Diversity in primary palate ontogeny of amniotes revealed with 3D imaging

The amniote primary palate encompasses the upper lip and the nasal cavities. During embryonic development, the primary palate forms from the fusion of the maxillary, medial nasal and lateral nasal prominences. In mammals, as the primary palate fuses, the nasal and oral cavities become completely separated. Subsequently, the tissue demarcating the future internal nares (choanae) thins and becomes the bucconasal membrane, which eventually ruptures and allows for the essential connection of the oral and nasal cavities to form. In reptiles (including birds), the other major amniote group, primary palate ontogeny is poorly studied with respect to prominence fusion, especially the formation of a bucconasal membrane. Using 3D optical projection tomography, we found that the prominences that initiate primary palate formation are similar between mammals and crocodilians but distinct from turtles and lizards, which are in turn similar to each other. Chickens are distinct from all non-avian lineages and instead resemble human embryos in this aspect. The majority of reptiles maintain a communication between the oral and nasal cavities via the choanae during primary palate formation. However, crocodiles appear to have a transient separation between the oral and nasal cavities. Furthermore, the three lizard species examined here, exhibit temporary closure of their external nares via fusion of the lateral nasal prominences with the frontonasal mass, subsequently reopening them just before hatching. The mechanism of the persistent choanal opening was examined in chicken embryos. The mesenchyme posterior/dorsal to the choana had a significant decline in proliferation index, whereas the mesenchyme of the facial processes remained high. This differential proliferation allows the choana to form a channel between the oral and nasal cavities as the facial prominences grow and fuse around it. Our data show that primary palate ontogeny has been modified extensively to support the array of morphological diversity that has evolved among amniotes.     

Recent insights into the morphological diversity in the amniote primary and secondary palates

The assembly of the upper jaw is a pivotal moment in the embryonic development of amniotes. The upper jaw forms from the fusion of the maxillary, medial nasal, and lateral nasal prominences, resulting in an intact upper lip/beak and nasal cavities; together called the primary palate. This process of fusion requires a balance of proper facial prominence shape and positioning to avoid craniofacial clefting, whilst still accommodating the vast phenotypic diversity of adult amniotes. As such, variation in craniofacial ontogeny is not tolerated beyond certain bounds. For clarity, we discuss primary palatogenesis of amniotes into in two categories, according to whether the nasal and oral cavities remain connected throughout ontogeny or not. The transient separation of these cavities occurs in mammals and crocodilians, while remaining connected in birds, turtles and squamates. In the latter group, the craniofacial prominences fuse around a persistent choanal groove that connects the nasal and oral cavities. Subsequently, all lineages except for turtles, develop a secondary palate that ultimately completely or partially separates oral and nasal cavities. Here, we review the shared, early developmental events and highlight the points at which development diverges in both primary and secondary palate formation. Developmental Dynamics 244:1457–1468, 2015. © 2015 Wiley Periodicals, Inc.     

Ontogenetic variation in human nasal morphology

Internal nasal cavity morphology has long been thought to reflect respiratory pressures related to heating and humidifying inspired air. Yet, despite the widely recognized importance of ontogeny in understanding climatic and thermoregulatory adaptations, most research on nasal variation in modern and fossil humans focuses on static adult morphology. This study utilizes cross-sectional CT data of three morphologically distinct samples (African, European, Arctic) spanning from infancy to adulthood (total n = 321). Eighteen landmarks capturing external and internal regions of the face and nose were subjected to generalized Procrustes and form-space principal component analyses (separately conducted on global and individual samples) to ascertain when adult-specific nasal morphology emerges during ontogeny. Across the global sample, PC1 (67.18% of the variation) tracks age-related size changes regardless of ancestry, while PC2 (6.86%) differentiates between the ancestral groups irrespective of age. Growth curves tracking morphological changes by age-in-years indicate comparable growth trajectories across all three samples, with the majority of nasal size and shape established early in ontogeny (<5 years of age). Sex-based trends are also evident, with females exhibiting a more truncated growth period than males, particularly for nasal height dimensions. Differences are also evident between the anterior and posterior nose, with the height and breadth dimensions of the anterior nasal aperture and nasal cavity showing differential ontogenetic patterns compared to the choanae. Cumulatively, these results suggest that multiple selective pressures influence human nasal morphology through ontogenetic processes, including metabolic demands for sufficient oxygen intake and climatic demands for adequate intranasal air conditioning.     

New Interpretation of the Palate of Pterosaurs

On the basis of a new, three‐dimensionally preserved specimen of the Early Jurassic pterosaur Dorygnathus banthensis we present a reinterpretation of the pterosaur palate. The hard palate is formed by the extensive palatal plate of the maxilla and not by the palatine as has been generally reconstructed. This palatal plate of the maxilla emarginates the choana rostrally and rostrolaterally as in other archosaurs and lepidosaurs. The longitudinally elongate and dorsoventrally flat palatine in Dorygnathus is an isolated bone caudal to the palatal plate of the maxilla and morphologically and topographically it resembles that of crocodilians and birds, respectively. The palatine separates the choana laterally from the suborbital fenestra demonstrating the homologous nature of the (primary) choana in all archosaurs and lepidosaurs. Our study indicates that in basal pterosaurs the pterygo–ectopterygoid fenestra existed caudal to the suborbital fenestra, which became confluent with the adductor chamber in pterodactyloids thereby increasing the relative size of the adductor chamber and hence the mass of the jaw adductors. The choana in basal pterosaurs was relatively small compared with the interpterygoid vacuity. With increasing rostroventral inclination of the quadrates in more derived pterosaurs, the interpterygoid vacuity was reduced considerably, whereas the choana increased in size. This exceptional Dorygnathus specimen also shows a hitherto unknown pair of fenestrae situated at the palatal contact of the premaxilla–maxilla and might represent the aperture for the vomeronasal organ. Anat Rec, 2010. © 2009 Wiley‐Liss, Inc.