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The interaction among evolutionary forces in the pathogenic fungus Mycosphaerella graminicola.

The population genetic dynamic of a species is driven by interactions among mutation, migration, drift, mating system, and selection, but it is rare to have sufficient empirical data to estimate values for all of these forces and to allow comparison of the relative magnitudes of these evolutionary forces. We combined data from a mark-release-recapture experiment, extensive population surveys, and computer simulations to evaluate interactions among these evolutionary forces in the pathogenic fungus Mycosphaerella graminicola. The results from these studies showed that, on average, the immigration rate was 0.027, the fraction of outcrossing individuals was 0.035, and the selection coefficient associated with immigrants was 0.106 each generation. We also estimated that effective population sizes for this fungus were larger than 24,000 and the mutation rate for the RFLP markers used in surveys and field experiments was approximately 4 x 10(-5). Computer simulations based on these estimates indicate that, on average, the global population of M. graminicola has reached equilibrium. Population genetic parameters including number of alleles, gene diversity, and population subdivision estimated from the computer simulations were surprisingly close to empirical estimates. Simulations also revealed that random drift is the major evolutionary force decreasing genetic variation in this fungus, followed by natural selection. The major force adding to genetic variation was mutation, followed by gene flow and sexual recombination. Gene flow played the leading role in decreasing population subdivision while natural selection was the major factor increasing population subdivision.

Ascomycota↗

Phylogenetic analysis of Phytophthora species based on mitochondrial and nuclear DNA sequences.

A molecular phylogenetic analysis of the genus Phytophthora was performed, 113 isolates from 48 Phytophthora species were included in this analysis. Phylogenetic analyses were performed on regions of mitochondrial (cytochrome c oxidase subunit 1; NADH dehydrogenase subunit 1) and nuclear gene sequences (translation elongation factor 1alpha; beta-tubulin) and comparisons made to test for incongruence between the mitochondrial and nuclear data sets. The genus Phytophthora was confirmed to be monophyletic. In addition, results confirm that the classical taxonomic grouping as described by [Waterhouse (1963)] does not reflect true phylogenetic relations. Phytophthora species were redistributed into 8 clades, providing a more accurate representation of phylogenetic relationships within the genus Phytophthora. The evolution and transition of morphological, pathogenic, and reproductive traits was inferred from the cladogram generated in this study. Mating system was inferred to be a homoplasious trait, with at least eight independent transitions from homothallism to heterothallism observed.

Biological Evolution↗

Low-level exposure to organophosphate pesticides leads to restrictive lung dysfunction.

Apart from symptomology, there are very few reports on lung function following exposure to low levels of organophosphate (OP) pesticides in man. Twenty-five occupationally exposed farmers and 22 environmentally exposed freshwater fishermen were evaluated between and during OP spray seasons. Forty marine fishermen living away from agricultural areas were recruited as a control group. Forced vital capacity (FVC) and forced expiratory volume in the first second (FEV(1)) were measured by spirometry. Haemoglobin corrected erythrocyte acetylcholinesterase (AChE) levels were measured during and between (baseline estimation) spray seasons using a portable WHO-approved Test-mate system (EQM Research, Ohio). FVC ratio was lower in the farmers as compared to the controls (P<0.001) between exposure seasons. In the farmers, FVC ratio decreased further during the exposure season (P=0.023). FEV(1) was lower in the farmers as compared to the controls in both periods (P<0.05). In the fishermen, the decrease in ratios of FVC and FEV(1) following exposure to pesticides was not significant. FEV(1)/FVC ratios were similar in the three groups between (P=0.988) and during (P=0.159) exposure periods. Following exposure to OPs, AChE levels dropped 12.75% in the farmers (P<0.001) and 5.62% in the freshwater fishermen (P=0.001). Occupational exposure to OP results in restrictive lung dysfunction, a phenomenon not observed following environmental exposure.

Acetylcholinesterase↗

Orchid diversity: an evolutionary consequence of deception?

The Orchidaceae are one of the most species-rich plant families and their floral diversity and pollination biology have long intrigued evolutionary biologists. About one-third of the estimated 18,500 species are thought to be pollinated by deceit. To date, the focus has been on how such pollination evolved, how the different types of deception work, and how it is maintained, but little progress has been made in understanding its evolutionary consequences. To address this issue, we discuss here how deception affects orchid mating systems, the evolution of reproductive isolation, speciation processes and neutral genetic divergence among species. We argue that pollination by deceit is one of the keys to orchid floral and species diversity. A better understanding of its evolutionary consequences could help evolutionary biologists to unravel the reasons for the evolutionary success of orchids.

Journal Article↗

High levels of LH and testosterone in a tropical seabird with an elaborate courtship display.

For birds breeding in temperate areas, territoriality and courtship co-occur during spring and are mirrored by elevated plasma levels of luteinizing hormone (LH) and testosterone. By contrast, several tropical species show no major changes in LH and testosterone throughout the year and plasma levels are often low. This study describes changes in body condition, LH and testosterone levels throughout breeding in a tropical seabird, the magnificent frigatebird (Fregata magnificens). Although bi-parental care is the rule in frigatebirds, these seabirds are unique among seabirds in having a lek-like mating system where males perform an elaborate courtship display during which an extravagant sexual ornament (gular pouch) is inflated. Body condition peaked during the displaying stage in males and declined gradually in both sexes, being the lowest during chick-rearing. LH and testosterone titers were the highest in displaying males with a fully inflated gular pouch (means: 23.6 ng/ml for LH and 6.2 ng/ml for testosterone) and remained low (means: 3.8 ng/ml for LH and 0.3 ng/ml for testosterone) during incubation and chick-rearing in both sexes. Body condition was positively related to testosterone levels in displaying males. In male frigatebirds, high levels of testosterone are not associated with aggression but are probably required to display the inflated gular pouch and to perform an elaborate courtship display. Because males invest in incubation and chick-rearing, testosterone has to return to low levels, to permit parental care. This hormonal pattern differs greatly with that of the few tropical seabirds studied so far and is probably related to the unique reproductive strategy of frigatebirds.

Aggression↗

Plasma steroid hormones in two Arctic-breeding shorebirds: monogamy versus polygyny.

Steroid hormones are predicted to vary with mating systems, degree of paternal care and aggression, as proposed in the "challenge hypothesis." We measured plasma concentrations of testosterone (T) and 5alpha-dihydrotestosterone (DHT) in two high Arctic breeding and closely related shorebird species, the polygynous pectoral sandpiper (Calidris melanotos) and the monogamous semipalmated sandpiper (Calidris pusilla) to examine whether the hormonal findings corroborate the predictions of the challenge hypothesis. In both species, males showed significantly higher levels of DHT and T than females, but in pectoral sandpipers median T levels were 34 times greater in males than in females, whereas in semipalmated sandpiper there was only a 4.9-fold difference. T and DHT concentrations correlated in semipalmated sandpipers and in male, but not in female, pectoral sandpipers. In semipalmated sandpipers, androgen levels were highest in the beginning of the breeding season and then declined, whereas male pectoral sandpipers showed extremely high androgen levels which were sustained throughout the breeding season. In both species, androgen levels were independent of body condition. Several incubating male semipalmated sandpipers had high circulating T levels, suggesting that low T is not required to exhibit paternal care. Our results are consistent with the challenge hypothesis in that androgen concentrations were higher throughout the breeding season in the polygynous compared to the monogamous species. Our study also supports previous studies suggesting that the short breeding season in the high Arctic and the importance of male care to nest success may be factors leading to behavioral T insensitivity.

Animals↗

Variability of GnRH secretion in two goby species with socially controlled alternative male mating tactics.

Male reproductive phenotypic plasticity related to environmental-social conditions is common among teleost fish. In several species, males adopt different mating tactics depending on their size, monopolizing mates when larger, while parasitizing dominant male spawns when smaller. Males performing alternative mating tactics are often characterized by a strong dimorphism in both primary and secondary reproductive traits. According to studies on sex-changing species and on species where only one male morph is reproductively active, male alternative phenotypes are expected to vary also in gonadotropin-releasing hormone (GnRH) neurons in forebrain preoptic area (POA). Here, we compared the intra- and inter-sexual variations in number and size of GnRH neurons, along with gonads and male accessory structure investment, in two goby species, the grass goby, Zosterisessor ophiocephalus, and the black goby, Gobius niger, characterized by male alternative mating phenotypes. In both species, older and larger males defend nests, court and perform parental care, while younger and smaller ones try to sneak territorial male spawning. We found that grass goby and black goby have different patterns of GnRH expression. Grass goby presents a clear intra-sexual dimorphism in GnRH expression, related to the occurrence of alternative mating tactics, while in the black goby, only inter-sexual differences are observed. The inter- and intra-specific variability in the GnRH neurons in these two goby species is discussed in light of the differences in migratory behavior, nest type, and mating system.

Analysis of Variance↗

The influence of self-fertilization and pairing on life-history traits in the freshwater snail Bulinus forskalii (Gastropoda, planorbidae).

This study analyses the basic reproductive biology of the schistosonie-vector snail Bulinus forskalli. This hermaphroditic species can self-fertilize. Variation of the mating system was analysed in two populations from Cameroon in a three-step experiment. (i) The fecundity (number of eggs and of egg capsules) of isolated virgin individuals from both populations was followed over a month, as well as the survival of offspring from these individuals. Estimates of these parameters were intermediate. Our results also indicate that inbreeding depression has a maximum value of about one-half in both populations. No difference was observed between populations for both the overall fecundity of adults and survival of offspring. (ii) Individuals were then paired either within, or between populations over three successive days, and the number of copulations recorded. Copulations were observed in all situations, including between-population pairs. No significant dlifference in the number of copulations per pair was detected. However, this value decreased with time, and reciprocal copulations were significantly less numerous between than within populations. (iii) Individuals that successfully copulated as female in (ii) were isolated, and their fecundity was followed over 3 weeks. The fecundity of individuals that had been maintained isolated over the whole experiment served as a control. No significant difference was on the whole detected. Our results collectively suggest that the individuals from the two populations studied are rather selfers, confirming that B. forskalii is a preferentially selfing species. There is also some pre-zygotic isolation against crosses between populations.

Animals↗

Are subordinates always stressed? A comparative analysis of rank differences in cortisol levels among primates.

Among primate species there is pronounced variation in the relationship between social status and measures of stress physiology. An informal meta-analysis was designed to investigate the basis of this diversity across different primate societies. Species were included only if a substantial amount of published information was available regarding both social behavior and rank-related differences in stress physiology. Four Old World and three New World species met these criteria, including societies varying from small-group, singular cooperative breeders (common marmoset and cotton top tamarin) to large-troop, multi-male, multi-female polygynous mating systems (rhesus, cynomolgus, talapoin, squirrel monkeys, and olive baboon). A questionnaire was formulated to obtain information necessary to characterize the stress milieu for individuals in particular primate societies. We standardized cortisol values within each species by calculating the ratio of basal cortisol concentrations of subordinates to those of dominants in stable dominance hierarchies and expressing the ratio as a percentage (relative cortisol levels). The meta-analysis identified two variables that significantly predicted relative cortisol levels: subordinates exhibited higher relative cortisol levels when they (1). were subjected to higher rates of stressors, and (2). experienced decreased opportunities for social (including close kin) support. These findings have important implications for understanding the different physiological consequences of dominant and subordinate social status across primate societies and how social rank may differ in its behavioral and physiological manifestations among primate societies.

Aggression↗

The influence of intensity of infection by a trematode parasite on the reproductive biology of Gammarus insensibilis (Amphipoda).

Following the behavioural alterations induced by the trematode Microphallus papillorobustus Rankin 1940 (Trematoda, Microphallidae) on its second intermediate host, the amphipod Gammmarus insensibilis, infected individuals are likely to mate among themselves. We investigated the influence of parasite intensity on the reproductive biology of infected hosts. In the mating system of amphipods, males compete severely for access to females and large males have greater ability to obtain large and more fecund females. We showed that the null hypothesis of random pair formation according to parasite intensity could not be rejected. In addition, infected males obtained females of the expected size according to their own sizes, whatever their parasite intensities. However, in both males and females, the parasite intensity increased the intermoult duration. Because size and reproductive success are strongly correlated in amphipods, we discuss the influence of this process on host fitness.

Animals↗

A site-specific DNA endonuclease specified by one of two ORFs encoded by a group I intron in Dictyostelium discoideum mitochondrial DNA.

The second intron (DdOX1/2.2) of Dictyostelium discoideum cytochrome oxidase subunit 1/2 fused gene has two free-standing ORF genes (Dd ai2a and Dd ai2b) in a loop, which have similar amino acid sequences and are homologous to aI4 DNA endonuclease (I-SceII) of Saccharomyces cerevisiae. To elucidate the functions of these ORFs, we cloned the ORFs into an expression vector and introduced the composite vectors into E. coli. The expression of Dd ai2a in E. coli caused growth inhibition and degradation of the E. coli genomic DNA. To determine whether Dd ai2a protein is a homing type DNA endonuclease, the ability to cleave the homing site of its intron in vivo was examined. Dd ai2a cleaved only one strand of intronless DNA sequence at the site which coincides with the I-SceII cleavage recognition site. We suppose that Dd ai2a functions actually as a homing type DNA endonuclease in D. discoideum mitochondria by virtue of other factors. To obtain further information about the relationship between the existence of introns and the mating system, we carried out in vitro self-splicing assay and polymerase chain reaction analysis using 13 strains of the cellular slime mold.

Animals↗

Active immunization of ewes against ovine placental lactogen increases birth weight of lambs and milk production with no adverse effect on conception rate.

In two experiments, 16 Booroola-Assaf and 35 Assaf ewe-lambs were actively immunized at 5 months of age against recombinant ovine placental lactogen (oPL). At 9 months of age, the ewe-lambs were mated for the first time and then introduced into a frequent mating-system. Anti-oPL antibody titers, reproductive performance, maternal serum levels of oPL during pregnancy, lamb birth weight and milk production of the ewes were followed in the immunized ewes and in their non-immunized control counterparts. All the immunized ewes developed anti-oPL antibodies, which interfered with oPL bioactivity in an in vitro cell proliferation assay. Conception rates did not differ (P>0.05) between immunized and non-immunized ewes. Abundant antibody-bound non-active oPL detected in sera of immunized ewes by western blotting indicated enhanced oPL production by the placenta following immunization. An increase (P<0.02) in serum oPL bioactivity, but not immunoreactivity, was observed in the immunized ewes in late gestation relative to control ewes. The average litter size was 1.83 and 1.32 lambs born per ewe lambing in the first and second experiments, respectively. Average birth weights of lambs born to the immunized ewes were higher (P<0.01) than for lambs born to control ewes by 10, 17 and 39% for those born as singles, twins and triplets, respectively. Immunized ewes produced 19 and 33% more milk (P<0.02) than the control ewes in the first 3.5 months of the first and second lactations, respectively. These findings do not suggest a role for oPL in maternal recognition of pregnancy, but they strongly suggest important roles for oPL in fetal growth and mammogenesis. Immunization of ewes against oPL may thus represent a novel practical technique for enhancing birth weights of lambs born to prolific sheep, as well as milk production by both dairy and mutton ewes.

Animals↗

Genome evolution: sex and the transposable element.

Mating systems are thought to play an important role in determining the fate of genomic parasites such as transposable elements. This is supported by recent studies which indicate that asexual genomes may be structured very differently to those of sexual species.

Animals↗

Female preferences in a fish genus without female mate choice.

The evolution and the adaptive logic (if any) of female mate choice are subjects of lively debate. Whereas most researchers believe that females have evolved to recognize signs of male 'quality' (the ability to provide females or their offspring with direct or indirect genetic or material benefits), there is intriguing evidence that males can evolve to appeal to pre-existing female preferences. Evidence for these pre-existing biases is often ambiguous because phylogenetic reconstructions have usually failed to establish conclusively whether the female preference or the favored male traits evolved first. This potential difficulty is minimal in the mosquitofish genus Gambusia, none of whose 45 species appears to have a female-choice mating system in the wild, and none of which shows the male behavioral and morphological traits that are characteristic of female choice. Nevertheless, in an experimental situation in the laboratory, female Gambusia holbrooki readily chose between models of males and demonstrated significant and reliable preferences for a variety of exaggerated male traits that are not seen in their species or their genus. Other morphological alterations were not preferred. The latent willingness of females to choose traits in a genus without such traits and without evident female choice in the wild is remarkable and may indicate a pre-existing bias in females that is ready to drive male evolution, should the social system or the ecological variables that control it change.

Animals↗

Genetic diversity and population structure of Plasmodium falciparum isolates from Dakar, Senegal, investigated from microsatellite and antigen determinant loci.

We investigated the genetic diversity and the population structure of 32 Plasmodium falciparum blood sample isolates (25 from Dakar city and suburbs and seven from other localities in Senegal) with two different types of molecular markers, 19 microsatellite and four antigenic determinant loci. Under the same technical procedure, microsatellite loci showed a mean number of alleles greater than that of antigenic loci. Both markers revealed that 15.6% of blood samples were multi-infected. Mean expected heterozygosity calculated from microsatellites and antigens was similar, 0.74 and 0.70, respectively. Significant linkage disequilibrium was observed from microsatellite loci and antigenic determinant loci. This suggests a non-panmictic structure for this sample that could be explained by two non-exclusive hypotheses: (i) a particular mating system (i.e. clonality), and/or (ii) a population structure in P. falciparum (i.e. Wahlund effect). Urban samples could have been drawn from a heterogeneous set of foci with different level of parasitic transmission. Moreover, no relationship was found between multilocus genotypes and different parameters (i.e. age, sex and blood group of parasitized patients; number of trophozoites per microliter of blood). The results are discussed taking into account recently published studies on malaria population biology.

Adolescent↗

Implications of an individualistic lifestyle for species conservation: lessons from jealous beasts.

Polecat populations show a very low genetic diversity and a high inbreeding coefficient. Furthermore, the estimate of effective population size is alarmingly low. Polecats Mustela putorius populations are structured into scattered breeding sub-units usually made up of one male and two females, according to a polygynous mating system. Because a strict spatio-temporal segregation was observed between males and females, we propose to call individualistic such species. We suggest that the solitary habits of individualistic species may result in or worsen a high inbreeding and exacerbate their conservation issue, a crucial perspective for critically endangered species such as the European mink.

Animals↗

Adaptive explanation in socio-ecology: lessons from the Equidae.

Socio-ecological explanations for intra- and interspecific variation in the social and spatial organization of animals predominate in the scientific literature. The socio-ecological model, developed first for the Bovidae and Cervidae, is commonly applied more widely to other groups including the Equidae. Intraspecific comparisons are particularly valuable because they allow the role of environment and demography on social and spatial organization to be understood while controlling for phylogeny or morphology which confound interspecific comparisons. Feral horse (Equus caballus Linnaeus 1758) populations with different demography inhabit a range of environments throughout the world. I use 56 reports to obtain 23 measures or characteristics of the behaviour and the social and spatial organization of 19 feral horse populations in which the environment, demography, management, research effort and sample size are also described. Comparison shows that different populations had remarkably similar social and spatial organization and that group sizes and composition, and home range sizes varied as much within as between populations. I assess the few exceptions to uniformity and conclude that they are due to the attributes of the studies themselves, particularly to poor definition of terms and inadequate empiricism, rather than to the environment or demography per se. Interspecific comparisons show that equid species adhere to their different social and spatial organizations despite similarities in their environments and even when species are sympatric. Furthermore, equid male territoriality has been ill-defined in previous studies, observations presented as evidence of territoriality are also found in non-territorial equids, and populations of supposedly territorial species demonstrate female defence polygyny. Thus, territoriality may not be a useful categorization in the Equidae. Moreover, although equid socio-ecologists have relied on the socio-ecological model derived from the extremely diverse Bovidae and Cervidae for explanations of variation in equine society, the homomorphic, but large and polygynous, and monogeneric Equidae do not support previous socio-ecological explanations for relationships between body size, mating system and sexual dimorphism in ungulates. Consequently, in spite of the efforts of numerous authors during the past two decades, functional explanations of apparent differences in feral horse and equid social and spatial organization and behaviour based on assumptions of their current utility in the environmental or demographic context remain unconvincing. Nevertheless, differences in social cohesion between species that are insensitive to intra- and interspecific variation in habitat and predation pressure warrant explanation. Thus, I propose alternative avenues of inquiry including testing for species-specific differences in inter-individual aggression and investigating the role of phylogenetic constraints in equine society. The Equidae are evidence of the relative importance of phylogeny and biological structure, and unimportance of the present-day environment, in animal behaviour and social and spatial organization.

Adaptation, Physiological↗

Genetic differentiation of neutral markers and quantitative traits in predominantly selfing metapopulations: confronting theory and experiments with Arabidopsis thaliana.

The comparison of the genetic differentiation of quantitative traits (QST) and molecular markers (FST) can inform on the strength and spatial heterogeneity of selection in natural populations, provided that markers behave neutrally. However, selection may influence the behaviour of markers in selfing species with strong linkage disequilibria among loci, therefore invalidating this test of detection of selection. We address this issue by monitoring the genetic differentiation of five microsatellite loci (FST) and nine quantitative traits (QST) in experimental metapopulations of the predominantly selfing species Arabidopsis thaliana, that evolved during eight generations. Metapopulations differed with respect to population size and selection heterogeneity. In large populations, the genetic differentiation of neutral microsatellites was much larger under heterogeneous selection than under uniform selection. Using simulations, we show that this influence of selection heterogeneity on FST can be attributable to initial linkage disequilibria among loci, creating stronger genetic differentiation of QTL than expected under a simple additive model with no initial linkage. We found no significant differences between FST and QST regardless of selection heterogeneity, despite a demonstrated effect of selection on QST values. Additional data are required to validate the role of mating system and linkage disequilibria in the joint evolution of neutral and selected genetic differentiation, but our results suggest that FST/QST comparisons can be conservative tests to detect selection in selfing species.

Arabidopsis↗