Vitellogenic and embryogenic activity of the microtubule disruptor vinblastine following ingestion by the wasp Bracon hebetor

The reproductive performance of Bracon hebetor females was adversely affected by the consumption of sub-lethal doses of vinblastine. While all oögenic cell types present in the gonads at the time of treatment displayed various degrees of fecundity and/or fertility depression, the transitional cells proved to be the least susceptible to vinblastine damage. Vitellogenically active oöcytes were most sensitive to vinblastine. In these oöcytes development was blocked at or prior to the terminal growth phase. Oviposited eggs which did arise from the exposed vitellogenic oöcytes were few in number and characterized by aberrant morphology. Embryogenic effects were predominantlydue to pre-blastoderm formation damage and most pronounced in oöcytes exposed during the most advanced stages of gonadal development, late vitellogenesis through meiotic metaphase I. Reduced egg hatchability also occurred in exposed undifferentiated oögonial cells but the effect was less severe than that seen in the more mature oögenic cells. All observed effects could be accounted for by vinblastine's selective interference with microtubule-dependent processes. Fecundity effects were most closely associated with oöcyte cortex and possibly follicular cell damage which prevented vitellogenic growth beyond the terminal growth phase. Fertility effects were caused by the inhibition of early embryonic karyokinesis with the most plausible target being mitotic spindle formation.     

Evidence that the ovaries of Musca domestica do not maintain cyclicity by regulating the corpus allatum.

When the female housefly retains eggs, vitellogenesis in the penultimate oöcytes is suppressed during continued protein feeding. Allatectomy of gravid females or of females with developing oöcytes did not prevent maturation of a second batch of eggs. This result does not support the claim of Adams (1970) that an oöstatic hormone, produced by ovaries with retained eggs, inactivates the corpus allatum (CA) and thereby prevents development of the next batch of eggs.The report of Adams and Hintz (1969) that the CA regulates egg maturation depends on their removal of ring gland, which they incorrectly refer to as ‘allatectomy’. In the present report, removal of the ring gland from 1 day old females suppressed egg development, whereas removal of the CA alone did not. Therefore, sufficient CA hormone for reproduction was secreted within 24 hr of emergence, and it was the removal of the corpus cardiacum, and not the CA, that had prevented egg development in the experiments of Adams and Hintz.     

Reproductive allocation by the blow fly Lucilia sericata in response to protein limitation

Adult females of the anautogenous blow fly Lucilia sericata (Meigen) (Diptera: Calliphoridae) were fed standardized meals of liquidized liver, in quantities shown to range around the minimum required to initiate yolk deposition. Females from each feeding regime were dissected at daily intervals between 4 and 11 days of age; the number and stage of development of all oöcytes were recorded. Once an initial threshold quantity of protein was ingested, yolk deposition was initiated in all available oöcytes. Subsequently, one of two distinct developmental pathways was followed: arrested development in all oöcytes at an early stage of yolk deposition, or more extensive yolk deposition followed by progressive oösorbtion and the maturation of small batches of eggs. The proportion of females showing oösorbtion relative to arrested follicular development increased with increasing protein meal size, suggesting that the difference in response may be triggered by a second protein threshold, either side of which the arrested development or oösorbtion pathways are followed. The behaviour observed may reflect strategies to maximize reproductive output in this short‐lived, resource‐limited insect species. Flies that display arrested development may have sufficient protein to mature few if any complete eggs, but may subsequently be able to mature a full egg‐batch if they obtain further protein meals; this possibility is offset by the risk of death before finding such a meal. Flies that show oöcyte development and oösorbtion produce smaller egg batches more quickly and hence have a higher probability of achieving at least some reproductive output. By initiating yolk deposition in all oöcytes, female L. sericata retain the potential to adopt either developmental pathway, depending on subsequent protein intake.     

Ovarian dynamics in Heliconius butterflies: Correlations among daily oviposition rates,egg weights,and quantitative aspects of oögenesis

Both oviposition and ovarian morphology were studied in individual butterflies of the neotropical genus Heliconius, which as adults ingest amino acids from pollen, live up to 6 months, and have continuous oögenesis. Among female Heliconius charitonius, daily oviposition correlates directly with the total number of oöcytes developing in the ovaries. The calculated time required for complete growth of one oöcyte is, however, reasonably constant among individuals of a species with widely varying oviposition rates. Thus, within a species, butterflies laying more eggs per day are not necessarily maturing each at a faster rate, but are processing more oöcytes simultaneously in their ovaries. A further correlation between oviposition rate and adult size suggests that in H. charitonius both ovarian capacity and daily egg production are determined ultimately by extent and/or quality of larval nutrition. Among other Heliconius species, those producing larger eggs generally take longer to make them, but also may develop more oöcytes at the same time in their ovaries. Finally, maximal volume attained by the cap of seven nurse cells associated with each growing oöcyte appears constant within a species and among species is directly proportional to average mature egg weight.     

Factors responsible for the initiation of a second oöcyte maturation cycle in the ovoviviparous cockroach Nauphoeta cinerea

The factors responsible for the initiation of a second oöcyte maturation cycle were investigated by measuring oöcyte growth, vitellogenin titre, and corpus allatum activity after injection of juvenile hormone and/or removal of the egg-case from pregnant females and by performing ovary and corpus allatum transplant experiments.Egg-case removal in late pregnancy results in immediate oöcyte growth, whereas in early pregnancy oöcyte growth is resumed only after a lapse of time, even after injection of juvenile hormone. This, however, induces an immediate increase in the haemolymph vitellogenin titre. A single injection of 2 or 10 μg of juvenile hormone II first stimulates some oöcyte growth after this lapse of time and later activates the corpora allata, which in turn leads to completion of oöcyte maturation. A repeat injection of 10 μg stimulates continuous oöcyte growth without activating the corpora allata. In the presence of an egg case, activation of the corpora allata is suppressed, even after injection of 2 μg of juvenile hormone III, and the oöcytes do not grow. Injection of higher doses stimulates oöcyte growth and leads to expulsion of the egg case in up to 95% of the females. This, however, is not a direct consequence of the increase in size of the ovaries. Ovary transplant experiment show that in young pregnant females the second generation of oöcyte is not yet competent for growth and that ovaries which are competent can mature in young pregnant females, treated with juvenile hormone, whose egg case has been removed.The results are summarized in a model demonstrating the various factors involved in regulating corpus allatum activity in oöcyte maturation and pregnancy and after application of juvenile hormone. We prepose that the corpus allatum activating effect of exogenous juvenile hormone is mediated by the growing oöcyte and that this activation can be suppressed by the continuous presence of exogenous juvenile hormone.     

Isolement,action biologique et evolution des substances paragoniales contenues dans le spermatophore d'Acanthoscelides obtectus (Coleoptère)

In Acanthoscelides obtectus, some male secretions deposited in the spermatophore during mating reach the blood of the females and stimulate oögenesis. Water extracts from spermatophores injected into a female abdomen stimulate oögenesis but do not influence egg-laying or sexual receptivity. After column chromatograph of spermatophores, aqueous extracts on Sephadex G 25 Coarse, G 25 Superfine, and G 15, an active fraction has been isolated. This injected into the abdomen of virgin females stimulates oögenesis at low concentrations, but it is toxic at higher concentrations. This fraction was examined by paper electrophoresis at low voltage and then chromatographed on G 10 Sephadex. Two peaks were obtained: the first corresponds to the paragonial substance A which stimulates oögenesis at 0,2 10^?3 μg/μl concentration. The second contains the paragonial substance B. At a 0,3 10^?3 ug/μl concentration this substance is toxic. First this toxicity inhibits oögenesis and then causes the death of most females at higher concentrations. The toxic effect appears 2 or 3 days after injection. These two substances are purified on paper chromatography and the biological activities are contained in a zone of R_f 0.25 to 0.45 (paragonial substance A) and in a zone of 0.16–0.30 R_f (paragonial substance B).The paragonial substances disappear from the spermatophore after mating. Aqueous extracts of spermatophores obtained 6 hr after mating do not stimulate oögenesis and do not have any toxic effect. The chemical nature of these both fractions is not yet determined because the quantity of extracts obtained at the end of the purification is very low.The action of both paragonial substances is similar to the action of hormones. The paragonial substances influence unknown receptors at low concentration after a latent period. The origin of the paragonial B substance was not determined, but this substance which inhibits oögenesis at low concentrations could be an antagonist of paragonial A substance.     

Effects of electrocoagulation of cerebral neurosecretory cells and implantation of corpora allata on oöcyte development in Locusta migratoria

The implantation of active corpora allata into intact Locusta females during growth accelerates pre-vitellogenic oöcyte growth and vitellogenesis. Localised stimulation of yolk deposition follows the implantation of active corpora allata between the ovarioles demonstrating a gonadotrophic rôle for the corpus allatum hormone. Electrocoagulation of the median neurosecretory cells of the brain prevents vitellogenesis whilst pre-vitellogenic oöcyte growth occurs normally. Implantation of active corpora allata into females with ablated cerebral neurosecretory cells promotes vitellogenesis in a proportion of test animals although mature oöcytes are never produced.It is suggested that the rôle of the median neurosecretory cells during egg development in Locusta is primarily concerned with the activation and maintenance of activity of the corpora allata. The corpus allatum hormone acts both metabolically and gonadotrophically.     

Hormonal aspects of oögenesis in the flies Phormia regina and Sarcophaga bullata

The corpora allata (CA) and median neurosecretory cells (MNC) of Phormia regina and Sarcophaga bullata become active with increasing age of the fly, on a diet of sugar alone. To prevent or retard oögenesis the CA or MNCs must be removed shortly after emergence, with subsequent protein meals. Topical JH application partially compensates for CA or MNC removal. This shows that the MNC activate the CA, and not vice versa. The trauma of either operation slightly depresses egg development.Injection of ecdysone into both species in the stage of initial yolk deposition causes the primary oöcytes to degenerate. This leads to development of the penultimate oöcytes. Older and younger egg stages are not sensitive to ecdysone. In P. regina the application of JH to females with developing primary oöcytes stimulates yolk deposition in the penultimate oöcytes.     

Inhibition of oöcyte development during pregnancy in the cockroach Eublaberus posticus

The corpora allata are inhibited during pregnancy in ovoviviparous Eublaberus posticus, and yolk is not deposited in the basal oöcytes for the entire or almost the entire gestation period.Precocious oöcyte development occurs if the oötheca is removed but this can be prevented by substituting a plastic oötheca for the true egg case in the uterus. Implantation of a uterus containing an oötheca into the abdomen of a female whose oötheca is removed does not prevent precocious oöcyte development even though many of the eggs in the implant grow and stretch the donor uterus. These experiments argue against the hypothesis that an ‘agent’ from the uterine eggs or stretched uterus inhibits the activity of the corpora allata (CA), and supports the hypothesis that inhibition from the uterus is mechanical.Cyclical activity of neurosecretory cells in certain abdominal ganglia in one species of ovoviviparous cockroach has been correlated with the cyclical inhibition of the oöcytes during pregnancy. Mechanoreceptors are found in the uteri of several ovoviviparous species including Eublaberus.In Eublaberus transecting the nerve cord between various ganglia in pregnant females only results in a marked decrease in the percentage of famales showing precocious oöcyte development when the nerves posterior to the sixth abdominal ganglion are severed. However, the results are the same if these nerves are severed after removing the oötheca. It is suggested that pressure of the oötheca on mechanoreceptors in the uterus, or cessation of pressure (after removal of the oötheca), result in sensory information being transmitted to the last abdominal ganglion which affect the CA, perhaps indirectly by controlling the activity of the neurosecretory cells in various abdominal ganglia.     

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.     

The role of factors from the head in the regulation of egg development in the mosquito Aedes aegypti

The relationships between the release of factors from the head after blood-feeding, subsequent levels of ecdysteroids and vitellin, and the ultimate maturation of eggs in Aedes aegypti were investigated. Females were decapitated at various times after a blood meal, at 20 or 48 h after feeding the animals were dissected and divided into two groups, those with arrested oöcytes (yolk length < 100 μm) and those with maturing oöcytes (yolk length > 100 μm). These yolk lengths correspond with the levels of oöcyte growth believed to accompany the proposed initiation and promotion phases of egg development. Animals dissected at 20 h were assayed for ecdysteroid by radioimmunoassay; those dissected at 48 h were assayed for vitellin by rocket immunoelectrophoresis.Non-blood-fed unoperated females contained 8% as much ecdysteroid as blood-fed controls and no measurable vitellin. Females with arrested oöcytes (< 100 μm) were obtained only if decapitations were performed before 8 h; these females had about 20% of the ecdysteroids and 8% of the vitellogenin normally found in blood-fed animals. Females decapitated between 2 and 8 h with maturing oöcytes contained 50–60% as much ecdysteroid and vitellin as blood-fed unoperated controls. Normal ecdysteroid and vitellin levels were reached only when decapitations were delayed for 12 and 24 h, respectively. The number of developing oöcytes was also decreased by early decapitation and was closely correlated with vitellin levels.We conclude that the egg development neurosecretory hormone is released twice, once before 8 h and once after 8 h, to control ecdysteroid levels. We also suggest the presence of other factors from the head that control vitellin levels, the number of developing oöcytes, and the early growth of the oöcyte (initiation).     

Systematic and palaeoecological significance of the first record of Pygocephalomorpha females bearing oöstegites (Malacostraca,Peracarida) from the lower Permian of southern Brazil

Malacostracan crustaceans of the fossil order Pygocephalomorpha are conspicuous elements in brackish to freshwater faunas in the upper Carboniferous of Laurentia and lower Permian of Gondwana. A pronounced sexual dimorphism is recognizable within its members, with Pygocephalomorpha females bearing oöstegites: modified epipodites that hold a brood pouch where the eggs develop until juvenile stage, with no planktonic dispersal of larvae. The preservation of oöstegites is quite rare and is described here for the first time from the American continents. The specimens, ventrally preserved Liocaris huenei females bearing oöstegites, come from the Irati Formation, a lower Permian sequence of the Paraná Basin, from an outcrop in southern Brazil. The oöstegites are similar to those previously described for Pygocephalomorpha, displaying seven pairs of overlapping epipodites that cover the whole cephalothorax. The scarcity of these structures in the fossil record may be explained by taphonomic bias (it requires ventrally preserved crustaceans) but more likely by palaeobiological aspects, such as preservation of an ovigerous moult. The record from the Irati Formation suggests that pygocephalomorphs had seasonal reproduction, with females maturing at the same time and acquiring oöstegites that would be lost after the release of juveniles. This record confirms the relationship of the Brazilian Liocaris Beurlen to the English Pygocephalus Huxley and South African Notocaris Broom pygocephalomorphs and provides an insight into its palaeoecological significance.     

The effects of starvation and subsequent feeding on juvenile hormone synthesis and oöcyte growth in Schistocerca americana gregaria

The effect of starvation on the synthesis of C₁₆ juvenile hormone (JH) and the growth of terminal oöcytes was assessed in Schistocerca americana gregaria at two times during adult life: before activation of the corpora allata and during the first gonotrophic cycle. In both groups, starvation resulted in a decline in JH synthesis within 2–3 days and rates of synthesis remained low throughout the experimental period. The growth rate of oöcytes which were not vitellogenic at the time of starvation was depressed whereas the percentage of resorption of vitellogenic oöcytes increased dramatically with starvation. Although the percentage of resorption increased in animals with vitellogenic oöcytes, some mature oöcytes were produced, particularly in animals in which the oöcytes were greater than 5 mm in length at the time of starvation. This suggests that oöcyte maturation can be divided into two distinct phases—an early phase of vitellogenesis associated with high rates of JH synthesis and a late phase, in oöcytes greater than 5 mm, associated with much lower rates of JH synthesis.Stimulation of JH synthesis by farnesenic acid in 5-day starved animals resulted in high rates of JH synthesis, indicating that starvation did not appreciably alter the enzymic activities of the final two stages in JH synthesis. Thus rate limitation did not occur at these stages.Feeding of 5-day starved animals resulted in a transient increase in the rate of JH synthesis. However, rates of JH synthesis and oöcyte growth remained subnormal throughout the observation period, suggesting that the effects of starvation cannot be entirely reversed by feeding. Thus starvation may decrease the reproductive potential of the females.     

Determinants of oöcyte degeneration in Drosophila melanogaster

During normal oögenesis in many insects some of the oöcytes fail to mature; instead they degenerate and are resorbed. In this work oöctte degeneration was investigated in Drosophila melanogaster females and found to be limited to early vitellogenic stages (stages 8–10). Even when retained for up to 18 days by females, mature (stage 14) oöcytes showed unaltered protein patterns after separation by SDS polyacrylamide electrophoresis, indicating that protein breakdown, which is characteristic of degeneration, does not occur in chorionated oöcytes.A number of environmental parameters were shown to influence the percentage of degenerating oöcytes in females. Strong responses as reflected by increased stage-8 and 9 oöcyte degeneration were found in females subjected to suboptimal (but not starvation) medium, virgin females, females mechanically unable to oviposit, and females unable to locate suitable oviposition sites. Little or no response was seen in females subjected to crowding, however, since all of these environmental parameters except adult crowding have been shown to decrease fecundity, and therefore the rate of oöcyte production, it is suggested that oöcyte degeneration is a strategy for decreasing the rate of oöcyte production in Drosophila.     

Flight during oviposition reduces maternal egg provisioning and influences offspring development in Pararge aegeria (L.)

As a result of increased habitat fragmentation in anthropogenic landscapes, flying insects may be required to travel over larger distances in search of resources such as suitable host plants for oviposition. The oögenesis–flight syndrome hypothesis predicts that physiological constraints caused by an overlap in the resources used by thoracic muscles during flight and during oögenesis (e.g. carbohydrates, lipids and water) result in a resource trade‐off, with any resources used during flight no longer available for reproduction. Increased flight costs could therefore potentially result in a decrease in maternal provisioning of eggs. In the present study, the speckled wood butterfly Pararge aegeria (L.) is used to investigate whether increased flight during oviposition results in changes in maternal investment in eggs and whether this contributes to variation in the development of offspring in subsequent life stages. Forcing females to fly during oviposition directly influences egg size and embryonic development time, and indirectly influences (through changes in egg size) egg hatching success and larval development time. These effects are mediated through ‘selfish maternal effects’, with mothers forced to fly maximizing their fecundity at the expense of investment to individual egg size. The present study demonstrates that a change in maternal provisioning as a result of increased flight during oviposition has the potential to exert nongenetic cross‐generational fitness effects in P. aegeria. This could have important consequences for population dynamics, particularly in fragmented anthropogenic landscapes.     

Host selection by Homalodisca vitripennis: the interplay between feeding, egg maturation, egg load, and oviposition

For some phytophagous insects, egg maturation may be dependent on adult feeding. Accordingly, rates of egg maturation may be dependent on the quality and quantity of available food sources. In turn, oviposition behavior could be affected by diet quality via changes in egg load (number of mature eggs carried by a female). Experiments were conducted to determine whether adult feeding may affect oviposition behavior of the glassy-winged sharpshooter, Homalodisca vitripennis. No-choice tests demonstrated that eggs accumulated in glassy-winged sharpshooter abdomens as time since last oviposition increased largely as a function of feeding plant species. In choice tests, glassy-winged sharpshooter females were observed most frequently on the plant species that imparted the greatest egg maturation rate in no-choice tests. Direct tests of the effects of egg load on glassy-winged sharpshooter oviposition behavior found that females were more likely to deposit eggs as egg load increased. Similarly, acceptance of a low-ranked oviposition plant species by female glassy-winged sharpshooters increased with egg load and time since last oviposition. The results indicate that adult feeding affected glassy-winged sharpshooter egg maturation, plant species varied in quality for providing nutrients for egg maturation, and egg load affected oviposition behavior. Thus, the quantity and quality of available feeding plant species may affect glassy-winged sharpshooter egg maturation rates, which in turn may affect the plant species female glassy-winged sharpshooters select for oviposition.     

Endocrine control of vitellogenesis in the harlequin bug, Dindymus versicolor

An active corpus allatum is essential for the maturation of eggs in the harlequin bug, Dindymus versicolor. The corpus allatum of virgin females remains virtually inactive and the oöcytes are resorbed once they have grown to the stage at which they become competent to incorporate yolk. Mating provides a stimulus which is essential for the initiation and maintenance of corpus allatum activity and, therefore, vitellogenesis. Corpus allatum activity (and vitellogenesis) can be induced in virgin females either by cutting the allatic nerves or by excision of a certain part of the protocerebrum. It is concluded that the corpus allatum of virgin females is inhibited nervously by the brain and that copulation provides a nervous stimulus which lifts this cerebral inhibition thereby permitting allatum activity.The median neurosecretory cells are not essential for vitellogenesis in Dindymus; however, it is suggested that they are necessary for maximum allatum activity. Evidence is provided to show that the growth and maintenance of the previtellogenic oöcytes are also under the control of the corpus allatum. A mechanism is described whereby the number of vitellogenic oöcytes in the ovarioles is maintained constant.     

Activity patterns and egg production in Coccophagus bartletti, an aphelinid parasitoid of scale insects

ABSTRACT.

• 1 Daily patterns of oviposition and host-feeding were examined in Coccophagus bartletti Annecke & Insley. Females began to host-feed and oviposit during the second or third day after emergence. Thereafter, both activities occurred regularly.
• 2 During long observation periods (5 h) most oviposition (93%) and host-feeding (90%) occurred within the first 3.5 h of wasps first encountering hosts. Experiments demonstrated that levels of activity were low for the rest of the day, and nocturnal oviposition occurred only if wasps had no alternative.
• 3 Dissection of female wasps that had been exposed to hosts, or withheld from them, for given periods of time, revealed that activity levels are governed by egg availability. Dispersal activity may also be influenced by the physiological state of the ovaries.
• 4 Production of a full complement of eggs (at 24±1°C (12 h L) and 18±1°C (12 h D)) took 48h or longer after host-feeding, and if wasps were withheld from hosts and provided with honey, the effects of egg resorption could be detected after about 10 days. Trends in oögenesis and oösorption in C. bartletti females seem not to conform with interpretations of oögenesis-oösorption cycles in other parasitoids.
• 5 The pattern of activity exhibited by C. bartletti females is not inflexible, but the major aspects mentioned above are species-specific. In general, information is needed about daily and hourly patterns of parasitoid oviposition and host-feeding before experiments are designed to test theories of parasitoid behaviour. Interpretation may otherwise rest on assumptions about their physiological condition.

     

On the relative importance of juvenile hormone and vitellogenin for oöcyte growth in the cockroach Nauphoeta cinerea

The corpora allata of castrated females of Nauphoeta grow only very slightly and do not reach a volume greater than that of the glands of normal females during gestation. These small corpora allata are, however, active and are responsible for the synthesis of vitellogenin (female specific protein) in large amounts. Besides vitellogenin the other haemolymph proteins are also synthesized and accumulated in the haemolymph in much higher concentrations than in normal females. Implanted oöcytes grow in castrated as well as in normal females at about the same rate until the tenth day of the oöcyte maturation period. Thereafter they only grow in castrated females. If castrated and normal females are decapitated, their protein content decreases. At the same time the growth stimulating capacity of their haemolymph decreases at a much faster rate. If oöcytes are implanted in castrated and decapitated females after 4 days they cannot grow any more although the vitellogenin titre of the haemolymph is still much higher than it is at any time in normal females. It can be concluded that vitellogenin alone cannot induce oöcyte growth and that juvenile hormone is necessary as well for vitellogenin synthesis as for its incorporation into the oöcytes. However, in insects rich in vitellogenin juvenile hormone leads to a more rapid oöcyte growth than in insects containing only small amounts of this protein.     

Meiotic parthenogenesis and heterochromatization in a soft scale,pulvinaria hydrangeae (Coccoidea: Homoptera)

Summary Meiotic parthenogenesis of a type not previously described was found in Pulvinaria hydrangeae Steinweden. During diakinesis 8 bivalents were formed. At prometaphase the spindle was tripolar but anaphase I was bipolar and normal. After completion of division of the primary oocyte, the following sequence occurred: 1. polar body I divided, usually into 3 products; 2. the secondary oocyte divided to yield the egg pronucleus and polar body II; 3. the egg pronucleus divided into its two haploid products; and 4. the second polar body divided. The products of the egg pronucleus fused while dividing to restore the diploid chromosome number; this division may be equated to the first cleavage division. The products of the polar bodies did not take part in the formation of the embryo proper or the mycetocytes.Among the embryos produced by females of two out of the three populations studied some of the embryos showed a heterochromatic chromosome set, characteristic of males in this and related families. The reproductive system of the females as well as the eggs did not contain any sperm; thus the male embryos were apparently produced parthenogenetically.The euchromatic and heterochromatic chromosome sets were genetically identical, since they both originated from the egg pronucleus by mitosis. The heterochromatization of one set but not the other might be due in part to a previous difference in their position in the cytoplasm.The females were completely homozygous yet they produced male and female embryos. Thus it appears that sex determination in the group does not depend on the segregation of genetic factors in either males or females.In addition to male and female embryos, three types of degenerating embryos were observed. It is believed that these embryos were formed by polyploid somatic cells which invaded abnormal eggs and embryos and took over development.