Unraveling the sperm bauplan: Relationships between sperm head morphology and sperm function in rodents

This is the author accepted manuscript. The final version is available from the Society for the Study of Reproduction via the DOI in this record.Rodents have spermatozoa with features not seen in other species. Sperm heads in many rodent species bear one or more apical extensions known as "hooks." The process by which hooks have evolved, together with their adaptive significance, are still controversial issues. In order to improve our understanding of the biological meaning of these sperm head adaptations, we analyzed hook curvature angles, hook length, and overall hook shape in muroid rodents by using geometric morphometrics. We also searched for relationships between hook design and measurements of intermale competition to assess whether postcopulatory sexual selection was an important selective force driving changes in this sperm structure. Finally, we sought possible links between aspects of sperm hook design and sperm velocity as a measure of sperm performance. Results showed that one hook curvature angle is under strong selective pressure. Similarly, hook length appears to be strongly selected by sexual selection, with this selective force also exhibiting a stabilizing role reducing intermale variation in this trait. The adaptive significance of changes in hook structure was supported by the finding that there are strong and significant covariations between hook dimensions and shape and between hook design and sperm swimming velocity. Overall, this study strongly suggests that postcopulatory sexual selection has an important effect on the design of the sperm head that, in turn, is important for enhancing sperm velocity, a function crucial to reaching the vicinity of the female gamete and winning fertilizations under competitive situations

A hominin first rib discovered at the Sterkfontein Caves, South Africa.

First ribs - the first or most superior ribs in the thorax - are rare in the hominin fossil record, and when found, have the potential to provide information regarding the upper thorax shape of extinct hominins. Here, we describe a partial first rib from Member 4 of the Sterkfontein Caves, South Africa. The rib shaft is broken away, so only the head and neck are preserved. The rib is small, falling closest to small-bodied Australopithecus first ribs (AL 288-1 and MH1). Given that it was recovered near the StW 318 femur excavation, which also represents a small individual, we suggest that the two may be associated. Three-dimensional geometric morphometric analyses were used to quantify the rib fragment morphology and compare it to extant hominoid and other fossil hominin ribs. While only the proximal end is preserved, our analyses show that South African Australopithecus share derived features of the proximal first rib more closely resembling A. afarensis and later hominins than great apes.NCS2016

Three-dimensional geometric morphometrics of thorax-pelvis covariation and its potential for predicting the thorax morphology: A case study on Kebara 2 Neandertal

The skeletal torso is a complex structure of outstanding importance in understanding human body shape evolution, but reconstruction usually entails an element of subjectivity as researchers apply their own anatomical expertise to the process. Among different fossil reconstruction methods, 3D geometric morphometric techniques have been increasingly used in the last decades. Two-block partial least squares analysis has shown great potential for predicting missing elements by exploiting the covariation between two structures (blocks) in a reference sample: one block can be predicted from the other one based on the strength of covariation between blocks. The first aim of this study is to test whether this predictive approach can be used for predicting thorax morphologies from pelvis morphologies within adult Homo sapiens reference samples with known covariation between the thorax and the pelvis. The second aim is to apply this method to Kebara 2 Neandertal (Israel, ∼60 ka) to predict its thorax morphology using two different pelvis reconstructions as predictors. We measured 134 true landmarks, 720 curve semilandmarks, and 160 surface semilandmarks on 60 3D virtual torso models segmented from CT scans. We conducted three two-block partial least squares analyses between the thorax (block 1) and the pelvis (block 2) based on the H. sapiens reference samples after performing generalized Procrustes superimposition on each block separately. Comparisons of these predictions in full shape space by means of Procrustes distances show that the male-only predictive model yields the most reliable predictions within modern humans. In addition, Kebara 2 thorax predictions based on this model concur with the thorax morphology proposed for Neandertals. The method presented here does not aim to replace other techniques, but to rather complement them through quantitative prediction of a virtual 'scaffold' to articulate the thoracic fossil elements, thus extending the potential of missing data estimation beyond the methods proposed in previous works

Covariation between the shape and mineralized tissues of the rib cross section in Homo sapiens, Pan troglodytes and Sts 14

Objectives: Studying rib torsion is crucial for understanding the evolution of the hominid ribcage. Interestingly, there are variables of the rib cross section that could be associated with rib torsion and, consequently, with the morphology of the thorax. The aim of this research is to conduct a comparative study of the shape and mineralized tissues of the rib cross section in different hominids to test for significant differences and, if possible, associate them to different thoracic morphotypes.Materials and Methods: The sample consists of the rib cross sections at the midshaft taken from 10 Homo sapiens and 10 Pan troglodytes adult individuals, as well as from A. africanus Sts 14. The shape of these rib cross sections was quantified using geometric morphometrics, while the mineralized tissues were evaluated using the compartmentalization index. Subsequently, covariation between both parameters was tested by a Spearman's ? test, a permutation test and a linear regression.Results: Generally, P. troglodytes individuals exhibit rib cross sections that are rounder and more mineralized compared to those of H. sapiens. However, the covariation between both parameters was only observed in typical ribs (levels 3-10). Although covariation was not found in the rib cross sections of Sts 14, their parameters are closer to P. troglodytes.Discussion: On the one hand, the differences observed in the rib cross sections between H. sapiens and P. troglodytes might be related to different degrees of rib torsion and, consequently, to different thoracic 3D configurations. These findings can be functionally explained by considering their distinct modes of breathing and locomotion. On the other hand, although the rib cross sections belonging to Sts 14 are more similar to those of P. troglodytes, previous publications determined that their overall morphology is closer to modern humans. This discrepancy could reflect a diversity of post-cranial adaptations in Australopithecus

Preconditioned iterative methods for convection diffusion and related boundary value problems

AbstractWe develop and analyze preconditioners for the iterative solution of the system of equations arising from the discretization of multi-dimensional singularity perturbed boundary value problems. This includes a class of convection diffusion models. The choice of preconditioner is crucial for the efficient solution of the system of equations. In particular, it is necessary to choose a preconditioner that substantially reduces the condition number κ both for small grid size h and for large values of the parameter K multiplying the convection terms. A class of preconditioners is analyzed that is inexpensive to implement and for which κ = 0(1) as h→0 and κ = (1 + K12) as K → ∞ for some convection diffusion problems with positive definite symmetric part. This result is then used to develop an algorithm with work estimate 0(1 + K12as K → ∞ for a more general class of convection diffusion problems including those with indefinite symmetric part. Numerical experiments using a symmetric multigrid preconditioner demonstrate the effectiveness of the numerical method even for large problems

Assessing thoraco‐pelvic covariation in Homo sapiens and Pan troglodytes: A 3D geometric morphometric approach

Objectives Understanding thoraco‐pelvic integration in Homo sapiens and their closest living relatives (genus Pan) is of great importance within the context of human body shape evolution. However, studies assessing thoraco‐pelvic covariation across Hominoidea species are scarce, although recent research would suggest shared covariation patterns in humans and chimpanzees but also species‐specific features, with sexual dimorphism and allometry influencing thoraco‐pelvic covariation in these taxa differently. Material and Methods N = 30 adult H. sapiens and N = 10 adult Pan troglodytes torso 3D models were analyzed using 3D geometric morphometrics and linear measurements. Effects of sexual dimorphism and allometry on thoraco‐pelvic covariation were assessed via regression analyses, and patterns of thoraco‐pelvic covariation in humans and chimpanzees were computed via Two‐Block Partial Least Squares analyses. Results Results confirm the existence of common aspects of thoraco‐pelvic covariation in humans and chimpanzees, and also species‐specific covariation in H. sapiens that is strongly influenced by sexual dimorphism and allometry. Species‐specific covariation patterns in chimpanzees could not be confirmed because of the small sample size, but metrics point to a correspondence between the most caudal ribs and iliac crest morphology that would be irrespective of sex. Conclusions This study suggests that humans and chimpanzees share common aspects of thoraco‐pelvic covariation but might differ in others. In humans, torso integration is strongly influenced by sexual dimorphism and allometry, whilst in chimpanzees it may not be. This study also highlights the importance not only of torso widths but also of torso depths when describing patterns of thoraco‐pelvic covariation in primates. Larger samples are necessary to support these interpretations

Anatomical Network Comparison of Human Upper and Lower, Newborn and Adult, and Normal and Abnormal Limbs, with Notes on Development, Pathology and Limb Serial Homology vs. Homoplasy

How do the various anatomical parts (modules) of the animal body evolve into very different integrated forms (integration) yet still function properly without decreasing the individual's survival? This long-standing question remains unanswered for multiple reasons, including lack of consensus about conceptual definitions and approaches, as well as a reasonable bias toward the study of hard tissues over soft tissues. A major difficulty concerns the non-trivial technical hurdles of addressing this problem, specifically the lack of quantitative tools to quantify and compare variation across multiple disparate anatomical parts and tissue types. In this paper we apply for the first time a powerful new quantitative tool, Anatomical Network Analysis (AnNA), to examine and compare in detail the musculoskeletal modularity and integration of normal and abnormal human upper and lower limbs. In contrast to other morphological methods, the strength of AnNA is that it allows efficient and direct empirical comparisons among body parts with even vastly different architectures (e.g. upper and lower limbs) and diverse or complex tissue composition (e.g. bones, cartilages and muscles), by quantifying the spatial organization of these parts-their topological patterns relative to each other-using tools borrowed from network theory. Our results reveal similarities between the skeletal networks of the normal newborn/adult upper limb vs. lower limb, with exception to the shoulder vs. pelvis. However, when muscles are included, the overall musculoskeletal network organization of the upper limb is strikingly different from that of the lower limb, particularly that of the more proximal structures of each limb. Importantly, the obtained data provide further evidence to be added to the vast amount of paleontological, gross anatomical, developmental, molecular and embryological data recently obtained that contradicts the long-standing dogma that the upper and lower limbs are serial homologues. In addition, the AnNA of the limbs of a trisomy 18 human fetus strongly supports Pere Alberch's ill-named "logic of monsters" hypothesis, and contradicts the commonly accepted idea that birth defects often lead to lower integration (i.e. more parcellation) of anatomical structures

Developmental pathways inferred from modularity, morphological integration and fluctuating asymmetry patterns in the human face

Facial asymmetries are usually measured and interpreted as proxies to developmental noise. However, analyses focused on its developmental and genetic architecture are scarce. To advance on this topic, studies based on a comprehensive and simultaneous analysis of modularity, morphological integration and facial asymmetries including both phenotypic and genomic information are needed. Here we explore several modularity hypotheses on a sample of Latin American mestizos, in order to test if modularity and integration patterns difer across several genomic ancestry backgrounds. To do so, 4104 individuals were analyzed using 3D photogrammetry reconstructions and a set of 34 facial landmarks placed on each individual. We found a pattern of modularity and integration that is conserved across sub-samples difering in their genomic ancestry background. Specifcally, a signal of modularity based on functional demands and organization of the face is regularly observed across the whole sample. Our results shed more light on previous evidence obtained from Genome Wide Association Studies performed on the same samples, indicating the action of diferent genomic regions contributing to the expression of the nose and mouth facial phenotypes. Our results also indicate that large samples including phenotypic and genomic metadata enable a better understanding of the developmental and genetic architecture of craniofacial phenotypes

Primate modularity and evolution: first anatomical network analysis of primate head and neck musculoskeletal system

Network theory is increasingly being used to study morphological modularity and integration. Anatomical network analysis (AnNA) is a framework for quantitatively characterizing the topological organization of anatomical structures and providing an operational way to compare structural integration and modularity. Here we apply AnNA for the first time to study the macroevolution of the musculoskeletal system of the head and neck in primates and their closest living relatives, paying special attention to the evolution of structures associated with facial and vocal communication. We show that well-defined left and right facial modules are plesiomorphic for primates, while anthropoids consistently have asymmetrical facial modules that include structures of both sides, a change likely related to the ability to display more complex, asymmetrical facial expressions. However, no clear trends in network organization were found regarding the evolution of structures related to speech. Remarkably, the increase in the number of head and neck muscles – and thus of musculoskeletal structures – in human evolution led to a decrease in network density and complexity in humans

Response to Comment on "The growth pattern of Neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain)".

This is the author’s version of the work. It is posted here by permission of the AAAS for personal use, not for redistribution. The definitive version was published in Rosas, A., et al. (2018). "Response to Comment on “The growth pattern of Neandertals, reconstructed from a juvenile skeleton from El Sidrón (Spain)”." Science 359(6380). on Science 09 Mar 2018: Vol. 359, Issue 6380, eaar3820, DOI: 10.1126/science.aar3820The comment by DeSilva challenges our suggestion that brain growth of the El Sidrón J1 Neandertal was still incomplete at 7.7 years of age. Evidence suggests that endocranial volume is likely to represent less than 90% adult size at El Sidrón as well as Neandertal male plus Krapina samples, in line with further evidence from endocranial surface histology and dural sinus groove size