Cadaveric dissection of connection between accessory hemiazygos vein and left brachiocephalic vein.

PURPOSE: The azygos system of veins has many anatomical variations that can impact mediastinal and vascular interventions. While radiological reports on these are of great clinical value, this study is among the first to present a high-quality cadaveric dissection of a rare anatomical variant to supplement previously published radiologic studies. The azygos venous system consists of the azygos vein (AV), hemiazygos vein (HAV), and the accessory hemiazygos vein (AHAV), which develop from the last portion of the posterior cardinal veins. The normal anatomical configuration includes drainage of the posterior intercostal veins, vertebral vein, esophageal veins, HAV, and AHAV to an unpaired right-side AV at the level of the 8th/9th thoracic vertebra. The reported incidence of AHAV draining directly into the left brachiocephalic vein is 1-2%. METHODS: An adult formalin-fixed 70-year-old female cadaver was dissected as part of a medical gross anatomy elective course. RESULTS: Gross documentation of a direct connection of the HAV to the AHAV with the AHAV draining into the left brachiocephalic vein. CONCLUSION: It is important to note the variations of the azygos system to avoid confusion with a potential pathology such as mediastinal masses. Understanding of the rare variant reported here could be useful in the prevention of iatrogenic bleeding from the misplacement of venous catheters and help facilitate radiological diagnosis in the incidence of venous clot formation.

The nasopharynx as a window to half a billion years of evolutionary change to the upper respiratory tract.

The nasopharynx is a region at the nexus of several vital physiological systems, including the nasal cavity, oral cavity, braincase, middle ear, and cervical vertebrae. It has undergone pronounced morphological change over the course of tetrapod, mammalian, and human evolution. However, despite its place in evolutionary history, the nasopharynx has received relatively little attention. This special issue focuses on "the evolution, development, and functional morphology of the nasopharynx and its boundaries." Topics covered here include evolutionary developmental biology (or evo-devo), nasopharyngeal adaptions in bats, the importance of the nasopharynx and adjacent structures over the course of human evolution, normal development, middle ear morphology, clinical importance, and the study of the nasopharynx throughout history. Contributions to this special issue range among reviews and syntheses, descriptive analyses, phylogenetic analysis, traditional morphometrics, three-dimensional geometric morphometrics, and computational fluid dynamics. Here, we discuss the central importance of the nasopharynx as can be seen through vertebrate paleontology and comparative morphology. It is via the composite evolutionary history of the nasopharyngeal boundaries that our origins may be better understood, starting with the derivation of the choanae from the median olfactory pit of jawless fish nearly half a billion years ago to the basicranial flexion and facial reduction that distinguish Homo sapiens from all other living mammals. Indeed, the nasopharynx must be acknowledged for its importance in the processes of encephalization and acquisition of speech that have become the hallmark of our species.

Nasopharyngeal morphology contributes to understanding the "muddle in the middle" of the Pleistocene hominin fossil record.

The late archeologist Glynn Isaac first applied the term "muddle in the middle" to a poorly understood period in the Middle Pleistocene human fossil record. This study uses the nasopharyngeal boundaries as a source of traits that may inform this unclear period of human evolution. The nasopharynx lies at the nexus of several vital physiological systems, yet relatively little is known about its importance in human evolution. We analyzed a geographically diverse contemporary Homo sapiens growth series (n = 180 adults, 237 nonadults), Homo neanderthalensis (La Chapelle aux Saints 1, La Ferrassie 1, Forbes Quarry 1, Monte Circeo 1, and Saccopastore 1), mid-Pleistocene Homo (Atapuerca 5, Kabwe 1, Petralona 1, and Steinheim 1), and two Homo erectus sensu lato (KNM-ER 3733 and Sangiran 17). Methods include traditional (Analysis 1) and 3D geometric morphometric analysis (Analysis 2). H. erectus exhibited tall, narrow nasopharyngeal shape, a robust, ancestral morphology. Kabwe 1 and Petralona 1 plotted among H. sapiens in Analysis 2, exhibiting relatively shorter and vertical cartilaginous Eustachian tubes and vertical medial pterygoid plates. Atapuerca 5 and Steinheim 1 exhibited horizontal vomeral orientation similar to H. neanderthalensis, indicating greater relative soft palate length and anteroposterior nasopharynx expansion. They may exhibit synapomorphies with H. neanderthalensis, supporting the accretionary hypothesis. Species-level differences were found among H. sapiens and H. neanderthalensis, including relatively longer dilator tubae muscles and extreme facial airorhynchy among Neanderthals. Furthermore, H. neanderthalensis were autapomorphic in exhibiting horizontal pterygoid plate orientation similar to human infants, suggesting that they may have had inferiorly low placement of the torus tubarius and Eustachian tube orifice on the lateral nasopharyngeal wall in life. This study supports use of osseous nasopharyngeal boundaries both for morphological characters and understanding evolution of otitis media susceptibility in living humans.

Identification of critical windows in early development of human upper respiratory tract and middle ear disease.

Otitis media (OM) or middle ear disease is a prevalent pediatric condition generally related to early growth of the cartilaginous Eustachian tube (CET). This study used a developmental series of dry crania to reconstruct CET and dilator tubae (DT, the muscle opening the CET) morphology. Timing and directionality of CET and upper respiratory tract (URT) growth were investigated. Traditional and 3D geometric morphometrics (GM) were used to assess bony landmarks on the crania. The series was divided using dental eruption into seven growth stages ranging from before eruption of deciduous dentition (approximately the first 6 postnatal months) to eruption of the first permanent maxillary molar (after approximately 6 years). Bony endpoints of the CET and DT were used to calculate their morphology. GM analysis showed substantial shape differences between newborns, early infants, and all later developmental stages. Univariate measures showed the largest growth change between birth and 6 months. Subsequently, CET morphology changed little in the latter half of year 1, instead maturing gradually until approximately 3 years whereas DT relative length and orientation finish growth by the end of year 1. Incongruence in slower CET growth and faster DT growth could impact CET function between 6 and 12 months and be a contributing factor of OM. Tubal aeration may improve after this time when both CET and DT morphology mature, coinciding with clinically reported drop-off in ear infections.

The Nasal Complex of a Semiaquatic Artiodactyl, the Moose (Alces alces): Is it a Good Evolutionary Model for the Ancestors of Cetaceans?

Among Cetartiodactyla, cetaceans are the only obligate aquatic dwellers. Given morphological similarities between cetacean relatives such as Indohyus (the best represented Eocene raoellid artiodactyl) with other, later artiodactyls, any crown artiodactyl that engages in aquatic behaviors is of interest as an evolutionary model for the adaptations that accompanied the origins of cetaceans. The American moose (Alces alces) is the only non-cetacean artiodactyl to engage in aquatic foraging and, other than Hippopotamus, is distinctive in its diving behaviors. This study surveyed the soft and hard tissue nasal morphology of Alces alces to assess phylogenetic polarity and the presence of adaptations for diving and feeding in fresh water habitats. A fresh dissection of the facial musculature and nasal cavity was performed on one subadult male individual and osteological analyses were also performed on dry crania. This species was analyzed alongside fossil crania of Cervalces (its presumed ancestor), other cervids (e.g., Odocoileus virginianus, the white tail deer; Dama dama, the fallow deer), a bovid (Bos taurus, domestic cattle), and a carnivoran (Ursus americanus, the American black bear). A fresh dissection of the facial musculature and nasal anatomy of one fallow deer specimen was also performed for comparison with the moose. Results indicate that Alces alces exhibited a primitive configuration of maxillolabial muscles and, like Dama, exhibited a series of subcutaneous fibrous tissues connecting these muscles to skin. Alces and Dama, however, both exhibited autapomorphies in the soft tissue anatomy of the external nares. The former possessed a series of muscles that act to constrict the anterior nares, likely during diving. Extremely large fibrofatty pads that were perforated by muscle tendon supported their alar fold. Internally, a double-scrolled maxilloturbinal occupied nearly the entire volume of the anterior nasal cavity and protruded beyond the rim of the piriform aperture in dry crania. Dama had long, thin muscles taking origin on their nasal conchae and inserting onto the alar fold. Yet, despite these anterior nasal autapomorphies, the ethmoturbinal patterns of all observed cervids and the one bovid all appeared primitive with a posteroinferiorly oriented array of ethmoturbinals in close contact with a relatively straight cribriform plate, a macrosmatic condition. These differed from the curved cribriform plate of Ursus whose posterior nasal anatomy appeared hyper-macrosmatic. Indohyus exhibits no skeletal sign of a fleshy proboscis such as an enlarged piriform aperture or shortened nasal bones. Thus, there is little evidence that the early ancestors of cetaceans engaged in prolonged bouts of diving for aquatic foods but more probably were surface swimmers traveling between terrestrial food sources or fleeing predators. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 302:667-692, 2019. © 2018 Wiley Periodicals, Inc.

Cranial Indicators Identified for Peak Incidence of Otitis Media.

Acute otitis media (AOM) is one of the most common pediatric conditions worldwide. Peak age of occurrence for AOM has been identified within the first postnatal year and it remains frequent until approximately six postnatal years. Morphological differences between adults and infants in the cartilaginous Eustachian tube (CET) and associated structures may be responsible for development of this disease yet few have investigated normal growth trajectories. We tested hypotheses on coincidence of skeletal growth changes and known ages of peak AOM occurrence. Growth was divided into five dental eruption stages ranging from edentulous neonates (Stage 1) to adults with erupted third maxillary molars (Stage 5). A total of 32 three-dimensional landmarks were used and Generalized Procrustes Analysis was performed. Next, we performed principal components analysis and calculated univariate measures. It was found that growth change in Stage 1 was the most rapid and comprised the largest amount of overall growth in upper respiratory tract proportions (where time is represented by the natural logarithmic transformation of centroid size). The analysis of univariate measures showed that Stage 1 humans did indeed possess the relatively shortest and most horizontally oriented CET's with the greatest amount of growth change occurring at the transition to Stage 2 (eruption of deciduous dentition at five postnatal months, commencing peak AOM incidence) and ceasing by Stage 3 (approximately six postnatal years). Skeletal indicators appear related to peak ages of AOM incidence and may contribute to understanding of a nearly ubiquitous human disease. Anat Rec, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 300:1721-1740, 2017. © 2017 Wiley Periodicals, Inc.

Toward Understanding the Mammalian Zygoma: Insights From Comparative Anatomy, Growth and Development, and Morphometric Analysis.

The zygoma, or jugum, is a cranial element that was present in Mesozoic tetrapods, well before the appearance of mammals. Although as an entity the zygoma is a primitive retention among mammals, it has assumed myriad configurations as this group diversified. As the zygoma is located at the intersection of the visual, respiratory, and masticatory apparatuses, it is potentially of great importance in systematic, phylogenetic, and functional studies focused on this region. For example, the facial component of the zygoma and its contribution to a postorbital bar (POB) appear to be relevant to the systematics of a number of mammalian subclades, and the formation of a bony postorbital septum (POS) that separates the orbit from the infratemporal fossa is unique to, and thus potentially phylogenetically significant for uniting anthropoid primates, while the zygoma itself appears to serve to resist tension and bending forces during mastication. In order to better understand the zygoma in the context of its contributions to the circumorbital region, we documented its morphological expression in specimens representing 10 orders of mammals. Since the presence of a POB and of a POS has long been used to justify uniting extant primates and anthropoid primates as respective clades, and because postorbital closure (POC) is morphologically more complex than a POB, we provide detail necessary to address these claims. Our taxically broad overview also allowed us to provide for the first time definitions of configurations that can be applied to future studies. Using a different, but also taxically broad sample of mammals, and of primates in particular, we performed two geometric morphometric analyses that were geared toward testing long-held interpretations of the functional role of the zygoma, especially with regard to mastication and in the context of orbital frontation (to which the zygoma contributes). Further, overall, zygomatic morphology tends not to scale with allometry, sexual dimorphism, or angle of orbital convergence, but it does contribute to unique patterns of intraspecies variation. Anat Rec, 300:76-151, 2017. © 2016 Wiley Periodicals, Inc.

Three-dimensional geometric morphometric analysis of the nasopharyngeal boundaries and its functional integration with the face and external basicranium among extant hominoids.

The nasopharynx is a centrally located but understudied upper respiratory tract component. This study tested hypotheses related to the functional integration of the nasopharyngeal boundaries with the facial skeleton and external basicranium over the course of development in humans and nonhuman hominoids. It was hypothesized that facial morphology (width, length, and kyphosis) is related to nasopharyngeal width and choanal morphology, whereas relative external basicranial proportions are related to nasopharyngeal depth. Human infants were used as models of extreme orthognathy and external basicranial retroflexion, whereas nonhuman hominoids were used to model greater relative prognathism and external basicranial retroflexion. Both of these groups were contrasted against adult humans, who exhibit both extreme orthognathy and external basicranial flexion. Three-dimensional landmark coordinate data were collected from age-graded series of Homo, Pan, Gorilla, Pongo, and Hylobates. Generalized Procrustes Analysis was performed, and multivariate shape differences were evaluated via principal components analysis. Additionally, linear measures were extracted from the Procrustes-corrected sets of landmark data. Results indicate that human adults are indeed distinct from all groups in possessing a relatively shallow nasopharyngeal roof and shorter, more flexed external basicranial axis. Human adults and infants both exhibit greater relative choanal and nasopharyngeal width. Nonhuman hominoid faces tended to become airorhynch into adulthood, whereas humans exhibited the opposite trend. When pooling all the hominoids, facial width and palate length were strongly correlated with choanal and nasopharyngeal width, whereas facial kyphosis was strongly correlated with choanal orientation. The hypotheses were supported as the results indicated a morphologic relationship among nasopharyngeal boundaries, the facial skeleton, and the external basicranium.

The nasal complex of Neanderthals: an entry portal to their place in human ancestry.

Neanderthals are one of the most intensely studied groups of extinct humans, as aspects of their phylogeny and functional morphology remain controversial. They have long been described as cold adapted but recent analyses of their nasal anatomy suggest that traits formerly considered adaptations may be the result of genetic drift. This study performs quantitative and qualitative analysis of aspects of the nasal complex (NC) in Neanderthals and other later Pleistocene fossils from Europe and Africa. A geographically diverse sample of modern human crania was used to establish an anatomical baseline for populations inhabiting cold and tropical climates. Nasofrontal angle, piriform aperture dimensions, and relative maxillary sinus volume were analyzed along with qualitative features of the piriform aperture rim. Results indicate that Neanderthals and other later Pleistocene Homo possessed NC's that align them with tropical modern humans. Thus comparison of Neanderthal nasal morphology with that of modern humans from cold climates may not be appropriate as differences in overall craniofacial architecture may constrain the narrowing of the piriform apertures in Neanderthals. They retain primitively long, low crania, large maxillary sinuses, and large piriform aperture area similar to mid-Pleistocene Homo specimens such as Petralona 1 and Kabwe 1. Adaptation to cold climate may have necessitated other adaptations such as bony medial projections at the piriform aperture rim and, potentially, midfacial prognathism. Nasal complex components of the upper respiratory tract remain a critical but poorly understood area that may yet offer novel insight into one of the greatest continuing controversies in paleoanthropology.

Consequences of evolution: is rhinosinusitis, like otitis media, a unique disease of humans?

We hypothesize that if otitis media is most likely primarily a human disease due to consequences of evolution, rhinosinusitis may also be limited to humans for similar reasons. If otitis media, with its associated hearing loss, occurred in animals in the wild, they probably would have been culled out by predation. Similarly, if rhinosinusitis occurred regularly in animals, they likely would have suffered from severely decreased olfactory abilities, crucial for predator avoidance, and presumably would likewise have been selected against evolutionarily. Thus, both otitis media and rhinosinusitis-common conditions particularly in infants and young children-appear to be essentially human conditions. Their manifestation in our species is likely due to our unique evolutionary trajectory and may be a consequence of adaptations, including adaptations to bipedalism and speech, loss of prognathism, and immunologic and environmental factors.

Frontal fusion: collapse of another anthropoid synapomorphy.

We test the hypothesis that the fused interfrontal suture of anthropoids is a uniquely distinguishing feature and a derived characteristic indicative of their monophyletic origin. Our survey of nonanthropoid primates and several archontan families indicates frontal fusion is widespread. It is most variable (fused, open or partially fused) inter- and intra-specifically among strepsirhines. The frontal bone is more commonly fused in living lemuroids and indrioids than among lorisoids. It appears to be fused regularly among Eocene adapids. Among nonanthropoid haplorhines, the interfrontal is fused in Tarsius, even in neonates and invariably in adults, probably also in all fossil tarsiiforms preserving the frontal bone, and in the late Eocene protoanthropoid Rooneyia. The plesiadapiform pattern remains uncertain, but fusion is ubiquitous among living tree shrews, colugos and bats. Distributional evidence implies that interfrontal fusion was present in the last common ancestor (LCA) of haplorhine primates and possibly in the LCA of euprimates as well. Anthropoids, therefore, cannot be defined cladistically by interfrontal fusion, not out of concern for homoplasy but because it is probably a primitive feature inherited from other taxa related to anthropoids. Fusion of the large anthropoid frontal bone, which was extended anteriorly to roof the orbits and expanded laterally in connection with a wide forebrain in the LCA of anthropoids and protoanthropoids, may have been preadaptive to the evolution of the postorbital septum. The zygomatico-frontal suture of the septum may provide an alternative mechanism for dissipating the calvarial strains of mastication formerly taken up by an open interfrontal suture.