Sexual Generation Research Articles

The reproductive system of the male and oviparous female of a model organism-the pea aphid, Acyrthosiphon pisum (Hemiptera, Aphididae).

The structure of the reproductive system of the sexual generation—males and oviparous females—of the pea aphid Acyrthosiphon pisum (Harris) (Hemiptera, Aphididae), a serious pest of cultivated plants of Fabaceae, was investigated. For the first time we describe the morphology, histology and ultrastructure of the reproductive system in both morphs of the sexual generation of aphids within one species, using light and fluorescent microscopy, as well as transmission and scanning electron microscopy. The results revealed that males have testes composed of three follicles fused by the upper ends of the vasa efferentia, the vasa deferentia run independently, the accessory glands are asymmetric and the ejaculatory duct shortened. Oviparous females have ovaries composed of seven ovarioles each. The lateral oviducts join to a short common oviduct connected with the unpaired spermatheca and paired accessory glands. Yolky eggs with an aggregation of symbiotic bacteria at the posterior pole are produced. Histologically, the components of genital tracts are broadly similar: the epithelial cells of the walls of the vasa deferentia and accessory glands of the male and oviparous female have secretory functions which correlate with the age of the studied morphs. We also found symbiotic bacteria within the vasa deferentia epithelial cells in males and within the cells of the lateral oviducts of females. Because the pea aphid is listed among the 14 species that are of the greatest economic importance, our results will be useful for managing aphid populations, protecting plants and ensuring global food security.

Heterogony in Cycloneuroterus (Hymenoptera: Cynipidae: Cynipini) From Rearing Experiments and DNA Barcoding.

Heterogony was confirmed in the cynipid genus Cycloneuroterus Melika and Tang in rearing experiments with DNA barcoding. These experiments involved Cycloneuroterus gilvus Tang and Melika, which was previously only described from the sexual generation adult. The first rearing experiment was conducted using unidentified asexual generation females collected from Quercus gilva Blume, and gall formation by the sexual generation offspring was confirmed on folded or unfolded young leaves of Q. gilva. The second experiment was conducted using sexual generation males and females reared from the leaf galls collected from Q. gilva, and gall formation by the asexual generation offspring was observed on leaves of Q. gilva. Based on the morphological features of the sexual generation adults and galls, this species was identified as C. gilvus. The species identity of wasp specimens of sexual and asexual generations used in the rearing experiments was cross-checked using DNA barcoding with the partial sequences of the cytochrome c oxidase subunit I (COI) region (658 bp). The asexual generation adult and gall of C. gilvus are described based on these results. The importance of ‘closing the life cycle,’ in this case a demonstration of heterogony, in oak gall wasps (Cynipini) is discussed.

Molecular identification of Andricus species (Hymenoptera: Cynipidae) inducing various oak galls in Central Zagros of Iran

This study uses an integrated approach to address the taxonomic status of six different and problematic oak galls and their inducing wasps sampled from two sites in the Central Zagros Mountains (Lorestan province) in western Iran. Our aim was to establish whether morphologically similar but different galls are induced by the same or distinct gall-inducers. The gall wasp specimens were identified morphologically to species level, and their genomic DNA was extracted. We used PCR and Sanger sequencing to amplify three fragments comprising cytochrome oxidase subunit I (COI), cytochrome b (cytB), and a multi-gene fragment of ribosomal DNA (rDNA) including partial 5.8S, complete internal transcribed spacer 2 (ITS2), and partial 28S rRNA. We found that a pair of structurally similar but differently coloured galls are induced by the sexual generation of Andricus grossulariae, while another similar pair are induced by the asexual generation of A. sternlichti. In contrast, we found that two similar galls that differ in some structural details and in developmental phenology are induced by two closely related but different gall wasps; one is the sexual generation of A. cecconii, while the second is a new but closely related sexual generation Andricus sp.

Effect of Neuroterus quercusbaccarum (L.) galls on physiological and biochemical response of Quercus robur leaves.

Gall formation is associated with multiple changes in plant cells, which still requires a better understanding. In this study, galls caused by sexual generation (♀♂) of Neuroterus quercusbaccarum (L.) (Hymenoptera: Cynipidae) on pedunculate oak trees (Quercus robur L.) were used as a model. Cytoplasmic membrane condition, concentration of hydrogen peroxide (H2O2), the activity of antioxidant enzymes and amino acid decarboxylase as well as chlorophyll fluorescence parameters were determined. Changes in physiological and biochemical parameters were analyzed in foliar tissues with galls and gall tissues themselves and compared to control. The presence of galls on oak leaves caused an increase of lipid peroxidation level. A significant decline in H2O2 and TBARS content with the reduction of guaiacol peroxidase (GPX) and ascorbate peroxidase (APX) activity were observed in gall tissues. The activity amino acid decarboxylase, i.e., LDC, ODC and TyDC varied between samples, which may affect the content of amino acids. The presence of N. quercusbaccarum galls caused an insignificant increase of the chlorophylls, carotenoids and anthocyanin contents, while the content of pigments and their ratios in gall tissues was extremely low. Moreover, photosynthetic parameters (F0, Fm, Fv/Fm, Y, qP) were significantly decreased. Data generated in this study indicate that the development of N. quercusbaccarum galls on pedunculate oak leaves has a negative effect on host plant related to the disruption of cell membrane integrity, disturbance of photosynthesis and reduction of the antioxidant potential of the host plant.

Nuclear arms races: Experimental evolution for mating success in the mushroom-forming fungus Schizophyllum commune.

When many gametes compete to fertilize a limited number of compatible gametes, sexual selection will favour traits that increase competitive success during mating. In animals and plants, sperm and pollen competition have yielded many interesting adaptations for improved mating success. In fungi, similar processes have not been shown directly yet. We test the hypothesis that sexual selection can increase competitive fitness during mating, using experimental evolution in the mushroom-forming fungus Schizophyllum commune (Basidiomycota). Mating in mushroom fungi occurs by donation of nuclei to a mycelium. These fertilizing ‘male’ nuclei migrate through the receiving ‘female’ mycelium. In our setup, an evolving population of nuclei was serially mated with a non-evolving female mycelium for 20 sexual generations. From the twelve tested evolved lines, four had increased and one had decreased fitness relative to an unevolved competitor. Even though only two of those five remained significant after correcting for multiple comparisons, for all five lines we found a correlation between the efficiency with which the female mycelium is accessed and fitness, providing additional circumstantial evidence for fitness change in those five lines. In two lines, fitness change was also accompanied by increased spore production. The one line with net reduced competitive fitness had increased spore production, but reduced fertilisation efficiency. We did not find trade-offs between male reproductive success and other fitness components. We compare these findings with examples of sperm and pollen competition and show that many similarities between these systems and nuclear competition in mushrooms exist.

New species of Dryocosmus Giraud gallwasps from California (Hymenoptera: Cynipidae: Cynipini) galling Chrysolepis Hjelmq. (Fagaceae).

Four species of Dryocosmus cynipid gallwasps are now known to induce galls on Chrysolepis in California and Oregon. Two new species, Dryocosmus demartinii Melika, Nicholls Stone and Dryocosmus juliae Melika, Nicholls Stone are described. Males of the sexual generation of D. rileypokei plus adults of the asexual generation of this species are both described for the first time. A detailed description of D. castanopsidis is given for the first time. In addition to descriptions we provide diagnoses and information on biology and host associations for all four species. All the newly described taxa are supported by morphological and molecular data.

Three Species of Andricus Hartig (Hymenoptera: Cynipidae) Inducing Similar Galls in East Asia, with Description of a New Species and the Asexual Generation of A. Pseudocurvator, and Redescription of A. Moriokae

The sexual generation of Andricus wuhanensis Ide, Abe, Su and Zhu, new species is described from Mainland China. The gall of this species is similar to the sexual generation galls of A. pseudocurvator Tang and Melika and A. moriokae Monzen, but the adult wasps of the three species differ morphologically. The asexual generation gall of A. pseudocurvator was previously reported as that of A. moriokae from Japan, but the adult morphology has not yet been described; therefore, we here describe the asexual generation adult of A. pseudocurvator for the first time. Andricus pseudocurvator is here first recorded from Mainland China and Japan.

The oak gall wasps of Israel (Hymenoptera, Cynipidae, Cynipini)-diversity, distribution and life history.

Oak-galling wasps (Hymenoptera: Cynipidae, Cynipini) have been studied in Israel only superficially so far, despite interest in the Israeli fauna given its location at the southern edge of distribution range of the wasps and their oak hosts. We provide the first comprehensive review of the Israeli fauna of oak cynipids based on an extensive survey of the five naturally occurring oak species in Israel. We report 53 cynipid species, 37 of which are reported from Israel for the first time, 10 are currently known only from this country, and 9 are undescribed. With 27 species, Andricus is by far the biggest genus in Israel, followed by Cerroneuroterus, Neuroterus Plagiotrochus, and Pseudoneuroterus with 4 species each. Andricus megalucidus is synonymized under A. cecconii and Cerroneuroterus cerrifloralis is synonymized under C. lanuginosus. The sexual generations of Andricus cecconii, A. coriariformis, A. coriarius, A. miriami, Cerroneuroterus lanuginosus and Pseudoneuroterus macropterus are reported here for the first time. We recorded 65 gall types, with bud galls being the most common and conspicuous, followed by leaf and catkin galls. We provide illustrated keys for all Israeli species based on their galls, as well as information on host associations, life history, phenology and distribution patterns where available. 21 cynipid species are associated with Quercus ithaburensis and 27 species are associated with Q. boissieri, whereas Q. libani, Q. calliprinos and Q. cerris host much fewer species. Most species are currently known from either their sexual or asexual generation while only 24% of them are known from both. Mount Hermon was found to be an important hotspot, hosting about half of the Israeli cynipid fauna, and species richness generally declines from the Golan Heights southwards to the Judean Mountains. Cynipid species that are associated with oaks of sections Cerris and Quercus in Europe are mostly associated with the same sections in Israel.

Effects of Transgenerational Plasticity on Morphological and Physiological Properties of Stoloniferous Herb Centella asiatica Subjected to High/Low Light.

Environmentally induced transgenerational plasticity can increase success of progeny and thereby be adaptive if progeny experiences the similarly parental environment. The ecological and evolutionary significance of transgenerational plasticity in plant has been studied mainly in the context of sexual generations. A pot experiment using the stoloniferous herb Centella asiatica was conducted to investigate the effects of high/low light treatment experienced by parental ramets (F0 generation) on morphological and physiological properties of offspring ramets (F2 generation) as well as growth performance. Light environment experienced by parental ramets (F0 generation) significantly influenced petiole length, specific petiole length, internode length of stolon, leaf area, specific leaf area (SLA), leaf nitrogen and chlorophyll contents, potential maximum net photosynthetic rate (Pmax) in offspring ramets subjected to parental or non-parental environments even after they were detached from the parental ramets. Potential maximum net photosynthetic rate (Pmax) of offspring ramets (F2 generation) from parental ramets (F0 generation) subjected to low light treatment was significantly greater than that of offspring ramets (F2 generation) from parental ramets (F0 generation) subjected to high light treatment. Potential maximum net photosynthetic rate (Pmax) of offspring ramets (F2 generation) subjected to parental light environment was greater than that of offspring ramets (F2 generation) subjected to non-parental light environment. The greatest biomass accumulation and total stolon length were observed in offspring ramets (F2 generation) subjected to low light treatment as parental ramets (F0 generation) experienced. When parental ramets (F0 generation) were subjected to low light treatment, biomass accumulation and total stolon length of offspring ramets (F2 generation) experiencing parental light environment were significantly greater than those of offspring ramets (F2 generation) experiencing non-parental light environment. Opposite pattern was observed in offspring ramets (F2 generation) from parental ramets subjected to high light treatment. Our work provides evidence that transgenerational plasticity through both morphological and physiological flexibility was triggered across vegetative generations for stoloniferous herb C. asiatica subjected to high/low light treatment. The transgenerational plasticity can allow offspring ramets to present adaptive phenotype early without lag time in response to the current environment. Thus, it is very important for clonal plants in adapting temporally and spatially heterogeneous habitats.

Host-mediated RNA interference targeting acuticular protein gene impaired fecundity inthe green peach aphid Myzus persicae.

The green peach aphid (Myzus persicae) is a devastating sap-sucking insect pest that damages many host plants worldwide and causes billions of dollars of crop losses. Induction of RNA interference (RNAi) through oral feeding of small interfering RNA (siRNA) has been demonstrated in aphids. Therefore, host-mediated delivery of double-stranded RNA (dsRNA) specific to vital structural genes of aphids has been envisaged as a tool for the development of resistance against this aphid species. Cuticular protein (CP) senses seasonal photoperiodism and drives a shift from clonal to sexual generation in aphids. Thus, attenuation of CP gene expression is likely to result in a different reproductive orientation in aphids and thereby affect their fecundity. A gene encoding CP in M. persicae has been targeted for RNAi-mediated knockdown. Transgenic Arabidopsis expressing dsRNA homologous to the MyCP gene was developed. The dsRNA-transgenics produced gene-specific siRNAs fed by aphids infesting the transgenics. A reverse transcription-quantitative polymerase chain reaction (RT-qPCR) study revealed an attenuated level of transcripts of the CP gene in aphid nymphs reared on the transgenic plants. Decreased expression of the CP gene resulted in a noticeable decline in aphid fecundity on the transgenic Arabidopsis plants. Increasing genetic resistance is the only sustainable way of minimizing the use of toxic agrochemicals to protect plants. Host-mediated RNAi of important insect genes has been proposed as a potential avenue for developing crop resistance against insect pests. This study demonstrated the potential of MyCP dsRNA in developing RNAi-based resistance to M. persicae. RNAi-mediated resistance is expected to be more durable compared with other transgenic strategies. © 2018 Society of Chemical Industry.

‘Closing the Life Cycle’ of Andricus quercuslanigera (Hymenoptera: Cynipidae)

While the alternation of asexually and sexually reproducing generations is common among the oak gall wasps (Hymenoptera: Cynipidae: Cynipini), it has been hypothesized that the diversity of taxa displaying this unique life cycle is underestimated because either 1) the alternative generation has not yet been described or 2) each generation is currently described as two distinct species and should be collapsed into one heterogonic organism (referred to as ‘closing the life cycle’). Through field observations, experimental rearing, morphological identification, laboratory behavioral assays, and genetic analysis, we demonstrate heterogony in the cynipid species Andricus quercuslanigera (Ashmead 1881) (Hymenoptera: Cynipidae), which was previously only described from the asexual generation. We confirm that the asexual generation, which develops in ‘fuzzy’ galls on the central vein on the underside of leaves on live oaks in southeast Texas, Quercus virginiana, represents only one generation in a bivoltine life cycle that alternates with a newly discovered sexual generation that develops in galls on catkins on the same host. Our study highlights the need for detailed inspections of the life cycles of unisexual gall wasp species and we discuss the closure of the A. quercuslanigera life cycle in light of recent advances in the study of the ecology and evolution of heterogony in the Cynipidae.

Diversity and distribution of Wolbachia in relation to geography, host plant affiliation and life cycle of a heterogonic gall wasp

BackgroundThe maternally inherited endosymbiont Wolbachia is widespread in arthropods and nematodes and can play an important role in the ecology and evolution of its host through reproductive manipulation. Here, we survey Wolbachia in Belonocnema treatae, a widely distributed North American cynipid gall forming wasp that exhibits regional host specialization on three species of oaks and alternation of sexually and asexually reproducing generations. We investigated whether patterns of Wolbachia infection and diversity in B. treatae are associated with the insect’s geographic distribution, host plant association, life cycle, and mitochondrial evolutionary history.ResultsScreening of 463 individuals from 23 populations including sexual and asexual generations from all three host plants across the southern U.S. showed an average infection rate of 56% with three common Wolbachia strains: wTre1–3 and an additional rare variant wTre4. Phylogenetic analysis based on wsp showed that these strains are unrelated and likely independently inherited. We found no difference in Wolbachia infection frequency among host plant associated populations or between the asexual and sexual generations, or between males and females of the sexual generation. Partially incomplete Wolbachia transmission rates might explain the occurrence of uninfected individuals. A parallel analysis of the mitochondrial cytochrome oxidase I gene in B. treatae showed high mtDNA haplotype diversity in both infected and uninfected populations suggesting an ancestral infection by Wolbachia as well as a clear split between eastern and western B. treatae mtDNA clades with a sequence divergence of > 6%. The strain wTre1 was present almost exclusively in the western clade while wTre2 and wTre3 occur almost exclusively in eastern populations. In contrast, the same strains co-occur as double-infections in Georgia and triple-infections in two populations in central Florida.ConclusionsThe diversity of Wolbachia across geographically and genetically distinct populations of B. treatae and the co-occurrence of the same strains within three populations highlights the complex infection dynamics in this system. Moreover, the association of distinct Wolbachia strains with mitochondrial haplotypes of its host in populations infected by different Wolbachia strains suggests a potential role of the endosymbiont in reproductive isolation in B. treatae.

The Challenges of Genome-Wide Studies in a Unicellular Eukaryote With Two Nuclear Genomes.

We present here methods to study a eukaryotic microorganism with two nuclear genomes, both originating from the same zygotic genome. Paramecium, like other ciliates, is characterized by nuclear dimorphism, which is the presence of two types of nuclei with distinct organization and functions in the same cytoplasm. The two diploid germline micronuclei (MIC) undergo meiosis and fertilization to transmit the genetic information across sexual generations. The highly polyploid somatic macronucleus (MAC) contains a reduced version of the genome optimized for gene expression. Reproducible programmed DNA elimination of about 30% of the complexity of the 100Mb MIC genome occurs during development of the MAC along with endoreplication to 800 copies. Large regions that contain transposable elements and other repeats are eliminated, and short single copy remnants of transposable elements, which often interrupt coding sequences, are precisely excised to restore functional open reading frames. Genome-wide studies of this process require access to MIC DNA which has long been impossible. The breakthrough with respect to this technical obstacle came with development of a MIC purification protocol involving a critical step of flow cytometry to sort nuclei representing only 0.5% of total genomic DNA. Here, we provide a step-by-step protocol and important tips for purifying nuclei, and present the methods developed for downstream analysis of NGS data.

Do facultative symbionts affect fitness of pea aphids in the sexual generation?

AbstractMany aphids carry one or more facultative symbiotic bacteria which can provide a variety of fitness benefits for their hosts. They have chiefly been investigated in asexually reproducing aphids, with studies of the sexual generation limited to investigation of transmission rates and the potential for sex ratio manipulation. The effects of two facultative symbionts on the mating success of pea aphids, Acyrthosiphon pisum Harris (Hemiptera: Aphididae, Macrosiphini), were investigated in a no‐choice experiment. We compared the fitness of aphids with natural infections of either Regiella insecticola Moran et al. or Hamiltonella defensa Moran et al. with that of aphids from genetically identical cured lines. Female fecundity was unaffected by the presence of facultative symbionts. However, females mated to males cured of H. defensa laid fewer eggs on average than females mated to males carrying H. defensa; in one case the percentage of melanized eggs (eggs that were either not fertilized or in which early death occurred) was also smaller with cured compared to naturally infected males. In addition, males with H. defensa suffered higher mortality during the experiment than cured males. Four of the aphid lines used also hosted Spiroplasma infections, a symbiont previously reported to cause male‐killing in pea aphids. Despite this, three of the Spiroplasma‐infected lines produced males, two at high numbers. We conclude that removing a natural symbiont infection may have a negative fitness effect on male aphids in some aphid clones, whereas sexual females from the same clones are largely unaffected.

REZIM SEKSUALITAS DAN AGAMA SKETSA POLITIK TUBUH PEREMPUAN DALAM ISLAM

This paper examines the various issues of sexuality in Islamic discourse. The discussion in this study attempts to examine the discourse of female sexuality at the beginning of Prophet's prophethood and then evolves over the development of the age to give birth to various inequalities and injustices against women in the sexual realm. There are at least three issues concerning sexual injustice in women; first, the Islamic tradition in the post-Prophetical Jurisprudence which places women as 'male sexual needs ministers' and 'sexual generations'. Second, the tendency of female body consumerism in modern industrial civilization. Third, local traditions in certain cultures still attach stereotypes to women as 'passengers' of male social glory.

REZIM SEKSUALITAS DAN AGAMA SKETSA POLITIK TUBUH PEREMPUAN DALAM ISLAM

This paper examines the various issues of sexuality in Islamic discourse. The discussion in this study attempts to examine the discourse of female sexuality at the beginning of Prophet's prophethood and then evolves over the development of the age to give birth to various inequalities and injustices against women in the sexual realm. There are at least three issues concerning sexual injustice in women; first, the Islamic tradition in the post-Prophetical Jurisprudence which places women as 'male sexual needs ministers' and 'sexual generations'. Second, the tendency of female body consumerism in modern industrial civilization. Third, local traditions in certain cultures still attach stereotypes to women as 'passengers' of male social glory.

Sexual morphs of Pterocomma tremulae Börner, 1940 (Aphididae, Aphidinae) with description of male reproductive system.

Paper presents the first description of the so far unknown sexual generation of Pterocomma tremulae (Aphididae, Aphidinae): oviparous female and alate male. It also provides detailed description of the male reproductive system. Discussion focuses on comparative analysis of male reproductive system with other aphid groups and possible importance of its structure in resolving phylogenetic interrelationships within the genus Pterocomma. A key is provided to the known males of European Pterocomma species.

THE PERCEPTIONS OF AGING AMONG GAY AND LESBIAN OLDER ADULTS: SOCIAL EXCLUSION OR INCLUSION?

Ageism and heterosexism as social exclusion processes can contribute to the social invisibility of gay and lesbian older adults. This contribution wishes to shed a light on the perceptions of aging and on the place of older gay and lesbian people in society, as perceived by gay and lesbian older adults. It is based on the results of semi-structured individual interviews conducted with 22 gay and lesbian older adults aged 60 and over. The participant’s discourses reveal, among others, two elements: a lack of social recognition and visibility of gay and lesbian older adults in society as well as the willingness to assert themselves and take their place. The discussion will propose a reflection of the intersection between sexual orientation, age and generation to examine the dual process of social exclusion, due to ageism and heterosexism, and of social inclusion.

Transgenerational seasonal timer for suppression of sexual morph production in the pea aphid, Acyrthosiphon pisum

Many aphid species switch reproductive modes seasonally, with the sexual generations appearing in autumn. Sexual generations are induced by short days. It has been reported that the appearance of sexual morphs is suppressed by a transgenerational factor (a seasonal timer) over several generations after hatching from overwintered eggs. The present study examined whether the seasonal timer measures the number of days from hatching or the number of generations from hatching using the pea aphid, Acyrthosiphon pisum Harris (Homoptera: Aphididae). Effects of temperature and photoperiod on the seasonal timer were also examined by successive rearing. The ability to produce sexual morphs was strongly suppressed in stem mothers (the foundress generation), and gradually recovered over successive generations produced during a few months. The duration for which the seasonal timer could function depended on the number of days from hatching and temperature, but not on photoperiod or the number of generations from hatching. We thus showed in a single study that the seasonal timer of the pea aphid has all the physiological characteristics shown in separate studies in different aphid species.

Independent life history evolution between generations of bivoltine species: a case study of cyclical parthenogenesis.

Successive generations of bi- and multivoltine species encounter differing biotic and abiotic environments intra-annually. The question of whether selection can independently adjust the relationship between body size and components of reproductive effort within successive generations in response to generation-specific environmental variation is applicable to a diversity of taxa. Herein, we develop a conceptual framework that illustrates increasingly independent life history adjustments between successive generations of taxa exhibiting complex life cycles. We apply this framework to the reproductive biology of the gall-forming insect, Belonocnema treatae (Hymenoptera: Cynipidae). This bivoltine species expresses cyclical parthenogenesis in which alternating sexual and asexual generations develop in different seasons and different environments. We tested the hypotheses that ecological divergence between the alternate generations is accompanied by generational differences in body size, egg size, and egg number and by changes in the relationships between body size and these components of reproductive effort. Increased potential reproductive effort of sexual generation B. treatae is attained by increased body size and egg number (with no trade-off between egg number and egg size) and by a significant increase in the slope of the relationship between body size and potential fecundity. These generation-specific relationships, interpreted in the context of the model framework, suggest that within each generation selection has independently molded the relationships relating body size to potential fecundity and potential reproductive effort in B. treatae. The conceptual framework is broadly applicable to comparisons involving the alternating generations of bi- and multivoltine species.