黑胸散白蚁下唇腺的解剖和扫描电镜观察

【目的】通过研究黑胸散白蚁Reticulitermes chinensis下唇腺解剖构造及其在不同品级个体间的分化,为进一步探索口交哺和品级分化机制提供参考。【方法】通过显微解剖观察,了解黑胸散白蚁下唇腺的形态构造及其在不同品级个体间的分化;通过扫描电镜观察,了解下唇腺的结构及神经支配;通过饮水实验,研究工蚁饮水及下唇腺囊的贮水功能。【结果】黑胸散白蚁下唇腺是左右对称结构,每侧构造由一个下唇腺腺泡群、一个下唇腺囊及相关的导管组成。每侧导管分别开口于舌基部下方。蚁后的下唇腺最发达,在下唇腺大小、下唇腺囊大小、腺泡大小及腺泡数量4方面均显著高于其他品级个体;兵蚁、工蚁、有翅成虫的下唇腺也较为发达,相互之间这4个参数的差异性不显著。扫描电镜观察发现,工蚁下唇腺腺泡由主导管和分支导管相连,腺泡外侧有神经分布。黑胸散白蚁工蚁有饮水习性,获得的水贮存在下唇腺囊内。【结论】黑胸散白蚁不同品级个体间,蚁后的下唇腺最发达,有翅成虫的下唇腺也比较发达,说明蚁后除行使生殖职能外,还承担交哺育幼等职能。黑胸散白蚁工蚁有饮水习性,下唇腺囊有贮水功能。     

Caste-specific expression of genetic variation in the size of antibiotic-producing glands of leaf-cutting ants

Social insect castes represent some of the most spectacular examples of phenotypic plasticity, with each caste being associated with different environmental conditions during their life. Here we examine the level of genetic variation in different castes of two polyandrous species of Acromyrmex leaf-cutting ant for the antibiotic-producing metapleural gland, which has a major role in defence against parasites. Gland size increases allometrically. The small workers that play the main role in disease defence have relatively large glands compared with larger workers, while the glands of gynes are substantially larger than those of any workers, for their body size. The gland size of large workers varies significantly between patrilines in both Acromyrmex echinatior and Acromyrmex octospinosus. We also examined small workers and gynes in A. echinatior, again finding genetic variation in gland size in these castes. There were significant positive relationships between the gland sizes of patrilines in the different castes, indicating that the genetic mechanism underpinning the patriline variation has remained similar across phenotypes. The level of expressed genetic variation decreased from small workers to large workers to gynes. This is consistent with the hypothesis that there is individual selection on disease defence in founding queens and colony-level selection on disease defence in the worker castes.     

Chemical and behavioural studies of the trail-following pheromone in the leaf-cutting ant Atta opaciceps,Borgmeier (Hymenoptera: Formicidae)

To understand the significance of the trail pheromone used in chemical communication of the leaf-cutting ants Atta opaciceps we investigated, under laboratory conditions, the trail-following behaviour of different castes. We observed a clear behavioural discrimination of conspecific venom gland extract of foraging ants from those of other species. Additionally, we determined the pheromone composition of A. opaciceps venom gland secretion using a two-dimensional gas chromatography coupled with mass spectrometry. Chemical analyses revealed the presence of three nitrogen-containing compounds, identified as 2,5-dimethylpyrazine, 3-ethyl-2,5-dimethylpyrazine and methyl 4-methylpyrrole-2-carboxylate (M4MPC). Four different bioassays performed with workers from different castes of A. opaciceps suggested that the trail pheromone elicits the trail-following behaviour in conspecifics of all castes, but the foragers respond more strongly to their own pheromone than to that of other castes (gardeners, generalists and soldiers). In addition, A. opaciceps foragers follow the trails made with the venom gland extracts of the unrelated Acromyrmex subterraneus subterraneus foragers as well as they follow the trails made with their own venom gland extract. M4MPC was identified to be the most abundant and the most behaviourally active component of the venom gland extract of A. opaciceps foragers.     

Comparative morphology of cephalic exocrine glands among castes of the black ant Lasius niger

The glandular system is crucially involved in main aspects of ant social life. The function of glands has been primarily studied in the workers (the non-reproductive individuals in a colony). In contrast, little information is available on queens (the reproductive females in a colony) or males in spite of the obvious functional differences between these castes. Here we report a comparison of the general morphology of the mandibular, propharyngeal and postpharyngeal glands between the three castes of the black ant Lasius niger. The analysis clearly shows that all these cephalic glands differ in relative size between castes and suggests a link between gland structure and its behavioral role in queens, workers and males. In particular, males present a hypertrophied mandibular gland. This is consistent with the fact that these glands might be the source of the sex pheromone in this caste. By contrast, queens exhibited the most developed postpharyngeal glands. This is consistent with the production of particular cues by queens for workers to help them to distinguish between reproductive and non-reproductive females. Finally, the propharyngeal glands were most developed in the worker caste and of similar relative size in males and queens. Their function is still enigmatic.     

The leg exocrine system inSerritermes serrifer (Hagen, 1858), phylogenetic implications (Isoptera: Serritermitidae)

Summary The leg exocrine gland was examined in two species of Neotropical termites. Scanning microscopy studies showed a set of pores on the ventral surface of the first and second tarsomeres in all legs ofSerritermes serrifer. InHeterotermes tenuis these pores are present on a sunken plate in all castes. To date, this gland has been observed only in Rhinotermitid species. The presence of leg exocrine gland provides additional evidence supporting a close phylogenetic relationship between the Serritermitidae and Rhinotermitidae.     

Physiology as a caste-defining feature

Division of labour is a key factor in the ecological success of social insects. Groups of individuals specializing on a particular behaviour are known as castes and are usually distinguished by morphology or age. Physiology plays a key role in both these types of caste, in either the developmental physiology which determines morphology, or the temporal changes in physiology over an insect’s life. Physiological correlates of morphological or temporal caste include differences in gland structure, secretory products, leanness, neuroanatomy and neurochemistry. However, purely physiological castes could also occur. Physiological castes are discrete groups of same-age same-size individuals with particular physiological competencies, or groups of individuals with similar physiology crossing age or size groups. A stable physiological caste occurs in the monomorphic Pharaoh’s ant, where some ants can detect old pheromone trails and retain this specialization over time. These ants differ physiologically from other workers, and the differences arise before eclosion. More temporary physiological castes occur in the ant Ectatomma where brood care specialists have more developed ovarioles than other same-aged workers, and in the honeybee where nurses, wax-workers and soldiers all differ physiologically from same-aged nestmates. Physiology is an important aspect of caste, not only in its contribution to age-related and morphological castes, but also in its own right as a caste grouping factor. While age and morphological differences make caste structures accessible for study, more cryptic physiological castes may play just as important a role in division of labour. Received 19 December 2007; revised 24 July and 18 September 2008; accepted 19 September 2008.     

Survey of the exocrine system in Protanilla wallacei (Hymenoptera,Formicidae)

We studied the exocrine system of both workers and ergatoid queens of Protanilla wallacei using light, scanning and transmission electron microscopy. Our survey revealed the presence of 26 glands, of which 6 had never been found before in ants. Five of these represent novel discoveries for social insects in general. The overall novel discoveries comprise an epithelial stipes gland, a pharyngeal wall gland, a central petiole gland, a lateral postpetiole gland and a foot-sole gland in the hindleg pretarsi. The intramandibular epithelial gland was already reported in some bees previously, but is now for the first time also reported in ants. The exocrine system of workers and ergatoid queens is very similar, with only the spermathecal gland showing an obvious difference. This is in line with the limited anatomical as well as behavioural difference between both castes in Protanilla compared to the situation in Leptanilla.     

Identification of major royal jelly proteins in the brain of the honeybee Apis mellifera

The consumption of royal jelly (RJ) determines the differences between castes and behavioral development in the honeybee Apis mellifera. However, it is not known whether the proteins of RJ are related to these differences, or which proteins are responsible for the changes. To understand the functions of RJ proteins that are present in other tissues of the bee, in addition to hypopharyngeal gland, we used a polyclonal antibody anti-MRJP1 to investigate the presence of this protein in nervous system of honeybee. This study showed the presence of three polypeptides (p57, p70 and p128) in specific tissues of bee brain. Mushroom body, optic lobe and antennal lobe neuropils all contained proteins recognized by anti-MRJP1. Proteomic analysis showed that the three polypeptides are correlated with proteins of the MRJP family. p57 is correlated with MRJP1, p70 with MRJP3, while p128 may be an oligomeric form or a new polypeptide. Immunostaining of the brain and hypopharyngeal gland revealed differential expression of MRJPs in various brain regions and in different honeybee castes and subcastes. The identification and localization of these MRJPs contribute to the elucidation of the biological roles of this protein family.     

Exocrine glands in the legs of the social wasp Vespula vulgaris

This study brings a survey of the exocrine glands in the legs of Vespula vulgaris wasps. We studied workers, males, virgin queens as well as mated queens. A variety of 17 glands is found in the different leg segments. Among these, five glands are novel exocrine structures for social insects (trochanter-femur gland, ventrodistal tibial gland, distal tibial sac gland, ventral tibial gland, and ventral tarsomere gland). Most leg glands are present in the three leg pairs of all castes. This may indicate a mechanical function. This is likely for the numerous glands that occur near the articulation between the various leg segments, where lubricant production may be expected. Other possible functions include antenna cleaning, acting as a hydraulic system, or pheromonal. Further research including leg-related behavioural observations and chemical analyses may help to clarify the functions of these glandular structures in the legs.     

Differences in cellulose digestive systems among castes in two termite lineages

Abstract.  Termites (Isoptera) are eusocial insects and express polyphenism. Soldiers have specialized morphology for colony defense, but their feeding activity is dependent on other colony members. To determine differences in cellulose degradation between soldier and worker termites, enzymatic activity and cellulase gene expression, as well digestive tract histology, are examined in two phylogenetically distant species. In Hodotermopsis sjostesti (family Termopsidae) , endo-β-1,4-glucanase activity is identified in the salivary glands, whereas β-glucosidase activity is identified in salivary glands and hindgut. The relative expression levels of endo-β-1,4-glucanase genes in soldiers are significantly lower than in workers. Thin sections of salivary gland of workers and soldiers are different in H. sjostedti . In Nasutitermes takasagoensis (family Termitidae), the endo-β-1,4-glucanase activity is restricted to the midgut in four tested castes (i.e. three types of workers and soldier). Examination of activity per termite reveals the highest activity in minor workers and the lowest activity in major workers and soldiers. The β-glucosidase activity is also concentrated on the midgut in all four castes. The relative expression level of the endo-β-1,4-glucanase gene does not correspond with its activity in the midgut. In thin sections prepared from N. takasagoensis , the folds and pulvillus in the gizzards, and cuticle structure of soldiers are less developed compared with the other three worker castes. The differences in digestive system among termite castes in terms of caste development in each species are discussed.     

Armed reproductives: Evolution of the frontal gland in imagoes of Termitidae

The frontal gland of termites is a structure without any equivalent among other animals. Although this gland is well known in soldiers, it received almost no attention in other castes. Recently, we described it in imagoes of Rhinotermitidae and Serritermitidae. In order to provide a complete picture of the evolution of this gland in termite imagoes, we studied it in additional 34 species of Termitidae, representing 7 of the 8 subfamilies. The frontal gland of these species is formed by class 1 secretory cells only, and occurs in two basic shapes: epithelial with reservoir in Foraminitermitinae and Macrotermitinae, and epithelial without reservoir in all other subfamilies. The size variability of the gland is high, not only among Termitidae subfamilies, but also within subfamilies. Our data suggest that the ancestral form of the frontal gland is epithelial with reservoir, as found in Rhinotermitidae, Serritermitidae, and basal Termitidae. The reduction of the reservoir occurred at least two times and the gland was lost two times independently: in Protermes sp. and in Microtermes toumodiensis (both Macrotermitinae).     

蜜蜂上颚腺及其分泌物研究进展

上颚腺是蜜蜂重要的外分泌腺体,其分泌物是维系蜂群社会性结构的重要物质。蜂王和工蜂上颚腺分泌物合成均以硬脂酸为合成前体,但在脂肪酸的β-氧化过程中表现出级型差异性,导致分泌物组分比例不同。蜂王上颚腺分泌物以9-羰基-2癸烯酸(9-ODA)为主,有吸引工蜂和雄蜂、抑制工蜂卵巢发育等作用;工蜂上颚腺分泌物以10-羟基-2癸烯酸(10-HDA)和10-羟基癸酸(10-HDAA)为主,是蜂王浆的重要组成部分。同时,这种具备典型级型差异的分泌物组成又具有级型间可塑性,在不同蜂种间也存在区别。近年来在转录水平和蛋白水平的一些研究进一步揭示了级型间差异的分子基础。针对蜜蜂上颚腺及其分泌物的研究在蜜蜂生物学、行为学和蜂产品质量控制等方面具有重要的意义。本文通过总结国内外相关研究进展,旨在为上颚腺分泌物的作用机制、生物合成机制等领域的进一步深入研究提供借鉴。     

Caste and colony variations in the chemical composition of the cephalic secretions of the African weaver ant, Oecophylla longinoda

ABSTRACT. Considerable differences were found in the composition of the volatile cephalic chemical of the castes of Oecophylla longinoda. The mandibular glands of minor workers contain a series of primary alcohols, predominantly 1-hexanol, 1-octanol and 1-nonanol, together with nerol and geraniol. Hexanal, 2-butyl-2-octenal and 3-undecanone (components of the mandibular gland secretion of major workers, and which are important stimuli in alarm communication) are absent. Major workers are attracted and arrested by high concentrations of nerol, which is part of a possible mechanism for communication within the nest. The heads of males contain a pungent secretion, which may be used in defence; the major constituents are five aliphatic acids. The quantities of both main and trace constituents of the mandibular gland secretion of major workers were found to vary between colonies collected in different areas of West Africa.     

Comparative Study of the Labial Gland Secretion in Termites (Isoptera)

Labial glands are present in all castes and developmental stages of all termite species. In workers, their secretion contains a food-marking pheromone and digestive enzymes, while soldier secretion plays a defensive role. However, these functions were studied only in a limited set of species, and do not allow drawing general conclusions. Hence, we have investigated the chemical composition of the labial gland extracts from soldiers and workers in 15 termite species belonging to 6 families using an integrative approach based on proteomic and small-molecule profiling. We confirmed the presence of hydroquinone and cellulase in the labial glands of workers, and we identified new toxic compounds in soldiers and workers of several species. Our results highlight the dual role of labial gland secretion, i.e. the defensive role in soldiers and workers of several termite species, and the digestive function in workers.     

Morpho-functional comparison of the Dufour gland in the female castes of the Amazon ant <Emphasis Type="Italic">Polyergus rufescens</Emphasis> (Hymenoptera,Formicidae)

The Dufour gland is crucially involved in main aspects of the parasite habit of the slave-making ant Polyergus rufescens, i.e. slave-raids and host colony usurpation. Workers use chemicals from this gland as recruitment signals during raid organization, while newly-mated queens use its secretion to appease residents during host nest invasion. Here, we report a comparison of the general morphology and fine structure of the Dufour gland in the female castes of P. rufescens: queens, ergatogynes (intermediate forms), and workers. The analysis clearly shows the link between gland structure and its behavioural role in queens and workers. In particular, queens present a hypertrophied gland with an extended lumen and a thin epithelium no more active in secretory function. This is consistent with the fact that usurper queens use the Dufour gland contents only during the short phase of host nest penetration. Contrary to adult queens, the cytoplasmic organization of the Dufour gland epithelium of raiders is typical for a tissue with secretory activity (abundance of mitochondria, free ribosomes, strands of smooth endoplasmic reticulum and a Golgi apparatus). This is consistent with the continuous raiding activity performed by workers throughout their adult life. The biology of ergatogynes is still an enigmatic matter. Their Dufour gland is intermediate in shape and size between that of queens and workers. It presents a fairly thick epithelium with features that are typical of a quite active secretory tissue.     

Not only soldiers have weapons: evolution of the frontal gland in imagoes of the termite families Rhinotermitidae and Serritermitidae

Background

The frontal gland is a unique adaptation of advanced termite families. It has been intensively studied in soldiers with respect to its anatomy and chemistry, with numerous novel compounds being discovered within the tremendous richness of identified products. At the same time, the presence of the frontal gland in non-soldier castes received only negligible attention in the past.

Principal Findings

Here, we report on the development of the frontal gland in alate imagoes of 10 genera and 13 species of Rhinotermitidae and Serritermitidae, in order to shed light on the evolution and function of this gland in imagoes. All investigated species possess a frontal gland. In most cases, it is well-developed and equipped with a sac-like reservoir, located in the postero-dorsal part of cranium, but reaching as far as the seventh abdominal segment in some Rhinotermitinae. The only exception is the genus Psammotermes, in which the gland is very small and devoid of the reservoir.

Conclusions

Our direct observations and comparisons with soldiers suggest a defensive role of the gland in imagoes of all studied species. This functional analogy, along with the anatomic homology between the frontal gland in soldiers and imagoes, make it likely that the gland appeared once during the early evolution of rhinotermitid ancestors, and remained as a defensive organ of prime importance in both, soldiers and imagoes.     

白蚁信息素研究进展

白蚁是最古老的社会性昆虫, 其社会性的维持需要信息素的相互作用。本文回顾了近年来国内外白蚁信息素研究的最新进展, 内容涉及白蚁踪迹信息素、 性信息素、 告警信息素和促食信息素的功能、 化学成分及产生信息素的外分泌腺。白蚁分泌信息素的腺体主要有背板腺、 腹板腺、 后腹板腺、 额腺和唾腺。绝大多数白蚁信息素是挥发性物质。白蚁在化学通讯上存在节俭策略, 即同一种化合物由不同的白蚁种类的不同外分泌腺分泌, 可具有不同的功能。总结了各类信息素在白蚁物种间、 同一物种的品级间和性别间的异同和作用方式, 强调了白蚁信息素的反应阈值、 最佳浓度、 有效期和物种特异性对其功能的影响。目前对白蚁信息素的研究尚处于起步阶段, 其研究成果对等翅目系统发育研究和白蚁防治具有重要的意义。文章最后展望了白蚁信息素在白蚁防治上的应用前景。     

Effects of consanguineous marriages on fertility among three endogamous groups of Andhra Pradesh

Abstract

For the present study three castes representing three socioeconomic levels have been selected from Andhra Pradesh, South India. Members of the wealthier castes married consanguineously more frequently than did the members of the lower‐ranking castes. The consanguineous unions among all the castes are found to be more fertile than are nonconsanguineous unions. Indeed, fertility increases with degree of relationship of the spouses. Number of pregnancies and live births are significantly (P = 0.01) higher among consanguineous unions (among all the castes). However, the difference in the number of surviving offspring between consanguineous and nonconsanguineous unions is not significant among the wealthier castes. This suggests that child mortality is higher among the offspring of consanguineous unions in spite of their greater wealth.     

La glande mandibulaire du Termite a cou jaune (Calotermes flavicollis)

Résumé La glande mandibulaire deCalotermes flavicollis est constituée de deux types distincts de cellules: des grandes cellules sécrétrices dont chacune déverse son produit de sécrétion à l'extérieur par l'intermédiaire d'un canalicule, et des petites cellules centrales. Au moment de la mue, les canalicules sont rejetés, et les petites cellules élaborent les nouveaux canalicules effectuant une migration centrifuge, puis dégénèrent.Dès l'éclosion, la glande mandibulaire est différenciée. Au cours du développement post-embryonnaire, elle augmente de taille, sans que son anatomie varie. Entre les différentes castes, les variations sont peu co nsidérables, mais le développement de la glande est maximum chez les sexués fonctionnels, imaginaux et néoténiques.

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Summary The mandibular gland ofCalotermes flavicollis consists of two distinct types of cells: large secretory cells each of which pours its secretion to the exterior by means of a small duct, and small central cells. At moulting time, the ducts are discarded and the small cells form fresh ducts and migrate to the periphery, after which they degenerate.The mandibular gland is differenciated at the time of hatching. During post-embryonic development it increases in size without any change in anatomy. The differences between the castes are, of little significance, though the development of the gland is at its maximum among functional reproductives, imagoes or neoteinics.

     

Electron microscopic study of the mandibular glands of Kalotermes flavicollis Fabr. (Isoptera; Calotermitidae)

Summary The mandibular glands of Kalotermes were examined in different castes. They showed sexual dimorphism in the soldiers and primary reproductives. Moreover, in female soldiers and queens, mandibular gland cells contained numerous crystalline structures of mitochondrial origin. The role of these glands (secretion of saliva or pheromone) is discussed.

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Résumé Les glandes mandibulaires de Calotermes sont étudiées dans les différentes castes. Elles présentent un dimorphisme sexuel chez les soldats et les sexués. Après la mue imaginale, les cellules glandulaires ont toujours une activité sécrétrice. De plus, chez les soldats femelles et les reines, elles contiennent de nombreuses formations cristallines d'origine mitochondriale. Le rôle de ces glandes (sécrétion de salive ou de phéromone) est discuté.