Unfertilized Females Research Articles

A cytological revisit on parthenogenetic Artemia and the deficiency of a meiosis-specific recombinase DMC1 in the possible transition from bisexuality to parthenogenesis.

Although parthenogenesis is widespread in nature and known to have close relationships with bisexuality, the transitional mechanism is poorly understood. Artemia is an ideal model to address this issue because bisexuality and "contagious" obligate parthenogenesis independently exist in its congeneric members. In the present study, we first performed chromosome spreading and immunofluorescence to compare meiotic processes of Artemia adopting two distinct reproductive ways. The results showed that, unlike conventional meiosis in bisexual Artemia, meiosis II in parthenogenic Artemia is entirely absent and anaphase I is followed by a single mitosis-like equational division. Interspecific comparative transcriptomics showed that two central molecules in homologous recombination (HR), Dmc1 and Rad51, exhibited significantly higher expression in bisexual versus parthenogenetic Artemia. qRT-PCR indicated that the expression of both genes peaked at the early oogenesis and gradually decreased afterward. Knocking-down by RNAi of Dmc1 in unfertilized females of bisexual Artemia resulted in a severe deficiency of homologous chromosome pairing and produced univalents at the middle oogenesis stage, which was similar to that of parthenogenic Artemia, while in contrast, silencing Rad51 led to no significant chromosome morphological change. Our results indicated that Dmc1 is vital for HR in bisexual Artemia, and the deficiency of Dmc1 may be correlated with or even possibly one of core factors in the transition from bisexuality to parthenogenesis.

Deterministic and Stochastic Fractional Order Model for Lesser Date Moth

In this paper, a deterministic and stochastic fractional order model for lesser date moth (LDM) using mating disruption and natural enemies is proposed and analysed. The interaction between LDM larvae, fertilized LDM female, unfertilized LDM female, LDM male and the natural enemy is investigated. In order to clarify the characteristics of the proposed deterministic fractional order model, the analysis of existence, uniqueness, non-negativity and boundedness of the solutions of the proposed fractional-order model are examined. In addition, some sufficient conditions are obtained to ensure the local and global stability of equilibrium points. The occurrence of local bifurcation near the equilibrium points is investigated with the help of Sotomayor’s theorem. Numerical simulations are conducted to illustrate the properties of the proposed fractional order model with respect to the intrinsic growth rate of the LDM larvae, natural enemy’s mortality rate, predation rate, sex pheromone trap parameter, fractional order and environmental noise. The impact of mating disruption on lesser date moth is demonstrated. Also, a numerical approximation method is developed for the proposed stochastic fractional-order model.

Male age and sexual experience affect male mating behavior in the hawthorn spider mite, Amphitetranychus viennensis.

As is reported, in species with first-male sperm precedence, male age and previous sexual experience play crucial roles in male mating behavior. In the hawthorn spider mite, Amphitetranychus viennensis Zacher, previous studies showed that only females that copulated for the first time could achieve fertilization. Based on this, the effects of male age and mating history on male mate choice and male mate competition were investigated. It was confirmed that males could distinguish virgins from fertilized females but they were unable to discriminate between virgins and unfertilized females. Interestingly, the copulation duration of males mated with fertilized females was much shorter than that of males mated with virgins or unfertilized females. Additionally, for male mating choice, males of all ages and more experienced males preferred 5-day-old virgin females, whereas only less experienced males preferred 1-day-old virgin females. In male-male competition, 3-day-old males were more competitive and obtained more copulations compared with others. Copula duration was closely related to male age. Though no significant differences were observed in mating competition between virgin and mated males, copula duration of males in first copulation was the longest and gradually shortened in subsequent copulations. In all, this investigation demonstrated that male age and sexual experience affected male mate choice and male-male competition, leading to further insight into the influences of male age and sexual experience on the reproductive fitness of both sexes.

Phenology of Antler Casting and Occurance of Late-Breeding in Nebraska White-Tailed Deer

White-tailed deer (Odocoileus virginianus) in Nebraska primarily breed in November (stimulated by photoperiod), but this season extends into December and January for unfertilized females and healthy fawns reaching the appropriate breeding weight by winter. Variation in the typical mating period can be attributed to geographic region, skewed sex ratios, and fawns reaching sexual maturity during their first year of birth. During April of 2019, we observed late season breeding by a male white-tailed deer in the central Platte River valley and documented late antler casting for this same late-copulating male. An additional male was observed in April of 2020 still retaining antlers within the region. To understand the irregularity of these observations relative to the central Platte River valley, we documented observations of early and late antler casting utilizing long-term cast antler collection and camera trap data. We were able to establish a baseline for antler casting phenology within this region to denote a shift in average casting timeframe (more than one month later than previously recorded in this region) and document early and the latest known antler casting within the literature. Herein, we detail the variation in antler casting phenology for white-tailed deer in this region, and the latest known occurrence of natural breeding in the state. Our findings help us better understand the life history of the Nebraska white-tailed deer population and can assist state wildlife biologists and property managers as they assess seasonal harvest regulations and adaptively manage the changing resource.

ORGANIZATION AND MONITORING OF FOREST PESTS WITH THE USE OF PHEROMONES IN THE CONDITIONS OF THE STATE ENTERPRISE «ZARICHANSKE FORESTRY»

ORGANIZATION AND MONITORING OF FOREST PESTS WITH THE USE OF PHEROMONES IN THE CONDITIONS OF THE STATE ENTERPRISE «ZARICHANSKE FORESTRY»

Evolutionary Patterns of Sex-Biased Genes in Three Species of Haplodiploid Insects.

Females and males often differ obviously in morphology and behavior, and the differences between sexes are the result of natural selection and/or sexual selection. To a great extent, the differences between the two sexes are the result of differential gene expression. In haplodiploid insects, this phenomenon is obvious, since males develop from unfertilized zygotes and females develop from fertilized zygotes. Whiteflies of the Bemisia tabaci species complex are typical haplodiploid insects, and some species of this complex are important pests of many crops worldwide. Here, we report the transcriptome profiles of males and females in three species of this whitefly complex. Between-species comparisons revealed that non-sex-biased genes display higher variation than male-biased or female-biased genes. Sex-biased genes evolve at a slow rate in protein coding sequences and gene expression and have a pattern of evolution that differs from those of social haplodiploid insects and diploid animals. Genes with high evolutionary rates are more related to non-sex-biased traits—such as nutrition, immune system, and detoxification—than to sex-biased traits, indicating that the evolution of protein coding sequences and gene expression has been mainly driven by non-sex-biased traits.

Nutritional symbionts of a putative vector, Xyleborus bispinatus, of the laurel wilt pathogen of avocado, Raffaelea lauricola

Ambrosia beetles subsist on fungal symbionts that they carry to, and cultivate in, their natal galleries. These symbionts are usually saprobes, but some are phytopathogens. Very few ambrosial symbioses have been studied closely, and little is known about roles that phytopathogenic symbionts play in the life cycles of these beetles. One of the latter symbionts, Raffaelea lauricola, causes laurel wilt of avocado, Persea americana, but its original ambrosia beetle partner, Xyleborus glabratus, plays an uncertain role in this pathosystem. We examined the response of a putative, alternative vector of R. lauricola, Xyleborus bispinatus, to artificial diets of R. lauricola and other ambrosia fungi. Newly eclosed, unfertilized females of X. bispinatus were reared in no-choice assays on one of five different symbionts or no symbiont. Xyleborus bispinatus developed successfully on R. lauricola, R. arxii, R. subalba and R. subfusca, all of which had been previously recovered from field-collected females of X. bispinatus. However, no development was observed in the absence of a symbiont or on another symbiont, Ambrosiella roeperi, recovered from another ambrosia beetle, Xylosandrus crassiusculus. In the no-choice assays, mycangia of foundress females of X. bispinatus harbored significant colony-forming units of, and natal galleries that they produced were colonized with, the respective Raffaelea symbionts; with each of these fungi, reproduction, fecundity and survival of the beetle were positively impacted. However, no fungus was recovered from, and reproduction did not occur on, the A. roeperi and no symbiont diets. These results highlight the flexible nature of the ambrosial symbiosis, which for X. bispinatus includes a fungus with which it has no evolutionary history. Although the “primary” symbiont of the neotropical X. bispinatus is unclear, it is not the Asian R. lauricola.

Wolbachia Infections Responsible for Thelytoky in Dryinid Wasps. The Case of Gonatopus bonaerensis Virla (Hymenoptera: Dryinidae).

We studied the occurrence of Wolbachia in the parasitoid Gonatopus bonaerensis Virla (Hymenoptera: Dryinidae). In order to verify the existence of natural infections in the parasitoid, a field survey was conducted. Identification of Wolbachia was performed on the basis of 16S rDNA, wsp_F1, and wsp_R1-sequences. After the detection of the bacteria, infected specimens of G. bonaerensis were treated with a solution of tetracycline. In Tucumán, parasitoids hold Wolbachia endosymbiont, which seems to control the wasp's reproduction in the nature turning it into thelytokous. The symbiont was identified as the Wolbachia sp. wRi strain. The cure of infected unfertilized females determined the normal arrhenotokous parthenogenesis and the production of male offspring. As a consequence of this procedure, the male of G. bonaerensis is described for the first time.

Aspectos reprodutivos do rotífero Asplanchna do Brasil

Biological aspects of the brazilian retifer Asplanchna sp., reared in the laboratory, were studied and the mean curve of development and its connections with the generation time of amictic females were established. The mean generation time was around 22 hr; thus, the fust offspring of an amictic female were born, on the average, almost at the end of its active growth period. Studies on the longevity of amictic females and on the fecundity in amictic and mictic females were also carried out. The amictic females showed a mean longevity of 90.30 hr and a mean fecundity of 17 female offspring. On the other hand, an unfertilized female produced an average of 13 male offspring or, if fertilized, 4 resting-eggs.

Sex Ratio and Body Mass of Adult Herbivorous Beetles Depend on Time of Occurrence and Light Conditions.

Body mass and sex ratio (F/M) of folivorous insects are easily measured parameters that are commonly used to assess the effect of food quality, living conditions, and preferences on the selection of favourable sites for offspring. A study was conducted on the polyphagous beetle, Gonioctena quinquepunctata (a pest of the native Prunus padus and alien P. serotina) and on the monophagous beetle, Altica brevicollis coryletorum (a pest of Corylus avellana). Both species have a similar life cycle with emergence of current-year adults in summer, and reproduction of 1-year-old insects in spring. A. brevicollis coryletorum feeds primarily on sunlit shrubs, while G. quinquepunctata prefers shaded leaves. The present study assessed the effect of time of occurrence (insect age) on body mass in both sexes and on the sex ratio F/M, taking into account the influence of light conditions associated with their favoured food source (sunlit vs. shaded leaves). We hypothesized that a change in body mass in current-year insects would be determined by the amount of consumed food, while the sex ratio would be stable, when in 1-year-old insects females would die shortly after oviposition, while males would be active for a prolonged time. Results confirmed the hypothesis that changes in mass of current-year beetles was determined by the amount of food intake. We also found that in spring, unfertilized females coexist with fertilized ones and that the latter females live for some time after oviposition; resulting in fluctuations of the mean mass for females. In both species, 1-year-old beetles were heavier than current-year. The preference of A. brevicollis coryletorum for sunlit leaves results in a higher body weight than in G. quinquepunctata in both seasons. The data are consistent and indicate seasonal fluctuations in body mass and changes in the sex ratio in 1-year-old beetles, due to the entrance into their reproductive period.

Female multiple matings and male harassment and their effects on fitness of arrhenotokous Thrips tabaci (Thysanoptera: Thripidae).

Although it is generally assumed that one or a few matings are sufficient to maximize female fitness and that mating is generally assumed to be costly to females, multiple matings of females have been reported across a wide and taxonomically diverse set of animals. Here, we investigated female mating frequency and male harassment rate in arrhenotokous Thrips tabaci. In addition, the cost to females of mating, multiple matings, and male harassment to females was evaluated. We found that T. tabaci females mated multiple times during their lifetime and were subjected to a high rate of male harassment at all the ages we tested. Mating was costly to females in terms of reducing longevity and delaying the initiation of egg laying, although mating did not affect the survivorship and longevity of males. Furthermore, continual exposure to males also resulted in a fitness cost to mated females in terms of delayed egg production and reduced fecundity. Virgin females of arrhenotokous thrips produce only male progeny whereas mated females of arrhenotokous thrips produce males from unfertilized eggs and females from fertilized eggs. However, multiple matings did not allow females to fertilize a larger proportion of their eggs to increase the female offspring ratio. Our study demonstrates the conflicts between the occurrence of multiple matings and the cost of sexual activities. This raises questions about the evolution of multiple matings and polyandry in this species. Furthermore, these findings suggest that such phenomena may occur in other animal species and influence the evolution of their mating systems.

The evolutionary dynamics of major regulators for sexual development among Hymenoptera species.

All hymenopteran species, such as bees, wasps and ants, are characterized by the common principle of haplodiploid sex determination in which haploid males arise from unfertilized eggs and females from fertilized eggs. The underlying molecular mechanism has been studied in detail in the western honey bee Apis mellifera, in which the gene complementary sex determiner (csd) acts as primary signal of the sex determining pathway, initiating female development by csd-heterozygotes. Csd arose from gene duplication of the feminizer (fem) gene, a transformer (tra) ortholog, and mediates in conjunction with transformer2 (tra2) sex-specific splicing of fem. Comparative molecular analyses identified fem/tra and its downstream target doublesex (dsx) as conserved unit within the sex determining pathway of holometabolous insects. In this study, we aim to examine evolutionary differences among these key regulators. Our main hypothesis is that sex determining key regulators in Hymenoptera species show signs of coevolution within single phylogenetic lineages. We take advantage of several newly sequenced genomes of bee species to test this hypothesis using bioinformatic approaches. We found evidences that duplications of fem are restricted to certain bee lineages and notable amino acid differences of tra2 between Apis and non-Apis species propose structural changes in Tra2 protein affecting co-regulatory function on target genes. These findings may help to gain deeper insights into the ancestral mode of hymenopteran sex determination and support the common view of the remarkable evolutionary flexibility in this regulatory pathway.

Половой и возрастной состав популяций мермитид (nematoda, mermithida) – полостных паразитов беспозвоночных животных

It was found that the higher the number of individuals of parasites per unit area, the less evident dissocia-tion of males and females within subpopulations. With an overall paucity of Mermithidae dissociation of genders is a common occurrence. Dissociation of individuals by gender within subpopulations in separate time intervals allows the bias of different generations of parasites, as unfertilized females and non-coupling males accumulate in the open environment for several years. As a result, in subpopulations there is rather complicated composition of age’s groups from individuals of more than one generation.

Unmatedness promotes the evolution of helping more in diplodiploids than in haplodiploids.

The predominance of haplodiploidy (where males develop from unfertilized haploid eggs and females from fertilized diploid eggs) among eusocial species has inspired a body of research that focuses on the possible role of relatedness asymmetries in the evolution of helping and eusociality. Previous theory has shown that in order for relatedness asymmetries to favor the evolution of helping, there needs to be variation in sex ratios among nests in the population (i.e., split sex ratios). In haplodiploid species, unmated females can produce a brood of all males, and this is considered the most likely mechanism for split sex ratios at the origin of helping. In contrast, in diploidiploids unmatedness means total reproductive failure. We compare the effect of unmatedness on selection for male and female helping in haplodiploids and diplodiploids. We show that in haplodiploids, unmatedness promotes helping in females but not in males within the empirical range. In diplodiploids, unmatedness promotes helping by both sexes, and the effect is stronger than in haplodiploids, all else being equal. Our study highlights the need to consider interactions between ecological and genetic factors in the evolution of helping and eusociality.

Exploring the function of red colouration in female spiny-footed lizards (Acanthodactylus erythrurus): patterns of seasonal colour change

In spiny-footed lizards (Acanthodactylus erythrurus), adult females (but not males) show conspicuous red colouration in the tail and hind legs. To investigate the function of this red colouring and proximal causes of seasonal colour change, we captured adult females before the reproductive season and kept them in captivity in one of the three following situations: with a male and fertilization possible, with a male and fertilization impossible, or with another female (fertilization also impossible). Colour was quantified using spectrophotometry. Red colouration increased shortly before the onset of reproduction, but faded during the breeding season and became whitish (light buff-gray) in all cases. Both fertilized and unfertilized females laid eggs or were gravid after two months of the experiment, but while fertilized females laid mostly fertile eggs, unfertilized females only laid infertile eggs. Both egg formation and colour change might be triggered by abiotic factors, although female characteristics also play a role, since heavier females changed colour and laid eggs earlier. Females interacting freely with a male were darker at the end of the breeding season than females separated from the male, indicating that fertilization or physical contact might also have an effect on colouration. Colour change patterns found in this study suggest that female red colouration might have a mating-related function, but do not support a courtship rejection function for the red colour. However, whitish colouration resulting from red fading might signal gravidity in this species. Future experimental manipulation of female colouration will be needed to test these hypotheses.

Life Cycle of Tetranychus macfarlanei Baker and Pritchard (Acari: Tetranychidae): A New Pest of Medicinal Plants in West Bengal

The life cycle of Tetranychus macfarlanei Baker and Pritchard was studied on two different medicinal plants, Clitoria ternatea L. and Justicia adhatoda L. Nees, in BOD at 32.5 °C and 75 % RH during April 2007 to May 2007. Observations towards duration of different stages like egg, larva, protonymph, deutonymph, adult, total life cycle, preoviposition, oviposition, postoviposition periods, longevity of female and male, and fecundity, sex ratio were recorded. Total developmental period of T. macfarlanei from egg to adult was 6.4 ± 0.37 (Mean ± SE) and 10.6 ± 0.56 days on C. ternatea and J. adhatoda, respectively. On C. ternatea, the fecundity in case of fertilized and unfertilized female were 91.6 ± 11.61 and 80 ± 21.64 eggs, respectively and longevity of fertilized and unfertilized female was 16.4 ± 1.44 and 8.6 ± 2.32 days, respectively. The corresponding figure on J. adhatoda for fecundity in case of fertilized and unfertilized female were 39 ± 2.85 and 19.8 ± 3.90 eggs, respectively and for longevity was 16 ± 0.37 and 11 ± 086 days, respectively. Among the two hosts, C. ternatea appears to be more preferred to J. adhatoda because life cycle was completed in shorter time and fecundity and female longevity were for longer duration.

Are individuals from thelytokous and arrhenotokous populations equally adept as biocontrol agents? Orientation and host searching behavior of a fruit fly parasitoid

Hymenopteran parasitoids generally reproduce by arrhenotoky, in which males develop from unfertilized eggs and females from fertilized eggs. A minority reproduce by thelytoky, in which all-female broods are derived from unfertilized eggs. Thelytokous populations are potentially of interest for augmentative biological control programs since the exclusive production of females could significantly lower the costs of mass rearing. Behavioral traits are a major component of parasitoid efficacy. Here, we examined orientation and host searching behavior in thelytokous and arrhenotokous populations of the fruit fly parasitoid Odontosema anastrephae Borgmeier (Hymenoptera: Figitidae). Orientation behavior to various odorant sources was studied in a two-choice olfactometer. No major differences were found between thelytokous and arrhenotokous wasps for this behavior. However, when host-searching behaviors were analyzed, some differences were found. Thelytokous females arrived sooner, foraged longer, and remained longer on non-infested guavas than arrhenotokous females. Individuals of both forms exhibited similar stereotyped behavioral sequences vis-a-vis guava treatments, with only slight deviations detected. Our results suggest that individuals from selected thelytokous and arrhenotokous O. anastrephae populations have similar abilities to search for tephritid larvae, supporting the use of thelytokous strains for augmentative releases.

Biology of Leipothrix dipsacivagus (Acari: Eriophyidae), a candidate for biological control of invasive teasels (Dipsacus spp.)

The present study describes key aspects of the biology of Leipothrix dipsacivagus, an eriophyid mite that is under study as a biological control candidate of Dipsacus fullonum and D. laciniatus (Dipsacaceae). Preliminary host-specificity tests have shown that it can develop and reproduce only on Dipsacus spp. (teasels). Studies were conducted in a laboratory at 26 ± 2(o)C with 16 h of light per day. Mites for the stock colony were collected from D. laciniatus in Klokotnitsa, Bulgaria and reared on rosettes of D. laciniatus in the laboratory. Unfertilized L. dipsacivagus females reared in isolation from the juvenile stage produced male offspring only, while progeny of fertilized females were of both sexes, suggesting arrhenotokous parthenogenesis with haplodiploid sex determination. Experiments were designed to compare male progeny from fertilized females to males from unfertilized females and to compare males and females from fertilized females. Male progeny of virgin mothers had significantly longer durations of active immature stages and total egg-to-adult period than male progeny of fertilized females. Female progeny had significantly longer durations of egg incubation, active immature stages and egg-to-adult period than male progeny from fertilized mothers. Adult longevity was significantly greater in females than in males. Fertilized females produced significantly more eggs per day and overall than virgin females. The results of this study suggest that fertilization status of L. dipsacivagus females can affect both their own fecundity and the development of their male progeny.

Observations on sex ratio and behavior of males in Xyleborinus saxesenii Ratzeburg (Scolytinae, Coleoptera).

Strongly female-biased sex ratios are typical for the fungalfeeding haplodiploid Xyleborini (Scolytinae, Coleoptera), and are a result of inbreeding and local mate competition (LMC). These ambrosia beetles are hardly ever found outside of trees, and thus male frequency and behavior have not been addressed in any empirical studies to date. In fact, for most species the males remain undescribed. Data on sex ratios and male behavior could, however, provide important insights into the Xyleborini’s mating system and the evolution of inbreeding and LMC in general.In this study, I used in vitro rearing methods to obtain the first observational data on sex ratio, male production, male and female dispersal, and mating behavior in a xyleborine ambrosia beetle. Females of Xyleborinus saxesenii Ratzeburg produced between 0 and 3 sons per brood, and the absence of males was relatively independent of the number of daughters to be fertilized and the maternal brood sex ratio. Both conformed to a strict LMC strategy with a relatively precise and constant number of males. If males were present they eclosed just before the first females dispersed, and stayed in the gallery until all female offspring had matured. They constantly wandered through the gallery system, presumably in search of unfertilized females, and attempted to mate with larvae, other males, and females of all ages. Copulations, however, only occurred with immature females. From galleries with males, nearly all females dispersed fertilized. Only a few left the natal gallery without being fertilized, and subsequently went on to produce large and solely male broods. If broods were male-less, dispersing females always failed to found new galleries.

Infrequent sporophyte production maintains a female‐biased sex ratio in the unisexual clonal moss Hylocomium splendens

Summary 1. Sex ratios in unisexual bryophytes are most often female biased, whereas male‐biased sex ratios predominate in unisexual seed plants. This ‘bryophyte paradox’, i.e. that sex ratios are biased in favour of the sex associated with the highest reproductive costs, has remained unexplained. 2. Analysis of sex‐ratio patterns via the influence of sex distribution on population growth rates (λ) has not previously been carried out for bryophytes. We used this method to model how variation in sex ratio and sporophyte frequency influences λ in the clonal bryophyte Hylocomium splendens. We obtained λ by matrix modelling of synthetic experimental populations derived from demographic field data, using a linear two‐sex model. 3. In our set of experimental populations λ varied between 1.13 and 1.27 in response to variation in sex ratio and sporophyte frequency, with the highest λ obtained for the combination of a very low sporophyte frequency and a slightly female‐biased sex ratio. 4. Our results explain the female‐biased sex ratio of H. splendens by the slightly lower survival of and production of vegetative offspring by males than by non‐sporophytic females. 5. Synthesis. According to our models, female dominance is the predicted outcome of low to moderate fertilization success and male performance intermediate between that of sporophytic and of non‐sporophytic females. Our results therefore explain how a female‐biased sex ratio can be maintained despite higher costs of reproduction in females than in males. In dioecious bryophytes, males and females must grow in close contact for fertilization to take place. Better performance of male ramets than of the female ramets they fertilize also explains how male clones can expand into female clones. A similar performance hierarchy of males and females may occur in unisexual clonal seed plants, but more efficient fertilization systems by pollination prevents the selective advantage of unfertilized females from being realized. This explains why vascular plant populations tend to be male biased. We hypothesise difference in fertilization distance range between sperm and pollen as a simple explanation why ramet level sex ratios are in general male dominated in clonal seed plants and female dominated in clonal bryophytes.