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[Data on the biological cycle of the ascaridoid Ophidascaris trichuriformis, parasite of snakes].

Ophidascaris trichuriformis is an ascaridoid commonly found in some species of Colubridae snakes. Similarly to what has been observed with several ascaridoid species, in O. trichuriformis the larvae, when still inside the egg-shells, reach the 3th snake stage around the 8th day of incubation in cultures kept at around 25 degrees C. Tentative assays to infect mice, snakes, toads, fish, and tadpoles of Hyla fuscovaria and Bufo sp. was made through the administration of embryonated eggs. Eggs didn't even eclode in mice, snakes or toads in such experiments; while in fish, although the eggs ecloded, its larvae were eliminated after the 4th day. In H. fuscovaria and Bufo sp. tadpoles, larvae were liberated from egg-shells and their development was observed up to the 36th day. They grew from the 6th hour to the above mentioned day, after which larvae were longer in Bufo sp. than in H. fuscovaria. By the 36th day distinction of sexes was possible through examination of genital primordia whose morphology and location differed in male and female larvae. From the 30th day on, following the recognizable beginning of metamorphosis in anuran amphibians and going up to the 65th day after experimental infection, an intense growth of the larvae infecting Bufo sp. was observed; the growth of such larvae was negligible between the 65th and the 85th days. Five snakes (Waglerophis merremii) were given per os larvae obtained from Bufo sp. after the 65th day of infection; these snakes died on the 53th, 54th, 60th, 110th and 116th days after their own infection, respectively, and in each of then 4th stage O. trichuriformis larvae were found in the stomach. The gastric location of such larvae in each snake was identical to the one in which adult worms of this same species are regularly found in natural infections: all of them had the middle part of the body embedded into the stomach wall while both anterior and posterior extremities kept free towards the lumen of the organ.

Animals↗

Surface area to volume relationships of snakes support the use of allometric scaling for calculating dosages of pharmaceuticals.

Metabolic scaling is used to estimate pharmaceutical dosages for snakes because few pharmacokinetic studies have determined optimum dosages. The question arose as to the validity of the assumption of metabolic scale among snakes because they have such a cylindrical shape. The surface area to volume relationships of a group of animals is a major portion of the metabolic scaling exponent of 0.75. An assumption inherent in scaling pharmaceutical dosages for snakes is that there is not a shape change with a size change, or that scaling of the surface area to volume has an exponent of 0.66. One hundred forty snakes, 106 Colubridae and 34 Boiidae, were weighted, their volume was determined by water displacement, and the snakes were measured. Geometric formulas were used to calculate surface area and volume of each snake. Surface area to volume relationship was nearly isometric, with a scaling exponent of 0.64, supporting the use of metabolic scaling for estimating pharmaceutical doses.

Animals↗

Sarcocystis singaporensis: studies on host specificity, pathogenicity, and potential use as a biocontrol agent of wild rats.

Host specificity and pathogenicity of Sarcocystis singaporensis were investigated as a prerequisite to a subsequent application of the parasite as a biocontrol agent of wild rats in Egypt. After inoculation of 7 snake species comprising the families Elapidae, Viperidae, Colubridae, and Boidae with sarcocysts, sporocyst development was only observed in a reticulated python. Among amphibians, reptiles, and rodents that orally received various sporocyst doses in the laboratory, 2 x 10(4) sporocysts or more were lethal to roof rats Rattus rattus frugivorous, brown rats Rattus norvegicus, and bandicoot rats Nesokia indica. Sarcocysts developed in Rattus spp. and Nile grass rats Arvicanthis niloticus. Subsequently, the pathogenicity of S. singaporensis was tested under natural control situations offering bait pellets containing high amounts of sporocysts to a free-living population of roof rats, which was monitored by indirect census baiting commonly used in rodenticide evaluation. Ten days after consumption of the bait pellets, the infected population collapsed, leading to a control success of 73%. A negative control population, which received a placebo, remained stable. These data demonstrate for the first time that S. singaporensis can be used as a biocontrol agent of wild rats. However, an immunization experiment with roof rats in the laboratory showed that these are capable of mounting a rapid specific immune response resulting in survival of acute sarcocystosis.

Amphibians↗

Organization of the ophidian amygdala: chemosensory pathways to the hypothalamus.

Although recent studies in squamate reptiles have importantly clarified how chemical information is processed in the reptilian brain, how the amygdala relays chemosensory inputs to the hypothalamus to influence chemically guided behaviors is still poorly documented. To identify these chemosensory pathways, the amygdalo-hypothalamic projections, intra-amygdaloid circuitry and afferents from the lateral cortex (LC) to the amygdala were investigated by injecting conjugated dextran-amines into the hypothalamus, amygdala, and LC of garter snakes. The amygdala was divided into olfactory recipient (ventral anterior and external amygdalae), vomeronasal recipient (nucleus sphericus, NS, and medial amygdala, MA), and nonchemosensory (e.g., posterior dorsal ventricular ridge, PDVR, and dorsolateral amygdaloid nucleus, DLA) subdivisions. Rostroventral (LCrv) and dorsocaudal subdivisions of the LC were distinguished. In addition to receiving afferents from the main olfactory bulb, the olfactory amygdala receives afferents from NS and projects to the NS, PDVR, and dorsal hypothalamus. The NS has only a minor projection to the lateral hypothalamus, whereas the MA, which receives afferents from the LCrv and NS, has projections to the ventromedial hypothalamic (VMH) and lateral posterior hypothalamic nuclei. Among the nonchemosensory amygdaloid structures, the PDVR receives afferents from the LCrv and the olfactory amygdala and projects to the VMH, whereas DLA receives afferents from the LCrv and NS, and projects to the periventricular hypothalamus. These results substantially clarify the olfactory and vomeronasal tertiary connections and demonstrate that parts of the nonchemosensory amygdala play a major role in relaying chemosensory information to the hypothalamus.

Amygdala↗

Pinealectomy, melatonin, and courtship behavior in male red-sided garter snakes (Thamnophis sirtalis parietalis).

Activation of courtship behavior in male red-sided garter snakes is independent of androgens. Only exposure to extended periods of low temperature with subsequent warming stimulates courtship in males. The pineal gland is thought to transduce temperature as well as photoperiodic information in reptiles. Therefore, we explored the relationship of the pineal and melatonin to sexual behavior in this species. Pinealectomy of male garter snakes disrupted sexual behavior upon emergence from a 17-week period of low temperature in approximately 60% of treated individuals in each of the 3 years of study. However, 40% of the males were unaffected by the pinealectomy, engaging in vigorous courtship. Administration of exogenous, chronic melatonin did not significantly modulate the effect of pinealectomy. Upon pinealectomy in the autumn (before hibernation), plasma levels of melatonin fell. However, upon emergence from hibernation, melatonin levels in pinealectomized (PINX) and sham-treated (SHAM) animals were equivalent, indicating extrapineal source(s) of melatonin. However, PINX males did not exhibit a diel cycle in melatonin levels upon emergence. Instead, melatonin remained elevated through the subsequent 24-hr period. SHAMs did exhibit a diel cycle. Ten days after emergence, PINX animals either had a disrupted/abnormal melatonin cycle and were non-courters or had a cycle similar to SHAM males and courted. Therefore, a normal diel cycle of melatonin appeared necessary for the proper expression of courtship behavior. These results suggest that the pineal in snakes 1) is part of a complex, multi-oscillator system as it is in birds and lizards and 2) may play a role in maintaining polymorphism in timing of reproductive behavior.

Acclimatization↗

Three-dimensional ultrastructure of the surface of the tongue of the rat snake, Elaphe climacophora.

BACKGROUND: Many studies have been performed to clarify the relationship between behavioral performance of the tongue and Jacobson's organ. The purpose of the present study was to examine the ultrastructural features of the surface of the tongue of the rat snake, Elaphe climacophora, and to delineate the functional relationship between the tongue and Jacobson's organ from a morphological perspective. METHODS: The three-dimensional ultrastructure of the surface of the tongue of the rat snake Elaphe climacophora was investigated by scanning electron microscopy. RESULTS: Most of the surface of the bifurcated apex of the tongue was relatively smooth. Dome-shaped, hemispherical bulges or microfacets were compactly arranged on the epithelial cell surface over this entire region. Intercellular borders were clearly recognizable as striations. These features were almost the same as those of the dorsal surface of the transitional area between the bifurcated lingual apex and the anterior part of the lingual body. In the posterior half of the lingual body, no microfacets were seen at all. Both microridges and microvilli were compactly distributed on cell surfaces. CONCLUSIONS: No evidence was obtained from our ultrastructural analysis for an important role of the lingual apex in the vomeronasal system. By contrast, the epithelial surface of the body of the tongue appeared suitable for retaining stimulating compounds.

Animals↗

Contribution of the vertebral artery to cerebral circulation in the rat snake Elaphe obsoleta.

Blood supplying the brain in vertebrates is carried primarily by the carotid vasculature. In most mammals, cerebral blood flow is supplemented by the vertebral arteries, which anastomose with the carotids at the base of the brain. In other tetrapods, cerebral blood is generally believed to be supplied exclusively by the carotid vasculature, and the vertebral arteries are usually described as disappearing into the dorsal musculature between the heart and head. There have been several reports of a vertebral artery connection with the cephalic vasculature in snakes. We measured regional blood flows using fluorescently labeled microspheres and demonstrated that the vertebral artery contributes a small but significant fraction of cerebral blood flow (approximately 13% of total) in the rat snake Elaphe obsoleta. Vascular casts of the anterior vessels revealed that the vertebral artery connection is indirect, through multiple anastomoses with the inferior spinal artery, which connects with the carotid vasculature near the base of the skull. Using digital subtraction angiography, fluoroscopy, and direct observations of flow in isolated vessels, we confirmed that blood in the inferior spinal artery flows craniad from a point anterior to the vertebral artery connections. Such collateral blood supply could potentially contribute to the maintenance of cerebral circulation during circumstances when craniad blood flow is compromised, e.g., during the gravitational stress of climbing.

Angiography↗

Testing an inference of function from structure: snake vertebrae do the twist.

The zygapophyses and zygosphene-zygantrum articulations of snake vertebrae are hypothesized to restrict or eliminate vertebral torsion. This hypothesis is apparently based solely on the inference of function from structure, despite the limitations of such inferences, as well as contradictory observations and measurements. In this study, I observed and measured axial torsion in gopher snakes, Pituophis melanoleucus. To examine the structural basis of axial torsion, I measured the vertebral articulation angles along the body and the insertion angles of five epaxial muscles. To examine torsion in a natural behavior, I digitized video images and measured the degree of apparent axial torsion during terrestrial lateral undulation. Finally, I measured the mechanical capacity of the vertebral joints for actual torsion over intervals of 10 vertebrae in fresh, skinned segments of the trunk. Vertebral articulation angles vary up to 30 degrees and are associated with variation in torsional capacity along the trunk. The freely crawling P. melanoleucus twisted up to 2.19 degrees per vertebra, which produced substantial overall torsion when added over several vertebrae. The vertebral joints are mechanically capable of torsion up to 2.89 degrees per joint. Therefore, despite the mechanical restriction imposed by the complex articulations, vertebral torsion occurs in snakes and appears to be functionally important in several natural behaviors. Even in cases in which mechanical function appears to be narrowly constrained by morphology, specific functions should not be inferred solely from structural analyses.

Animals↗

Environmental and seasonal adaptations of the adrenocortical and gonadal responses to capture stress in two populations of the male garter snake, Thamnophis sirtalis.

Stress and reproduction are generally thought to work in opposition to one another. This is often manifested as reciprocal relationships between glucocorticoid stress hormones and sex steroid hormones. However, seasonal differences in how animals respond to stressors have been described in extreme environments. We tested the hypothesis that garter snakes, Thamnophis sirtalis, with limited reproductive opportunities will suppress their hormonal stress response during the breeding season relative to conspecifics with an extended breeding season. The red-sided garter snake, T.s. parietalis, of Manitoba, Canada, has a brief breeding season during which males displayed no change in either plasma levels of testosterone or corticosterone, which were both elevated above basal levels, in response to capture stress. During the summer, capture stress resulted in increased plasma corticosterone and decreased testosterone. During the fall, when mating can also occur, males exhibited a significant decrease in testosterone but no increase in corticosterone in response to capture stress. The red-spotted garter snake, T.s. concinnus, of western Oregon, has an extended breeding season during which males displayed a stress response of increased plasma corticosterone and decreased testosterone levels. The corticosterone response to capture stress was similar during the spring, summer, and fall. In contrast, the testosterone response was suppressed during the summer and fall when gametogenesis was occurring. These data suggest that male garter snakes, in both populations, seasonally adapt their stress response but for different reasons and by potentially different mechanisms. J. Exp. Zool. 289:99-108, 2001.

Adaptation, Physiological↗

Neural substrates for tongue-flicking behavior in snakes.

Snakes deliver odorants to the vomeronasal organ by means of tongue-flicks. The rate and pattern of tongue-flick behavior are altered depending on the chemical context. Accordingly, olfactory and vomeronasal information should reach motor centers that control the tongue musculature, namely, the hypoglossal nucleus (XIIN); however, virtually nothing is known about the circuits involved. In the present work, dextran amines were injected into the tongue of garter snakes (Thamnophis sirtalis) to identify the motoneurons of the XIIN. Tracers were then delivered into the XIIN to identify possible afferents of chemical information. Large injections into the XIIN yielded retrograde labeling in two chemosensory areas: the medial amygdala (MA) and the lateral posterior hypothalamic nucleus (LHN). Smaller injections only yielded labeled neurons in the LHN. In fact, the MA, which receives afferents from the accessory olfactory bulb, the rostroventral lateral cortex, and the nucleus sphericus, projects to the LHN. Injections into the MA did not show terminal labeling in the XIIN but in an area lateral to it. However, injections into the LHN gave rise not only to labeled fibers in the XIIN but also to retrograde labeling in the MA, thus confirming the chemosensory input to LHN. Injecting different fluorescent tracers into the tongue and into the LHN corroborated the projection from the LHN to the XIIN. The present report investigates further connections of the olfactory and vomeronasal systems and describes the afferent connections to XIIN in a nonmammalian vertebrate. The circuit for tongue-flicking behavior described herein should be evaluated using functional studies.

Amygdala↗

Evolutionary transformations of the fetal membranes of viviparous reptiles: a case study of two lineages.

The reptilian placenta is a composite structure formed by a functional interaction between extraembryonic membranes and the maternal uterus. Study of placental structure of squamate reptiles over the past century has established that each of the multiple independent origins of placentation, which characterize the reproductive diversity of squamates, has resulted from the evolutionary transformation of these homologous structures. Because each evolutionary transformation is an independent novel relationship between maternal and embryonic tissues, the resulting placentae are not homologous, even though the individual components may be. The evolution of reptilian placentation should reveal much about evolutionary patterns and mechanisms because similar structural-functional systems have been transformed along parallel trajectories on multiple occasions. We compared extraembryonic membrane and placental development and pattern of embryonic nutrition in thamnophiine snakes and Pseudemoia lizards in the context of recent hypotheses of phylogenetic relationships. Two primary types of placentation, chorioallantoic and yolk sac, evolved in each lineage. Smooth, highly vascular regions of chorioallantoic placentation are indistinguishable homoplasies that evolved in parallel, likely to facilitate respiratory exchange. The yolk sac placenta of each lineage is specialized for histotrophic nutrient transfer, yet composition of these structures differs because of variation in the ancestral snakes and lizards. In addition, the omphalopleure that contributes to yolk sac placentation persists to later embryonic stages compared to oviparous outgroups, but the two lineages have evolved different structures that prevent replacement of the omphalopleure by the allantois. Each lineage has also evolved unique structural specializations of the chorioallantoic placenta.

Allantois↗

Prenatal sex ratios and expression of sexually dimorphic traits in three snake species.

Variation in intrauterine exposure to hormones associated with variation in the sex of litter mates has well-established and far-reaching effects on sexual development in some mammals. Research on this phenomenon in reptiles is scant, but suggests that lizards may follow the mammalian model whereas snakes may be affected differently. We examined sex-specific expression of four sexually dimorphic traits (tail length, head length, ventral scale count, swimming speed) in three species of snakes (Nerodia sipedon, Thamnophis sirtalis, T. sauritus) relative to litter sex ratios. We found little evidence that traits in either sex were masculinized or feminized in response to variation in litter sex ratio. The one significant result appeared best explained as a statistical artifact attributable to a single litter. Our results indicate that snakes are different from the one lizard studied to date. Unlike previous suggestions that prenatal hormonal mechanisms operate differently in snakes and lizards, however, the difference appears to be that development of sexually dimorphic traits in lizards is affected by litter sex ratios whereas in snakes it is not.

Animals↗