Sexual dimorphism in the growth of the cranium

The major sexual dimorphisms in body size appear at puberty but, by then, 95% of the growth of the cranium is completed. As sexual dimorphism in the cranium is as great as for other parts of the body, this suggests that it must appear at an earlier age, and that cranium/body size ratios for the two sexes will vary during growth. Results from a longitudinal study of Montreal children are used to investigate this phenomenon. The effect is expressed quantitatively by proportional growth and growth velocity curves, based on the final size of boys, which show that the dimorphism indeed makes an early appearance. The data are also analyzed on an age scale relative to the ages of peak growth velocity in stature, derived from the individual growth curves. This shows that although there is a minor pubertal spurt in growth for the external cranial dimensions of boys, it contributes relatively little to the final dimorphism in cranial size. To summarize this aspect of growth, an index of cephalization is calculated: head length × head width/stature. Cross-sectional standards for the change of the mean index with age show a linear decline for boys and girls until puberty, with a constant difference between them. After puberty, the index becomes equal in the two sexes. Individual development curves for the index are however not linear.     

Linear versus geometric morphometric approaches for the analysis of head shape dimorphism in lizards

Differences between the sexes may arise because of differences in reproductive strategy, with females investing more in traits related to reproductive output and males investing more in traits related to resource holding capacity and territory defence. Sexual dimorphism is widespread in lizards and in many species males and females also differ in head shape. Males typically have bigger heads than females resulting in intersexual differences in bite force. Whereas most studies documenting differences in head dimensions between sexes use linear dimensions, the use of geometric morphometrics has been advocated as more appropriate to characterize such differences. This method may allow the characterization of local shape differences that may have functional consequences, and provides unbiased indicators of shape. Here, we explore whether the two approaches provide similar results in an analyses of head shape in Tupinambis merianae. The Argentine black and white tegu differs dramatically in body size, head size, and bite force between the sexes. However, whether the intersexual differences in bite force are simply the result of differences in head size or whether more subtle modifications (e.g., in muscle insertion areas) are involved remains currently unknown. Based on the crania and mandibles of 19 lizards with known bite force, we show intersexual differences in the shape of the cranium and mandible using both linear and geometric morphometric approaches. Although both types of analyses showed generally similar results for the mandible, this was not the case for the cranium. Geometric morphometric approaches provided better insights into the underlying functional relationships between the cranium and the jaw musculature, as illustrated by shape differences in muscle insertion areas not detected using linear morphometric data. J. Morphol. 275:1016–1026, 2014. © 2014 Wiley Periodicals, Inc.     

Testing the cranial evolutionary allometric ‘rule’ in Galliformes

Recent comparative studies have indicated the existence of a common cranial evolutionary allometric (CREA) pattern in mammals and birds, in which smaller species have relatively smaller faces and bigger braincases than larger species. In these studies, cranial allometry was tested using a multivariate regression between shape (described using landmarks coordinates) and size (i.e. centroid size), after accounting for phylogenetic relatedness. Alternatively, cranial allometry can be determined by comparing the sizes of two anatomical parts using a bivariate regression analysis. In this analysis, a slope higher or lower than one indicates the existence of positive or negative allometry, respectively. Thus, in those species that support the CREA ‘rule’, positive allometry is expected for the association between face size and braincase size, which would indicate that larger species have disproportionally larger faces. In this study, I applied these two approaches to explore cranial allometry in 83 Galliformes (Aves, Galloanserae), ranging in mean body weight from 30 g to 2.5 kg. The multivariate regression between shape and centroid size revealed the existence of a significant allometric pattern resembling CREA, whereas the second analysis revealed a negative allometry for beak size and braincase size (i.e. contrary to the CREA ‘rule’, larger galliform species have disproportionally shorter beaks than smaller galliform species). This study suggests that the presence of CREA may be overestimated when using cranium size as the standard measurement.     

Identification of basilosaurid archaeocetes (Mammalia,Cetacea) collected in Egypt by Richard Markgraf (1901–1916)

Chicago Field Museum FMNH P-12343 includes a cranium and dentary of a late Eocene archaeocete whale. These were acquired from the commercial fossil dealer Friedrich Kranz of Bonn in 1914–1915, after being collected by Richard Markgraf in Fayum, Egypt. Both were regarded as a single specimen identified in museum records as Prozeuglodon osiris. However, the cranium and dentary differ in preservation and in size, and cannot be parts of the same individual specimen nor belong to the same species. Comparison with other Fayum archaeocetes indicates that the cranium is Stromerius nidensis from the Qasr el-Sagha Formation, and the dentary is Dorudon atrox from the underlying Birket Qarun Formation. Fossils collected by Markgraf are found in museums all over the world, but he was neither a paleontologist nor stratigrapher, and identifications and ages of fossils he sold commercially cannot be accepted uncritically.     

Ontogenetic relationships between cranium and mandible in coyotes and hyenas

Developing animals must resolve the conflicting demands of survival and growth, ensuring that they can function as infants or juveniles while developing toward their adult form. In the case of the mammalian skull, the cranium and mandible must maintain functional integrity to meet the feeding needs of a juvenile even as the relationship between parts must change to meet the demands imposed on adults. We examine growth and development of the cranium and mandible, using a unique ontogenetic series of known‐age coyotes (Canis latrans), analyzing ontogenetic changes in the shapes of each part, and the relationship between them, relative to key life‐history events. Both cranial and mandibular development conform to general mammalian patterns, but each also exhibits temporally and spatially localized maturational transformations, yielding a complex relationship between growth and development of each part as well as complex patterns of synchronous growth and asynchronous development between parts. One major difference between cranium and mandible is that the cranium changes dramatically in both size and shape over ontogeny, whereas the mandible undergoes only modest shape change. Cranium and mandible are synchronous in growth, reaching adult size at the same life‐history stage; growth and development are synchronous for the cranium but not for the mandible. This synchrony of growth between cranium and mandible, and asynchrony of mandibular development, is also characteristic of a highly specialized carnivore, the spotted hyena (Crocuta crocuta), but coyotes have a much less protracted development, being handicapped relative to adults for a much shorter time. Morphological development does not predict life‐history events in these two carnivores, which is contrary to what has been reported for two rodent species. The changes seen in skull shape in successive life‐history stages suggest that adult functional demands cannot be satisfied by the morphology characterizing earlier life‐history stages. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Patterns of growth in the presacral vertebral column of the leopard gecko (Eublepharis macularius)

Postnatal growth patterns within the vertebral column may be informative about body proportions and regionalization. We measured femur length, lengths of all pre‐sacral vertebrae, and lengths of intervertebral spaces, from radiographs of a series of 21 Eublepharis macularius, raised under standard conditions and covering most of the ontogenetic body size range. Vertebrae were grouped into cervical, sternal, and dorsal compartments, and lengths of adjacent pairs of vertebrae were summed before analysis. Femur length was included as an index of body size. Principal component analysis of the variance‐covariance matrix of these data was used to investigate scaling among them. PC1 explained 94.19% of total variance, interpreted as the variance due to body size. PC1 differed significantly from the hypothetical isometric vector, indicating overall allometry. The atlas and axis vertebrae displayed strong negative allometry; the remainder of the vertebral pairs exhibited weak negative allometry, isometry or positive allometry. PC1 explained a markedly smaller amount of variance for the vertebral pairs of the cervical compartment than for the remainder of the vertebral pairs, with the exception of the final pair. The relative standard deviations of the eigenvalues from the PCAs of the three vertebral compartments indicated that the vertebrae of the cervical compartment were less strongly integrated by scaling than were the sternal or dorsal vertebrae, which did not differ greatly between themselves in their strong integration, suggesting that the growth of the cervical vertebrae is constrained by the mechanical requirements of the head. Regionalization of the remainder of the vertebral column is less clearly defined but may be associated with wave form propagation incident upon locomotion, and by locomotory changes occasioned by tail autotomy and regeneration. Femur length exhibits negative allometry relative to individual vertebral pairs and to vertebral column length, suggesting a change in locomotor requirements over the ontogenetic size range.     

Evolution of sexual size dimorphism in birds: test of hypotheses using blue tits in contrasted Mediterranean habitats

Many hypotheses, either sex‐related or environment‐related, have been proposed to explain sexual size dimorphism in birds. Two populations of blue tits provide an interesting case study for testing these hypotheses because they live in contrasting environments in continental France and in Corsica and exhibit different degree of sexual size dimorphism. Contrary to several predictions, the insular population is less dimorphic than the continental one but neither the sexual selection hypothesis nor the niche variation hypothesis explain the observed patterns. In the mainland population it is advantageous for both sexes to be large, and males are larger than females. In Corsica, however, reproductive success was greater for pairs in which the male was relatively small, i.e. pairs in which sexual size dimorphism is reduced. The most likely explanation is that interpopulation differences in sexual size dimorphism are determined not by sex‐related factors, but by differences in sex‐specific reproductive roles and responses to environmental factors. Because of environmental stress on the island as a result of food shortage and high parasite infestations, the share of parents in caring for young favours small size in males so that a reduced sexual size dimorphism is not the target of selection but a by‐product of mechanisms that operate at the level of individual sexes.     

Growth of the skull in the mutant dwarf mouse

By following the development of individual mice from birth to 200 days of age, it was found that the skull of the homozygous mutant (dwdw) was 5% shorter than that of the normal phenotype at birth and some 30% shorter at 200 days of age. During post-natal growth of the skull, the cranium was least altered but the facial skeleton was greatly affected. The differences seen in different parts of the skull are attributed to the peak growth velocities of the various parts occurring at different times.     

Analytic power and sample size calculation for the genotypic transmission/disequilibrium test in case‐parent trio studies

Case‐parent trio studies considering genotype data from children affected by a disease and their parents are frequently used to detect single nucleotide polymorphisms (SNPs) associated with disease. The most popular statistical tests for this study design are transmission/disequilibrium tests (TDTs). Several types of these tests have been developed, for example, procedures based on alleles or genotypes. Therefore, it is of great interest to examine which of these tests have the highest statistical power to detect SNPs associated with disease. Comparisons of the allelic and the genotypic TDT for individual SNPs have so far been conducted based on simulation studies, since the test statistic of the genotypic TDT was determined numerically. Recently, however, it has been shown that this test statistic can be presented in closed form. In this article, we employ this analytic solution to derive equations for calculating the statistical power and the required sample size for different types of the genotypic TDT. The power of this test is then compared with the one of the corresponding score test assuming the same mode of inheritance as well as the allelic TDT based on a multiplicative mode of inheritance, which is equivalent to the score test assuming an additive mode of inheritance. This is, thus, the first time the power of these tests are compared based on equations, yielding instant results and omitting the need for time‐consuming simulation studies. This comparison reveals that these tests have almost the same power, with the score test being slightly more powerful.     

Three-dimensional geometric morphometric analysis of cranio-facial sexual dimorphism in a Central European sample of known sex

This article presents an approach for estimating the sexual dimorphism of adult crania using three-dimensional geometric morphometric methods. The study sample consisted of 139 crania of known sex (73 males and 66 females) belonging to persons who lived during the first half of the 20th century in Bohemia. The three-dimensional co-ordinates of 82 ecto-cranial landmarks and 39 semi-landmarks covering the midsagittal curve of the cranial vault were digitised using a MicroScribe G2X contact digitiser. The purposes of the investigation were to define the regions of the cranium where sexual dimorphism is most pronounced and to investigate the effectiveness of this method for determining sex from the shape of the cranium. The results demonstrate that it is better to analyse apportionable parts of the cranium rather than the cranium as a whole. Significant sexual differences (significance was determined using multivariate analysis of variance) were noted in the shape of the midsagittal curve of the vault, upper face, the region of the nose, orbits, and palate. No differences were recorded either in the shape of the cranium as a whole or in the regions of the base and the neurocranium. The greatest accuracy in determining sex was found in the region of the upper face (100% of study subjects correctly classified) and the midsagittal curve of the vault (99% of study subjects correctly classified).     

云南西瓦古猿头骨的初步研究

本文对1978年12月在云南禄丰石灰坝煤窑发现的第一个古猿头骨化石作了描述和比较,归属为云南西瓦古猿类型。由于它在形态上保留一些原康修尔猿的性质和显示出较多的相似于猩猩的特征,所以作者认为它可能代表后两者的中间环节。同时,作者又发现这个头骨具有接近于南方古猿粗壮种的性质,因此认为云南西瓦古猿与南方古猿粗壮类型可能有亲缘关系。     

The dating of Lantian man and his significance for analyzing trends in human evolution

The Lantian fossil hominid cranium from Southern Shensi Province, China, provides the earliest record of Homo erectus in northern east Asia, and is morphologically the most primitive specimen in the entire world. Importantly, the Kungwangling Lantian cranium (calvarium plus face), with associated stone tools in good geologic and paleontological context, is demonstrably both earlier and more primitive than the Choukoutien I remains. Faunal and palynological evidence support a mid-Mosbachium equivalent age (some 700,000 years). These facts are not recognized in the original Chinese reports. The Chenchiawo Lantian mandible, like the Choukoutien I remains, is attributable to the Holstein-equivalent in China (some 300,000 years ago), and therefore should no longer be temporally associated with the Kungwangling Lantian cranium. However, that the mandible may be morphologically associated with either calls attention to the relative independence of the mandible in human evolution. A comparative study of some modern Mongoloid populations in which very large mandibles may or may not be associated with a scaphoid keel or sagittal elevation depending upon the size and shape of the cranium demonstrates the relative autonomy of the mandible. Continuing selection pressure for a masticatory complex with large jaws provides another point of continuity between East Asian fossil and modern Mongoloid hunting populations such as Eskimos and Aleuts. A number of morphological features of the cranium, especially vault thickness, cranial capacity and reinforcement system, conform to expectation and confirm a general trend of reduction in vault thickness and reinforcement system with increase in cranial capacity over time within the single human species.     

The Lake Ndutu cranium and early Homo sapiens in Africa

The partial cranium from Lake Ndutu, near Olduvai Gorge in northern Tanzania, has generally been viewed as Homo erectus, although points of similarity to H. sapiens have also been recognized. Bones of the vault are in fact quite thick, and the cranium is small. Length and breadth dimensions are comparable to those of earlier H. erectus from Koobi Fora and Ileret, and the Ndutu individual is more similar in size to O.H. 12 than to O.H. 9. Unfortunately, the facial skeleton and frontal bone are very incomplete, and little useful information can be obtained from these parts of the existing reconstruction. The parietals are also damaged, but the left temporal is more satisfactorily preserved, and the occiput is nearly complete. Occipital morphology, mastoid shape, and characteristics of the glenoid cavity and tympanic plate probably provide the best available guide to affinities of the Ndutu hominid. In many of these features the cranium resembles Broken Hill, Elandsfontein, and other African fossils referred to archaic H. sapiens. There are some similarities to modern humans also, but no ties to the Neanderthals of Europe. Allocation of Ndutu to an African subspecies of H. sapiens seems most appropriate, even if the pattern of relationships between such archaic populations and recent humans is still unclear.     

Segmentation of the Vertebrate Head

Historical views of head segmentation are reviewed. The concensusis that the head is segmented essentially in terms of myomeres,and that other organs have responded in varying degrees to this.From the various lines of reasoning a model of the primitivevertebrate is generated. This model denies the tunicate originof the vertebrates—rather it identifies amphioxus as mostlike the ancestral vertebrate. The vertebrate head is made upof a preoral segment plus four other segments. Because of sclerotomites,the head extends through five and a half segments. The nasalorgans and eyes are preoral structures while the ear is locatedbetween segments three and four. The occipital portion of thehead skeleton is formed from the posterior half of the fifthsegment and the anterior half of the sixth; it is vertebra-likein structure. This "segment" is much altered as a result ofthe multiplication of the visceral pouches and is often viewedas the fusion product of several segments. Thus the idea ofcorrespondence between somite and visceral segments posteriorto the second branchial arch is rejected. In some fishes, additionalvertebrae are added to the posterior part of the cranium andthis can be observed in development. The bony cranium of thevertebrate appears to partially reflect segmentation; its componentssuggest a vertebra-like developmental influence in operation.Study of the shark head has contributed much to our knowledgeof this area.     

Evolution of the dragonfly head-arresting system

The arrester or fixation system of the head in adult Odonata is unique among arthropods. This system involves the organs of two body segments: the head and the neck. It consists of a skeleton–muscle apparatus that sets the arrester parts in motion. The parts comprise formations covered with complicated microstructures: fields of microtrichia on the rear surface of the head and post-cervical sclerites of the neck. The arrester immobilizes the head during feeding or when the dragonfly is in tandem flight. Thus, it may serve as an adaptation to save the head from violent mechanical disturbance and to stabilize gaze in a variety of behavioural situations. This study shows the evolutionary trend of the arrester in the order Odonata by using scanning electron microscopy and measurements of arrester structures in 227 species from 26 odonate families. The arrester design occurring in the Epiophlebiidae, Gomphidae, Neopetaliidae, Petaluridae and Chlorogomphinae is suggested to be the basic one. Two convergent pathways of head-arrester evolution among Zygoptera and Anisoptera are proposed. The possible functional significance of the arrester system is discussed.     

柳江人身体大小和形状——体重、身体比例及相对脑量的分析

发现于广西柳江的更新世晚期人类化石除1具完整的头骨外,还包含有右侧髋骨、骶骨、两段股骨及若干件椎骨。根据各方面的特征分析,初步认定这些化石属于同一个体。这一有利条件为我们比较准确地获取与该个体身体大小和形状有关的指标数据提供了可能。本文通过对柳江人头骨及复原骨盆的测量,计算了柳江人的身高、体重、身体比例、相对脑量等。在此基础上分析了柳江人的身体大小和形状。本研究发现:柳江人化石所代表的个体具有适应温暖气候环境的纤细型身体比例,代表相对脑量的EQ指数5.602大于金牛山、山顶洞等中国更新世中、晚期化石人类,而与包括港川人在内的更新世末期及现代人类的EQ指数接近。柳江人体重52.0kg小于金牛山、山顶洞、尼安德特人等生活在高纬度地区的化石人类,而与港川、非洲的KNM-ER3883、KNM-ER3733等生活在温暖环境的古人类接近。作者认为这些发现除说明柳江人生活的气候环境外,还提示柳江人身体大小、比例及相对脑量与更新世末期及现代人类接近。     

高粱对丝黑穗病的抗性及遗传研究

1981—1985年在人工接种的条件下,对1239份高粱品种资源,进行了抗丝黑穗病性鉴定。与此同时,用17个抗性不同的品种系,按照不完全双列杂交设计,进行了高粱对丝黑穗病的抗性遗传研究。结果表明,高粱对丝黑穗病的抗性遗传方式因品种而异。有的品种系具数量性状遗传特点,有的则具有质量性状遗传特点。抗病性属数量性状遗传的品种系,其抗性主要是受加性基因控制。     

The Role of the Genotype in the Development of the Neurophysiological Mechanisms Involved in the Spatial Integration of Neocortex Bioelectric Activity

The contribution of genetic factors into the formation of the neurophysiological mechanisms determining the systemic organization of cortical activity has been estimated in 12 pairs of monozygotic (MZ) and 5 pairs of dizygotic (DZ) twins (18–25 years of age). The similarity of individual patterns of the spatial interaction of cortical bioelectric activity measured for different combinations of EEG statistical relationships from 16 monopolar leads has been estimated in each pair of twins and in each of the 544 pairs of unrelated subjects in both groups. The results of the study suggest a high population invariance and small genetic and phenotypic variations of the morphofunctional systems constituting the main neurophysiological mechanisms of general cerebral integration. Brainstem and subcortical regulatory structures play the leading role in the integration of different brain regions into an organized system. Apparently, the formation of these structures during ontogeny follows the same general pattern in all individuals, because deviation from it is likely to affect the fundamental monomorphic characters of the species. The formation of neocortical interregional connections may be expected to exhibit greater individual variation, with the roles of hereditary and environmental factors in the formation of long and relatively short intercortical interactions being different. Apparently, the individual variation of the long intra- and interhemispheric fiber pathways forming the specific morphological framework of the neocortex is largely determined by the genotype. However, the intercentral interactions mediated by short corticocortical connections that are formed in the course of the vital activity of an individual are likely to be mainly determined by the external and internal environments.     

Variation in the cranium shape of wall lizards (Podarcis spp.): effects of phylogenetic constraints, allometric constraints and ecology

We used geometric morphometrics to explore the influence of phylogenetic and allometric constraints as well as ecology on variation in cranium shape in five species of monophyletic, morphologically similar Podarcis lizards (Podarcis erhardii, Podarcis melisellensis, Podarcis muralis, Podarcis sicula and Podarcis taurica). These species belong to different clades, they differ in their habitat preferences and can be classified into two distinct morphotypes: saxicolous and terrestrial. We found (i) no phylogenetic signal in cranium shape, (ii) diverging allometric slopes among species, and (iii) a significant effect of habitat on cranium shape. The saxicolous species (P. erhardii and P. muralis) had crania with elongated parietals, elongated cranium bases, shortened anterior parts of the dorsal cranium, reduced chambers of the jaw adductor muscles and larger subocular foramina. These cranial features are adaptations that compensate for a flattened cranium, dwelling on vertical surfaces and seeking refuge in crevices. The crania of the terrestrial species (P. melisellensis, P. sicula and P. taurica) tended to be more elongate and robust, with enlarged chambers of the jaw adductor muscle, reduced skull bases and shortened parietals. Terrestrial species exhibited more variation in cranium shape than saxicolous species. Our study suggests that shape variation in Podarcis sp. lizards is largely influenced by ecology, which likely affects species-specific patterns of static allometry.     

Paternally inherited alleles in male body parts of an ant (<Emphasis Type="Italic">Diacamma</Emphasis> sp.) sex mosaic: implication for androgenetic male production in the Hymenoptera

Sex mosaicism, also called gynandromorphism, refers to an accidental phenomenon in dioecious organisms (mainly animals) in which an individual phenotype includes both female and male characteristics. Despite the rarity of this phenomenon, elucidating the mechanisms of naturally occurring sex mosaicism should deepen our understanding of diverse sex determination and differentiation systems in nature. We report the results of a genetic study of a sex mosaic individual of the ant Diacamma sp. from Japan’s Okinawa Island. Parentage analysis using microsatellite markers revealed that the female and male parts of the sex mosaic showed different inheritance patterns: female parts had alleles consistent with their biparental inheritance, whereas most of the male parts had alleles consistent with their paternal inheritance (i.e., androgenesis). We discuss plausible cytogenetic mechanisms that gave rise to the male parts of this individual: polyspermy and the subsequent independent cleavage by a surplus sperm pronucleus, and maternal genome elimination after fertilization of an ovule. Moreover, we hypothesize that the androgenetically produced males found in some Hymenoptera might share the same underlying cytogenetic mechanism with hymenopteran sex mosaicism.