Development of the nasal capsule cartilages was studied in seven Geoffroy's tamarins (Saguinus geoffroyi), including one fetus, five neonates and one infant. Four additional postnatal specimens of the genus were studied (one 5-month-old and three adults) to determine the magnitude of postnatal expansion of the paranasal sinuses. Alcian blue histochemistry and osteopontin immunohistochemistry were employed in selected subadult specimens to characterize cartilage matrix. The fetal S. geoffroyi possesses a continuous nasal capsule, including a zona anularis; the primordial maxillary sinuses are surrounded by cartilage. Secondary pneumatization is in progress in all older specimens, which have sinuses that are more than twofold larger compared to that of the fetus. Results indicate that extensive ossification of the middle part of the nasal capsule (pars intermedia) is occurring in the perinatal timeframe, forming portions of the ethmoid bone. Anteriorly, the nasal capsule comprises isolated fragments in perinatal specimens, which are fewer and smaller in the infant and in a 5-month-old S. midas, and nearby multinucleate cells suggest that osteoclasts break apart these initially continuous elements. Fragments of the pars intermedia and the tectum nasi are found transiently between mucosa and the sites of secondary pneumatization. The maxillary sinus mucosa is highly vascular in most perinatal specimens. Histochemical and immunohistochemical findings show that cartilage of endochondral bones and non-ossifying parts are distinct in the perinatal time period. These results indicate that breakdown of the capsular cartilage precedes secondary pneumatization as previously suggested. There are portions of the cartilage of the recessus maxillaris and tectum nasi that transiently block mucosa from interfacing directly with bone. Vascularization may play a role in the breakdown of cartilages as well as the onset of secondary pneumatization. Since cartilage has the capacity to produce substances that trigger angiogenesis and bone resorption, further detailed characterization of the cartilage bordering sites of secondary pneumatization is merited. Anat Rec, 291:1397-1413, 2008.(c)2008 Wiley-Liss, Inc.

The air filled cavities of paranasal sinuses are thought by some to appear opportunistically in spatial "gaps" within the craniofacial complex. Anthropoid primates provide excellent natural experiments for testing this model, since not all species possess a full complement of paranasal sinuses. In this study, two genera of monkeys (Saguinus and Cebuella) which form maxillary sinuses (MS) as adults were compared to squirrel monkeys (Saimiri spp.), in which a MS does not form. Using microCT and histomorphometric methods, the spatial position of paranasal spaces was assessed and size of the adjacent dental sacs was measured. In Saguinus, secondary pneumatization is underway perinatally, and the sinus extends alongside deciduous premolars (dp). The MS overlaps all permanent molars in the adult. In Saimiri, the homologous space (maxillary recess) extends no farther posterior than the first deciduous premolar at birth and extends no farther than the last premolar in the adult. Differences in dental size and position may account for this finding. For example, Saimiri has significantly larger relative dp volumes, and enlarged orbits, which encroach on the internasal space to a greater degree when compared to Saguinus. These factors limit space for posterior expansion of the maxillary recess. These findings support the hypothesis that secondary pneumatization is a novel, opportunistic growth mechanism that removes "unneeded" bone. Moreover, paranasal spaces occur in association with semiautonomous skeletal elements that border more than one functional matrix, and the spatial dynamics of these units can act as a constraint on pneumatic expansion of paranasal spaces. Anat Rec, 2009.(c) 2009 Wiley-Liss, Inc.

Development of the nasal capsule cartilages was studied in seven Geoffroy's tamarins (Saguinus geoffroyi), including one fetus, five neonates and one infant. Four additional postnatal specimens of the genus were studied (one 5-month-old and three adults) to determine the magnitude of postnatal expansion of the paranasal sinuses. Alcian blue histochemistry and osteopontin immunohistochemistry were employed in selected subadult specimens to characterize cartilage matrix. The fetal S. geoffroyi possesses a continuous nasal capsule, including a zona anularis; the primordial maxillary sinuses are surrounded by cartilage. Secondary pneumatization is in progress in all older specimens, which have sinuses that are more than twofold larger compared to that of the fetus. Results indicate that extensive ossification of the middle part of the nasal capsule (pars intermedia) is occurring in the perinatal timeframe, forming portions of the ethmoid bone. Anteriorly, the nasal capsule comprises isolated fragments in perinatal specimens, which are fewer and smaller in the infant and in a 5-month-old S. midas, and nearby multinucleate cells suggest that osteoclasts break apart these initially continuous elements. Fragments of the pars intermedia and the tectum nasi are found transiently between mucosa and the sites of secondary pneumatization. The maxillary sinus mucosa is highly vascular in most perinatal specimens. Histochemical and immunohistochemical findings show that cartilage of endochondral bones and non-ossifying parts are distinct in the perinatal time period. These results indicate that breakdown of the capsular cartilage precedes secondary pneumatization as previously suggested. There are portions of the cartilage of the recessus maxillaris and tectum nasi that transiently block mucosa from interfacing directly with bone. Vascularization may play a role in the breakdown of cartilages as well as the onset of secondary pneumatization. Since cartilage has the capacity to produce substances that trigger angiogenesis and bone resorption, further detailed characterization of the cartilage bordering sites of secondary pneumatization is merited. Anat Rec, 291:1397-1413, 2008.(c)2008 Wiley-Liss, Inc.

Ventral view of a computer-generated, 3D reconstruction of the nasal capsule in a 16 mm CRL Microcebus murinus. The position of the eyes is lateral to the nasal capsule. See Smith et al., Anatomical Record 291:895-915.

Turbinals (scroll bones, turbinates) are projections from the lateral wall of the nasal fossa. These bones vary from simple folds to branching scrolls. Conventionally, maxilloturbinals comprise the respiratory turbinals, whereas nasoturbinals and ethmoturbinals comprise olfactory turbinals, denoting the primary type of mucosa that lines these conchae. However, the first ethmoturbinal (ETI) appears exceptional in the variability of it mucosal covering. Recently, it was suggested that the distribution of respiratory versus olfactory mucosae varies based on body size or age in strepsirrhine primates (lemurs and lorises). The present study was undertaken to determine how the rostrocaudal distribution of olfactory epithelium (OE) versus non-OE scales relative to palatal length in strepsirrhines. Serially sectioned heads of 20 strepsirrhines (10 neonates, 10 adults) were examined for presence of OE on ETI, rostral to its attachment to the nasal fossa wall (lateral root). Based on known distances between sections of ETI, the rostrocaudal length of OE was measured and compared to the length lined solely by non-OE (primarily respiratory epithelium). In 13 specimens, the total surface area of OE versus non-OE was calculated. Results show that the length of non-OE scales nearly isometrically with cranial length, while OE is more negatively allometric. In surface area, a lesser percentage of non-OE exists in smaller species than larger species and between neonates and adults. Such results are consistent with recent suggestions that the olfactory structures do not scale closely with body size, whereas respiratory structures (e.g., maxilloturbinals) may scale close to isometry. In primates and perhaps other mammals, variation in ETI morphology may reflect dual adaptations for olfaction and endothermy. Anat Rec, 2007.(c) 2007 Wiley-Liss, Inc.

The ontogeny of the nasolacrimal ducts (NLD) and canals (NLC) are investigated in strepsirrhine and haplorhine primates. Developmental series of serially sectioned fetal, perinatal and adult specimens, in combination with juvenile and adult skulls subjected to high-resolution computed tomography, reveal that the vertical NLC and NLD of adult tarsiers and anthropoids are produced by the degeneration of a more horizontal anterior arm of the NLD that is present only transiently in haplorhines, but is maintained throughout life in strepsirrhines. This degeneration manifests as an 'unzipping' of the anterior arm by means of progressive enlargement (in a rostral direction) of a caudally placed opening of the NLD (at the base of the vertical NLC), followed by breakdown of the resulting epithelial groove. The similar mode by which the anterior arm of the membranous NLD degenerates in tarsiers and anthropoids strongly suggests that the conditions in these two taxa are homologous, and provides additional evidence for a monophyletic Haplorhini. The functional relationship between the nasolacrimal duct and the vomeronasal organ is reviewed in light of this evidence, and it is suggested that these changes in the haplorhine NLD were functionally linked to the development of anatomical haplorhinism of the oronasal complex.

The paranasal sinuses remain elusive both in terms of function and in the proximate mechanism of their development. The present study sought to describe the maxillary sinuses (MSs) in three species of callitrichid primates at birth, a time when secondary pneumatization occurs rapidly in humans. The MSs were examined in serially sectioned and stained slides from the heads of two Callithrix jacchus, one Leontopithecus rosalia, and two Saguinus geoffroyi. Specimens were examined microscopically regarding the distribution of osteoclasts and osteoblasts along the osseous boundaries of the MS and other parts of the maxillary bone. Selected sections were immunohistochemically evaluated for the distribution of osteopontin (OPN), which facilitates osteoclast binding. Taken together, OPN immunoreactivity and bone cell distribution suggested trends of bone resorption/deposition that were consistent among species for the superior (roof) and inferior (floor) boundaries of the MS. Expansion at the roof and floor of the MS appeared to correspond to overall vertical midfacial growth in callitrichids. Much more variability was noted for the lateral (alveolar) and medial (nasal walls) of the MS. Unlike the other species, the nasal wall of Saguinus was static and mostly composed of inferior portions of the nasal capsule that were undergoing endochondral ossification. The variation seen in the alveolar walls may relate to the presence or absence of adjacent structures, although it was noted that adjacency of deciduous molars influenced medial drift of the alveolar wall in Saguinus but not Leontopithecus. The results of this study are largely consistent with the "structural" or "architectural" hypothesis of sinus formation with respect to vertical MS enlargement, and the variable cellular/OPN distribution found along the nasal and alveolar walls was evocative of Witmer's (J Vert Paleontol 1997;17:1-73) epithelial hypothesis in revealing that most expansion occurred in regions unopposed by adjacent structures.

Interpretations of the postcranial anatomy of East African early and middle Miocene large-bodied hominoids (e.g., Proconsul, Afropithecus, Turkanapithecus, Nacholapithecus) have suggested that these diverse primates utilized positional behaviors dominated by arboreal quadrupedalism. Preliminary descriptions of the Equatorius africanus partial skeleton (KNM-TH 28860) and other forelimb specimens, however, have argued that this animal relied more on terrestrial locomotion compared to its contemporaries, possibly similar to extant large papionin monkeys. In this paper, we reevaluate this interpretation by examining intrinsic hand proportions based on the lengths of the third proximal phalanx and fifth metacarpal in Equatorius in reference to a large sample of extant catarrhine primate taxa. We focused on the lengths of these hand bones because the ratio between phalanx and metacarpal lengths has been previously documented to discriminate terrestrial from arboreal mammalian taxa, including primates. The Equatorius hand displays semi-terrestrial hand proportions with a relatively shorter proximal phalanx compared to most arboreal monkeys. Its proximal phalanx, however, is relatively longer than those of habitually terrestrial monkeys (e.g., Theropithecus, Papio). Accordingly, although Equatorius retains some arboreal quadrupedal characteristics, these results corroborate the previous inference that it engaged in more terrestrial locomotion than earlier Miocene apes such as Proconsul. We suggest that the postcranial skeleton of Equatorius evinces the earliest signs of semi-terrestriality in the hominoid fossil record. It is likely that the terrestrial specialization utilized by living hominoids, e.g., knuckle-walking, evolved separately.

The remarkable progress in characterizing the human genome sequence, exemplified by the Human Genome Project and the HapMap Consortium, has led to the perception that knowledge and the tools (e.g., microarrays) are sufficient for many if not most biomedical research efforts. A large amount of data from diverse studies proves this perception inaccurate at best, and at worst, an impediment for further efforts to characterize the variation in the human genome. Because variation in genotype and environment are the fundamental basis to understand phenotypic variability and heritability at the population level, identifying the range of human genetic variation is crucial to the development of personalized nutrition and medicine. The Human Variome Project (HVP; http://www.humanvariomeproject.org/) was proposed initially to systematically collect mutations that cause human disease and create a cyber infrastructure to link locus specific databases (LSDB). We report here the discussions and recommendations from the 2008 HVP planning meeting held in San Feliu de Guixols, Spain, in May 2008.

The extent of fusion in facial sutures has implications for topics ranging from biomechanics to phylogeny reconstruction. An unfortunate limitation of studying sutural fusion in skeletal specimens is that it is difficult to assess whether apparently patent sutures are in fact fused internally. Both histology and microcomputed tomography (CT) are potential tools for solving this, but relatively few studies have attempted to discern the limits of micro CT for visualization of microanatomical structures. We examined microanatomical aspects of facial sutures in adult cadaveric samples from captive bushbabies. Premaxillary and nasopremaxillary sutures were examined in serially sectioned snouts of four greater bushbabies (Otolemur garnettii) and four lesser bushbabies (Galago moholi). Sections containing sutures with osseous bridging were rated as "fused," and the presence or absence of grooves on the external side was recorded. One bushbaby was studied using micro CT prior to physical sectioning. O. garnettii and two of the G. moholi show multiple foci of fusion. Histological examination confirmed that sutural fusion is limited to the internal surface in numerous sections, resulting in an external notch. Such points of internal fusion could be clearly visualized in raw CT slices. The presence of such notches suggests that external examination can underestimate the degree of suture fusion. Thus, microanatomical evidence may be needed to fully assess biomechanical correlates and phylogenetic interpretations based on fusion of facial sutures. Our results also indicate micro CT may be a useful tool to obtain this evidence. Am J Phys Anthropol 2009.(c) 2009 Wiley-Liss, Inc.

The orbicularis oris muscle plays a role in the production of primate facial expressions and vocalizations, nutrient intake, and in some non-human primates it is used as a prehensile, manipulative tool. As the chimpanzee (Pan troglodytes) is the closest living relative of humans, a comparison of the orbicularis oris muscle between these species may increase our understanding of the morphological specializations related to the differing functional demands of their lips and the factors responsible for their divergent evolution. To this end, this study compares the microanatomy of the mid-line upper fibers of the orbicularis oris muscle between chimpanzees and humans. A mid-line portion of the orbicularis oris muscle was harvested from the upper lips of three chimpanzee and five human cadavers. The sampled blocks included the area between the lateral borders of the nasal alar cartilages in both species. Each sample was processed for paraffin histology, sectioned and stained with a variety of protocols. Sections were examined for fiber direction and relative thickness of muscle layers. Ratios of cross-sectional connective tissue area vs. cross-sectional muscle tissue area, muscle fiber diameter and relative dermal thickness were calculated for each species. In both species, a clear pars marginalis layer was recognized, contrary to previous reports that only humans possess this layer. In chimpanzees, the relative fiber diameter and relative amount of muscle tissue (i.e. based on ratio of connective tissue area : muscle tissue area) were significantly (P < 0.05) greater than in humans. In contrast, measurements of relative dermal thickness showed that humans have a greater average dermal thickness of the upper lip than chimpanzees. Taken together, these results suggest that both human and chimpanzee orbicularis oris muscle upper fibers meet the specific functional demands associated with their divergent vocal and facial display repertoires, the development of human speech, and the use of the upper lip as a prehensile tool in chimpanzees.