The evolution of mammalian brain size.

Relative brain size has long been considered a reflection of cognitive capacities and has played a fundamental role in developing core theories in the life sciences. Yet, the notion that relative brain size validly represents selection on brain size relies on the untested assumptions that brain-body allometry is restrained to a stable scaling relationship across species and that any deviation from this slope is due to selection on brain size. Using the largest fossil and extant dataset yet assembled, we find that shifts in allometric slope underpin major transitions in mammalian evolution and are often primarily characterized by marked changes in body size. Our results reveal that the largest-brained mammals achieved large relative brain sizes by highly divergent paths. These findings prompt a reevaluation of the traditional paradigm of relative brain size and open new opportunities to improve our understanding of the genetic and developmental mechanisms that influence brain size.

The macroevolutionary consequences of phenotypic integration: from development to deep time.

Phenotypic integration is a pervasive characteristic of organisms. Numerous analyses have demonstrated that patterns of phenotypic integration are conserved across large clades, but that significant variation also exists. For example, heterochronic shifts related to different mammalian reproductive strategies are reflected in postcranial skeletal integration and in coordination of bone ossification. Phenotypic integration and modularity have been hypothesized to shape morphological evolution, and we extended simulations to confirm that trait integration can influence both the trajectory and magnitude of response to selection. We further demonstrate that phenotypic integration can produce both more and less disparate organisms than would be expected under random walk models by repartitioning variance in preferred directions. This effect can also be expected to favour homoplasy and convergent evolution. New empirical analyses of the carnivoran cranium show that rates of evolution, in contrast, are not strongly influenced by phenotypic integration and show little relationship to morphological disparity, suggesting that phenotypic integration may shape the direction of evolutionary change, but not necessarily the speed of it. Nonetheless, phenotypic integration is problematic for morphological clocks and should be incorporated more widely into models that seek to accurately reconstruct both trait and organismal evolution.

Three-dimensional shape variation of talar surface morphology in hominoid primates.

The hominoid foot is of particular interest to biological anthropologists, as changes in its anatomy through time reflect the adoption of terrestrial locomotion, particularly in species of Australopithecus and Homo. Understanding the osteological morphology associated with changes in whole foot function and the development of the plantar medial longitudinal foot arch are key to understanding the transition through habitual bipedalism in australopithecines to obligate bipedalism and long-distance running in Homo. The talus is ideal for studying relationships between morphology and function in this context, as it is a major contributor to the adduction-abduction, plantar-dorsal flexion and inversion-eversion of the foot, and transmits all forces encountered from the foot to the leg. The talar surface is predominantly covered by articular facets, which have different quantifiable morphological characters, including surface area, surface curvature and orientation. The talus also presents challenges to the investigator, as its globular shape is very difficult to quantify accurately and reproducibly. Here we apply a three-dimensional approach using type 3 landmarks (slid semilandmarks) that are geometrically homologous to determine overall talar shape variations in a range of living and fossil hominoid taxa. Additionally, we use novel approaches to quantify the relative orientations and curvatures of talar articular facets by determining the principal vectors of facet orientation and fitting spheres to articular facets. The resulting metrics are analysed using phylogenetic regressions and principal components analyses. Our results suggest that articular surface curvatures reflect locomotor specialisations with, in particular, orangutans having more highly curved facets in all but the calcaneal facet. Similarly, our approach to quantifying articular facet orientation appears to be effective in discriminating between extant hominoid species, and may therefore provide a sound basis for the study of fossil taxa and evolution of bipedalism in Australopithecus and Homo.

Brain reorganization, not relative brain size, primarily characterizes anthropoid brain evolution.

Comparative analyses of primate brain evolution have highlighted changes in size and internal organization as key factors underlying species diversity. It remains, however, unclear (i) how much variation in mosaic brain reorganization versus variation in relative brain size contributes to explaining the structural neural diversity observed across species, (ii) which mosaic changes contribute most to explaining diversity, and (iii) what the temporal origin, rates and processes are that underlie evolutionary shifts in mosaic reorganization for individual branches of the primate tree of life. We address these questions by combining novel comparative methods that allow assessing the temporal origin, rate and process of evolutionary changes on individual branches of the tree of life, with newly available data on volumes of key brain structures (prefrontal cortex, frontal motor areas and cerebrocerebellum) for a sample of 17 species (including humans). We identify patterns of mosaic change in brain evolution that mirror brain systems previously identified by electrophysiological and anatomical tract-tracing studies in non-human primates and functional connectivity MRI studies in humans. Across more than 40 Myr of anthropoid primate evolution, mosaic changes contribute more to explaining neural diversity than changes in relative brain size, and different mosaic patterns are differentially selected for when brains increase or decrease in size. We identify lineage-specific evolutionary specializations for all branches of the tree of life covered by our sample and demonstrate deep evolutionary roots for mosaic patterns associated with motor control and learning.

Calculating the axes of rotation for the subtalar and talocrural joints using 3D bone reconstructions.

Orientation of the subtalar joint axis dictates inversion and eversion movements of the foot and has been the focus of evolutionary and clinical studies for a number of years. Previous studies have measured the subtalar joint axis against the axis of the whole foot, the talocrural joint axis and, recently, the principal axes of the talus. The present study introduces a new method for estimating average joint axes from 3D reconstructions of bones and applies the method to the talus to calculate the subtalar and talocrural joint axes. The study also assesses the validity of the principal axes as a reference coordinate system against which to measure the subtalar joint axis. In order to define the angle of the subtalar joint axis relative to that of another axis in the talus, we suggest measuring the subtalar joint axis against the talocrural joint axis. We present corresponding 3D vector angles calculated from a modern human skeletal sample. This method is applicable to virtual 3D models acquired through surface-scanning of disarticulated 'dry' osteological samples, as well as to 3D models created from CT or MRI scans.

Allometric shape vector projection: a new method for the identification of allometric shape characters and trajectories applied to the human astragalus (talus).

The surface morphology of the human astragalus (talus) is difficult to represent accurately using landmarks as it is essentially globular in shape. Advances in laser scanning technology allow fast and accurate capture of bone surface morphology. However, methodologies to utilise these new accurate 3D data have not been fully developed. The present study uses canonical sampling of whole surface morphology attained through laser scanning and for the first time applies the technique to analysis of bone morphology. We introduce a new technique for identifying allometric shape characters in whole bone surface morphology. In a sample of adult human astragalus the new technique is successful in identifying and isolating intra-specific allometric shape characters in a bone which typically lacks landmarks and has, consequently, proved difficult to analyse using traditional 3D morphometric methods.

Nails and claws in primate evolution.

The issue of whether nails or claws were present on the digits of the last common ancestor of living primates is central to the understanding of the ecological context in which the order originated. Two lines of evidence are available, the shape (claw, nail, toilet-claw) and the histological structure (one or two horny strata). Here we review the existing data regarding the shape and histological structure of cheirideal appendages in primates and present new information from a wide range of living primates. We demonstrate the presence of a typical toilet-claw in Daubentonia madagascariensis and discuss its consequences, since the alleged lack of such structures in this species has long obscured the issue. The general view that primate nails, with the exception of those in New World primates, consist of only one layer is disproved by the presence of two distinct strata in the nails of the feet of three out of seven catarrhine species examined, as well as in Lemur catta. The combined new and old data indicate that the last common ancestor of the extant primates had lost the typical mammalian claws of its ancestors and developed nails on all pedal digits except digit II, which bore a toilet-claw. All nails as well as the toilet-claw originally consisted of two layers. We present a new hypothesis regarding the adaptational significance of these changes.

Identification of kinetochores and DNA synthesis in micronuclei induced by mitomycin C and colchicine in Chinese hamster ovary cells.

The presence of kinetochore and DNA synthesis in micronuclei (MN) induced in Chinese hamster ovary (CHO) cells by clastogenic and aneuploidogenic substances such as mitomycin C (MMC) and colchicine was determined by immunofluorescence technique using CREST antikinetochore antibodies and anti-bromodeoxyuridine (BrdUrd) antibodies. A cytofluorimetric analysis was also performed. Colchicine significantly increased micronucleated cells at least up to 96 h from the end of treatment. As expected, among colchicine-induced micronucleated cells the majority contained at least one CREST + MN. MMC induced a significant increase in micronucleated cells up to 120 h from the end of treatment and the great majority of MN lacked kinetochore fluorescence, indicating that MMC-induced MN were derived from acentric fragments. However, colchicine and MMC at 48 and 72 h from the end of treatment, induced a significant increase of CREST- and CREST + MN, respectively, suggesting an induction of clastogenicity by colchicine and aneuploidy by MMC. The clastogenic effect of colchicine after 48 h was also confirmed by the presence of chromatid fragments in metaphase cells. A cytofluorimetric analysis indicated that, as expected, colchicine and MMC interfere with the G2/M and S phases, respectively; however, a slight interference of colchicine with the S phase was also observed. DNA synthesis was present in MN and it was in most cases synchronous with synthesis in the main nucleus. The frequency of cells with MN in S phase observed in untreated or MMC-treated cells is in agreement with the proportion of cells without MN showing DNA synthesis. On the contrary, the frequency of cells with MN in S phase observed in colchicine-treated cells was significantly lower than that observed in control and MMC-treated cells.