Morphological respiratory diffusion capacity of the lungs of ball pythons (Python regius)

This study aims at a functional and morphological characterization of the lung of a boid snake. In particular, we were interested to see if the python's lungs are designed with excess capacity as compared to resting and working oxygen demands. Therefore, the morphological respiratory diffusion capacity of ball pythons (Python regius) was examined following a stereological, hierarchically nested approach. The volume of the respiratory exchange tissue was determined using computed tomography. Tissue compartments were quantified using stereological methods on light microscopic images. The tissue diffusion barrier for oxygen transport was characterized and measured using transmission electron micrographs. We found a significant negative correlation between body mass and the volume of respiratory tissue; the lungs of larger snakes had relatively less respiratory tissue. Therefore, mass-specific respiratory tissue was calculated to exclude effects of body mass. The volume of the lung that contains parenchyma was 11.9±5.0mm(3)g(-1). The volume fraction, i.e., the actual pulmonary exchange tissue per lung parenchyma, was 63.22±7.3%; the total respiratory surface was, on average, 0.214±0.129m(2); it was significantly negatively correlated to body mass, with larger snakes having proportionally smaller respiratory surfaces. For the air-blood barrier, a harmonic mean of 0.78±0.05μm was found, with the epithelial layer representing the thickest part of the barrier. Based on these findings, a median diffusion capacity of the tissue barrier ( [Formula: see text] ) of 0.69±0.38ml O(2)min(-1)mmHg(-1) was calculated. Based on published values for blood oxygen concentration, a total oxygen uptake capacity of 61.16mlO(2)min(-1)kg(-1) can be assumed. This value exceeds the maximum demand for oxygen in ball pythons by a factor of 12. We conclude that healthy individuals of P. regius possess a considerable spare capacity for tissue oxygen exchange.     

The effect of meal composition on specific dynamic action in burmese pythons (Python molurus)

We quantified the specific dynamic action (SDA) resulting from the ingestion of various meal types in Burmese pythons (Python molurus) at 30 degrees C. Each snake was fed a series of experimental meals consisting of amino acid mixtures, simple proteins, simple or complex carbohydrates, or lipids as well as meals of whole animal tissue (chicken breast, beef suet, and mouse). Rates of oxygen consumption were measured for approximately 4 d after feeding, and the increment above standard metabolic rate was determined and compared to energy content of the meals. While food type (protein, carbohydrate, and lipid) had a general influence, SDA was highly dependent on meal composition (i.e., amino acid composition and carbohydrate structure). For chicken breast and simple carbohydrates, the SDA coefficient was approximately one-third the energetic content of the meal. Lard, suet, cellulose, and starch were not digested and did not produce measurable SDA. We conclude that the cost of de novo protein synthesis is an important component of SDA after ingestion of protein meals because (1) simple proteins, such as gelatin and collagen, did not stimulate levels of SDA attained after consumption of complete protein, (2) incomplete mixtures of amino acids failed to elicit the SDA of a complete mixture, and (3) the inhibition of de novo protein synthesis with the drug cycloheximide caused a more than 70% decrease in SDA. Stomach distension and mechanical digestion of intact prey did not cause measurable SDA.     

Autonomic control of heart rate during orthostasis and the importance of orthostatic-tachycardia in the snake Python molurus

Orthostasis dramatically influences the hemodynamics of terrestrial vertebrates, especially large and elongated animals such as snakes. When these animals assume a vertical orientation, gravity tends to reduce venous return, cardiac filling, cardiac output and blood pressure to the anterior regions of the body. The hypotension triggers physiological responses, which generally include vasomotor adjustments and tachycardia to normalize blood pressure. While some studies have focused on understanding the regulation of these vasomotor adjustments in ectothermic vertebrates, little is known about regulation and the importance of heart rate in these animals during orthostasis. We acquired heart rate and carotid pulse pressure (P _PC) in pythons in their horizontal position, and during 30 and 60° inclinations while the animals were either untreated (control) or upon muscarinic cholinoceptor blockade and a double autonomic blockade. Double autonomic blockade completely eradicated the orthostatic-tachycardia, and without this adjustment, the P _PC reduction caused by the tilts became higher than that which was observed in untreated animals. On the other hand, post-inclinatory vasomotor adjustments appeared to be of negligible importance in counterbalancing the hemodynamic effects of gravity. Finally, calculations of cardiac autonomic tones at each position revealed that the orthostatic-tachycardia is almost completely elicited by a withdrawal of vagal drive.     

Experimentally derived salinity tolerance of hatchling Burmese pythons (Python molurus bivittatus) from the Everglades,Florida (USA)

In a laboratory setting, we tested the ability of 24 non-native, wild-caught hatchling Burmese pythons (Python molurus bivittatus) collected in the Florida Everglades to survive when given water containing salt to drink. After a one-month acclimation period in the laboratory, we grouped snakes into three treatments, giving them access to water that was fresh (salinity of 0, control), brackish (salinity of 10), or full-strength sea water (salinity of 35). Hatchlings survived about one month at the highest marine salinity and about five months at the brackish-water salinity; no control animals perished during the experiment. These results are indicative of a “worst-case scenario”, as in the laboratory we denied access to alternate fresh-water sources that may be accessible in the wild (e.g., through rainfall). Therefore, our results may underestimate the potential of hatchling pythons to persist in saline habitats in the wild. Because of the effect of different salinity regimes on survival, predictions of ultimate geographic expansion by non-native Burmese pythons that consider salt water as barriers to dispersal for pythons may warrant re-evaluation, especially under global climate change and associated sea-level-rise scenarios.     

Functional morphology and patterns of blood flow in the heart of Python regius

Brightness‐modulated ultrasonography, continuous‐wave Doppler, and pulsed‐wave Doppler‐echocardiography were used to analyze the functional morphology of the undisturbed heart of ball pythons. In particular, the action of the muscular ridge and the atrio‐ventricular valves are key features to understand how patterns of blood flow emerge from structures directing blood into the various chambers of the heart. A step‐by‐step image analysis of echocardiographs shows that during ventricular diastole, the atrio‐ventricular valves block the interventricular canals so that blood from the right atrium first fills the cavum venosum, and blood from the left atrium fills the cavum arteriosum. During diastole, blood from the cavum venosum crosses the muscular ridge into the cavum pulmonale. During middle to late systole the muscular ridge closes, thus prohibiting further blood flow into the cavum pulmonale. At the same time, the atrio‐ventricular valves open the interventricular canal and allow blood from the cavum arteriosum to flow into the cavum venosum. In the late phase of ventricular systole, all blood from the cavum pulmonale is pressed into the pulmonary trunk; all blood from the cavum venosum is pressed into both aortas. Quantitative measures of blood flow volume showed that resting snakes bypass the pulmonary circulation and shunt about twice the blood volume into the systemic circulation as into the pulmonary circulation. When digesting, the oxygen demand of snakes increased tremendously. This is associated with shunting more blood into the pulmonary circulation. The results of this study allow the presentation of a detailed functional model of the python heart. They are also the basis for a functional hypothesis of how shunting is achieved. Further, it was shown that shunting is an active regulation process in response to changing demands of the organism (here, oxygen demand). Finally, the results of this study support earlier reports about a dual pressure circulation in Python regius. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Effects of meal size, clutch, and metabolism on the energy efficiencies of juvenile Burmese pythons, Python molurus

We explored meal size and clutch (i.e., genetic) effects on the relative proportion of ingested energy that is absorbed by the gut (apparent digestive efficiency), becomes available for metabolism and growth (apparent assimilation efficiency), and is used for growth (production efficiency) for juvenile Burmese pythons (Python molurus). Sibling pythons were fed rodent meals equaling 15%, 25%, and 35% of their body mass and individuals from five different clutches were fed rodent meals equaling 25% of their body mass. For each of 11–12 consecutive feeding trials, python body mass was recorded and feces and urate of each snake was collected, dried, and weighed. Energy contents of meals (mice and rats), feces, urate, and pythons were determined using bomb calorimetry. For siblings fed three different meal sizes, growth rate increased with larger meals, but there was no significant variation among the meal sizes for any of the calculated energy efficiencies. Among the three meal sizes, apparent digestive efficiency, apparent assimilation efficiency, and production efficiency averaged 91.0%, 84.7%, and 40.7%, respectively. In contrast, each of these energy efficiencies varied significantly among the five different clutches. Among these clutches production efficiency was negatively correlated with standard metabolic rate (SMR). Clutches containing individuals with low SMR were therefore able to allocate more of ingested energy into growth.     

Retained caseous yolk sac in a Burmese python (Python molurus bivittatus)

Retained yolk sacs are common in the domestic chicken and account for considerable morbidity and mortality during late embryonic development and within the first 10 days of life. What is believed to be the first recorded instance of a retained caseous yolk sac and its successful surgical removal from a Burmese python (Python molurus bivittatus) is reported. The snake experienced no post-operative complications and continues to be well 16 mo following surgery.     

Effects of meal size, clutch, and metabolism on the energy efficiencies of juvenile Burmese pythons, Python molurus.

We explored meal size and clutch (i.e., genetic) effects on the relative proportion of ingested energy that is absorbed by the gut (apparent digestive efficiency), becomes available for metabolism and growth (apparent assimilation efficiency), and is used for growth (production efficiency) for juvenile Burmese pythons (Python molurus). Sibling pythons were fed rodent meals equaling 15%, 25%, and 35% of their body mass and individuals from five different clutches were fed rodent meals equaling 25% of their body mass. For each of 11-12 consecutive feeding trials, python body mass was recorded and feces and urate of each snake was collected, dried, and weighed. Energy contents of meals (mice and rats), feces, urate, and pythons were determined using bomb calorimetry. For siblings fed three different meal sizes, growth rate increased with larger meals, but there was no significant variation among the meal sizes for any of the calculated energy efficiencies. Among the three meal sizes, apparent digestive efficiency, apparent assimilation efficiency, and production efficiency averaged 91.0%, 84.7%, and 40.7%, respectively. In contrast, each of these energy efficiencies varied significantly among the five different clutches. Among these clutches production efficiency was negatively correlated with standard metabolic rate (SMR). Clutches containing individuals with low SMR were therefore able to allocate more of ingested energy into growth.     

蟒的人工饲养初探

本文对2001～2007年先后饲养于成都动物园的14条蟒的食性、行为和护理等进行了观察。分析结果为,蟒行为活动受季节和气温变化影响大,夏季活动与休息的比例为1∶9;冬季活动与休息的比例仅1∶60。人工饲养的适宜温度为25～30℃,相对湿度为50%～90%;安全越冬的基础温度为20～22℃,相对湿度60%～90%。可以自主采食兔子、鸡、鸭和大白鼠等活体动物,而人工填食可用牛肉、精猪肉。易患寄生虫、口腔炎、感冒并继发肺炎等疾病,人工饲养难度较大。     

Postprandial morphological response of the intestinal epithelium of the Burmese python (Python molurus)

The postprandial morphological changes of the intestinal epithelium of Burmese pythons were examined using fasting pythons and at eight time points after feeding. In fasting pythons, tightly packed enterocytes possess very short microvilli and are arranged in a pseudostratified fashion. Enterocyte width increases by 23% within 24 h postfeeding, inducing significant increases in villus length and intestinal mass. By 6 days postfeeding, enterocyte volume had peaked, following as much as an 80% increase. Contributing to enterocyte hypertrophy is the cellular accumulation of lipid droplets at the tips and edges of the villi of the proximal and middle small intestine, but which were absent in the distal small intestine. At 3 days postfeeding, conventional and environmental scanning electron microscopy revealed cracks and lipid extrusion along the narrow edges of the villi and at the villus tips. Transmission electron microscopy demonstrated the rapid postprandial lengthening of enterocyte microvilli, increasing 4.8-fold in length within 24 h, and the maintaining of that length through digestion. Beginning at 24 h postfeeding, spherical particles were found embedded apically within enterocytes of the proximal and middle small intestine. These particles possessed an annular-like construction and were stained with the calcium-stain Alizarine red S suggesting that they were bone in origin. Following the completion of digestion, many of the postprandial responses were reversed, as observed by the atrophy of enterocytes, the shortening of villi, and the retraction of the microvilli. Further exploration of the python intestine will reveal the underlying mechanisms of these trophic responses and the origin and fate of the engulfed particles.     

Thermoregulation in pythons. IV Thermoregulation in the Papuan-New Guinean pythons within the genera Python, Liasis and Chondropython

The thermal preference, voluntary maxima, and thermal control were studied in the Papuan-New Guinean pythons Python spilotus, P. boeleni, Liasis albertisii, L. papuanus, L. boa, and Chondropython viridis. Preferred temperature data were similar for all species and mean preferred head temperatures were lower than those for the body. In general preferred and voluntary means were higher for the Papuan-New Guinean pythons than those found for Australian species and likely reflect adaptation to hotter tropical environments.     

Effect of feeding on circulating micronutrient concentrations in the Burmese python (Python molurus).

Burmese pythons (Python molurus) regulate digestive performance and metabolism with the ingestion of each meal. To explore the python's postprandial responses, we monitored the concentrations of blood micronutrients and homocysteine during fasting and for 15 days after feeding. Plasma folate concentrations peaked with a 270% increase over fasting levels 3 days after feeding, whereas plasma B-12 peaked with a 66% increase within 1 day. Erythrocyte folate concentrations were highest 15 days after feeding with a 44% increase. The major plasma folate was 5-methyltetrahydrofolate during fasting and was non-5-methyltetrahydrofolate during digestion, whereas erythrocytes contained polyglutamyl forms of non-5-methyltetrahydrofolate. Plasma homocysteine concentrations peaked with a 56% increase 3 days after feeding, and were markedly greater than those of mammals. Plasma zinc and copper did not change significantly. Plasma zinc concentrations were 20 times greater than plasma copper and approximately 30 times higher than those of mammals. Pythons showed a significant postprandial decline of 25% in hematocrit. Plasma pyridoxal 5'-phosphate (coenzyme form of vitamin B-6) was not detected probably due to its tight protein binding. Most micronutrient concentrations appear to plateau 3 days after feeding, suggesting that pythons have relatively rapid homeostasis of micronutrients despite the ingestion of large meals.     

Effects of temperature on the metabolic response to feeding in Python molurus

As ectothermic vertebrates, reptiles undergo diurnal and seasonal changes in body temperature, which affect many biological functions. In conjunction with a general review regarding the effects of temperature on digestion in reptiles, we describe the effects of various temperatures (20-35 degrees C) on the metabolic response to digestion in the Burmese python (Python molurus). The snakes were fed mice amounting to 20% of their body weight and gas exchange (oxygen uptake and CO(2) production) were measured until digestion had ended and gas exchange returned to fasting levels. Elevated temperature was associated with a faster and larger metabolic increase after ingestion, and the time required to return to fasting levels was markedly longer at low temperature. The factorial increase between fasting oxygen consumption (VO(2)) and maximal VO(2) during digestion was, however, similar at all temperatures studied. Furthermore, the integrated SDA response was not affected by temperature suggesting the costs associated with digestion are temperature-independent. Other studies on reptiles show that digestive efficiency is only marginally affected by temperature and we conclude that selection of higher body temperatures during digestion (postprandial thermophilic response) primarily reduces the time required for digestion.     

Metabolism,temperature relations,maternal behavior,and reproductive energetics in the ball python (Python regius)

Summary Thermogenic incubation has been documented in two large species of pythons, but the phenomenon has not been studied in small species with concomitantly large heat transfer coefficients. We describe behavior, metabolic rates, mass changes, and temperature relations for adult ball pythons (Python regius), the smallest member of the genus, during the reproductive cycle. Egg and hatchling metabolism and hatchling growth rates were also examined.Rates of oxygen consumption ( ) of both gravid and non-gravid snakes showed typical ectothermic responses to changing ambient temperature (T _a). TheQ ₁₀ forT _a's of 20–35°C was 2.2–2.3. The of gravid females was significantly greater than that of non-gravid snakes at allT _a. Maximum oxygen consumption ( max) during forced exercise was about 12 times resting atT _a=30°C.Eggs (5–6 per female) were laid in April. Total clutch mass was approximately 32% of the females' pre-oviposition mass. After oviposition, mother snakes coiled tightly around their clutches and remained in close attendance until the eggs hatched in June. Sudden decreases inT _a elicited abrupt but transient 2- to 4-fold increases in the of incubating females. Similar responses were not observed in non-incubating snakes. The steady-state of incubating females was independent ofT _a. In no case was body temperature (T _b) elevated more than a few tenths of a degree aboveT _a in steady-state conditions.The of developing eggs increased sigmoidally through the 58–70 day incubation period. Total oxygen consumption during incubation atT _a=29.2°C was about 3.61 per egg. Young snakes quadrupled their mass during their first year of growth.Compared to larger python species which are endothermic during incubation, ball pythons have similar aerobic scopes and higher mass-specific max. However, effective endothermy in ball pythons is precluded by high thermal conductance and limited energy stores.     

Modeling baroreflex regulation of heart rate during orthostatic stress

During orthostatic stress, arterial and cardiopulmonary baroreflexes play a key role in maintaining arterial pressure by regulating heart rate. This study presents a mathematical model that can predict the dynamics of heart rate regulation in response to postural change from sitting to standing. The model uses blood pressure measured in the finger as an input to model heart rate dynamics in response to changes in baroreceptor nerve firing rate, sympathetic and parasympathetic responses, vestibulo-sympathetic reflex, and concentrations of norepinephrine and acetylcholine. We formulate an inverse least squares problem for parameter estimation and successfully demonstrate that our mathematical model can accurately predict heart rate dynamics observed in data obtained from healthy young, healthy elderly, and hypertensive elderly subjects. One of our key findings indicates that, to successfully validate our model against clinical data, it is necessary to include the vestibulo-sympathetic reflex. Furthermore, our model reveals that the transfer between the nerve firing and blood pressure is nonlinear and follows a hysteresis curve. In healthy young people, the hysteresis loop is wide, whereas, in healthy and hypertensive elderly people, the hysteresis loop shifts to higher blood pressure values, and its area is diminished. Finally, for hypertensive elderly people, the hysteresis loop is generally not closed, indicating that, during postural change from sitting to standing, baroreflex modulation does not return to steady state during the first minute of standing.     

Cardiovascular actions of python bradykinin and substance P in the anesthetized python, Python regius

The cardiovascular actions of python bradykinin (BK) and substance P (SP) have been investigated in the anesthetized ball python, Python regius. Bolus intra-arterial injections of python BK (0.03-3 nmol/kg) produced concentration-dependent increases in arterial blood pressure, heart rate (HR), and cardiac output concomitant with small decreases in systemic resistance and stroke volume. Intra-arterial injection of 3 nmol/kg python BK produced a tenfold increase in circulating concentration of norepinephrine, but epinephrine levels did not change. BK-induced tachycardia was attenuated (>90%) by the beta-adrenergic receptor antagonist sotalol, and the hypertensive response was attenuated (>70%) by the alpha-adrenergic receptor antagonist prazosin, indicating that effects of python BK are mediated at least in part by activation of the extensive network of adrenergic neurons present in vascular tissues. Bolus intra-arterial injections of python SP in the range 0. 01-30 pmol/kg produced concentration-dependent decreases in arterial blood pressure and systemic peripheral resistance concomitant with increases in cardiac output and stroke volume but with only minor effects on HR. The data suggest that kinins play a physiologically important role in cardiovascular regulation in the python.     

A new species of Hepatozoon (Apicomplexa: Adeleorina) from Python regius (Serpentes: Pythonidae) and its experimental transmission by a mosquito vector

Hepatozoon ayorgbor n. sp. is described from specimens of Python regius imported from Ghana. Gametocytes were found in the peripheral blood of 43 of 55 snakes examined. Localization of gametocytes was mainly inside the erythrocytes; free gametocytes were found in 15 (34.9%) positive specimens. Infections of laboratory-reared Culex quinquefasciatus feeding on infected snakes, as well as experimental infection of juvenile Python regius by ingestion of infected mosquitoes, were performed to complete the life cycle. Similarly, transmission to different snake species (Boa constrictor and Lamprophis fuliginosus) and lizards (Lepidodactylus lugubris) was performed to assess the host specificity. Isolates were compared with Hepatozoon species from sub-Saharan reptiles and described as a new species based on the morphology, phylogenetic analysis, and a complete life cycle.     

Discrimination of integumentary prey chemicals and strike-induced chemosensory searching in the ball python,Python regius

Experimental tests show that the ball python (Python regius) has the ability to discriminate prey chemicals from control substances by tongue-flicking and exhibits a poststrike elevation in tongue-flicking rate (PETF). Prey chemical discrimination was revealed by significantly higher number of tongue-flicks and tongue-flick attack score in response to integumental chemicals from mice than to cologne or distilled water and by a higher frequency of biting in response to prey than control chemicals. PETF was indicated by higher tongue-flicking rates after biting than in several control conditions. Concurrent movements of the body suggest the operation of strike-induced chemosensory searching (SICS). Ecological factors affecting responses to prey chemicals, including defensive behaviors and characteristics of foraging behavior related to reliance on different sensory modalities, are discussed. The presence of PETF and SICS in a henophidian snake and in scleroglossan lizards suggests that these behaviors are plesiomorphic in snakes.