Developmental origins and homologies of the hyracoid dentition

Understanding the origins of morphological specializations in mammals is a key goal in evolutionary biology. It can be accomplished by studying dental homology, which is at the core of most evolutionary and developmental studies. Here, we focused on the evolution and development of the specialized dentition of hyraxes for which dental homologies have long been debated, and could have implications on early placental evolution. Specifically, we analysed dental mineralization sequences of the three living genera of hyraxes and 17 fossil species using X‐ray computed microtomography. Our results point out the labile position of vestigial upper teeth on jaw bones in extant species, associated with the frequently unusual premolar shape of deciduous canines over 50 Ma of hyracoid evolution. We proposed two evolutionary and developmental hypotheses to explain these original hyracoid dental characteristics. (a) The presence of a vestigial teeth on the maxilla in front of a complex deciduous canine could be interpreted as extra‐teeth reminiscent of early placental evolution or sirenians, an order phylogenetically close to hyracoids and showing five premolars. (b) These vestigial teeth could also correspond to third incisors with a position unusually shifted on the maxilla, which could be explained by the dual developmental origin of these most posterior incisors and their degenerated condition. This integrative study allows discussion on the current evolutionary and developmental paradigms associated with the mammalian dentition. It also highlights the importance of nonmodel species to understand dental homologies.     

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.     

Crown dimensions of deciduous teeth from pre-Columbian Peru

The mean mesiodistal and buccolingual diameters are presented for the deciduous teeth from Pre-Columbian Peru. Generally, the deciduous teeth from Pre-Columbian Peru are larger in most dimensions than the deciduous teeth of modern populations of European descent and smaller than those of modern Australian aboriginals. Differences in crown dimensions between the Pre-Columbian Peruvian deciduous teeth and those from Prehistoric Inamgaon and Mesolithic Europe are inconsistent. However, the maxillary and mandibular second molars are larger in the Peruvian population while the incisors are smaller. Since comparative data for prehistoric deciduous teeth are rare, this paper adds valuable data to the growing literature on deciduous crown dimensions.     

Increasing human tooth length between birth and 5.4 years

Most previous studies of tooth development have used fractional stages of tooth formation to construct growth standards suitable for aging juvenile skeletal material. A simple alternative for determining dental age is to measure tooth length throughout development. In this study, data on tooth length development are presented from 63 individuals of known age at death, between birth and 5.4 years, from an archeological population recovered from the crypt of Christ Church, Spitalfields, London. Isolated developing teeth (304 deciduous, 269 permanent) were measured in millimeters and plotted against individual age. Regression equations to estimate age from a given tooth length, are presented for each deciduous maxillary and mandibular tooth type and for permanent maxillary and mandibular incisors, canines, and first permanent molars. Data on the earliest age of root completion of deciduous teeth and initial mineralization and crown completion of some permanent teeth in this sample are given, as well as the average crown height and total tooth length from a small number of unworn teeth. This method provides an easy, quantitative and objective measure of dental formation appropriate for use by archeologists and anthropologists. © 1993 Wiley-Liss, Inc.     

Homologies of the anterior teeth in Indriidae and a functional basis for dental reduction in primates

In a recent paper Schwartz ('74) proposes revised homologies of the deciduous and permanent teeth in living lemuriform primates of the family Indriidae. However, new evidence provided by the deciduous dentition ofAvahi suggests that the traditional interpretations are correct, specifically: (1) the lateral teeth in the dental scraper of Indriidae are homologous with the incisors of Lemuridae and Lorisidae, not the canines; (2) the dental formula for the lower deciduous teeth of indriids is 2.1.3; (3) the dental formula for the lower permanent teeth of indriids is 2.0.2.3; and (4) decrease in number of incisors during primate evolution was usually in the sequence I3, then I2, then I1. It appears that dental reduction during primate evolution occurred at the ends of integrated incisor and cheek tooth units to minimize disruption of their functional integrity.     

Formation of anterior maxillary teeth in 0-3-year-old children with cleft lip and palate and prenatal risk factors for delayed development

Formation of deciduous and permanent maxillary incisors and canines was studied in 361 children with cleft lip and palate. Altogether, 704 occlusal X rays of the children aged from 2 weeks to 41 months were assessed using a 13-stage scale. The early timing of tooth development was in both dentitions close to that reported for healthy children. However, developmental asymmetry was noted between contralateral teeth on the cleft and noncleft sides. In the unilateral cleft lip and palate group, every tenth cleft-side deciduous central incisor, deciduous canine, and permanent central incisor was delayed compared to the corresponding contralateral tooth. The formation timing was also compared within 11 prenatal and hereditary variables. Risk factors for delayed development were low placental weight, short gestation, mother's use of drugs during pregnancy, first in birth order, cleft with multiple additional malformations, and relatives having malformations other than oral clefts. However, the associations between a risk factor and tooth development were weak.     

Morphological features of Jat dentition

Observations on morphological characters of milk and permanent teeth, based on 648 pairs of dental casts of 356 male and 292 female Jat children of Haryana (India) are reported. Deciduous teeth show high frequencies of bilateral winging of maxillary central incisor, Carabelli's cusp of maxillary second molar, and deflecting wrinkle of mandibular second molar. Reduction of maxillary molar cusps is more marked in males than in females. Y pattern is very common in deciduous molars. Permanent teeth have high frequencies of grooved cingulum of incisors, cingular nodule of lateral incisors and canines, and distal accessary ridge of canines. Low frequencies of Carabelli's cusp and winging are also common. The tendency towards faintly developed shovelling in milk incisors occurs more often than in the permanent teeth.     

Tooth eruption in Reeves' muntjac (Muntiacus reevesi) and its use as a method of age estimation (Mammalia: Cervidae)

The sequence of tooth eruption and replacement in Reeves' muntjac was determined from captive animals of known age. Pronounced sexual dimorphism is shown by the permanent upper canine which in the male is large, tusk-like and is used as a weapon. The upper canine was the first deciduous tooth to be replaced in males, at approximately 21 weeks of age, compared with 53–57 weeks in the female. The permanent mandibular teeth erupted in the order: molars, first and second incisors, premolars, third incisor and canine. The maxillary teeth erupted in the order: first molar, canine (in male), second and third molars, canine (in female), premolars. The full complement of 34 functional permanent teeth was attained by 83–92 weeks of age.     

Prevalence and age at development of enamel hypoplasias in Mexican children

Enamel hypoplasias, deficiencies in enamel thickness resulting from disturbances during the secretory phase of enamel development, are generally believed to result from nonspecific metabolic and nutritional disruptions. However, data are scare on the prevalence and chronological distributions. of hypoplasias in populations experiencing mild to moderate malnutrition. The purpose of this article is to present baseline data on the prevalences and chronological distributions of enamel hypoplasias, by sex and for all deciduous and permanent anterior teeth, in 300 5 to 15-year-old rural Mexican children. Identification of hypoplasias was aided by comparison to a published standard (Federation Dentaire Internationale: Int. Dent. J. 32(2):159-167, 1982). The location of defects, by transverse sixths of tooth crowns, was used to construct distributions of defects by age at development. One or more hypoplasias were detected in 46.7% (95% CI = 40.9-52.5%) of children. Among the unworn and completely erupted teeth, the highest prevalence of defects was found on the permanent maxillary central incisors (44.4% with one or more hypoplasias), followed by the permanent maxillary canine (28.0%) and the remaining permanent teeth (26.2 to 22.2%) Only 6.1% of the completely erupted and unworn deciduous teeth were hypoplastic. The prevalence of enamel defects on the permanent teeth was up to tenfold greater than that found in studies of less marginal populations that used the FDI method. The prevalence of defects in transverse zones suggests a peak frequency of hypoplasias during the second and third years for the permanent teeth, corresponding to the age at weaning in this group. In the deciduous teeth, a smaller peak occurs between 30 and 40 weeks post gestation. The frequency of defects after three years of age is slightly higher in females than males, suggesting a sex difference in access to critical resources.     

Supernumerary deciduous incisors and the order of eruption of the incisor teeth in the albino ferret

A radiological study of the developing dentition of the domestic albino ferret has revealed the presence of four upper and three lower deciduous functionless incisor teeth in each half of the jaw. The presence of the additional upper incisor tooth can only be accounted for in two ways:
(1) It arose as the result of a gene mutation within the closed colony.
(2) It is indicative of the fact that at some early date in the evolution of the Carnivora, four deciduous incisors were functional in each half of the jaw.     

Dentition development and budding morphogenesis

The development of functional teeth in the mouse has been widely used as a model to study general mechanisms of organogenesis. Compared with other mammals, in which three incisors, one canine, four premolars, and three molars may occur even in each dental quadrant, the mouse functional dentition is strongly reduced. It comprises only one incisor separated from three molars by a toothless gap diastema at the location of the missing teeth. However, mouse embryos also develop transient vestigial dental primordia between the incisor and molar germs in both the upper and lower jaws. These rudimental structures regress, and epithelial apoptosis is involved in this process. The existence of the vestigial dental structures allowed a better assessment of the periodicity in the mouse dentition, which extends opportunities for the interpretation of molecular data on tooth development. We compared the dentition development with tentative models of budding morphogenesis in other epithelial appendages lungs and feathers. We suggested how developmental control by signaling molecules, including bone morphogenetic protein (Bmp), sonic hedgehog (Shh), and fibroblast growth factor (Fgf), can be similarly involved during budding morphogenesis of dentition and other epithelial appendages. We propose that epithelial apoptosis plays an important role in achieving specific features of dentition, whose development involves both budding and its more complex variant branching. The failure of segregation of the originating buds supports the participation of the concrescence of several tooth primordia in the evolutionary differentiation of mammalian teeth.     

Longitudinal analysis of deciduous tooth emergence: IV. Covariate effects in Japanese children

Emergence of the deciduous teeth is generally considered to be robust to moderate environmental insults, malnutrition, and disease. Consequently, deciduous tooth emergence has been used to assess growth and development and for age estimation in children. In this paper, we examine the way in which nutritional status and other covariates affect deciduous tooth emergence in a sample of 114 Japanese children born in Tokyo in 1914 and 1924. Parametric survival analysis was used to quantify the effects of nutritional status, breastfeeding behavior, and sex on the hazard of deciduous tooth emergence. Children of poor nutritional status exhibited significantly delayed emergence of all deciduous teeth, with effects that ranged from 14-29% increases in mean emergence times. Children of medium nutritional status exhibited increases in mean emergence times of 5-9% for the canines and lower molars, and 13-17% for the incisors. Partial breastfeeding had no effect on tooth emergence, but children who were not breastfed at all showed delayed emergence of the upper incisors. No significant sex differences in emergence were found. The findings contradict the idea that moderate malnutrition has little effect on deciduous tooth emergence. Furthermore, nutritional differences may account for some of the observed differences among populations in the timing of tooth emergence.     

Precise chronology of differentiation of developing human primary dentition

While correlation of developmental stage with embryonic age of the human primary dentition has been well documented, the available information regarding the differentiation timing of the primary teeth was largely based on the observation of initial mineralization and varies significantly. In this study, we aimed to document precise differentiation timing of the developing human primary dentition. We systematically examined the expression of odontogenic differentiation markers along with the formation of mineralized tissue in each developing maxillary and mandibular teeth from human embryos with well-defined embryonic age. We show that, despite that all primary teeth initiate development at the same time, odontogenic differentiation begins in the maxillary incisors at the 15th week and in the mandibular incisors at the 16th week of gestation, followed by the canine, the first primary premolar, and the second primary premolar at a week interval sequentially. Despite that the mandibular primary incisors erupt earlier than the maxillary incisors, this distal to proximal sequential differentiation of the human primary dentition coincides in general with the sequence of tooth eruption. Our results provide an accurate chronology of odontogenic differentiation of the developing human primary dentition, which could be used as reference for future studies of human tooth development.     

Variation in elemental intensities among teeth and between pre- and postnatal regions of enamel

Microspatial analyses of the trace element composition of dental enamel are made possible using laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS). Fine spatial resolution, multielement capabilities, and minimal sample destruction make this technique particularly well-suited for documenting the distribution of elements in sequentially calcifying layers of enamel. Because deciduous enamel forms from week 13 in utero up to 9 months postnatally (thereafter essentially becoming inert), the application of LA-ICP-MS allows for the retrospective measurement of prenatal and early postnatal trace-element uptake during a critical period of child development. In this study, we compared intra- and intertooth intensities of 25Mg, 57Fe, 66Zn, 68Zn, 88Sr, 138Ba, and 208Pb via LA-ICP-MS of 38 exfoliated deciduous incisors and canines donated by 36 participants in the Solís Valley Mexico Nutrition Collaborative Research Support Program (NCRSP). Pre- and postnatal comparisons within teeth showed significant increases (P < 0.001) and greater variation in the abundance of all isotopes in postnatal enamel, with the exception of a decrease in 25Mg (P < 0.001) and constant values for 88Sr (P = 0.681). Conversely, comparisons by tooth type and mouth quadrant revealed few significant differences between teeth of the same individual. We argue that more variation in the trace element composition of teeth occurs across developmental areas within a tooth than among different teeth of the same person. This study further demonstrates that sequentially calcifying areas of enamel have different chemical concentrations. The results support the use of microspatial analyses of enamel for understanding changes in nutrition, pollution, and residence.     

Enamel hypoplasias as indicators of developmental stress in pongids and hominids

This paper reports the results from a study of enamel hypoplasia in the deciduous dentition of free-living Liberian chimpanzees (Pan troglodytes verus). The entire study collection includes 280 specimens (278 skulls plus two unassociated mandibles), of which 70 are young enough to retain some decidous teeth. Among the subsample of infants, the total frequency of hypoplasia summed over all teeth in any individual reaches 80%, being expressed in the form of pits rather than transverse lines. Hypoplasia of a linear form was common in the permanent dentition, attaining a frequency of 46.7% on maxillary central incisors and 69.7% on mandibular canines.     

On the development of the deciduous teeth in the common seal (Phoca vitulina)

Information on the development of teeth in seals is rare. In this study, histological details are presented from fetuses and several postnatal stages of the common seal (Phoca vitulina), showing that the deciduous teeth undergo a very rapid, short-term, and somewhat incomplete development. This means that the permanent dentition can and also has to ‚out-pace’ the deciduous dentition, because Phoca vitulina, contrary to fur seals, has a very short suckling period after which there is an abrupt change to normal feeding (crabs, fishes).     

Dentition of a mesolithic population from Wadi Halfa, Sudan

The dentition of a Mesolithic population (8,000–11,000 years old) from Wadi Halfa, Sudan, can be characterized as morphologically complex, massive and relatively free from pathology except that associated with extreme wear. Maxillary incisors show shoveling. All of the maxillary molars show some development of the hypocone. Supernumerary cusps appear frequently. Almost one-half of the second lower molars observed show distally located third cusps. Over one-half of the maxillary third molars show an extra cusp. A high percentage of mandibular molars show six cusps. Overall the teeth from this population compare favorably in size with those of an Australian aborigine tribe and are slightly larger than the teeth of the Neanderthaloids from Skūhl. The teeth show wear of a degree greater than that found in present day Eskimo or Australian aborigine groups. This data may be interpreted as indicating that this Mesolithic group was subjected to rigorous selective pressures favoring large and/or morphologically complex teeth. This pressure was apparently intensive wear, presumably caused by the inclusion of large amounts of grit in the diet through the eating of vegetable food macerated on coarse grinding stones.     

Human tooth development,tooth length and eruption; a study of British archaeological dentitions

Abstract

Human teeth erupt during root growth but few studies report the relationship between fractions of root development and eruption levels. The aim of this study was to assess root stages of deciduous and early erupting permanent teeth (maxillary and mandibular central incisors and first molars) at eruption levels and relate this to root fraction and tooth length. The sample consisted of 620 modern human skeletal remains with developing teeth. Tooth stage (based on Moorrees crown and root stages) and eruption levels of all developing teeth were assessed where possible. Tooth length of isolated teeth was measured. The distribution of root stage at eruption levels was calculated. Results showed that root stage at alveolar eruption was less variable than at partial eruption. Most teeth (72% of 138) at alveolar eruption were at root a quarter (R¼) whereas teeth at partial eruption were at R¼ or root half (R½) (38 and 50% of 128 respectively). These findings suggest that the active phase of eruption is probably a rapid process and occurs during the first half of root growth.     

Arch form,tooth size,and occlusomandibular kinesis in the Ceboidea

Correlations between dental morphology, arch configuration, and jaw movement patterns were quantitatively investigated in 23 ceboid species to elucidate integrative aspects of occlusal functional anatomy in an adaptive and evolutionary context. Differential maxillary-mandibular arch widths are primary in guiding lateral jaw movements. These movements are characterized according to their associated condylar shifts as either predominantly translatory or rotational. Predominantly translatory movements result from peripheral contact relationships between maxillary arches which are considerably wider posteriorly than their opposing mandibular arches. The greatest degree of mandibular movement is in the molar region in functional association with wide “primitive” maxillary molars, narrow mandibular molars, constricted maxillary intercanine widths, and narrow maxillary incisors. In contrast, predominantly rotational masticatory jaw movements result from differential arch widths which are greatest in the maxillary canine and incisor regions. Here most jaw movement is in the anterior segment and this is reflected in small maxillary-mandibular molar width differences, a high degree of premolarization, wide-set maxillary canine teeth, and wide maxillary incisors. Possible selectional factors in the putative evolution of rotational predominance in mastication from the more primitive translatory pattern are discussed.     

Prenatal dental development in the black howler monkey (Alouatta caraya)

Prenatal dental development was investigated in 19 black howler monkey fetuses that ranged in crown-rump length from 84 mm to 170.5 mm. During this period of fetal development, all of the deciduous teeth and the first permanent molars were found to pass through various stages of crown and root formation. The deciduous incisors, canines, and first molars each calcified from a single calcification center. The second deciduous molars each calcified from two separate calcification centers. And, the third deciduous and first permanent molars each calcified from four separate calcification centers. The morphogenetic events involved in the establishment of the molar crown patterns were found to differ markedly from those of their counterparts among catarrhine primates. In all cases, however, the calcification sequence was found to be directly related to the morphology of the developing crown.