The fine structure of colleterial glands in two cockroaches and three termites, including a detailed study of Cryptocercus punctulatus (Blattaria, Cryptocercidae) and Mastotermes darwiniensis (Isoptera, Mastotermitidae)

The colleterial glands of insects are organs associated with the female genital apparatus. In cockroaches, these glands produce secretions that cover two parallel rows of eggs during oviposition, and in oviparous species, these secretions become the tanned, sculpted, rigid outer casing of the ootheca. The goal of this study was to compare the gross anatomy of the colleterial glands and the ultrastructure of their component tubules in the phylogenetically significant genera Cryptocercus (Blattaria) and Mastotermes (Isoptera). Recent studies indicate that cockroaches in the genus Cryptocercus are the sister group of termites, and Mastotermes is the only termite known to produce a cockroach-like ootheca. One additional oviparous cockroach, Therea, and two additional termites, Zootermopsis and Pseudacanthotermes, were also examined. As in other cockroaches, the colleterial glands of Cryptocercus and Therea are asymmetrical, with a well developed bipartite left gland and a smaller right gland. In the termites Mastotermes, Zootermopsis, and Pseudacanthotermes, the colleterial glands are composed of a well-developed, paired, anterior gland and a small posterior gland; histological staining and cytological evidence suggest that these are homologues of the left and the right colleterial glands of cockroaches, respectively. At the ultrastructural level, colleterial gland tubules are made of cells belonging to a modified class 1 type cell in the cockroaches, in Mastotermes, and in Zootermopsis; the latter lays its eggs singly, without a surrounding ootheca-like structure. In the advanced termite Pseudacanthotermes, the tubules are made of secretory units belonging to the class 3 cell type. This study demonstrates that the cytological characteristics of colleterial glands in basal termites are similar to those of cockroaches, whether the termite secretes an oothecal casing that covers two parallel rows of eggs, as in Mastotermes, or lays its eggs singly, as in Zootermopsis. The function of colleterial glands in non-mastotermitid termites is unknown.     

The structure and possible mode of functioning of the female reproductive system in Nasonia vitripennis (Hymenoptera: Pteromalidae)

Associated with the female insect reproductive system are a number of glands. Those present in Nasonia vitripennis are described, their microanatomy examined and where possible this is linked with their function during the process of drilling into the host puparium, feeding on the host fluids and oviposition. The structures dealt with are the ovaries, spermatheca, alkaline gland, colleterial gland and the acid gland with reservoir.     

Genetic variation in patch time allocation in a parasitic wasp

1. The intra-patch experience acquired by foraging parasitoid females has often been considered to have a strong influence on their tendency to leave a patch, and thus on their total patch residence time. Most studies that have been performed on this subject suggest that the patch-leaving rules observed are adaptive because they enable the females to adjust their patch residence time to local environmental conditions.
2. Considering a behavioural rule as being adaptive supposes that it has been progressively settled by natural selection, and thus that there is, in the population, genetic variation on which the natural selection could act.
3. Therefore, this study aimed to discover whether there was indeed genetic variability in the patch-leaving decision rules in a population of the egg parasitoid species Telenomus busseolae , which attacks patches of its hosts, the eggs of Sesamia nonagrioides . Different wasp families were compared using the isofemale lines method, and the behavioural records were analysed by means of a modified version of the Cox's proportional hazards model proposed by Haccou et al . (1991 ) and Hemerik, Driessen & Haccou (1993 ).
4. The results obtained show that T. busseolae females increase their tendency to leave the patch after each successful oviposition. Each host rejection also led to an increase in the tendency to leave the patch, but this effect was smaller when host rejections were observed between two ovipositions occurring in rapid succession. Subsequent visits to the patch also increased the patch-leaving tendency.
5. Genetic variability was found in both the global patch-leaving tendency and in the effect that successful ovipositions and host rejections have on this tendency.
6. The adaptive and evolutionary consequences of these results are discussed.     

Control of oötheca formation and oviposition in Blattaria

The brain is not required for oviposition in five species of Blaberidae; the control centre for formation, 90° counterclockwise rotation, and retraction of the oötheca lies in the abdomen. A similar centre controls oötheca formation and 90° clockwise rotation in Blattella germanica (Blattellidae). It is suggested that during oviposition, abdominal proprioceptors or musculature contribute nervous information to the last abdominal ganglion. Nerve impulses are presumably integrated in the last abdominal ganglion and transmitted to the colleterial glands, oviducts, and ovipositor. In Periplaneta americana (Blattidae), the brain is needed for initiating egg case formation, but it is unnecessary once the process has begun. The results suggest a divergence of control centres for oviposition between the Blaberoidea and Blattoidea.     

Cellular Differentiation in a Highly Specialized Insect Secretory Organ

The colleterial glands of insects are accessory reproductive structures which produce secretions that are applied to eggs after fertilization and which serve a number of protective functions. The colleterial glands of lepidoptera are of particular interest in the study of the events of cellular differentiation because they undergo rapid development, generally during the pupal adult transformation, and contain highly specialized cells which produce large amounts of a restricted variety of secretory products. The extreme specialization of these organs facilitates parallel studies of differentiation at the biochemical and morphological level. This communication describes the changes in the ultrastructure of cells which will form the protein-secreting segment of the colleterial gland of the moth Actias luna during the period of transition from the undifferentiated state to the acquisition of secretory ability.
An initial stage of general cellular proliferation by mitosis in the presumptive colleterial tissue mass is followed by differentiation of the cells in the absence of further mitosis. Four distinctive cell types are recognized during the phase of differentiation. These types include a chitogenous cell which forms the chitin lining of the main duct, and three cells which cooperate in the formation of a secretory apparatus. Cell A forms two temporary flagella-like structures which assist in the formation of a ductule, which eventually leads from the secretory cell to the main duct. Near the end of the differentiative phase, Cell A degenerates and is phagocitized by Cell B. Cell B becomes the actual secretory element, and acquires cytoplasmic features such as extensive rough endoplasmic reticulum and Golgi apparatus which are associated with synthesis and secretion of protein. The final element, Cell C, remains associated with the ductule which it helps to construct and which traverses its cytoplasm.     

A note on the accessory glands of the reproductive system of the scolopendromorph centipede, Cormocephalus anceps anceps Porat

The function of the reproductive accessory glands has not before been investigated. Their prominence arouses interest in their possible function. A preliminary histological, histochemical, and biochemical analysis has been undertaken in order to start to answer this question. No definite conclusions can, however, be made from the results obtained. From evidence accumulated on the behaviour of the sexes and the peaks of secretion production it seems likely that the product of the female glands would be used to coat the eggs with a colleterial substance. The male Geophilomorpha, Scolopendromorpha, and Lithobiomorpha all spin webs on which they place a spermatophore during courtship, and all three groups possess two well developed pairs of accessory glands which may be responsible for the production of web material. The Scutigeromorpha, on the other hand, only possess a single vestigeal pair of glands and do not spin a web. It is suggested that the accessory glands of the male produce web material although this still remains unproven.     

Proteome analysis of the silkworm (Bombyx mori. L) colleterial gland during different development stages

The silkworm, Bombyx mori, colleterial gland developed very slowly until 2 days before emergence, then markedly enlarged due to the accumulation of a glue-like substances (mainly including 85% water and 11% proteins). However, the No glue (Ng) mutant female moth secreted only very little glue-like substance and laid loose eggs naturally. High-resolution two-dimensional polyacrylamide gel electrophoresis, followed by computer-assisted analysis, was used to screen the secretory region of colleterial gland protein patterns during different development stages to find quantitative and qualitative difference in protein expression during the pupae and moth stages. More than 700 protein spots were resolved in different developmental stages from the secretory region of the glands and most of the proteins were distributed in the mass range from 30 to 70 kD with pH 4-8. Through comparison and analysis, it was found that 3 proteins were only expressed in the later pupae stage (one or two days before emergence) and moth stage. Furthermore, these proteins were not expressed in the Ng mutant especially actin. There was a great variation of some protein expression volume during the development. Protein spots that changed more than 1.5-fold in expression level (relative to day 9), including 6 spots that were down-regulated and 2 spots that were up-regulated in expression were excised for identification by matrix-assisted laser desorption/ionization-time of flight mass spectrometry. Results indicated that actins that participated or regulated the exocytosis of colleterial gland and other differentially expressed proteins might be related to colleterial gland development or the secretion of a glue-like substance.     

柞蚕和柳蚕雌蛾粘液腺及内容物的研究对比分析

本实验测定了柳蚕卵表面的胶着物质的含量以及粘液腺的形状、大小,并对粘液腺的内容物进行了SDS一聚丙烯酰胺凝胶电泳分析。结果表明：柳蚕卵表面的胶着物质的含量为1．11mg／卵,与柞蚕（1．01mg／卵）相当。柞蚕、柳蚕雌蛾的粘液腺整体呈较细长的管状结构且几乎无枝状部分枝,柳蚕粘液腺的干状部长约为2．0cm,柞蚕为1．8cm,二者相近。柳蚕粘液腺的内容物经SDS-聚丙烯酰胺凝胶电泳分析出现了7条蛋白质谱带,分子量分别为114kD,92kD,78kD,53kD,50kD,23kD,12kD,与柞蚕存在差别。     

Ecdysterone antagonism,mimicry, and synergism of juvenile hormone action on the Monarch butterfly reproductive tract

Previous research on Monarch butterflies has shown that juvenile hormone (JH) stimulates the development of the ovary and certain reproductive glands of both sexes. Ecdysterone injections into intact Monarchs demonstrate that low doses of this hormone inhibit ovarian development, and higher doses stimulate the male and female reproductive glands. In addition, experiments using neckligatured adults show that ecdysterone stimulates the reproductive glands of both sexes, in the apparent absence of JH, with the most pronounced effect being observed on the female colleterial gland. Other studies with neck-ligatured animals demonstrate that ecdysterone also synergizes with JH on the female gland and all three male glands. The feasibility of using Monarch reproductive glands for studies on the mode of action and interaction of JH and ecdysterone, and the possibility of a rôle of ecdysterone in the normal regulation of Monarch oögenesis, are discussed.     

Ultrastructure of the accessory glands of gene's organ in the cattle tick, Boophilus

The organization and ultrastructure of the accessory glands of the cattle tick, Boophilus microplus, are described. The glands consist of two groups of acinar cells situated on either side of Gene's organ. A single acinus consists of from eight to 12 cells and each cell is connected via an individual duct to pores on the dorsal surface of the mouthparts. The position of these pores is such that the secretion of the accessory glands is incorporated into the egg wax during oviposition. Each gland cell has striking quantities of smooth endoplasmic reticulum and numerous Golgi dictyosomes and appears to produce a secretion that is lipoidal in nature. Each cell secretes into its own individual lumen and is connected to a cuticular pore by a duct cell.     

The fine structure of the colleterial glands of Hyalophora cecropia (Lepidoptera)

The structure of cells in the colleterial glands of the Cecropia silkmoth was examined. Morphologically and functionally the gland is divided into two regions, a tubular one in which columnar protein-synthesizing cells are located, and an expanded region in which flattened cells with very different structure are most prominent. The fine structure of the latter cells which are presumed to secrete a phenolic glucoside, closely resembles that of cells described in the colleterial glands of orthopterans. The protein-secreting cells have many features normally associated with pancreatic acinar, and other cells of similar function. Among these are extensive rough endoplasmic reticulum, an elaborate Golgi complex, and a modest number of mitochondria. Other features which are less usual in cells of this type are an elaborate secretory apparatus consisting of a cuticular tubule inserted into a microvilli-lined cavity at the apical end of the cell, and large numbers of cytolysomes, myelin figures, and lipid droplets. A chitogenous cell with a very distinct and specific type of ultrastructure is found associated with the secretory cell. This cell type is attached to the cuticular elements of the gland, and the main features of its cytoplasm are extensive bundles of microtubules which presumably serve as supportive elements for the secretory cells.     

Chemical stimuli from corn plants affect host selection and oviposition behavior of Sesamia nonagrioides (Lepidoptera: Noctuidae)

The role of chemical stimuli in host selection and oviposition of Sesamia nonagrioides (Lef) was studied in the laboratory using eight corn hybrids. Female oviposition preference was evaluated on plants of the eight hybrids, their extracts in pentane and methanol, and volatiles collected by steam distillation in ether. Results from two-choice bioassays indicate various degrees of female preference to the hybrids. Four hybrids received more eggs than control plants, whereas the remainder received fewer eggs. Relatively similar results were obtained when filter paper sticks baited with pentane and steam distillation extracts were evaluated. Methanol extracts from all eight hybrids received fewer eggs than the methanol controls. Results showed that pentane and steam distillation extracts contain chemicals that elicit female oviposition response, while methanol extracts may contain chemicals that deter or repel females to oviposition.     

The role of the accessory reproductive glands and genital ducts in egg pod formation in female Schistocerca gregaria

Removal of the accessory reproductive glands in female Schistocerca gregaria does not prevent the formation of eggplugs as expected. The egg pods formed are incomplete, with froth in the egg plug partially lacking. The extent of collapse is dependent on the time of removal of the glands. These results suggest that the genital ducts (egg calyces and lateral oviducts) play a significant role in the production of egg pod material, and the accessory glands probably participate in the process. Ligation of the glands has the same effect on egg pod structure; secretion accumulates in the glands indicating that, although the contribution made to the egg pod by the glands is small, they are not vestigial and are not acting simply as reservoirs for secretions produced distally. Cautery of the Comstock-Kellogg glands produces no visible change in egg pod structure or in method of formation, and thus their function remains undetermined. In the absence of the ventral median pouch egg masses are laid without the plug. A thin layer of nonvacuolated material is smeared over the walls of the oviposition hole, suggesting that the secretions produced in the upper regions of the genital ducts are added to or modified by an additional secretion from this gland, before being released to the exterior.     

L'ultrastruttura delle ghiandole della ooteca in Ortotteri Acridoidei,Blattoidei e Mantoidei

Riassunto La presente ricerca ultrastrutturistica è stata condotta sulle ghiandole che secernono l'ooteca nella femmina degli Ortotteri Acridoidei, dei Blattoidei e dei Mantoidei. Questi tre gruppi sono i soli, fra gli Ortotteroidei, che secernono ooteche indurite e complesse, ma mentre nei Blattoidei e Mantoidei alla secrezione presiedono le primitive ghiandole colleteriche, che sboccano nell'ovidutto impari, negli Acridoidei, che sono di più recente comparsa e (fra gli Ortotteri) i più recentemente evoluti, la secrezione è opera di due diverticoli degli ovidutti pari (ghiandole pseudocolleteriche) che sostituiscono le vere colleteriche perdute lungo la filogenesi dell'ordine Orthoptera. La ultrastruttura delle ghiandole pseudocolleteriche è infatti quella di un tipico organo di recente acquisizione: le due ghiandole sono uguali fra loro, e composte di cellule simili per la intera lunghezza, che secernono tutti i prodotti costituenti l'ooteca: enzimi, proteine strutturali, polisaccaridi.Le due ghiandole colleteriche degli Ortotteroidei più antichi, i Blattoidei, sono invece diverse fra di loro: la sinistra differenzia due tratti secretori: uno ricco di ergastoplasma (che elabora proteine strutturali) ed uno di mitocondri (che elabora una fenolossidasi); la destra pure due tratti, uno con prevalente ergastoplasma (secretore di -glucosidasi) ed uno con abbondanti Golgi e granuli glucidici (elaboratore di un polisaccaride).Nei Mantoidei vi sono 5 ghiandole: una coppia, costituite da numerosi tubuli, che elaborano, da un abbondante ergastoplasma, un secreto protidico e glucidico; una seconda coppia (ramificate e suddivise ciascuna in due diversi tratti) che secernono a monte una ossidasi (sono simili al tratto ricco di mitocondri della sinistra di Periplaneta) e a valle una glucosidasi (sono simili al tratto ricco di ergastoplasma della destra di Periplaneta). La quinta ghiandola, impari, ha i caratteri degli organi secretori di glucidi e protidi e secerne Calcio che viene aggiunto al secreto.

---

Summary This paper deals with the ultrastructure of the glands that secrete the ootheca in the female of three groups of the super order Orthopteroidea: Acridoidea, Blattoidea and Mantoidea. Among the Orthopteroidea these three are the only groups that secrete hardened and complex oothecae. Whereas in the Blattoidea and Mantoidea they are secreted by the primitive colleterial glands that open into the unpaired oviduct, in the more recently evolved Acridoidea the secretion is the product of two loops of the paired oviducts (pseudocolleterial glands). These take the place of the colleterial glands lost in the phylogeny of the order. In fact, the ultrastructure of pseudocolleterial glands is typical of recently acquired organs. The two glands are alike and are composed of the same type of cells throughout. These cells secrete all the products that make up the ootheca: enzymes, structural proteins, and polysaccharides.The two colleterial glands of the most ancient of these three groups of Orthopteroidea, the Blattoidea, are diverse. In the left-hand one, zones can be differentiated. One, rich in ergastoplasm, produces structural proteins and the other, rich in mitochondria, produces a phenoloxidase. The gland on the right also consists of zones: one, rich in ergastoplasm, that secretes -glucoxidase, and the other, with abundant Golgi elements and glucidic granules, that elaborates a polysaccharide.In the Mantoidea there are five glands. One pair is made up of numerous tubules that produce a glucidic and proteinaceous secretion from copious ergastoplasm. In a second pair each gland is branched and subdivided into two diverse parts. The zone at the origin of the gland (like the left-hand zone of Periplaneta rich in mitochondria) secretes an oxidase, whereas the distal portion (like the right-hand zone of Periplaneta, rich in ergastoplasm) secretes a glucoxidase. The fifth gland, unpaired, has the characteristics of an organ that elaborates carbohydrates and proteins and releases calcium, which is added to the secretion.

---

---

Research performed under C.N.R. contract.     

Multiple forms of arylalkylamine N-acetyltransferase (NAT) from cockroach female colleterial glands and activity changes during oocyte maturation

Arylalkylamine N-acetyltransferase (NAT) from the female colleterial glands of Periplaneta americana showed activity peaks at pH 6.0 and 9.5 and the pH profile changed during oogenesis. The left gland contained higher activity than the right gland but the right gland also contained recognizable activity. The patterns in activity change depended on the substrate used, tryptamine (TN) or serotonin (5-HT). When TN was used as the substrate, the alkaline peak was higher than the acidic peak. In contrast, when 5-HT was used, the acidic peak was much higher than the alkaline peak. This suggests that at least two NATs are present in this species that are specific to pH and substrate species. Of the four combinations of the two pH ranges and two substrate indolamines, the enzyme activity that showed a similar change to the oocyte maturation was obtained in the combination of pH 6.0 and TN. TN was actually detected in the colleterial glands by fluorescent measurements according to Hess and Uderfriend [J. Pharmacol. Exp., 127 (1959) 175-177]. It peaked on the 6th day of emergence, which corresponded to the first rise of oocyte length and yolk accumulation, whereas a small peak appeared in the phase of the second rise. TN, or more likely N-acetyl TN, may therefore be involved in the regulation of oocyte maturation which could be a novel mechanism in oocyte maturation.     

Adult emergence rhythm of the egg-parasitoid Telenomus busseolae

The adult emergence rhythm of Telenomus busseolae, an egg parasitoid of Sesamia nonagrioides, was examined when parasitoids were exposed to different light-dark regimes. Most of the adult parasitoids emerged throughout the whole period of the photoperiodic cycle. Peak male emergence occurred 2–5 hours earlier than that of females. Adult emergence was asynchronous in continuous darkness or light. However, regimes of alternative light and dark phases such as L4:D20, L8:D16, L12:D12, L16:D8 and L20:D4 h generated a population rhythm with a period length of 24 hours. The peak of the emergence activity moves from the scotophase to the middle of the photophase with an increase of the photophase from 4 to 20 h. Rhythmical activity of adults was synchronised within 2 cycles when immature stages of parasitoid grow under continuous light conditions (LL) and then transferred to L12:D12. Moreover, emergence rhythm persisted and continued in a free-run with a period length of less than 24 hours by transferring a rhythmic culture from L12:D12 h to LL or RR (continuous red light) conditions, indicating the existence of a circadian rhythm. The ecological implications of the expression rhythm relate to better survival of the parasitoids.     

Histology and functional morphology of the female reproductive tract of the tortoise Gopherus polyphemus

The oviducts of 25 tortoises (Gopherus polyphemus) were examined by using histology and scanning electron microscopy to determine oviductal functional morphology. Oviductal formation of albumen and eggshell was of particular interest. The oviduct is composed of 5 morphologically distinct regions; infundibulum, uterine tube, isthmus, uterus, and vagina. The epithelium consists of ciliated cells and microvillous secretory cells throughout the oviduct, whereas bleb secretory cells are unique to the infundibulum. The epithelium and endometrial glands of the uterine tube histologically resemble those of the avian magnum which produce egg albumen and may be functionally homologous. The isthmus is a short, nonglandular region of the oviduct and appears to contribute little to either albumen or eggshell formation. The uterus retains the eggs until oviposition and may form both the fibrous and calcareous eggshell. The endometrial glands are histologically similar to the endometrial glands of the isthmus of birds, which are known to secrete the fibers of the eggshell. These glands hypertrophy during vitellogenesis but become depleted during gravidity. The uterine epithelium may supply "plumping water" to the egg albumen as well as transport calcium ions for eggshell formation. The vagina is extremely muscular and serves as a sphincter to retain the eggs until oviposition. Sperm are found within the oviductal lumen and endometrial glands from the posterior tube to the anterior uterus throughout the reproductive cycle. This indicates sperm storage within the female tract, although the viability and reproductive significance of these sperm are unknown.     

Short-range cues mediate parasitoid searching behavior on maize: The role of oviposition-induced plant synomones

Host searching by egg parasitoids faces a main constraint due to low detectability of cues from host eggs. Therefore egg parasitoids have developed distinctive strategies by exploiting cues that originate from non-target instars of the host and/or from plants. The scelionid Telenomus busseolae is specialized on concealed eggs of Sesamia nonagrioides and other noctuid stemborers. In this paper we show that oviposition by S. nonagrioides induces changes in the cues present on maize leaf surface, which arrest naïve females of T. busseolae. The induction appears to be systemic as the parasitoid also responds to leaves and leaf portions that are not directly affected by ovipositing females. Such oviposition-induced, short-range, plant synomones, acting in sequence with the kairomonal cues from scales left on the plant by the ovipositing host female, significantly increases parasitoid efficiency during host finding. The elicitor of plant response originates from the host female reproductive system, being contained both in the host’s ovarian eggs and in the colleterial gland secretion. Induction starts 24 h after oviposition and lasts at least till 72 h. The ecological role of this oviposition-induced plant synomone in host searching by T. busseolae is discussed.     

Electroantennogram, wind tunnel and field responses of male Mediterranean corn borer, Sesamia nonagrioides, to several blends of its sex pheromone components

The electrophysiological and behavioural responses of male Sesamia nonagrioides Lefèbvre (Lepidoptera: Noctuidae) to several synthetic blends of its sex pheromone components were investigated. Electroantennogram (EAG) recordings showed the greatest response to a 77:8:10:5 blend of the four compounds found in the female glands: (Z)-11-hexadecen-1-yl acetate, (Z)-11-hexadecen-1-ol, (Z)-11-hexadecenal and dodecyl acetate. A logistic equation was fitted to the dose-response data and differences in relative EAG responses were analysed by deriving five parameters of this equation. Flight responses of males were also evaluated in a wind tunnel where the proposed blend (77:8:10:5) also elicited the best response. Field tests confirmed the laboratory findings as the 77:8:10:5 blend captured more S. nonagrioides males than other pheromone mixtures and showed more selectivity for male S. nonagrioides over other noctuid species, Discestra trifolii Rottenberg and Lacanobia oleracea L., but not for Mythimna unipuncta Haworth, a species with a sex pheromone composition almost identical to that of S. nonagrioides.     

Histochemical analysis of accessory genital gland secretions in female Pimpla turionellae L. (Hymenoptera: Ichneumonidae)

The secretions produced by the 3 accessory glands of the female genital tract of Pimpla turionellae were investigated histochemically. The uterus gland secretion was identified as a mixture of hyaluronic acid and lipoprotein, that of the poison gland as a neutral mucoprotein. The product of the alkaline gland (Dufour's gland) consists of lecithin and a cholesterol ester. Pimpla females, as parasites of Lepidoptera pupae, inject these secretions into the host hemocoel during oviposition, thus preventing hemocytic encapsulation of their eggs. The physiological functions of the secretions are briefly discussed.