Population differences in host use by a seed-beetle: local adaptation, phenotypic plasticity and maternal effects

For insects that develop inside discrete hosts, both host size and host quality constrain offspring growth, influencing the evolution of body size and life history traits. Using a two-generation common garden experiment, we quantified the contribution of maternal and rearing hosts to differences in growth and life history traits between populations of the seed-feeding beetle Stator limbatus that use a large-seeded host, Acacia greggii, and a small-seeded host, Pseudosamanea guachapele. Populations differed genetically for all traits when beetles were raised in a common garden. Contrary to expectations from the local adaptation hypothesis, beetles from all populations were larger, developed faster and had higher survivorship when reared on seeds of A. greggii (the larger host), irrespective of their native host. We observed two host plant-mediated maternal effects: offspring matured sooner, regardless of their rearing host, when their mothers were reared on P. guachapele (this was not caused by an effect of rearing host on egg size), and females laid larger eggs on P. guachapele. This is the first study to document plasticity by S. limbatus in response to P. guachapele, suggesting that plasticity is an ancestral trait in S. limbatus that likely plays an important role in diet expansion. Although differences between populations in growth and life history traits are likely adaptations to their host plants, host-associated maternal effects, partly mediated by maternal egg size plasticity, influence growth and life history traits and likely play an important role in the evolution of the breadth of S. limbatus’ diet. More generally, phenotypic plasticity mediates the fitness consequences of using novel hosts, likely facilitating colonization of new hosts, but also buffering herbivores from selection post-colonization. Plasticity in response to novel versus normal hosts varied among our study populations such that disentangling the historical role of plasticity in mediating diet evolution requires the consideration of evolutionary history.     

Effects of temperature on reproductive output, egg provisioning, juvenile hormone and vitellogenin titres in the butterfly Bicyclus anynana

Environmentally induced phenotypic plasticity is common in nature. Hormones, affecting multiple traits and signaling to a variety of distant target tissues, provide a mechanistic link between environments, genes and trait expression, and may therefore well be involved in the regulation phenotypic plasticity. Here, we investigate whether in the tropical butterfly Bicyclus anynana temperature-mediated plasticity in egg size and number, with fewer but larger eggs produced at lower temperatures and vice versa, is under control of juvenile hormone, and whether different temperatures cause differences in egg composition. Female B. anynana butterflies showed the expected response to temperature, however, we found no evidence for an involvement of juvenile hormone. Neither haemolymph JH II and JH III titres nor vitellogenin levels differed across temperatures. The smaller eggs produced at the higher temperature contained relatively higher amounts of water, free carbohydrates and proteins, but relatively lower amounts of lipids. While these smaller eggs had a lower absolute energy content, total reproductive investment was higher at the higher temperature (due to a higher fecundity). Overall, our study indicates that temperature-mediated plasticity in reproduction in B. anynana is mechanistically related to a biophysical model, with oocyte production (differentiation) and oocyte growth (vitellogenesis) having differential temperature sensitivities.     

Developmental plasticity and maternal effects of reproductive characteristics in the frog,Bombina orientalis

Summary Life history theory suggests that reproductive characteristics such as ovum size and clutch size should be well buffered against vararies of the environment. However, studies which demonstrate environmental sensitivity of reproductive characteristics are increasing in number, as are studies which find that maternal effects are responsible for much of the variation in developmental and growth rates in embryonic and larval fish and amphibians. The data reported here demonstrate that the environment, in terms of temperature and food availability that a specific individual encounters during vitellogenesis, exerts a strong influence on both egg size and number. Warmer temperatures and less food decrease ovum size, while colder temperatures and less food decrease clutch size. The variation in ovum size that is induced by the environment can exert a strong influence on variation in offspring development and growth and serve as an excellent model for studies on the evolution of developmental plasticity.     

Reproductive response of Euterpina acutifrons in two estuaries of the Basque Country (Bay of Biscay) with contrasting nutritional environment

The reproductive response involving egg number and size was studied for the egg carrying copepod Euterpina acutifrons in two estuaries with different nutritional environment (Bilbao and Mundaka). To characterise the available food for copepods, the concentration of particulate organic matter (POM), the combined concentration (PGL) and proportion of particulate protein, carbohydrate and lipids and the concentration of chlorophyll a were measured in the <20 μm seston fraction. Data on female size (prosome length) and abiotic factors, such as temperature, salinity and oxygen saturation in water, were also obtained to relate them to reproductive and nutritional variables. Correlation and stepwise regression analyses indicated that temperature via female size was the main factor controlling brood volume in a temporal context, whilst the nutritional environment had a secondary role. Nevertheless, the nutritional environment caused noticeable between-estuary differences in the reproductive response of this species. ANOVA tests showed that E. acutifrons clutch size and brood volume were significantly bigger in the estuary of Bilbao, which was richer in both total (POM) and labile (PGL) organic particulate matter than the estuary of Mundaka. In contrast, egg size, female size and phytoplanktonic food quantity did not differ significantly between estuaries. These results suggest that the organic enrichment with particles of detritical-heterotrophic origin enhanced E. acutifrons clutch size and brood volume in Bilbao. Egg size was closely related to phytoplankton in both estuaries, but a clear trade-off between egg size and number per clutch was observed only in Mundaka. Thus, we suggest that the reduction of egg number in order to increase egg size might happen only under food-limiting conditions, but not in organically rich and quite constant nutritional environments such as found those in Bilbao.     

Fate of maternal conjugated ecdysteroids during embryonic development in Locusta migratoria

Newly laid eggs of Locusta migratoria contain impressively high concentrations of conjugated 2-deoxyecdysone and conjugated ecdysone of maternal origin. These molecules are metabolized during embryonic development, the changes concerning not only the ecdysteroid genins but also the conjugating moieties. In the present paper the fates of the maternal conjugates were followed during embryogenesis in the eggs. The conjugates were separated both by silica gel TLC and reverse-phase HPLC and measured, before and after hydrolysis, by RIA. Fluctuations of radioactive ecdysteroid conjugates were also investigated in eggs laid by females subjected to massive injections of tritiated cholesterol. The results are discussed in relation to recent data on identification of ecdysteroid conjugates in Locusta and a model for the sequences of metabolic events leading from maternal ecdysteroid conjugates to the embryonic ecdysteroids is proposed.     

Nutrients in fruit increase fertility in wild-caught females of large and long-lived Euphaedra species (Lepidoptera, Nymphalidae)

Fruit-feeding butterflies can experience a more nutrient rich adult diet than nectar-feeding species, and can be expected to use these nutrients for egg production. Here we compare life span, and reproduction parameters of wild-caught females of large and long-lived species on either a sucrose or a mashed banana diet. With small sample sizes per species, but rich longitudinal data for each individual, we examined the longitudinal reproduction pattern, egg size and hatchability of these butterflies in captivity. Diet significantly affected mortality in captivity in a time-dependent manner. On average, we found that butterflies fed mashed banana laid 1.855 times more eggs than those fed sugar. They laid significantly more eggs when they laid and conserved egg size with age while butterflies fed sucrose showed significantly declining egg sizes. Egg hatchability was not significantly affected by diet. Long pre-oviposition periods, significantly smaller first eggs, and absence of age at capture effects on intensity of reproduction indicate low reproduction rates in the field that are due to low food availability. With our small sample sizes, we did not detect significant differences between the species in their response to the diet treatments.     

Neuropeptide F and the corn earworm, Helicoverpa zea: A midgut peptide revisited

The neuropeptide Y family of peptides is implicated in the regulation of feeding across a broad range of animals, including insects. Among vertebrates, neuropeptide Y exerts its actions mainly centrally, whereas peptide YY and pancreatic polypeptide arise from digestive tissues. Among invertebrates, neuropeptide F (NPF) is the sole counterpart of the NPY family. Shared features of NPF sequences derived for Lepidoptera indicate that the midgut peptide (Hez-MP-I) of the corn earworm, Helicoverpa zea, characterized more than a decade ago, is a carboxyl fragment of a full-length NPF. An antibody to Hez-MP-I was used to characterize the peptide's distribution in tissues of larvae, pupae, and adults. Immunostaining demonstrated NPF-related material both in nervous tissues and in abundant endocrine cells of the midgut. Radioimmunoassay of Hez-MP-I in the head, midgut and hemolymph of fifth instar larvae revealed concentration changes corresponding to development and feeding state. As with the vertebrate homologs, NPF may arise both centrally and peripherally to modulate the physiology of feeding and digestion of Lepidoptera.     

Egg size and the evolution of phenotypic plasticity in larvae of the echinoid genus Strongylocentrotus

Planktotrophic larvae grow by utilizing energy obtained from food gathered in the plankton. Morphological plasticity of feeding structures has been demonstrated in multiple phyla, in which food-limited larvae increase feeding structure size to increase feeding rates. However, before larvae can feed exogenously they depend largely on material contained within the egg to build larval structures and to fuel larval metabolism. Thus, the capacity for plasticity of feeding structures early in development may depend on egg size. Using the congeneric sea urchins Strongylocentrotus franciscanus and S. purpuratus, which differ in egg volume by 5-fold, I tested whether the degree of expression of feeding structure (larval arm length) plasticity is correlated with differences in the size of the egg. I experimentally manipulated egg size of S. franciscanus (the larger-egged species) by separating blastomeres at the 2-cell stage to produce half-sized larvae. I reared half-size and normal-size larvae under high and low food treatments for 20 days. I measured arm and body lengths at multiple ages during development and calculated the degree of plasticity expressed by larvae from all treatments. Control and unmanipulated S. franciscanus larvae (from ∼ 1.0 nl eggs) had significantly longer arms relative to body size and a significantly greater degree of plasticity than half-sized S. franciscanus larvae (from < 0.18 nl eggs), which in turn expressed a significantly greater degree of plasticity than S. purpuratus larvae (from ∼ 0.3 nl eggs). These results indicate that egg size affects larval arm length plasticity in the genus Strongylocentrotus; larger eggs produce more-plastic larvae both in an experimental and a comparative context. However, changes in egg size alone are not sufficient to account for evolved differences in the pattern of plasticity expressed by each species over time and may not be sufficient for the evolutionary transition from feeding to non-feeding.     

Negative effects of elevated testosterone on female fecundity in zebra finches

Although factors influencing androgen deposition in the avian egg and its effects on nestling fitness are recently receiving considerable attention, little is known about the potential costs of high testosterone levels in the females. Our study aimed at determining the effect of injections of testosterone (T) in female zebra finches (Taeniopygia guttata), on clutch size, egg mass, yolk mass, and yolk androgen content. Females were given a single bolus injection of T in a range of doses after laying the first egg. Results show that administration of T negatively affected clutch size; the strength of this effect increased with increasing doses of T. Females injected with the highest testosterone dose showed suppressed oviposition of the third and the fourth eggs. Interestingly, testosterone administration made females produce eggs with relatively large yolks, suggesting that T may mediate the trade-off between number and size of eggs. Testosterone injection resulted in elevated levels of androgen in the eggs, in contrast to control clutches, which showed a decreasing pattern of androgen concentration along the laying sequence. We conclude that high androgen investment in eggs may be limited by physiological requirements of the ovulatory process.     

Interactive effects of time and vegetation on reproduction of redshanks (Tringa totanus) breeding in Wadden Sea salt marshes

As shown for various species, nesting waders are non-randomly distributed on wetlands and preferentially select riparian nest-sites adjacent to limnic or marine waterbodies. Studying the redshank Tringa totanus, we tested the hypotheses that, in a coastal wader species which conceals its clutch in vegetation, predation and hatching success are affected by vegetation zonation, and that breeding in lower salt marsh areas has negative consequences for reproduction. We further predicted effects of timing of breeding and breeding experience/age of adults potentially reflected by egg biometrics both on nest-site selection and reproduction. Effects of vegetation, space, time and individual quality on hatching success of redshanks were studied in the German part of the Wadden Sea. Dominant plant species, vertical vegetation structure and nest concealment varied significantly between nests. Variation in nest concealment was relatively low: about 90% of clutches were classified as being well concealed. This variation was explainable by vegetation structure but not by vegetation composition at the nest-site, distance to shoreline, and time of clutch initiation. Vertical vegetation structure varied by dominant plant species but not by distance to shoreline and time of clutch initiation. Hatching success of clutches was low (10.6%) due to high predation (daily predation rate: 7.4%). Hatching success and duration of clutch survival were negatively and predation positively related to the date of clutch initiation. Furthermore, negative relationships were found between egg size and predation and duration of survival, respectively. We assume that concealed nests, early breeding and breeding experience diminish predation in salt marsh breeding redshanks. Thus, redshank reproduction appears to be affected by interactive effects of timing of breeding and vegetation facilitating early breeding. In contrast to open-nesting species, breeding in riparian habitats next to waterbodies may be disadvantageous for species breeding concealed in vegetation if these are covered by less structured vegetation.     

Heritability estimates and maternal effects on tarsus length in pied flycatchers,Ficedula hypoleuca

Tarsus length has previously been shown to have an additive genetic component in some pied flycatcher populations. In addition to estimation of additive genetic variation by means of repeatability analyses at various ontogenetic stages and degrees of genetic resemblance, we explore in this paper variation among parent-offspring regressions between sexes and years, as well as the influence of hatching date, ectoparasite abundance, egg volume, and male and female condition on the tarsus length of their offspring. Mother-offspring regressions gave heritability estimates consistently higher than father-offspring regressions, although variation among years was large and both types of estimates yielded lower heritability values than those estimated by means of full-sib resemblance. This indicates that common environmental effects were inflating heritability estimates. There existed maternal effects via egg size, larger eggs fledging chicks with larger tarsi. Mean tarsus length of broods decreased with hatching date and, independently, with high loads of ectoparasitic, blood-feeding mites (Acari). The maternal effect via egg size persisted into the adulthood, and confounded the interpretation of differences between heritability slopes. We address the method of examining differences in parent-offspring regressions as a shorthand for estimating extra-pair copulation (EPC) rates. In our population, this method would give an EPC rate of 0–59%, depending on whether the analysis is performed with fledglings recruited to the breeding population or with offspring at the nest.     

Reproductive activity and physiological status of the calanoid copepods Calanus helgolandicus and Calanoides carinatus under food-limiting conditions

The reproductive activity and the physiological state of the calanoid copepods Calanus helgolandicus and Calanoides carinatus were investigated off the coast of NW Spain during autumn to evaluate the effect of short food resources on both populations. Phytoplankton biomass was low, and neither phytoplankton size distribution nor composition was suitable to support high reproductive rates. Accordingly, egg production rates (EPR) were much lower than maximum rates for both species, pointing to food limitation. The reproductive index (RI), which represents the proportion of females with mature gonads, was < 50% at each of the three zones into which the sampling area was divided (coast, shelf and ocean). Potential recruitment rates were very low except at some nearshore stations, where the highest concentrations of chlorophyll-a (Chl-a), diatoms, dinoflagellates and large cells were found. EPR of C. helgolandicus and C. carinatus were correlated with phytoplankton biomass and unaffected by temperature. Phytoplankton carbon ingestion explained ca. 50% of the variability in EPR for both species. At most of the stations, herbivory was insufficient to cover the carbon requirements for reproduction and respiration, so females probably fed on heterotrophic prey to meet their demands. However, given the low fecundity observed, this omnivorous diet did not seem to be optimum for reproduction, and a severe food limitation is thus suggested. Furthermore, the high C/N values measured point to a notable lipid storage, but given the low EPR found, lipid reserves were probably invested into female maintenance rather than into gonad maturation. C. helgolandicus and C. carinatus populations did not mirror phytoplankton biomass distribution, but they correlated well when considering only copepodites V (CV). The CV could be preparing for the overwintering, storing lipid reserves to ensure a successful diapause, and they could also be advected by the poleward current detected during the study. Females showed a diel feeding rhythm, with highest ingestion rates during night. From our results, it follows that C. helgolandicus and C. carinatus females did not perform diel vertical migrations. We suggest that this behaviour is likely due to the food-limiting conditions, which make it more advantageous to remain at the surface during daytime.     

Reproductive parameters in relation to food supply in the whiskered tern ( Chlidonias hybrida )

It is generally accepted that breeding terns are sensitive to food supply and that their reproductive effort could be substantially affected by the availability and access to resources. In this study we examined reproductive parameters in the whiskered tern Chlidonias hybrida in relation to food supply during the courtship feeding period (food brought by males to females) over a 2-year period (2004–2005). We also studied whether the condition of the nestlings [body condition index (BCI)] was related to a proxy of the reproductive investment of the adults (the clutch size) during a season of food shortage. Behavioural observations showed a decrease in the intensity of male courtship feeding between years (2004 > 2005), and a strong shift in the relative abundance of the two prey groups (invertebrates/fish; invertebrate prey dropped from 88.0 to 49.3%) brought by males. This change in food delivery rates did not result in a delay in laying, but there was a significant difference in mean clutch size between years (2.71 ± 0.49 eggs in 2004 and 2.05 ± 0.78 eggs in 2005) without any within-year variation in relation to the laying dates. The egg size (volume and length) was related to the year (2004 < 2005), suggesting a trade-off in the quantity and the quality of eggs between the two seasons. We also found no evidence that the investment in a large clutch affected nestling BCI in the course of the food shortage season. Since many pairs (about 60%) interrupted breeding during the incubation stage, we assumed that parents that succeeded in rearing nestlings in these conditions were probably ‘high-quality’ individuals. Our results therefore showed that whiskered terns are sensitive to the varying food conditions they experienced throughout the courtship period. The diversity of prey types could be a key factor in the reproductive investment of this tern species.     

Interactions between size and temperature influence fecundity and longevity of a tortricid moth,Zeiraphera canadensis

Females of Zeiraphera canadensis Mut. & Free., the spruce bud moth, were reared in the laboratory at constant and alternating temperatures, and in an outdoor insectary, to (1) determine the effects of temperature, age and size on several reproductive parameters and, (2) to test the hypothesis that body size-temperature interactions influence longevity and realized fecundity. Egg maturation was linearly related to age and large moths developed eggs at a higher rate than small ones. Mcan lifetime oviposition rate reached a maximum and remained stable at temperatures 20° C while the mean lifetime rate of egg maturation increased linearly with temperature, indicating that higher temperatures adversely affect oviposition. The production of nonviable eggs increased with age but also with temperature, suggesting high temperature (25° C) reduces egg quality and/or hinders fertilization. The realized fecundity and longevity of females reared under an alternating temperature regime (mean 20° C) was significantly less than that of females reared at constant 20° C. Similar realized fecundity, longevity and mean lifetime oviposition rates for females reared at temperatures alternating between 10 and 25° C (mean 20° C) and those at constant 25° C reflected the inability of females to recover from elevated diurnal temperatures. Longevity was positively related to female body size at constant 15 and 20° C but the relationships were negative for moths exposed to diurnal temperatures equal to or exceeding 25° C. Due to the reduced longevity of large moths at high temperatures, linear regressions between size and realized fecundity were only significant at constant temperatures 20° C. At higher temperatures, the size-fecundity relationship became curvilinear as a result of the diminished reproductive output of large individuals. Reduced fecundity and longevity of large females at high temperatures may have been due to elevated internal temperatures of large-bodied moths. Large females in a controlled-environment chamber maintained at 25° C developed an internal temperature excess (i.e. temperature above ambient) of nearly 2° C while small-bodied females exceeded ambient by only 0.3° C. However, when held at 20° C, the temperature excess of large-bodied moths was much less than 1° C and small-bodied females did not differ from ambient. Such interactions between temperature and body size suggest that there should be stabilizing selection toward moderate-sized individuals and may explain the absence of size-related effects on fecundity and longevity previously reported for several other lepidopterans.     

The interactive effects of temperature,food level and maternal phenotype on offspring size in Daphnia magna

Invertebrate offspring are usually larger in colder environments. To test for possible effects of covariates (e.g. maternal phenotype and feeding conditions) on this pattern, we performed a laboratory experiment to look at the effect of temperature on newborn weight in the planktonic crustacean Daphnia magna. Three tempèratures (12°C, 16°C and 22°C) and two food levels (10,000 cells ml^–1 and 150,000 cells ml^–1) were used, and offspring were examined from the first five clutches of mothers that had been maintained under the constant experimental conditions for three generations. Preliminary analysis suggested that newborn weight was significantly affected by temperature although patterns in the data were not clear cut. In addition, the covariates mother weight and clutch size were positively and negatively correlated with newborn weight, respectively; and later clutches tended to contain heavier offspring. Therefore, in an effort to control for the effects of the covariates, repeated-measures analysis of covariance was performed using ratio values of newborn weight/mother weight (relative newborn weight) as the dependent variable, clutch size as the covariate and clutch number as the repeated measures term. Now, temperature as a main effect in an ANCOVA model did not significantly influence relative newborn weight. The repeatedmeasure term clutch number also became nonsignificant, indicating that when differences in mother weight due to age were accounted for there were no overall differences in relative newborn weight between clutches from a particular mother. Temperature effects on relative newborn weight were only significant as part of interaction terms with food concentration and with clutch number. Thus there were different weight responses to temperature within food levels, and between clutch numbers within food levels. Under the low-food conditions newborn were heaviest at 16°C, lightest at 12°C and intermediate at 22°C. Conversely, under the high-food condition newborn were lightest at 16°C, heaviest at 12°C and again intermediate at 22°C. However, newborn tended to be heavier under the low food condition, and food concentration was highly significant as a main effect. Mother growth rate showed no significant relationship with newborn weight. It is concluded that direct temperature effects on relative newborn weight are marginal and nonsignificant. Temperature effects through interactions with food concentration and clutch number are important determinants of newborn weight, but relatively speaking account for only a small proportion of observed variance in newborn weight (25%), compared with the direct effect of food concentration (67%).     

Genetic and maternal influences on life history plasticity in milkweed bugs (Oncopeltus): response to temperature

This study was designed to examine life history flexibility arising from phenotypic plasticity in response to temperature and from maternal effects in response to reproductive diapause in a temperate zone population of the milkweek bug (Oncopeltus fasciatus). We employed a split-family, first-cousin, full-sib design with siblings reared at different temperatures in order to quantify phenotypic plasticity, maternal effects, and variation for each. The following traits were analyzed: development time, age at first reproduction, longevity, early-life fecundity, and wing length. We found both life history plasticity and maternal effects on life history traits which tend to enhance the colonizing ability of offspring born to mothers that have undergone reproductive diapause. We were unable to demonstrate additive genetic variation for plasticity for any of the traits, while for development time and wing length we found variation due to non-additive genetic or common-environmental sources. We were also unable to demonstrate additive genetic variation for maternal effects, although variation may exist at low levels that are difficult to detect using cousin-families. The apparent lack of variation in this population would constrain evolution of life history flexibility even though considerable flexibility exists in the phenotype.     

Territorial behaviour of beaugregory damselfish (Stegastes leucostictus) in response to egg predators

To test the hypothesis that territorial male beaugregory damselfish (Stegastes leucostictus) can assess the degree of threat from egg predators, the aggressive behaviour of both males guarding eggs and males without eggs was recorded when they were presented with either one or five individuals of the wrasse egg predator Thalassoma bifasciatum. Presentations were made in a clear plastic bottle 0.5 m from artificial beaugregory breeding sites in the field. Two measures of aggressive behaviour (downhead threats and time spent within 10 cm of the presentation bottle) were significantly greater for multiple wrasse presentations than for single wrasse presentations, while a third measure (chase attempts) was not. In contrast, chase attempts were significantly greater for males guarding eggs than for males without eggs, but there was no significant difference in the level of downhead threats or bottle proximity time for this comparison. Levels of the three aggressive measures were not correlated with either the amount of eggs or the number of egg clutches present. These results indicate that territorial male beaugregories respond to an increase in the number of intruding egg predators by increasing the levels of low-cost threat behaviour, while the presence (but not the amount) of eggs results in an increase in the amount of higher cost attack behaviour. These experiments, therefore, support the hypothesis that male beaugregories can assess the degree of threat from egg predators, and indicate that they use this assessment to alter the strategies used in territorial defence.     

Cortical anchorages and cell type segregations of maternal postplasmic/PEM RNAs in ascidians

Ascidian postplasmic/PEM RNAs constitute a large class of cortical maternal RNAs which include developmental determinants (macho-1 and pem-1). We have analyzed the localization, cortical anchorage and cell type segregation of postplasmic/PEM RNAs in Ciona intestinalis and Phallusia mammillata using very high-resolution fluorescent in situ hybridization. We also compared RNAs extracted from whole oocytes and from isolated cortices using microarrays and localized RNAs possessing clusters of xCACx motifs in their 3′UTRs.Based on these combined approaches we conclude that: (1) the vast majority of the 39 postplasmic/PEM RNAs (including vasa) are localized in the egg cortex. (2) Many postplasmic/PEM RNAs 3′UTR are enriched in xCACx motifs, allowing us to identify 2 novel postplasmic/PEM RNAs (PSD and MnK). (3) Postplasmic/PEM RNAs anchored to cortical Endoplasmic Reticulum (cER) and those associated with granules have different cell destinations.We propose that there are 2 distinct categories of postplasmic/PEM RNAs on the basis of their cortical anchorages and cell destinations: (1) macho-1-like postplasmic/PEM RNAs anchored to cER which segregate into somatic B8.11 cells. (2) vasa-like postplasmic/PEM RNAs associated with granules which in addition to B8.11 cells segregate into B8.12 germ cells.