Shape variation in modern human upper premolars.

Morphological variation in modern human dentition is still an open field of study. The understanding of dental shape and metrics is relevant for the advancement of human biology and evolution and is thus of interest in the fields of dental anthropology, as well as human anatomy and medicine. Of concern is also the variation of the inner aspects of the crown which can be investigated using the tools and methods of virtual anthropology. In this study, we explored inter- and intra-population morphometric variation of modern humans' upper third and fourth premolars (P3s and P4s, respectively) considering both the inner and outer aspects of the crown, and discrete traits. We worked by means of geometric morphometrics on 3D image data from a geographically balanced sample of human populations from five continents, to analyse the shape of the dentinal crown, and the crown outline in 78 P3s and 76 P4s from 85 individuals. For the study of dental traits, we referred to the Arizona State University Dental Anthropology System integrated with more recent classification systems. The 3D shape variation of upper premolar crowns varied between short and mesio-distally broad, and tall and mesio-distally narrow. The observed shape variation was independent from the geographical origin of the populations, and resulted in extensive overlap. We noted a high pairwise correlation (r1 = 0.83) between upper P3s and P4s. We did not find any significant geographic differences in the analysed non-metric traits. Our outcomes thus suggest that geographical provenance does not play a determinant role in the shaping of the dental crown, whose genesis is under strict genetic control.

Robert M. Ricketts and Rudolf Slavicek: dentistry by the rules of nature.

The stomatognathic structures act as a complex and integrated system, thereby accomplishing several essential functions of the body. Aside from participating in food digestion, they are key for respiration and swallowing and play a central role in social interaction and stress management. The lifeworks of Robert M. Ricketts (1920-2003), an American orthodontist, and Rudolf Slavicek (1928-2022), an Austrian prosthodontist, were centered on this understanding. Both were educated in the time of gnathology, functional dentistry, and cephalometry and were ready to challenge conventional knowledge and traditions, leading toward innovation. As untiring clinicians, researchers, and mentors, they were fully invested in the study of the stomatognathic system, considering its morphology, dynamics, growth patterns, evolution, and interactions with the body and mind. Based on their extensive knowledge of the masticatory system, they advanced dentistry both with theoretical notions and by implementing new diagnostic and therapeutic concepts, thus reinforcing the idea of dentistry as a medical discipline requiring interdisciplinary effort. Their heritage is represented by numerous publications, discoveries, and inventions that inspire the dental community to follow their exemplary approach to the individualized care of patients. Their knowledge and passion are further passed on through their students. As part of their legacy, they prepared the ground for new research aimed at fostering advancements in occlusion medicine, hence supporting education in oral health.

Initial therapeutic approaches for orofacial myofascial pain: three pilot studies.

OBJECTIVE: Myofascial pain diminishes the stomatognathic function and hinders clinical diagnosis. Therefore, initial pain reduction is crucial before definitive treatment. Here, the clinical validity of non-pharmaceutical therapies, including the Aqualizer® splint, physiotherapy, and dry-needle acupuncture was comparatively assessed. METHODS: Myofascial pain patients (n = 28; 20-65 years old) were examined through a visual analog scale, and intra- and extra-oral muscle palpation. Mandibular maximum opening and neck mobility were also evaluated. Changes in parameters through time were analyzed via the Kruskal-Wallis test, while the Friedman test and dot-plots were used for comparative therapies assessment. General patient improvement was represented via an isometric Principal Component. RESULTS: The Aqualizer® and physiotherapy resulted in improvement of all parameters except for mouth opening. Acupuncture improved extra-oral muscle pain and neck mobility. CONCLUSION: The Aqualizer®, physiotherapy, and oral acupuncture are effective initial pain therapies. Among all, physiotherapy provided the greatest benefits, followed by the Aqualizer®.

Premolar Extraction Affects Mandibular Kinematics.

OBJECTIVES: The practice of premolar extraction in orthodontics is controversial for its potential detrimental effects on the stomatognathic system. However, the ways in which premolar extraction affects mandibular function are still poorly understood. The purpose of this study was to assess the influence of premolar extraction on mandibular kinematics by evaluating axiographic tracings. MATERIALS AND METHODS: Forty-five orthodontically treated patients with premolar teeth extraction were compared with 45 paired untreated controls, selected for the absence of malocclusions. Systematic three-dimensional axiographic recordings of the mandibular movements were performed for protrusive-retrusive movements and speech. The transversal deviations and length of the movements were recorded for both sides along with the rotation angle during speech. STATISTICAL ANALYSIS: Differences between the axiographic variables were analyzed via the permutation test and Wilcoxon rank-sum test. Linear regression was performed to test whether axiographic parameters were predictive of group affiliation. Dot plots were used to explore the distribution of each of the axiographic outcomes, and isometric principal component analysis to assess the differences between the cumulative effects of premolar extraction on jaw motion. RESULTS: The mandibular lateral translation in protrusion-retrusion and speech, the amount of rotation as well as the length of mandibular movements during speech were significantly higher in the treated subjects than in the controls, while retral stability did not differ. The linear regression yielded significant results for the mandibular lateral translation in protrusion-retrusion. The isometric principal component analysis showed higher values of the axiographic variables for 11 out of 45 individuals in the study sample compared with the control group. CONCLUSIONS: Premolar extraction altered mandibular kinematics in at least 25% of the cases within our sample, and the transversal discrepancy between protrusive and retrusive tracings was even predictive of group affiliation. These results support the notion that the routine practice of premolar extraction as part of the orthodontic treatment should be discouraged. It is compelling to perform further studies to assess whether a disrupted kinematics of the mandible is associated to temporomandibular disorders.

Dynamic finite-element simulations reveal early origin of complex human birth pattern.

Human infants are born neurologically immature, potentially owing to conflicting selection pressures between bipedal locomotion and encephalization as suggested by the obstetrical dilemma hypothesis. Australopithecines are ideal for investigating this trade-off, having a bipedally adapted pelvis, yet relatively small brains. Our finite-element birth simulations indicate that rotational birth cannot be inferred from bony morphology alone. Based on a range of pelvic reconstructions and fetal head sizes, our simulations further imply that australopithecines, like humans, gave birth to immature, secondary altricial newborns with head sizes smaller than those predicted for non-human primates of the same body size especially when soft tissue thickness is adequately approximated. We conclude that australopithecines required cooperative breeding to care for their secondary altricial infants. These prerequisites for advanced cognitive development therefore seem to have been corollary to skeletal adaptations for bipedal locomotion that preceded the appearance of the genus Homo and the increase in encephalization.

Centric relation: A matter of form and substance.

The recent review article by Zonnenberg, Türp and Greene 'Centric relation critically revisited - What are the clinical implications'? opens an important debate by addressing topics of central relevance in Dentistry, namely the relationship between occlusion and the condyle-to-glenoid-fossa position, and the need for diagnostic assessment and therapeutic alteration of the condylar position in orthodontic patients. Zonnenberg, Türp and Greene concluded that the mandibular condyle is correctly situated in most orthodontic patients. Thus, in their view, orthodontists can disregard this aspect during treatment, and rely on the plastic properties of the masticatory supporting structures, while aiming at finishing the cases in a good occlusal relationship. We think that this approach fails to consider that biological variation of the stomatognathic structures can also be pathological and that, as dental occlusion determines condylar relative position within the glenoid fossa, changes in the occlusion are likely to alter the original condylar-to-glenoid-fossa relation. Hence, we claim that whenever the occlusal relationship must be changed, the clinician should carefully monitor the condyle position and the mandibular function to prevent possible iatrogenic effects. To advance the discourse on the topic, we invite Zonnenberg, Türp and Greene to clarify their definition of 'average patient' and their interpretation of 'full-mouth orthodontic and orthognathic treatment', their understanding of 'biologically acceptable condylar relationship', their justification of maximum intercuspation as reference position, the extent to which they think it is safe to rely on the TMJ resilience, and finally their alternative to centric relation in the treatment of patients needing condylar repositioning.

Sex classification using the human sacrum: Geometric morphometrics versus conventional approaches.

The human pelvis shows marked sexual dimorphism that stems from the conflicting selective pressures of bipedal locomotion and parturition. The sacrum is thought to reflect this dimorphism as it makes up a significant portion of the pelvic girdle. However, reported sexual classification accuracies vary considerably depending on the method and reference sample (54%-98%). We aim to explore this inconsistency by quantifying sexual dimorphism and sex classification accuracies in a geographically heterogeneous sample by comparing 3D geometric morphometrics with the more commonly employed linear metric and qualitative assessments. Our sample included 164 modern humans from Africa, Europe, Asia, and America. The geometric morphometric analysis was based on 44 landmarks and 56 semilandmarks. Linear dimensions included sacral width, corpus depth and width, and the corresponding indices. The qualitative inspection relied on traditional macroscopic features such as proportions between the corpus of the first sacral vertebrae and the alae, and sagittal and coronal curvature of the sacrum. Classification accuracy was determined using linear discriminant function analysis for the entire sample and for the largest subsamples (i.e., Europeans and Africans). Male and female sacral shapes extensively overlapped in the geometric morphometric investigation, leading to a classification accuracy of 72%. Anteroposterior corpus depth was the most powerful discriminating linear parameter (83%), followed by the corpus-area index (78%). Qualitative inspection yielded lower accuracies (64-76%). Classification accuracy was higher for the Central European subsample and diminished with increasing geographical heterogeneity of the subgroups. Although the sacrum forms an integral part of the birth canal, our results suggest that its sex-related variation is surprisingly low. Morphological variation thus seems to be driven also by other factors, including body size, and sacrum shape is therefore likely under stronger biomechanical rather than obstetric selection.

The morphological consequences of segmentation anomalies in the human sacrum.

OBJECTIVES: Despite the high frequency of segmentation anomalies in the human sacrum, their evolutionary and clinical implications remain controversial. Specifically, inconsistencies involving the classification and counting methods obscure accurate assessment of lumbosacral transitional vertebrae. Therefore, we aim to establish more reliable morphological and morphometric methods for differentiating between sacralizations and lumbarizations in clinical and paleontological contexts. MATERIALS AND METHODS: Using clinical CT data from 145 individuals aged 14-47 years, vertebral counts and the spatial relationship between the sacrum and adjoining bony structures were assessed, while the morphological variation of the sacrum was assessed using geometric morphometrics based on varied landmark configurations. RESULTS: The prevalence of lumbosacral and sacrococcygeal segmentation anomalies was 40%. Lumbarizations and sacralizations were reliably distinguishable based on the spatial relationship between the iliac crest and the upward or downward trajectory of the linea terminalis on the sacrum. Different craniocaudal orientations of the alae relative to the corpus of the first sacral vertebra were also reflected in the geometric morphometric analyses. The fusion of the coccyx (32%) was frequently coupled with lumbarizations, suggesting that the six-element sacra more often incorporate the coccyx rather than the fifth lumbar vertebra. CONCLUSIONS: Our approach allowed the consistent identification of segmentation anomalies even in isolated sacra. Additionally, our outcomes either suggest that homeotic border shifts often affect multiple spinal regions in a unidirectional way, or that sacrum length is highly conserved perhaps due to functional constraints. Our results elucidate the potential clinical, biomechanical, and evolutionary significance of lumbosacral transitional vertebrae.

Sex determination accuracy using the human sacrum in a Central European sample.

The sacrum is often used for sex estimation of human remains. However, reported sexual classification accuracies based on the sacrum produce highly discrepant estimates ranging from 54% to 98% depending on the method and reference sample. Here, we therefore aim to evaluate the sacrum's efficacy in differentiating males from females by comparing three different approaches in a homogeneous Central European sample (n = 58, 27 males and 31 females) obtained from the 19th century Weisbach collection. Specifically, we investigated the sacrum by 1) a qualitative visual inspection, 2) traditional linear metrics, namely, sacral width, corpus width, and the associated corporo-basal index, and 3) geometric morphometrics (GM) using a 3D configuration of 100 landmarks and semilandmarks. Classification accuracies for the qualitative approach ranged from 69% to 81%. The investigated quantitative methods based on linear dimensions led to comparable classification accuracies of 62% for sacral width and 78% for the corporo-basal index. However, absolute corpus width had a stronger discriminative power (86%), similar to the PC scores from shape space (87%). Unexpectedly, the GM approach, which considered the shape of the entire sacrum, did not classify better than linear variables. This was exceeded only when sacrum shape was combined with corpus width or another measure of size, like the natural logarithm of the centroid size, yielding a classification accuracy of 95%. The male and female group means differed mainly in the general height-to-width relationship and corpus-to-alae proportions, corroborating patterns previously described in the literature. Our results suggest that the sacrum is markedly less effective than the hipbone for sex determination, which is at odds with its central position within the pelvic girdle and thus its supposed obstetric relevance.

Response to Comment on "A Middle Pleistocene Homo from Nesher Ramla, Israel".

Marom and Rak claim, on the basis of a few mandibular features, that the Nesher Ramla (NR) Homo is a Neanderthal. Their comments lack substance and contribute little to the debate surrounding the evolution of Middle Pleistocene Homo. Limitations and preconceptions in their study prevented them from achieving resolution beyond a dichotomous interpretation of the NR as either a Neanderthal or a modern human.

New insights on hip bone sexual dimorphism in adolescents and adults using deformation-based geometric morphometrics.

Morphological variation of the human pelvis, and particularly the hip bone, mainly results from both female-specific selective pressure related to the give birth of large-headed newborns, and constraints in both sexes for efficient bipedal locomotion, abdominal stability, and adaptation to climate. Hip bone morphology has thus been extensively investigated using several approaches, although the nuances of inter-individual and sex-related variation are still underappreciated, and the effect of sex on ontogenetic patterns is debated. Here, we employ a landmark-free, deformation-based morphometric approach to explore variation in modern human hip bone shape and size from middle adolescence to adulthood. Virtual surface models of the hip bone were obtained from 147 modern human individuals (70 females and 77 males) including adolescents, and young and mature adults. The 3D meshes were registered by rotation, translation, and uniform scaling prior to analysis in Deformetrica. The orientation and amplitude of deviations of individual specimens relative to a global mean were assessed using Principal Component Analysis, while colour maps and vectors were employed for visualisation purposes. Deformation-based morphometrics is a time-efficient and objective method free of observer-dependent biases that allows accurate shape characterisation of general and more subtle morphological variation. Here, we captured nuanced hip bone morphology revealing ontogenetic trends and sex-based variation in arcuate line curvature, greater sciatic notch shape, pubic body and rami length, acetabular expansion, and height-to-width proportions of the ilium. The observed ontogenetic trends showed a higher degree of bone modelling of the lesser pelvis of adolescent females, while male variation was mainly confined to the greater pelvis.

Morphological variation of the deciduous second molars in the Baka Pygmies.

The Baka Pygmies are known for their short stature resulting from a reduced growth rate during infancy. They are peculiar also for their teeth erupt earlier than in any other African population, and their posterior dentition is larger than in non-Pygmy populations. However, the Baka's dental morphology, like several other aspects of their biology, is still understudied. Here, we explore the variation of the Baka's deciduous upper and lower second molars (dm2s) in comparison to a geographically heterogeneous human sample by means of 3D geometric morphometrics and analysis of dental traits. Our results show that the different populations largely overlap based on the shape of their dm2s, especially the lower ones. Their distal region and the height of the dentinal crown differ the most, with the Baka showing the most extreme range of variation. Upper and lower dm2s covary to a great extent (RV = 0.82). The Baka's and South Americans' dm2s were confirmed among the largest in our sample. Despite the Baka's unique growth pattern, long-lasting isolation, and extreme dental variation, it is not possible to distinguish them from other populations based on their dm2s' morphology only.

The obstetrical dilemma hypothesis: there's life in the old dog yet.

The term 'obstetrical dilemma' was coined by Washburn in 1960 to describe the trade-off between selection for a larger birth canal, permitting successful passage of a big-brained human neonate, and the smaller pelvic dimensions required for bipedal locomotion. His suggested solution to these antagonistic pressures was to give birth prematurely, explaining the unusual degree of neurological and physical immaturity, or secondary altriciality, observed in human infants. This proposed trade-off has traditionally been offered as the predominant evolutionary explanation for why human childbirth is so challenging, and inherently risky, compared to that of other primates. This perceived difficulty is likely due to the tight fit of fetal to maternal pelvic dimensions along with the convoluted shape of the birth canal and a comparatively low degree of ligamentous flexibility. Although the ideas combined under the obstetrical dilemma hypothesis originated almost a century ago, they have received renewed attention and empirical scrutiny in the last decade, with some researchers advocating complete rejection of the hypothesis and its assumptions. However, the hypothesis is complex because it presently captures several, mutually non-exclusive ideas: (i) there is an evolutionary trade-off resulting from opposing selection pressures on the pelvis; (ii) selection favouring a narrow pelvis specifically derives from bipedalism; (iii) human neonates are secondarily altricial because they are born relatively immature to ensure that they fit through the maternal bony pelvis; (iv) as a corollary to the asymmetric selection pressure for a spacious birth canal in females, humans evolved pronounced sexual dimorphism of pelvic shape. Recently, the hypothesis has been challenged on both empirical and theoretical grounds. Here, we appraise the original ideas captured under the 'obstetrical dilemma' and their subsequent evolution. We also evaluate complementary and alternative explanations for a tight fetopelvic fit and obstructed labour, including ecological factors related to nutrition and thermoregulation, constraints imposed by the stability of the pelvic floor or by maternal and fetal metabolism, the energetics of bipedalism, and variability in pelvic shape. This reveals that human childbirth is affected by a complex combination of evolutionary, ecological, and biocultural factors, which variably constrain maternal pelvic form and fetal growth. Our review demonstrates that it is unwarranted to reject the obstetrical dilemma hypothesis entirely because several of its fundamental assumptions have not been successfully discounted despite claims to the contrary. As such, the obstetrical dilemma remains a tenable hypothesis that can be used productively to guide evolutionary research.

Sacrum morphology supports taxonomic heterogeneity of "Australopithecus africanus" at Sterkfontein Member 4.

The presence of multiple Australopithecus species at Sterkfontein Member 4, South Africa (2.07-2.61 Ma), is highly contentious, and quantitative assessments of craniodental and postcranial variability remain inconclusive. Using geometric morphometrics, we compared the sacrum of the small-bodied, presumed female subadult Australopithecus africanus skeleton Sts 14 to the large, alleged male adult StW 431 against a geographically diverse sample of modern humans, and two species of Pan, Gorilla, and Pongo. The probabilities of sampling morphologies as distinct as Sts 14 and StW 431 from a single species ranged from 1.3 to 2.5% for the human sample, and from 0.0 to 4.5% for the great apes, depending on the species and the analysis. Sexual dimorphism and developmental or geologic age could not adequately explain the differences between StW 431 and Sts 14, suggesting that they are unlikely to be conspecific. This supports earlier claims of taxonomic heterogeneity at Sterkfontein Member 4.

The dental remains from the Early Upper Paleolithic of Manot Cave, Israel.

This study presents the dental remains discovered at Manot Cave (MC), Western Galilee, Israel. The cave contains evidence for human occupation during the Early Upper Paleolithic period (46-33 ka) mainly of Early Ahmarian (∼46-42 ka) and Levantine Aurignacian (∼38-34 ka) cultural levels. Six teeth (three deciduous and three permanent) were found at the site, of which four could be thoroughly analyzed. The morphology of the teeth was qualitatively described and analyzed using traditional and geometric morphometric methods. A large comparative sample was used in order to assess the morphological affiliation of the Manot specimens with other Homo groups. The results provided equivocal signals: the upper first premolar (MC-9 P3) is probably modern human; the upper deciduous second molar (MC-10 dm2) and the upper second permanent molar (MC-8 M2) might be modern humans; the lower second deciduous molar (MC-7 dm2) might be Neanderthal. Owing to the small sample size and the almost total lack of distinctive characteristics, our outcome could not supply conclusive evidence to address the question of whether Manot Aurignacian population came from Europe or descended from the local Ahmarian population.

The partial skeleton StW 431 from Sterkfontein - Is it time to rethink the Plio-Pleistocene hominin diversity in South Africa?

The discovery of the nearly complete Plio-Pleistocene skeleton StW 573 Australopithecus prometheus from Sterkfontein Member 2, South Africa, has intensified debates as to whether Sterkfontein Member 4 contains a hominin species other than Australopithecus africanus. For example, it has recently been suggested that the partial skeleton StW 431 should be removed from the A. africanus hypodigm and be placed into A. prometheus. Here we re-evaluate this latter proposition, using published information and new comparative data. Although both StW 573 and StW 431 are apparently comparable in their arboreal (i.e., climbing) and bipedal adaptations, they also show significant morphological differences. Surprisingly, StW 431 cannot be unequivocally aligned with either StW 573 or other hominins from Sterkfontein commonly attributed to A. africanus (nor with Paranthropus robustus and Australopithecus sediba). This finding, together with considerations about the recent dating of Plio-Pleistocene hominin-bearing sites in South Africa and palaeoecological/palaeoclimatic conditions, raises questions whether it is justified to subsume hominins from Taung, Makapansgat and Sterkfontein (and Gladysvale) within a single taxon. Given the wealth of fossil material and analytical techniques now available, we call for a re-evaluation of the taxonomy of South African Plio-Pleistocene hominins. Such an endeavour should however go beyond the current (narrow) focus on establishing an A. africanus-A. prometheus dichotomy.

Accuracy of transferring analog dental casts to a virtual articulator.

STATEMENT OF PROBLEM: Complex digital workflows have been developed to create virtual dental patients. Direct and indirect digital methods are available for transferring analog patient information to virtual articulators. The direct method consists solely of digital workflows. The indirect method combines analog steps and digital procedures, representing an intermediate solution between the analog and direct digital approach. Studies that have investigated the overall accuracy of the virtual working space are sparse. PURPOSE: The purpose of this clinical study was to investigate the accuracy of the virtual dental space using the indirect digital workflow. MATERIAL AND METHODS: Mounted gypsum casts of 18 patients were used for indirect scanning. The maxillary casts were mounted in their skull-related position with a kinematic facebow. The mandibular casts were mounted in centric relation to the maxillary casts. The obtained digitized casts were transferred to a virtual articulator. An occlusal analysis was performed both in the analog and virtual environments, and the coordinates of matching analog and virtual contact points were measured. The trueness and precision of the indirect transferring procedure were assessed. RESULTS: A total of 194 analog points was considered in the reference. Ninety-three percent of all analog points matched a virtual correspondent, and 96% of the analog first contacts between the casts were also present as first contacts in the virtual space. The trueness of the data transfer, corresponding to the spatial distance between the matching analog and virtual points, was 0.55 ±0.31 mm. The maximum recorded deviation was 1.02 mm. CONCLUSIONS: The correspondence between the number and position of analog and virtual contacts was high. The mean absolute deviation of the matching point-pairs was better than that reported for the direct digital method. Under the conditions described, the virtual dental space created with the indirect digital method can be reliably used for virtual occlusal analysis in clinical practice.

Variation of 3D outer and inner crown morphology in modern human mandibular premolars.

OBJECTIVES: This study explores the outer and inner crown of lower third and fourth premolars (P3 , P4 ) by analyzing the morphological variation among diverse modern human groups. MATERIALS AND METHODS: We studied three-dimensional models of the outer enamel surface and the enamel-dentine junction (EDJ) from µCT datasets of 77 recent humans using both an assessment of seven nonmetric traits and a standard geometric morphometric (GM) analysis. For the latter, the dental crown was represented by four landmarks (dentine horns and fossae), 20 semilandmarks along the EDJ marginal ridge, and pseudolandmarks along the crown and cervical outlines. RESULTS: Certain discrete traits showed significantly different regional frequencies and sexual dimorphism. The GM analyses of both P3 s and P4 s showed extensive overlap in shape variation of the various populations (classification accuracy 15-69%). The first principal components explained about 40% of shape variance with a correlation between 0.59 and 0.87 of the features of P3 s and P4 s. Shape covariation between P3 s and P4 s expressed concordance of high and narrow or low and broad crowns. CONCLUSIONS: Due to marked intragroup and intergroup variation in GM analyses of lower premolars, discrete traits such as the number of lingual cusps and mesiolingual groove expression provide better geographic separation of modern human populations. The greater variability of the lingual region suggests a dominance of functional constraints over geographic provenience or sex. Additional information about functionally relevant aspects of the crown surface and odontogenetic data are needed to unravel the factors underlying dental morphology in modern humans.