Deep-sea origin and depth colonization associated with phenotypic innovations in scleractinian corals.

The deep sea (>200 m) is home to a surprisingly rich biota, which in some cases compares to that found in shallow areas. Scleractinian corals are an example of this - they are key species in both shallow and deep ecosystems. However, what evolutionary processes resulted in current depth distribution of the marine fauna is a long-standing question. Various conflicting hypotheses have been proposed, but few formal tests have been conducted. Here, we use global spatial distribution data to test the bathymetric origin and colonization trends across the depth gradient in scleractinian corals. Using a phylogenetic approach, we infer the origin and historical trends in directionality and speed of colonization during the diversification in depth. We also examine how the emergence of photo-symbiosis and coloniality, scleractinian corals' most conspicuous phenotypic innovations, have influenced this process. Our results strongly support an offshore-onshore pattern of evolution and varying dispersion capacities along depth associated with trait-defined lineages. These results highlight the relevance of the evolutionary processes occurring at different depths to explain the origin of extant marine biodiversity and the consequences of altering these processes by human impact, highlighting the need to include this overlooked evolutionary history in conservation plans.

Ancestral State Reconstruction Using BayesTraits.

The fossil record is the best evidence of the characteristics of extinct species, but only a narrow range of traits fossilize or survive the fossilization process. Lacking fossil or other evidence about the past, ancestral states can be reconstructed. Three pieces of information are combined when reconstructing ancestral states: extant or known trait values (data); the evolutionary history, linking the species of interest (phylogeny); and the evolutionary model of trait change. These reconstructed ancestral states can be interpreted as our best guess as to the route evolution took, given the distribution of the trait across species, the relationships among them, and our model of evolution. Because the information we use to reconstruct the past is often not known without error, uncertainty about their true values should be accounted for when reconstructing ancestral states. In this chapter we describe how ancestral states can be reconstructed using a Bayesian framework implemented in the software BayesTraits to account for uncertainty in the phylogenetic tree and the model of evolution.

General statistical model shows that macroevolutionary patterns and processes are consistent with Darwinian gradualism.

Macroevolution posed difficulties for Darwin and later theorists because species' phenotypes frequently change abruptly, or experience long periods of stasis, both counter to the theory of incremental change or gradualism. We introduce a statistical model that accommodates this uneven evolutionary landscape by estimating two kinds of historical change: directional changes that shift the mean phenotype along the branches of a phylogenetic tree, and evolvability changes that alter a clade's ability to explore its trait-space. In mammals, we find that both processes make substantial independent contributions to explaining macroevolution, and are rarely linked. 'Watershed' moments of increased evolvability greatly outnumber reductions in evolutionary potentials, and large or abrupt phenotypic shifts are explicable statistically as biased random walks, allowing macroevolutionary theory to engage with the language and concepts of gradualist microevolution. Our findings recast macroevolutionary phenomena, illustrating the necessity of accounting for a variety of evolutionary processes simultaneously.

Genes underlying the evolution of tetrapod testes size.

BACKGROUND: Testes vary widely in mass relative to body mass across species, but we know very little about which genes underlie and contribute to such variation. This is partly because evidence for which genes are implicated in testis size variation tends to come from investigations involving just one or a few species. Contemporary comparative phylogenetic methods provide an opportunity to test candidate genes for their role in phenotypic change at a macro-evolutionary scale-across species and over millions of years. Previous attempts to detect genotype-phenotype associations across species have been limited in that they can only detect where genes have driven directional selection (e.g. brain size increase). RESULTS: Here, we introduce an approach that uses rates of evolutionary change to overcome this limitation to test whether any of twelve candidate genes have driven testis size evolution across tetrapod vertebrates-regardless of directionality. We do this by seeking a relationship between the rates of genetic and phenotypic evolution. Our results reveal five genes (Alkbh5, Dmrtb1, Pld6, Nlrp3, Sp4) that each have played unique and complex roles in tetrapod testis size diversity. In all five genes, we find strong significant associations between the rate of protein-coding substitutions and the rate of testis size evolution. Such an association has never, to our knowledge, been tested before for any gene or phenotype. CONCLUSIONS: We describe a new approach to tackle one of the most fundamental questions in biology: how do individual genes give rise to biological diversity? The ability to detect genotype-phenotype associations that have acted across species has the potential to build a picture of how natural selection has sculpted phenotypic change over millions of years.

Temperature thresholds of ecosystem respiration at a global scale.

Ecosystem respiration is a major component of the global terrestrial carbon cycle and is strongly influenced by temperature. The global extent of the temperature-ecosystem respiration relationship, however, has not been fully explored. Here, we test linear and threshold models of ecosystem respiration across 210 globally distributed eddy covariance sites over an extensive temperature range. We find thresholds to the global temperature-ecosystem respiration relationship at high and low air temperatures and mid soil temperatures, which represent transitions in the temperature dependence and sensitivity of ecosystem respiration. Annual ecosystem respiration rates show a markedly reduced temperature dependence and sensitivity compared to half-hourly rates, and a single mid-temperature threshold for both air and soil temperature. Our study indicates a distinction in the influence of environmental factors, including temperature, on ecosystem respiration between latitudinal and climate gradients at short (half-hourly) and long (annual) timescales. Such climatological differences in the temperature sensitivity of ecosystem respiration have important consequences for the terrestrial net carbon sink under ongoing climate change.

Nothing Wrong with the Analysis of Clades in Comparative Evolutionary Studies: A Reply to Poe et al.

In a recent paper, Poe et al. assert that scientists should abandon clade-based approaches, particularly those using named taxonomic ranks. Poe et al. attempt to demonstrate that clade selection can have effects on the results of evolutionary analyses but unfortunately fall short of making any robust conclusions. Here, we demonstrate that the assertions made by Poe et al. have two important flaws: (i) an erroneous view of modern phylogenetic comparative methods; and (ii) a lack of statistical rigor in their analyses. We repeat Poe et al.'s analysis but using appropriate phylogenetic comparative approaches. We demonstrate that results remain consistent regardless of the clade definition. We go on to discuss the value of taxonomic groupings and how they can provide meaningful units of comparison in evolutionary study. Unlike the disheartening suggestion to abandon the use of clades, scientists can instead continue to use phylogenetic " corrections" that are already the standard for most comparative evolutionary analyses. [Comparative methods; evolution; phylogeny; taxonomy.].

150 million years of sustained increase in pterosaur flight efficiency.

The long-term accumulation of biodiversity has been punctuated by remarkable evolutionary transitions that allowed organisms to exploit new ecological opportunities. Mesozoic flying reptiles (the pterosaurs), which dominated the skies for more than 150 million years, were the product of one such transition. The ancestors of pterosaurs were small and probably bipedal early archosaurs1, which were certainly well-adapted to terrestrial locomotion. Pterosaurs diverged from dinosaur ancestors in the Early Triassic epoch (around 245 million years ago); however, the first fossils of pterosaurs are dated to 25 million years later, in the Late Triassic epoch. Therefore, in the absence of proto-pterosaur fossils, it is difficult to study how flight first evolved in this group. Here we describe the evolutionary dynamics of the adaptation of pterosaurs to a new method of locomotion. The earliest known pterosaurs took flight and subsequently appear to have become capable and efficient flyers. However, it seems clear that transitioning between forms of locomotion2,3-from terrestrial to volant-challenged early pterosaurs by imposing a high energetic burden, thus requiring flight to provide some offsetting fitness benefits. Using phylogenetic statistical methods and biophysical models combined with information from the fossil record, we detect an evolutionary signal of natural selection that acted to increase flight efficiency over millions of years. Our results show that there was still considerable room for improvement in terms of efficiency after the appearance of flight. However, in the Azhdarchoidea4, a clade that exhibits gigantism, we test the hypothesis that there was a decreased reliance on flight5-7 and find evidence for reduced selection on flight efficiency in this clade. Our approach offers a blueprint to objectively study functional and energetic changes through geological time at a more nuanced level than has previously been possible.

Evolutionary Timeline and Genomic Plasticity Underlying the Lifestyle Diversity in Rhizobiales.

Members of the order Rhizobiales include those capable of nitrogen fixation in nodules as well as pathogens of animals and plants. This lifestyle diversity has important implications for agricultural and medical research. Leveraging large-scale genomic data, we infer that Rhizobiales originated as a free-living ancestor ∼1,500 million years ago (Mya) and that the later emergence of host-associated lifestyles broadly coincided with the rise of their eukaryotic hosts. In particular, the first nodulating lineage arose from either Azorhizobium or Bradyrhizobium 150 to 80 Mya, a time range in general concurrent with the emergence of legumes. The rates of lifestyle transitions are highly variable; nodule association is more likely to be lost than gained, whereas animal association likely represents an evolutionary dead end. We searched for statistical correlations between gene presence and lifestyle and identified genes likely contributing to the transition and adaptation to the same lifestyle in divergent lineages. Among the genes potentially promoting successful transitions to major nodulation lineages, the nod and nif clusters for nodulation and nitrogen fixation, respectively, were repeatedly acquired during each transition; the fix, dct, and phb clusters involved in energy conservation under micro-oxic conditions were present in the nonnodulating ancestors; and the secretion systems were acquired in lineage-specific patterns. Our study data suggest that increased eukaryote diversity drives lifestyle diversification of bacteria and highlight both acquired and preexisting traits facilitating the origin of host association.IMPORTANCE Bacteria form diverse interactions with eukaryotic hosts. This is well represented by the Rhizobiales, a clade of Alphaproteobacteria strategically important for their large diversity of lifestyles with implications for agricultural and medical research. To investigate their lifestyle evolution, we compiled a comprehensive data set of genomes and lifestyle information for over 1,000 Rhizobiales genomes. We show that the origins of major host-associated lineages in Rhizobiales broadly coincided with the emergences of their host plants/animals, suggesting bacterium-host interactions as a driving force in the evolution of Rhizobiales We further found that, in addition to gene gains, preexisting traits and recurrent losses of specific genomic traits may have played underrecognized roles in the origin of host-associated lineages, providing clues to genetic engineering of microbial agricultural inoculants and prevention of the emergence of potential plant/animal pathogens.

Rapid decreases in relative testes mass among monogamous birds but not in other vertebrates.

Larger testes produce more sperm and therefore improve reproductive success in the face of sperm competition. Adaptation to social mating systems with relatively high and low sperm competition are therefore likely to have driven changes in relative testes size in opposing directions. Here, we combine the largest vertebrate testes mass dataset ever collected with phylogenetic approaches for measuring rates of morphological evolution to provide the first quantitative evidence for how relative testes mass has changed over time. We detect explosive radiations of testes mass diversity distributed throughout the vertebrate tree of life: bursts of rapid change have been frequent during vertebrate evolutionary history. In socially monogamous birds, there have been repeated rapid reductions in relative testes mass. We see no such pattern in other monogamous vertebrates; the prevalence of monogamy in birds may have increased opportunities for investment in alternative behaviours and physiologies allowing reduced investment in expensive testes.

Dominant words rise to the top by positive frequency-dependent selection.

A puzzle of language is how speakers come to use the same words for particular meanings, given that there are often many competing alternatives (e.g., "sofa," "couch," "settee"), and there is seldom a necessary connection between a word and its meaning. The well-known process of random drift-roughly corresponding in this context to "say what you hear"-can cause the frequencies of alternative words to fluctuate over time, and it is even possible for one of the words to replace all others, without any form of selection being involved. However, is drift alone an adequate explanation of a shared vocabulary? Darwin thought not. Here, we apply models of neutral drift, directional selection, and positive frequency-dependent selection to explain over 417,000 word-use choices for 418 meanings in two natural populations of speakers. We find that neutral drift does not in general explain word use. Instead, some form of selection governs word choice in over 91% of the meanings we studied. In cases where one word dominates all others for a particular meaning-such as is typical of the words in the core lexicon of a language-word choice is guided by positive frequency-dependent selection-a bias that makes speakers disproportionately likely to use the words that most others use. This bias grants an increasing advantage to the common form as it becomes more popular and provides a mechanism to explain how a shared vocabulary can spontaneously self-organize and then be maintained for centuries or even millennia, despite new words continually entering the lexicon.

Transitions between foot postures are associated with elevated rates of body size evolution in mammals.

Terrestrial mammals have evolved various foot postures: flat-footed (plantigrady), tiptoed (digitigrady), and hooved (unguligrady) postures. Although the importance of foot posture on ecology and body size of mammalian species has been widely recognized, its evolutionary trajectory and influence on body size evolution across mammalian phylogeny remain untested. Taking a Bayesian phylogenetic approach combined with a comprehensive dataset of foot postures in 880 extant mammalian species, we investigated the evolutionary history of foot postures and rates of body size evolution, within the same posture and at transitions between postures. Our results show that the common ancestor of mammals was plantigrade, and transitions predominantly occurred only between plantigrady and digitigrady and between digitigrady and unguligrady. At the transitions between plantigrady and digitigrady and between digitigrady and unguligrady, rates of body size evolution are significantly elevated leading to the larger body masses of digitigrade species (∼1 kg) and unguligrade species (∼78 kg) compared with their respective ancestral postures [plantigrady (∼0.75 kg) and digitigrady]. Our results demonstrate the importance of foot postures on mammalian body size evolution and have implications for mammalian body size increase through time. In addition, we highlight a way forward for future studies that seek to integrate morphofunctional and macroevolutionary approaches.

Evaluation of theranostic perspective of gold-silica nanoshell for cancer nano-medicine: a numerical parametric study.

Using gold-silica nanoshell as a reference nano-agent, this work has performed preliminary numerical parametric study to investigate the feasibility and if feasible the efficiency of using a single nano-agent to achieve theranostic goals. In total, seven generics of gold-silica nanoshells have been tested including the R[50,10] (radius of the silica core is 50 nm and thickness of the gold shell is 10 nm), R[40,15], R[55,25], R[40,40], R[75,40], R[104,23], and R[154,24] nanoshells. A planar tissue model has been constructed as the platform for parametric study. For mathematical modeling, radiant transport equation (RTE) has been applied to describe the interactions among laser lights, the hosting tissue, and the hosted nanoshells and Penne's bio-heat equation has been applied to describe the hyperthermia induced by such interactions. Effects of different nanoshell generics on the diffuse reflectance signal and hyperthermia temperature transition have been simulated, basing on which the potential of a certain nanoshell generic as theranostic nano-agent has been evaluated. It has been found that it is highly feasible for gold-silica nanoshells to be engineered for theranostic purpose and nanoshell generics that are preferentially scattering should be explored for good theranostic candidates. On the condition that nanoshell generic with the right optical properties has been located, a moderate nanoshell retention in the target tissue site is already sufficient to induce effective theranostic effects, which indicates that theranostic nano-medicine might not have a stringent requirement for the delivery technique. Among nanoshells that have been tested, the R[55,25] nanoshell seems to be a promising candidate as theranostic nano-agent. Further testing on it is highly recommended. Nanoshells that are preferentially absorbing such as the R[50,10] and R[40,15] nanoshells are efficient photothermal agent and could be used for therapeutic purpose only. However, it is not recommended that preferentially absorbing nanoshells being used for theranostic purpose due to possible negative effects such nanoshells might bring to the diffuse reflectance signal.

Dinosaurs reveal the geographical signature of an evolutionary radiation.

Dinosaurs dominated terrestrial ecosystems across the globe for over 100 million years and provide a classic example of an evolutionary radiation. However, little is known about how these animals radiated geographically to become globally distributed. Here, we use a biogeographical model to reconstruct the dinosaurs' ancestral locations, revealing the spatial mechanisms that underpinned this 170-million-year-long radiation. We find that dinosaurs spread rapidly initially, followed by a significant continuous and gradual reduction in their speed of movement towards the Cretaceous/Tertiary boundary (66 million years ago). This suggests that the predominant mode of dinosaur speciation changed through time with speciation originally largely driven by geographical isolation-when dinosaurs speciated more, they moved further. This was gradually replaced by increasing levels of sympatric speciation (species taking advantage of ecological opportunities within their existing environment) as terrestrial space became a limiting factor. Our results uncover the geographical signature of an evolutionary radiation.

Apportioning bacterial carbon source utilization in soil using 14 C isotope analysis of FISH-targeted bacterial populations sorted by fluorescence activated cell sorting (FACS): 14 C-FISH-FACS.

An unresolved need in microbial ecology is methodology to enable quantitative analysis of in situ microbial substrate carbon use at the population level. Here, we evaluated if a novel combination of radiocarbon-labelled substrate tracing, fluorescence in situ hybridisation (FISH) and fluorescence-activated cell sorting (FACS) to sort the FISH-targeted population for quantification of incorporated radioactivity (14 C-FISH-FACS) can address this need. Our test scenario used FISH probe PSE1284 targeting Pseudomonas spp. (and some Burkholderia spp.) and salicylic acid added to rhizosphere soil. We examined salicylic acid-14 C fate (mineralized, cell-incorporated, extractable and non-extractable) and mass balance (0-24 h) and show that the PSE1284 population captured ∼ 50% of the Nycodenz extracted biomass 14 C. Analysis of the taxonomic distribution of the salicylic acid biodegradation trait suggested that PSE1284 population success was not due to conservation of this trait but due to competitiveness for the added carbon. Adding 50KBq of 14 C sample-1 enabled detection of 14 C in the sorted population at ∼ 60-600 times background; a sensitivity which demonstrates potential extension to analysis of rarer/less active populations. Given its sensitivity and compatibility with obtaining a C mass balance, 14 C-FISH-FACS allows quantitative dissection of C flow within the microbial biomass that has hitherto not been achieved.

The deep history of the number words.

We have previously shown that the 'low limit' number words (from one to five) have exceptionally slow rates of lexical replacement when measured across the Indo-European (IE) languages. Here, we replicate this finding within the Bantu and Austronesian language families, and with new data for the IE languages. Number words can remain stable for 10 000 to over 100 000 years, or around 3.5-20 times longer than average rates of lexical replacement among the Swadesh list of 'fundamental vocabulary' items. Ordinal evidence suggests that number words also have slow rates of lexical replacement in the Pama-Nyungan language family of Australia. We offer three hypotheses to explain these slow rates of replacement: (i) that the abstract linguistic-symbolic processing of 'number' links to evolutionarily conserved brain regions associated with numerosity; (ii) that number words are unambiguous and therefore have lower 'mutation rates'; and (iii) that the number words occupy a region of the phonetic space that is relatively full and therefore resist change because alternatives are unlikely to be as 'good' as the original word.This article is part of a discussion meeting issue 'The origins of numerical abilities'.

A cautionary note on the use of Ornstein Uhlenbeck models in macroevolutionary studies.

Phylogenetic comparative methods are increasingly used to give new insights into the dynamics of trait evolution in deep time. For continuous traits the core of these methods is a suite of models that attempt to capture evolutionary patterns by extending the Brownian constant variance model. However, the properties of these models are often poorly understood, which can lead to the misinterpretation of results. Here we focus on one of these models - the Ornstein Uhlenbeck (OU) model. We show that the OU model is frequently incorrectly favoured over simpler models when using Likelihood ratio tests, and that many studies fitting this model use datasets that are small and prone to this problem. We also show that very small amounts of error in datasets can have profound effects on the inferences derived from OU models. Our results suggest that simulating fitted models and comparing with empirical results is critical when fitting OU and other extensions of the Brownian model. We conclude by making recommendations for best practice in fitting OU models in phylogenetic comparative analyses, and for interpreting the parameters of the OU model.

Bantu expansion shows that habitat alters the route and pace of human dispersals.

Unlike most other biological species, humans can use cultural innovations to occupy a range of environments, raising the intriguing question of whether human migrations move relatively independently of habitat or show preferences for familiar ones. The Bantu expansion that swept out of West Central Africa beginning ∼5,000 y ago is one of the most influential cultural events of its kind, eventually spreading over a vast geographical area a new way of life in which farming played an increasingly important role. We use a new dated phylogeny of ∼400 Bantu languages to show that migrating Bantu-speaking populations did not expand from their ancestral homeland in a "random walk" but, rather, followed emerging savannah corridors, with rainforest habitats repeatedly imposing temporal barriers to movement. When populations did move from savannah into rainforest, rates of migration were slowed, delaying the occupation of the rainforest by on average 300 y, compared with similar migratory movements exclusively within savannah or within rainforest by established rainforest populations. Despite unmatched abilities to produce innovations culturally, unfamiliar habitats significantly alter the route and pace of human dispersals.

Adaptive evolution toward larger size in mammals.

The notion that large body size confers some intrinsic advantage to biological species has been debated for centuries. Using a phylogenetic statistical approach that allows the rate of body size evolution to vary across a phylogeny, we find a long-term directional bias toward increasing size in the mammals. This pattern holds separately in 10 of 11 orders for which sufficient data are available and arises from a tendency for accelerated rates of evolution to produce increases, but not decreases, in size. On a branch-by-branch basis, increases in body size have been more than twice as likely as decreases, yielding what amounts to millions and millions of years of rapid and repeated increases in size away from the small ancestral mammal. These results are the first evidence, to our knowledge, from extant species that are compatible with Cope's rule: the pattern of body size increase through time observed in the mammalian fossil record. We show that this pattern is unlikely to be explained by several nonadaptive mechanisms for increasing size and most likely represents repeated responses to new selective circumstances. By demonstrating that it is possible to uncover ancient evolutionary trends from a combination of a phylogeny and appropriate statistical models, we illustrate how data from extant species can complement paleontological accounts of evolutionary history, opening up new avenues of investigation for both.

Detecting regular sound changes in linguistics as events of concerted evolution.

BACKGROUND: Concerted evolution is normally used to describe parallel changes at different sites in a genome, but it is also observed in languages where a specific phoneme changes to the same other phoneme in many words in the lexicon­a phenomenon known as regular sound change. We develop a general statistical model that can detect concerted changes in aligned sequence data and apply it to study regular sound changes in the Turkic language family. RESULTS: Linguistic evolution, unlike the genetic substitutional process, is dominated by events of concerted evolutionary change. Our model identified more than 70 historical events of regular sound change that occurred throughout the evolution of the Turkic language family, while simultaneously inferring a dated phylogenetic tree. Including regular sound changes yielded an approximately 4-fold improvement in the characterization of linguistic change over a simpler model of sporadic change, improved phylogenetic inference, and returned more reliable and plausible dates for events on the phylogenies. The historical timings of the concerted changes closely follow a Poisson process model, and the sound transition networks derived from our model mirror linguistic expectations. CONCLUSIONS: We demonstrate that a model with no prior knowledge of complex concerted or regular changes can nevertheless infer the historical timings and genealogical placements of events of concerted change from the signals left in contemporary data. Our model can be applied wherever discrete elements­such as genes, words, cultural trends, technologies, or morphological traits­can change in parallel within an organism or other evolving group.