Enamel Extension Rates Research Articles

Early onset of enamel formation in Baka pygmy’s deciduous canines

ObjectiveOur aim was to evaluate by enamel microstructure analysis two hypotheses that would explain the early dental eruption in the Bakaparticularity, a shorter crown formation time and/or earlier onset of crown formation. DesignDeciduous canines corresponds to the best teeth to perform the analysis of enamel microstructure. Longitudinal ground sections of 21 deciduous canines from 12 individuals were studied with transmitted light microscopy. Cross-striations, striaes of Retzius (SR) and the neonatal line (NNL) enable to establish the prenatal crown formation time (preCFT), the postnatal crown formation time (postCFT), the crown formation time (CFT) as well as the daily secretion rate (DSR) and the enamel extension rate (EER) and their variation along crown formation. ResultsThe DSR and the EER in the Baka are similar than in other populations with an average DSR of 3.26 µm and EER of 18.18 µm. The preCFT was 154 days, the postCFT 265 days and CFT 419 days. Comparison with other population does not show difference in CFT. However, the preCFT and the postCFT differ, the first is higher and the second lower in the Baka than in other populations. Furthermore, the number of prenatal areas of enamel was greater in the Baka. ConclusionOur analysis suggests that the Baka does not distinguish by a different CFT but the onset of crown formation is earlier than in other groups. Therefore, the early dental eruption in the Baka results from an earlier onset of crown formation.

A histological study of enamel developmental defects in a chacma baboon (Papio ursinus) incisor

Physiological stress disrupts normal growth creating visible grooves on the enamel surface (i.e., linear enamel hypoplasia or LEH). Hypoplastic defects often, but not always, co-occur with internal accentuated lines (AL). Monkeys reportedly exhibit fewer enamel defects than hominoids as their presumably faster-growing teeth produce shallower LEH defects that are harder to macroscopically identify. In this case study of a chacma baboon (Papio ursinus) incisor, we assessed whether AL are matched by LEH defects; how enamel extension rates and striae angles relate to the surface distribution of LEH defects; and whether striae angles are shallow and rates of enamel extension fast compared to hominoid anterior teeth. We found a higher prevalence of internal AL (N = 48) compared to external LEH defects (N = 10), which co-occurred in all instances of LEH. However, 79.2% of AL defects do not co-occur with LEH defects. The spatial distribution of AL is more consistent, ranging from 3-10/decile, while LEH defects occur mainly in the midcrown and cervical regions. This incisor exhibits faster extension rates (mean = 23.6 µm/day) and shallower striae angles (11-16°) compared to hominoids, likely creating shallower LEH defects and contributing to the discrepancy between AL and LEH defects.

Wild boar versus domestic pig-Deciphering of crown growth in porcine second molars.

Based on the previously established periodicity of enamel growth marks, we reconstructed crown growth parameters of mandibular second molars from two wild boar and two domestic pigs of the Linderöd breed. Body weight gain and progression of dental development were markedly faster in the domestic pigs than the wild boar. While the final crown dimensions of the M2 did not differ between domestic pigs and wild boar, mean crown formation time (CFT) of this tooth was considerably shorter in the domestic pigs (162 days) than in the wild boar (205 days). The difference in CFT was mainly attributable to a higher enamel extension rate (EER) in the domestic pig. Generally, EER was highest in the cuspalmost deciles of the length of the enamel-dentine-junction and markedly dropped in cervical direction, with lowest values occurring in the cervicalmost decile. In consequence, the cuspal half of the M2 crown was formed about three times faster than the cervical half. In contrast to the EER, no marked difference in daily enamel secretion rate (DSR) was recorded between domestic pigs and wild boar. The duration of enamel matrix apposition as well as linear enamel thickness in corresponding crown portions was only slightly lower in the domestic pigs than the wild boar. Thus, the earlier completion of M2 crown growth in the domestic pig was mainly achieved by a higher EER and not by an increased DSR. The more rapid recruitment of secretory ameloblasts in the course of molar crown formation of domestic pigs compared to wild boar is considered a side-effect of the selection for rapid body growth during pig domestication.

Molar form, enamel growth, and durophagy in Cercocebus and Lophocebus

AbstractObjectivesTo test the hypothesis that differences in crown structure, enamel growth, and crown geometry in Cercocebus and Lophocebus molars covary with differences in the feeding strategies (habitual vs. fallback durophagy, respectively) of these two genera. Relative to Lophocebus molars, Cercocebus molars are predicted to possess features associated with greater fracture resistance and to differ in enamel growth parameters related to these features.Materials and MethodsSample proveniences are as follows: Cercocebus atys molars are from the Taï Forest, Ivory Coast; Lophocebus albigena molars are from a site north of Makoua, Republic of Congo; and a Lophocebus atterimus molar is from the Lomako Forest, Democratic Republic of Congo. For μCT scans on which aspects of molar form were measured, sample sizes ranged from 5 to 35 for Cercocebus and 3 to 12 for Lophocebus. A subsample of upper molars was physically sectioned to measure enamel growth variables.ResultsPartly as a function of their larger size, Cercocebus molars had significantly greater absolute crown strength (ACS) than Lophocebus molars, supporting the hypothesis. Greater crown heights in Cercocebus are achieved through faster enamel extension rates. Also supporting the hypothesis, molar flare and proportional occlusal basin enamel thickness were significantly greater in Cercocebus. Relative enamel thickness (RET), however, was significantly greater in Lophocebus.DiscussionIf ACS is a better predictor of fracture resistance than RET, then Cercocebus molars may be more fracture resistant than those of Lophocebus. Greater molar flare and proportional occlusal basin thickness might also afford Cercocebus molars greater fracture resistance.

Fossil Primate Teeth as an Archive for High Resolution Ontogenic, Dietary and Palaeoclimate Record

ABSTRACT Rhythmic incremental growth of daily or longer duration is faithfully recorded in the tooth enamel of all the mammals including those of primates. Just a single well preserved fossil primate tooth can yield a variety of invaluable information, ranging from gestation period, age at weaning, age at death, prenatal and postnatal diet, ecology, water intake, and even body and brain size of an individual, that lived millions of years ago. Till a few years back fossil teeth of rare primates were physically sectioned to count these rhythmic growth lines for extracting ontogenic information such as, crown formation times, daily secretion rates, rhetzius line periodicity and enamel extension rates. Now all this is changing with the application of non-invasive X-Ray Synchroton micro-tomography. Further, laser ablation based stable carbon and oxygen isotope analyses of these growth layers are showing immense potential for yielding high resolution dietary, ecological and climate data, with minimal visible damage to the very rare fossil specimens.

Variation in the timing of enamel formation in modern human deciduous canines

ObjectivesDeciduous canines are now used increasingly in archaeological and forensic studies to establish the time of birth and as a retrospective source of trace elements incorporated into enamel before and after birth. However, data on the variability of deciduous enamel formation times are scarce. Our objectives were to use daily incremental markings to estimate daily secretion rates, the timing of prenatal, postnatal and total enamel formation and any changes in enamel coverage or prism and stria orientation that occur during enamel formation. Materials and MethodsLongitudinal ground sections of 81 deciduous canines were studied with transmitted light microscopy. High-resolution digital images were imported from an Olympus VS-120 virtual slide scanning system into a geographic information system (ArcGIS, ESRI USA) for quantitative and statistical analyses of linear, angular and area measurements of buccal enamel. ResultsDaily rates of enamel secretion close to the EDJ were faster than in permanent enamel (3.23 μm/day, SD = 0.54). Prism and stria angles subtended to the EDJ both increased through crown formation. Enamel coverage was low in the cusp and cervix but maximal ∼150 days after birth. The mean prenatal enamel formation time was 118 days (range 60−150, SD, 29.2, n = 24). The overall mean postnatal enamel formation time was 319 days (range 210–420, SD 50.6, n = 67). ConclusionsDaily enamel secretion rates compared well with previous studies of deciduous enamel, however, enamel extension rates in deciduous cuspal enamel were notably lower. The variability of both prenatal and postnatal deciduous enamel formation times was greater than previously reported.

Faster growth corresponds with shallower linear hypoplastic defects in great ape canines

Deeper or more ‘severe’ linear enamel hypoplasia (LEH) defects are hypothesized to reflect more severe stress during development, but it is not yet clear how depth is influenced by intrinsic enamel growth patterns. Recent work documented inter- and intraspecific differences in LEH defect depth in extant great apes, with mountain gorillas having shallower defects than other taxa, and females having deeper defects than males. Here, we assess the correspondence of inter- and intraspecific defect depth and intrinsic aspects of enamel growth: enamel extension rates, outer enamel striae of Retzius angles, and linear enamel thickness. Thin sections of great ape canines (n = 40) from Gorilla beringei beringei, Gorilla gorilla gorilla, Pan troglodytes, and Pongo spp. were analyzed. Enamel extension rates were calculated within deciles of enamel-dentine junction length. Linear enamel thickness and the angle of intersection between striae of Retzius and the outer enamel surface were measured in the imbricational enamel. Mountain gorillas have faster enamel extension rates and shallower striae angles than the other taxa examined. Mountain gorillas have thinner imbricational enamel than western lowland gorillas and orangutans, but not chimpanzees. In the combined-taxon sample, females exhibit larger striae angles and thicker imbricational enamel than males. Enamel extension rates are highly negatively correlated with striae angles and LEH defect depth. Enamel growth variation corresponds with documented inter- and intraspecific differences in LEH defect depth in great ape canines. Mountain gorillas have shallower striae angles and faster extension rates than other taxa, which might explain their shallow LEH defect morphology and the underestimation of their LEH prevalence in previous studies. These results suggest that stressors of similar magnitude and timing might produce defects of different depths in one species or sex vs. another, which has implications for interpretations of stress histories in hominins with variable enamel growth patterns.

Dental histology of late Miocene hipparionins compared with extant Equus, and its implications for Equidae life history

Hipparionins were a dominant element of the late Miocene faunas of Europe; however, their biology and ecology remain incompletely understood. In this paper, we explore the pace of life history of different-sized hipparionin horses, using dental histology, and compare it with extant equids. In doing so we consider (i) the size diversity of hipparionins, (ii) their generally smaller size compared to extant equids, and (iii) the allometric coupling between size and life history. In particular, we reconstruct the dental growth in lower first/second molars and in later-formed lower third molars for three hipparionin taxa: two dwarf species (Hipparion periafricanum and H. gromovae), and a larger species (H. concudense). We also analyze dental growth in an extant zebra (Equus quagga) for comparative purposes. Our results reveal that, within each species, there are differences in enamel growth parameters between the first/second molars and third molars. These results illustrate the differences in the developmental timing of these teeth and the existence of a relationship between dental growth parameters with somatic growth. We also find that hipparionin teeth grow at slower rates and tend to erupt later in time than in extant Equus. Dwarf hipparionins, moreover, exhibit lower enamel extension rates than the larger species, but similar formation and eruption times. Considering the link between dental development and life history, these results suggest a slower pace of growth of selected hipparionins compared to extant equids, and a further slower life history than expected for their size in the two dwarf forms.

Enamel formation and growth in non-mammalian cynodonts.

The early evolution of mammals is associated with the linked evolutionary origin of diphyodont tooth replacement, rapid juvenile growth and determinate adult growth. However, specific relationships among these characters during non-mammalian cynodont evolution require further exploration. Here, polarized light microscopy revealed incremental lines, resembling daily laminations of extant mammals, in histological sections of enamel in eight non-mammalian cynodont species. In the more basal non-probainognathian group, enamel extends extremely rapidly from cusp to cervix. By contrast, the enamel of mammaliamorphs is gradually accreted, with slow rates of crown extension, more typical of the majority of non-hypsodont crown mammals. These results are consistent with the reduction in dental replacement rate across the non-mammalian cynodont lineage, with greater rates of crown extension required in most non-probainognathians, and slower crown extension rates permitted in mammaliamorphs, which have reduced patterns of dental replacement in comparison with many non-probainognathians. The evolution of mammal-like growth patterns, with faster juvenile growth and more abruptly terminating adult growth, is linked with this reduction in dental replacement rates and may provide an additional explanation for the observed pattern in enamel growth rates. It is possible that the reduction in enamel extension rates in mammaliamorphs reflects an underlying reduction in skeletal growth rates at the time of postcanine formation, due to a more abruptly terminating pattern of adult growth in these more mammal-like, crownward species.

Reconstructing molar growth from enamel histology in extant and extinct Equus

The way teeth grow is recorded in dental enamel as incremental marks. Detailed analysis of tooth growth is known to provide valuable insights into the growth and the pace of life of vertebrates. Here, we study the growth pattern of the first lower molar in several extant and extinct species of Equus and explore its relationship with life history events. Our histological analysis shows that enamel extends beyond the molar’s cervix in these mammals. We identified three different crown developmental stages (CDS) in the first lower molars of equids characterised by different growth rates and likely to be related to structural and ontogenetic modifications of the tooth. Enamel extension rate, which ranges from ≈400 μm/d at the beginning of crown development to rates of ≈30 μm/d near the root, and daily secretion rate (≈17 μm/d) have been shown to be very conservative within the genus. From our results, we also inferred data of molar wear rate for these equids that suggest higher wear rates at early ontogenetic stages (13 mm/y) than commonly assumed. The results obtained here provide a basis for future studies of equid dentition in different scientific areas, involving isotope, demographic and dietary studies.

New regression formula to estimate the prenatal crown formation time of human deciduous central incisors derived from a Roman Imperial sample (Velia, Salerno, Italy, I-II cent. CE).

The characterization and quantification of human dental enamel microstructure, in both permanent and deciduous teeth, allows us to document crucial growth parameters and to identify stressful events, thus contributing to the reconstruction of the past life history of an individual. Most studies to date have focused on the more accessible post-natal portion of the deciduous dental enamel, even though the analysis of prenatal enamel is pivotal in understanding fetal growth, and reveals information about the mother’s health status during pregnancy. This contribution reports new data describing the prenatal enamel development of 18 central deciduous incisors from the Imperial Roman necropolis of Velia (I-II century CE, Salerno, Italy). Histomorphometrical analysis was performed to collect data on prenatal crown formation times, daily secretion rates and enamel extension rates. Results for the Velia sample allowed us to derive a new regression formula, using a robust statistical approach, that describes the average rates of deciduous enamel formation. This can now be used as a reference for pre-industrial populations. The same regression formula, even when daily incremental markings are difficult to visualize, may provide a clue to predicting the proportion of infants born full term and pre-term in an archaeological series.

Perikymata numbers and enamel extension rates in the incisors of three archaeological modern human populations from two caves located in Spain: Maltravieso Cave (Cáceres) and Mirador Cave (Burgos)

Abstract A number of authors have studied the variability of total perikymata counts on permanent incisors in different modern human populations, finding that some populations show a significantly lower number of perikymata than others. However, little is known about the overall variability of these traits in archaeological populations of modern humans. Our aim was to broaden the available data on total perikymata counts on incisors, and to further discuss enamel extension rate variability in modern human incisors. Our sample consisted of 15 incisors from three different archaeological populations belonging to the Holocene: two populations from Mirador Cave (Sierra de Atapuerca, Burgos) assigned to two different chronological periods, and a population from Maltravieso Cave (Caceres). We refer to these populations as the MTV–MIR sample. Perikymata were counted from several images obtained using an environmental scanning electron microscope (ESEM). Our results suggest the total number of perikymata of the MTV–MIR incisors fall most comfortably within the distribution of counts reported for modern Europeans and Inuit rather than those representing southern African samples of incisors. Furthermore, the percentage of the perikymata numbers in the last five deciles into which the crowns were divided (the cervical half) did not show statistically significant differences when compared to the other modern human populations. This suggests that enamel extension rates among modern human incisors share a common pattern, which might differ from that present in other species. More data from other modern and archaeological sub-Saharan African samples are needed to test whether low perikymata counts are exclusive to the southern African group or are indeed truly representative of sub-Saharan African populations as a whole today and in the past.

Enamel thickness and enamel growth in Oreopithecus: Combining microtomographic and histological evidence

Oreopithecus bambolii, a large-bodied fossil ape, lived in the Tusco-Sardinian archipelago during the Late Miocene, until ca. 6.7Ma. Its dentition, an apparent blend of hominoid and cercopithecoid-like features, has been a matter of discussion since its first description, in 1872. While the height and sharpness of its molar cusps recall some Cercopithecidae, Oreopithecus is currently considered by many as more likely related to dryopithecines. Here, we use microtomographic-based quantitative imaging and histological evidence to link outer and inner tooth structural morphology with enamel development in Oreopithecus permanent teeth. The material consists of 14 teeth/crowns from the sites of Baccinello and Casteani, in Tuscany, and Fiume Santo, in Sardinia. In particular, we add to the record of 2–3D of molar enamel thickness topographic variation and enamel-dentine junction morphology, and using high-resolution replicas of the outer crown and ground sections, comparatively assess molar growth trajectory (crown formation times and enamel extension rates). Our results shed new light on dental development of this “enigmatic anthropoid” and provide additional evidence concerning the still debated question of its evolutionary history.

Environmental change and seasonal behavior of mastodons in the Great Lakes region inferred from stable isotope analysis

We investigate seasonal variations in the diet and drinking water of four Great Lakes mastodon (Mammut americanum) specimens using stable isotope analysis of serially sampled inner-enamel bioapatite structural carbonate (δ13Csc, δ18Osc), and previously published bulk analyses. Isotopic analyses and thin section measurements showed that mastodon tooth enamel extension rates (~12–4mm/yr, decreasing toward the cervix) were lower than those of mammoths or modern elephants. Mastodons had distinct and highly regular seasonal variations in δ13Csc and δ18Osc, which we interpret in the context of local glacial history and vegetation changes. Seasonal variations in δ18O were large but variations in δ13C were small, and may have been obscured if coarser sampling methods than our inner-enamel sampling approach were used. Thus, our approach may be particularly useful for understanding relatively small seasonal changes in δ13C within C3 environments. The seasonal patterns, though not entirely conclusive, suggest that the Ontario mastodons did not migrate over very long distances. Rather, the climate and seasonal dietary patterns of mastodons within the region changed over time, from ~12,400 to 10,400 14CyrBP (~15,000 – 12,000calyrBP). Insights gained using these methods can contribute to a better understanding of megafaunal extinctions and Paleoamerican lifeways.

Ecological and life-history correlates of enamel growth in ruminants (Artiodactyla)

Enamel incremental markings are widely used to reconstruct growth patterns of extinct mammals. However, the likely existence of an allometric relationship between dental morphology and enamel growth suggests that caution is required when making life-history inferences based on these microstructures. In the present study, we aimed to explore the potential of using enamel growth rate as a reliable proxy of the pace of life in fossil species. We sectioned 24 permanent first lower molars from 19 extant and two fossil ruminant species. By using polarized light microscopy, we measured the two parameters that determine enamel growth rates: daily secretion rate (DSR) and extension rate, as quantified by enamel formation front (EFF) angle. These parameters were regressed against body mass, hypsodonty index, and relative age at first reproduction (relative to body mass) as a proxy for the species' pace of life, using phylogenetic generalized least squares analyses. Our results indicate that DSR conforms to the allometric relationship because it is positively correlated with hypsodonty. By contrast, enamel extension rate is strongly related to the pace of life. These findings suggest that the two mechanisms of enamel growth might be subject to different selective forces. The application in two fossil species provides evidence that EFF angle is a reliable proxy of the life history of extinct mammals. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112, 657–667.

Characterization of enamel incremental markings and crown growth parameters in minipig molars.

We studied the structure and periodicity of regular incremental markings in third molar enamel of minipigs. Light microscopy of ground sections revealed the presence of incremental markings matching the description of laminations. Their number within the section planes closely paralleled crown formation time (CFT) in days reported for minipig third molars, thereby indicating the daily nature of laminations. Spacing of consecutive laminations increased from lowest values in the inner to highest values in the outer enamel, where mean daily secretion rates of about 20 µm were recorded. Mean enamel extension rates determined for deciles along the enamel-dentin junction varied between highest values (155 µm/day) in the most cuspally located and lowest values (19 µm/day) in cervical enamel. Backscattered electron imaging in the SEM revealed the presence of thin, regularly spaced hypermineralized incremental lines in the outer enamel portion. These lines exhibited the same spacing as the laminations and were, thus, likewise regarded as daily incremental markings. Between two successive daily incremental markings, subdaily growth marks were discernible in light microscopic and in BSE-SEM images. These subdaily growth marks closely resembled the (daily) prism-cross striations of human enamel. Supra-daily growth marks were not identified in the minipig enamel. The results of this study parallels previous findings in sheep enamel. It is cautioned that CFT of ungulate teeth may be considerably overestimated if the periodicity established for growth marks in human enamel is uncritically transferred to the analysis of morphologically similar growth marks in ungulate enamel.

Lines of evidence-incremental markings in molar enamel of Soay sheep as revealed by a fluorochrome labeling and backscattered electron imaging study.

We studied the structural characteristics and periodicities of regular incremental markings in sheep enamel using fluorochrome injections for vital labeling of forming enamel and backscattered electron imaging in the scanning electron microscope. Microscopic analysis of mandibular first molars revealed the presence of incremental markings with a daily periodicity (laminations) that indicated successive positions of the forming front of interprismatic enamel. In addition to the laminations, incremental markings with a sub-daily periodicity were discernible both in interprismatic enamel and in enamel prisms. Five sub-daily increments were present between two consecutive laminations. Backscattered electron imaging revealed that each sub-daily growth increment consisted of a broader and more highly mineralized band and a narrower and less mineralized band (line). The sub-daily markings in the prisms of sheep enamel morphologically resembled the (daily) prisms cross striations seen in primate enamel. Incremental markings with a supra-daily periodicity were not observed in sheep enamel. Based on the periodicity of the incremental markings, maximum mean daily apposition rates of 17.0 µm in buccal enamel and of 13.4 µm in lingual enamel were recorded. Enamel extension rates were also high, with maximum means of 180 µm/day and 217 µm/day in upper crown areas of buccal and lingual enamel, respectively. Values in more cervical crown portions were markedly lower. Our results are in accordance with previous findings in other ungulate species. Using the incremental markings present in primate enamel as a reference could result in a misinterpretation of the incremental markings in ungulate enamel. Thus, the sub-daily growth increments in the prisms of ungulate enamel might be mistaken as prism cross striations with a daily periodicity, and the laminations misidentified as striae of Retzius with a supra-daily periodicity. This would lead to a considerable overestimation of crown formation times in ungulate teeth.

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.

Mammoth tooth enamel growth rates inferred from stable isotope analysis and histology

Mammoth (Mammuthus sp.) teeth are relatively abundant in Quaternary deposits from Eurasia and North America, and their isotopic compositions can be used to reconstruct past seasonal patterns in precipitation, diet, and migration. Strategies for collecting and interpreting such data, however, are strongly dependent on growth rates, which can vary among species, individuals, and within teeth. In this study, we use histological and isotopic measurements to determine enamel growth rates for a Columbian mammoth (Mammuthus columbi) tooth in two directions. Using histology, the growth rate through the enamel thickness (ET; perpendicular to the height of the tooth) is estimated at 0.8 to 1.5mm/yr. Isotopic sampling through the innermost 0.36mm of the ET recovered less than half a period of variation (i.e., half an inferred year of growth), which is consistent with the histological estimate for ET growth rate. A combination of histological and isotopic measurements suggests that the enamel extension rate (growth in the height of the tooth) is 13–14mm/yr. Knowledge of enamel growth rates should improve the design and interpretation of future isotopic studies of mammoth teeth. The combination of histological and isotopic measurements may also prove useful in determining growth rates for other extinct taxa.

Enamel hypoplasia in molars of sheep and goats, and its relationship to the pattern of tooth crown growth.

Enamel is the most highly mineralized and durable tissue of the mammalian body. As enamel does not undergo remodeling or repair, disturbances of enamel formation leave a permanent record in the tissue that can be used for life history reconstruction. This study reports light and scanning electron microscope findings on hypoplastic enamel defects, and on the chronology of crown growth in the molars of sheep and goats. A marked reduction of enamel extension rates in cervical compared with more cuspal crown portions of sheep and goat molars was recorded, with formation of the cervical 25% of the crown taking about the same time as that of the upper 75% of the crown. This explains the more frequent occurrence of enamel hypoplasia in cervical compared with upper and middle crown portions. Regarding the identification of hypoplastic enamel defects by external inspection, our results suggest a dependence on the type of defect and the associated presence of smaller or larger amounts of coronal cementum. Defects considered to reflect a slight to moderate impairment of secretory ameloblast function can normally be correctly diagnosed as they are not occluded by thick layers of cementum. In contrast, defects denoting a severe impairment of enamel matrix secretion can typically not be correctly identified because they are occluded by large amounts of cementum, so that neither depth nor extension of the defects can be assessed on external inspection. In these cases, microscopic analysis of tooth sections is required for a correct diagnosis of the hypoplastic enamel defects.