Oldest pathology in a tetrapod bone illuminates the origin of terrestrial vertebrates

The origin of terrestrial tetrapods was a key event in vertebrate evolution, yet how and when it occurred remains obscure, due to scarce fossil evidence. Here, we show that the study of palaeopathologies, such as broken and healed bones, can help elucidate poorly understood behavioural transitions such as this. Using high-resolution finite element analysis, we demonstrate that the oldest known broken tetrapod bone, a radius of the primitive stem tetrapod Ossinodus pueri from the mid-Viséan (333 million years ago) of Australia, fractured under a high-force, impact-type loading scenario. The nature of the fracture suggests that it most plausibly occurred during a fall on land. Augmenting this are new osteological observations, including a preferred directionality to the trabecular architecture of cancellous bone. Together, these results suggest that Ossinodus, one of the first large (>2m length) tetrapods, spent a significant proportion of its life on land. Our findings have important implications for understanding the temporal, biogeographical and physiological contexts under which terrestriality in vertebrates evolved. They push the date for the origin of terrestrial tetrapods further back into the Carboniferous by at least two million years. Moreover, they raise the possibility that terrestriality in vertebrates first evolved in large tetrapods in Gondwana rather than in small European forms, warranting a re-evaluation of this important evolutionary event.

The discovery of Foxl2 paralogs in chondrichthyan, coelacanth and tetrapod genomes reveals an ancient duplication in vertebrates

The Foxl2 (forkhead box L2) gene is an important member of the forkhead domain family, primarily responsible for the development of ovaries during female sex differentiation. The evolutionary studies conducted previously considered the presence of paralog Foxl2 copies only in teleosts. However, to search for possible paralog copies in other groups of vertebrates and ensure that all predicted copies were homolog to the Foxl2 gene, a broad evolutionary analysis was performed, based on the forkhead domain family. A total of 2464 sequences for the forkhead domain were recovered, and subsequently, 64 representative sequences for Foxl2 were used in the evolutionary analysis of this gene. The most important contribution of this study was the discovery of a new subgroup of Foxl2 copies (ortholog to Foxl2B) present in the chondrichthyan Callorhinchus milii, in the coelacanth Latimeria chalumnae, in the avian Taeniopygia guttata and in the marsupial Monodelphis domestica. This new scenario indicates a gene duplication event in an ancestor of gnathostomes. Furthermore, based on the analysis of the syntenic regions of both Foxl2 copies, the duplication event was not exclusive to Foxl2. Moreover, the duplicated copy distribution was shown to be complex across vertebrates, especially in tetrapods, and the results strongly support a loss of this copy in eutherian species. Finally, the scenario observed in this study suggests an update for Foxl2 gene nomenclature, extending the actual suggested teleost naming of Foxl2A and Foxl2B to all vertebrate sequences and contributing to the establishment of a new evolutionary context for the Foxl2 gene. © 2013 Macmillan Publishers Limited All rights reserved.

Computer simulation of rod-coil block copolymers and tetrapod,polymer mixtures

In this thesis I present a new coarse-grained model suitable to investigate the phase behavior of rod-coil block copolymers on mesoscopic length scales. In this model the rods are represented by hard spherocylinders, whereas the coil block consists of interconnected beads. The interactions between the constituents are based on local densities. This facilitates an efficient Monte-Carlo sampling of the phase space. I verify the applicability of the model and the simulation approach by means of several examples. I treat pure rod systems and mixtures of rod and coil polymers. Then I append coils to the rods and investigate the role of the different model parameters. Furthermore, I compare different implementations of the model. I prove the capability of the rod-coil block copolymers in our model to exhibit typical micro-phase separated configurations as well as extraordinary phases, such as the wavy lamellar state, percolating structuresrnand clusters. Additionally, I demonstrate the metastability of the observed zigzag phase in our model. A central point of this thesis is the examination of the phase behavior of the rod-coil block copolymers in dependence of different chain lengths and interaction strengths between rods and coil. The observations of these studies are summarized in a phase diagram for rod-coil block copolymers. Furthermore, I validate a stabilization of the smectic phase with increasing coil fraction.rnIn the second part of this work I present a side project in which I derive a model permitting the simulation of tetrapods with and without grafted semiconducting block copolymers. The effect of these polymers is added in an implicit manner by effective interactions between the tetrapods. While the depletion interaction is described in an approximate manner within the Asakura-Oosawa model, the free energy penalty for the brush compression is calculated within the Alexander-de Gennes model. Recent experiments with CdSe tetrapods show that grafted tetrapods are clearly much better dispersed in the polymer matrix than bare tetrapods. My simulations confirm that bare tetrapods tend to aggregate in the matrix of excess polymers, while clustering is significantly reduced after grafting polymer chains to the tetrapods. Finally, I propose a possible extension enabling the simulation of a system with fluctuating volume and demonstrate its basic functionality. This study is originated in a cooperation with an experimental group with the goal to analyze the morphology of these systems in order to find the ideal morphology for hybrid solar cells.

Crustaceans and fish abundances and species at and around artificially introduced tetrapod fields in the southern North Sea, 2010-2011

In the coming decades, artificial defence structures will increase in importance worldwide for the protection of coasts against the impacts of global warming. However, the ecological effects of such structures on the natural surroundings remain unclear. We investigated the impact of experimentally introduced tetrapod fields on the demersal fish community in a hard-bottom area in the southern North Sea. The results indicated a significant decrease in fish abundance in the surrounding area caused by migration effects towards the artificial structures. Diversity (HB) and evenness (E) values exhibited greater variation after the introduction of the tetrapods. Additionally, a distinct increase in young-of-the-year (YOY) fish was observed near the structures within the second year after introduction. We suggest that the availability of adequate refuges in combination with additional food resources provided by the artificial structures has a highly species-specific attraction effect. However, these findings also demonstrate that our knowledge regarding the impact of artificial structures on temperate fish communities is still too limited to truly understand the ecological processes that are initiated by the introduction of artificial structures. Long-term investigations and additional experimental in situ work worldwide will be indispensable for a full understanding of the mechanisms by which coastal defence structures interact with the coastal environment.

Crustaceans and fish abundances and species at and around artificially introduced tetrapod fields in the southern North Sea, 2009

The micro-scale spatial distribution patterns of a demersal fish and decapod crustacean assemblage were assessed in a hard-bottom kelp environment in the southern North Sea. Using quadrats along line transects, we assessed the in situ fish and crustacean abundance in relation to substratum types (rock, cobbles and large pebbles) and the density of algae. Six fish and four crustacean species were abundant, with Ctenolabrus rupestris clearly dominating the fish community and Galathea squamifera dominating the crustacean community. Differences in the substratum types had an even stronger effect on the micro-scale distribution than the density of the dominating algae species. Kelp had a negative effect on the fish abundances, with significantly lower average densities in kelp beds compared with adjacent open areas. Averaged over all of the substrata, the most attractive substratum for the fish was large pebbles. In contrast, crustaceans did not show a specific substratum affinity. The results clearly indicate that, similar to other complex systems, significant micro-scale species-habitat associations occur in northern hard-bottom environments. However, because of the frequently harsh environmental conditions, these habitats are mainly sampled from ships with sampling gear, and the resulting data cannot be used to resolve small-scale species-habitat associations. A detailed substratum classification and community assessment, often only possible using SCUBA diving, is therefore important to reach a better understanding of the functional relationships between species and their environment in northern temperate waters, knowledge that is very important with respect to the increasing environmental pressure caused by global climate change.

Australian lungfish neurohypophysial hormone genes encode vasotocin and [Phe2]mesotocin precursors homologous to tetrapod-type precursors

In view of the well-established role of neurohypophysial hormones in osmoregulation of terrestrial vertebrates, lungfishes are a key group for study of the molecular and functional evolution of the hypothalamo-neurohypophysial system. Here we report on the primary structure of the precursors encoding vasotocin (VT) and [Phe2]mesotocin ([Phe2]MT) of the Australian lungfish, Neoceratodus forsteri. Genomic sequence analysis and Northern blot analysis confirmed that [Phe2]MT is a native oxytocin family peptide in the Australian lungfish, although it has been reported that the lungfish neurohypophysis contains MT. The VT precursor consists of a signal peptide, VT, that is connected to a neurophysin by a Gly-Lys-Arg sequence, and a copeptin moiety that includes a Leu-rich core segment and a glycosylation site. In contrast, the [Phe2]MT precursor does not contain a copeptin moiety. These structural features of the lungfish precursors are consistent with those in tetrapods, but different from those in teleosts where both VT and isotocin precursors contain a copeptin-like moiety without a glycosylation site at the carboxyl terminals of their neurophysins. Comparison of the exon/intron organization also supports homology of the lungfish [Phe2]MT gene with tetrapod oxytocin/MT genes, rather than with teleost isotocin genes. Moreover, molecular phylogenetic analysis shows that neurohypophysial hormone genes of the lungfish are closely related to those of the toad. The present results along with previous morphological findings indicate that the hypothalamo-neurohypophysial system of the lungfish has evolved along the tetrapod lineage, whereas the teleosts form a separate lineage, both within the class Osteichthyes.

Early steps of paired fin development in zebrafish compared with tetrapod limb development.

The development of zebrafish paired fins and tetrapod forelimbs and hindlimbs show striking similarities at the molecular level. In recent years, the zebrafish, Danio rerio has become a valuable model for the study of the development of vertebrate paired appendages and several large-scale mutagenesis screens have identified novel fin mutants. This review summarizes recent advances in research into zebrafish paired fin development and highlights features that are shared with and distinct from limb development in other main animal models.

First Triassic tetrapod (Sauropterygia, Nothosauridae) from Castilla y León: evidence of an unknown taxon for the Spanish record

Several vertebrae of a sauropterygian specimen have been recovered in Fuencaliente de Medinaceli (Soria Province, Castilla y León, Spain). The remains come from Middle–Upper Triassic Muschelkalk Facies. This finding represents the first documented evidence of a Triassic tetrapod in Castilla y León. The vertebrae belong to Nothosaurus, a sauropterygian genus found in Europe, Middle East, North of Africa and China. This genus is poorly-known in the Iberian record. The new remains constitute the first evidence of the species Nothosaurus giganteus, or a related taxon, in the Iberian Peninsula. This study reveals the occurrence of at least two species of the sauropterygian Nothosaurus in the Spanish record.

Avian diversification patterns across the K-Pg boundary: Influence of calibrations, datasets, and model misspecification

Birds represent the most diverse extant tetrapod clade, with ca. 10,000 extant species, and the timing of the crown avian radiation remains hotly debated. The fossil record supports a primarily Cenozoic radiation of crown birds, whereas molecular divergence dating analyses generally imply that this radiation was well underway during the Cretaceous. Furthermore, substantial differences have been noted between published divergence estimates. These have been variously attributed to clock model, calibration regime, and gene type. One underappreciated phenomenon is that disparity between fossil ages and molecular dates tends to be proportionally greater for shallower nodes in the avian Tree of Life. Here, we explore potential drivers of disparity in avian divergence dates through a set of analyses applying various calibration strategies and coding methods to a mitochondrial genome dataset and an 18-gene nuclear dataset, both sampled across 72 taxa. Our analyses support the occurrence of two deep divergences (i.e., the Palaeognathae/Neognathae split and the Galloanserae/Neoaves split) well within the Cretaceous, followed by a rapid radiation of Neoaves near the K-Pg boundary. However, 95% highest posterior density intervals for most basal divergences in Neoaves cross the boundary, and we emphasize that, barring unreasonably strict prior distributions, distinguishing between a rapid Early Paleocene radiation and a Late Cretaceous radiation may be beyond the resolving power of currently favored divergence dating methods. In contrast to recent observations for placental mammals, constraining all divergences within Neoaves to occur in the Cenozoic does not result in unreasonably high inferred substitution rates. Comparisons of nuclear DNA (nDNA) versus mitochondrial DNA (mtDNA) datasets and NT- versus RY-coded mitochondrial data reveal patterns of disparity that are consistent with substitution model misspecifications that result in tree compression/tree extension artifacts, which may explain some discordance between previous divergence estimates based on different sequence types. Comparisons of fully calibrated and nominally calibrated trees support a correlation between body mass and apparent dating error. Overall, our results are consistent with (but do not require) a Paleogene radiation for most major clades of crown birds.

Shape Effect on Electronic and Photovoltaic Properties of CdS Nanocrystals

Changes in electronic and photovoltaic properties of semiconductor nanocrystals predominantly due to changes in shape are discussed here. Cadmium sulfide (CdS) semiconductor nanocrystals of various shapes (tetrapod, tetrahedron, sphere and rod) obtained using an optimized solvothermal process exhibited a mixed cubic (zinc blende) and hexagonal (wurtzite) crystal structure. The simultaneous presence of the two crystal phases in varying amounts is observed to play a pivotal role in determining both the electronic and photovoltaic properties of the CdS nanocrystals. Light to electrical energy conversion efficiencies (measured in two-electrode configuration laboratory solar cells) remarkably decreased by one order in magnitude from tetrapod -> tetrahedron -> sphere -> rod. The tetrapod-CdS nanocrystals, which displayed the highest light to electrical energy conversion efficiency, showed a favorable shift in position of the conduction band edge leading to highest rate of electron injection (from CdS nanocrystal to the wide band gap semiconductor viz, titanium dioxide, TiO2) and lowest rate of electron-hole recombination (higher free electron lifetimes).

A ecologia em miniatura: o menor tetrápodo do Brasil, Brachycephalus didactylus (Izecksohn, 1971) (Anura: Brachycephalidae) em uma área de Mata Atlântica no Espírito Santo, Sudeste do Brasil

O processo de miniaturização em anuros pode ter implicações ecológicas como, por exemplo, a redução no tamanho da ninhada produzida pelas fêmeas, o aumento do tamanho relativo dos ovos e o desenvolvimento direto. O consequente reduzido tamanho da boca tende também a constituir um fator limitante no consumo de presas por estes pequenos anuros, tornando acessível ao consumo apenas uma gama restrita de presas diminutas. Assim, estudamos parâmetros da ecologia de uma população do anuro Brachycephalus didactylus, em uma área de Mata Atlântica no sul do Espírito Santo, objetivando ampliar o conhecimento sobre tais parâmetros que compreenderam o dimorfismo sexual, algumas características da reprodução, a dieta e a relação com a disponibilidade de atrópodos no folhiço e uso do microhabitat. A amostragem dos dados ocorreu durante um ano, nas estações seca e chuvosa. Brachycephalus didactylus possuiu dimorfismo sexual, com fêmeas sendo maiores do que os machos. O maior tamanho atingido pelas fêmeas provavelmente resulta da seleção intrassexual atuando nas fêmeas para favorecer um maior investimento na massa dos ovos, já que a espécie possui um tamanho de ninhada consideravelmente reduzido (um a dois ovos). A dieta de B. didactylus foi composta por um espectro relativamente amplo de presas (17 itens), tendo elevadas proporções de pequenas presas como Acari e Collembola. O consumo de Formicidae (formigas) foi evitado por B. didactylus, apesar de essa presa ser a mais abundante no folhiço, o que categorizou o anuro como um "especialista em não comer formigas" ("non-ant specialist", sensu Toft 1980a). O reduzido tamanho da boca nas fêmeas de B. didactylus limitou o tamanho máximo de presas passíveis de serem ingeridas pelo anuro e restringindo-o ao consumo de presas pequenas, com maiores indivíduos consumindo um número maior de presas. Por fim, B. didactylus apresentou uma utilização essencialmente horizontal do hábitat, sendo mais frequentemente encontrado sobre o folhiço no chão da mata.

A graphene-based large area surface-conduction electron emission display

The fabrication and functionality of a 21 cm graphene-based transverse electron emission display panel is presented. A screen-printed triode edge electron emission geometry has been developed based on chemical vapor deposited (CVD) graphene supported on vertically aligned carbon nanotubes (CNT) necessary to minimize electrostatic shielding induced by the proximal bulk substrate. Integrated ZnO tetrapod electron scatterers have been shown to increase the emission efficiency by more than 90%. Simulated electron trajectories validate the observed emission characteristics with driving voltages less than 60 V. Fabricated display panels have shown real-time video capabilities that are hysteresis free (<0.2%), have extremely stable lifetimes (<3% variation over 10 h continuous operation) in addition to rapid temporal responses (<1 ms). © 2013 Elsevier Ltd. All rights reserved.

Pressure behaviour of the UV and green emission bands in ZnO micro-rods

The pressure behavior of the ultraviolet (UV) and green emission bands in ZnO tetrapod-like micro-rods has been investigated at 300 and 70 K, respectively. The pressure coefficient of the UV band at 300 K is 24.5 meV/GPa, consistent with that of the band gap of bulk ZnO. However, the pressure coefficient of the green band is 25 meV/GPa, far larger than previous literature reports. The green band in this work originates from Cu-related emission, as confirmed by the fine structure observed in the spectra at 10 K. The pressure coefficients of four phonon replicas of the free exciton emission (FX) at 70 K are 21.0, 20.2, 19.8, and 19.3 meV/GPa, respectively. The energy shift rate of the FX emission and the LO phonon energies is then determined to be 21.4 and 0.55 meV/GPa. The pressure coefficient of the neutral donor bound exciton ((DX)-X-0) transition is 20.5 meV/GPa, only 4% smaller than that of FX. This confirms that the (DX)-X-0 emission corresponds to excitons bound to neutral shallow donors. (C) 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.

Fabrication of Novel Superhydrophobic Surfaces and Water Droplet Bouncing Behavior - Part 1: Stable ZnO-PDMS Superhydrophobic Surface with Low Hysteresis Constructed Using ZnO Nanoparticles

A superhydrophobic surface has many advantages in micro/nanomechanical applications, such as low adhesion, low friction and high restitution coefficient, etc. In this paper, we introduce a novel and simple route to fabricate superhydrophobic surfaces using ZnO nanocrystals. First, tetrapod-like ZnO nanocrystals were prepared via a one-step, direct chemical vapor deposition (CVD) approach. The nanostructured ZnO material was characterized by scanning electron microscope (SEM) and X-ray diffraction (XRD) and the surface functionalized by aminopropyltriethoxysilane (APS) was found to be hydrophobic. Then the superhydrophobic surface was constructed by depositing uniformly ZnO hydrophobic nanoparticles (HNPs) on the Poly(dimethylsiloxane) (PDMS) film substrate. Water wettability study revealed a contact angle of 155.4 +/- 2 degrees for the superhydrophobic surface while about 110 degrees for pure smooth PDMS films. The hysteresis was quite low, only 3.1 +/- 0.3 degrees. Microscopic observations showed that the surface was covered by micro- and nano-scale ZnO particles. Compared to other approaches, this method is rather convenient and can be used to obtain a large area superhydrophobic surface. The high contact angle and low hysteresis could be attributed to the micro/nano structures of ZnO material; besides, the superhydrophobic property of the as-constructed ZnO-PDMS surface could be maintained for at least 6 months. (C) Koninklijke Brill NV, Leiden, 2010