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829 results for “Evolvability”
Do meristic characters used in phylogenetic analysis evolve in an ordered manner?
The use of ordered characters in phylogenetic analysis has been inconsistent through research history. It has become more widespread in recent years, and some have advocated that all characters representing continuous or meristic traits should be ordered as a matter of course. Here, using the example of dental evolution, we examine two factors that may impact on whether meristic characters actually evolve in an ordered manner: the regulatory hierarchy governing the development of teeth that allows large sections of the entire tooth-row to be supressed in a single transition, and regionalisation of the tooth row where different modules have a degree of independence in their evolution. These are studied using both empirical and simulated data. Models of evolution of such characters are examined over molecular phylogenies to see if ordered or unordered models fit best. Simulations of tooth-row evolution are designed to incorporate changes in region size and multiple levels developmental control to supress individual regions or the entire row. The empirical analyses show that in a clade with largely homodont dentition the characters evolve in an ordered manner, but if dentition is heterodont with distinct regionalisation their evolution better fits an unordered model. In the simulations, even if teeth are added and removed from the tooth row in an ordered manner, dividing the row into independently evolving modules can lead to characters covering multiple modules better fitting an unordered model of evolution. Adding the ability to suppress regions or the entire tooth row has a variable effect depending on the rates of suppression relative to the rates of addition and subtraction of individual teeth. We therefore advise not following a single policy when deciding whether to order meristic traits, but to base the decision on the evolution and developmental biology of the clade under study.
Increased time sampling in an evolve-and-resequence experiment with outcrossing Saccharomyces cerevisiae reveals multiple paths of adaptive change
<p>"Evolve and resequence" (E&R) studies combine experimental evolution and whole-genome sequencing to interrogate the genetics underlying adaptation. Due to ease of handling, E&R work with asexual organisms like bacteria can employ optimized experimental design, with large experiments and many generations of selection. By contrast, E&R experiments with sexually reproducing organisms are more difficult to implement, and design parameters vary dramatically among studies. Thus, efforts have been made to assess how these differences, such as number of independent replicates, or size of experimental populations, impact inference. We add to this work by investigating the role of time sampling – the number of discrete timepoints sequence data is collected from evolving populations. Using data from an E&R experiment with outcrossing <i>Saccharomyces cerevisiae </i>in which populations were sequenced 17 times over ~540 generations, we address the following questions: (i) do more timepoints improve the ability to identify candidate regions underlying selection? And (ii) does high-resolution sampling provide unique insight into evolutionary processes driving adaptation? We find that while time sampling does not improve the ability to identify candidate regions, high-resolution sampling does provide valuable opportunities to characterize evolutionary dynamics. Increased time sampling reveals three distinct trajectories for adaptive alleles: one consistent with classic population genetic theory (i.e. models assuming constant selection coefficients), and two where trajectories suggest more context-dependent responses (i.e. models involving dynamic selection coefficients). We conclude that while time sampling has limited impact on candidate region identification, sampling 8 or more timepoints has clear benefits for studying complex evolutionary dynamics.</p>
The Evolving Nature of Developers' Contributions in Open Source Projects
<p>Vídeo com a apresentação do artigo The Evolving Nature of Developers’ Contributions in Open Source Projects</p> <p>Autores: Talita Orfanó, Mariza Bigonha, Kecia Ferreira</p>
Temperature-mediated plasticity in incubation schedules is unlikely to evolve to buffer embryos from climatic challenges in a seasonal songbird
<p>Phenotypic plasticity is hypothesised to facilitate adaptive responses to challenging conditions, such as those resulting from climate change. However, the key predictions of this 'rescue hypothesis', that variation in plasticity exists and can evolve to buffer unfavourable conditions, remain rare. Here, we investigate among-female variation in temperature-mediated plasticity of incubation schedules and consequences for egg temperatures using the chestnut-crowned babbler (<i>Pomatostomus ruficeps</i>) from temperate regions of inland south-eastern Australia. Given phenological advances in this seasonal breeder and thermal requirements of developing embryos (> ~25 °C, optimally ~38 °C), support for evolutionary rescue - perhaps paradoxically - requires that plasticity serves to buffer embryos more from sub-optimally low temperatures. We found significant variation in the duration of incubation bouts (mean ± SD = 27 ± 22 min) and foraging bouts (mean ± SD = 17 ± 11 min) in this maternal-only incubator. However, these patterns arose because of variation in the extent to which mothers increased on- and off-bout durations when temperatures (0-36 °C) were more favourable rather than unfavourable as required under rescue. In addition, there was a strong positive intercept-slope correlation in on-bout durations, indicating that those with stronger plastic responses incubated more at average temperatures (~19 °C). Combined, these effects reduced the functional significance of plastic responses: an individual's plasticity was neither associated with daily contributions to incubation (i.e. attentiveness) nor average egg temperatures. Our results highlight that despite significant among-individual variation in environmental-sensitivity, plasticity in parental care traits need not evolve to facilitate buffering against unfavourable conditions.</p>
Data from: The impact of gene expression variation on robustness and evolvability of a developmental gene regulatory network
Regulatory interactions buffer development against genetic and environmental perturbations, but adaptation requires phenotypes to change. We investigated the relationship between robustness and evolvability within the gene regulatory network underlying development of the larval skeleton in the sea urchin Strongylocentrotus purpuratus. We find extensive variation in gene expression in this network throughout development in a natural population, some of which has a heritable genetic basis. Switch-like regulatory interactions predominate during early development, buffer expression variation, and may promote the accumulation of cryptic genetic variation affecting early stages. Regulatory interactions during later development are typically more sensitive (linear), allowing variation in expression to affect downstream target genes. Variation in skeletal morphology is associated primarily with expression variation of a few, primarily structural, genes at terminal positions within the network. These results indicate that the position and properties of gene interactions within a network can have important evolutionary consequences independent of their immediate regulatory role.
Data from: Remarkable life history polymorphism may be evolving under divergent selection in the silverleaf sunflower
Substantial intraspecific variation in life history is rare and potentially a signal of incipient ecological speciation, if variation is driven by geographically heterogenous natural selection. We present the first report of extensive life history polymorphism in Helianthus argophyllus, the silverleaf sunflower, and examine evidence for its evolution by divergent selection. In 18 populations sampled from across the species range and grown in a common garden, most quantitative traits covaried such that individuals could be assigned to two distinct life history syndromes: tall and late flowering with small initial flowerheads, or short and early flowering with larger initial flowerheads. Helianthus argophyllus exhibits regional genetic structure, but this population structure does not closely correspond with patterns of phenotypic variation. The early-flowering syndrome is primarily observed in populations from coastal barrier islands, while populations from the nearby mainland coast, although geographically and genetically close, are primarily late flowering. Additionally, several traits are more differentiated among regions than expected based on neutral genetic divergence (QST > FST), including the first principal component score corresponding with life history syndrome. This discordance between patterns of phenotypic and genetic variation suggests that divergent selection is driving genetic differences in life history across the species range. If so, the silverleaf sunflower may be in early stages of ecological speciation.
Data from: Core genes evolve rapidly in the long-term evolution experiment with Escherichia coli
Bacteria can evolve rapidly under positive selection owing to their vast numbers, allowing their genes to diversify by adapting to different environments. We asked whether the same genes that evolve rapidly in the long-term evolution experiment with Escherichia coli (LTEE) have also diversified extensively in nature. To make this comparison, we identified ~2000 core genes shared among 60 E. coli strains. During the LTEE, core genes accumulated significantly more nonsynonymous mutations than flexible (i.e., noncore) genes. Furthermore, core genes under positive selection in the LTEE are more conserved in nature than the average core gene. In some cases, adaptive mutations appear to modify protein functions, rather than merely knocking them out. The LTEE conditions are novel for E. coli, at least in relation to its evolutionary history in nature. The constancy and simplicity of the environment likely favor the complete loss of some unused functions and the fine-tuning of others.
Data from: Highest plasticity of carbon concentrating mechanisms in earliest evolved phytoplankton
Phytoplankton photosynthesis strongly relies on the operation of carbon‐concentrating mechanisms (CCMs) to accumulate CO2 around their carboxylating enzyme ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RuBisCO). Earlier evolved phytoplankton groups were shown to exhibit higher CCM activities to compensate for their RuBisCO with low CO2 specificities. Here, we tested whether earlier evolved phytoplankton groups also exhibit a higher CCM plasticity. To this end, we collected data from literature and applied a Bayesian linear meta‐analytic model. Our results show that with elevated pCO2, photosynthetic CO2 affinities decreased strongest and most consistent for the earlier evolved groups, i.e., cyanobacteria and dinoflagellates, while CO2‐dependent changes in affinities for haptophytes and diatoms were smaller and less consistent. In addition, responses of maximum photosynthetic rates toward elevated pCO2 were generally small and inconsistent across species. Our results demonstrate that phytoplankton groups with an earlier origin possess a high CCM plasticity, whereas more recently evolved groups do not, which likely results from evolved differences in the CO2 specificity of RuBisCO.
Data from: Gene expression plasticity evolves in response to colonization of freshwater lakes in threespine stickleback
Phenotypic plasticity is predicted to facilitate individual survival and/or evolve in response to novel environments. Plasticity that facilitates survival should both permit colonization and act as a buffer against further evolution, with contemporary and derived forms predicted to be similarly plastic for a suite of traits. On the other hand, given the importance of plasticity in maintaining internal homeostasis, derived populations that encounter greater environmental heterogeneity should evolve greater plasticity. We tested the evolutionary significance of phenotypic plasticity in coastal British Columbian postglacial populations of threespine stickleback (Gasterosteus aculeatus) that evolved under greater seasonal extremes in temperature after invading freshwater lakes from the sea. Two ancestral (contemporary marine) and two derived (contemporary freshwater) populations of stickleback were raised near their thermal tolerance extremes, 7°C and 22°C. Gene expression plasticity was estimated for more than 14 000 genes. Over five thousand genes were similarly plastic in marine and freshwater stickleback, but freshwater populations exhibited significantly more genes with plastic expression than marine populations. Furthermore, several of the loci shown to exhibit gene expression plasticity have been previously implicated in the adaptive evolution of freshwater populations, including a gene involved in mitochondrial regulation (PPARAa). Collectively, these data provide molecular evidence that highlights the importance of plasticity in colonization and adaptation to new environments.
Data from: The NBS-LRR architectures of plant R-proteins and metazoan NLRs evolved in independent events
There are intriguing parallels between plants and animals, with respect to the structures of their innate immune receptors, that suggest universal principles of innate immunity. The cytosolic nucleotide binding site–leucine rich repeat (NBS-LRR) resistance proteins of plants (R-proteins) and the so-called NOD-like receptors of animals (NLRs) share a domain architecture that includes a STAND (signal transduction ATPases with numerous domains) family NTPase followed by a series of LRRs, suggesting inheritance from a common ancestor with that architecture. Focusing on the STAND NTPases of plant R-proteins, animal NLRs, and their homologs that represent the NB-ARC (nucleotide-binding adaptor shared by APAF-1, certain R gene products and CED-4) and NACHT (named for NAIP, CIIA, HET-E, and TEP1) subfamilies of the STAND NTPases, we analyzed the phylogenetic distribution of the NBS-LRR domain architecture, used maximum-likelihood methods to infer a phylogeny of the NTPase domains of R-proteins, and reconstructed the domain structure of the protein containing the common ancestor of the STAND NTPase domain of R-proteins and NLRs. Our analyses reject monophyly of plant R-proteins and NLRs and suggest that the protein containing the last common ancestor of the STAND NTPases of plant R-proteins and animal NLRs (and, by extension, all NB-ARC and NACHT domains) possessed a domain structure that included a STAND NTPase paired with a series of tetratricopeptide repeats. These analyses reject the hypothesis that the domain architecture of R-proteins and NLRs was inherited from a common ancestor and instead suggest the domain architecture evolved at least twice. It remains unclear whether the NBS-LRR architectures were innovations of plants and animals themselves or were acquired by one or both lineages through horizontal gene transfer.
Data from: Evolved genetic and phenotypic differences due to mitochondrial-nuclear interactions
The oxidative phosphorylation (OxPhos) pathway is responsible for most aerobic ATP production and is the only pathway with both nuclear and mitochondrial encoded proteins. The importance of the interactions between these two genomes has recently received more attention because of their potential evolutionary effects and how they may affect human health and disease. In many different organisms, healthy nuclear and mitochondrial genome hybrids between species or among distant populations within a species affect fitness and OxPhos functions. However, what is less understood is whether these interactions impact individuals within a single natural population. The significance of this impact depends on the strength of selection for mito-nuclear interactions. We examined whether mito-nuclear interactions alter allele frequencies for ~11,000 nuclear SNPs within a single, natural Fundulus heteroclitus population containing two divergent mitochondrial haplotypes (mt-haplotypes). Between the two mt-haplotypes, there are significant nuclear allele frequency differences for 349 SNPs with a p-value of 1% (236 with 10% FDR). Unlike the rest of the genome, these 349 outlier SNPs form two groups associated with each mt-haplotype, with a minority of individuals having mixed ancestry. We use this mixed ancestry in combination with mt-haplotype as a polygenic factor to explain a significant fraction of the individual OxPhos variation. These data suggest that mito-nuclear interactions affect cardiac OxPhos function. The 349 outlier SNPs occur in genes involved in regulating metabolic processes but are not directly associated with the 79 nuclear OxPhos proteins. Therefore, we postulate that the evolution of mito-nuclear interactions affects OxPhos function by acting upstream of OxPhos.
Data from: Female resistance to sexual coercion can evolve to preserve the indirect benefits of mate choice
Sexual conflict over the indirect benefits of mate choice may arise when traits in one sex limit the ability of the other sex to freely choose mates but when these coercive traits are not necessarily directly harmful (i.e. forced fertilization per se). While we might hypothesize that females can evolve resistance in order to retain the indirect, genetic benefits (reflected in offspring attractiveness) of mating with attractive males, up to now it has been difficult to evaluate potential underlying mechanisms. Traditional theoretical approaches do not usually conceptually distinguish between female preference for male mating display and female resistance to forced fertilization, yet sexual conflict over indirect benefits implies the simultaneous action of all of these traits. Here we present an integrative theoretical framework that draws together concepts from both sexual selection and sexual conflict traditions, allowing for the simultaneous coevolution of displays and preferences, and of coercion and resistance. We demonstrate that it is possible for resistance to coercion to evolve in the absence of direct costs of mating to preserve the indirect benefits of mate choice. We find that resistance traits that improve the efficacy of female mating preference can evolve as long as females are able to attain some indirect benefits of mating with attractive males, even when both attractive and unattractive males can coerce. These results reveal new evolutionary outcomes that were not predicted by prior theories of indirect benefits or sexual conflict.
Data from: Trunk dental tissue evolved independently from underlying dermal bony plates but is associated to surface bones in living odontode-bearing catfish
Although oral dental tissue is a vertebrate attribute, trunk dental tissue evolved in several extinct vertebrate lineages but is rare among living species. The question of which processes trigger dental-tissue formation in the trunk remains open, and would shed light on odontogenesis evolution. Extra-oral dental structures (odontodes) in the trunk are associated with underlying dermal bony plates, leading us to ask whether the formation of trunk bony plates is necessary for trunk odontodes to emerge. To address this question, we focus on Loricarioidei: an extant, highly diverse group of catfish whose species all have odontodes. We examined the location and cover of odontodes and trunk dermal bony plates for all six loricarioid families and 17 non-loricarioid catfish families for comparison. We inferred the phylogeny of Loricarioidei using a new 10-gene dataset, eight time-calibration points, and noise-reduction techniques. Based on this phylogeny, we reconstructed the ancestral states of odontode and bony plate cover, and find that trunk odontodes emerged before dermal bony plates in Loricarioidei. Yet we discovered that when bony plates are absent, other surface bones are always associated with odontodes, suggesting a link between osteogenic and odontogenic developmental pathways, and indicating a remarkable trunk odontogenic potential in Loricarioidei.
Data from: The thermal dependency of locomotor performance evolves rapidly within an invasive species
Biological invasions can stimulate rapid shifts in organismal performance, via both plasticity and adaptation. We can distinguish between these two proximate mechanisms by rearing offspring from populations under identical conditions, and measuring their locomotor abilities in standardized trials. We collected adult cane toads (Rhinella marina) from invasive populations that inhabit regions of Australia with different climatic conditions. We bred those toads, and raised their offspring under common-garden conditions before testing their locomotor performance. At high (but not low) temperatures, offspring of individuals from a hotter location (northwestern Australia) outperformed offspring of conspecifics from a cooler location (northeastern Australia). This disparity indicates that, within less than 100 years, thermal performance in cane toads has adapted to the novel abiotic challenges that cane toads have encountered during their invasion of tropical Australia.
Data from: Genetic diversity, population structure and sex-biased dispersal in three co-evolving species
Genetic diversity and spatial structure of populations are important for antagonistic coevolution. We investigated genetic variation and population structure of three closely related European ant species: the social parasite Harpagoxenus sublaevis and its two host species Leptothorax acervorum and Leptothorax muscorum. We sampled populations in 12 countries and analyzed eight microsatellite loci and an mtDNA sequence. We found high levels of genetic variation in all three species, only slightly less variation in the host L. muscorum. Using a newly introduced measure of differentiation (Jost's DEST), we detected strong population structuring in all species and less male-biased dispersal than previously thought. We found no phylogeographic patterns that could give information on post-glacial colonization routes - northern populations are as variable as more southern populations. We conclude that conditions for Thompson's geographic mosaic of coevolution are ideal in this system: all three species show ample genetic variation and strong population structure.
Data from: Do convergent ecomorphs evolve through convergent morphological pathways? Cranial shape evolution in fossil hyaenids and borophagine canids (Carnivora, Mammalia)
Cases of convergent evolution, particularly within ecomorphological contexts, are instructive in identifying universally adaptive morphological features across clades. Tracing of evolutionary pathways by which ecomorphological convergence takes place can further reveal mechanisms of adaptation, which may be strongly influenced by phylogeny. Ecomorphologies of carnivorous mammals represent some of the most outstanding cases of convergent evolution in the Cenozoic radiation of mammals. This study examined patterns of cranial shape change in the dog (Canidae) and hyena (Hyaenidae) families, in order to compare the evolutionary pathways that led to the independent specialization of bone-cracking hypercarnivores within each clade. Geometric morphometrics analyses of cranial shape in fossil hyaenids and borophagine canids provided evidence for deep-time convergence in morphological pathways toward the independent evolution of derived bone-crackers. Both clades contained stem members with plesiomorphic generalist/omnivore cranial shapes, which evolved into doglike species along parallel pathways of shape change. The evolution of specialized bone-crackers from these doglike forms, however, continued under the constraint of a full cheek dentition and restriction on rostrum length reduction in canids, but not hyaenids. Functionally, phylogenetic constraint may have limited borophagine canids to crack bones principally with their carnassial instead of the third premolar as in hyaenids, but other cranial shape changes associated with durophagy nevertheless evolved in parallel in the two lineages. Size allometry was not a major factor in cranial shape evolution in either lineage, supporting the interpretation of functional demands as drivers for the observed convergence. The comparison between borophagines and hyaenids showed that differential effects of alternative functional "solutions" that arise during morphological evolution may be multiplied with processes of the "macroevolutionary ratchet" already in place to further limit the evolutionary pathways available to specialized lineages.
Data from: Directional dispersal has not evolved during the cane toad invasion
1. The ability to disperse along a consistent compass heading strongly affects the rate and efficiency of an animal's displacement, and thus is under selection at the expanding edge of a biological invasion. 2. We used radiotelemetry to assess whether the dispersal direction of cane toads (Rhinella marina) changed as a function of time since invasion, by comparing (i) toads at a single site monitored annually for 10 years subsequent to toad arrival; (ii) toads collected from sites across the species' invaded range in Australia, and radiotracked at a common site; and (iii) the offspring of those transported toads that were reared in captivity under common-garden conditions. 3. The first of these data sets showed non-random directionality, indicating strong spatial sorting operating on this trait: toads moved in a north-westerly direction for the first 6 years post-invasion, but in random directions thereafter. Despite the evidence for trait sorting, no consistent directionality was seen in toads relocated from populations with different invasion histories nor in their offspring. Why do we see no evolutionary shifts? 4. Dispersal directionality of the offspring was not correlated with that of their parents, arguing against a genetic basis to this behavioural trait. Thus, while an expanding invasion front creates an evolutionary pressure for animals to move in a specific direction, evolution of this trait has not occurred in this system because directionality is not heritable. 5. The observed north-westerly movements of toads at the invasion front were due to simple density differentials: in the first few years, most toads arriving at our study site originated from earlier-colonized (and hence denser) populations to the south-east.
Data from: Soil biota and chemical interactions promote co-existence in co-evolved grassland communities
1. Plant populations can exhibit local adaptation to their abiotic environment, such as climate and soil properties, as well as biotic components such as the chemical signatures of dominant plant species and mutualistic and pathogenic microbial populations. While patterns of local adaptation in individual species are widely recorded, the importance of microevolutionary processes for plant community assembly and function is poorly understood. 2. Here we examined how a history of long-term co-existence, and thus potential for local co-adaptation, influenced the process of plant community assembly. Soil inocula and seeds of eight plant species were collected from three calcareous grasslands with a long history of grazing within a single geographical region. Mesocosm communities were established using local genotypes from a single site or an artificial mixture of genotypes from two different sites. To investigate the role of root exudates and local ("home") and non-local ("away") soil biota as mediators of plant species co-existence, the population origin treatment was combined with the addition of activated carbon, which is known to adsorb exudates from soil, and sterilisation of soil inocula. Individual-, species- and mesocosm-level responses were measured over the course of three growing seasons. 3. We found that root exudates promoted seedling survival, species co-existence and productivity in assemblages of genotypes originating from the same community but had a weak impact in mixed, novel communities. Soil biota promoted the growth of subordinate forbs and restrained the growth of dominant graminoids, particularly in communities composed of local genotypes. The effects of population origin were significant in the first two years of the experiment but were not detectable in the third year when interbreeding and new seedling establishment took place. Plant genotypes coupled with "home" microbial inoculum experienced a stronger reduction in growth compared with genotypes exposed to "away" inoculum, indicating that plants experienced home-field disadvantage in interactions with soil biota. 4. Synthesis. Our study demonstrates that the mechanisms of initial grassland community assembly depend on community history, with belowground chemical interactions and plant interactions with soil biota becoming stronger drivers of dynamics in established and potentially co-evolved communities.
Data from: Evolving from static to dynamic signals: evolutionary compensation between two communicative signals
Signals that convey related information may impose selection on each other, creating evolutionary links between different components of the communicative repertoire. Here, we ask about the consequences of the evolutionary loss of one signal (a colour patch) on another (a motion display) in Sceloporus lizards. We present data on male lizards of four species: two pairs of sister taxa representing two independent evolutionary losses of the static colour patch (S. cozumelae and S. parvus; S. siniferus and S. merriami). Males of the two species that have undergone an evolutionary loss of blue-belly patches (S. cozumelae, S. siniferus) were less active than their blue-bellied sister taxa (S. parvus, S. merriami), consistent with the suggestion that the belly patches were lost to reduce conspicuousness of species with high predation pressure. In contrast, the headbob display appears to have become more, rather than less, conspicuous over evolutionary time. The colour patch is exhibited primarily during aggressive encounters, whereas headbob displays are multifunction signals used in several different contexts, including aggressive encounters. Males of species that have lost the colour patch produced more motion displays, and the structure of those motion displays were more similar to those produced during combat. In both evolutionary episodes, a static colour signal appears to have been replaced by dynamic motion displays that can be turned off in the presence of predators and other unwanted receivers. The predominant pattern is one of evolutionary compensation and interactions between multiple signals that convey related information.
Data from: C4 anatomy can evolve via a single developmental change
C4 photosynthesis boosts productivity in warm environments. Paradoxically, this complex physiological process evolved independently in numerous plant lineages, despite requiring specialized leaf anatomy. The anatomical modifications underlying C4 evolution have previously been evaluated through interspecific comparisons, which capture numerous changes besides those needed for C4 functionality. Here, we quantify the anatomical changes accompanying the transition between non-C4 and C4 phenotypes by sampling widely across the continuum of leaf anatomical traits in the grass Alloteropsis semialata. Within this species, the only trait that is shared and specific to C4 individuals is an increase in vein density, driven specifically by minor vein development. The minor veins are genetically determined, and their multiple effects facilitate C4 function. For species with the necessary anatomical preconditions, developmental proliferation of veins can therefore be sufficient to produce a functional C4 leaf anatomy, creating an evolutionary entry point to complex C4 syndromes that can become more specialized.
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Allen Brain Atlas
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Annotated Behaviour and Observability Dataset (ABODe)
ABODe is a University of Edinburgh DataShare dataset for behavior classification in group-housed mice using home-cage video, identities, bounding boxes, ground-plate positions, and annotator labels.
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