Ovary structure in a presocial insect,Elasmucha grisea (Heteroptera,Acanthosomatidae)

First generation egg clusters of Elasmucha grisea are more closely guarded than second generation clusters. The ovaries of this species are structured to enhance this behavior. The population of E. grisea from S-W Poland breeds in the spring (May–June) and late summer (July–August). The second generation clutches contain fewer eggs and are destroyed 3–4 days after oviposition by predators and parasitoids.The ovary structure in the studied species differs from that found in other Heteroptera. The average number of ovarioles per ovary is 24 while in the other investigated species the number of ovarioles per ovary is 6–7. Lateral oviducts are elongated and the ovarioles are arranged in a pennate pattern. Each ovariole contains only one growing ovarian follicle. Differentiation of the ovarioles and ovarian follicles is synchronised thus enabling simultaneous oviposition. A comparative analysis of the ovary structure during the life cycle, particularly the presence of atresive ovarian follicles in the ovarioles of egg- and nymph guarding females, as well as the shape and structure of the apical part of the tropharium all support the hypothesis of cooperation between females in E. grisea. A similar ovary structure has been observed in the Coccoidea (Hemiptera, Homoptera) which indicates presocial behavior.     

Oogenesis and ovarian morphology in pollinating and non-pollinating fig wasps: evidence from adult and immature stages

Hymenopteran insects have meroistic polytrophic ovaries characterised by trophocytes associated with oocytes inside the follicles. In pro-ovigenic species, all oocytes mature before emergence and no trace of oogenesis is visible in adult females. Pro-ovigeny is a rare condition among Hymenoptera, but common in pollinating fig wasps. In the present investigation, we studied adult and pupa females of three fig wasp species with different trophic strategies. We demonstrated that females of Pegoscapus aerumnosus and Idarnes spp. have an unusual ovarian organisation (i.e. each ovariole has only one mature egg and no oocyte) that has led to misleading interpretation of fig wasp reproductive anatomy. The ovaries of these studied species have several ovarioles, recognisable by the presence of nuclei of tunica propria cells surrounding them. Each adult wasp ovariole had one mature egg. None of the pupae had mature eggs, but all of them had follicles with oocytes in different developmental stages. The studied fig wasps are pro-ovigenic, irrespective of their trophic strategy, since there were no signs of ovigeny in adult females. We discuss ecological and phylogenetic factors that could play a role in fig wasps reproductive strategies.     

Role of colour and shape stimuli in host-enhanced oogenesis in the walnut fly, Rhagoletis juglandis

This study aimed to quantify effects of the host plant on oogenesis in the walnut-husk-infesting fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), and to assess the role of physical cues in those effects.
In laboratory assays, the presence of fruit was manipulated independently of the presence of foliage for newly emerged females. After eight days, in each of two trials, females with fruit were found to have significantly higher egg loads than females without fruit. Foliage presence had little effect.
In a second experiment, females held with fruit or a fruit model (plastic yellow sphere of a size similar to fruit) had significantly higher egg loads than females held with neither fruit nor model. Egg loads of females with fruit were not significantly different from those of females with models.
In a third experiment, females were held with spheres of various colours or no sphere at all. Females with yellow or green spheres (similar to the colour of walnut fruit) had significantly higher egg loads than females with black, blue or red spheres of other colours or females without spheres.
In a fourth experiment, females held with spheres had significantly higher egg loads than females held with cubes of equivalent surface area or females held without a model.
Finally, cohorts of newly emerged females held with yellow spheres or without spheres were sampled periodically. In the sphere treatment, mean egg load increased sharply from negligible levels between days 8 and 10. The pattern was similar in the no-sphere treatment, although the increase in egg load appeared to occur a day later.
From these experiments, we conclude that physical host fruit stimuli known to be important in host selection in Rhagoletis flies, including colour and shape, also enhance oogenesis in the first egg maturation cycle, and that enhancement of oogenesis via these stimuli requires neither nutritional input from the fruit nor prior egg deposition.     

Reproductive characteristics of the flower breeding Drosophila hibisci Bock (Drosophilidae) along a latitudinal gradient in eastern Australia: relation to flower and habitat features

For many insect species, egg and larval substrate characteristics are significantly correlated with interspecific differences in female reproductive allocation and egg size-number tradeoffs. We tested the hypothesis that a similar pattern occurred within the Australian drosophilid, Drosophila hibisci, that is restricted throughout its life cycle to flowers of species in the genus Hibiscus. These plants occur as small, isolated, normally monospecific stands that should facilitate differentiation of the fly populations in relation to specific oviposition and larval substrates. Data from 38 sites ranging from 20.8̀ to 34.4̀ S latitude in eastern Australia indicated no relationship between female body size, egg size, or ovariole numbers and floral size or mass among four species of Hibiscus. However, the flies did show a latitudinal cline in ovariole number that was independent of floral variation. Females averaged 15–20 ovarioles per female in the south (32–34̀ S latitude) and 10–12 ovarioles in the north (21–22̀ S latitude). The increase in ovariole number with latitude was due to divergence in the ovariole number of the largest females. In contrast, small females in the north and south had the same number of ovarioles. Reproductive allocation of female flies in the northern region was less than females in the southern region. The latitudinal divergence in ovariole number was not associated with habitat differences (density of trees, density of flies and beetles), nor with differences in floral characteristics (flower weight, petal length, yeast species present). Short term weather patterns in daily temperature and rainfall preceding collections pardy explain the variation in ovariole number. These observations in conjunction with preliminary genetic results suggest the cline is associated with genetic differences that interact with environmental determinants such as the temperature during larval development.     

The ovaries of aphids (Hemiptera,Sternorrhyncha, Aphidoidea): morphology and phylogenetic implications

The ovaries of aphids belonging to the families Eriosomatidae, Anoeciidae, Drepanosiphidae, Thelaxidae, Aphididae, and Lachnidae were examined at the ultrastructural level. The ovaries of these aphids are composed of several telotrophic ovarioles. The individual ovariole is differentiated into a terminal filament, tropharium, vitellarium, and pedicel (ovariolar stalk). Terminal filaments of all ovarioles join together into the suspensory ligament, which attaches the ovary to the lobe of the fat body. The tropharium houses individual trophocytes and early previtellogenic oocytes termed arrested oocytes. Trophocytes are connected with the central part of the tropharium, the trophic core, by means of broad cytoplasmic processes. One or more oocytes develop in the vitellarium. Oocytes are surrounded by a single layer of follicular cells, which do not diversify into distinct subpopulations. The general organization of the ovaries in oviparous females is similar to that of the ovaries in viviparous females, but there are significant differences in their functioning: (1) in viviparous females, all ovarioles develop, whereas in oviparous females, some of them degenerate; (2) the number of germ cells per ovariole is usually greater in females of the oviparous generation than in females of viviparous generations; (3) in oviparous females, oocytes in the vitellarium develop through three stages (previtellogenesis, vitellogenesis, and choriogenesis), whereas in viviparous females, the development of oocytes stops after previtellogenesis; and (4) in the oocyte cytoplasm of oviparous females, lipid droplets and yolk granules accumulate, whereas in viviparous females, oocytes accrue only lipid droplets. Our results indicate that a large number of germ cells per ovariole represent the ancestral state within aphids. This trait may be helpful in inferring the phylogeny of Aphidoidea.     

Age determination in the blackfly Simulium woodi, a vector of onchocerciasis in Tanzania

Abstract. Age-dependent changes in the ovarioles of Simulium woodi (Diptera: Simuliidae) and the age structure of a wild population of this species in the Amani hills of north-eastern Tanzania were studied using the ovarian oil injection technique. Contrary to previous theories, egg sacs degenerate completely and do not form dilatations. The physiological age of the females could be determined only by gonotrophic dilatations formed from degenerating follicles during gonotrophic cycles. In individual females, the proportions of ovarioles with degenerating follicles were 3–54% (mean 22%) in the first cycle and 8–61% (mean 36%) in the second gonotrophic cycle. Agonotrophic degenerating follicles occurred in 2% of ovarioles in 12% of females. Additional information with regard to the degree of parity is given by the level of granulation in the basal body of each ovariole, a group of six to eight cells in the calyx wall enclosed by the end of the ovariolar sheath. Their granulation progressively increased in intensity following each subsequent ovulation.     

The roles of cell size and cell number in determining ovariole number in Drosophila

All insect ovaries are composed of functional units called ovarioles, which contain sequentially developing egg chambers. The number of ovarioles varies between and within species. Ovariole number is an important determinant of fecundity and thus affects individual fitness. Although Drosophila oogenesis has been intensively studied, the genetic and cellular basis for determination of ovariole number remains unknown. Ovariole formation begins during larval development with the morphogenesis of terminal filament cells (TFCs) into stacks called terminal filaments (TFs). We induced changes in ovariole number in Drosophila melanogaster by genetically altering cell size and cell number in the TFC population, and analyzed TF morphogenesis in these ovaries to understand the cellular basis for the changes in ovariole number. Increasing TFC size contributed to higher ovariole number by increasing TF number. Similarly, increasing total TFC number led to higher ovariole number via an increase in TF number. By analyzing ovarian morphogenesis in another Drosophila species we showed that TFC number regulation is a target of evolutionary change that affects ovariole number. In contrast, temperature-dependent plasticity in ovariole number was due to changes in cell-cell sorting during TF morphogenesis, rather than changes in cell size or cell number. We have thus identified two distinct developmental processes that regulate ovariole number: establishment of total TFC number, and TFC sorting during TF morphogenesis. Our data suggest that the genetic changes underlying species-specific ovariole number may alter the total number of TFCs available to contribute to TF formation. This work provides for the first time specific and quantitative developmental tools to investigate the evolution of a highly conserved reproductive structure.     

Structure and development of the telotrophic ovariole in ensign scale insects (Hemiptera, Coccomorpha: Ortheziidae)

The paired ovaries of young larva of the 3rd instar of Orthezia urticae are filled with numerous germ cell clusters that can be regarded as ovariole anlagen. Germ cells (cystocytes) belonging to one cluster form a rosette, in the centre of which a polyfusome occurs. Staining with rhodamine-phalloidin has revealed that polyfusomes contain numerous microfilaments. The number of cystocytes per cluster is not stable and varies considerably. The ovaries of older larva become elongated with numerous young ovarioles protruding into the body cavity. The ovarioles are not subdivided into the tropharium and vitellarium. In this stage germ cells differentiate into oocytes and trophocytes (nurse cells). The ovaries of adult females are composed of about 20 (Newsteadia floccosa) or 30 (O. urticae) ovarioles. Their trophic chambers contain trophocytes and arrested oocytes. In the vitellarium, at the given moment, only one oocyte develops. It has been observed that after maturation of the first egg the arrested oocytes may develop.     

Structure of the female reproductive system of the lac insect,Kerria chinensis (Sternorrhyncha,Coccoidea: Kerridae)

The ovaries of female lac insects, Kerria chinensis Mahd (Sternorrhyncha: Coccoidea: Kerridae), at the last nymphal stage are composed of several balloon‐like clusters of cystocytes with different sizes. Each cluster consists of several clusters of cystocytes arranging in rosette forms. At the adult stage, the pair of ovaries consists of about 600 ovarioles of the telotrophic‐meroistic type. An unusual feature when considering most scale insects is that the lateral oviducts are highly branched, each with a number of short ovarioles. Each ovariole is subdivided into an anterior trophic chamber (tropharium) containing six or seven large trophocytes and a posterior vitellarium harbouring one oocyte which is connected with the trophic chamber via a nutritive cord. No terminal filament is present. Late‐stage adult females show synchronized development of the ovarioles, while in undernourished females, a small proportion of ovarioles proceed to maturity.     

The ovaries of Mecoptera: basic similarities and one exception to the rule

Two entirely different types of ovaries (ovarioles) have been described in mecopterans. In the representatives of Meropeidae, Bittacidae, Panorpodidae and Panorpidae the ovarioles are of the polytrophic-meroistic type. Four regions: a terminal filament, germarium, vitellarium and ovariole stalk can be distinguished in the ovarioles. The germaria house numerous germ cell clusters. Each cluster arises as a result of 2 consecutive mitoses of a cystoblast and consists of 4 sibling cells. The oocyte always differentiates from one of the central cells of the cluster, whereas the remaining 3 cells develop into large, polyploid nurse cells. The vitellaria contain 7-12 growing egg chambers (= oocyte-nurse cell complexes). In contrast, the ovaries of the snow flea, Boreus hyemalis, are devoid of nurse cells and therefore panoistic (secondary panoistic). The ovarioles are composed of terminal filaments, vitellaria and ovariole stalks only; in adult females functional germaria are absent. Histochemical tests suggest that amplification of rDNA takes place in the oocyte nuclei. Resulting dense nucleolar masses undergo fragmentation into multiple polymorphic nucleoli. The classification of extant mecopterans as well as the phylogenetic relationships between Mecoptera and Siphonaptera are discussed in the context of presented data.     

Age‐related changes in the development of oocytes and testicular follicles after emergence in Stavsolus japonicus (Plecoptera)

Abstract Female stoneflies oviposit several times during the adult stage of their life cycle. The number of eggs within the deposited egg masses decreases at successive ovipositions. To clarify the reason for this decrease and to determine the conditions of testicular follicles, the patterns of development of oocytes and testicular follicles on different days after emergence are investigated in the systellognathan species Stavsolus japonicus (Okamoto) (Perlodidae). The size of the mature oocytes in the ovariole peaks a few weeks after emergence but decreases to the lowest level by 35 days after emergence. Several maturing oocytes can be observed in the ovarioles of individuals a few weeks after emergence but only one mature oocyte is observed at 35 days after emergence. The decreased number of eggs laid per mass by older individuals may therefore be due to the lower maturation of all the ovarioles.     

Distribution of cytoskeletal elements in the ovarioles of Mutilla sp. (Insecta,Hymenoptera)

The ovaries of Mutilla sp., as those of other hymenopterans, consist of meroistic-polytrophic ovarioles. Within each ovariole, a terminal filament, a germarium, and a vitellarium can be distinguished. The germaria contain numerous dividing and/or differentiating groups (clusters) of germ cells. The vitellaria are composed of several, linearly arranged, ovarian follicles; each follicle consists of an oocyte and a group of nurse cells. Distribution of cytoskeletal elements (microfilaments and microtubules) throughout the ovarioles of Mutilla sp. has been studied on whole mount preparations stained with rhodamine-conjugated phalloidin and FITC-labelled anti-tubulin.     

Ontogenesis of the ovary in a moth midge,Tinearia alternata Say (Diptera: Psychodidae)

In a psychodid, Tinearia alternata, the initial differentiation of the polytrophic ovary occurs during the early larval stages. Early in development, each ovary anlage is a solid organ subdivided into three distinct zones: the cortex houses germ cells and somatic interstitial tissue, while two other somatic regions will give rise to the oviduct calyx and anterior part of the lateral oviduct. Germ cell cluster formation precedes the development of ovarioles. Each ovariole houses only one functional egg chamber. All ovarioles within paired ovaries are developmentally synchronized. In the larval ovaries, the newly formed egg chambers and then the ovarioles are intermingeled with and surrounded by the somatic interstitial tissue of the ovary cortex. The interstitial cells give rise to all the somatic elements of the ovarioles. In the pupal ovaries, the remaining interstitial tissue degenerates; thus, the ovarioles protrude into the body cavity. The ovaries in psychodids develop relatively large and swollen oviduct calyxes that are equivalent to receptaculum seminis (spermatheca). The morphological differentiation of germ cells within the egg chambers starts during late larval/early pupal stages. Nurse cell nuclei contain prominent nucleoli and polytene chromosomes. Oocyte growth results from accumulation of yolk and then, in the final stages of oogenesis, from an inflow of cytoplasm from the nurse cells. J. Morphol. 236:167–177, 1998. © 1998 Wiley-Liss, Inc.     

两种寄生蜂的卵巢结构与卵成熟特点初步观察

对斜纹夜蛾的内寄生蜂双斑侧沟茧蜂MicroplitisbicoloratusChen和南美斑潜蝇的外寄生蜂潜蝇姬小蜂Diglyhusisaea(Walker)的卵巢结构进行了研究。双斑侧沟茧蜂的2个卵巢管底部膨大有大量的卵,通过凹陷的颈部与萼区相连接,在萼区里大量成熟待产的卵浸泡在萼液中。产卵量随产卵时间延长而降低,在日产卵实验中,日产卵量202(±114SE),从1000至1800每2h的产卵量分别占日产卵量的38,32,17和13%;在一生产卵量实验中,9d产卵期,每头雌峰一生平均产卵量为502(±78SE)粒,第1d产卵量占一生总产卵量的42%。潜蝇姬小蜂的每个卵巢由3条卵巢管组成,每条输卵管中仅有1粒成熟卵,整个卵巢中只有6粒成熟卵,每次产卵数不超过6粒。结果表明,M.bicoloratus是早熟卵(proovigenic)类昆虫;D.isaea是同步卵(synovigenic)类昆虫。     

Phase polymorphism in Schistocerca gregaria: Reproductive parameters

Solitarious phase Schistocerca gregaria were selected according to the criterion of the occurrence of an extra stadium during larval development. The proportion of locusts undergoing an extra ecdysis increased within one generation of rearing under isolated conditions but it failed to reach 100% throughout an observation period encompassing six generations. A higher proportion of female locusts exhibited an extra stadium indicating sexual dimorphism in this phase character.The number of eggs per pod produced by solitarious females was larger than their gregarious counterparts. This was shown to be due to an increase in the number of ovarioles and a relative decrease in the proportion of non-functional oöcytes in solitary females, confirming earlier work.The inverse relationship between ovariole numbers and oöcyte size characterizing locust phase polymorphism was examined. Vitellin content of ovaries containing mature oöcytes was similar in gregarious and solitarious females. However, vitellin content per oöcyte was less in the latter suggesting that the increase in ovariole number may occur at the expense of oöcyte size and vitellin content.     

Egg Load and Body Size of Lab-cultured Cotesia marginiventris

The egg load of lab-cultured Cotesia marginiventris (Cresson) (Hymenoptera: Braconidae), a solitary koinobiont endoparasitoid of noctuid caterpillars, was determined in this study. Information on egg load may provide clues to more efficient in vivo rearing of C. marginiventris. I tested the hypothesis that egg load, defined as the number of mature oöcytes (i.e., fully chorionated eggs) found in adult females, was related to body size. Cotesia marginiventris females possessed two ovaries and two ovarioles per ovary; mature eggs were found in ovaries and oviducts. Newly-emerged females held an average of 149 mature eggs. Immature eggs were slightly visible in the distal portions of the ovarioles; they were not counted. Egg load was marginally related to body size (i.e., hind tibia length). The results of this study suggest that (1) body size can sometimes predict egg load or potential fecundity of lab-cultured C. marginiventris and (2) an efficient rearing system that exploits the potential fecundity of C. marginiventris might involve using young females and allowing them to oviposit in new hosts, each day, for up to a week.     

Inhibition of reproduction and embryogenesis in the firebrat, Thermobia domestica, by juvenile hormone analogues

Application of compounds with juvenile hormone (JH) activity to reproducing females may result in shortening of the ovipositor and occasionally in some other changes in appearance. Severe defects occur in the ovaries: the differentiation of oögonia and prefollicular cells seems to be hindered and the mature oöcytes are resorbed. The affected ovarioles diminish and in extreme cases tear into the germarium, which remains attached to the terminal filament, and the rest of the ovariole, which shrinks into a small rudiment adjacent to the oviduct. The development of eggs deposited by treated females is often lethally affected. Disorders in embryogenesis also occur in the eggs contaminated with active substances at any time between deposition and 2 days before hatching. The substances interrupt the normal course of embryogenesis but the embryos survive for some time and may develop into strange creatures; in one case, two embryos were found within a single egg shell. One out of 27 tested substances was active in amounts lower than 1 ng/egg.     

Reproductive characteristics of Drosophila hibisci in the Northern Territory, Australia

Reproductive traits of Drosophila hibisci collected at 18 sites in the Northern Territory (NT) of Australia in May, 1998, as well as at two sites in north Queensland, in June, 1998, were compared to those from earlier work on a cline in ovariole number in D. hibisci along the east coast of Australia. The flies in the NT were considerably smaller, but had more ovarioles than comparably-sized flies on the east coast. Although the flies on the east coast showed an increasing number of ovarioles in populations at increasing distances from the equator, these new populations, both on the east coast and in the NT, reversed this trend, producing a generally U-shaped pattern of ovariole number with latitude among all populations. The northernmost and southernmost populations allocate more to ovariole numbers than populations in intermediate latitudes. Ovariole number is closely related to body size of females in all populations, but the regression coefficient is small at intermediate latitudes and increases at the northern and southern ends of the distribution. Egg volumes primarily varied with body size of the female (positive) and number of ovarioles per female (negatively), producing a generally inverted U-shaped pattern of egg volumes with latitude. Reproductive allocation patterns, but not thorax size or ovariole number, varied significantly in two samples taken 10 days apart at one NT site. This variation probably results from environmental differences across generations of developing larvae and is consistent with our earlier suggestion of substantial effects of the environment, primarily rainfall and temperature, on reproductive allocation in D. hibisci.     

Effects of female body size on lifetime fecundity of <Emphasis Type="Italic">Monochamus urussovii</Emphasis> (Coleoptera: Cerambycidae)

To analyze the relationship between female body size and lifetime fecundity (LF) in Monochamus urussovii (Fischer), 17 pairs of adults were reared on Picea jezoensis (Sieb. et Zucc.) Carrière twig sections and were allowed to oviposit on Abies sachalinensis (Fr. Schmidt) Masters bolts at 25 °C and 16:8 h light:dark. The means of life span and LF were 42.5 days and 28.2 eggs for fertile and sterile females combined, respectively. The mean lengths of preoviposition and oviposition periods (OPs) of fertile females were 12.1 and 28.1 days, respectively. The potential fecundity was estimated to be 72.1. Path analysis showed that female body size, i.e., mass and elytral length, had positive effects on LF through the oviposition rate but not through the OP. Ovariole number was 24 and did not vary according to body size, indicating a higher egg productivity per ovariole in large females than in small ones. Higher egg productivity per ovariole in large females is true for each of three other synovigenic lamiine beetles with an almost fixed number of ovarioles. The body size-dependent egg productivity per ovariole is considered to be one of the reasons for the body size–LF correlation in synovigenic species with a fixed ovariole number.