A critique of the «increasing population density effect»

The «increasing population density effect» (IPDE) proposed by Macchiarelli and Bondioli (1986) represents one of the most recent attempts to account for dental reduction in modern human populations. Under the IPDE, the marked reduction in tooth size observed in post-Pleistocene human groups is seen as merely a side-effect of a more general reduction in body size, which resulted from an increase in population density. The model is dependent upon a strong correlation between tooth size and body size, which after numerous attempts has yet to be convincingly demonstrated in humans. This paper argues that the IPDE neglects the negative consequences on individual fitness of teeth which are too large to fit into diminishing jaws or are more susceptible to pathology, and that the worldwide reduction of tooth size is the result of selection for smaller teeth due to shifts to a softer and/or more cariogenic diet. Although increased population density may have intensified this selective effect by decreasing general fitness by lowering resistance to oral infections, it was not the primary cause of dental reduction. All proposed mechanisms of dental reduction, however, are in need of additional testing, and possible directions are offered for future studies of the phenomenon.     

Neandertal dental asymmetry and the probable mutation effect

The purpose of this paper is to show that if Brace's hypothesis concerning the role of random mutations in structural reduction (PME) is accepted for the sake of argument, then it cannot be used to explain hominid tooth size reduction since the Mousterian. Specifically it is shown that Brace's Hypothesis implies a smaller variance in dental asymmetry for Neandertal than modern man. Variance analysis reveals that, to the contrary, Neandertal dentitions are more variable than a large sample of modern man's. Finally it is shown that the sequence of dental asymmetry per tooth class is approximately the same in Neandertal as in modern man.     

Posterior dental size reduction in hominids: The Atapuerca evidence

In order to reassess previous hypotheses concerning dental size reduction of the posterior teeth during Pleistocene human evolution, current fossil dental evidence is examined. This evidence includes the large sample of hominid teeth found in recent excavations (1984–1993) in the Sima de los Huesos Middle Pleistocene cave site of the Sierra de Atapuerca (Burgos, Spain). The lower fourth premolars and molars of the Atapuerca hominids, probably older than 300 Kyr, have dimensions similar to those of modern humans. Further, these hominids share the derived state of other features of the posterior teeth with modern humans, such as a similar relative molar size and frequent absence of the hypoconulid, thus suggesting a possible case of parallelism. We believe that dietary changes allowed size reduction of the posterior teeth during the Middle Pleistocene, and the present evidence suggests that the selective pressures that operated on the size variability of these teeth were less restrictive than what is assumed by previous models of dental reduction. Thus, the causal relationship between tooth size decrease and changes in food-preparation techniques during the Pleistocene should be reconsidered. Moreover, the present evidence indicates that the differential reduction of the molars cannot be explained in terms of restriction of available growth space. The molar crown area measurements of a modern human sample were also investigated. The results of this study, as well as previous similar analyses, suggest that a decrease of the rate of cell proliferation, which affected the later-forming crown regions to a greater extent, may be the biological process responsible for the general and differential dental size reduction that occurred during human evolution. © 1995 Wiley-Liss, Inc.     

中国古人类牙齿尺寸演化特点及东亚直立人的系统地位

对中国境内不同时代人类牙齿测量数据的发析显示,中国古人类牙齿尺寸的总体演化趋势与世界其它地区人类一致,呈缩小变化,其中一个表现特点是中国直立人与早期智人在牙齿尺寸上不差别不大,但作者根据对中国古人类化石形态特征,生存年代等方面的综合分析认为中国直立人在化石形态,生存年代等方面均与智人有明显的不同,取消直立人,将其并入智人意见的证据还是不够充分的,目前仍宜将直立人与智人作为人属内两个不同的种来看待,     

Phenotypic evolution in prehistoric Ohio Amerindians: natural selection versus random genetic drift in tooth size reduction

Many anthropologic investigations involve measurement and analysis of polygenic skeletal and dental traits in prehistoric populations from which genetic details cannot be inferred. However, population genetics concepts can be applied productively to analyses of phenotypic variation in prehistoric human populations. One potentially useful approach, derived from basic quantitative genetics (Lande 1976, p. 314), models the effects of natural selection and random genetic drift on the evolution of the average phenotype in a population. We apply this model to the problem of dental size reduction in three prehistoric Amerindian populations from Ohio. Conversion of mean log-transformed buccolingual diameters for six permanent teeth (maxillary and mandibular I1, M1, and M2) to phenotypic standard deviation units reveals significant size reduction in the maxillary teeth only. By assuming 40 generations (t) between the 2 populations and a narrow heritability (h2) range of 0.30-0.70, the estimated minimum selective mortality required to produce the reductions is 1.8 deaths per 100 persons per generation. Given the same t and h2 values, the effective population size (Ne) needed to reject the neutral hypothesis (i.e., random genetic drift) with 95% confidence is approximately 150. Because paleodemographic and ethnographic studies suggest minimum effective sizes of this magnitude for these populations, we tentatively reject random genetic drift and conclude that selective mortality is most probably responsible for the maxillary tooth size reduction observed.     

Metric dental change in the European upper paleolithic and mesolithic.

Evolutionary trends for dental reduction are presented for European Upper Paleolithic and Mesolithic samples. The analysis demonstrates that the greatest decrease in tooth size occurs between the two divisions of the Upper Paleolithic, while little and insignificant change characterizes the Late Upper Paleolithic/Mesolithic transition. Trends for tooth size over this period indicate that (1) human evolution does not stop with the appearance of "anatomically modern Homo sapiens," (2) changes in tooth size fluctuate with increases in the efficiency and complexity of cultural systems, and (3) the Early Upper Paleolithic sample should be considered transitional between Wurm II European Neanderthals and later Upper Paleolithic and Mesolithic groups.     

A new perspective of human origin and dispersals derived from the microevolution of teeth

Recent genetic studies have heightened the expectation that the origin of modern humans will be defined, but one clear vision has yet to be developed. The study of teeth has historically been an informative means to help define human dispersals. Quantitative tooth data is presented encompassing worldwide human populations. A null hypothesis phylogeny developed from the multivariate analysis of the microevolution of the dental phenotype was interpreted to be broadly in accord with the dominant interpretation of genetic, archaeological, and other dental data by showing that the first division in the dispersion of extant humanity was within sub-Sahara Africans; i.e., San, and Western Africans and Bantu. This "out-of-Africa" interpretation of the graphical results suggests that the first modern human African emigrants not to go extinct were Southeast Asian Negritos. All Eurasians then emerged and expanded through a series of extinct antecedent populations branching from the short lineage extending from Negritos to Australian aborigines. Caucasoids were the first group to fission from this stock. Under this hypothesis, the next to have emerged were antecedent Southeast Asians, from which present Southeast Asians and then antecedent east Central Asians then diverged. Independently, people from the region of Mongolia and all Native Americans arose as daughter populations from antecedent east Central Asians. The broad outline of humanity studied here cannot disprove the equally explanatory protean multiregional hypotheses, but with the inclusion of hominids and further modern human populations either parts of the multiregional hypothesis or the outlined more linear evolutionary scenario likely can be refuted.     

Tooth Size Variation in Prehistoric India

Permanent dental crown dimensions are largely unknown for living and prehistoric populations of South Asia. This paper describes permanent tooth crown measurements and indices for the Late Chalcolithic skeletal series from Inamgaon (1700-700 B.C.) in western India. These data are compared with tooth size data for prehistoric populations in India and both living and prehistoric populations outside the South Asian subcontinent. In summed cross-sectional area, the Inamgaon (1,218 mm²) permanent teeth are most similar to prehistoric skeletal series from Mahurjhari, India, Non Nok Tha, Thailand, and Bellan Bandi Palassa, Sri Lanka. The Inamgaon total crown area is only 3.1% smaller than the figure reported for early Neolithic skeletons from Mehr-garh, Pakistan, indicating the relatively large size of the Inamgaon dentition. Large tooth size at Inamgaon is interpreted as a biological adaptation to coarse dietary items, basic food preparation methods, and a mixed economy that included hunting wild game and collecting wild fruit. Dental indices of "ethnic" significance are perhaps better interpreted as indicators of masticatory stress and differential dental reduction. Maxillary and mandibular third molars show minimal reduction in crown size, but mandibular second molar teeth are distinctly reduced in size. The Incisor Breadth Index may indicate mixed genetic ancestry of the Inamgaon people or similar dietary stresses in genetically different populations.     

Dental agenesis in the Dariusleut Hutterite Brethren: comparisons to selected Caucasoid population surveys

We report the results of a study of the prevalence of nonsyndromic dental agenesis among a sample of 208 individuals (105 females, 103 males) between the ages of 15 and 29 years from a North American religious and genetic isolate, the Dariusleut Hutterites of Western Canada. Direct examination of dental casts, oral examination reports, dental treatment records, and a limited number of dental radiographs reveals congenital absence and/or obvious morphometric reduction of at least one tooth (excluding third molars) in 98 subjects (55 females, 43 males), yielding a prevalence estimate of approximately 47%. This estimate is nearly four times those reported for nonisolate Caucasoid populations of European descent and substantially higher than the elevated prevalences observed in several other isolated populations. Although the prevalence of dental agenesis in the Dariusleut is indeed high, neither the incidence of bilateral agenesis (exhibited at least once in 58% of affected dentitions), number of affected teeth per person (mean, 2.4), morphologic tooth classes affected, or combinations of tooth classes affected ostensibly distinguish them from other populations with similar geographic origins. We conclude that the dental agenesis observed in this North American genetic isolate does not represent a private polymorphism or rare developmental variant. Consequently, the results of further study in these Dariusleut Brethren will be directly relevant to critically testing as yet unresolved hypotheses for the mode of gene action and the relative contributions of hereditary and environmental factors to the reduction of tooth numbers in human dentitions.     

A genetic model of dental reduction through the probable mutation effect

A simulation approach is used in order to elucidate the nature of the hypothesized “probable mutation effect” as it applies to dental reduction in man. Computer-generated simulations of the accumulation of mutations in a human gene pool show the results of the proposed model under the influence of various parameters, as well as illustrating the nature of such genetic change through time. This approach supports a polygenic model of the probable mutation effect as a viable hypothesis for an explanation of the dental reduction which has occurred in some human populations over the last 40,000 years.     

Sexual dimorphism and human tooth size differences

Current male/female differences in tooth size are due to the male/female differences in body bulk that exist in any given human population. These differences are residues of the sexual dimorphism that was maintained for adaptive reasons during the Middle Pleistocene. Late in the Pleistocene the development of food processing techniques led to the reduction of both male and female dental dimensions. Dental sexual dimorphism, however, was maintained until the very end of the Pleistocene when the hunting of large game animals by crude techniques was replaced by a focus on great numbers of small game caught by more sophisticated means and by an increasing utilization of plant foods. The subsequent reduction in dimorphism represents the actions of the Probable Mutation Effect operating under conditions of relaxed selection. The conclusion offered is that the smallest degree of sexual dimorphism visible in the modern world is to be found among those populations that are separated by the greatest interval of time from precursors who depended for their survival on a Pleistocene big game hunting mode of subsistence.     

Biomechanical association of dental and temporomandibular pathology in a medieval Nubian population

An analysis of the relationship between oral pathology and degenerative change at the temporomandibular joint (TMJ) was undertaken on an archaeological sample of 122 adult crania from the Medieval site of Kulubnarti in Sudanese Nubia. The crania were sorted into 2 groups: those demonstrating clearly visible bony changes at the joint (TMJ+) and those without visible change (TMJ-). These groups were compared according to 1) age; 2) sex; 3) active dental pathologies (abscesses, caries, partial socket resorption); 4) tooth loss with complete socket resorption; and 5) dental attrition. No statistically significant association was evident between degenerative change at the TMJ and age, active dental pathologies, or dental attrition; however, sex differences and posterior tooth loss with complete socket resorption revealed a significant correspondence to degenerative TMJ changes. Both of these factors agree with the clinical literature and with biomechanical models (most notably that of Hylander) based upon modern populations. Furthermore, the results support the contention that paleopathological conditions can be analyzed from a clinical and functional biomechanical perspective.     

Dental reduction in post-pleistocene Nubia

Tooth size changes among Nubian archaeological populations dating from the Mesolithic through the Christian era, a period of approximately 12,000 years, are analyzed. Standard length and breadth dimensions of all permanent teeth from several cultural horizons are combined to form three large samples: Mesolithic, 10000-70000 B.C.; Agriculturalist, 3300-1100 B.C. (A-group, C-group, Pharaonic); and Intensive Agriculturalist, A.D. 0-1400 (Meroitic, X-Group, Christian). Such information not only fills a void in the knowledge of Nubian skeletal biology, but also provides a much needed African reference point for the comparison of tooth size data among anatomically modern Homo sapiens from various regions of the world. Changes in mean tooth size and associated t-tests reveal strong and significant reduction in dental size between the Mesolithic and Agriculturalist samples, followed by a continued although diminished trend of reduction for only the molar teeth between the two Agriculturalist groups. These patterns are best observed by examining tooth breadths, which are considered as the most reliable indicator of tooth size. Previously published odontometrics of the Nubian Mesolithic are briefly compared to the findings of this diachronic analysis of Nubian dental change.     

Wear Fast,Die Young: More Worn Teeth and Shorter Lives in Iberian Compared to Scottish Red Deer

Teeth in Cervidae are permanent structures that are not replaceable or repairable; consequently their rate of wear, due to the grinding effect of food and dental attrition, affects their duration and can determine an animal''s lifespan. Tooth wear is also a useful indicator of accumulative life energy investment in intake and mastication and their interactions with diet. Little is known regarding how natural and sexual selection operate on dental structures within a species in contrasting environments and how these relate to life history traits to explain differences in population rates of tooth wear and longevity. We hypothesised that populations under harsh environmental conditions should be selected for more hypsodont teeth while sexual selection may maintain similar sex differences within different populations. We investigated the patterns of tooth wear in males and females of Iberian red deer (Cervus elaphus hispanicus) in Southern Spain and Scottish red deer (C. e. scoticus) across Scotland, that occur in very different environments, using 10343 samples from legal hunting activities. We found higher rates of both incisor and molar wear in the Spanish compared to Scottish populations. However, Scottish red deer had larger incisors at emergence than Iberian red deer, whilst molars emerged at a similar size in both populations and sexes. Iberian and Scottish males had earlier tooth depletion than females, in support of a similar sexual selection process in both populations. However, whilst average lifespan for Iberian males was 4 years shorter than that for Iberian females and Scottish males, Scottish males only showed a reduction of 1 year in average lifespan with respect to Scottish females. More worn molars were associated with larger mandibles in both populations, suggesting that higher intake and/or greater investment in food comminution may have favoured increased body growth, before later loss of tooth efficiency due to severe wear. These results illustrate how independent selection in both subspecies, that diverged 11,700 years BP, has resulted in the evolution of different longevity, although sexual selection has maintained a similar pattern of relative sex differences in tooth depletion. This study opens interesting questions on optimal allocation in life history trade-offs and the independent evolution of allopatric populations.     

Asynchronous dentofacial development and dental crowding: a cross-sectional study in a contemporary sample of children in France

Background

The causes of dental crowding are not fully understood, but it may result from an evolutionary trend towards reduced facial volume, without a proportional reduction in tooth sizes. Most previous studies conducted among modern humans have revealed a very low or non-existent correlation between tooth size and jaw size. Cross-comparison between dental age and facial skeletal age could help to provide better knowledge of the dynamic process of dental crowding. The primary objective of this research was to study the synchronism of dental maturation and skeletal facial growth in a sample of modern children living in France. The secondary objective was to assess the link between dentofacial asynchronism and dental crowding.

Results

The random sample comprised 28 subjects (16 girls, 12 boys). Mean chronological age was 13.5 years (±2.1; range 9.2–17.6). Mean dental age was 14.2 years (±2.8; range 7.5–17) and mean facial skeletal age was 12.8 years (±2.6, range 7–22). In the estimations of dental age and facial skeletal age, there was no evidence of systematic bias. There were 10 subjects (9 girls, 1 boy) with asynchronous dentofacial development. Finally, there were 13 subjects (8 girls, 5 boys) with dental crowding. A significant association was found between delayed facial skeletal growth/advanced dental maturation and dental crowding (P = 0.01).

Conclusions

Dental maturation and facial growth are not necessarily synchronous. Further understanding of the interactions between dental maturation and facial growth could have crucial implications in biological anthropology, as well as for the clinical practice of orthodontists. From an anthropological perspective, this study suggests that asynchronous dentofacial development could, at least partially, explain the frequency of dental crowding in modern populations.     

Regions of lower crossing over harbor more rare variants in African populations of Drosophila melanogaster

A correlation between diversity levels and rates of recombination is predicted both by models of positive selection, such as hitchhiking associated with the rapid fixation of advantageous mutations, and by models of purifying selection against strongly deleterious mutations (commonly referred to as "background selection"). With parameter values appropriate for Drosophila populations, only the first class of models predicts a marked skew in the frequency spectrum of linked neutral variants, relative to a neutral model. Here, we consider 29 loci scattered throughout the Drosophila melanogaster genome. We show that, in African populations, a summary of the frequency spectrum of polymorphic mutations is positively correlated with the meiotic rate of crossing over. This pattern is demonstrated to be unlikely under a model of background selection. Models of weakly deleterious selection are not expected to produce both the observed correlation and the extent to which nucleotide diversity is reduced in regions of low (but nonzero) recombination. Thus, of existing models, hitchhiking due to the recurrent fixation of advantageous variants is the most plausible explanation for the data.     

Heterochronic quantitative microevolution: Dental divergence in aboriginal Americans

Studies of tooth crown morphology alone have proven valuable in defining human population differentiation. We test the hypothesis that quantitative comparisons of more complex whole tooth structure may prove informative in understanding human diversity. Three disparate populations in Native American genetic history were compared: Kodiak Island Western Eskimos, Peruvian Inca Amerindians, and Southeast Asians. Enamel depth (an increasing gradient extended from Southeast Asians to the Inca) and root parameters were the most discriminating variables. The observed microevolution appears to result from variation in timing of known X-linked, Y-linked, and autosomal genes that affect either ameloblast or odontoblast differentiation. The dental traits were sexually dimorphic, the effect being more pronounced in aboriginal Americans, with male teeth having robust roots and thin enamel compared to female. Southeast Asians were isometrically related. The prominence of sexual dimorphism and the importance of sex-linked genes in the determination of the dental phenotypes suggest that sexual selection was one evolutionary force acting on early Asian populations. Subsequently, the selection appears to have been relaxed in Southeast Asians. Observed divergence of tooth shape among the populations, i.e., differences in the appropriation process of tooth primordia, was mainly the consequence of genetic drift modulating heterochronic regulators of homeotic genes. © 1996 Wiley-Liss, Inc.     

Dental agenesis as evidence of possible genetic isolation in the colobine monkey,Rhinopithecus roxellana

In a sample of 24 skulls of the rare colobine monkey of China,Rhinopithecus roxellana, agenesis of the permanent upper third premolar was observed in 8 specimens (33%). Two of the individuals (8% of the sample) showed unilateral agenesis, while six (25%) showed bilateral agenesis. All of the affected individuals appear to have originated from two areas in central Sichuan Province in China, Moupin (Baoxing) and Wen Chuan, that lie approximately 120 km apart on the western rim of the Sichuan Basin. In this species agenesis of the upper third premolar was generally accompanied by rotation of the upper fourth premolar, but not by any other observable variation in tooth size, morphology, or number. Further, dental agenesis in this species appears to have had no effect craniofacial morphology. In certain human populations, a high prevalence of dental agenesis has been associated with small population effects and genetic isolation. Premolar agenesis inR. roxellana can probably be traced to a similar origin. Reconstruction of the evolutionary history of the species indicates that a severe reduction of the geographic range of the species occurred as a result of climatic deterioration during the Late Pleistocene. Although large numbers of populations appear to have become extinct, others survived to give rise to the modern populations of the species that inhabit China today. The two populations showing a high prevalence of premolar agenesis appear to have originated from one that passed through a population bottleneck and suffered the consequences of founder effect in the Late Pleistocene. This interpretation is supported by evidence of dental agenesis in a population of an insectivore species from the same region and by the fact that premolar agenesis is not found in any of the other species ofRhinopithecus. There is no evidence to support the interpretation that dental agenesis inR. roxellana is due to natural selection, mutation or an evolutionary force other than a small population effect.     

The role of deleterious mutations in allopatric speciation

Allopatric speciation is often assumed to occur as a consequence of adaptive divergence between two isolated populations. However, there are some scenarios in which reproductive isolation can be favored due to accumulated unconditionally deleterious mutations. If deleterious mutations have synergistic epistatic effects, it is shown here that the average fitness of recombinants between two parental lines with a given number of fixed mutations is lower than that of the parents in both the F1 and F2 generations. If individual mutations are only slightly deleterious, then they will tend to fixation at a high enough rate to cause lower hybrid fitness. If the fitness effects of mutation give rise to antagonistic epistasis, the hybrids tend to have a higher average fitness than the parental lines, suggesting a possible scenario for the origin of hybrid vigor. The other model of deleterious mutations investigated is the accumulation of knockout mutants in a duplicated gene family. While neutral in the parental lines, upon contact the F1 and later generations have a significant probability of carrying double knockouts. Under this scenario, selection may also favor reproductive isolation between the two lines. Even when the selection coefficients generated are too low to drive speciation, epistatic interactions between deleterious mutations offer a possible explanation for both outbreeding depression and hybrid vigor.     

A Common Variation in EDAR Is a Genetic Determinant of Shovel-Shaped Incisors

Shovel shape of upper incisors is a common characteristic in Asian and Native American populations but is rare or absent in African and European populations. Like other common dental traits, genetic polymorphisms involved in the tooth shoveling have not yet been clarified. In ectodysplasin A receptor (EDAR), where dysfunctional mutations cause hypohidrotic ectodermal dysplasia, there is a nonsynonymous-derived variant, 1540C (rs3827760), that has a geographic distribution similar to that of the tooth shoveling. This allele has been recently reported to be associated with Asian-specific hair thickness. We aimed to clarify whether EDAR 1540C is also associated with dental morphology. For this purpose, we measured crown diameters and tooth-shoveling grades and analyzed the correlations between the dental traits and EDAR genotypes in two Japanese populations, inhabitants around Tokyo and in Sakishima Islands. The number of EDAR 1540C alleles in an individual was strongly correlated with the tooth-shoveling grade (p = 7.7 × 10⁻¹⁰). The effect of the allele was additive and explained 18.9% of the total variance in the shoveling grade, which corresponds to about one-fourth of the heritability of the trait reported previously. For data reduction of individual-level metric data, we applied a principal-component analysis, which yielded PC1-4, corresponding to four patterns of tooth size; this result implies that multiple factors are involved in dental morphology. The 1540C allele also significantly affected PC1 (p = 4.9 × 10⁻³), which denotes overall tooth size, and PC2 (p = 2.6 × 10⁻³), which denotes the ratio of mesiodistal diameter to buccolingual diameter.