Parthenogenesis in Hexapoda: Entognatha and non‐holometabolous insects

In hexapods, unlike the majority of animals, development without fertilization is a common phenomenon. They evolved a striking diversity of unisexual reproductive types that include a variety of modes starting from spontaneous parthenogenesis in females to the production of impaternate males with different variants in between. Many reports about parthenogenetic species have accumulated over time. Here, we present a review of various parthenogenetic hexapod groups with a particular focus on their chromosome systems and ploidy level. We show that conclusions about the reproductive mode often lack solid evidence and sometimes inefficiently demonstrate how parthenogenesis is maintained in corresponding groups. In this review, basal hexapods (Protura, Collembola, Diplura), primarily wingless insect groups (‘Apterygota’) and non‐holometabolous insects are listed with references to a variety of their unisexual reproductive modes.     

Sexual functionality of Leptopilina clavipes (Hymenoptera: Figitidae) after reversing Wolbachia-induced parthenogenesis

Females infected with parthenogenesis-inducing Wolbachia bacteria can be cured from their infection by antibiotic treatment, resulting in male production. In most cases, however, these males are either sexually not fully functional, or infected females have lost the ability to reproduce sexually. We studied the decay of sexual function in males and females of the parasitoid Leptopilina clavipes. In western Europe, infected and uninfected populations occur allopatrically, allowing for an investigation of both male and female sexual function. This was made by comparing females and males induced from different parthenogenetic populations with those from naturally occurring uninfected populations. Our results indicate that although males show a decay of sexual function, they are still able to fertilize uninfected females. Infected females, however, do not fertilize their eggs after mating with males from uninfected populations. The absence of genomic incompatibilities suggests that these effects are due to the difference in mode of reproduction.     

The evolution of developmental gene networks: lessons from comparative studies on holometabolous insects

Recent comparative studies have revealed significant differences in the developmental gene networks operating in three holometabolous insects: the beetle Tribolium castaneum, the parasitic wasp Nasonia vitripennis and the fruitfly Drosophila melanogaster. I discuss these differences in relation to divergent and convergent changes in cellular embryology. I speculate on how segmentation gene networks have evolved to operate in divergent embryological contexts, and highlight the role that co-option might have played in this process. I argue that insects represent an important example of how diversification in life-history strategies between lineages can lead to divergence in the genetic and cellular mechanisms controlling the development of homologous adult structures.     

Manipulation of oviposition choice of the parasitoid wasp, Encarsia pergandiella, by the endosymbiotic bacterium Cardinium

Reproductive manipulations of hosts by maternally inherited bacterial endosymbionts often result in an increase in the proportion of infected female hosts in the population. When this involves the conversion of incipient males to genetic or functional females, it presents unique difficulties for symbionts invading hosts with sex-specific reproductive behaviours, such as the autoparasitic Encarsia pergandiella. In sexual forms of this species, female eggs are laid in whitefly nymphs and male eggs are laid in conspecific or heterospecific parasitoids developing within the whitefly cuticle. Further, eggs laid in the 'wrong' host do not ordinarily complete development. This study explored the role of a bacterial symbiont, Cardinium, in manipulating oviposition behaviour in a thelytokous population of E. pergandiella. Oviposition choice was measured by the number and location of eggs deposited by both infected and uninfected adult waSPS in arenas containing equal numbers of hosts suitable for the development of male and female waSPS. Uninfected waSPS included antibiotic-treated female waSPS and (untreated) daughters of antibiotic-treated female waSPS. The choices of waSPS in the thelytokous population treatments were compared with those of a conspecific sexual population. We found that offspring of antibiotic-cured thelytokous waSPS reverted to the behaviour of unmated sexual waSPS, laying their few eggs almost exclusively in hosts appropriate for male eggs. Infected thelytokous waSPS distributed their eggs approximately evenly between host types, much like mated sexual female waSPS. The antibiotic-treated female waSPS exhibited choices intermediate to waSPS in the other two treatments. The change in the observed behaviour appears sufficient to allow invasion and persistence of Cardinium in sexual populations. Lastly, our results suggest a reduction in host discrimination as a possible mechanism by which Cardinium influences this change.     

A general theory of genital homologies for the Hexapoda (Pancrustacea) derived from skeletomuscular correspondences,with emphasis on the Endopterygota

No consensus exists for the homology and terminology of the male genitalia of the Hexapoda despite over a century of debate. Based on dissections and the literature, genital skeletomusculature was compared across the Hexapoda and contrasted with the Remipedia, the closest pancrustacean outgroup. The pattern of origin and insertion for extrinsic and intrinsic genitalic musculature was found to be consistent among the Ectognatha, Protura, and the Remipedia, allowing for the inference of homologies given recent phylogenomic studies. The penis of the Hexapoda is inferred to be derived from medially-fused primary gonopods (gonopore-bearing limbs), while the genitalia of the Ectognatha are inferred to include both the tenth-segmental penis and the ninth-segmental secondary gonopods, similar to the genitalia of female insects which comprise gonopods of the eighth and ninth segments. A new nomenclatural system for hexapodan genitalic musculature is presented and applied, and a general list of anatomical concepts is provided. Novel and refined homologies are proposed for all hexapodan orders, and a series of groundplans are postulated. Emphasis is placed on the Endopterygota, for which fine-grained transition series are hypothesized given observed skeletomuscular correspondences.     

Influence of microbe-associated parthenogenesis on the fecundity ofTrichogramma deion andT. pretiosum

Microbe-associated parthenogenesis (thelytoky) has been discovered in nineTrichogramma species, parasitoids of mainly lepidopteran eggs. Parthenogenetic and bisexual conspecifics co-occur in many field populations. As an initial step to understand the dynamics of these two reproductive strategies we studied the effect of microbe-associated parthenogenesis on fecundity. The fecundity of two parthenogenetic isofemale lines ofT. pretiosum and one ofT. deion was compared with bisexual lines derived from them by antibiotic treatment. In all three cases parthenogenetic females were less fecund over their lifetime than bisexual females. Also, parthenogenetic females produced fewer daughters in two cases and in one case a similar number of daughters as their respective bisexual counterparts. The lack of mating and insemination was excluded as an explanation for the reduced fecundity of parthenogenetic females, because mated and virgin parthenogenetic females produce the same number of offspring. Antibiotic treatment can also be excluded because females of field-collected bisexual line treated with antibiotics produced the same number of offspring as untreated females. The reduced fecundity of parthenogenetic females was caused by a lower number of eggs being laid rather than by a greater developmental mortality. Parthenogenetic females produced less daughters than bisexual females when host availability was not limiting, but when host availability was severely limited, parthenogenetic females produced more daughters than the bisexual females.     

Parthenogenesis in Tubificidae

The ability of tubificids to reproduce parthenogenetically following or in place of bisexual reproduction has been proved. During parthenogenesis the spermatogenesis ceases at an early stage of sexual cell development, which, together with some peculiarities of the structure of the sexual system, rules out the possibility of self-fertilization in the worms. For two years of cultivation the life cycles of 3 parthenogenetic generations in Tubifex tubifex and 2 in Limnodrilus hoffmeisteri were traced.     

浙江金华棉花粉蚧种群的产两性孤雌生殖

棉花粉蚧是近年来入侵我国的重要害虫,关于该虫在我国的入侵种群的生殖方式缺乏详细研究。本文在室内条件下对棉花粉蚧浙江金华种群的生殖方式进行了观察。结果显示孤雌生殖的棉花粉蚧可以产下雌性和雄性后代。孤雌生殖的棉花粉蚧的产卵前期为14.8 d,显著长于两性生殖的6.9 d;孤雌生殖的棉花粉蚧所产后代为348.3头,显著少于两性生殖的棉花粉蚧的494.1头;孤雌生殖的棉花粉蚧的雌虫总历期为57.2 d,明显长于两性生殖的32.8 d。但这2种生殖方式产下后代的性比差异并不显著,孤雌生殖的为1.1,而两性生殖的为1.2。该棉花粉蚧种群的生殖方式为兼性产两性孤雌生殖。     

A quantitative study of worker reproduction in queenright colonies of the Cape honey bee, Apis mellifera capensis

Reproduction by workers is rare in honey bee colonies that have an active queen. By not producing their own offspring and preventing other workers from producing theirs, workers are thought to increase their inclusive fitness due to their higher average relatedness towards queen-produced male offspring compared with worker-produced male offspring. But there is one exception. Workers of the Cape honey bee, Apis mellifera capensis, are able to produce diploid female offspring via thelytokous parthenogenesis and thus produce clones of themselves. As a result, worker reproduction and tolerance towards worker-produced offspring is expected to be more permissive than in arrhenotokous (sub)species where worker offspring are male. Here we quantify the extent to which A. m. capensis workers contribute to reproduction in queenright colonies using microsatellite analyses of pre-emergent brood. We show that workers produced 10.5% of workers and 0.48% of drones. Most of the workers' contribution towards the production of new workers coincided with the colonies producing new queens during reproductive swarming.     

No evidence for Wolbachia‐induced parthenogenesis in the social Hymenoptera

In some parasitoid wasps, infection by the micro‐organism Wolbachia leads to asexual reproduction. Within the Hymenoptera, the limits of distribution of parthenogenesis inducing Wolbachia have not yet been established. To address this issue, we screened all known thelytokous social hymenopteran species using a PCR assay. None was infected, and therefore we conclude that worker thelytoky evolves independently of Wolbachia in ants and bees. This supports the previously proposed hypothesis that a sex determining system based on heterozygosity may form a proximate limitation to Wolbachia‐induced parthenogenesis.     

Gene flow between sexual and asexual strains of parasitic wasps: a possible case of sympatric speciation caused by a parthenogenesis-inducing bacterium

Sympatric speciation is strictly defined as the emergence of two species from a population in which mating has been random with respect to the place of birth of the mating partners. Mathematical models have shown that sympatric speciation is possible, but very few examples have been documented in nature. In this article, we demonstrate that arrhenotokous and thelytokous strains of a parasitic wasp, Neochrysocharis formosa, speciated sympatrically through infection by a symbiotic bacterium Rickettsia for the following reasons: First, Rickettsia infection was detected in all of the thelytokous strains collected throughout Japan. Second, the arrhenotokous and thelytokous strains have been collected sympatrically. Third, crossing experiments between the two strains did not result in fertilized offspring. In addition, the two strains were genetically isolated at the nuclear and mitochondrial genes. Fourth, the two strains showed a sister relationship in nuclear 28S rRNA gene. Finally, thelytokous females treated with antibiotics produced Rickettsia-free male offspring of the same reproductive form as arrhenotokous females indicating that the thelytokous strain could have speciated sympatrically from an individual of the arrhenotokous strain.     

Antibiotic curing of parthenogenesis in Eretmocerus mundus (Australian parthenogenic form)

The thelytokous parasitoid, Eretmocerus mundus (Australian parthenogenetic form) was infected with a species of Group B Wolbachia. Antibiotic treatment of the parasitoid cured parthenogenesis and lead to male production and reduced egg hatching by virgin females. For each generation infected females produced more progeny than cured females. The descendants of individuals treated with antibiotic were in most cases free of a detectable infection. Both treated females and descendant females mated and received sperm, but these were not used as mated and unmated cured females produced offspring with a similar sex ratio. Males produced as a result of the antibiotic treatment were at least partially functional as they produced sperm and copulated with and inseminated females. However, no sexual line could be established.     

Evolutionary contrasts in insects: nutritional advantages of holometabolous development

ABSTRACT. Two insects, one holometabolous and one hemimetabolous, are compared and contrasted in respect to feeding efficiency, growth and respiration. An examination of cuticle production in these and other species leads to the conclusion that the nutritional burden of cuticle production in hemimetabolous forms has a significant effect on efficiency, and it is speculated that greater nutritional efficiency in holometabolous forms has been an important evolutionary advantage.     

The life cycle of Micromalthus debilisLeConte (1878) (Coleoptera: Archostemata: Micromalthidae): historical review and evolutionary perspective

Micromalthus debilis 33 ), has one of the most bizarre life cycles of any metazoan. Reproduction is typically by thelytokous, viviparous, larviform females, but there is also a rare arrhenotokous phase. The active first‐instar (triungulin) larva develops into a legless, feeding (cerambycoid) larva. This form either pupates, leading to a diploid adult female, or develops into any of three subsequent types of reproductive paedogenetic forms: (1) a thelytokous female that produces triungulins via viviparity; (2) an arrhenotokous female that produces a single egg that develops into the short‐legged (curculionoid) larva, eventually devouring its mother and becoming a haploid adult male; or (3) an amphitokous female that can follow either of the two above reproductive pathways. We speculate that Micromalthus is dependent on maternally transmitted bacteria for the ability to digest rotting wood, and that these bacteria are senescent in males, causing males to be obligately cannibalistic. Obligate male cannibalism, in turn, would have dramatically increased the cost of males, and have created a strong selective advantage for cyclic thelytoky and the other features of the Micromalthus life cycle that minimize the role of the male.     

哺乳动物孤雌干细胞与孤雌生殖

未受精的孤雌胚胎衍生的孤雌胚胎干细胞(parthenogenetic embryonic stem cells,pESCs),具有与胚胎干细胞(embryonic stem cells,ESCs)相似的多向分化和自我更新能力,且具备来.源广泛、获取高效及低致瘤性等优势,因此成为近年来的研究热点.该文概述了pESCs特殊...     

Parthenogenesis in an elasmobranch in the presence of conspecific males

Parthenogenesis has been observed in several elasmobranch species, primarily in public aquaria. The majority of cases of parthenogenesis have occurred either when females were held without males or once a male was removed from a female's habitat. Here we report a second instance of parthenogenesis in a zebra shark female that was housed with conspecific mature males. This study calls into question the conditions under which elasmobranch females undergo parthenogenesis.