Embryonic pattern of cartilaginous head development in the European toad, Bufo bufo.

The craniofacial skeleton of vertebrates is a major innovation of the whole clade. Its development and composition requires a precisely orchestrated sequence of chondrification events which lead to a fully functional skeleton. Sequential information on the precise timing and sequence of embryonic cartilaginous head development are available for a growing number of vertebrates. This enables a more and more comprehensive comparison of the evolutionary trends within and among different vertebrate clades. This comparison of sequential patterns of cartilage formation enables insights into the evolution of development of the cartilaginous head skeleton. The cartilaginous sequence of head formation of three basal anurans (Xenopus laevis, Bombina orientalis, Discoglossus scovazzi) was investigated so far. This study investigates the sequence and timing of larval cartilaginous development of the head skeleton from the appearance of mesenchymal Anlagen until the premetamorphic larvae in the neobatrachian species Bufo bufo. Clearing and staining, histology, and 3D reconstruction enabled the tracking of 75 cartilaginous structures and the illustration of the sequential changes of the skull as well as the identification of evolutionary trends of sequential cartilage formation in the anuran head. The anuran viscerocranium does not chondrify in the ancestral anterior to posterior direction and the neurocranial elements do not chondrify in posterior to anterior direction. Instead, the viscerocranial and neurocranial development is mosaic-like and differs greatly from the gnathostome sequence. Strict ancestral anterior to posterior developmental sequences can be observed within the branchial basket. Thus, this data is the basis for further comparative developmental studies of anuran skeletal development.

Basicranial Modular Organization. A Study in the Araucanian Horse of Colombia.

The skull is divided into neurocranium and splanchnocranium, and its variation allows ecomorphological studies to learn about possible evolutionary and adaptive characteristics. The basicranial organization of the neurocranium and splanchnocranium modules was studied in a sample of 31 skulls from adult Araucanian horses by means of 2D geometric morphometric techniques. The neurocranium and splanchnocranium modules on the ventral aspect were analyzed separately using a set of 31 landmarks. The RV coefficient (the multivariate analog of a correlation) was estimated to analyze the independence of these two parts, as well as their morphological integration, using a two-block analysis of least squares. The study results confirm the modular development of the neurocranium and the splanchnocranium, the former being more stable than the latter as well as low morphological integration between the two. The development between both parties is structured in a modular way but allows relative independence. Now it would be interesting for future studies to add muscles (those that connect the cranial parts, but also the cervical), the hyoid apparatus, and the ossicles of the internal ear and the jaw and analyze if they behave as integrated modules between them. Since this research has been conducted at the subspecific breed level, it could be plausible that in other breeds, this integrative development was different.

A Geometric Morphometric Study on Sexual Dimorphism in Viscerocranium.

The level of sexual dimorphism manifested by human bones is an important factor for development of effective sex estimation methods. The aim of the study was to investigate the sexual dimorphism in the size and shape of the viscerocranium using geometric morphometric techniques. It also aimed to explore the sex differences in distinct viscerocranial regions and to establish the most dimorphic region with regard to size and shape. Computed tomography images of 156 males and 184 females were used in the study. Three-dimensional coordinates of 31 landmarks were acquired. Five landmark configurations were constructed from the viscerocranium and its orbital, nasal, maxillary, and zygomatic region. Generalized Procrustes superimposition, principal component analysis, and discriminant analysis were applied to each configuration. The significance of the sex differences in size and shape was assessed and significant differences were found in all configurations. The highest accuracy was obtained from both shape and size of the whole viscerocranium. Based on size only, the highest accuracy was achieved by the nasal region. The accuracy based on shape was generally low for all configurations, but the highest result was attained by the orbital region. Hence, size is a better sex discriminator than shape.

Genetic Interaction of Thm2 and Thm1 Shapes Postnatal Craniofacial Bone.

Ciliopathies are genetic syndromes that link skeletal dysplasias to the dysfunction of primary cilia. Primary cilia are sensory organelles synthesized by intraflagellar transport (IFT)-A and B complexes, which traffic protein cargo along a microtubular core. We have reported that the deletion of the IFT-A gene, Thm2, together with a null allele of its paralog, Thm1, causes a small skeleton with a small mandible or micrognathia in juvenile mice. Using micro-computed tomography, here we quantify the craniofacial defects of Thm2-/-; Thm1aln/+ triple allele mutant mice. At postnatal day 14, triple allele mutant mice exhibited micrognathia, midface hypoplasia, and a decreased facial angle due to shortened upper jaw length, premaxilla, and nasal bones, reflecting altered development of facial anterior-posterior elements. Mutant mice also showed increased palatal width, while other aspects of the facial transverse, as well as vertical dimensions, remained intact. As such, other ciliopathy-related craniofacial defects, such as cleft lip and/or palate, hypo-/hypertelorism, broad nasal bridge, craniosynostosis, and facial asymmetry, were not observed. Calvarial-derived osteoblasts of triple allele mutant mice showed reduced bone formation in vitro that was ameliorated by Hedgehog agonist, SAG. Together, these data indicate that Thm2 and Thm1 genetically interact to regulate bone formation and sculpting of the postnatal face. The triple allele mutant mice present a novel model to study craniofacial bone development.

Sequence of chondrocranial development in basal anurans-Let's make a cranium.

BACKGROUND: The craniofacial skeleton is an evolutionary innovation of vertebrates. Due to its complexity and importance to protect the brain and aid in essential functions (e.g., feeding), its development requires a precisely tuned sequence of chondrification and/or ossification events. The comparison of sequential patterns of cartilage formation bears important insights into the evolution of development. Discoglossus scovazzi is a basal anuran species. The comparison of its chondrocranium (cartilaginous neuro- & viscerocranium) development with other basal anurans (Xenopus laevis, Bombina orientalis) will help establishing the ancestral pattern of chondrification sequences in anurans and will serve as basis for further studies to reconstruct ancestral conditions in amphibians, tetrapods, and vertebrates. Furthermore, evolutionary patterns in anurans can be studied in the light of adaptations once the ancestral sequence is established. RESULTS: We present a comprehensive overview on the chondrocranium development of D. scovazzi. With clearing and staining, histology and 3D reconstructions we tracked the chondrification of 44 elements from the first mesenchymal Anlagen to the premetamorphic cartilaginous head skeleton and illustrate the sequential changes of the skull. We identified several anuran and discoglossoid traits of cartilage development. In D. scovazzi the mandibular, hyoid, and first branchial arch Anlagen develop first followed by stepwise addition of the branchial arches II, III, and IV. Nonetheless, there is no strict anterior to posterior chondrification pattern within the viscerocranium of D. scovazzi. Single hyoid arch elements chondrify after elements of the branchial arch and mandibular arch elements chondrify after elements of the branchial arch I. CONCLUSIONS: In Osteichthyes, neurocranial elements develop in anterior to posterior direction. In the anurans investigated so far, as well as in D. scovazzi, the posterior parts of the neurocranium extend anteriorly, while the anterior parts of the neurocranium, extend posteriorly until both parts meet and fuse. Anuran cartilaginous development differs in at least two crucial traits from other gnathostomes which further supports the urgent need for more developmental investigations among this clade to understand the evolution of cartilage development in vertebrates.

Developmental changes of the facial skeleton from birth to 18 years within a South African cohort (A computed tomography study).

Skeletal remains are often found on a crime scene in which a forensic anthropologist is then consulted to create a biological profile, which includes the estimation of age, sex, ancestry and stature. The viscerocranium plays an important role in the formation of a biological profile. However, to utilise the viscerocranium for age estimation, population specific normative data and knowledge of the development of the viscerocranium is required. Therefore, this study aimed to investigate the developmental changes from birth to 18 years of age of the facial skeleton of individuals from a South African cohort. This study comprised of 239 computed tomography (CT) scans (128 males; 111 females). The viscerocranium was subdivided into five regions viz.: orbital, nasal, midfacial, maxillary and mandibular. The linear parameters in each region were correlated to age to identify the developmental growth patterns of the viscerocranial regions according to male and female. The measurements which displayed the highest correlations with age were used to develop formulas which could be used for age estimation. The results of this study showed that the measurements in the orbital, midfacial, maxillary and mandibular regions experienced rapid growth between 0 and 5 years of age, with the nasal region increasing steadily over time. It was noted that males displayed overall larger measurements than females except for the anterior interorbital distance and both right and left zygomatic arch lengths (ZAL). Although only the left orbital height, nasal aperture height and mandible width displayed statistically significant size differences according to sex (p ≤ 0.05). The measurements which showed the highest correlations to age were the zygomatic arch distance (r = 0.8842, p < 0.001), ZAL (right: r = 0.8929, p < 0.001; left: r = 0.8656, p < 0.001) and the mandible width (r = 0.8444, p < 0.001). Formulas were derived for the measurements that could be used to forensically estimate age within a subadult cohort.

Diferencias morfométricas en el hueso palatino de la oveja (Ovis aries) y la cabra (Capra hircus) / Morphometric palatine differences between sheep (Ovis aries) and goat (Capra hircus)

RESUMEN: Un hueso craneal muy poco estudiado en anatomía veterinaria comparada es el palatino. En nuestro estudio, realizamos la comparación de este hueso entre la oveja (Ovis aries L., 1758) y la cabra (Capra hircus L., 1758), a partir de una muestra de 36 y 17 cráneos respectivamente, y utilizando métodos de morfometría geométrica. Se eligieron un total de 32 puntos (4 hitos y 28 semi-hitos) para analizar la lámina horizontal del hueso palatino. Se reflejaron diferencias estadísticamente significativas tanto para el tamaño como para la forma. En Capra se denota una clara expansión lateral del hueso y rostral y una contracción central, con un acercamiento relativo de los forámenes palatinos, mientras que en Ovis los forámenes están más lateralmente alejados. A nuestro parecer, esta mayor anchura y longitud de la lámina horizontal en Capra se explicaría por una mayor capacidad de frotación del alimento en relación a Ovis.

SUMMARY: Palatine is a cranial bone very less studied in comparative veterinary anatomy. In our study, we performed the comparison of this bone between sheep (Ovis aries L., 1758) and goat (Capra hircus L., 1758), from a sample of 36 and 17 skulls respectively, using methods of geometric morphology. A total of 32 points (4 landmarks and 28 semi-landmarks) were chosen to analyse the horizontal lamina of the palatine bone. Statistically significant differences were reflected for both size and shape. Capra denotes a clear lateral expansion of the bone and rostral and a central contraction, with a relative approach of palatine foramina, while in Ovis foramina are more laterally distant. In our view, this greater width of the horizontal lamina in Capra would be explained by a greater capacity to rub the food compared to Ovis.

Fibroblast Growth Factor Receptors Function Redundantly During Zebrafish Embryonic Development.

Fibroblast growth factor (Fgf) signaling regulates many processes during development. In most cases, one tissue layer secretes an Fgf ligand that binds and activates an Fgf receptor (Fgfr) expressed by a neighboring tissue. Although studies have identified the roles of specific Fgf ligands during development, less is known about the requirements for the receptors. We have generated null mutations in each of the five fgfr genes in zebrafish. Considering the diverse requirements for Fgf signaling throughout development, and that null mutations in the mouse Fgfr1 and Fgfr2 genes are embryonic lethal, it was surprising that all zebrafish homozygous mutants are viable and fertile, with no discernable embryonic defect. Instead, we find that multiple receptors are involved in coordinating most Fgf-dependent developmental processes. For example, mutations in the ligand fgf8a cause loss of the midbrain-hindbrain boundary, whereas, in the fgfr mutants, this phenotype is seen only in embryos that are triple mutant for fgfr1a;fgfr1b;fgfr2, but not in any single or double mutant combinations. We show that this apparent fgfr redundancy is also seen during the development of several other tissues, including posterior mesoderm, pectoral fins, viscerocranium, and neurocranium. These data are an essential step toward defining the specific Fgfrs that function with particular Fgf ligands to regulate important developmental processes in zebrafish.

Larval development of the Mexican Snook, Centropomus poeyi (Teleostei: Centropomidae)

We document for the first time the early ontogeny of Centropomus poeyi based on captive raised material representing 0-19 days posthatch (dph). The achievement of early developmental landmarks (i.e., yolk-sac depletion, flexion, development of fins) and changes in pigmentation are described (1.4 mm NL-10.6 mm SL; 0-19 dph) and documented for a subset of individuals using high quality photographs. The ontogeny of the viscerocranium is also described (2.4 mm NL-10.6 mm SL; 6-19 dph). Development in C. poeyi occurs over a short period with attainment of the juvenile stage (i.e., full complement of fin rays present in each fin) occurring by 6.9 mm SL. The ontogeny of external pigmentation in C. poeyi is marked by two trends throughout growth: (1) a decrease in pigmentation dorsally; and (2) an increase in pigmentation ventrally along the midline. Development of the viscerocranium begins with the appearance of the maxilla and dentary in individuals of 2.4 mm NL, coinciding with the depletion of the yolk-sac. By 10.6 mm SL all bones of the viscerocranium are present and teeth are present on all teeth-bearing bones of the adult. Aspects of early development in C. poeyi are compared with the congeners C. undecimalis and C. parallelus.(AU)

Se documentó por primera vez la ontogenia temprana de Centropomus poeyi basada en material obtenido de cautiverio a partir de embriones (0 días después de la eclosión, dde) hasta los 19 dde. Se registró la aparición de puntos de referencia en el desarrollo temprano (p.ej. consumo de saco vitelino, flexión, desarrollo de aletas) y los cambios de pigmentación han sido descritos (1.4 mm NL-10.6 mm SL; 0-19 dde), siendo documentado para un subgrupo de individuos utilizando fotografías de alta calidad. El desarrollo del viscerocráneo también fue descrita (2.4 mm NL-10.6 mm SL; 6-19 dde). El desarrollo en C. poeyi sucede en un periodo corto de tiempo hasta alcanzar una etapa juvenil (p.ej., aparición completa de los radios presentes en cada aleta), lo que se observa a los 6.9 mm SL. La pigmentación externa en C. poeyi esta marcada por dos tendencias en el crecimiento: (1) un descenso en la pigmentación dorsal; y (2) un incremento en la pigmentación ventral a lo largo de la línea media. El desarrollo del viscerocráneo comienza con el aparecimiento del maxilar y el dentario en individuos de 2.4 mm NL, lo cual coincide con el consumo del saco vitelino. A los 10.6 mm SL todos los huesos del viscerocráneo están presentes como se observa en los dientes en los adultos. Algunos aspectos del desarrollo temprano en C. poeyi son comparados con los congéneros C. undecimalis y C. parallelus.(AU)

Craniometric characteristics and cranial indices of Polish heath sheep rams ­ extended data / Características craneométricas e índices craneales de ovejas heath polacas - datos ampliados

The study was conducted on the mouflon-derived horned sheep of the preserved Polish Heath Sheep breed. In this study, basic skull measurements of 24 skulls of the rams of different age: 1 day, 90 days, 210 days, 270 days, 3 and 6 years, were analyzed. The weight of each skull was determined, as well as 43 craniometric characteristics were measured, and cranial cavity volume and viscerocranial and neurocranial indices were calculated. From birth to the age of 6 years, the greatest length of the skull, the condylobasal length and the length of the cheek-tooth row increased two-fold. To day 210 of age, the bony housing for eyeball and its organs grew intensively. To day 270 of age, the largest increase was characteristic of the elements of the angular length of the neurocranium and its breadth. The structural elements characterizing both the breadth and the length of the viscerocranium developed to the age of 6 years. A significant (P0.05), decreasing with age, value of the cephalic index was observed, being mostly affected by development of the squamous part of the frontal bone with its orbital part and the orbital margin.

El estudio se realizó en la raza preservada de ovejas Heath, derivadas del Muflón. En este estudio se analizaron las mediciones craneales básicas de 24 cráneos de carneros de diferentes edades: 1 día, 90 días, 210 días, 270 días, 3 y 6 años. Se determinó el peso de cada cráneo, se midieron 43 características craneométricas y se calculó el volumen de la cavidad craneal y los índices viscerocraneales y neurocraneales. Desde el nacimiento hasta la edad de 6 años, la longitud más grande del cráneo, la longitud del cóndilo y la longitud de la hilera del diente de la mejilla aumentó dos veces. A los 210 días de edad, la cubierta ósea para el bulbo ocular y sus órganos creció intensivamente. Al día 270 de edad, el mayor incremento fue la longitud angular del neurocráneo y su amplitud. Los elementos estructurales que caracterizan tanto el ancho como la longitud del viscerocráneo se desarrollaron a la edad de 6 años. Se observó un valor significativo (P0,05), decreciente con la edad, del índice cefálico, siendo afectado principalmente por el desarrollo de la parte escamosa del hueso frontal con su parte orbitaria y el margen orbital.

The human brain and face: mechanisms of cranial, neurological and facial development revealed through malformations of holoprosencephaly, cyclopia and aberrations in chromosome 18.

The study of inborn genetic errors can lend insight into mechanisms of normal human development and congenital malformations. Here, we present the first detailed comparison of cranial and neuro pathology in two exceedingly rare human individuals with cyclopia and alobar holoprosencephaly (HPE) in the presence and absence of aberrant chromosome 18 (aCh18). The aCh18 fetus contained one normal Ch18 and one with a pseudo-isodicentric duplication of chromosome 18q and partial deletion of 18p from 18p11.31 where the HPE gene, TGIF, resides, to the p terminus. In addition to synophthalmia, the aCh18 cyclopic malformations included a failure of induction of most of the telencephalon - closely approximating anencephaly, unchecked development of brain stem structures, near absence of the sphenoid bone and a malformed neurocranium and viscerocranium that constitute the median face. Although there was complete erasure of the olfactory and superior nasal structures, rudiments of nasal structures derived from the maxillary bone were evident, but with absent pharyngeal structures. The second non-aCh18 cyclopic fetus was initially classified as a true Cyclops, as it appeared to have a proboscis and one median eye with a single iris, but further analysis revealed two eye globes as expected for synophthalmic cyclopia. Furthermore, the proboscis was associated with the medial ethmoid ridge, consistent with an incomplete induction of these nasal structures, even as the nasal septum and paranasal sinuses were apparently developed. An important conclusion of this study is that it is the brain that predicts the overall configuration of the face, due to its influence on the development of surrounding skeletal structures. The present data using a combination of macroscopic, computed tomography (CT) and magnetic resonance imaging (MRI) techniques provide an unparalleled analysis on the extent of the effects of median defects, and insight into normal development and patterning of the brain, face and their skeletal support.

An attempt to determine the size of biometric differences in the skull of two colour variants of American mink (Neovison vison) / Un intento de determinar la magnitud de las diferencias biométricas del cráneo de dos variedades de color del visón americano (Neovison vison)

This study aimed at answering the question whether production of new colour variants of American mink in mink farms using mutations may entail changes in skull morphology and relationships between the bone elements building it. Analyses were made on the skulls of 56 eight-month-old males and females of two American mink colour variants (standard Brown and mutant Sapphire) from the same farm. Mean values, standard deviations and coefficients of variation were determined for carcass weight, cranial and mandibular weights and 7 dorsal surface, 8 lateral surface and 11 basal surface traits of the skull. The values of 24 cranial and mandibular indices and the values of sexual size dimorphism (SSD), i.e. a coefficient describing differences between sexes, were calculated. It was demonstrated that mutant colour variants of American mink may be a significant source of variation (P0.05 and P0.01) for some traits of skull morphology and relationships between respective bone elements of viscerocranium and neurocranium.

El objetivo de este estudio fue responder a la pregunta de si la producción de nuevas variedades de color del visón americano en granjas mediante mutaciones puede causar cambios en la morfología del esqueleto de la cabeza y en las relaciones mutuas de los elementos óseos que lo construyen. Los estudios se realizaron en 56 machos y hembras de ocho meses de dos variedades de color del visón americano (bronce estándar y zafiro por mutación) derivados de la misma granja. Se determinaron valores medios, DE y coeficiente de variación para peso corporal, del cráneo y la mandíbula, además de 7 rasgos de la superficie del dorso, 8 de la superficie lateral y 11 características de la base del cráneo. Fueron calculados los valores de 24 índices craneales y mandibulares, junto al valor de la magnitud del dimorfismo sexual, i.e. un coeficiente de las diferencias entre los sexos. Se demostró que las variedades mutantes de color del visón pueden ser fuente significativa (p0,05 y p0,01) de variación para algunas características morfológicas del esqueleto de la cabeza y relaciones entre elementos óseos del víscero y neurocráneo.

Biometry of the skull of wild and farm long-tailed chinchilla (chinchilla laniger, molina, 1782) / Biometría del cráneo de la chinchilla de cola larga (chinchilla laniger, molina, 1782) salvaje y de granja

This study aimed at presenting the values of 47 metric traits and 20 cranial indices of the skull of 291 mature farm chinchillas and comparing these data with those being determined on 32 chinchilla skull specimens from the Natural History Museum in London. Measurements of the viscerocranium, neurocranium and mandible parameters were taken. No normal distribution of these traits was observed. The values of selected Spearman's rank correlation coefficients were calculated. It was found that most cranial traits of the farm chinchillas showed statistically significantly higher values (P0.01) when compared to those being determined on the skulls of museum specimens. The effect of the farm environment, in which the farm chinchillas had been kept for many generations, was a likely reason for these differences.

El objetivo fue presentar los valores de 47 caracteres métricos y 20 índices craneales del cráneo de 291 chinchillas de granja maduras y comparar estos datos con los determinados sobre 32 cráneos de chinchilla obtenidas desde el Museo de Historia Natural de Londres. Se tomaron mediciones de los parámetros del viscerocráneo, neurocráneo y de la mandíbula. No se observó una distribución normal de estos rasgos. Se calcularon los valores de los coeficientes de correlación de Spearman seleccionado. Se encontró que los rasgos craneales de las chinchillas de granja mostraron valores significativamente superiores (P 0,01) en comparación con los especímenes de museo. El efecto del entorno agrícola, en el que las chinchillas de granja se habían mantenido durante muchas generaciones, podría ser una razón probable para estas diferencias.