Tooth crown heights, tooth wear, sexual dimorphism and jaw growth in hominoids

The aim of this review is to bring together data that link tooth morphology with tooth function and tooth growth: We aim to show how the microanatomy of hominoid teeth is providing evidence about rates of tooth growth that are likely to be a consequence of both masticatory strategy and social behaviour. First, we present data about incisor and molar tooth wear in wild short chimpanzees that demonstrate how crown heights are likely to be related to relative tooth use in a broad sense. Following this we review recent studies that describe the microanatomy of hominoid tooth enamel and show how these studies are providing evidence about tooth crown formation times in hominoids, as well as improving estimates for the age at death of certain juvenile fossil hominids. Next, we outline what is known about the mechanisms of tooth growth in the sexually dimorphic canine teeth of chimpanzees and compare these patterns of growth with tooth growth patterns in the canines of three fossil hominids from Laetoli, Tanzania. Finally, we discuss how selection pressures that operate to increase or reduce the size of anterior teeth interact with jaw size. We argue that the space available to grow developing teeth in the mandibles of juvenile hominoids is determined by the growth patterns of the mandibles, which in turn reflect masticatory strategy. The consequences of selection pressure to grow large or small anterior teeth are likely to be reflected in the times at which these teeth are able to emerge into occlusion.     

The growth of the freshwater crayfish A ustropotamobius pallipes in Northumbria

SUMMARY. Growth increments, moult frequency and growth rates of individuals in a population of the freshwater crayfish Austropotamobius pallipes were followed for a period of 3 years during a mark-recapture study. There was an inverse relationship between body size and the growth increment relative to body size. The absolute increment increased from the juvenile to the young adult stage and thereafter declined with increasing body size.
Differences between the adult male, reproductive female, non-reproductive female and juvenile subpopulations in the size of growth increments are reported and growth increments during the two major moult periods of each year are compared. The effect of the endoparasite Thelohania contejeani and of chela regeneration was to reduce slightly the growth increments.
Instantaneous growth rates declined throughout life in both sexes, but adult males maintained higher growth rates than adult females, particularly in the largest size class studied. The effect of reproduction on growth appeared to be most severe in the smallest breeding females. Males maintained the tendency to moult twice annually to a larger size than females. A simplified construction of the relationship between size and age in the population is presented for each sex. The oldest animals were estimated to be at least 11 years old.     

Evolution of morphogenesis in 360‐million‐year‐old conodont chordates calibrated in days

SUMMARY Highly rhythmic increments of crown tissue are identifiable in conodont oral apparatus elements from the Late Devonian of the Holy Cross Mountains, Poland; individual laminae being of thickness comparable with daily increments of vertebrate tooth enamel and fish otoliths. Abundant occurrence of such specimens enables bed‐by‐bed (stratophenetic) studies of the process of evolution at the population level and quantitative presentation of the evolution of ontogeny in the sampled geological section covering several million years. The morphologic transformation is expressed as expansion of a juvenile asymmetry to later stages of the ontogeny and in decrease of the mature element width, which was due to a change of the mineral tissue secretion rate. It was not just a simple extension of a juvenile character into the later stage of the ontogeny (heterochrony) but rather a true developmental novelty. The evolution was gradual and very slow. The proposed quantitative approach to growth increments in the mineral skeleton of ancient chordates introduces real‐time units to evolutionary developmental studies connected with direct paleontological evidence on the course of evolution.     

Evaluating the annual nature of juvenile rings in Bolivian tropical rainforest trees

Knowledge on juvenile tree growth is crucial to understand how trees reach the canopy in tropical forests. However, long-term data on juvenile tree growth are usually unavailable. Annual tree rings provide growth information for the entire life of trees and their analysis has become more popular in tropical forest regions over the past decades. Nonetheless, tree ring studies mainly deal with adult rings as the annual character of juvenile rings has been questioned. We evaluated whether juvenile tree rings can be used for three Bolivian rainforest species. First, we characterized the rings of juvenile and adult trees anatomically. We then evaluated the annual nature of tree rings by a combination of three indirect methods: evaluation of synchronous growth patterns in the tree- ring series, ¹⁴C bomb peak dating and correlations with rainfall. Our results indicate that rings of juvenile and adult trees are defined by similar ring-boundary elements. We built juvenile tree-ring chronologies and verified the ring age of several samples using ¹⁴C bomb peak dating. We found that ring width was correlated with rainfall in all species, but in different ways. In all, the chronology, rainfall correlations and ¹⁴C dating suggest that rings in our study species are formed annually.     

The effect of unerupted permanent tooth crowns on the distribution of masticatory stress in children

Human mothers wean their children from breast milk at an earlier developmental stage than do ape mothers, resulting in human children chewing solid and semi-solid foods using the deciduous dentition. Mechanical forces generated by chewing solid foods during the post-weaning period travel through not only the deciduous teeth, but also the enamel caps of the developing permanent teeth within the maxilla and mandible, which are not present in the adult face. The effects of mechanical stress propagating through these very stiff structures have yet to be examined. Based on a heuristic model, we predicted that the enamel of the embedded developing teeth would act to reduce stresses in the surrounding bony elements of the juvenile face. We tested this hypothesis by simulating occlusal loading in a finite element (FE) model of a child's cranium with a complete set of deciduous teeth and the first permanent molars embedded in the bony crypt in the maxilla. We modeled bone and enamel with appropriate material properties and assessed the effect of embedding high-stiffness enamel structures on stress distribution in the juvenile face. Against expectation, the presence of unerupted enamel caps does not affect the magnitude or location of stresses in the juvenile face. Our results do not support the hypothesis that the unerupted secondary teeth act to moderate stresses in the juvenile face.     

Coordinating growth and maturation - insights from Drosophila

Adult body size in higher animals is dependent on the amount of growth that occurs during the juvenile stage. The duration of juvenile development, therefore, must be flexible and responsive to environmental conditions. When immature animals experience environmental stresses such as malnutrition or disease, maturation can be delayed until conditions improve and normal growth can resume. In contrast, when animals are raised under ideal conditions that promote rapid growth, internal checkpoints ensure that maturation does not occur until juvenile development is complete. Although the mechanisms that regulate growth and gate the onset of maturation have been investigated for decades, the emerging links between childhood obesity, early onset puberty, and adult metabolic disease have placed a new emphasis on this field. Remarkably, genetic studies in the fruit fly Drosophila melanogaster have shown that the central regulatory pathways that control growth and the timing of sexual maturation are conserved through evolution, and suggest that this aspect of animal life history is regulated by a common genetic architecture. This review focuses on these conserved mechanisms and highlights recent studies that explore how Drosophila coordinates developmental growth with environmental conditions.     

Contributions of juvenile and adult diet to the lifetime reproductive success and lifespan of a spider

Food availability can vary widely for animals in nature and can have large effects on growth, reproduction and survival. While the consequences of food limitation for animals have been extensively studied, significant questions still remain including how ontogenetic variation in food availability contributes to lifetime reproductive success. We tested the effects of juvenile and adult food limitation on the lifetime reproductive success and lifespan of bridge spiders, Larinioides sclopetarius. Food availability was manipulated (low or high) over the entire juvenile and adult stage in a full‐factorial design and reproductive output and lifespan were measured. Juvenile and adult food limitation both reduced lifetime egg and hatchling production with effect sizes that were not significantly different from each other. Unlike some other arthropods, where juvenile food limitation reduces fecundity by reducing adult body size, body size was not affected by juvenile diet in bridge spiders. Clutch size was also significantly reduced by both juvenile and adult food limitation. The effect of adult diet on clutch size was stronger than that of juvenile diet. Juvenile and adult food limitation both extended total lifespan, and adult food limitation extended adult longevity (i.e. time from maturation to death). However, juvenile food limitation decreased adult longevity, in contrast to what would be predicted by dietary or caloric restriction. Compensatory feeding and growth are widely recognized mechanisms through which animals can ameliorate some of the negative effects of periods of food limitation. Yet our results combined with studies of a range of other species suggest that there may be lasting consequences of juvenile food limitation on lifetime reproductive success that cannot be compensated for by adult feeding in some species.     

Otolith microstructure pattern in Oreochromis niloticus (L.)

The microstructure of otoliths from young and old Oreochromis niloticus (L.) were studied. Otoliths were prepared histologically except for those from newly hatched fish. Hatching results in the formation of a check in the otoliths, which appeared 1 day later. Other checks are rare in juvenile otoliths but common in adult otoliths. Faint and non-daily increments were observed within the hatching check. After hatching, increments were deposited daily. Sub-daily increments were faint and narrow, they were present in the area along the dorso-ventral axis of the otolith but did not continue into the lateral region. Discontinuous zones in the medial area appeared different from those in the lateral area. New growth centres were not only found in the juvenile fish otoliths, but also in adult fish otoliths.     

Life histories have a history: effects of past and present conditions on adult somatic growth rates in wild Trinidadian guppies

1.?Environmental conditions in the present, more recent past and during the juvenile stage can have significant effects on adult performance and population dynamics, but their relative importance and potential interactions remain unexplored. 2.?We examined the influence of food availability at the time of sampling, 2?months prior and during the juvenile stage on adult somatic growth rates in wild Trinidadian guppies (Poecilia reticulata). 3.?We found that food availability during both the early and later parts of an individual's ontogeny had important consequences for adult growth strategies, but the direction of these effects differed among life stages and their magnitude, in some cases, depended on food levels experienced during other life stages. Current food levels and those 2?months prior to growth measurements had positive effects on adult growth rate; though, food levels 2?months prior had a greater effect on growth than current food levels. In contrast, the effects of food availability during the juvenile stage were higher in magnitude but opposite in direction to current food levels and those 2?months prior to growth rate measurements. Individuals recruiting under low food levels grew faster as adults than individuals recruiting during periods of high food availability. There was also a positive interaction between food levels experienced during the juvenile stage and 2?months prior such that the effects of juvenile food level diminished as the food level experienced 2?months prior increased. 4.?These results suggest that the similar conditions occurring at different life stages can have different effects on short- and long-term growth strategies of individuals within a population. They also demonstrate that, while juvenile conditions can have lasting effects on adult performance, the strength of that effect can be dampened by environmental conditions experienced as an adult. 5.?A simultaneous consideration of past events in both the adult and juvenile stage may therefore improve predictions for individual- and population-level responses to environmental change.     

Age and growth of Benthosema pterotum (Alcock, 1890) (Myctophidae) in the Oman Sea

Juvenile and adult specimens of Benthosema pterotum (skinnycheek lanternfish) were collected during several surveys conducted on the Iranian continental shelf of the Oman Sea. Age was estimated by enumeration of growth increments in sagittae otolith sections on the assumption of their daily deposition. Three distinct growth zones in otolith microstructure (central, middle, and external) were defined. These three zones presumably represent increments deposited during successive life history stages, characterized by a different migratory behavior and depth occurrence. The number of increments in the central zone of the B. pterotum otolith (26.8 on average) was thus far one of the lowest in myctophid species studied. A negative correlation between the number of increments in the central and middle zones was observed. This might suggest a functional relation between these two periods of early life history, when fewer larvae in the epipelagic layers may be compensated by a longer non‐migratory behavior of metamorphosis larvae and early juveniles. The maximum number of growth increments in B. pterotum otoliths, i.e. 315, indicated a short lifespan of probably <1 year. The relationship between otolith length and standard length was described by linear regression model (r² = 0.902), and between the body length and weight as an isometric growth in 274 juvenile and adult specimens (r² = 0.922). The growth model estimated only for juveniles and adults was curvilinear (SL = 1.150 × Age^0.616; r² = 0.868). Back‐calculated hatch dates might suggest a spawning season of B. pterotum from May to September, but due to the limited number of investigated specimens a prolonged spawning cannot be excluded. Further trials are needed to measure the population parameter dynamics.     

Dental health and lifestyle in four early mediaeval juvenile populations: Comparisons between urban and rural individuals, and between coastal and inland settlements

Dental stress markers such as enamel hypoplasia and caries are suitable indicators of population health and lifestyle, although they must be recorded and interpreted carefully. To date, they have been predominantly studied in adult samples, whereas juvenile remains are also affected by these lesions. In this study, dental enamel hypoplasia and caries were both evaluated on 613 non-adult individuals from four early mediaeval Moravian and Frankish skeletal series, who had experienced contrasting environments and lifestyles. The aim of this study is to assess the relationship between these biological traits and living conditions, and how this is manifested in the juvenile dental remains.Significant differences between populations were found in stress markers, dental lesions and the way these were manifested. Exposure to stressful conditions varies between urban and rural populations and is related to age groups. Although the children under investigation seem to have had different diets, it is difficult to distinguish the biological contribution (different enamel susceptibility) from the lifestyle contribution (different food, environment) in the formation of caries. Moreover, such studies must be interpreted carefully due to the possibility of intra- and inter-observer errors and the subjectivity of the scoring techniques.Nevertheless, this study also demonstrates that results of an investigation of juvenile skeletal remains can be as informative as a study of adults and that juvenile skeletons can be included in large bioarchaeological population studies.     

Ontogeny of canine dimorphism in extant hominoids.

Many behavioral and ecological factors influence the degree of expression of canine dimorphism for different reasons. Regardless of its socioecological importance, we know virtually nothing about the processes responsible for the development of canine dimorphism. Our aim here is to describe the developmental process(es) regulating canine dimorphism in extant hominoids, using histological markers of tooth growth. Teeth preserve a permanent record of their ontogeny in the form of short- and long-period incremental markings in both enamel and dentine. We selected 52 histological sections of sexed hominoid canine teeth from a total sample of 115, from which we calculated the time and rate of cuspal enamel formation and the rate at which ameloblasts differentiate along the future enamel-dentine junction (EDJ) to the end of crown formation. Thus, we were able to reconstruct longitudinal growth curves for height attainment in male and female hominoid canines. Male hominoids consistently take longer to form canine crowns than do females (although not significantly so for our sample of Homo). Male orangutans and gorillas occasionally take up to twice as long as females to complete enamel formation. The mean ranges of female canine crown formation times are similar in Pan, Gorilla, and Pongo. Interspecific differences between female Pan canine crown heights and those of Gorilla and Pongo, which are taller, result from differences in rates of growth. Differences in canine crown heights between male Pan and the taller, more dimorphic male Gorilla and Pongo canines result both from differences in total time taken to form enamel and from faster rates of growth in Gorilla and Pongo. Although modern human canines do not emerge as significantly dimorphic in this study, it is well-known that sexual dimorphism in canine crown height exists. Larger samples of sexed modern human canines are therefore needed to identify clearly what underlies this.     

The evolution of adult pollen feeding did not alter postembryonic growth in Heliconius butterflies

For many animals, the availability and provision of dietary resources can vary markedly between juvenile and adult stages, often leading to a temporal separation of nutrient acquisition and use. Juvenile developmental programs are likely limited by the energetic demands of many adult tissues and processes with early developmental origins. Enhanced dietary quality in the adult stage may, therefore, alter selection on life history and growth patterns in juvenile stages. Heliconius are unique among butterflies in actively collecting and digesting pollen grains, which provide an adult source of essential amino acids. The origin of pollen feeding has therefore previously been hypothesized to lift constraints on larval growth rates, allowing Heliconius to spend less time as larvae when they are most vulnerable to predation. By measuring larval and pupal life‐history traits across three pollen‐feeding and three nonpollen‐feeding Heliconiini, we provide the first test of this hypothesis. Although we detect significant interspecific variation in larval and pupal development, we do not find any consistent shift associated with pollen feeding. We discuss how this result may fit with patterns of nitrogen allocation, the benefits of nitrogenous stores, and developmental limitations on growth. Our results provide a framework for studies aiming to link innovations in adult Heliconius to altered selection regimes and developmental programs in early life stages.     

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.     

The Evolution of Life Histories in Holo-anhydrobiotic Animals: A First Approach

The life histories of holo-anhydrobiotic animals differ fromthose of all other organisms by a regular or irregular entranceinto an ametabolic state induced by desiccation. Such ametabolicperiods will arrest growth and reproduction completely and thusaffect primary life history parameters dramatically. The selectiveforces and the genetic and physiological trade-offs acting onanhydrobiotic animals are to a large extent unknown. Assuminglow growth rates and low juvenile to adult survival, generaltheoretical models on life history responses to stress predictthat anhydrobiotic animals will be selected for a high degreeof iteroparity, with low fecundity, large egg size, and lowtotal reproductive investment. A high degree of variabilityin growth and reproduction should create a selective force inthe same direction. Although basic empirical data on life historyparameters are very scarce, available observations seem to beconsistent with this prediction.     

Winter weather affects asp viper Vipera aspis population dynamics through susceptible juveniles

Detailed studies on mammals and birds have shown that the effects of climate variation on population dynamics often depend on population composition, because weather affects different subsets of a population differently. It is presently unknown whether this is also true for ectothermic animals such as reptiles. Here we show such an interaction between weather and demography for an ectothermic vertebrate by examining patterns of survival and reproduction in six populations of a threatened European snake, the asp viper ( Vipera aspis ), over six to 17 years. Survival was lowest among juvenile and highest among adult snakes. The estimated annual probability for females to become gravid ranged from 26% to 60%, and was independent of whether females had reproduced in the year before or not. Variation in juvenile survival was strongly affected by winter temperature, whereas adult survival was unaffected by winter harshness. A matrix population model showed that winter weather affected population dynamics predominantly through variation in juvenile survival, although the sensitivity of the population growth rate to juvenile survival was lower than to adult survival. This study on ectothermic vipers revealed very similar patterns to those found in long-lived endothermic birds and mammals. Our results thus show that climate and life history can interact in similar ways across biologically very different vertebrate species, and suggest that these patterns may be very general.     

Size-selective mortality of Sea of Okhotsk pink salmon in the ocean in the winter and spring

The mortality of Sea of Okhotsk pink salmon in the winter and spring varies significantly from year to year, which complicates forecasts of its arrival in the following year based on data on the downstream migration of fry and surveys in the fall. The size-selective mortality of pink salmon was studied and the possibility of using the size and weight parameters of juveniles for predicting their return was evaluated through measurements of scale increments in juvenile pink salmon that were caught in the southern Sea of Okhotsk in the fall of 2007 and 2008 and in fish of these year classes that came back to spawn. In the 2007 year class, which had a low overwinter survival level in the ocean, the average scale increments for the first year of life were considerably smaller than those in adult fish that returned to the spawning grounds. In the pink salmon of 2008, which had a very high level of overwinter survival, the values of scale increments in juveniles and adults were similar. This confirms the hypothesis of a critical size and a critical period, according to which slowly growing juveniles that do not accumulate enough energy reserves for summer are eliminated in the winter to a greater extent as compared to fast-growing fish. Correlation analysis revealed a significant negative relationship between the size and weight of juvenile pink salmon and their mortality in the ocean. After conducting further and more extensive studies this will allow using the size parameters of juvenile pink salmon as one of predictors of its return for the year following the fall surveys. These results emphasize how important it is to take the size and growth rate of juvenile salmon into account when forecasting their return.     

Lineage-specific gene duplication and loss in human and great ape evolution

Given that gene duplication is a major driving force of evolutionary change and the key mechanism underlying the emergence of new genes and biological processes, this study sought to use a novel genome-wide approach to identify genes that have undergone lineage-specific duplications or contractions among several hominoid lineages. Interspecies cDNA array-based comparative genomic hybridization was used to individually compare copy number variation for 39,711 cDNAs, representing 29,619 human genes, across five hominoid species, including human. We identified 1,005 genes, either as isolated genes or in clusters positionally biased toward rearrangement-prone genomic regions, that produced relative hybridization signals unique to one or more of the hominoid lineages. Measured as a function of the evolutionary age of each lineage, genes showing copy number expansions were most pronounced in human (134) and include a number of genes thought to be involved in the structure and function of the brain. This work represents, to our knowledge, the first genome-wide gene-based survey of gene duplication across hominoid species. The genes identified here likely represent a significant majority of the major gene copy number changes that have occurred over the past 15 million years of human and great ape evolution and are likely to underlie some of the key phenotypic characteristics that distinguish these species.     

Ontogeny and phylogeny of the pelvis in Gorilla, Pongo, Pan, Australopithecus and Homo

To examine the evolutionary differences between hominoid locomotor systems, a number of observations concerning the growth of the pelvis among the great apes as compared to modern and fossil hominids are reported. We are interested in the size and shape of the coxal bones at different developmental stages across species that may elucidate the relationship between ontogeny and phylogeny (i.e., heterochrony) in the hominoid pelvis. Our hypotheses are: (1) do rates of absolute growth differ?, (2) do rates of relative growth differ?, and (3) does heterochrony explain these differences? Bivariate and multivariate analyses of pelvic dimensions demonstrate both the diversity of species-specific ontogenetic patterns among hominoids, and an unequivocal separation of hominids and the great apes. Heterochrony alone fails to account for the ontogenetic differences between hominids and the great apes. Compared to recent Homo,Australopithecus can be described as 'hyper-human' from the relative size of the ischium, and short but broad ilium. Australopithecus afarensis differs from Australopithecus africanus by its relatively long pubis. In multivariate analyses of ilium shape, the most complete coxal bone attributed to Homo erectus, KNM-ER 3228, falls within the range of juvenile and adult Australopithecus, whereas Broken Hill falls within the range of modern Homo, suggesting that the modern human ilium shape arose rather recently. Among the great apes, patterns of pelvic ontogeny do not exclusively separate the African apes from Pongo.     

On thick and thin enamel in hominoids

Martin (1983, 1985) reviewed the significance of enamel thickness in hominoid evolution. He studied cut faces of hominoid teeth using the scanning electron microscope and related enamel prism packing patterns to both enamel formation rates and enamel thickness, although he did not present primary data on formation rates, which he summarised as being either “fast” or “slow.” Martin concluded that thick enamel formed at a fast rate represented the ancestral condition in the human and great ape clade. Thin enamel in African apes reflected a secondary reduction in secretion rates, with outer enamel being formed at a slow rate. The present study on ground sections of great ape and human teeth, using polarised light microscopy, was designed to measure the spacing between incremental growth lines in enamel, including striae of Retzius and prism cross striations, to determine rates of enamel formation in hominoids. Measurements on stria spacing showed that striae generally diverged as they passed outwards through enamel in all taxa. Cross-striation spacings also increased from inner to outer enamel. Secretion rates did not fall into two exclusive categories but varied, giving a spectrum of values generally increasing from within outwards at any one crown level and reducing in cervical enamel. There was no evidence for a reduction in enamel formation rates in outer enamel among African apes. These findings cast doubt on the proposition that the common ancestor of great apes and man had thick enamel formed at a fast rate. It is possible that thin enamel was the primitive condition, in which case thick enamel in humans and in Sivapithecus is derived, suggesting that thick enamel on low cusped teeth evolved on more than one occasion.