Termite queens have disproportionately more DNA in their fat body cells: reproductive division of labor and endoreduplication

Increases in DNA content caused by endoreduplication are widely observed in the metabolically active tissues of plants and animals. During egg production, insect females synthesize very large amounts of vitellogenin in their fat bodies, and female fat bodies of some insects become polyploid to accelerate vitellogenin production. Social insects have developed reproductive division of labor, wherein queens lay most of the eggs while other individuals have reduced fertility and undertake tasks required for maintaining the colony. Therefore, only queens are engaged in vitellogenin synthesis for egg production in social insects. Here, we show that termite queens have disproportionately more DNA in their fat body cells. Our DNA content analysis using flow cytometry demonstrated that more cells contained 4C‐DNA than 2C‐DNA in the fat bodies of Reticulitermes speratus queens. This high level of endoreduplication was not found in the fat body cells of other castes. This caste‐dependent doubling of DNA content in fat body cells suggests that termites exploit endoreduplication to boost egg production, in conjunction with the development of reproductive division of labor. This study highlights nuclear polyploidization as an adaptive strategy in social insects.     

Nutritional status and mortality of highland children in Nepal: Impact of sociocultural factors

This study investigates the nutritional status of native children in the highlands of Nepal (1,700–3,000 m) and explores the relationship between child mortality and surviving children's nutritional status. A random sample of 145 households from 11 villages in the Koshi Hill Zone in east Nepal was surveyed, and the nutritional status of the 438 children <14 years of age living in these households was assessed by means of anthropometry. We found a severe growth retardation in the Nepalese children compared to lowland reference groups as well as to highland children from the Andes. Child mortality and altitude are not significantly different between higher (Brahman and Chetri) and lower (Baisya and Sudra) caste households. A lower caste status and higher altitude of the household is associated with a significantly better nutritional status in offspring. In multiple regression analyses, improved nutritional status in children is significantly associated with lower caste (P = 0.001), higher altitude (P = 0.009), and less crowding (P = 0.001) but not with sibling mortality (P = 0.11). We thus conclude that nutritional status of children in households in the highlands of Nepal is associated with the household's socioeconomic status and altitude but not with mortality among siblings. © 1996 Wiley-Liss, Inc.     

乳白蚁不同品级的比例

乳白蚁Coptotermes在湖北宜昌地区每年11月～次年3月收集到的巢群,每年品级比例基本平衡。品级头比的数据一律来自1月份的巢群;品级重比是11月～次年3月的巢群。品级的比例分别为：工蚁占81．70％、干重比69．45％;兵蚁占7．50％、干重比7．69％;生殖蚁占10．80％、干重比26．76％。     

Body Size Shapes Caste Expression,and Cleptoparasitism Reduces Body Size in the Facultatively Eusocial Bees <Emphasis Type="Italic">Megalopta</Emphasis> (Hymenoptera: Halictidae)

We used the facultatively social sweat bee Megalopta genalis (Halictidae) to test whether body size is associated with social caste. Behavioral observations showed that non-reproductive foragers were significantly smaller than reproductive nest mate queens, and foragers were also smaller than presumed pre-dispersal reproductives. Moreover, among females from field-collected nests without behavioral observations, relative body size correlated with relative ovary size. Reproductive status is not a direct result of body size, as body size was not significantly associated with either ovary size or fecundity among both solitary and social reproductives. Reproductive status is apparently an outcome of social competition for reproductive dominance, and status is influenced by size relative to nest mates. Our study is the first to demonstrate an association of body size with caste expression in a facultatively social species with relatively weak seasonal constraints on independent nesting. Larvae of a parasitic fly (Fiebrigella sp., Chloropidae) consume pollen provisions stored in nest cells of M. genalis and M. ecuadoria. We tested whether fly parasitism of M. genalis reduces body size. Parasitized females are significantly smaller as adults than their unparasitized nestmates. This reduction is of a similar magnitude to the size differences between castes, and has the potential to shape host reproductive options by influencing competition with nest mates. We present data on the prevalence of parasitism from four collections of M. genalis and two collections of M. ecuadoria from Barro Colorado Island, Panama, and La Selva, Costa Rica.     

Larval cannibalism and worker-induced separation of larvae in <Emphasis Type="Italic">Hypoponera</Emphasis> ants: a case of conflict over caste determination?

Reproductive altruism and cooperative brood care are key characteristics of eusocial insects and reasons for their ecological success. Yet, Hymenopteran societies are also the stage for a multitude of intracolonial conflicts. Recently, a conflict between adult and larval colony members over caste fate was described and evidence for overt conflict was uncovered in several bee species. In theory, diploid larvae of many Hymenopteran species should experience strong fitness benefits, if they would be able to change their developmental pathway towards the queen caste. However, larval self-determination potential is low in most advanced eusocial Hymenopterans, because workers often control larval food intake and queenworker caste dimorphisms are generally high. In the ant genus Hypoponera, larvae actively feed on food provided by workers and here we show extremely low queenworker size differences in these ants: the lowest in H. opacior, where fertile wingless (intermorphic) queens weigh on average only 13% more than workers. Thus, slightly better nutrition during development might change the fate of a Hypoponera larva from a completely sterile worker to a fertile queen. One possibility to obtain extra food for Hypoponera larvae with their well-developed mandibles would be to cannibalise adjacent larvae. Indeed, we observed frequently larval cannibalism in ant nests. Yet, adult workers apparently try to prohibit larval cannibalism by carefully separating larvae in the nest. Larvae, which were experimentally brought into close contact, were rapidly set apart. Workers further sorted larvae according to size and responded swiftly to decreasing food levels, by increasing inter-larval distance. Still, an experimental manipulation of the larval cannibalism rate in H. schauinslandi failed to provide conclusive evidence for the link between larval cannibalism and caste development. Hence, further experiments are needed to determine whether the widespread larval cannibalism in Hypoponera and the untypical brood distribution can be explained by an overt caste conflict. Received 18 December 2006; revised 2 August and 20 September 2007; accepted 21 September 2007.     

Lipid stores, ovary development, and brain gene expression in Polistes metricus females

In order to gain insights into the mechanistic basis of caste and behavioral differences in Polistes paper wasps, we examined abdominal lipid stores and ovary development in Polistes metricus females in four groups: foundresses, queens, workers, and gynes. Queens had the largest ovaries, followed by foundresses, workers, and gynes. Gynes had 6x higher lipid stores than the other groups, and lipid stores were lower in foragers (foundresses, workers) than non-foragers (queens, gynes). Lipid levels and ovary development were negatively correlated across the four groups, but removing gynes from the analysis revealed a significant positive correlation for foundresses, workers, and queens, suggesting different energy allocation strategies for gynes vs. other groups. Expression levels of 9 genes (including three in the insulin pathway), examined in a previous study, correlated with either lipid stores or ovary development. These correlative results suggest important relationships between nutrition, reproduction, and division of labor in primitively social insects. We also show that it is possible to assign P. metricus females to one of the four female groups on the basis of wing wear (an indicator of foraging experience), lipid stores, and ovary development, which can facilitate caste-specific collections for future studies. Received 23 May 2008; revised 6 November 2008; accepted 20 November 2008.     

Artificial selection on ant female caste ratio uncovers a link between female‐biased sex ratios and infection by Wolbachia endosymbionts

Social insect sex and caste ratios are well‐studied targets of evolutionary conflicts, but the heritable factors affecting these traits remain unknown. To elucidate these factors, we carried out a short‐term artificial selection study on female caste ratio in the ant Monomorium pharaonis. Across three generations of bidirectional selection, we observed no response for caste ratio, but sex ratios rapidly became more female‐biased in the two replicate high selection lines and less female‐biased in the two replicate low selection lines. We hypothesized that this rapid divergence for sex ratio was caused by changes in the frequency of infection by the heritable bacterial endosymbiont Wolbachia, because the initial breeding stock varied for Wolbachia infection, and Wolbachia is known to cause female‐biased sex ratios in other insects. Consistent with this hypothesis, the proportions of Wolbachia‐infected colonies in the selection lines changed rapidly, mirroring the sex ratio changes. Moreover, the estimated effect of Wolbachia on sex ratio (~13% female bias) was similar in colonies before and during artificial selection, indicating that this Wolbachia effect is likely independent of the effects of artificial selection on other heritable factors. Our study provides evidence for the first case of endosymbiont sex ratio manipulation in a social insect.     

Polymorphism in the neotropical termite Serritermes serrifer

Serritermitidae (Isoptera) is a small and little known Neotropical termite family which includes only two genera: Glossotermes and Serritermes. Despite the lack of detailed studies, it has been assumed that these termites have a true worker caste. A recent study revealed that Glossotermes has a linear development pathway and lacks true workers. Here, we present a study of the polymorphism of Serritermes serrifer Hagen & Bates, a species endemic to the Cerrado ecoregion of central Brazil which lives as an inquiline inside nests of Cornitermes spp. A morphometric analysis was performed based on measurements taken of 11 body parts of 544 specimens of immatures, worker‐like individuals, soldiers, and alates. Sex of specimens was determined by dissection and examination of the seventh sternite. A principal component analysis (PCA) was used to evaluate morphological changes during development. Contrary with previous information from the literature and similar to Glossotermes, Serritermes shows a linear development pathway with two larval instars, two sizes of pseudergates, and a single nymphal morph. Pseudergates apparently undergo stationary molts. Sex ratio among pseudergates is male‐biased, but not as strongly as in Glossotermes. Typical colonies have a single physogastric primary queen and a single primary king. Ergatoid reproductives are relatively rare and some female ergatoids may become strongly physogastric. Nymphoid reproductives were not found. All soldiers are male and bear well‐developed testes.     

Relationship of queen number and worker size in polygyne colonies of the fire antSolenopsis invicta

Summary We examined the relationship between queen number and worker size in colonies of the fire antSolenopsis invicta. Worker size in monogyne colonies was significantly greater than in polygyne colonies; furthermore, polygyne colonies snowed a strong negative linear relationship between queen number and worker size. Higher queen pheromone level and/or decreased food availability accompanying an increase in queen number likely play important roles in producing the observed patterns.