Evolution of eusociality and the soldier caste in termites: a validation of the intrinsic benefit hypothesis

In termites the evolution of reproductive altruism is not based on a particularly high relatedness between nestmates. For the evolution and maintenance of the ancestral sterile soldier caste, the benefits generated by the soldiers' presence must compensate the loss of the soldiers' reproductive potential. To study the impact of soldiers on colony's fitness, we manipulated the proportion of soldiers to nonsoldiers in colonies of the dry-wood termite Cryptotermes secundus.'Soldier-less' colonies were obtained by removing soldiers and inhibiting their development with an extract of soldier heads. The colonies were set up for 1 year in experimental nests in the field. 'Soldier-less' colonies produced fewer soldiers. The reduction of soldiers neither affected colony survival nor helper growth, but fewer dispersing sexuals were produced in 'soldier-less' than in control colonies. This confirms what was only supposed so far, that in termites soldiers are maintained for their intrinsic benefit to cost ratio.     

Reproductive schedule and factors affecting soldier production in the eusocial bamboo aphid Pseudoregma bambucicola (Homoptera, Aphididae)

Summary: The reproductive characteristics of the soldier-producing aphid Pseudoregma bambucicola were studied in Kagoshima, Southern Japan, to know the factors affecting soldier production of eusocial aphids. The soldier proportion in aphid colonies was highest from October to November. In some large colonies, soldiers were observed in all seasons except in July when colony size was relatively small. Multiple regression analysis showed that the colony size was a principal factor affecting soldier proportion throughout a year. Other social or environmental factors such as aphid composition, host plant conditions and predator abundance were not always significant. Rearing experiments revealed that large colonies (̿,000 individuals) produced soldiers in almost all seasons while small colonies (<1,000) never produced any soldiers. The caste-production schedule of adult females was examined in the field. When solitary females produced both castes, they usually produced normal nymphs first and then soldiers. Females from large colonies tended to produce more soldiers in the earlier period of their lifetime, whereas females from newly established small colonies produced no or only a few soldiers at later times. The average number of soldiers and normal nymphs produced consecutively by a single female was >10 and >20, respectively. Because they have a small number of ovarioles (<15 on average), females should alter caste production within the same ovarioles according to changes in environmental conditions. Artificial removal or introduction of predators and reduction of colony size did not affect soldier production over two successive generations, revealing maternal effects on soldier production. Females cannot shift caste production quickly in response to changes in predator abundance and colony size. This is probably due to early developmental determination of castes within the mother's body.     

Two closely related species of desert carpenter ant differ in individual-level allocation to fat storage

Comparison of closely related species that differ in their life histories is a powerful method for studying the underlying physiological mechanisms contributing to life-history variation. I investigated whether two closely related members of the Camponotus festinatus species complex of desert carpenter ants, C. nr. festinatus Desert Light and C. nr. festinatus Desert Dark, differed in their life-history tactics with respect to fat storage. Newly mated queens were collected in the field, and colonies were reared under common conditions in the laboratory for 2 yr before sampling. I show that the two species differ in fat storage at the individual level. While the basic scaling relationship between lean mass and fat content did not differ between the two species, Dark workers and soldiers stored significantly more fat per unit lean mass than Light workers or soldiers. There were no significant demographic differences in the proportions of workers or soldiers involved in fat storage between the two species, although there was a trend toward Light colonies having a greater proportion of soldiers storing large amounts of fat. There was also no significant difference in the total amount of fat stored by the two species at the colony level. The detection of strong individual-level effects but no colony-level effects was likely due to the low statistical power of colony-level analyses. Showing that these two closely related species differ in fat storage at the individual level in a common environment demonstrates their utility as a model for understanding the physiological and behavioral mechanisms regulating life-history variation in fat storage in ants.     

Conflicting Notions on Violence and PTSD in the Military: Institutional and Personal Narratives of Combat-Related Illness

Research indicates that soldiers struggling with PTSD under-utilize mental health care. Quantitative studies of barriers to care point to the importance of soldiers’ beliefs about mental health and mental health interventions in their care-seeking behavior, yet these studies still struggle to understand the particular beliefs involved and the ways they impact care-seeking behavior. This preliminary study makes a start in examining these questions through qualitative literature analysis. It maps out dominant messages surrounding PTSD in military mental health interventions, and explores how they can both shape and conflict with soldiers’ personal notions. It does so by analyzing these messages and notions as institutional and personal (illness) narratives. Institutional military PTSD-narratives, which draw on mainstream scientific and clinical models, appear to communicate contradictory notions on the meanings of violence and its psychological consequences, often without acknowledging these contradictions. As such, these narratives seem to shape struggles of soldiers, both within themselves and with the military institution. The identified conflicts indicate, contrary to the individualizing and decontextualizing focus of dominant PTSD-understandings, that soldiers’ struggles also have social and moral dimensions. This has important implications for both research into PTSD-interventions and understandings of PTSD as such.     

海军指战员心理健康状况研究现状

我国军人的心理健康状况从80年代后期开始有了较为系统的研究。海军指战员是社会的特殊群体,健康的心理素质可以使海军指战员保持战斗力,更好的适应国防现代化的需要。海军指战员在一段较长的时间内,整日面对单一枯燥的环境,承受巨大工作压力,加之其他一些影响因素,难免产生诸多心理问题。因此,探讨职业特殊环境对海军指战员心理健康造成的影响,对海军指战员人格结构及心理健康状况进行评价,用以指导海军官兵在特殊环境中提高心理素质,避免不良影响,最终提高部队战斗力具有十分重要的意义。通过查阅文献了解海军特殊职业环境对士兵心理健康影响的研究现状,总结当前应用的各种评价工具、异常心理状况、影响因素及其危害,分析海军心理健康研究方面存在的不足与亟待解决的问题,为今后的研究提供参考。     

海军指战员心理健康状况研究现状

我国军人的心理健康状况从80年代后期开始有了较为系统的研究。海军指战员是社会的特殊群体,健康的心理素质可以使海军指战员保持战斗力,更好的适应国防现代化的需要。海军指战员在一段较长的时间内,整日面对单一枯燥的环境,承受巨大工作压力,加之其他一些影响因素,难免产生诸多心理问题。因此,探讨职业特殊环境对海军指战员心理健康造成的影响,对海军指战员人格结构及心理健康状况进行评价,用以指导海军官兵在特殊环境中提高心理素质,避免不良影响,最终提高部队战斗力具有十分重要的意义。通过查阅文献了解海军特殊职业环境对士兵心理健康影响的研究现状,总结当前应用的各种评价工具、异常心理状况、影响因素及其危害,分析海军心理健康研究方面存在的不足与亟待解决的问题,为今后的研究提供参考。     

Antifungal activity in thrips soldiers suggests a dual role for this caste

The social insect soldier is perhaps the most widely known caste, because it often exhibits spectacular weapons, such as highly enlarged jaws or reinforced appendages, which are used to defend the colony against enemies ranging in size from wasps to anteaters. We examined the function of the enlarged forelimbs of soldiers (both male and female) of the eusocial, gall-inhabiting insect Kladothrips intermedius, and discovered that they have little impact on their ability to repel the specialized invading thrips Koptothrips species. While the efficacy of the enlarged forelimb appears equivocal, we show that soldiers secrete strong antifungal compounds capable of controlling the specialized insect fungal pathogen, Cordyceps bassiana. Our data suggest that these thrips soldiers have evolved in response to selection by both macro- and micro-organisms. While it is unknown whether specialized fungal pathogens have been major selective agents in the evolution of the soldier caste in general, they were probably present when sociality first evolved and may have been the primordial enemies of social insects.     

Armor and anesthesia: exposure, feeling, and the soldier's body

For many civilians, the high-tech weapons, armor, and military medicine with which U.S. soldiers are equipped present an image of lethal capacity and physical invulnerability. But, as this article explores, soldiers themselves just as often associate the life-sustaining technology of modern warfare with feelings that range from a pragmatic ambivalence about exposure to harm all the way to profoundly unsettling vulnerability. This article, based on fieldwork among soldiers and military families at the U.S. Army's Ft. Hood, examines sensory and affective dimensions of soldiers' intimate bodily relationships with the technologies that alternately or even simultaneously keep them alive and expose them to harm. I argue that modern military discipline and technology conspire to cultivate soldiers as highly durable, capable, unfeeling, interchangeable bodies, or what might be called, after Susan Buck-Morss (1992), anesthetic subjects. But for soldiers themselves, their training, combat environment, protective gear, and weapons are a rich font of both emotional and bodily feeling that exists in complex tension with the also deeply felt military imperative to carry on in the face of extreme discomfort and danger.     

Could Work Be a Source of Behavioural Disorders? A Study in Horses

Stress at work, as shown by a number of human studies, may lead to a variety of negative and durable effects, such as impaired psychological functioning (anxiety, depression…). Horses share with humans this characteristic of working on a daily basis and are submitted then to work stressors related to physical constraints and/or more “psychological” conflicts, such as potential controversial orders from the riders or the requirement to suppress emotions. On another hand, horses may perform abnormal repetitive behaviour (“stereotypies”) in response to adverse life conditions. In the present study, we investigated whether the type of work the horses are used for may have an impact on their tendency to show stereotypic behaviour (and its type) outside work. Observations in their box of 76 horses all living in the same conditions, belonging to one breed and one sex, revealed that the prevalence and types of stereotypies performed strongly depended upon the type of work they were used for. The stereotypies observed involved mostly mouth movements and head tossing/nodding. Work constraints probably added to unfavourable living conditions, favouring the emergence of chronic abnormal behaviours. This is especially remarkable as the 23 hours spent in the box were influenced by the one hour work performed every day. To our knowledge, this is the first evidence of potential effects of work stressors on the emergence of abnormal behaviours in an animal species. It raises an important line of thought on the chronic impact of the work situation on the daily life of individuals.     

Association between caste and genotype in the termite Mastotermes darwiniensis Froggatt (Isoptera: Mastotermitidae)

Abstract Termite workers and soldiers differ markedly in their morphology and behaviour. We sought evidence for genetic influences on caste determination in the giant northern termite, Mastotermes darwiniensis , by investigating if workers and soldiers from the same colony differed genetically. The genotypes of 795 termites from 11 distinct colonies were assayed at six polymorphic microsatellite loci. We found that the multilocus genotypes of workers and soldiers from 8 of the 11 colonies did not differ significantly. Thus, the majority of the data provided no evidence for a genetic association with caste in workers and soldiers of M. darwiniensis . However, the genotype frequencies of workers and soldiers from three colonies differed, suggesting that genotype is occasionally associated with caste in this species. The genetic differentiation of castes within these colonies could reflect differences in the propensities of termites with distinct genotypes to develop into particular castes and provide a selective advantage to colonies headed by multiple reproductives.     

Endocrine control of cuticular hydrocarbon profiles during worker-to-soldier differentiation in the termite Reticulitermes flavipes

The social organization of termites, unlike that of other social insects, is characterized by a highly plastic caste system. With the exception of the alates, all other individuals in a colony remain at an immature stage of development. Workers in particular remain developmentally flexible; they can switch castes to become soldiers or neotenics. Juvenile hormone (JH) is known to play a key role in turning workers into soldiers. In this study, we analyzed differences in cuticular hydrocarbon (CHC) profiles among castes, paying particular attention to the transition of workers to soldiers, in the subterranean termite species Reticulitermes flavipes. CHCs have a fundamental function in social insects as they serve as cues in inter- and intraspecific recognition. We showed that (1) the CHC profiles of the different castes (workers, soldiers, nymphs and neotenics) are different and (2) when workers were experimentally exposed to a JH analog and thus induced to become soldiers, their CHC profiles were modified before and after the worker-presoldier molt and before and after the presoldier-soldier molt.     

Social biology and sex ratios of the eusocial gall-inducing thrips Kladothrips hamiltoni

1. Thrips comprise the only order besides Hymenoptera where females are diploid and males are haploid. This makes them useful insects for studying the roles of kin selection and ecology in social evolution. 2. Kladothrips hamiltoni is one of six species of Australian gall-inducing thrips that have been identified as eusocial. Galls are initiated by a single foundress, who rears her brood and remains within the enclosed gall for life. The adults of both sexes of her first brood cohort are morphologically distinct from the second generation, comprising a nondispersing soldier caste. The foundress and sib-mated soldiers jointly produce a second, dispersing generation, approximately 60–80% of which are produced by the soldiers. Mean per capita egg production of female soldiers is less than 33% that of the foundress. 3. Adult eclosion of soldiers is protandrous but the overall sex ratio of the soldiers lacks bias (52% male). Protandry of soldiers increases the probability that female soldiers will be inseminated soon after their eclosion and therefore lay fertilised, female eggs. The lack of bias could be due to a balance between local resource competition and local mate competition. Gender-specific defensive behaviour of soldiers with their enemies may also be important in explaining this unexpected sex ratio. 4. The dispersing generation has an overall extreme female bias (5.6% male). Soldier incest increases relatedness between females more than between males, such that the foundress is more related to her granddaughters than her daughters, and female soldiers are more related to their daughters than their sons (assuming within-gall relatedness < 1). A female bias in the offspring of soldiers should be preferred by both the foundress and soldiers as they are more related to soldier-produced dispersing females than any other thrips in the gall. Female bias in the dispersing generation will also reduce local mate competition between males. Both soldier incest and local mate competition may therefore contribute to the extreme female bias in the dispersing generation. 5. Selection pressures for sociality in gall-inducing thrips appear to be more similar to those in gall-inducing aphids and naked mole rats than to those in Hymenoptera.     

Toxicity, repellency, and horizontal transmission of fipronil in the formosan subterranean termite (Isoptera: Rhinotermitidae)

Toxicity of fipronil was evaluated against field-collected Coptoteres formosanus Shiraki. In topical application assays, fipronil was highly effective against both workers and soldiers at very low doses. Acute toxicity after 24 h was significantly greater in workers than in soldiers. The LD50s were 2.59- and 2.91-fold greater with soldiers than with workers from the two tested colonies. The LD50s of fipronil at 72 h after treatment were <2.0 ng/insect, with no significant differences regarding the tested workers/soldiers or colonies. Treated soldiers placed with untreated workers significantly increased worker mortality. However, there was no significant horizontal transmission of fipronil from treated workers to untreated soldiers. Fipronil at rates of 0.063% or less showed no repellency, whereas sand treatments of 0.125% fipronil were repellent to termite workers.     

Studies on cold pressor response of subjects who suffered from frostbite at high altitude

Seven lowlander soldiers who had 1st to 3rd degree of frostbite and 8 lowlander soldiers who recovered from high altitude pulmonary oedema were tested for cold pressor response at 260 m, 26°C, 6.8°C and 4,085 m, 26°C, 6.8°C. 8 lowlander soldiers (Control subjects) of comparable age group were tested for their cold pressor response at 260 m, 26°C and 3,300 m, 26°C. Another group of control subjects from the laboratory were also tested at 2,121 m, 26°C. There was a highly significant decrease in cold pressor response of the frostbite subjects at 260 m, 26°C and a very significant increase at 260 m, 6.8°C as compared to non-frostbite subjects. The subjects who recovered from high altitude pulmonary oedema did not show significant differences as compared with the control subjects. The results suggest certain basic physiological differences in regulation of supply of blood to the extremities under condition of general and local cold exposure.     

Panic escape polyethism in worker and soldier Coptotermes formosanus (Isoptera: Rhinotermitidae)

Termites were the first animal to form societies. Two hundred million years of evolution provide for a multitude of innate social behaviors that can be experimentally dissected. These fine‐tuned patterns of behavior are especially interesting when observing group decision making in the panic mode. In this study, we examined behavioral patterns of termites under panic conditions to gain insight into how an escape flow self‐organizes. One hundred worker and 10 soldier Coptotermes formosanus were released into agar plates. After a disturbance was created most workers followed each other and ran along the wall of dishes, thus forming a unidirectional escape flow, whereas soldiers showed a significantly higher frequency of moving to the center of the arena or on periphery of the escape flow as compared to workers. Agonistic behavior was usually observed as soldiers moved to center or periphery. This is the first report on the behavioral repertoire of termites when panicked, with details on the behavioral polymorphism of workers and soldiers during an escape.     

The impact of kleptoparasitic invasions on the evolution of gall-size in social and solitary Australian Acacia thrips

Many species of gall-inducing Acacia thrips are attacked by kleptoparasitic thrips who enter the gall, destroy the occupants, and then use the gall for producing their own offspring. The hypothesis tested here is that pressure exerted by ldeptoparasites （genus Koptothrips） not only provoked the evolution of soldiers in the gall-inducing clade, but have also influenced the evolution of gall size and morphology. Various size dimensions of invaded galls were compared to those of uninvaded galls using data from six gall-inducing species and their kleptoparasites. For the non-social gall-inducing species （K. ellobus and K. nicholsoni） invaded galls showed no significant size differences from galls that had not been invaded. For the four social gall-inducingspecies （K. habrus, K. intermedius, K. waterhousei and K. morrisi） invaded galls were significantly narrower and/or shorter than uninvaded galls. Galls of social species that had not been invaded and contained adult soldiers were significantly larger than galls where soldiers were still at a larval stage, suggesting that gall size is related to gall age in these species. An hypothesis is proposed that links the timing of invasion by kleptoparasites to size of the host gall： induction of a smaller gall by host founders will reduce the period of vulnerability to invasion （before soldiers become adults） for social thrips by allowing foundresses in these smaller galls to begin laying soldierdestined eggs relatively sooner.     

Current status of vivax malaria among civilians in Korea

A result of national malaria surveillance in Korean civilians was described. Since a case of indigenous vivax malaria was detected in 1993, a total of 2,198 cases was confirmed by blood smear up to 1997. Of them, 1,548 cases were soldiers serving in the demilitarized zone (DMZ), while 650 cases were civilians. Number of civilian cases was 3 in 1994, 19 in 1995, 71 in 1996, and 557 in 1997. Of them, 239 were ex-soldiers who discharged after military service in the prevalent areas such as Paju, Yonchon, Kimpo, Kangwha, Tongduchon in Kyonggi-do and Chorwon in Kangwon-do while 308 patients were civilian residents in the prevalent areas. Seventy-two patients, living nationwide, had a history of visiting the prevalent areas during transmission season. Only 32 civilian patients denied any relation with the prevalent areas. As a whole, a half of the civilian cases was diagnosed when living in non-prevalent areas. Male patients in their twenties was the highest in number. Annual parasite index is steadily elevated in residents living in the prevalent areas. Monthly incidence showed an unimodal distribution, forming a peak in August. Ex-soldiers exhibited a delayed incubation ranging from 153 to 452 days (279 ± 41 days). The time required for diagnosis was shortened from 23.6 days in 1995 to 13.7 days in 1997. Although the current epidemic of vivax malaria started as a border malaria, it seems highly probable that vivax malaria is established in the local areas and responsible for at least a part of transmission.     

Human ecology and parasitic infections. 1. The effect of occupation on the prevalence of parasitic infections in Calabar, Nigeria.

One thousand six hundred people belonging to three different occupational groups were randomly selected. Blood, urine and stool specimens were collected from them and processed for the detection of any parasitic infections. The results show 28.5% infection rate with one or more species of intestinal parasites; 50.7% in the school children, 17.3% among the farmers, and 12% among the soldiers. Results of blood examination showed 5.3%, 8.0% and 2.0% infection rates for P. falciparum in the school pupils, farmers and soldiers respectively, while 31.3% of the three groups were infected with the microfilaria of Dipetalonema perstans, and Loa loa. These were found in 60.7% of the farmers and none at all in the other two groups. Urine examination yielded no positive cases of urinary schistosomiasis and only one case of Trichomonas vaginalis. These results reflect the endemicity of the respective parasites, the degree of their sanitary awareness and their exposure frequencies to the pathogens as a result of their daily activities.     

Head-plug defense in a gall aphid

The aphid Astegopteryx sp. forms a banana-bunch shaped gall consisting of several subgalls on Styrax benzoides in northern Thailand, and completes its life cycle on the tree, without migrating to secondary hostplants. We found that its soldiers had sclerotic, protruded heads with many spine-like setae, and that several soldiers cooperate to plug the ostiole of the subgall with these heads. Of 173 ostioles examined in the field, 90.8 % were plugged with no space among the guarding soldiers. Many eggs and sexuals were found within subgalls guarded by soldiers, and a number of males were found trying to intrude into these subgalls. However, they were blocked by guarding soldiers, and it was no easy task for them to intrude into subgalls. The same was true for some soldiers that had rushed out of the subgall. Guarding soldiers often prevented outside soldiers from coming back into the subgall. These findings suggest an interesting possibility that guarding soldiers might consequently select still active, reusable soldiers and strong males for sexual females in their subgall. Received 6 March 2005; revised 5 and 20 July 2005; accepted 27 July 2005.