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The monoptychic 'apocrine' scent glands of the sternal region of two adult male Tupaia belangeri were studied by transmission and scanning electron microscopy, in order to assess the modes of release of their secretory products. In segments of the secretory tubules with a narrow lumen the epithelial cells are columnar and are firmly connected to each other by desmosomes and junctional complexes. Myoepithelial cells are interspersed between the secretory epithelium and the basement membrane. The cytoplasm of the secretory epithelial cells contains granular and agranular endoplasmic reticulum, mitochondria, Golgi apparatus, lysosomes and secretory granules. The free surface of the secretory epithelium is furnished with densely arranged microvilli. The apices of the cells protrude as dome-shaped extensions into the glandular lumen. At the tip of these extensions cellular processes of irregular shape are found. The surface of these processes shows no microvilli. They contain densely packed and dilated cisternae of agranular endoplasmic reticulum and free ribosomes. The constriction of the base of these processes and the occasional observation of a 'demarcation membrane' between a process and the secretory cell indicate that these secretory processes become detached from the secretory cells according to the apocrine mode of extrusion. Within the glandular lumen they break down and form secretion. Parallel to this apocrine extrusion the same cells produce secretory granules, the diameter of which measures 300-900 nm. The contents of these granules are released into the glandular lumen by exocytosis according to the eccrine mode of secretion. This resembles the situation in other mammals in which monoptychic glands release their secretory products by means of both apocrine and eccrine extrusion. Therefore monoptychic skin glands should not be classified into apocrine and eccrine glands. Our results corroborate Schaffer's (1927, 1940) more general classification of exocrine glands according to the nature of the secretory epithelium into monoptychic and polyptychic glands.
The scent gland and ordinary sebaceous gland of Syrian hamsters and Mongolian gerbils were studied using lectin histochemistry. Male scent glands were larger in size than those of female in both species, but in histochemistry there was no distinct difference between both sexes. Histologically, the scent gland of Syrian hamsters had a strong resemblance to ordinary sebaceous glands, compared to that of Mongolian gerbils. Since the duct of Syrian hamsters was very short, it is difficult to distinguish the duct from the infundibulum. The duct and infundibulum of Mongolian gerbils were larger in diameter than those of Syrian hamsters. The duct widely opened to dermis. Seven lectins (ConA, RCA I, PNA, SBA, UEA-I, DBA and WGA) were used in this study. All of them labelled the differentiated cells of acinus in Syrian hamsters, although SBA did not label those cells in Mongolian gerbils. The cells of the duct in Syrian hamsters and Mongolian gerbils were not labelled with SBA. Ordinary sebaceous glands in both species were labelled with all seven lectins. Lectin-binding patterns of scent glands and sebaceous glands were similar in Syrian hamsters. While, in Mongolian gerbils, SBA-binding patterns were different between scent and sebaceous glands. Histologically, the scent gland of Syrian hamsters resembles the sebaceous gland. Therefore, there is a possibility that the scent gland of Syrian hamsters still provides some properties similar to sebaceous gland.
The scent gland tumor of the Syrian hamster is induced by exogenous androgen and estrogen. Microscopic nodules are induced normally in old males by endogenous androgen. The histogenesis of the scent gland tumor is complex and not completely understood. In this study microscopic preneoplastic nodules and macroscopic tumors were studied by light and electron microscopy, and the macroscopic tumors were grown in tissue culture on collagen-coated coverslips and on sponge foam matrices by the organ culture method. The cultures were fed with an unfiltered fetal calf serum-bovine serum ultrafiltrate medium, which contained endogenous androgen-estrogen, 110-100 pg and could maintain growth without additional androgen-estrogen. Exogenous androgen-estrogen was also added to some cultures. Scent gland tumors grown in organ culture contained cells of two shapes, spindle and ovoid arranged in cords. Cultures on coverslips showed radiating outgrowths of spindle cells suggesting either mesenchymal or Schwann cells. By electron microscopy, both in vivo and in vitro preneoplastic and tumor samples contained cells with segments of basal lamina, micropinocytotic vesicles, and junctional complexes. These features were similar to those of poorly differentiated experimental malignant rat schwannomas maintained in similar in vitro systems. Tumors grown in vivo and in vitro were associated with collagen fibrils with a periodicity ranging from 400 to 1075 A. The evidence reported in this paper suggests that one component of the scent gland tumor is an androgen-estrogen-induced poorly differentiated schwannoma.
The presence of the scent glands were examined macroscopically and histologically in the Japanese vole, Microtus montebelli and the Hungarian vole, Microtus arvalis. A pair of scent glands were observed on the buttocks of the Japanese vole as the oval nodules with yellowish colored skin. The size of the glands was larger in males than in females. Histologically, the glands were composed of the aggregation of many acini of sebaceous glands. The epidermis above the scent glands were thicker than that in the other areas. In the Hungarian vole, no such gland could be detected with macroscopical and histological examinations.
The poison (Zn3P2) bait mixed with preputial gland and cheek gland extract was highly effective as compared to poison bait mixed without the scent gland extract, in increasing the food consumption and mortality rate in rats. Among these two scent glands, preputial gland extract was more effective than cheek gland extract in increasing the bait acceptance. The scent gland extract mixed with poison bait was capable of improving the poison bait acceptance. A 10% concentration of scent gland extract was more effective than 5% concentration in acceptance of poison bait. The results suggest efficacy of scent gland in improving the poison bait acceptance.
A marking-like behavior (defined by authors), a marking behavior, and growth of the scent glands were observed in young Mongolian gerbils of an inbred strain. In males and females, a marking-like behavior, in which animals rub their abdominal scent glands on the floor, began to be seen at the age of 19 days and could be seen in almost all the gerbils at 22 days of age during the suckling period. The frequency of this behavior was highest at 60 days of age (males: 17.9/10 min, females: 15.4/10 min) and there was no sex difference. Marking behavior, in which animals rub their abdominal scent glands on small protruding objects, began to be seen at the age of 40 days in males and 50 days in females. The frequency of this behavior tended to increase until 90 days of age in males (13.7/10 min), but the levels were low (2.5-5.0/10 min) in females. The values in the male group therefore tended to be higher than that in the female group. Macroscopic scent gland pads were clearly observed at the age of 30 days in males, but not until 45 days of age in females. At the age of 45-90 days, the length of the scent gland pad in males and females was 2.1-2.8 and 1.6-1.7 cm, respectively and the width was 0.3-0.5 in males and 0.2-0.3 cm in females. During this period, the length and depth of the pads in males were significantly greater than those in females (p < 0.05). Histological examination of the structure of the scent glands after the age of 45 days showed that the development of clusters of acinar cells in females occurred much later than that in males, but the basic structure of these glands was similar in both sexes. These results suggest that the marking-like behavior was manifested although during the period when the scent glands had not yet developed, whereas true marking behavior first occurred when the glands were moderately well developed.
The histology of the cloacal scent gland, or anal gland, was examined from a diverse group of 50 snakes. Extensive interspecific morphological variation was observed in the general structure of the gland and the glandular epithelium. Morphological variants were quantitatively scored from eight features: lobate nature of the gland; septa in the glandular epithelium; glandular epithelium having a rough or irregular outer surface; glandular epithelium having a rough or irregular inner surface; thickness of the glandular epithelium; PAS reactivity in the glandular epithelium; Periodic Acid-Schiff reactivity in the secretory product; and relative size of the cloacal scent gland. The distribution of the encoded morphological variation was compared to an established phylogeny for snakes and used as the basis for a cluster analysis. In both cases there was no apparent relationship between phylogeny and the morphological variation of the cloacal scent gland.
The genus Didelphis (Marsupialia, Didelphidae) has the unique capacity of supporting both multiplication cycles of Trypanosoma cruzi simultaneously; besides the intracellular forms, the epimastigotes can be found multiplying and differentiating abundantly in the lumen of the scent glands. The biological significance of the life cycle of T. cruzi within the scent glands of Didelphis marsupialis, as well as its contribution to the epidemiology of the disease, is presently unclear. In order to clarify the mechanisms involved in the colonization of this singular habitat by T. cruzi, as well as to understand its biological role, we have carried out a serological and parasitological follow-up of both natural and experimental infections of young and adult opossums. Although all natural infections were stable and long lasting, no infected scent glands were found, indicating that the stability of the systemic infections does not depend on the presence of flagellates in the scent gland. In 84% of the experimentally infected animals the colonization of the scent glands was preceded by a period of patent parasitemia. Parasitism of the scent glands was essentially permanent and bilateral, and its maintenance was independent of circulating parasites. Moreover, the course of the infection differed depending on the source (scent glands versus axenic culture-derived) of the metacyclic forms. Our results suggest that parasitism of the SG of D. marsupialis is most likely a secondary acquisition, a step toward independence from the insect vector, similarly to what is accepted for Trypanosoma equiperdum.
Gas chromatographic-mass spectrometric analyses of the scent gland secretions of Siro duricorius and S. exilis (Opiliones, Cyphophthalmi, Sironidae) revealed a set of 24 components, comprising a series of saturated and unsaturated methyl ketones (C11-C15) and four naphthoquinones. Whereas the scent gland secretions of S. duricorius, collected in Austria, and S. exilis from USA were qualitatively nearly indistinguishable (with the exception of acetophenone that was specific to S. duricorius), they distinctly differed in their relative quantitative compositions: major components of the secretion of S. duricorius were 7-tridecen-2-one, tridecan-2-one, undecan-2-one, 1,4-naphthoquinone, 6-methyl-1,4-naphthoquinone (tentatively identified only), and 4-chloro-1,2-naphthoquinone. In contrast, in S. exilis a compound tentatively identified as 6-methyl-4-chloro-1,2-naphthoquinone was present in large amounts (in S. duricorius a trace component), whereas undecan-2-one only occurred in minor quantities. Secretion profiles of juveniles and adults (both sexes) of each species showed high correspondence. This is the first report on the chemistry of scent gland secretions of the opilionid suborder Cyphophthalmi. 4-Chloro-1,2-naphthoquinone was identified as a new exocrine product of arthropods, whereas 1,4-naphthoquinone and the tentatively identified 6-methyl-1,4-naphthoquinone are known constituents of exocrine secretions from one species of palpatorid opilionids, Phalangium opilio. In contrast, all ketones identified were new for opilionid scent glands, although similar ketones are characteristic of scent gland secretions of palpatorid genera Leiobunum and Hadrobunus. With regard to the near-basic position of Cyphophthalmi in currently proposed phylogenetic trees of Opiliones, naphthoquinones and ketones from Siro may represent the condition ancestral to the (derived) naphthoquinone- and ketone-rich secretions in phalangid Palpatores.
Extrabuccal infralabial secretion outlets (ILOs), periodically absent and heretofore non-described, have been found in nineteen species representing three of the eight psammophine snake genera. These infralabial outlets are thought to be non-existent in the other five genera. A few psammophines representing four genera, among which three of the five without ILOs, are characterised by their 'self-rubbing' behaviour, for which they employ an external narial valve bearing a secretion outlet of a special nasal gland. Such a valve is here reported for representatives of all eight genera. It is proposed that all psammophines perform self-rubbing to enable subsequent chemical marking of substrate and conspecific individuals. The newly found infralabial outlets, however, serve for direct chemical marking of conspecifics. This direct marking seems to substitute for a function of both the special nasal gland and the cloacal scent glands during social activity of the snakes in trees or shrubs. This hypothesis and the relationship of the observed traits with the presumed advantages of a thin and short hemipenis are discussed. A taxonomic implication is suggested.
Marking behavior, marking-like behavior [3], and changes of the scent glands were observed in aged Mongolian gerbils. In Experiment 1, changes in the marking and marking-like behavior with aging were evaluated in adult male and female Mongolian gerbils of an inbred strain aged 6 to 36 months. The frequency of marking behavior in males was significantly higher than females throughout the observation period except at 36 months of age. On the other hand, frequency of marking-like behavior in males, but not in females decreased with aging, significantly. In Experiment 2, changes of the scent gland in adult males and females aged 6 to 36 months were morphologically evaluated. Macroscopic examination revealed an increase in the size length and width of the glands of males aged 12 months and females aged 6 months. Histologically the glands of all the males and females aged 6 months developed moderately or well. Some of the 12-month-old males and females showed acinar atrophy of the glands, and all the females aged 18 months or more had highly atrophied scent glands. From these results, we concluded that there is no relationship between the changes of marking behavior and those of the scent glands in aged male Mongolian gerbils, and assume that marking behavior in aged animals does not have an important meaning as marking. In Experiment 3, marking and marking-like behavior in castrated adult Mongolian gerbils aged 16 weeks were observed. The result showed that marking behavior, not marking-like behavior was inhibited after castration. From these findings, we consider that generally marking behavior in Mongolian gerbils consists of androgen-dependent marking behavior and androgen-independent marking behavior (marking-like behavior).
Previous research [J. Neurosci. 21 (2001) 5832-5840] showed that ferrets of both sexes require olfactory signals to identify opposite-sex mating partners at a distance. The present experiments assessed the contributions of anal scent gland and urinary odorants to these preferences. Sexually experienced, ovohysterectomized female and castrated male ferrets were injected daily with estradiol benzoate and testosterone propionate, respectively. When tested in an airtight Y-maze, subjects of both sexes preferred to approach volatile odors emitted from opposite- versus same-sex stimulus ferrets that were anesthetized and placed in the goal boxes, regardless of whether the anal scent glands of stimulus ferrets had been surgically removed or left intact. Subjects of each sex showed an equal preference to approach volatile odors emitted from anesthetized opposite-sex ferrets that were scent-gland intact as opposed to descented. Female subjects preferred to approach volatile anal scent gland odorants, as well as urinary odorants from male, as opposed to female conspecifics. Male subjects preferred to approach volatile anal scents from females versus males; however, males showed no preference for female over male urinary odorants. Our results suggest that anal scent gland odorants are sufficient, but not required, for mate recognition in the ferret. Instead, a combination of body odorants including, but not restricted to, those derived from anal scent gland secretions apparently underlie olfactory sex discrimination and partner preference in this carnivore.
This study examined the dynamics of colonization of Trypanosoma cruzi in the scent glands of the opossum Didelphis marsupialis following direct inoculation with 10(5) epimastigotes of isolate G-49 (an opossum-derived strain). One, three, and five days, 1 month, and 1 year after inoculation, scent glands were fixed for analysis using brightfield and electron microscopies. One day after inoculation the parasites, mainly as epimastigotes, were randomly distributed into the lumen. From the third day on, the parasites still in the form of epimastigotes tended to concentrate closer to the epithelium. The flagellates reached the definitive distribution pattern on the fifth day, when they formed huge clusters deep into the foveae. In samples collected 1 month and 1 year after inoculation, the ratio of epimastigotes:trypomastigotes was 1:1, with epimastigotes predominating near the epithelium and trypomastigotes far from it. Our observations suggest that T. cruzi grows continuously in the scent glands and does not depend on adhesion to promote metacyclogenesis. Metacyclogenesis far from the epithelium seems to be an important selective advantage to both host and parasite, since it assures the elimination of the infective forms of the parasite when the host expels the glands' contents, which occurs in frightening situations or at times of stress. The morphological characteristics of infected and noninfected scent glands using transmission and scanning electron microscopies were also described.
Infants of gerbil mothers whose ventral scent gland has been excised were compared on a number of developmental indices with pups of mothers who had had an equivalent section of lateral skin removed. The removal of the mother's ventral scent gland retarded the offsprings' pattern of ultrasounding during the 1st 21 days of development. In addition, infants from such mothers showed a retarded righting reflex. Inhibitions in the developmental behavior of gerbil pups might have resulted from less effective heat transfer during mother-pup contact. Indeed, excision of the gland resulted in lower ventral temperatures. The duality of function of the gland as an odor source for communication and as a heat source for offspring development suggests integrated communicative and thermoregulatory processes.
Body odors contribute to mate recognition and sexual partner preference in many mammals, including ferrets. We used a habituation/dishabituation procedure to test whether sex steroid hormones influence whether ferrets will approach and investigate different concentrations of volatile anal scent gland odors from male and female conspecifics. When tested with high concentrations of anal scent gland secretions in oil vehicle, gonadectomized male and female ferrets that received no sex steroids reliably discriminated anal scents from male and female conspecifics. This discrimination most likely reflects gender recognition rather than individual recognition because gonadectomized, sex steroid-treated ferrets discriminated between anal scents of males and females but not between anal scents of individual males or females. Treatment with either the estrogen receptor agonist, estradiol benzoate (EB), or the androgen receptor agonist, 5-alpha dihydrotestosterone proprionate (DHTP), increased investigation of low concentrations of anal scent by gonadectomized ferrets. These data suggest that ferrets could use anal scent gland secretions in mate recognition and that seasonal increases in circulating sex steroid hormones increase ferrets' responsiveness to low concentrations of these odors.
A comparative histological study of the papilla and fundic region of the scent glands of Mephitis mephitis, M. macroura, Spilogale putorius. S. pygmaea and Conepatus mesoleucus was conducted. Both the papilla and fundic region consist of a mucosa and a muscularis. The entire organ is covered by a fibrous capsule. The glandular tissue has a compound tubulo-alveolar arrangement where both the alveoli and tubules are lined by simple cuboidal serous cells. The scent glands are highly modified, enlarged, apocrine sweat glands. Differences in both the papilla and fundic region occur between the species and involve epithelial keratinization, vacuolization of the mucosa and arrangement of secretory tissue in the gland.