Evolution of predator avoidance in a Daphnia population: evidence from the egg bank

Dormant propagule pools (egg banks) of zooplankton populations that accumulate in sediments provide biological archives of past conditions and enable the investigation of evolutionary changes in populations over relatively long periods of time (many decades). This study examined the egg bank of a Daphnia pulicaria population in a lake that has been stocked annually with rainbow trout (a zooplanktivore) since 1961. Resting eggs from sediments from the 1920s to 2001 (determined by ²¹⁰Pb dating) were hatched and established as isofemale clonal lines. The phototactic behavior (a proxy for vertical migration behavior) of clones was assessed in the presence and absence of fish kairomones to evaluate the hypothesis that the heavy and consistent level of predation imposed by the stocked trout would have selected for clones that are more negatively phototactic. In addition, exposure to kairomones was expected to induce stronger negative phototaxis for all clones relative to control conditions. The behavior of clones derived from resting eggs from the trout era (after 1961) was significantly more negatively phototactic than it was for pre-trout era clones. Kairomone exposure induced a more negative phototactic response in clones from both eras, but the response was much greater for the pre-trout era clones. These results suggest that the consistently high level of predation by trout over several decades has selected for a conservative (less plastic) vertical migration strategy in which Daphnia maintain a deep daytime distribution with or without the presence of chemical cues from fish predators.     

Genetic Analysis of Phototactic Behavior in DROSOPHILA SIMULANS

Phototaxis mazes have been employed to select photopositive and photonegative strains of Drosophila simulans. The results suggest that phototactic behavior in D. simulans, as in other Drosophila species, is a polygenic trait. Hybridization using divergent strains revealed that the genes controlling negative phototactic behavior in D. simulans are autosomal, as opposed to D. melanogaster in which negative phototactic behavior is known to be very strongly sex-linked.     

Genomic Architecture of Rapid Parallel Adaptation to Fresh Water in a Wild Fish

Rapid adaptation to novel environments may drive changes in genomic regions through natural selection. However, the genetic architecture underlying these adaptive changes is still poorly understood. Using population genomic approaches, we investigated the genomic architecture that underlies rapid parallel adaptation of Coilia nasus to fresh water by comparing four freshwater-resident populations with their ancestral anadromous population. Linkage disequilibrium network analysis and population genetic analyses revealed two putative large chromosome inversions on LG6 and LG22, which were enriched for outlier loci and exhibited parallel association with freshwater adaptation. Drastic frequency shifts and elevated genetic differentiation were observed for the two chromosome inversions among populations, suggesting that both inversions would undergo divergent selection between anadromous and resident ecotypes. Enrichment analysis of genes within chromosome inversions showed significant enrichment of genes involved in metabolic process, immunoregulation, growth, maturation, osmoregulation, and so forth, which probably underlay differences in morphology, physiology and behavior between the anadromous and freshwater-resident forms. The availability of beneficial standing genetic variation, large optimum shift between marine and freshwater habitats, and high efficiency of selection with large population size could lead to the observed rapid parallel adaptive genomic change. We propose that chromosomal inversions might have played an important role during the evolution of rapid parallel ecological divergence in the face of environmental heterogeneity in C. nasus. Our study provides insights into the genomic basis of rapid adaptation of complex traits in novel habitats and highlights the importance of structural genomic variants in analyses of ecological adaptation.     

Chromosome Studies in Wild Populations of D. MELANOGASTER

Chromosome studies of wild D. melanogaster populations from Missouri, Mississippi, Louisiana and Texas uncovered 58 inversions. Six were common and cosmopolitan; 52 were new, rare and generally endemic. In one of two Missouri populations tested, structurally heterozygous females carried significantly more sperm at capture than did the homozygotes. In both populations comparisons of wild sperms with the females carrying them indicated significant positive assortative mating and an excess production of homozygotes among the F₁ progeny. Wild females structurally heterozygous in up to three major autosomal arms showed no associated nondisjunctional egg lethality; those heterozygous in all four arms produced from 0% to 24% dead eggs, suggesting the presence of intrapopulational gene modifiers of meiosis. Texas populations supported on windfall citrus fruit showed a slight but significant difference in inversion frequencies between flies breeding on oranges and those breeding on grapefruit. Within these populations inversions were not distributed at random among individuals; rather there was an observed excess of individuals carrying intermediate numbers, and a deficiency of those carrying very few or very many inversions. While there was no significant linkage disequilibrium associated with this central tendency, there was a significant interchromosomal interaction: flies carrying inversions in chromosome 2 tended not to carry them in chromosome 3, and vice versa.     

EVOLUTIONARY POTENTIAL AND LOCAL GENETIC DIFFERENTIATION IN A PHENOTYPICALLY PLASTIC TRAIT OF A CYCLICAL PARTHENOGEN,DAPHNIA MAGNA

The frequent use of neutral markers to quantify genetic variation in natural populations emphasizes the role of stochastic events in explaining genetic differentiation between populations. Complementary studies on ecologically relevant traits are needed to assess the role of natural selection acting on adaptive variation in the development of local genetic differentiation. To test the hypothesis of local adaptation in the cyclical parthenogenetic species Daphnia magna, the phototactic behavior in the absence and presence of fish kairomone was assayed for 30 clones derived from resting eggs isolated from three habitats characterized by a different predation pressure by fish. Clones derived from populations in which fish are present were, on average, more negatively phototactic in and more responsive to the presence of fish kairomone than clones derived from a fishless habitat. In addition, the results show a high genetic diversity for the traits studied in all three gene pools investigated, indicating a high potential for microevolutionary changes in behavior of these Daphnia populations in the face of changes in predation pressure. The results of the present study indicate that working with large samples at the expense of having less precise estimates of genotypic values for specific genotypes may result in a loss of information with regard to the evolutionary potential of local populations.     

The Genetic Basis of Host Preference and Resting Behavior in the Major African Malaria Vector,Anopheles arabiensis

Malaria transmission is dependent on the propensity of Anopheles mosquitoes to bite humans (anthropophily) instead of other dead end hosts. Recent increases in the usage of Long Lasting Insecticide Treated Nets (LLINs) in Africa have been associated with reductions in highly anthropophilic and endophilic vectors such as Anopheles gambiae s.s., leaving species with a broader host range, such as Anopheles arabiensis, as the most prominent remaining source of transmission in many settings. An. arabiensis appears to be more of a generalist in terms of its host choice and resting behavior, which may be due to phenotypic plasticity and/or segregating allelic variation. To investigate the genetic basis of host choice and resting behavior in An. arabiensis we sequenced the genomes of 23 human-fed and 25 cattle-fed mosquitoes collected both in-doors and out-doors in the Kilombero Valley, Tanzania. We identified a total of 4,820,851 SNPs, which were used to conduct the first genome-wide estimates of “SNP heritability” for host choice and resting behavior in this species. A genetic component was detected for host choice (human vs cow fed; permuted P = 0.002), but there was no evidence of a genetic component for resting behavior (indoors versus outside; permuted P = 0.465). A principal component analysis (PCA) segregated individuals based on genomic variation into three groups which were characterized by differences at the 2Rb and/or 3Ra paracentromeric chromosome inversions. There was a non-random distribution of cattle-fed mosquitoes between the PCA clusters, suggesting that alleles linked to the 2Rb and/or 3Ra inversions may influence host choice. Using a novel inversion genotyping assay, we detected a significant enrichment of the standard arrangement (non-inverted) of 3Ra among cattle-fed mosquitoes (N = 129) versus all non-cattle-fed individuals (N = 234; χ², p = 0.007). Thus, tracking the frequency of the 3Ra in An. arabiensis populations may be of use to infer selection on host choice behavior within these vector populations; possibly in response to vector control. Controlled host-choice assays are needed to discern whether the observed genetic component has a direct relationship with innate host preference. A better understanding of the genetic basis for host feeding behavior in An. arabiensis may also open avenues for novel vector control strategies based on driving genes for zoophily into wild mosquito populations.     

Larval photoresponses of the polyclad flatworm Maritigrella crozieri (Platyhelminthes,Polycladida) (Hyman)

Phototaxis by larvae of the flatworm Maritigrella crozieri was used to determine spectral sensitivity, the ontogeny of the phototactic pattern, and the lowest light intensity to induce a directional response (intensity threshold). Adult M. crozieri live in shallow water with the tunicate Ecteinascidia turbinata and have a planktonic larval phase lasting longer than 3 weeks. The primary spectral sensitivity maximum was at 500 nm, which is probably an adaptation to the spectrum available underwater at twilight. The phototactic threshold changed with age, as dark-adapted, 1-week-old larvae had a threshold (1.84×10¹⁷ photons m⁻² s⁻¹) an order of magnitude higher than that of 3-week-old larvae. Flatworm larvae are relatively insensitive to light as compared to other invertebrate larvae. Young larvae were positively phototactic at high light intensities and negative at low, a pattern typical of a predator avoidance shadow response. In contrast, older larvae were only positively phototactic, which would be useful for transport to shallow-water adult habitats.     

The Genetic Content of Chromosomal Inversions across a Wide Latitudinal Gradient

There is increasing evidence regarding the role of chromosomal inversions in relevant biological processes such as local adaptation and speciation. A classic example of the adaptive role of chromosomal polymorphisms is given by the clines of inversion frequencies in Drosophila subobscura, repeatable across continents. Nevertheless, not much is known about the molecular variation associated with these polymorphisms. We characterized the genetic content of ca. 600 individuals from nine European populations following a latitudinal gradient by analysing 19 microsatellite loci from two autosomes (J and U) and the sex chromosome (A), taking into account their chromosomal inversions. Our results clearly demonstrate the molecular genetic uniformity within a given chromosomal inversion across a large latitudinal gradient, particularly from Groningen (Netherlands) in the north to Málaga (Spain) in the south, experiencing highly diverse environmental conditions. This low genetic differentiation within the same gene arrangement across the nine European populations is consistent with the local adaptation hypothesis for th evolutionof chromosomal polymorphisms. We also show the effective role of chromosomal inversions in maintaining different genetic pools within these inverted genomic regions even in the presence of high gene flow. Inversions represent thus an important barrier to gene flux and can help maintain specific allelic combinations with positive effects on fitness. Consistent patterns of microsatellite allele-inversion linkage disequilibrium particularly in loci within inversions were also observed. Finally, we identified areas within inversions presenting clinal variation that might be under selection.     

thyA as a Selection Marker in Construction of Food-Grade Host-Vector and Integration Systems for Streptococcus thermophilus

We constructed food-grade host-vector and integration systems for Streptococcus thermophilus by using a thymidylate synthase gene (thyA) as the selection marker. Two thyA genes, thyA_St and thyA_Lb, were cloned from S. thermophilus and Lactobacillus delbrueckii subsp. bulgaricus, respectively. Thymidine-requiring mutants of S. thermophilus were obtained after successive cultures in the presence of trimethoprim, and one of them, TM1-1, was used as the host. Food-grade vectors were constructed by using either thyA_St or thyA_Lb as the selection marker. Transformants of TM1-1 created by using these vectors were selected for thymidine autotrophy as efficiently as for erythromycin resistance. By using the host-vector system developed in this way, a foreign amylase gene (amyA) was expressed in TM1-1 and was also integrated into the chromosome by use of a temperature-sensitive integration vector constructed with thyA_Lb as the selection marker via a double-crossover event. The results obtained show that thyA is an efficient and safe selection marker for S. thermophilus that is suitable for food applications.     

Expression of Immunoglobulin Receptors with Distinctive Features Indicating Antigen Selection by Marginal Zone B Cells from Human Spleen

Marginal zone (MZ) B cells, identified as surface (s)IgM^highsIgD^lowCD23^low/−CD21⁺CD38⁻ B cells, were purified from human spleens, and the features of their V(D)J gene rearrangements were investigated and compared with those of germinal center (GC), follicular mantle (FM) and switched memory (SM) B cells. Most MZ B cells were CD27⁺ and exhibited somatic hypermutations (SHM), although to a lower extent than SM B cells. Moreover, among MZ B-cell rearrangements, recurrent sequences were observed, some of which displayed intraclonal diversification. The same diversifying sequences were detected in very low numbers in GC and FM B cells and only when a highly sensitive, gene-specific polymerase chain reaction was used. This result indicates that MZ B cells could expand and diversify in situ and also suggested the presence of a number of activation-induced cytidine deaminase (AID)-expressing B cells in the MZ. The notion of antigen-driven expansion/selection in situ is further supported by the VH CDR3 features of MZ B cells with highly conserved amino acids at specific positions and by the finding of shared (“stereotyped”) sequences in two different spleens. Collectively, the data are consistent with the notion that MZ B cells are a special subset selected by in situ antigenic stimuli.     

How 5000 independent rowers coordinate their strokes in order to row into the sunlight: Phototaxis in the multicellular green alga <Emphasis Type="Italic">Volvox</Emphasis>

Background

The evolution of multicellular motile organisms from unicellular ancestors required the utilization of previously evolved tactic behavior in a multicellular context. Volvocine green algae are uniquely suited for studying tactic responses during the transition to multicellularity because they range in complexity from unicellular to multicellular genera. Phototactic responses are essential for these flagellates because they need to orientate themselves to receive sufficient light for photosynthesis, but how does a multicellular organism accomplish phototaxis without any known direct communication among cells? Several aspects of the photoresponse have previously been analyzed in volvocine algae, particularly in the unicellular alga Chlamydomonas.

Results

In this study, the phototactic behavior in the spheroidal, multicellular volvocine green alga Volvox rousseletii (Volvocales, Chlorophyta) was analyzed. In response to light stimuli, not only did the flagella waveform and beat frequency change, but the effective stroke was reversed. Moreover, there was a photoresponse gradient from the anterior to the posterior pole of the spheroid, and only cells of the anterior hemisphere showed an effective response. The latter caused a reverse of the fluid flow that was confined to the anterior hemisphere. The responsiveness to light is consistent with an anterior-to-posterior size gradient of eyespots. At the posterior pole, the eyespots are tiny or absent, making the corresponding cells appear to be blind. Pulsed light stimulation of an immobilized spheroid was used to simulate the light fluctuation experienced by a rotating spheroid during phototaxis. The results demonstrated that in free-swimming spheroids, only those cells of the anterior hemisphere that face toward the light source reverse the beating direction in the presence of illumination; this behavior results in phototactic turning. Moreover, positive phototaxis is facilitated by gravitational forces. Under our conditions, V. rousseletii spheroids showed no negative phototaxis.

Conclusions

On the basis of our results, we developed a mechanistic model that predicts the phototactic behavior in V. rousseletii. The model involves photoresponses, periodically changing light conditions, morphological polarity, rotation of the spheroid, two modes of flagellar beating, and the impact of gravity. Our results also indicate how recently evolved multicellular organisms adapted the phototactic capabilities of their unicellular ancestors to multicellular life.

     

Genic VERSUS Chromosomal Variation in Natural Populations of DROSOPHILA SUBOBSCURA

Gametic frequencies in one mainland and one island population of D. subobscura were obtained by means of extracting wild chromosomes and subsequently analyzing them for inversions and allozymes. The high degree of cytological heterogeneity which characterizes these populations is not reflected in the genetic data. Two cases of non-random association were observed among eighteen pair-wise comparisons involving gene alleles and inversions to which the locus is linked. In both cases exchange of alleles at the locus is completely suppressed by the inversions. Four cases of linkage disequilibrium were detected among eighteen pairs of loci; two of them could best be explained as transient associations generated by random drift. The results suggest that disequilibria among enzyme loci are not widespread in natural populations—Populations with a lower degree of chromosomal variation are genetically as variable as populations with a higher degree of chromosomal variation. This observation does not support the hypothesis that selection in marginal homokaryotypic populations is for specialized homozygous genotypes.     

Tubby-tagged balancers for the Drosophila X and second chromosomes

We generated FM7a and CyO balancer chromosomes bearing a Tubby¹ (Tb¹) dominant transgene. Flies heterozygous for these FM7a and CyO derivatives exhibit a phenotype undistinguishable from that elicited by the Tb¹ mutation associated with the TM6B balancer. We tested two of these Tb-bearing balancers (FM7-TbA and CyO-TbA) for more than 30 generations and found that the Tb¹ transgene they carry is stable. Thus, these new Tb-tagged balancers are particularly useful for balancing lethal mutations and distinguish homozygous mutant larvae from their heterozygous siblings.     

The Genetics of DROSOPHILA SUBOBSCURA Populations. V. a Study of Linkage Disequilibrium in Natural Populations between Genes and Inversions of the E Chromosome

The genetics of Hk and Est-9 complex gene have been studied in Drosophila subobscura. While Hk alleles mendelize normally, Est-9 is a complex locus consisting of several very closely linked genes with active and silent alleles. Both genes are located on chromosome E; a detailed genetic map was constructed with the help of visible markers and inversions. Both Hk and Est-9 are included in or are very near to inversions of the E chromosome. While Hk does not show linkage disequilibrium either with Est-9 or inversions, Est-9 does show disequilibrium in two ways: both with inversions and between different Est-9 genes. All natural populations studied show the same kind of association between Est-9 gene combinations and inversions. It is argued that these results are better explained by selection than by neutrality.     

The advantage of factorial mating under selection is uncovered by deterministically predicted rates of inbreeding

Rates of inbreeding (ΔF) in selected populations were predicted using the framework of long-term genetic contributions and validated against stochastic simulations. Deterministic predictions decomposed ΔF into four components due to: finite population size, directional selection, covariance of genetic contribution of mates, and deviation of variance of family size from that expected from a Poisson distribution. Factorial (FM) and hierarchical (HM) mating systems were compared under mass and sib-index selection. Prediction errors were in most cases for ΔF less than 10% and for rate of gain less than 5%. ΔF was higher with index than mass selection. ΔF was lower with FM than HM in all cases except random selection. FM reduced the variance of the average breeding value of the mates of an individual. This reduced the impact of the covariance of contributions of mates on ΔF. Thus, contributions of mates were less correlated with FM than HM, causing smaller deviations of converged contributions from the optimum contributions. With index selection, FM also caused a smaller variance of number of offspring selected from each parent. This reduced variance of family size reduced ΔF further. FM increases the flexibility in breeding schemes for achieving the optimum genetic contributions.     

Association between polymorphism in the FTO gene and growth and carcass traits in pig crosses

Background

Independent studies have shown that several single nucleotide polymorphisms (SNP) in the human FTO (fat mass and obesity associated) gene are associated with obesity. SNP have also been identified in the pig FTO gene, among which some are associated with selected fat-deposition traits in F₂ crosses and commercial populations. In this study, using both commercial pig populations and an experimental Meishan × Pietrain F₂ population, we have investigated the association between one FTO SNP and several growth and carcass traits. Association analyses were performed with the FTO polymorphism either alone or in combination with polymorphisms in flanking loci.

Methods

SNP ({"type":"entrez-nucleotide","attrs":{"text":"FM244720","term_id":"209570741","term_text":"FM244720"}}FM244720:g.400C>G) in exon 3 of porcine FTO was genotyped by PCR-RFLP and tested for associations with some growth, carcass and fat-related traits. Proportions of genetic variance of four pig chromosome 6 genes (FTO, RYR1, LIPE and TGFB1) on selected traits were evaluated using single- and multi-locus models.

Results

Linkage analysis placed FTO on the p arm of pig chromosome 6, approximately 22 cM from RYR1. In the commercial populations, allele C of the FTO SNP was significantly associated with back fat depth and allele G with muscling traits. In the Meishan × Pietrain F₂ pigs, heterozygotes with allele C from the Pietrain sows and allele G from the Meishan boar were more significantly associated with fat-related traits compared to homozygotes with allele G from the Pietrain and allele G from the Meishan breed. In single- and multi-locus models, genes RYR1, TGFB1 and FTO showed high associations. The contribution in genetic variance from the polymorphism in the FTO gene was highest for back fat depth, meat area on the musculus longissimus lumborum et thoracis tissues and metabolite glucose-6-phosphate dehydrogenase.

Conclusions

Our results show that in pig, FTO influences back fat depth in the commercial populations, while in the Meishan × Pietrain F₂ pigs with a CG genotype, heterosis occurs for several fat-related traits.     

Chromosomal polymorphism in the populations of malaria vector mosquito Anopheles messeae at the south of Russian Plain

Zonal features of the geographic distribution of chromosomal inversions in the populations of An. messae at the south of the Russian Plain were examined. The An. messae was identified based on morphological characters and cytogenetically. The chromosomal inversions identified in populations of An. messae comprised XL₁, XL₄, 2R₁, 3R₁, and 3L₁. Inversion XL₄ was endemic and found with low frequency (2%) in Penza oblast and Moscow oblast. Based on the population karyotype structure similarity, on the territory of Russian Plain, three zones (southwestern, southeastern, and central) were identified. Central zone was characterized by higher levels of inversion polymorphism in all chromosomes, except arm 3L. In the two southern zones, high frequency of the XL₀ inversion, along with complete absence of homo- and heterozygotes for the 2R₁ inversion, and high proportion of the individuals with inversion 3L₁ was observed. Specific feature of southeastern zone was the increased frequency of hetero- and homozygotes for the 3R₁ and 3L₁ inversions. Zonal differences reflected adaptive character of chromosomal polymorphism and pointed to hierarchic organization of the species population structure.     

Inversion polymorphism in two Serbian natural populations of Drosophila subobscura: Analysis of long-term changes

To study whether inversions (or arrangements) by themselves or karyotypes are the main global warming adaptive target of natural selection, two Drosophila subobscura Serbian populations (Apatin and Petnica) were re-analyzed using different statistical approaches. Both populations were sampled in an approximately 15 years period: Apatin in 1994 and 2008 + 2009 and Petnica in 1995 and 2010. For all chromosomes, the four colleptions studied were in Hardy-Weinberg equilibrium. Thus, it seems that inversions (or arrangements) combined at random to constitute populations’ karyotypes. However, there were differences in karyotypic frequencies along the years, although they were significant only for Apatin population. It is possible to conclude that inversions (or arrangements) are likely the target of natural selection, because they presented long-term changes, but combine at random to generate the corresponding karyotypic combinations. As a consequence, the frequencies of karyotypes also change along time.     

Evaluating the phototactic behavior responses of the diamondback moth,Plutella xylostella,to some different wavelength LED lights in laboratory and field

Light traps equipped with light emitting diodes (LEDs) have been applied to manage some phototactic insect pests. The diamondback moth, Plutella xylostella, is a cosmopolitan insect pest to be seriously harmful to many cruciferous plants. The present research focused on evaluating the phototactic behavior responses of the moths to several wavelengths and photon flux densities of LED lights under laboratory and field conditions. The results from the laboratory showed that the highest phototactic behavior responses of P. xylostella moths were recorded for UV (380 nm) LED light under 1.5 µmol m⁻² s⁻¹ and 2.5 µmol m⁻² s⁻¹. The moths were more attracted to light traps equipped with 4 LEDs and 6 LEDs of 380 nm, respectively, between 20:00 and 22:00 than the other groups and night times in the field. The finding from the field was consistent with the results from the laboratory. We found that the 380 nm LED light results in the strongest attraction rate of the moths by 92.4 % and the moths caught in light trap with the UV LEDs was 1.62 times more than that with a black light. These data clearly demonstrate that P. xylostella moths have a high sensitivity to 380 nm, therefore, a 380 nm LED light trap could be useful for monitoring and controlling the moths.