The causes of porotic hyperostosis and cribra orbitalia: A reappraisal of the iron‐deficiency‐anemia hypothesis

Porosities in the outer table of the cranial vault (porotic hyperostosis) and orbital roof (cribra orbitalia) are among the most frequent pathological lesions seen in ancient human skeletal collections. Since the 1950s, chronic iron‐deficiency anemia has been widely accepted as the probable cause of both conditions. Based on this proposed etiology, bioarchaeologists use the prevalence of these conditions to infer living conditions conducive to dietary iron deficiency, iron malabsorption, and iron loss from both diarrheal disease and intestinal parasites in earlier human populations. This iron‐deficiency‐anemia hypothesis is inconsistent with recent hematological research that shows iron deficiency per se cannot sustain the massive red blood cell production that causes the marrow expansion responsible for these lesions. Several lines of evidence suggest that the accelerated loss and compensatory over‐production of red blood cells seen in hemolytic and megaloblastic anemias is the most likely proximate cause of porotic hyperostosis. Although cranial vault and orbital roof porosities are sometimes conflated under the term porotic hyperostosis, paleopathological and clinical evidence suggests they often have different etiologies. Reconsidering the etiology of these skeletal conditions has important implications for current interpretations of malnutrition and infectious disease in earlier human populations. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Age determination on long bones in a skeletal subadults sample (b-12 years)

The skeletal age on the basis of the diaphyseal length of long bones was assessed. To this aim a sample of subadults skeleton, dated to last century, coming from the cemetery of Bologna was studied. The sample is composed by 79 males and 70 females between 0 and 12 years, whose chronological age and sex are known. Some information can be obtained by the means, standard deviation and graphs of the specimens grouped in age classes. The comparison with other studies confirms the interest of using standards based on direct measurements on long bones of known age and similar to the skeletal populations under study.     

GENES WITH SOCIAL EFFECTS ARE EXPECTED TO HARBOR MORE SEQUENCE VARIATION WITHIN AND BETWEEN SPECIES

The equilibrium sequence diversity of genes within a population and the rate of sequence divergence between populations or species depends on a variety of factors, including expression pattern, mutation rate, nature of selection, random drift, and mating system. Here, we extend population genetic theory developed for maternal-effect genes to predict the equilibrium polymorphism within species and sequence divergence among species for genes with social effects on fitness. We show how the fitness effects of genes, mating system, and genetic system affect predicted gene polymorphism. We find that, because genes with indirect social effects on fitness effectively experience weaker selection, they are expected to harbor higher levels of polymorphism relative to genes with direct fitness effects. The relative increase in polymorphism is proportional to the inverse of the genetic relatedness between individuals expressing the gene and their social partners that experience the fitness effects of the gene. We find a similar pattern of more rapid divergence between populations or species for genes with indirect social effects relative to genes with direct effects. We focus our discussion on the social insects, organisms with diverse indirect genetic effects, mating and genetic systems, and we suggest specific examples for testing our predictions with emerging sociogenomic tools.     

Natural selection on protein polymorphism in the rodent genus Peromyscus: evidence from interlocus contrasts

The effects of natural selection are generally locus-specific, whereas migration, drift, and inbreeding are expected to have relatively uniform effects across the entire genome. This suggests that multilocus surveys of multiple populations can be used to distinguish selection from demographic effects. The purpose of this study was to test for evidence of selection on protein polymorphism in natural populations of mice in the genus Peromyscus. We analyzed published data from geographic surveys of allozyme variation and used a coalescent-based simulation model to identify specific loci that deviated from neutral expectations. Observed F(ST) values generally exhibited a remarkably close fit to the expected neutral distributions, indicating that the majority of loci are simply tracking stochastic demographic processes. A smaller number of loci exhibited highly significant departures from the expectations of the neutral model and thus appear to be tracking the direct or indirect effects of selection. Most departures from neutrality were characterized by F(ST) values that far exceeded neutral expectations and were therefore attributable to spatially varying selection. Interestingly, the albumin locus was implicated as a candidate gene for local adaptation in four different species of Peromyscus. The results also demonstrate that selection can severely bias marker-based estimates of neutral parameters.     

Muscle‐induced loading as an important source of variation in craniofacial skeletal shape

The shape of the craniofacial skeleton is constantly changing through ontogeny and reflects a balance between developmental patterning and mechanical‐load‐induced remodeling. Muscles are a major contributor to producing the mechanical environment that is crucial for “normal” skull development. Here, we use an F₅ hybrid population of Lake Malawi cichlids to characterize the strength and types of associations between craniofacial bones and muscles. We focus on four bones/bone complexes, with different developmental origins, alongside four muscles with distinct functions. We used micro‐computed tomography to extract 3D information on bones and muscles. 3D geometric morphometrics and volumetric measurements were used to characterize bone and muscle shape, respectively. Linear regressions were performed to test for associations between bone shape and muscle volume. We identified three types of associations between muscles and bones: weak, strong direct (i.e., muscles insert directly onto bone), and strong indirect (i.e., bone is influenced by muscles without a direct connection). In addition, we show that although the shape of some bones is relatively robust to muscle‐induced mechanical stimulus, others appear to be highly sensitive to muscular input. Our results imply that the roles for muscular input on skeletal shape extend beyond specific points of origin or insertion and hold significant potential to influence broader patterns of craniofacial geometry. Thus, changes in the loading environment, either as a normal course of ontogeny or if an organism is exposed to a novel environment, may have pronounced effects on skeletal shape via near and far‐ranging effects of muscular loading.     

Age and sex bias in the reconstruction of past population structures

Palaeodemographical studies are founded on the assumption that the sex and age distribution of the skeletal sample reflects the constitution of the original population. It is becoming increasingly clear, however, that the type and amount of information that may be derived from osteoarchaeological collections are related to the state of preservation of remains. This work proposes a new method to evaluate bone preservation, to identify age and sex biases in the preservation of human skeletal remains, and to assess whether differences in preservation patterns are more dependent on factors intrinsic or extrinsic to anatomical features of human bones. Three osteological collections and over 600 skeletons were observed. The state of preservation of human bones was assessed using three preservation indexes: the anatomical preservation index (API), the bone representation index (BRI), and the qualitative bone index (QBI). The results suggest that subadult skeletons are generally more poorly preserved and with bones less well-represented than adult skeletons. Among subadults, female and male skeletons have different patterns of preservation according to their age. This pattern of preservation depends on intrinsic anatomical properties of bones themselves, while external factors can only increase these differences in the state of preservation and representation of osseous remains. It is concluded from this that failure to recognize these differences may lead to misleading interpretations of paleodemography of past human populations.     

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.     

Phenotypic selection on leaf water use efficiency and related ecophysiological traits for natural populations of desert sunflowers

Plant water-use efficiency (WUE) is expected to affect plant fitness and thus be under natural selection in arid habitats. Although many natural population studies have assessed plant WUE, only a few related WUE to fitness. The further determination of whether selection on WUE is direct or indirect through functionally related traits has yielded no consistent results. For natural populations of two desert annual sunflowers, Helianthus anomalus and H. deserticola, we used phenotypic selection analysis with vegetative biomass as the proxy for fitness to test (1) whether there was direct and indirect selection on WUE (carbon isotope ratio) and related traits (leaf N, area, succulence) and (2) whether direct selection was consistent with hypothesized drought/dehydration escape and avoidance strategies. There was direct selection for lower WUE in mesic and dry H. anomalus populations, consistent with dehydration escape, even though it is the longer lived of the two species. For mesic H. anomalus, direct selection favored lower WUE and higher N, suggesting that plants may be “wasting water” to increase N delivery via the transpiration stream. For the shorter lived H. deserticola in the direr habitat, there was indirect selection for lower WUE, inconsistent with drought escape. There was also direct selection for higher leaf N, succulence and leaf size. There was no direct selection for higher WUE consistent with dehydration avoidance in either species. Thus, in these natural populations of two desert dune species higher fitness was associated with some combination direct and indirect selection for lower WUE, higher leaf N and larger leaf size. Our understanding of the adaptive value of plant ecophysiological traits will benefit from further consideration of related traits such as leaf nitrogen and more tests in natural populations.     

SEXUAL SELECTION BY THE HANDICAP MECHANISM

The handicap mechanism of sexual selection by female choice has been strongly criticized because it does not cause sexual selection to reinforce viability selection and it cannot account for the origin of mating preferences. However, several models indicate that the handicap mechanism can have important effects when operating in conjunction with Fisher's mechanism in polygynous populations. These models have been criticized because they require that fitness remains heritable indefinitely. I develop a simple haploid model of the handicap mechanism based on nonheritable variation in paternal investment, thus eliminating the problem of heritable fitness. This model produces the same evolutonary dynamics as both simple and quantitative genetic models of the handicap mechanism based on heritable fitness. If the parameters are such that Fisherian runaway selection does not occur in the null model (i.e., the polymorphic equilibria, which lie along the “Fisher line,” are stable), then the handicap mechanism turns the Fisher line into an evolutionary trajectory upon which all other trajectories converge. This occurs because Fisher's mechanism generates no net selection on female preference when the population is on the Fisher line, so that any additional source of selection (direct or indirect) on female choice causes the population to evolve deterministically along the Fisher line. This change in the evolutionary dynamics has the important consequence of eliminating the potential for rapid population divergence for mating systems via genetic drift along the Fisher line.     

Upper pleistocene fossil hominids from Sri Lanka

Between 1978 and 1983 hominid skeletal remains were collected from the cave sites of Batadomba lena and Beli lena Kitulgala in Sri Lanka (formerly Ceylon). These are the most ancient specimens of anatomically modern Homo sapiens found thus far in South Asia, radiocarbon dates placing them in the Upper Pleistocene. Morphometric analysis of the remains of some 38 individuals from the two sites indicates that these populations were characterized by medium stature, moderate to pronounced cranial and postcranial robusticity, medium-size permanent tooth crown measurements, prognathic alveolar facial proportions, and low incidence of osseous and dental pathological conditions. Comparisons of these ancient Sri Lankans with other prehistoric skeletal series from South Asia and elsewhere support the hypothesis that muscular-skeletal robusticity was a significant physical adaptation of earlier hunting-foraging populations. A trend towards reduction of sexual dimorphism and development of more gracile body form and smaller teeth appears to have accelerated with the socioeconomic transition to food-production strategies involving agriculture and pastoralism and refinement of technologies for food procurement and preparation, as documented by morphometric studies of later prehistoric inhabitants of South Asia.     

Rheumatoid arthritis "in the buff": erosive arthritis in defleshed bones

Examination of isolated bones from patients with unequivocal rheumatoid arthritis provides only a glimpse of the disease but has been the only "gold standard" for recognition of osseous lesions as compatible with the diagnosis of rheumatoid arthritis. Documentation of skeletal pathologic changes in confidently diagnosed individuals has been a major missing link in the transition from clinical to skeletal analysis. Availability of appropriate skeletal material from two patients with long-standing, well-documented rheumatoid arthritis provided the opportunity for acquiring such information. The osseous appearance, skeletal distribution, and distinguishing features of rheumatoid arthritis "in the buff" were delineated in two contemporary patients and in a skeletal population of 2,906 individuals. The preconceived notion of anticipated severity of disease has hereby been tested and found wanting. Severe lesions are not recognizable or distinguishable from artifact in relatively fragile rheumatoid arthritis-affected bones, whether freshly prepared or remotely sampled. Characterization of the nature and epidemiology of osseous alterations in two contemporary skeletal populations permitted the development of a standard for recognition of the disease in skeletal populations.     

GOOD GENES AND DIRECT SELECTION IN THE EVOLUTION OF MATING PREFERENCES

A model is used to study quantitatively the impact of a good genes process and direct natural selection on the evolution of a mating preference. The expression of a male display trait is proportional to genetic quality, which is determined by the number of deleterious mutations a male carries throughout his genome. Genetic variances and covariances, including the covariance between the preference and male trait that drives the good genes process, are allowed to evolve under an infinitesimal model. Results suggest that the good genes process generates only weak indirect selection on preferences, with an effective selection intensity of a few percent or less. If preferences are subject to direct natural selection of the intensity observed for other characters, the good genes process alone is not expected to exaggerate the male trait by more than a few phenotypic standard deviations, contrary to what is observed in highly sexually selected species. Good genes can, however, cause substantial exaggeration if preference genes are nearly selectively neutral. Alternatively, direct selection on preference genes, acting on mating behavior itself or on the genes' pleiotropic effects, can cause mating preferences and male display traits to be exaggerated by any degree. Direct selection of preference genes may therefore play an important role in species that show extreme sexual selection.     

脊椎动物古病理学研究现状及其中国中生代化石记录

中国长江流域沿岸的洞穴和裂隙堆积物中常含有属种丰富的更新世哺乳动物群, 其中小哺乳动物化石由于标本数量多且演化速率快, 因此在第四纪生物地层对比方面具有重要的作用。本文系统研究了产自长江下游南岸的安徽繁昌癞痢山东裂隙的鼠科化石, 包括: Apodemus cf. A. asianicus、Micromys minutus、Chiropodomys cf. C. primitivus、Vandeleuria sp.、Niviventer cf. N. preconfucianus、Leopoldamys sp.、Berylmys cf. B. bowersi 和 Rattus sp., 共计 8 属 8 种, 其中绝灭种和现生种的比例分别为 37.5%和 25%。依据化石属种的形态差异和演化水平, 东裂隙鼠科动物群的性质比安徽繁昌人字洞和重庆巫山龙骨坡都要明显进步, 比湖北建始龙骨洞的鼠科组合也显得稍进步, 兼具早—中更新世的特点, 但更偏向早更新世, 其地质时代很可能为早更新世晚期。此外, 本文还初步建立了长江流域早更新世鼠科动物群的层序, 从早到晚依次为: 安徽繁昌人字洞(距今约 220–256 万年)和重庆巫山龙骨坡(距今约 220–250 万年)—湖北建始龙骨洞(距今约 120–160 万年)—安徽繁昌癞痢山东裂隙(估计距今约 78–120 万年)。     

Paleopathologies of the vertebral column in medieval skeletons

Paleopathological data provide valuable information about health, longevity and mortality in earlier human populations. We investigated the incidence of spinal pathologies on 54 individuals (1045 vertebrae and 18 sacral bones) that belong to a medieval skeletal series discovered in the Dalheim monastery (Northwest Germany) and compared them with contemporary and recent populations. The skeletons were analyzed with anthropological methods (sex and age determination), by macroscopic inspection, and, if pathologies of the spine and the sacrum were visible, also by X-ray. We investigated evidence of trauma, specific and nonspecific infectious diseases, joint diseases, tumors, and congenital as well as metabolic disorders. Radiocarbon determination of four samples of different specimens was also undertaken revealing a historic dating of ca. 1050 AD. The most common pathological findings were degenerative changes of the spine found in 29 individuals (53.3%). Examples of infections of the spine were rare (0.8% of all vertebrae). There were no cases of traumatic injuries of the spine. The prevalence of spondylosis deformans, the most commonly found type of pathology was found to be higher in the lumbar region, in males as well as in individuals of low stature.     

贵州剑河寒武系乌溜阶褶颊虫类三叶虫Eosoptychopariaguizhouensis的蜕壳研究*

1 中国地质大学(北京)地球科学与资源学院, 北京 100083; 2 生物地质与环境地质国家重点实验室, 中国地质大学(北京), 北京 100083 提要古病理学是一门新兴的交叉学科, 涉及病理学、生物学、体质人类学、考古学等领域。近年来, 中生代古脊椎动物的研究案例日益增加, 在化石骨骼上发现了破坏、体积变化、变形、增生和肿瘤, 其提供的信息愈发引起学术界关注。中国的中生代脊椎动物化石记录相当丰富, 已经在恐龙类(包括: 鸟类)、有鳞类、似哺乳爬行类中发现了骨折、划咬伤及感染、椎体融合、关节炎及牙病等疾病记录。具有古病理学特征的标本尚有较大的发现与研究空间。本文介绍了病理学的主要研究方法和国际研究进展, 并以已发现的中国中生代脊椎动物化石的骨骼古病理学实例为基础, 分析讨论这些病变的致病原理、潜在的行为学信息及其病变对动物生存和生态的影响, 展望了该交叉学科的发展方向, 包括: 利用病理研究古生物的形态功能、社会性行为等。     

Treponematosis in regional and chronological perspective from central Gulf Coast Florida

Examination of five central Gulf Coast Florida archaeological skeletal series (n = 547) from the late prehistoric and early historic periods reveals the presence of pathological lesions characteristic of treponemal infection. These skeletal remains of native American populations represent a crucial time regarding hypotheses about the exchange of syphilis between the Old and New World or the mutation of one treponemal infection into another. Comparison of the lesions with those observed in modern studies of treponemal infection does not support an interpretation of veneral syphilis. The data from this study and from other archaeological skeletal series indicate the presence of a treponemal disease prior to European contact. The skeletal elements from Florida do not suggest that the disease changed dramatically following contact with Europeans. It is possible that cases of nonspecific postcranial pathology can be explained by the presence of this disease in the Florida populations. This study indicates that a treponemal disease was endemic throughout the region by at least A. D. 1000. © 1993 Wiley-Liss, Inc.     

Why are some snails visibly polymorphic,and others not?

Work on Cepaea land-snails since 1950 is surveyed, and various explanations for their visible polymorphism, including predator selection, the influence of sunlight and temperature, co-adaptation and linkage disequilibrium, and "area effects", are discussed. All of these can modify the genetic make-up of natural populations in particular circumstances, but none provide a satisfactory answer to the question of why some species are visibly polymorphic whereas others present a uniform external appearance. The likely explanation is that probably all species are genetically heterogeneous at numerous loci, as a result of heterozygote advantage, co-adaptation and other selective factors maintaining the different alleles in equilibrium, which may sometimes have visible effects on the phenotype. If these are positively disadvantageous, selection for epistatic genes will suppress the visible polymorphism, without affecting the underlying genetical heterogeneity, preserved by selection for other non-visible pleiotropic effects of the alleles involved. But this will not happen if the visibly distinct effects of these different allelomorphs are selectively more or less neutral. Many examples of polymorphism, including the so-called 'pseudo'-polymorphism, are therefore essentially non-adaptive so far as their visible manifestation is concerned, being maintained as balanced polymorphism by selection for non-visible pleiotropic effects of the genes involved.     

Multiple paternity in reptiles: patterns and processes

The evolution of female promiscuity poses an intriguing problem as benefits of mating with multiple males often have to arise via indirect, genetic, effects. Studies on birds have documented that multiple paternity is common in natural populations but strong evidence for selection via female benefits is lacking. In an attempt to evaluate the evidence more broadly, we review studies of multiple paternity in natural populations of all major groups of nonavian reptiles. Multiple paternity has been documented in all species investigated so far and commonly exists in over 50% of clutches, with particularly high levels in snakes and lizards. Marine turtles and lizards with prolonged pair-bonding have relatively low levels of multiple paternity but levels are nevertheless higher than in many vertebrates with parental care. There is no evidence that high levels of polyandry are driven by direct benefits to females and the evidence that multiple paternity arises from indirect genetic benefits is weak. Instead, we argue that the most parsimonious explanation for patterns of multiple paternity is that it represents the combined effect of mate-encounter frequency and conflict over mating rates between males and females driven by large male benefits and relatively small female costs, with only weak selection via indirect benefits. A crucial step for researchers is to move from correlative approaches to experimental tests of assumptions and predictions of theory under natural settings, using a combination of molecular techniques and behavioural observations.     

Comparison of child morbidity in two contrasting medieval cemeteries from Denmark

This study compares associations between demographic profiles, long bone lengths, bone mineral content, and frequencies of stress indicators in the preadult populations of two medieval skeletal assemblages from Denmark. One is from a leprosarium, and thus probably represents a disadvantaged group (Naestved). The other comes from a normal, and in comparison rather privileged, medieval community (AEbelholt). Previous studies of the adult population indicated differences between the two skeletal collections with regard to mortality, dental size, and metabolic and specific infectious disease. The two samples were analyzed against the view known as the "osteological paradox" (Wood et al. [1992] Curr. Anthropol. 33:343-370), according to which skeletons displaying pathological modification are likely to represent the healthier individuals of a population, whereas those without lesions would have died without acquiring modifications as a result of a depressed immune response. Results reveal that older age groups among the preadults from Naestved are shorter and have less bone mineral content than their peers from AEbelholt. On average, the Naestved children have a higher prevalence of stress indicators, and in some cases display skeletal signs of leprosy. This is likely a result of the combination of compromised health and social disadvantage, thus supporting a more traditional interpretation. The study provides insights into the health of children from two different biocultural settings of medieval Danish society and illustrates the importance of comparing samples of single age groups.     

Probable destructive meningioma in an archaeological adult male skull from Alaska

Several intracranial pathological conditions can affect the bones of the skull. The most common cause of these conditions is tumor, but infection and other diseases are also known to affect the bones of the skull. Distinguishing between the various causes of intracranial skeletal pathology in archaeological human remains is usually a challenging exercise, and a specific diagnosis will often be impossible. Meningiomas are tumors that arise in arachnoid tissues embedded in the outer layer of the dura. Because of this association, they occur almost exclusively in the skull and vertebral column. Usually meningiomas are slow-growing tumors that do not metastasize to other organs and tissues of the body. However, rare cases can be malignant and, even when meningiomas are benign, their presence and growth can adversely affect the nervous and vascular supply to other tissues in the skull and vertebral column. Their effect on adjacent bone tissue varies from stimulating bone-forming lesions to causing highly destructive lesions. A few examples of meningioma have been described in the paleopathological literature. Most of these cases are bone-stimulating meningiomas. The case presented here is a probable example of a highly destructive meningioma of the skull base, with unilateral extension into the left side of the cranium. This case is compared with a modern clinical case of destructive intracranial meningioma that was documented both radiographically and pathologically. Destructive meningiomas can be confused with other pathological conditions, including benign and malignant tumors. Criteria for differentiating the diagnostic options are reviewed.