Changes in the composition of the urine of yellow-vented bulbuls (Pycnonotus xanthopygos): the effects of ambient temperature, nitrogen, and water intake.

Uricotely (uric acid >50% of urinary nitrogen) in birds was once considered ubiquitous. However, Anna's hummingbirds (Calypte anna) have been shown to be an exception to this rule; under conditions of low ambient temperature (T(a)) and on a nitrogen-free diet, they increased their water intake and often became ammonotelic (ammonia >50% of urinary nitrogen). Our aim was to identify the effects of nitrogen intake, water intake, and T(a) on the ammonia excretion of yellow-vented bulbuls (Pycnonotus xanthopygos). We chose this predominantly frugivorous species because many of the characteristics of nectarivores that were used to explain increased ammonia excretion by C. anna are also characteristics of frugivorous birds. We assayed ureteral urine composition in eight yellow-vented bulbuls (P. xanthopygos), each randomly allocated a diet of 20% (0.6 M) sucrose solution supplemented by either 1.03 g/L or 7.23 g/L soy protein and held at a T(a) of either 28 degrees C or 10 degrees C. Food, and therefore water, intake rates varied with nitrogen intake but not with T(a). Food intake increased significantly with decreased nitrogen intake, while concentrations of all the excretory compounds in the urine (P<0.05) decreased; yet their proportions in the urine did not change significantly. The lower T(a) had no significant effect on food intake or on the concentration of uric acid. However, at 10 degrees C, the ammonia and urea concentrations increased (P<0.05), and this led to a significant increase in the proportion of ammonia in the urine. Our results demonstrate that, when bulbuls are exposed to low T(a), they are able to save energy by increasing the proportion of the ammonia in their urine.     

Sucrose hydrolysis does not limit food intake by Pallas's long-tongued bats

Nectarivorous bats include very dilute nectar in their natural diet, and recent work with Pallas's long-tongued bat Glossophaga soricina showed that sugar (energy) intake rate decreased at dilute sucrose solutions. However, chiropterophillous nectar is composed mainly of the hexoses glucose and fructose. Because bats fed hexose nectar would save the delay of hydrolyzing sucrose, we hypothesized that sugar intake rate should be higher on this diet than on sucrose nectar. We compared intake response in Pallas's long-tongued bats offered 1 : 1 glucose-fructose (hexose) and sucrose diets at 5%, 10%, 20%, 30%, and 40% (mass/volume) sugar solutions. We also tested the hypothesis that sucrose hydrolysis limits food intake in bats. Intake response was the same in bats fed both types of diet: sugar intake rate was lower in dilute solutions and then increased with sugar concentration. Similar intake responses in both diets indicate that sucrose hydrolysis alone does not limit food intake and support the idea that the burden of processing excess water in dilute solutions plays a major role.     

The bioenergetics of grass carp, Ctenopharyngodon idella (Val.): the influence of body weight, ration and dietary composition on nitrogenous excretion

Nitrogenous excretion by grass carp, Ctenopharyngodon idella (Val.), was measured in the form of ammonia and urea. Endogenous nitrogen excretion (ENE) was estimated as the daily rate of excretion by grass carp which had been starved for 2 days. ENE was scaled allometrically with body weight with weight exponents of 0.75 for ammonia, 0.63 for total nitrogen and 0.63 for the energy lost. The proportion of nitrogen attributable to urea was smaller than that attributable to ammonia and decreased from 25 to 12% as fish weight increased from 2 to over 10 g.
Linear relationships were found between daily rates of ammonia, total nitrogen and energy loss and daily rates of food intake. High carbohydrate and high lipid diets were not shown to have a protein-sparing action compared to a high protein diet. Differences in the amount of nitrogen excreted were explained by the differing nitrogen contents of the diets. Nitrogen budgets were erected and their implications discussed.     

Correlation of the plasma tyrosine to phenylalanine ratio with energy intake in self-selecting weanling rats

The relationship between changes in blood plasma amino acids and the quantity of protein and energy self-selected by the weanling rat, simultaneously offered two diets varying only in gluten (15 and 55%) concentration, was examined. Gluten and energy intakes were manipulated by additions of lysine, arginine or ammonia to gluten. In two experiments groups of ten weanling rats were fed the diets for a two week or four week period and food intake selection recorded. Blood samples were obtained between 0900–1100 hr at the end of the two week or four week period. Correlation coefficients of protein intake with the plasma TRP/NAA (Tyr+Phe+Leu+Val+Ile) ratios were ?0.97 and ?0.98, and of energy intake with TYR/PHE ratios were 0.77 and 0.70 in experiments 1 and 2, respectively. It is suggested that the plasma TRP/NAA and TYR/PHE ratios reflect the mechanisms regulating protein and energy intakes, respectively.     

Ammonia excretion increased and urea excretion decreased in urine of a new world nectarivorous bat with decreased nitrogen intake

We determined the effect of water and nitrogen intake on nitrogenous waste composition in the nectarivorous Pallas's long-tongued bat Glossophaga soricina (Phyllostomidae) to test the hypothesis that bats reduce excretion of urea nitrogen and increase the excretion of ammonia nitrogen as nitrogen intake decreases and water intake decreases. Because changes in urine nitrogen composition are expected only in animals whose natural diets are low in nitrogen and high in water content, we also measured maintenance nitrogen requirements (MNR). We hypothesized that, similar to other plant-eating vertebrates, nectarivorous bats have low MNR. Our nitrogen excretion hypothesis was partly proved correct. There was an increase in the proportion of N excreted as ammonia and a decrease in the proportion excreted as urea in low-nitrogen diets. The proportion of N excreted as ammonia and urea was independent of water intake. Most individuals were ureotelic (n = 28), and only a few were ureo-ammonotelic (n = 3) or ammonotelic (n = 2). According to our nitrogen requirement hypothesis, apparent MNR (60 mg kg(-0.75) d(-1)) and truly digestible MNR (54 mg N kg(-0.75) d(-1)) were low. A decrease in urea excretion in low-nitrogen diets may result from urea recycling from liver to the gut functioning as a nitrogen salvage system in nectarivorous bats. This mechanism probably contributes to the low MNR found in Pallas's long-tongued bats.     

乙醇浓度对圈养长舌果蝠摄食糖水量的影响

正在自然生境中,果蝠主要以果实或者花蜜为食,成熟果实或花蜜由于微生物发酵而含一定量的乙醇。乙醇在生态系统中是一种常见物质,它是生物厌氧代谢的终端产物之一(Van Waarde,1991)。一旦果实成熟,它们中的糖分就会被微生物或者自身发酵代谢成乙醇(Battcock and Azam-Ali,1998;Dudley,2000,2004)。果实中的乙醇浓度主要是由这些以糖分为能源的发酵微生物的扩增状态来决定     

Physiological constraint to food ingestion in a new world nectarivorous bat

We investigated the intake response of the nectarivorous Pallas's long-tongued bat Glossophaga soricina to different nectar concentrations to test the hypothesis that bats show compensatory feeding. Bats were offered sucrose solutions between 146 and 1,168 mmol L(-1). Contrary to our expectations, long-tongued bats did not show compensatory feeding, suggesting that volumetric food intake is physiologically constrained. Energy intake was lower at the most dilute solutions (146-584 mmol L(-1)) and then remained relatively constant at more concentrated diets (876 and 1,168 mmol L(-1)). The shape of the observed intake response was very similar to the one predicted by a model dependent on intestinal morphology and in vitro sucrase activity. However, the model predicted higher volumetric food intake at the lower concentrations tested, which suggests that the intestines of the bats were not functioning to their full capacity. Rates of sucrose hydrolysis and water processing probably constrain food intake in long-tongued bats as diets get more dilute.     

Seasonal dietary niche contraction in coexisting Neotropical frugivorous bats (Stenodermatinae)

Tropical dry forests are characterized by punctuated seasonal precipitation patterns that drive primary production and the availability of fruits, seeds, flowers, and insects throughout the year. In environments in which the quantity and quality of food resources varies seasonally, consumers should adjust their foraging behavior to maximize energy intake while minimizing overlap with competitors during periods of low food availability. Here, we investigated how the diets of frugivorous bats in tropical dry forests of NW Mexico varied in response to seasonal availability and how this affected dietary overlap of morphologically similar species. We performed stable isotope analyses to understand temporal and interspecific patterns of overall isotopic niche breadth, trophic position, and niche overlap in the diet of six frugivorous species of closely related New World leaf-nosed bats (family Phyllostomidae, subfamily Stenodermatinae). We estimated seasonal changes in resource abundance in two complementary ways: (a) vegetative phenology based on long-term remote sensing data and (b) observational data on food availability from previously published insect and plant fruiting surveys. In all species, there was a consistent pattern of reduced isotopic niche breadth during periods of low food availability. However, patterns of niche overlap varied between morphologically similar species. Overall, results from our study and others suggest that seasonal food availability likely determines overall dietary niche breadth in Phyllostomidae and that despite morphological specialization, it is likely that other mechanisms, such as opportunistic foraging and spatiotemporal niche segregation, may play a role in maintaining coexistence rather than simply dietary displacement.     

Foraging behavior adjustments related to changes in nectar sugar concentration in phyllostomid bats

Nectar-feeding bats regulate their food ingestion in response to changes in sugar concentration as a way to achieve a constant energy intake. However, their digestive capability to assimilate sugars can limit their total energy intake, particularly when sugar concentration in nectar is low. Our experimental study evaluated the effect that changes in sugar concentration of nectar have on the foraging behavior of the nectar-feeding bats Glossophaga soricina and Leptonycteris yerbabuenae in captivity. We measured foraging behavior and food intake when bats fed at different concentrations of sucrose (5, 15, 25 and 35%wt/vol.). To compensate for low-energy intake, both bat species reduced their flight time, and increased feeding time when sugar concentration decreased. Our results suggest that nectar-feeding bats in nature confront two scenarios with complementary ecological effects: 1) bats feeding on dilute nectars (i.e. ≤15%wt/vol.) should increase the number of flowers visited per night enhancing pollination, and 2) bats feeding on concentrated nectars could spend more time flying, including long- and short-distance-flights increasing food patch exploration for use during subsequent nights, and thus enhancing plant gene flow. Further studies on foraging behavior of nectarivorous bats under natural conditions are necessary to corroborate these hypotheses.     

Sugar assimilation and digestive efficiency in Wahlberg's epauletted fruit bat (Epomophorus wahlbergi)

Fruit- and nectar-feeding bats have high energy demands because of the cost of flight, and sugar is a good fuel because it is easily digested and absorbed. This study investigated the digestive efficiency of different sugars at different concentrations in Wahlberg's epauletted fruit bat (Epomophorus wahlbergi). We predicted that the sugar type and concentration would affect the total amount of solution consumed, while the total energy gained and the apparent assimilation efficiency would be high, irrespective of sugar type or concentration. Equicaloric solutions of two sugar types, glucose and sucrose, at low (10%), medium (15%) and high (25%) concentrations were offered in separate trials to bats. Total amount of solution consumed, total energy gained from each solution, and apparent assimilation efficiency, were measured. Bats had higher total volumetric intake of glucose and sucrose at the low concentrations than at the higher concentrations. However, bats maintained similar total energy intake on the respective glucose and sucrose concentrations. Bats were found to have high assimilation efficiencies on both glucose and sucrose irrespective of concentration. As bats used both sugars efficiently to maximize and maintain energy gain, it is expected that they feed opportunistically on fruit in the wild depending on temporal and spatial availability to obtain their energy requirements. Furthermore, fruit with high sucrose or glucose content will be consumed.     

Thermoregulation in the Angolan free-tailed bat Mops condylurus: A small mammal that uses hot roosts.

The Angolan free-tailed bat (Mops condylurus) uses roosts that often exceed 40 degrees C, an ambient temperature (Ta) that is lethal to many microchiropterans. We measured the physiological responses of this species at Ta's from 15 degrees to 45 degrees C. Torpor was commonly employed during the day at the lower Ta, but the bats generally remained euthermic at night, with a mean body temperature (Tb) of 35.2 degrees C. Metabolic rate reflected the pattern of Tb, increasing with falling Ta at night but decreasing during the day. Metabolic rate and evaporative losses were lower in torpid than in euthermic bats. Body temperature increased at each Ta >35 degrees C and was 43 degrees C at Ta of 45 degrees C. At Ta of 40 degrees C bats increased dry thermal conductance and evaporative heat loss compared to lower Ta. At 45 degrees C dry thermal conductance was lower than at 40 degrees C and evaporative heat loss was 132% of metabolic heat production. At high Ta there was only a slight increase in metabolic rate despite the employment of evaporative cooling mechanisms and an increase in Tb. Collectively our results suggest that M. condylurus is well suited to tolerate high Ta, and this may enable it to exploit thermally challenging roost sites and to colonise habitats and exploit food sources where less stressful roosts are limiting.     

The intake responses of three species of leaf-nosed Neotropical bats

Flower-visiting bats encounter nectars that vary in both sugar composition and concentration. Because in the new world, the nectars of bat-pollinated flowers tend to be dominated by hexoses, we predicted that at equicaloric concentrations, bats would ingest higher volumes of hexoses than sucrose-containing nectars. We investigated the intake response of three species of Neotropical bats, Leptonycteris curasoae, Glossophaga soricina and Artibeus jamaicensis, to sugar solutions of varying concentrations (292, 438, 584, 730, 876, and 1,022 mmol L⁻¹) consisting of either sucrose or 1:1 mixtures of glucose and fructose solutions. Bats did not show differences in their intake response to sucrose and 1:1 glucose–fructose solutions, indicating that digestion and absorption in bat intestines are designed under the principle of symmorphosis, in which no step is more limiting than the other. Our results also suggest that, on the basis of energy intake, bats should not prefer hexoses over sucrose. We used a mathematical model that uses the rate of sucrose hydrolysis measured in vitro and the small intestinal volume of bats to predict the rate of nectar intake as a function of sugar concentration. The model was a good predictor of the intake responses of L. curasoae and G. soricina, but not of A. jamaicensis.     

Changes in nectar concentration: how quickly do whitebellied sunbirds (Cinnyris talatala) adjust feeding patterns and food intake?

Nectarivorous birds encounter varying nectar concentrations while foraging on different food plants and must adjust their consumption to maintain constant energy intake. We determined how rapidly captive whitebellied sunbirds (Cinnyris talatala) adjust their volumetric intake and feeding patterns after changes in diet concentration. On four consecutive days, birds were fed sucrose diets alternating between a standard diet of 16% w/w and test diets of 2.5, 8.5, 16 or 30% w/w, respectively, for 1.5 h periods. Feeding events were recorded with an infrared photo-detection system and food intake and body mass were monitored continuously by electronic balances interfaced to a computer. Generally, birds demonstrated a measurable increase in feeding frequency and food intake within 10 min after a decrease in sucrose concentration. However, individuals responded differently to the most dilute diet (2.5%): while most increased their food intake, others stopped feeding for a short while, appearing to dislike this diet. Furthermore, the number and duration of feeding events increased in the first 5 min after the switch from 2.5% back to 16%, as the birds attempted to compensate for previous reduced sugar intake. Daily sugar intake was lower when birds alternated between 2.5 and 16% diets than on other test days, but birds were able to maintain body mass, presumably through behavioural adjustments.     

Effects of ethanol on food consumption and skin temperature in the Egyptian fruit bat (Rousettus aegyptiacus)

Since mammalian frugivores generally choose to eat ripe fruit in which ethanol concentration ([EtOH]) increases as the fruit ripens, we asked whether ethanol acts as an appetitive stimulant in the Egyptian fruit bat, Rousettus aegyptiacus, and also studied the effects of ethanol on their skin temperature (T(s)). We hypothesized that the responses of fruit bats to dietary ethanol are concentration dependent and tested the predictions that the bats' response is positive, i.e., they eat more when [EtOH] in the food is in the range found in naturally ripe fruit, while it negatively affects them at higher concentrations. We also tested the prediction that in winter, even when availability of fruit is low and thermoregulatory costs are high, ingestion of ethanol by fruit bats is low because assimilated ethanol reduces shivering thermogenesis and peripheral vasodilation; these, alone or together, are detrimental to the maintenance of body temperature (T(b)). In summer, captive bats offered food containing 0.1% ethanol significantly increased consumption over food with no ethanol; they did not change consumption when food contained 0.01, 0.3, or 0.5% ethanol; but significantly decreased consumption at higher levels of ethanol [EtOH], i.e., 1 and 2%. In winter, captive bats ate significantly less when their food contained 0.1% ethanol than when it contained 0, 0.3, or 0.5%. During summer, freshly caught bats ate significantly more ethanol-containing food than freshly caught bats in winter. Skin temperature (T(s)) in Egyptian fruit bats decreased significantly at an ambient temperature (T(a)) of 12 °C (winter conditions) after gavage with liquid food containing 1% ethanol. The effect was clearly temperature-dependent, since ethanol did not have the same effect on bats gavaged with food containing 1% or no ethanol at a T(a) of 25 °C (summer conditions). In conclusion, ethanol may act as an appetitive stimulant for Egyptian fruit bats at low concentrations, but only in summer. Bats are deterred by food containing [EtOH] corresponding to that in overripe, unpalatable fruit (1 and 2%). Furthermore, during winter, Egyptian fruit bats are deterred by ethanol-rich fruit, possibly due to the potential thermoregulatory consequences of ethanol consumption.     

A physiological perspective on nectar-feeding adaptation in phyllostomid bats

Nectar-feeding animals increase their food intake when nectar sugar concentration decreases. However, some species present physiological constraints that limit their energy intake when nectar is diluted. We hypothesized that gut capacities of bats affect the ability of these animals to acquire and store energy, modifying how they use food resources in the field. We measured the food intake and changes in body mass of the members of an assemblage of nectar-feeding bats (Choeronycteris mexicana, Leptonycteris yerbabuenae, and Glossophaga soricina) feeding on sucrose solutions of different concentrations (146, 292, 438, 584, 730, 876, and 1,022 mmol L(-1)). The three bat species presented differences in their food intake and their capacity to store energy. While C. mexicana was able to maintain a constant energy intake at all concentrations tested, G. soricina and L. yerbabuenae decreased their sugar/energy intake at the lowest sugar concentrations. Choeronycteris mexicana also increased body mass independent of sugar concentration, while G. soricina and L. yerbabuenae did not. On the basis of our results, we generated a model relating gut capacities and the use of food resources in the field. Our model's predictions and field data support the idea that digestive traits affect the way these animals use the food resources present in their environment.     

The possible roles of ethanol in the relationship between plants and frugivores: first experiments with egyptian fruit bats

In this paper we discuss how yeast, fungi ubiquitously presentin sugar-rich fruit, can influence the interaction between frugivoresand fleshy-fruited plants via ethanol. We suggest that plants,the seeds of which are mostly dispersed by vertebrates, exploitthe ethanol from alcoholic fermentation by yeast in their seeddispersal strategy. Moderate consumption of ethanol, i.e., atconcentrations close to those in naturally ripening fruit, byfrugivores may have beneficial short- and long-term effectsfor these potential dispersers, whereas consumption of largerquantities may have negative short- and long-term effects. Ethanolvapor emanating from palatable fruit may act as an odor cue,guiding bats and other frugivores to the fruit, and aiding themto assess its quality. In addition, we suggest that ingestedethanol may be an appetitive stimulant. We also evaluate thepossibility that ethanol within fruit may be used as a sourceof energy by frugivorous vertebrates. Our preliminary data indicatethat Egyptian fruit bats (Rousettus aegyptiacus) can use theodor of ethanol to assess food suitability, but also that itmay not serve as an attractant over short distances (i.e., <1m). Instead, ethanol is avoided at concentrations greater than1%, a value which might typically characterize overripe andotherwise unpalatable fruit. Our initial results further indicatethat Egyptian fruit bats significantly decrease their food consumptionif it contains 1 or 2% ethanol. Overall, ethanol may play diverseroles in the nutritional ecology and behavior of fruit-eatingbats, and in the interaction between frugivores and plants,in general.     

Homeostatic regulation of protein intake: in search of a mechanism

Free-living organisms must procure adequate nutrition by negotiating an environment in which both the quality and quantity of food vary markedly. Recent decades have seen marked progress in our understanding of neural regulation of feeding behavior. However, this progress has occurred largely in the context of energy intake, despite the fact that food intake is influenced by more than just the energy content of the diet. A large number of behavioral studies indicate that both the quantity and quality of dietary protein can markedly influence food intake. High-protein diets tend to reduce intake, low-protein diets tend to increase intake, and rodent models seem to self-select between diets in order to meet protein requirements and avoid diets that are imbalanced in amino acids. Recent work suggests that the amino acid leucine regulates food intake by altering mTOR and AMPK signaling in the hypothalamus, while activation of GCN2 within the anterior piriform cortex contributes to the detection and avoidance of amino acid-imbalanced diets. This review focuses on the role that these and other signaling systems may play in mediating the homeostatic regulation of protein balance, and in doing so, highlights our lack of knowledge regarding the physiological and neurobiological mechanisms that might underpin such a regulatory phenomenon.     

The sweet side of life: nectar sugar type and concentration preference in Wahlberg's epauletted fruit bat

Whether nectarivores or frugivores place selective pressure on the plants they feed on, in terms of nectar or fruit traits, is much debated. Globally sugar preferences, concentration preference and digestive ability of avian nectarivores have been extensively researched. In contrast, relatively little is known about mammalian nectarivores or frugivores in terms of these, particularly Old World species. Consequently effect of sugar type and concentration on food preference in Wahlberg's epauletted fruit bat Epomophorus wahlbergi was investigated. Pair-wise choice tests were conducted using equicaloric hexose and sucrose solutions at five different concentrations (5%-25%). It was expected that they would prefer hexose sugars as these are dominant in available indigenous fruits. However, bats preferred hexoses only when offered dilute (5%) concentrations. From 10% to 25% they showed a decrease in volume intake. Their body mass was generally higher and similar after feeding during the night with the exception of 5% concentration where the mean body mass decreased. When E. wahlbergi were offered a range of sucrose or hexose solutions (10%-25%) respectively, they showed no concentration preference in terms of total volume consumed, nor energy intake. These findings suggest that these fruit bats do not appear to act as a selective pressure on sugar composition in Old World fruit. In fruit bats with high energy requirements, dietary flexibility may be an advantage when faced with seasonal and unpredictable fruit availability.     

Does gut function limit hummingbird food intake?

Many nectar-feeding bird species decrease food intake when sugar concentration in food is increased. This feeding response can be explained by two alternative hypotheses: compensatory feeding and physiological constraint. The compensatory feeding hypothesis predicts that if birds vary intake to maintain a constant energy intake to match energy expenditures, then they should increase intake when expenditures are increased. Broad-tailed hummingbirds were presented with sucrose solutions at four concentrations (292, 584, 876, and 1,168 mmol L(-1)) and exposed to two environmental temperatures (10 degrees and 22 degrees C). Birds decreased volumetric food intake in response to sugar concentration. However, when they were exposed to a relatively sudden drop in environmental temperature and, hence, to an acute increase in thermoregulatory energy expenditures, they did not increase their rate of energy consumption and lost mass. These results support the existence of a physiological constraint on feeding intake. A simple chemical reactor model based on intestinal morphology and in vitro measurements of sucrose hydrolysis predicted observed intake rates closely. This model suggests that intestinal sucrose hydrolysis rates were near maximal and, thus, may have imposed limits to sugar assimilation. Although sugar assimilation was high (95%), the proportions of excreted sucrose, glucose, and fructose found in excreta differed significantly. The monosaccharides glucose and fructose were about eight and three times more abundant than sucrose, respectively. Broad-tailed hummingbirds are small high-altitude endotherms that face unpredictable weather and the energetic expense of premigratory fattening. Digestive processes have the potential to impose severe challenges to their energy budgets.     

An investigation of factors controlling food intake in Arctic charr, Salvelinus alpinus L.

Digestibility values were determined for diets of varying protein content fed to Arctic charr. The use of these values, in conjunction with food intake data made possible the study of the effects of various parameters on the control of food consumption. Dietary energy content may be more important than protein content in regulating consumption. The regulation of food intake may be partially mediated through factors influencing gastric emptying coupled with mechanisms monitoring dietary metabolizable energy.