 2–3 months, longer than reported for chimpanzees living in other habitats, which we attribute to heat stress and food shortage near shrinking waterholes. We conclude that canine teeth from a small mortality cohort of chimpanzees at Fongoli preserve a faithful record of dry season stress in an extreme environment. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.