Muscular function and skull growth in the laboratory rat (Rattus norvegicus)

Bilateral masseterectomy in newly-born rats results in a diminution in cranial weight and size. Such reduction affects the facial skeleton to a greater overall extent than the braincase, although in each region it is the dimensions of length that are affected to a greater extent that are those of height or width. Such contrasts are not dependent upon changes in body weight.
Removal of both temporal muscles—a smaller component of the masticatory musculature—results in little change in cranial proportions other than an increase in width of the braincase.
Such findings can be related, first, to contrasts in the timing of growth in the braincase and facial skeleton and, second, to the extent to which muscular function is reduced.     

中国原大羚化石的再研究

在研究新发现化石材料及对已有标本重新观察的基础上,讨论中国晚中新世至早上新世原大羚(?Protoryx)的系统分类位置。依据头骨、角心和牙齿等方面所具有的独特性状,建议把发现于中国北方曾归入?Protoryx的种类从旧大陆同时代或时代稍早的Protoryx Major, 1891中分出,另立2个新属:以?Protoryx yushensis Teilhard&Trassaert,1938为属型种的华北羚(Huabeitragus gen．nov．)和以?Protoryx shansiensis Bohlin,1935为属型种的粗壮羚(Macro- tragus gen．nov．)。华北羚体型中等大小,头骨窄长,颅轴与面轴夹角近直角,弯曲发生在角前的额部,眶上孔小,分开远,脑颅部短窄,间顶骨长方形,基枕骨方形,没有中纵沟或中嵴,角心长而纤细,基部靠近,向上分散度大,基部横断面近三角形,具前棱。Macrotragus头骨大而粗壮,其弯曲发生在角心基部之间的额面,弯曲度大于直角;角基之间的额中缝隆起呈脊;脸部宽;眶前窝浅;眶下孔后缘位于P3之上。眼眶向头骨两侧突出不明显,其前缘位于M3后缘之后;脑颅部短宽,呈简状,背面稍凸;枕面高、呈半圆形;基枕部为长方形或梯形,具中纵沟和弱的中纵脊;卵圆孔大,面向侧方;角心大而粗壮,内外侧扁,无棱,基部紧靠,向上的分散度小,基部横切面为椭圆形。无角后窝。前臼齿列退化。这两个属之间的主要不同在于前者的个体较小;角心较细弱,向上分散度大(前面观),具前棱,基部横切面呈次三角形;头骨窄,其弯曲位置靠前,弯曲接近90°;脑颅部窄;枕面平,为低矮的长方形;基枕部呈方形,面平,无中纵沟和中纵脊,前、后节结不发育和卵圆孔相对小等。在系统位置上,它们可能属于山羊亚科(Caprinae)。     

Brief communication: Endocranial volumes in an ontogenetic sample of chimpanzees from the Taï Forest National Park, Ivory Coast

Ontogenetic samples of endocranial volumes (EVs) from great apes and humans are critical for understanding the evolution of the brain growth pattern in the hominin lineage. However, high quality ontogenetic data are scarce, especially for nonhuman primates. Here, we provide original data derived from an osteological collection of a wild population of Pan troglodytes verus from the Taï Forest National Park, Ivory Coast. This sample is unique, because age, sex, and pedigree information are available for many specimens from behavioral observations in the wild. We scanned crania of all 30 immature specimens and 13 adult individuals using high-resolution computed tomography. We then created virtual casts of the bony braincase (endocasts) to measure EVs. We also measured cranial length, width, and height and attempted to relate cranial distances to EV via regression analysis. Our data are consistent with previous studies. The only neonate in the sample has an EV of 127 cm³ or 34% of the adult mean. EV increases rapidly during early ontogeny. The average adult EV in this sample is 378.7 ± 30.1 cm³. We found sexual dimorphism in adults; males seem to be already larger than females before adult EV is attained. Regressions on cranial width and multiple regression provide better estimates for EV than regressions on cranial length or height. Increasing the sample size and compiling more high quality ontogenetic data of EV will help to reconcile ongoing discussions about the evolution of hominin brain growth. Am J Phys Anthropol 147:319–325, 2012. © 2011 Wiley Periodicals, Inc.     

Ontogeny and diachronic changes in sexual dimorphism in the craniofacial skeleton of rhesus macaques from Cayo Santiago, Puerto Rico

Insight into the ontogeny of sexual dimorphism is important to our understanding of life history, ecology, and evolution in primates. This study applied a three-dimensional method, Euclidean Distance Matrix Analysis, to investigate sexual dimorphism and its diachronic changes in rhesus macaque (Macaca mulatta) skulls. Twenty-one landmarks in four functional areas of the craniofacial skeleton were digitized from macaques of known age and sex from the Cayo Santiago collections. Then, a series of mean form matrices, form difference matrices, and growth matrices were computed to demonstrate growth curves, rates and duration of growth, and sexual dimorphism within the neurocranium, basicranium, palate, and face. The inclusion of fully adult animals revealed a full profile of sexual dimorphism. Additionally, we demonstrate for the first time diachronic change in adult sexual dimorphism caused by extended growth in adult females. A quicker growth rate in males from ages 2 to 8 was offset by a longer duration of growth in adult females that resulted in diminished dimorphism between the ages of 8 and 15. Four functional areas showed different sex-specific growth patterns, and the rate and duration of growth in the anterior facial skeleton contributed most to the changing profiles of sexual dimorphism. The late maturation in size of the female facial skeleton corresponds to later and less complete fusion of facial sutures. The prolongation of growth in females is hypothesized to be an evolutionary response to high levels of intrasexual competition, as is found in other primate species such as common chimpanzees with similar colony structure and reproductive behavior. Further investigation is required to determine (1) if this phenomenon observed in craniofacial skeletons is linked to sexual dimorphism in body size, and (2) whether this diachronic change in sexual dimorphism is species specific. The changing profile of sexual dimorphism in adult rhesus macaques suggests caution in studying sexual dimorphism in fossil primate and human forms.     

Disproportionate sexual dimorphism in the human face

Analysis of facial dimensions of 86 young adults and their 76 parents indicates that a disproportionate sexual dimorphism exists in the ramus of the mandible, demonstrating a regional difference in growth response. The male ramus is on the average 14% longer than the female ramus, whereas other facial dimensions approximate an 8% sex difference. The findings have relevance to the analysis of skeletal remains and suggest the desirability of age specific discriminant function analysis for the sexing of adult mandibles.     

Roentgen cephalometric studies on skull development in rats. II. Normal and hypophysectomized males; sex differences

Growth of the skull in normal male rats of the Long-Evans strain has been studied by serial roentgen cephalometry (under anesthesia) from 27 to 280 days of age, and has been compared with the retardation following hypophysectomy and with growth data obtained on females in an earlier similar study. The dimensions measured represented major skull regions and segments, and in some instances allowed calculation of indices (e.g., neurocranial width/length ratio) which would show changing proportions. In general, the skull showed the more rapid and prolonged growth characteristic of male rats' skeletal development when compared with that of females. Viscerocranial (“facial”) length and width were notable in this respect. On the other hand, growth in neurocranial width and height ceased at a time and size much like that in females. Examination of the possibility that the adult female skull might correspond closely to that of an immature male disclosed that though this might be true for the neurocranium, facial dimensions showed distinctive sex differences. After hypophysectomy at 27 days of age males (as also females) showed marked reduction of growth but not complete cessation. Dimensional increases were between one-fifth and one-fourth of the normal gains. The ratios computed showed that the proportions of infancy were retained after early hypophysectomy.     

Postnatal growth of skull linear measurements of Cape Hare Lepus capensis in northern China: an analysis in an adaptive context

304 skulls of Cape hare (Lepus capensis) were collected from two climatically distinct localities in northern China. With eye lens weight as a continuous age variable, postnatal growth patterns of 25 cranial linear measurements in relation to sex, growth season and region were analysed to understand the morphological basis of life history adaptation. In almost all the skull measurements, no significant differences were found between either sex or growth seasons. Principal component analysis revealed that facial elements accounted for the greatest proportion of skull morphological variation. Von Bertalanffy function was applied to describe growth trajectories of the skull elements. Based on this model, the growth rates of skull elements and the age at which they reached a certain proportion (95%) of asymptotes were compared. The results showed that skull growth exhibited an allometric pattern, with neural components attaining their final size more rapidly (at about 2–3 months old in tympanic bulla and 4–6 months old in others) than did the facial, which continued to grow well into postnatal life (at 6–10 months old). The earlier establishment of neurocranial morphology was associated with a fully developed central nervous system, which may play a key role in improving the survival of animals during the early phase of life. There was a regional difference in developmental rate of the hare skull. For all the skull parameters, northern hares had a more rapid rate of cranial growth compared to the southern, i.e. skull elements of juveniles from northern population were relatively larger at comparable ages and achieved adult size 0.5–4.0 months earlier than those from the south. In adult hares, however, no significant regional differences in any of the skull parameters were present. Adaptive explanations for the regional difference in ontogenetic pattern of skull morphology include age‐specific thermoregulation constraint, season‐related food availability and age‐dependent predation pressure. Based on the findings of this study, it is suggested that the postnatal growing period represents a crucial time of life, and that improvement of survivorship when young by growth adaptation forms an important aspect of the hare's life history strategies. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78 , 343–353.     

Developmental changes in the facial morphology of the Borneo orangutan (Pongo pygmaeus): possible signals in visual communication

Orangutans display remarkable developmental changes and sexual differences in facial morphology, such as the flanges or cheek-pads that develop only on the face of dominant adult males. These changes suggest that facial morphology is an important factor in visual communication. However, developmental changes in facial morphology have not been examined in detail. We studied developmental changes in the facial morphology of the Borneo orangutan (Pongo pygmaeus) by observing 79 individuals of various ages living in the Sepilok Orangutan Rehabilitation Centre (SORC) in Malaysia and in Japanese zoos. We also analyzed photographs of one captive male that were taken over a period of more than 16 years. There were clear morphological changes that occurred with growth, and we identified previously unreported sexual and developmental differences in facial morphology. Light-colored skin around the eyes and mouth is most prominent in animals younger than 3 years, and rapidly decreases in area through the age of approximately 7 years. At the same time, the scattered, erect hairs on the head (infant hair) become thick, dense hairs lying on the head (adult hair) in both sexes. The results suggest that these features are infant signals, and that adult signals may include darkened face color, adult hair, whiskers, and a beard, which begin to develop after the age of approximately 7 years in both sexes. In females, the eyelids remain white even after 10 years, and turn black at around the age of 20; in males, the eyelids turn black before the age of 10. The whiskers and beards of adults are thicker in males than in females, and are fully developed before the age of 10 in males, while they begin to develop in females only after approximately 20 years. White eyelids and undeveloped whiskers and beards may be visual signals that are indicative of young adult females. Our results also show that the facial morphology of the unflanged male is similar to that of the adult female, although it has also been pointed out that unflanged males resemble younger individuals.     

Sexual dimorphism of cranial suture complexity in wild sheep (Ovis orientalis)

Sutural complexity (the degree of interdigitation) of 13 cranial sutures was compared between male and female wild sheep ( Ovis orientalis ) to investigate a morphological feature that is potentially important with respect to stress transmission in the skulls of males during fighting. Most facial sutures (four of six) were not sexually dimorphic, but two sutures, the maxillojugal and jugolacrimal, had greater complexity in males than in females, suggesting that significant forces may be transmitted through the facial region of rams, most likely during horn clashing. Most of the braincase sutures (five of seven) were more complex in males than in females, and different factors appear to underlie this sexual dimorphism. In females, increased complexity of sutures during ontogeny was predicted best by variables measuring growth of the skull, brain or face, while in males, changes in complexity were predicted best by variables representing mechanical loading and frontal bone growth.     

Changes in size of Baltic field voles over the last 50 years: are they really shrinking?

Using museum materials and recently trapped specimens of field voles (Microtus agrestis (Linnaeus, 1761)) from Lithuania and Estonia, we assessed temporal and latitudinal trends in body and skull size, comparing the periods 1980–1996 and 2014–2016. We measured four body and 23 skull characters, size-adjusting them using the geometric mean procedure. A pronounced decrease in the size of M. agrestis was noted in Estonia, where 23 out of 27 adjusted body and skull characters had decreased by up to 21.9%, with only the tail length, hind foot length, maximum height of mandibula excluding coronoid process and coronoid height of mandibula increasing significantly. Decreases were less marked in voles from Lithuania – most pronounced were a 6.1% decrease in adjusted body length, an 11.6% decrease in adjusted length of the braincase, a 3.85% decrease in the breadth of the braincase, measured at the widest part, a 2.9% decrease in condylobasal skull length and a 2.2% decrease in the height of the braincase. The coronoid height of the mandibula of Lithuanian individuals showed an 8.4% size increase. In both countries, the confounding effect of sex on the size changes of M. agrestis from 1980 to 2016 was much smaller than the effect of time period. Concluding, voles in Estonia became significantly smaller, while changes in the measured characters in Lithuania were heterogeneous.     

Sexual Dimorphism and Facial Growth Beyond Dental Maturity in Great Apes and Gibbons

The great apes and gibbons are characterized by extensive variation in degree of body size and cranial dimorphism, but although some studies have investigated how sexual dimorphism in body mass is attained in these species, for the majority of taxa concerned, no corresponding work has explored the full extent of how sexual dimorphism is attained in the facial skeleton. In addition, most studies of sexual dimorphism combine dentally mature individuals into a single “adult” category, thereby assuming that no substantial changes in size or dimorphism take place after dental maturity. We investigated degree and pattern of male and female facial growth in Pan troglodytes troglodytes, Pan paniscus, Gorilla gorilla gorilla, Pongo pygmaeus, and Hylobates lar after dental maturity through cross-sectional analyses of linear measurements and geometric mean values of the facial skeleton and age-ranking of individuals based on molar occlusal wear. Results show that overall facial size continues to increase after dental maturity is reached in males and females of Gorilla gorilla gorilla and Pongo pygmaeus, as well as in the females of Hylobates lar. In male Pongo pygmaeus, adult growth patterns imply the presence of a secondary growth spurt in craniofacial dimensions. There is suggestive evidence of growth beyond dental maturity in the females of Pan troglodytes troglodytes and Pan paniscus, but not in the males of those species. The results show the presence of statistically significant facial size dimorphism in young adults of Pan paniscus and Hylobates lar, and of near statistical significance in Pan troglodytes troglodytes, but not in older adults of those species; adults of Gorilla gorilla gorilla and Pongo pygmaeus are sexually dimorphic at all ages after dental maturity. The presence of sex-specific growth patterns in these hominoid taxa indicates a complex relationship between socioecological selective pressures and growth of the facial skeleton.     

Soft-tissue facial morphometry from 6 years to adulthood: a three-dimensional growth study using a new modeling

A recently introduced three-dimensional computerized system with landmark representation of the soft-tissue facial surface allows noninvasive and fast quantitative study of facial growth. The aims of the present investigation were (1) to quantify growth changes in soft-tissue facial morphology, (2) to evaluate sex differences in growth patterns, and (3) to provide reference data for selected angular and linear measurements that could be of interest for the objective analysis of maxillofacial surgery or orthodontic patients. The three-dimensional coordinates of 22 standardized facial landmarks were automatically collected by automated infrared photogrammetry using the three-dimensional facial morphometry method in a mixed longitudinal and cross-sectional study, in which 2023 examinations were obtained in 1348 healthy nonpatient subjects between 6 years of age and young adulthood. Selected parameters (angles, linear distances, and ratios) were calculated and averaged for age and sex. Male values were compared with female values by means of Student's t test. Within each age group, linear distances were significantly larger in boys than in girls (p < 0.05) with some exceptions coinciding with the earlier female growth spurt, whereas angular measurements did not show a corresponding sexual dimorphism. Linear distances in girls had almost reached adult dimensions in the 12-to-13-year-old age group, whereas in boys a large increase was still to occur. This was most evident in the middle third of the face, where both sexes showed almost the same dimension and amount of growth up to the age of 13, with significant differences afterward, boys being larger than girls. On the contrary, in the lower third of the face, significant differences occurred throughout the whole investigated period, boys being always larger than girls. The male versus female angular comparison reflected the differential timing in attainment of adult proportions. The three-dimensional facial morphometry method allowed the noninvasive evaluation of a large sample of nonpatient subjects, leading to the definition of three-dimensional normative data about facial soft tissues. The method could supplement more invasive radiographic evaluations, allowing frequent examinations of children and adolescents before and during treatment, as well as in the follow-up.     

A comparative study of adult facial morphology and its ontogeny in the fossil macaque Macaca majori from Capo Figari, Sardinia, Italy

This study examines the morphology of the face in the fossil macaque Macaca majori from Capo Figari (north-eastern Sardinia, Italy) in a comparative ontogenetic context. Thus, a fairly complete face from an adult representative of this fossil species is compared with 3 extant macaque species: Macaca sylvanus (of which species it is questioned whether it is a subspecies, M. sylvanus majori), Macaca mulatta and Macaca fascicularis. Additional incomplete subadult and adult specimens are also examined in order to compare their facial ontogeny with that of the same living species. The comparisons are based on facial landmark data and are undertaken using geometric morphometric methods. These studies indicate that the adult facial morphology and ontogeny of face size and shape in M. majori share much in common with extant macaque species. However, the adult M. majori face displays some unique morphological features, in particular with regard to lateral flaring and relative size of the zygomatic roots. From the study of a limited sample of fossils there is an indication that this flaring arises during postnatal growth, and in consequence the ontogeny of the face of this fossil species may be different from that of M. sylvanus and the other macaque species included in this analysis. From these studies, we conclude that M. majori shows differences in adult facial morphology and possibly in ontogeny from M. sylvanus compatible with a specific rather than subspecific distinction.     

似哺乳类爬行动物和哺乳类动物脑颅侧壁构造类型的演替

本文对从原始似哺乳爬行动物进化到高级兽类哺乳动物过程中脑颅侧壁所发生的形态演化进行了分析,根据构造上的不同,将这些动物的脑颅侧壁的构造方式划分为四个构造类型,代表四个进化阶段。     

Growth of nestling Hamerkops Scopus umbretta in central Mali

Data on growth of nestling Hamerkops Scopus umbretto were analysed with respect to year and season of hatching and to hatch sequence. Quantitative growth characters included weight, culmen, tarsus, tail, third outermost primary quill, standard wing-length and wing span. The growth constant K of weight was 0.179 and time t₁₀-t₉₀ was 21.8 days. Least-squares analyses showed differences in weight gain, culmenlength and tarsus-length related to year and month but not to sequence of hatching. Predictive equations for character against age are provided for all linear measurements. Characters which attained apparent asymptotes before fledging were weight and culmen and tarsus-length whilst other characters varied between 80 and 90% of reported adult size. Qualitative indications of growth discussed are crest development, colouration and the disappearance of the egg-tooth. The rainy and post-rains seasons appear to offer the Hamerkop the best chances of successful breeding and maximum growth rates.     

An experimental study of craniofacial growth in a heterotopic rat head transplant

In order to examine whether or not facial bones are themselves able to regulate their own growth, we devised a new experimental model in which we transplanted the whole head of an infant rat to the body of an isohistogeneic adult rat by means of microvascular anastomoses. The advantage of this model is that the transplanted head has neither scars nor any moving soft tissues that could modify growth around facial bones. Using this model, we conducted a study of the nasomaxillary region that has led us to conclude that facial bones do in fact regulate their own growth. The results also suggested that facial bone sutures play a more active role in the growth process than presently suspected.     

Correlations between the cross-sectional area of the jaw muscles and craniofacial size and shape

In adult human subjects, the correlations were determined between the cross-sectional areas of the jaw muscles (measured in CT scans) and a number of facial angles and dimensions (measured from lateral radiographs). Multivariate statistical analysis of the skeletal variables in a group of 50 subjects led to the recognition of six independent factors determining facial shape, i.e., cranial base length, lower facial height, cranial base flexure and prognathism, facial width, mandibular length, and upper facial height. In 29 of these subjects, the cross-sectional areas of the jaw muscles were determined, and correlations between these areas and the scores on the above-mentioned factors were calculated. It appeared that the cross-sectional areas of temporalis and masseter muscles correlated positively with facial width, whereas the areas of masseter and both pterygoid muscles did so with mandibular length. It has been shown experimentally that a decrease in jaw muscle size in various animals likewise has an effect on facial width and mandibular length. Our results therefore support the hypothesis that in man too the jaw muscles affect facial growth and partly determine the final facial dimensions. They also hint that the role of each muscle is different.     

Early development of the cephalic skeleton of Barbus barbus (Teleostei, Cyprinidae)

The inception, and development of the cephalic skeleton of Barbus barbus from hatching to 24 days passes through periods of fast and slow growth; these rates are not the same in different parts of the skull. Trabeculae, parachordal plates, Meckelian cartilages and hyposymplectics are present at hatching. Then the cartilaginous floor of the neurocranium develops, the pars quadrata, the hyoid bars and branchial arches elements appear shortly before the first movable dermal bones, the dentaries, maxillae and opercles. The first bone of the braincase to appear is the parasphenoid; other bones develop subsequently and at the same time: the angular, quadrate, interopercle and fifth ceratobranchial. Later the splanchnocranium continues to develop at a relatively fast rate while the neurocranium shows little growth. The braincase does not begin to close before the 24th day, nor do the first bones of the skull roof appear, while the bucco-pharyngeal apparatus is complete, having the adult shape. The early constitution of the latter structures seems to be linked with the mechanical demands of biological functions such as breathing and feeding.     

Gender-specific growth patterns of transversal body dimensions in Croatian children and youth (2 to 18 years of age)

In a cross-sectional study of growth, 5,260 healthy children of both sexes from Zagreb (Croatia) aged 2 to 18 years were measured. Six transversal body dimensions were studied: biacromial, transverse chest, antero-posterior chest, biiliocristal, bicondylar humerus and bicondylar femur diamters. A significant increase in body diameters has been observed until the age of 14 to 15 in girls and until the age of 16 in boys, showing that girls have a 1 to 2 years shorter period of growth. Compared to boys of the same age, they achieved larger amounts of final transversal bone size throughout the whole growth period. The most pronounced example was the knee diameter that in girls attained 95% of adult size as early as the age of 10. In both genders, the adult size is achieved earlier in widths of the extremities than in those of the trunk. The studied transversal body segments showed different growth dynamics, which is gender-specific. While sexual dimorphism in pelvic and shoulder diameters emerged with pubertal spurt, gender differences in chest and extremities' diameters started early in life. In all ages, boys had larger chest, elbow and knee diameters. In pubertal age boys gained a significantly larger biacromial diameter (from the age of 13 onwards), while girls exceeded them in biiliocristal diameter (from 10 to 14 years). The findings of gender differences were compared to those reported for other European populations and their growth patters were discussed comparing viewpoints.     

Diphenylpiperazines enhance regeneration after facial nerve injury

Immature rat facial motoneurons are very sensitive to injury with nearly 80% dying during the first week after axotomy. This motoneuron death is apoptotic, similar to that induced in neurons after tropic factor withdrawal. The diphenylpiperazines, flunarizine and cinnarizine, protect dorsal root ganglion neurons from death after withdrawal of trophic support, i.e., nerve growth factor withdrawal, in vitro. Similarly, the monoamine oxidase inhibitor, deprenyl, promotes survival of facial motoneurons after axotomy. These pharmacological agents were assessed both alone and in combination for their ability to prevent death in non-nerve growth factor dependent CNS motoneurons after facial nerve axotomy in newborn rats. Long-term experiments were done with the diphenylpiperazines to evaluate potential enhancement of regeneration. Facial nerve transection resulted in 78% neuronal loss in the injured compared with the contralateral, uninjured nucleus. Systemic administration of diphenylpiperazines for 1 week after facial nerve transection doubled the number of surviving motoneurons from 23% to 47%. Similar results were obtained with deprenyl. Combinations of diphenylpiperazines and deprenyl provide a similar degree of neuronal protection 1 week after injury as that obtained by either agent alone. We assessed the ability of diphenylpiperazines to protect facial motoneurons from death over a prolonged period and enhance subsequent regeneration. Motor neuron counts in rats treated with diphenylpiperazines for 1 month after injury and assessed 2 months later demonstrated long-term enhancement of neuronal protection with an increase of 45% in the number of horseradish peroxidase-labelled motoneurons. The diphenylpiperazines group had ～80% more regenerated myelinated axons in the distal facial nerve than the control group. Thus, diphenylpiperazine treatment during the first month after injury provides long-term protection of non-nerve growth factor dependent CNS motoneurons with subsequent potentiation of long-term facial nerve regeneration.