Compensatory evolution and the origins of innovations

Cryptic genetic sequences have attenuated effects on phenotypes. In the classic view, relaxed selection allows cryptic genetic diversity to build up across individuals in a population, providing alleles that may later contribute to adaptation when co-opted - e.g. following a mutation increasing expression from a low, attenuated baseline. This view is described, for example, by the metaphor of the spread of a population across a neutral network in genotype space. As an alternative view, consider the fact that most phenotypic traits are affected by multiple sequences, including cryptic ones. Even in a strictly clonal population, the co-option of cryptic sequences at different loci may have different phenotypic effects and offer the population multiple adaptive possibilities. Here, we model the evolution of quantitative phenotypic characters encoded by cryptic sequences, and compare the relative contributions of genetic diversity and of variation across sites to the phenotypic potential of a population. We show that most of the phenotypic variation accessible through co-option would exist even in populations with no polymorphism. This is made possible by a history of compensatory evolution, whereby the phenotypic effect of a cryptic mutation at one site was balanced by mutations elsewhere in the genome, leading to a diversity of cryptic effect sizes across sites rather than across individuals. Cryptic sequences might accelerate adaptation and facilitate large phenotypic changes even in the absence of genetic diversity, as traditionally defined in terms of alternative alleles

Salinity effects on competition between cryptic species of the nematode Rhabditis (Pellioditis) marina

Behind the morphological similarity of many species, a substantial hidden genetic diversity can be found. This cryptic diversity has been well documented in the marine nematode, Rhabditis (Pellioditis) marina, where several cryptic species occur sympatrically. Despite the growing knowledge about its cryptic diversity, little is known about potential differences in the ecology and functional roles of the different species. In this experiment four cryptic species of Rhabditis (Pellioditis) marina were reared together, starting from identical initial abundances at two different salinity concentrations to investigate differences in their environmental preferences and how they may affect the outcome of their competitive interactions. Every fifth day of the experiment one third of the adults was removed from the population for analyzing the genetic composition of the population; adult and juvenile population dynamics were also examined. The first results show a difference in population dynamics dependent on the salinity. At low salinity, a population crash occurred after 15 days in all the populations. Only half of the populations recovered from this bottleneck and reached higher abundances than before. Genetic analyses revealed that the composition of these populations consists solely of individuals of one cryptic species. This effect was totally absent in the populations reared at higher salinity. Two explanations are possible: (1) the different cryptic species have different survival rates at different salinities and (2) the competitive interactions between them differ at different salinities. We are currently identifying the sampled adults using a restriction fragment analysis to assess these explanations and to elucidate the effect of salinity on the coexistence of cryptic species of Rhabditis (Pellioditis) marina

Cryptic female choice favours sperm from major histocompatibility complex-dissimilar males

Cryptic female choice may enable polyandrous females to avoid inbreeding or bias offspring variability at key loci after mating. However, the role of these genetic benefits in cryptic female choice remains poorly understood. Female red junglefowl, Gallus gallus, bias sperm use in favour of unrelated males. Here, we experimentally investigate whether this bias is driven by relatedness per se, or by similarity at the major histocompatibility complex (MHC), genes central to vertebrate acquired immunity, where polymorphism is critical to an individual's ability to combat pathogens. Through experimentally controlled natural matings, we confirm that selection against related males' sperm occurs within the female reproductive tract but demonstrate that this is more accurately predicted by MHC similarity: controlling for relatedness per se, more sperm reached the eggs when partners were MHC-dissimilar. Importantly, this effect appeared largely owing to similarity at a single MHC locus (class I minor). Further, the effect of MHC similarity was lost following artificial insemination, suggesting that male phenotypic cues might be required for females to select sperm differentially. These results indicate that postmating mechanisms that reduce inbreeding may do so as a consequence of more specific strategies of cryptic female choice promoting MHC diversity in offspring

Differences in life-histories refute ecological equivalence of cryptic species and provide clues to the origin of bathyal Halomonhystera (Nematoda)

The discovery of morphologically very similar but genetically distinct species complicates a proper understanding of the link between biodiversity and ecosystem functioning. Cryptic species have been frequently observed to co-occur and are thus expected to be ecological equivalent. The marine nematode Halomonhystera disjuncta contains five cryptic species (GD1-5) that co-occur in the Westerschelde estuary. In this study, we investigated the effect of three abiotic factors (salinity, temperature and sulphide) on life-history traits of three cryptic H. disjuncta species (GD1-3). Our results show that temperature had the most profound influence on all life-cycle parameters compared to a smaller effect of salinity. Life-history traits of closely related cryptic species were differentially affected by temperature, salinity and presence of sulphides which shows that cryptic H. disjuncta species are not ecologically equivalent. Our results further revealed that GD1 had the highest tolerance to a combination of sulphides, high salinities and low temperatures. The close phylogenetic position of GD1 to Halomonhystera hermesi, the dominant species in sulphidic sediments of the Hakon Mosby mud volcano (Barent Sea, 1280 m depth), indicates that both species share a recent common ancestor. Differential life-history responses to environmental changes among cryptic species may have crucial consequences for our perception on ecosystem functioning and coexistence of cryptic species

Low genetic but high morphological variation over more than 1000 km coastline refutes omnipresence of cryptic diversity in marine nematodes

Background: The resilience of ecosystems to negative impacts is generally higher when high gene flow, species diversity and genetic diversity are present. Population genetic studies are suitable to investigate genetic diversity and estimate gene flow between populations. Seaweed beds form a dynamic shallow water ecosystem influenced by climate change and human exploitation, as such, seaweed beds are a particularly powerful model to investigate ecosystem resilience in coastal areas. We studied the population genetic structure of the new nematode species Paracanthonchus gynodiporata associated with seaweeds in northeastern Brazil. Nematodes are generally believed to have a limited dispersal capacity because of the lack of planktonic larvae. Yet, they can drift on seaweeds, and water currents might be a natural barrier for their dispersal. Populations of P. gynodiporata were sampled over more than 1000 km coastline in regions across major oceanic currents with and without historical exploitation of seaweed. Results: P. gynodiporata is described in an integrative way using mitochondrial and nuclear sequences and morphological data. The 3D model of the head region shows for the first time a detailed view of the ventrosublateral teeth, a character often overlooked in older taxonomic studies of the genus. A total of 17 mitochondrial COI haplotypes were found with one haplotype representing 63 to 83% of the frequencies in each population. AMOVA showed overall little population genetic structure (F-ST = 0.05204), and no genetic subdivision between the populations under the influence of the two different water currents were found. Effects of historical seaweed exploitation on population genetic diversity were not detected. In contrast, significant differences between populations were found in morphometric characters. This discrepancy in genetic and morphological differentiation between populations across 1000 km of coastline is surprising in view of the frequently observed presence of several cryptic species at small geographical scale in other macroalgal associated nematodes. Conclusions: Our results show that cryptic species are not omnipresent in marine nematode species, suggesting that nematodes associated with seaweeds have been able to disperse over large distances across well-known biogeographic barriers

Cryptic genetic diversity in the genus <i>Mesopodopsis</i> (Crustacea, Mysidacea)

Genetic biodiversity is dynamic, species and their environment undergo continuously evolutionary changes. The start of these evolutionary changes always takes place at the population level. Genetic (allele) variation is caused by mutation, intraspecific and interspecific recombination. The survival of these new alleles is determined by selection, genetic drift and gene flow. Since molecular techniques have become more accessible in diversity research, it became clear that genetic biodiversity is strongly underestimated. The existence of cryptic genetic diversity has been revealed in numerous marine taxa (Knowlton, 2000). Next to the dispersal potential of a species other factors (such as hydrodynamic currents, dispersal limitation by larval behaviour, selection by local adaptation and historical barriers) have an important influence on population genetic divergence. Mesopodopsis slabberi is one of the most common mysids along the European coasts. The lack of pelagic larvae and the broad geographic distribution of the species, makes it an interesting model organism for evolutionary research on patterns of cryptic speciation (Remerie et al., 2006). In the present study, the degree and causes of cryptic speciation within the species M. slabberi were further investigated. By analysing populations of different habitats (lagoons, surf zones, estuaria, marshes) and by focusing on the Atlantic-Mediterranean transition zone, we tried to create a better vision about the contemporary and historical phenomena and the ecological background behind the observed genetic diversity. The phylogeographic patterns among populations were analysed by means of DNA sequencing of a fragment of the mitochondrial cytochrome c oxidase subunit 1 (COI) and the 16S ribosomal RNA genes. We concluded that a high level of cryptic speciation was present within Mesopodopsis slabberi, with at least 6 cryptic lineages (2 Atlantic, 3 Mediterranean and 1 Black Sea). These levels of cryptic diversity are discussed in a contemporary and historical framework

Dispersal and gene flow in free-living marine nematodes

Dispersal and gene flow determine connectivity among populations, and can be studied through population genetics and phylogeography. We here review the results of such a framework for free-living marine nematodes. Although field experiments have illustrated substantial dispersal in nematodes at ecological time scales, analysis of the genetic diversity illustrated the importance of priority effects, founder effects and genetic bottlenecks for population structuring between patches <1 km apart. In contrast, only little genetic structuring was observed within an estuary (<50 km), indicating that these small scale fluctuations in genetic differentiation are stabilized over deeper time scales through extensive gene flow. Interestingly, nematode species with contrasting life histories (extreme colonizers vs persisters) or with different habitat preferences (algae vs sediment) show similar, low genetic structuring. Finally, historical events have shaped the genetic pattern of marine nematodes and show that gene flow is restricted at large geographical scales. We also discuss the presence of substantial cryptic diversity in marine nematodes, and end with highlighting future important steps to further unravel nematode evolution and diversity

The Plasmid Mobilome of the Model Plant-Symbiont Sinorhizobium meliloti: Coming up with New Questions and Answers

Rhizobia are Gram-negative Alpha- andBetaproteobacteria living in the underground that have theability to associate with legumes for the establishment ofnitrogen-fixing symbioses.Sinorhizobium melilotiinparticular—the symbiont ofMedicago,Melilotus, andTrigonellaspp.—has for the past decades served as a model organism forinvestigating, at the molecular level, the biology, biochemistry,and genetics of a free-living and symbiotic soil bacterium ofagricultural relevance. To date, the genomes of seven differentS. melilotistrains have been fully sequenced and annotated,and several other draft genomic sequences are also available(http://www.ncbi.nlm.nih.gov/genome/genomes/1004).The vast amount of plasmid DNA thatS. melilotifrequently bears(up to 45% of its total genome), the conjugative ability of some ofthose plasmids, and the extent of the plasmid diversity hasprovided researchers with an extraordinary system to investigatefunctional and structural plasmid molecular biology within theevolutionary context surrounding a plant-associated modelbacterium. Current evidence indicates that the plasmidmobilome inS. melilotiis composed of replicons varying greatlyin size and having diverse conjugative systems and propertiesalong with different evolutionary stabilities and biological roles.While plasmids carrying symbiotic functions (pSyms) are knownto have high structural stability (approaching that ofchromosomes), the remaining plasmid mobilome (referred to asthe non-pSym,functionally cryptic,oraccessorycompartment)has been shown to possess remarkable diversity and to be highlyactive in conjugation. In light of the modern genomic andcurrent biochemical data on the plasmids ofS. meliloti,the current article revises their main structural components,their transfer and regulatory mechanisms, and their potentialas vehicles in shaping the evolution of the rhizobial genome.Fil: Lagares, Antonio. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Biotecnología y Biología Molecular. Universidad Nacional de La Plata. Facultad de Ciencias Exactas. Instituto de Biotecnología y Biología Molecular; ArgentinaFil: Sanjuán Pinilla, Juan. Consejo Superior de Investigaciones Científicas. Estación Experimental del Zaidín; EspañaFil: Pistorio, Mariano. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Biotecnología y Biología Molecular. Universidad Nacional de La Plata. Facultad de Ciencias Exactas. Instituto de Biotecnología y Biología Molecular; Argentin

Discrimination, Crypticity, and Incipient Taxa in Entamoeba

Persistent difficulties in resolving clear lineages in diverging populations of prokaryotes or unicellular eukaryotes (protistan polyphyletic groups) are challenging the classical species concept. Although multiple integrated approaches would render holistic taxonomies, most phylogenetic studies are still based on single-gene or morphological traits. Such methodologies conceal natural lineages, which are considered “cryptic.” The concept of species is considered artificial and inadequate to define natural populations. Social organisms display differential behaviors toward kin than to nonrelated individuals. In “social” microbes, kin discrimination has been used to help resolve crypticity. Aggregative behavior could be explored in a nonsocial protist to define phylogenetic varieties that are considered “cryptic.” Two Entamoeba invadens strains, IP-1 and VK-1:NS are considered close populations of the same “species.” This study demonstrates that IP-1 and VK-1:NS trophozoites aggregate only with alike members and discriminate members of different strains based on behavioral and chemical signals. Combined morphological, behavioral/chemical, and ecological studies could improve Archamoebae phylogenies and define cryptic varieties. Evolutionary processes in which selection acted continuously and cumulatively on ancestors of Entamoeba populations gave rise to chemical and behavioral signals that allowed individuals to discriminate nonpopulation members and, gradually, to the emergence of new lineages; alternative views that claim a “Designer” or “Creator” as responsible for protistan diversity are unfounded