Functional morphology of the postpharyngeal gland of queens and workers of the ant Monomorium pharaonis (L.)

Eelen D., Børgesen L.W. and Billen J. 2006. Functional morphology of the postpharyngeal gland of queens and workers of the ant Monomorium pharaonis (L.). —Acta Zoologica (Stockholm) 87 : 101–111 The postpharyngeal gland (PPG) is unique to ants and is the largest exocrine gland in their head. In queens of the pharaoh's ant, Monomorium pharaonis, the gland contains approximately 15 finger‐like epithelial extensions on each side and opens dorsolaterally in the posterior pharynx. In these ants the PPG morphology varies considerably according to age and mating status. The epithelial thickness increases with age and reaches a maximum at 3 weeks in both virgin and mated queens. A considerable expansion of the lumen diameter occurs in both groups between 4 and 7 days. Virgin queens release their secretion into the gland lumen from an age of 7 days, whereas mated queens accumulate large amounts of secretion in their epithelium. The increasing epithelial thickness, together with the increasing lumen diameter, the presence of numerous inclusions in the epithelium and the release of secretion, are indicative for increasing gland activity. The gland ultrastructure indicates involvement in lipid metabolism and de novo synthesis of lipids. The PPG of workers consists of 12 finger‐like tubes at each side. There is a significant difference in epithelial thickness between nurses and repletes and between nurses and foragers. We suggest the PPG serves different purposes in pharaoh's ants: it is likely that the PPG of workers and virgin queens is used to feed larvae. In mated queens the gland probably plays a role in providing the queen with nutritious oils for egg production. The PPG may also function in signalling species nestmate and caste identity, as well as in the reproductive capacity of the queens.     

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.     

Queen number in a supercolony of the invasive garden ant, <Emphasis Type="Italic">Lasius neglectus</Emphasis>

Summary We have analysed the distribution of queens under stones at the core and at the periphery of a supercolony of Lasius neglectus that occupies 14 ha at Seva (NE Spain). Queens were not found alone, but rather within worker groups. Density at the center (mean ± s.d.: 1.38 ± 2.87 queens/stone; n = 100 stones; range 0–14) was not different from density at the periphery (1.18 ± 2.38; range 0–12). The estimate of the number of queens found under stones for the whole colony is about 35500. Egg-laying rates for queens from these two zones were obtained in the laboratory, at three different temperatures, and there were no differences detected. The presence of brood stages, from eggs to cocoons, was also similar in both zones. The homogeneous distribution of colony components may indicate that the area occupied by L. neglectus has already reached saturation. With a different technique – soil core extraction – we could estimate the density of workers in the soil: 800 workers per m². Soil cores had 6.28 ± 20.0 workers/core (range: 0–173), giving a rough estimate of 1.12 × 10⁸ workers in the soil, for the entire colony. Though few, some queens were also recovered from soil cores. Queen numbers for the supercolony, based on queens found in the soil, reaches the astounding level of 360000. Numbers are consistent with previous predictions.Received 10 September 2003; revised 1 December 2003; accepted 9 December 2003.     

Exocrine glands and the attractiveness of the ergatoid queen in the ponerine antMegaponera foetens

Summary Queens inMegaponera foetens are permanently wingless. In a colony from Kenya we observed that the ergatoid queen was often surrounded by a large number of workers, all conspicuously facing her. To investigate the nature of this attraction, we divided the colony in half. When the queen was transferred from one group to the other, a large court of workers formed immediately. When marked workers were transferred from the queenright group to the other, they attracted much attention and were licked repeatedly. These data suggest that the queen produces a chemical signal which attracts workers, and this signal is transferred onto the workers. In a histological investigation of queen and workers, we found many intersegmental glands in both castes. However, only the queen had a thick glandular epithelium lining her entire body, and this may be the source of her signal. The numerous erect setae covering the queen (but not the workers) may help to dispense the secretions. Our results confirm that ergatoid queens in the Ponerinae are morphologically specialized, despite their external similarity with workers.     

Queen power in relation to age and mating status in the Argentine antIridomyrmex humilis (Mayr)

Summary Queen power inIridomyrmex humilis during six arbitrarily chosen physiological stages of queens (virgin winged queens at the time of emergence, the same 4–5 days old, mated winged queens 5–6 days old, mated queens at the time of dealation, young egg-laying queens, and old egg-laying queens) was tested with regard to workers. Tested workers and queens were nestmates. The results were as follows. 1) Power of the old queens remained rather constant throughout the reproductive season. 2) Young queens were always less attractive than older ones. No changes were observed from the time of emergence to dealation. 3) At the time of egg-laying, these young queens became markedly attractive for workers but never as much as old egg-laying queens. Therefore, insemination and dealation do not trigger the increase in queen power. In contrast, oogenesis ending by egg-laying produces a significant increase in queen power.     

The fine structure of the planarianPolycelis tenuis ijima

Summary The fine structure of the pharynx is presented and demonstrates that the pharyngeal epithelial system is a continuous one. The epithelial lining of the pharyngeal cavity with its characteristic fibrous secretory bodies merges with the outer pharyngeal epithelium at the point of anchorage of the pharynx. A few of these cells are insunk, the nuclei occurring beneath the underlying muscular layers. The nature of the outer epithelium changes towards the free end of the pharynx; the cells become ciliated and in contents come to resemble the inner epithelium which it joins at the tip.The gut cells merge at a transitional zone with the inner pharyngeal epithelium and at this point both bear microvilli and contain rod-shaped apical bodies. Some of these cells are also insunk. Towards the mouth the epithelium shows a greater degree of insinking and exhibits microapocrine secretion. Both inner and outer epithelia bear sense receptors which are concentrated at the lip.At the point of pharyngeal insertion, the sub-epithelial tissue resembles planarian parenchyma, but is rich in gland cells. These glands open on to the outer epithelium especially towards the free end of the pharynx.This research was supported by the Scientific Research Council. Grant No. B/RG/086.     

The gas bladder of Argentina silus L., with special reference to the ultrastructure of the gas gland cells and the countercurrent vascular bundles

Summary The orifice between the two chambers of the gas bladder in Argentina silus is surrounded by a sphincter muscle. Gas analyses of the gas bladder contents of fish from 400 meters depth give 0–1% carbon dioxide and 9–72% oxygen. Micro-retia mirabilia form a countercurrent vascular system, and the arterial component has peripherally a sphincter mechanism. The function of the glandular layer of the anterior chamber remains uncertain, but the structure indicates secretion into blood capillaries. The lining epithelium of the anterior chamber may secrete some substance into blood or directly into the lumen, which may be involved in a secretory mechanism. This conclusion is not supported by our histochemical tests. The posterior chamber has no micro-retia and the blood vessels have a different origin from those of the anterior chamber. The blood vessels form a plexus of capillaries or sinuses in contact with the flat lining epithelium, thus allowing gases to pass freely by diffusion. — The muscular layers of both chambers are innervated by catecholamine-containing nerve fibres.The investigation was supported by grants from the Swedish Natural Research Council (No. 99-35) and by the Faculty of Mathematics and Science, University of Lund.     

On the Fine Structure of the Alimentary Tract of Cephalodiscus gracilis (Pterobranchia,Hemichordata)

The U-shaped alimentary tract of Cephalodiscus is of exclusively epithelial structure; on the basis of fine structural criteria the entire tract can be divided into two large subdivisions: an anterior one with mouth, mouth cavity, pharynx and oesophagus, and a posterior one with stomach and intestine. The anterior subdivision is built up of a relatively uniform, innervated, pseudostratified, ciliated epithelium with mucus cells which are concentrated in the initial parts of the mouth cavity. Cilia and mucus presumably constitute a mechanism transporting food particles into the stomach. In the area of the gill slits specific vacuolated cells occur which may lend rigidity to the walls of the slits. The gastric epithelium consists of prismatic cells characterized by, among others, large inclusion bodies, which may represent digestive vacuoles, small dense rod-shaped granules and an elaborate system of microridges, at the base of which abundant endocytotic vesicles occur. The dorsal gastric pouch contains cells rich in rough ER and secretory granules, probably containing digestive enzymes. Thus morphological evidence points both to intra- and extracellular digestion. The intestinal epithelium resembles that of the stomach, however, it is lower, its organelles are fewer and it bears, beside cilia, mainly microridges, which towards its distal end become sparse. Both in the gastric and intestinal epithelium small granulated cells have been found, which presumably represent endocrine cells.     

Yolk protein in the pharaoh's ant: Influence of larvae and workers on vitellogenin and vitellin content in queens

Summary Vitellin from the pharaoh's ant,Monomorium pharaonis (L.), was found to contain a single apoprotein with aM _r of 198.3 kDa. The protein is a glycoprotein exposing mannose containing carbohydrate groups.Antibodies to pharaoh's ant vitellin (v _t), raised in rabbits, were used to determine the head-, thorax-, and abdominal contents of vitellogenin and vitellin (v _g+v_t) in queens in relation to the presence or absence of larvae and workers using an ELISA test. Thev _g+v_t contents were also compared to the egg-laying rate and the weight of the queens.The results confirm the existence of a positive correlation betweenv _g+v_t contents in the queens and their access to larvae, probably related to the queens' preferential feeding on larval secretions. In queens without larvae the abdominalv _g+v_t contents declined in concordance with a low oviposition rate of 6–8 eggs/day. In spite of cessation of egg-laying within 24 hours after removal of both larvae and workers, the queens maintained basal contents ofv _g+vt. This may indicate that the presence of larvae is not only essential for the nutrition of the queens, but also for the uptake of vitellogenin in the growing oocytes. This additional stimultive factor may be based on the queens' response to primer pheromones liberated by the larvae. v _g+v_t could not be demonstrated in workers or larvae with the ELISA test. If anyv _g+vt is present in larvae and workers the amount is lower than the detection limit (1–2 ng/individual). This seems to rule out the possibility of transfer of proteins of this kind to the queens.     

Olfactory metamorphosis in the coastal tailed frog Ascaphus truei (Amphibia,Anura, Leiopelmatidae)

The structure of the olfactory organ in larvae and adults of the basal anuran Ascaphus truei was examined using light micrography, electron micrography, and resin casts of the nasal cavity. The larval olfactory organ consists of nonsensory anterior and posterior nasal tubes connected to a large, main olfactory cavity containing olfactory epithelium; the vomeronasal organ is a ventrolateral diverticulum of this cavity. A small patch of olfactory epithelium (the “epithelial band”) also is present in the preoral buccal cavity, anterolateral to the choana. The main olfactory epithelium and epithelial band have both microvillar and ciliated receptor cells, and both microvillar and ciliated supporting cells. The epithelial band also contains secretory ciliated supporting cells. The vomeronasal epithelium contains only microvillar receptor cells. After metamorphosis, the adult olfactory organ is divided into the three typical anuran olfactory chambers: the principal, middle, and inferior cavities. The anterior part of the principal cavity contains a “larval type” epithelium that has both microvillar and ciliated receptor cells and both microvillar and ciliated supporting cells, whereas the posterior part is lined with an “adult‐type” epithelium that has only ciliated receptor cells and microvillar supporting cells. The middle cavity is nonsensory. The vomeronasal epithelium of the inferior cavity resembles that of larvae but is distinguished by a novel type of microvillar cell. The presence of two distinct types of olfactory epithelium in the principal cavity of adult A. truei is unique among previously described anuran olfactory organs. A comparative review suggests that the anterior olfactory epithelium is homologous with the “recessus olfactorius” of other anurans and with the accessory nasal cavity of pipids and functions to detect water‐borne odorants. J. Morphol. 2011. © 2011 Wiley Periodicals, Inc.     

Exocrine glands of the ant Myrmoteras iriodum

This paper describes the morphological characteristics of nine major exocrine glands in workers of the formicine ant Myrmoteras iriodum. The elongate mandibles reveal along their entire length a conspicuous intramandibular gland, which contains both class‐1 and class‐3 secretory cells. The secretory cells of the mandibular glands show a peculiar appearance, with a branched end apparatus, which is unusual for ants. The other major glands (pro‐ and postpharyngeal gland, infrabuccal cavity gland, labial gland, metapleural gland, venom gland and Dufour gland) show common features for formicine ants. The precise function of the glands could not yet be experimentally demonstrated, and to clarify this will depend on the availability of live material of these enigmatic ants in future.     

Queen execution increases relatedness among workers of the invasive Argentine ant,Linepithema humile

Polygyny in social insects can greatly reduce within‐nest genetic relatedness. In polygynous ant species, potential rival queens in colonies with multiple queens are often executed by other queens, workers, or both. The Argentine ant, Linepithema humile, native to South America, forms a “supercolony” that is composed of a large number of nests and is considered to contribute to the ant's invasion success. Currently, four mutually antagonistic supercolonies are contiguously distributed within a small area of Japan. Here, we analyzed the genetic structure and relatedness within and among the four supercolonies using microsatellite markers to clarify how L. humile maintains its supercoloniality. The results of AMOVA and BASP, the F_ST values, and the existence of several private alleles indicated that the L. humile population in the Kobe area had a characteristic genetic structure. Within a given supercolony, there was significant genetic differentiation (F_ST) among workers collected in May and those collected in September. The significant deviation from Hardy–Weinberg equilibrium increased, and the relatedness among workers significantly increased from May to September in all supercolonies. This result suggested that the supercolonies replaced old queens with new ones during the reproductive season, thus supporting the plausibility of queen execution. From the perspective of kin selection, workers collectively eliminate queens, thereby increasing their own inclusive fitness. Restricted gene flow among supercolonies, together with mating with sib and queen execution, could help to maintain the unique social structure of L. humile, the distribution of which is expanding worldwide.     

Ontogeny of the digestive tract during larval development of yellowtail flounder: a light microscopic and mucous histochemical study

The histological development and mucous histochemistry of the alimentary tract in larval yellowtail flounder were studied using light microscopy. Samples were taken when the larvae were first offered food at 3 days post-hatch, then at 7, 10, 29, 36, and 46 days post-hatch, at which time they were metamorphosing. Regional partitioning of the digestive tract into the buccal cavity, pharynx, oesophagus, post-oesophageal swelling (PES), intestine, and rectum was complete by day 10. Goblet cells were present only in the buccal cavity, pharynx and intestine by day 7, but increased in number and distribution as development continued. By day 29, the posterior zone of the oesophagus had a marked increase in goblet cell density and mucosal folding. At the transition from oesophagus to PES/stomach stratified epithelium with goblet cells changed abruptly to a columnar epithelium with no goblet cells. Multicellular glands in the PES of 36-day larvae allowed it to be defined as a stomach. The distinct brush border of columnar epithelium and the presence of goblet cells characterize the intestine and rectum. All goblet cells throughout the digestive tract were strongly positive for acid mucins as was the luminal layer of the stratified epithelia lining the buccal cavity, pharynx and oesophagus. The PES/stomach epithelium stained weakly for neutral mucins. No mucin staining was associated with the gastric glandular epithelium. The brush borders of the intestine and rectum were strongly positive for combinations of neutral and acid mucins.     

Ultrastructure of the anterior alimentary tract of a monogenean, Diclidophora merlangi

The anterior alimentary tract of Diclidophora merlangi is composed of a complex series of morphologically distinct epithelia interconnected by septate desmosomes and penetrated by the openings of numerous unicellular glands. The mouth and buccal cavity are lined by an infolding of modified body tegument, distinguished by uniciliate sense receptors, buccal gland openings, and in the buccal region by a dense, spiny appearance. The prepharynx is covered by an irregularly folded epithelium and, for part of its length, by the luminal cytoplasm of the prepharyngeal gland cells. The epithelium is syncytial and pleiomorphic, and regional variation in structure is common. A separate epithelium invests the lips of the pharynx and its free surface is greatly amplified by numerous, dense lamellae of varying dimensions. The lip epithelium is continuous with cytoplasmic processes of cells located external to the pharynx. A further, distinct epithelium borders the pharynx lumen and is composed of discrete cytoplasmic units connected by short septate desmosomes. The oesophagus is lined by a modified caecal epithelium, lacking haematin cells, and, in places, is perforated by the openings of oesophageal gland cells; it is continuous with the syncytial connecting tissue of the gut caeca.     

Kin structure and colony male reproduction in the hornet<Emphasis Type="Italic"> Vespa crabro</Emphasis> (Hymenoptera: Vespidae)

We estimated queen mating frequency, genetic relatedness among workers, and worker reproduction in Vespa crabro flavofasciata using microsatellite DNA markers. Of 20 colonies examined, 15 contained queens inseminated by a single male, 3 colonies contained queens inseminated by two males, and 2 colonies contained queens inseminated by three males. The genetic relatedness among workers was estimated to be 0.73±0.003 (mean±SE). For this high relatedness, kin selection theory predicts a potential conflict between queens and workers over male production. To verify whether males are derived from queens or workers, 260 males from 13 colonies were genotyped at four microsatellite loci. We found that all of the males were derived from the queens. This finding was further supported by the fact that only 33 of 2,990 workers dissected had developed ovaries. These workers belonged to 2 of the 20 colonies. There was no relationship between queen mating frequency and worker reproduction, and no workers produced male offspring in any of the colonies. These results suggest that male production dominated by queens in V. crabro flavofasciata is possibly due to worker policing.     

Behavioral ecology of Japanese paper wasps,Polistes spp. IV. Comparison of ethograms between queens and workers ofP. chinensis antennalis in the ergonomic stage

Behavioral catalogues, transition matrices between 2 consecutive behavioral acts, and the frequency distribution of time spent in each behavioral act were described for queens and workers of a Japanese paper wasp,Polistes chinensis antennalis in the ergonomic stage (the period from the emergence of the first worker to the emergenece of the first reproductive). The results were; (1) The whole behavior repertoire of queens was completely included in that of workers. Workers oviposited frequently in queen-right colonies. (2) The proportion of off-nest activities to the number of total acts for workers was higher than that for queens. The proportion of time spent off the nest in workers was 5 times as high as that in queens. (3) There was no significant difference in the proportion of number of oviposition to the total number of acts between queens and workers. The oviposition rate was 0.06 eggs/h for workers and 0.37 eggs/h for queens. Queens made more dominance behavior and workers more subordinated behavior. Queens made abdomen wagging more frequently than workers. (4) For behavioral acts common in both queens and workers, the mean duration of a specific act was approximately equal for both. The mean duration of unsuccessful flesh collection for workers was 3.4 times as longs as that for queens. Workers did not search for flesh in 10.1% of duration of flesh collections in the field. (5) The pattern of transitions between 2 consecutive behavioral acts in workers was similar to that in queens. (6) Alien workers approaching a nest with a load were more often accepted by the nest occupants than alien workers with no load. Alien workers with no load were not accepted as often as a legitimate nestmates with no load. The reproductive division of labor between queens and workers inP. chinesis antennalis was incomplete. This suggests worker-queen conflict in different reproductive strategies among colony members. Several behaviors were consistent with the view that they are the behavioral expression of this conflict.     

Colony genetic structure in the mono- and polygynous sibling species of the antsCamponotus nawai andCamponotus yamaokai: DNA fingerprint analysis

Using the ant-derived probe (pMY7), we performed DNA fingerprinting in monogynous and polygynous sibling ant speciesCamponotus nawai andCamponotus yamaokai. In monogynousC. nawai, band-sharing probabilities were low between unrelated individuals (mean 0.09), but those and relatedness estimates were consistently high between workers of the same nest (mean 0.85 and 0.74–0.83, respectively), suggesting that the queen mated once and nestmate workers are super-sisters. It also suggested monoandry: that is, that all nestmate workers shared most of the bands which were considered to have derived from a male. In polygynousC. yamaokai, band-sharing probabilities were low between queens of different populations (mean 0.13), moderate between queens of different nests in the same population (mean 0.25), but very high between queens of the same nest (within-nest means were 0.84–0.96). These results suggest that nestmate queens are genetically closely related with each other. Relatedness estimates between colony members sometimes reached 1. This might result from successive intranidal mating (inbreeding or large Wahlund effect) and adoption of new queens into the natal nests.     

Structure of the Digestive Tube in the Holothurian Eupentacta fraudatrix (Holothuroidea: Dendrochirota)

In the holothurian Eupentacta fraudatrix,the gut wall exhibits trilaminar organization. It consists of an inner digestive epithelium, a middle layer of connective tissue, and an outer mesothelium (coelomic epithelium). The pharynx, esophagus, and stomach are lined with a cuticular epithelium composed of T-shaped cells. The lining epithelium of the intestine and cloaca lacks a cuticle and consists of columnar vesicular enterocytes. Mucocytes are also encountered in the digestive epithelium. The connective tissue layer is composed of a ground substance, which houses collagen fibers, amoebocytes, morula cells, and fibroblasts. The gut mesothelium is a pseudostratified epithelium, which is dominated by peritoneal and myoepithelial cells and also includes the perikarya and processes of the neurons of the hyponeural plexus and vacuolated cells.