The effect of sugar solution type,sugar concentration and viscosity on the imbibition and energy intake rate of bumblebees

Nectar is an essential resource for bumblebees and many other flower-visiting insects. The main constituents of nectar are sugars, which vary in both composition and concentration between plant species. We assessed the influence of sugar concentration, sugar solution viscosity and sugar solution composition on the imbibition and energy intake rate of bumblebees, Bombus impatiens Cresson (Hymenoptera: Apidae). To do this, we measured their rate of solution intake for 49 different sugar solution treatments, which varied in both sugar composition and concentration. In general, the imbibition rates of bumblebees were found to increase with increasing sugar concentration, probably due to their preference for high sugar concentrations, up to a concentration of 27% (w/w), at which point solutions reached a threshold viscosity of approximately 1.5–1.6 mPa.s. Above this threshold, the increasing viscosity of the solutions physically inhibited the imbibition rates of bees, and imbibition rate began to decrease as the concentration increased. Nevertheless, bumblebee energy intake rate increased with increasing concentration up to about 42–56%. Although we found that sugar solution composition had an impact on both imbibition and energy intake rate, its effect was not as straightforward as that of sugar concentration and viscosity.     

Fluid intake rates in ants correlate with their feeding habits

This study investigates the techniques of nectar feeding in 11 different ant species, and quantitatively compares fluid intake rates over a wide range of nectar concentrations in four species that largely differ in their feeding habits. Ants were observed to employ two different techniques for liquid food intake, in which the glossa works either as a passive duct-like structure (sucking), or as an up- and downwards moving shovel (licking). The technique employed for collecting fluids at ad libitum food sources was observed to be species-specific and to correlate with the presence or absence of a well-developed crop in the species under scrutiny. Workers of ponerine ants licked fluid food during foraging and transported it as a droplet between their mandibles, whereas workers of species belonging to phylogenetically more advanced subfamilies, with a crop capable of storing liquids, sucked the fluid food, such as formicine ants of the genus Camponotus. In order to evaluate the performance of fluid collection during foraging, intake rates for sucrose solutions of different concentrations were measured in four ant species that differ in their foraging ecology. Scaling functions between fluid intake rates and ant size were first established for the polymorphic species, so as to compare ants of different size across species. Results showed that fluid intake rate depended, as expected and previously reported in the literature, on sugar concentration and the associated fluid viscosity. It also depended on both the species-specific feeding technique and the extent of specialization on foraging on liquid food. For similarly-sized ants, workers of two nectar-feeding ant species, Camponotus rufipes (Formicinae) and Pachycondyla villosa (Ponerinae), collected fluids with the highest intake rates, while workers of the leaf-cutting ant Atta sexdens (Myrmicinae) and a predatory ant from the Rhytidoponera impressa-complex (Ponerinae) did so with the lowest rate. Calculating the energy intake rates in mg sucrose per unit time, licking was shown to be a more advantageous technique at higher sugar concentrations than sucking, whereas sucking provided a higher energy intake rate at lower sugar concentrations.     

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.     

Sugar preferences, absorption efficiency and water influx in a Neotropical nectarivorous passerine, the Bananaquit (Coereba flaveola)

Nectarivory has evolved repeatedly in a number of unrelated bird taxa throughout the world and nectar feeding birds, regardless of their taxonomic affiliation, display convergent foraging and food processing adaptations that allow them to subsist on weak sugar solutions. However, phylogeny influences sugar type preferences of nectarivores. We investigated sugar preferences, assimilation efficiency and water flux in a Neotropical honeycreeper, the Bananaquit (Coereba flaveola; Coerebidae), a member of a radiation of tanagers and finches. Bananaquits showed no preference for nearly equicaloric (25% w/v) sucrose, glucose, fructose or glucose-fructose mixtures in pair-wise choice tests. In agreement with this lack of preference, they were equally efficient at absorbing sucrose and both hexoses. Apparent assimilation efficiency of these sugars was around 97.5%. In pair-wise tests, Bananaquits displayed a strong preference for the most concentrated sucrose solution when the lowest concentration ranged from 276 to 522 mM. Between 522 and 1120 mM sucrose solution concentrations, Bananaquits were able to adjust their volumetric food intake in order to maintain a constant energy intake. At solution concentration of 276 mM, birds could not maintain their rate of energy intake by increasing food consumption enough. We consider that at low sugar concentrations, Bananaquits faced a physiological constraint; they were unable to process food at a fast enough rate to meet their energy needs. We also explored the possibility that dilute nectars might be essential to sustain high water needs of Bananaquits by allowing them to control osmolarity of the food. Between 276 and 1120 mM sucrose solution concentrations, average amount of free water drunk by Bananaquits was independent of food concentration. They drank very little supplementary water and did not effectively dilute concentrated nectars. The evidence suggests that water bulk of dilute nectars is a burden to Bananaquits.     

Metabolism of glucose and cellobiose by cellulolytic mesophilic Clostridium sp. strain H10.

The metabolism of strain H10, a cellulolytic mesophilic Clostridium sp., was studied on glucose and cellobiose as energy and carbon sources. The main products of fermentation of both sugars were acetate, lactate, and ethanol. At low sugar levels, molar growth yields were better for cellobiose than for glucose. In both cases, an inhibition of growth was observed between 1 and 2 g/liter and a total inhibition after the latter concentration. Inhibition was not the result of low pH due to acid formation; growth under static pH conditions was limited in the same way. On the other hand, acetate and lactate had no inhibitory effect when added at concentrations equal to the final titers. Concomitant with the inhibition of growth was a change in metabolic pathways for sugar concentrations between 1 and 2 g/liter, i.e., the production of lactate was higher. After complete inhibition of growth, an accumulation of carbohydrates which were neither glucose nor cellobiose was observed.     

Sugar preferences of some southern African nectarivorous birds

Three southern African nectarivorous passerine birds, Gurney's Sugarbird Promerops gur-neyi , the Malachite Sunbird Nectarinia famosa and the Black Sunbird Nectarinia amethystina , were tested to determine their hexose and sucrose preferences. All three species preferred sucrose when offered a choice of 0.25 M solutions of glucose, fructose and sucrose. However, when the concentrations were increased to 0.73 M, the three species showed no preference for any of the three sugars. The choice made at low concentrations (equivalent to the lower limit of the range of nectar concentrations of preferred nectar-producing plants) may reflect preference for the sugar with the highest energy reward. We also examined the proposition that birds offered a choice of different concentrations of one sugar would show ranked preferences and maximize their rate of energy return by selectivity. In contrast to expectations, Gurney's Sugarbird and the Malachite Sunbird showed no preference for the highest concentrations. We suggest that dietary choices in these species indicate the birds had either reached a limit where they had sufficient energy intake or were affected by post-ingestion constraints.     

Management of protein intake in the fruit fly Anastrepha fraterculus

This work tested if carbohydrates and proteins ingestion is regulated in the South American fruit fly, Anastrepha fraterculus, to optimize survival and reproduction. Adult food treatments were established by providing sugar and hydrolyzed yeast in various combinations either alone or mixed at a standard 3:1 ratio (sugar:hydrolyzed yeast). Individual food consumption was assessed and related to survival patterns. The effects of adult feeding on fecundity and fertility patterns were investigated in groups of flies. Sugar consumption was the lowest in the treatment where it was provided with hydrolyzed yeast at a fixed 3:1 ratio. Consumption of hydrolyzed yeast did not differ between this treatment and the one in which this solution was complemented with one solution of sugar. It seems that a mixture of sugar and hydrolyzed yeast at a fixed ratio of 3:1, respectively, restricts extra ingestion of sugar; most probably because of negative response of the fly to overconsumption of protein. Survival was affected by the treatments, being lower in those cases where protein was at the fixed ratio. Group experiments revealed that protein restriction expanded longevity and decreased egg production. In contrast, egg production was enhanced when flies were kept continuously with a mixture of yeast and sugar plus an extra source of sugar, and this was not in detriment of survival. Our results suggest that fixed sugar-protein ratios in which protein is in excess affects fitness components such as longevity and reproduction. These findings are discussed from a theoretical and applied perspective in the context of pest control by means of the sterile insect technique.     

Activities of plasma membrane-bound enzymes isolated from roots of spruce (Picea abies) grown in the presence of aluminium

In batch cultures of Petunia hybrida cv. Rosy Morn Fertile. one respiratory peak is usually observed shortly after subculturing. However, two types of peak respiration could be distinguished, one connected with the dilution process and one with sugar addition at low biomass concentrations. The dilution peak was observed when cells were diluted in medium without sugar, in the presence or absence of mannitol. The sugar peak occurred only after previous dilution of the cells and not when sugar is added at high biomass concentrations Apparently the existence of a dilute suspension is a prerequisite for the induction of the peak. The presence of sugar is not a prerequisite for the increased respiratory activity but it is necessary lor growth: however, growth is possible without the increase in respiration, as was shown by the addition of sugar to a culture with a high biomass concentration. The peak caused by dilution either in the presence or absence of sugar showed no significant differences in height. The height of the peak caused by sugar addition to a previously diluted cell suspension was correlated with the sugar concentration. The respiratory peak disappeared long before the end of the growth period; this decline of the respiratory rates was not connected to sugar or oxygen limitation. In a continuous culture of Petunia hybrida growing at low biomass concentration, the respiration was always at the high level as observed during the peak of batch culture. Growing at lower biomass concentrations might be more expensive for plant cell suspensions.     

Sugar preferences and digestive efficiency in an opportunistic avian nectarivore, the Dark-capped Bulbul Pycnonotus tricolor

It has recently been recognized that flowers pollinated by generalist opportunistic nectarivores tend to have different nectar properties to those pollinated by specialist nectarivores (including both hummingbirds and specialist passerines). While renewed interest in specialist avian nectarivore sugar preferences and digestive physiology has helped explain the concentrated sucrose-dominated nectar of plants they feed on, there has been little progress in understanding why generalist or occasional nectar-feeding birds tend to be associated with flowers that have dilute hexose-dominated nectar. We examined sugar preferences and assimilation efficiencies over a range of concentrations, and concentration preferences, in Dark-capped Bulbuls Pycnonotus tricolor, one of the more common occasional avian nectarivores in southern Africa. Dark-capped Bulbuls showed significant preference for hexose sugar solutions, irrespective of concentration, when given a choice between hexose and sucrose solutions in equicaloric pair-wise choice tests conducted at five different concentrations (5–25%). This contrasts with results from specialist nectarivore groups which generally show a significant concentration-dependant switch in preference from hexose at low concentrations to sucrose at high concentrations for equicaloric solutions. In addition, Dark-capped Bulbuls showed an unusual lack of preference for solutions of higher sugar concentration when simultaneously offered four solutions varying in concentration from 10 to 25%. Dark-capped bulbuls also showed a unique effect of concentration on sugar assimilation efficiency, assimilating relatively more energy on 5% diets than on 25% diets. Although able to assimilate sucrose effectively, assimilation rates of hexose sugars were marginally higher. These results shed new light on pollination systems involving occasional nectarivores and, in particular, help to explain the prevalence of low concentration hexose-dominated nectars in flowers pollinated by these birds.     

Increasing fermentation efficiency at high sugar concentrations by supplementing an additional source of nitrogen during the exponential phase of the tequila fermentation process

In the tequila industry, fermentation is traditionally achieved at sugar concentrations ranging from 50 to 100 g x L(-1). In this work, the behaviour of the Saccharomyces cerevisiae yeast (isolated from the juices of the Agave tequilana Weber blue variety) during the agave juice fermentation is compared at different sugar concentrations to determine if it is feasible for the industry to run fermentation at higher sugar concentrations. Fermentation efficiency is shown to be higher (above 90%) at a high concentration of initial sugar (170 g x L(-1)) when an additional source of nitrogen (a mixture of amino acids and ammonium sulphate, different than a grape must nitrogen composition) is added during the exponential growth phase.     

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.     

Selective choice of sucrose solution concentration by the hovering hawk mothMacroglossum stellatarum

To determine the preference of the hovering hawk mothMacroglossum stellatarum for different sugar concentrations, the foraging behavior of adults were analyzed under laboratory conditions. Six sucrose concentrations (range, 10–60%, w/w) were simultaneously offered in six artificial ab libitum feeders. The number of feeding bouts and the duration of each visit were automatically recorded and stored in a computer. Results showed that the frequency of visits to the feeders did not vary among the different solutions offered, but the gathered volume by the group attained a maximum at between 20 and 50% (w/w). Moths invested more time in front of the feeder with the more concentrated sugar solutions. It was assumed that factors different from maximizing energy intake, such as water balance and viscosity of concentrated nectars, have to be considered in order to understand the observed patterns of nectar choice.     

Conformational characteristics of the RNA subunit ApA from energy minimization studies

In an attempt to better our understanding of the conformational stabilities in RNAs, an intensive theoraticl study has been carried out on one of its dimeric subunits, ApA, using an improved set of atom-atom interaction energy parameters and an improved version of energy-minimization technique. The C(3′)0endo and the C(2′)-endo sugar ApA units were sperately considered and 38 probable conformations have been analyzed in each case. The total potential energy, comprising nonbonded, electrostatic, and torsional contributions, was minimized by varying all seven relevant dihedral angles simumtaneously. The result reveal that 17 conformations in the case of C(3′)-endo sugar ApA and 7 confomations in the case of C(2′)-endo sugar ApA unit, the lowest energy conformation corresponds to a nonhelical structure and the A-RNA and the Watson-Crick-yype conformations lie at energy levels of about 0.5 and 1.0 Kcal/mo., respectively, above the lowest energy found. For ApA with the lops of different types in the backbone and they all differ in energies by about 3.5 Kcal/mol with refrence to the lowest energy founs. It is noted that the order ofmprefrence of the base stacking is observed in the A-RNA and the Watson-Crick type conformers. The ApA unit with C(2′)-endo sugar is forced to assume phosphodiester conformations with large deviations fom the expected staggered conformations compared to the ApA unit with C(3′)-endo sugar. The result obtained for ApA are discussed with refrence to those previously obtained for the dApdA unit. Te theoretical predictions are compared with the experimental data on the tRNA^Phe crystal, as well as those on fibrous RNAs and RNa subunitlike crystal structures. This study brings out many important aspects of the conformational stability of ApA which have been missed by studies made by others on this system.     

Determinants of ligand binding affinity and cooperativity at the GLUT1 endofacial site

Cytochalasin B (CB) and forskolin (FSK) inhibit GLUT1-mediated sugar transport in red cells by binding at or close to the GLUT1 endofacial sugar binding site. Paradoxically, very low concentrations of each of these inhibitors produce a modest stimulation of sugar transport [ Cloherty, E. K., Levine, K. B., and Carruthers, A. ((2001)) The red blood cell glucose transporter presents multiple, nucleotide-sensitive sugar exit sites. Biochemistry 40 ((51)) 15549-15561]. This result is consistent with the hypothesis that the glucose transporter contains multiple, interacting, endofacial binding sites for CB and FSK. The present study tests this hypothesis directly and, by screening a library of cytochalasin and forskolin analogues, asks what structural features of endofacial site ligands determine binding site affinity and cooperativity. Like CB, FSK competitively inhibits exchange 3-O-methylglucose transport (sugar uptake in cells containing intracellular sugar) but noncompetitively inhibits sugar uptake into cells lacking sugar at 4 °C. This refutes the hypothesis that FSK binds at GLUT1 endofacial and exofacial sugar binding sites. Some forskolin derivatives and cytochalasins inhibit equilibrium [(3)H]-CB binding to red cell membranes depleted of peripheral proteins at 4 °C. Others produce a moderate stimulation of [(3)H]-CB binding when introduced at low concentrations but inhibit binding as their concentration is increased. Yet other analogues modestly stimulate [(3)H]-CB binding at all inhibitor concentrations applied. These findings are explained by a carrier that presents at least two interacting endofacial binding sites for CB or FSK. We discuss this result within the context of models for GLUT1-mediated sugar transport and GLUT1 quaternary structure, and we evaluate the major determinants of ligand binding affinity and cooperativity.     

Substrate conditions that influence the assays used for determining the beta-glucosidase activity of cellulolytic microorganisms

Culture filtrates from Trichoderma harzianum E58, T. reesei CL 847 and Penicillium sp. C 462 were assayed for beta-glucosidase activity using a range of substrates and sugar analysis methods. Although sugar analyses by the dinitrosalicylic acid (DNS) and Nelson-Somogyi methods gave a similar profile, when increasing concentrations of salicin were assayed, considerably higher values were obtained with the DNS assay. The salicin concentration used for the assay greatly influenced the final beta-glucosidase values with higher values obtained for T. harzianum E58 and T. reesei CL 847 at substrate concentrations of 1 mg/mL while optimum values for Penicillium sp. C 462 were obtained at substrate concentrations greater than 3 mg/mL. Low concentrations of salicin and p-nitro-phenyl-beta-D-glucopyranoside (PNPG) gave the same response as cellobiose. Cellobiose should be used at concentrations greater than 3.74 mg/mL to avoid substrate limitation of the beta-glucosidase assay.     

Kinetics of the enzymatic hydrolysis of cellulose

Enzymatic hydrolysis of cellulose for sugar production offers advantages of higher conversion, minimal by-product formation, low energy requirements, and mild operating conditions over other chemical conversions. The development of a kinetic model, based on observable, macroscopic properties of the overall system, is helpful in design and economic evaluation of processes for sugar conversion and ethanol production. A kinetic model is presented, incorporating enzyme adsorption, product inhibition, and considers a multiple enzyme and substrate system. This model was capable of simulating saccharification of a lignocellulosic material, rice straw, at high substrate (up to 333 g/L) and enzyme concentrations (up to 9.2 FPU/mL) that are common to proposed process designs.     

Thermoanaerobacter ethanolicus Growth and Product Yield from Elevated Levels of Xylose or Glucose in Continuous Cultures

The performance of Thermoanaerobacter ethanolicus was evaluated in continuous culture with media containing concentrations of xylose (8 to 20 g/liter) greater than those previously reported. The ethanol yield declined from to 0.42 to 0.29 g of ethanol per g of xylose consumed when input xylose was increased from 4 to 20 g/liter. Yields of both total C₂ and C₃ products from consumed xylose and of cell biomass from ATP produced declined as the input xylose concentration was increased, which was not the case when glucose was the substrate. This suggested that yeast extract functioned as a significant energy and carbon source for cells in fermentations of xylose but not of glucose. The feasibility of this interpretation was confirmed by (i) the calculation of the products theoretically obtainable from yeast extract and (ii) the observation of significant quantities of fermentation products in inoculated sugar-free media. Markedly different patterns of metabolism for the two sugar substrates were also evidenced by the cell yield for glucose being twice that of xylose at elevated sugar concentrations. It was noted that caution must be exerted when results obtained at low xylose concentrations are extrapolated to predict those which can be obtained at higher concentrations.     

The human erythrocyte sugar transporter presents two sugar import sites

The human erythrocyte sugar transporter presents sugar import (e2) and sugar export (e1) sites simultaneously. This study asks whether the sugar transporter exposes only one or multiple import sites. We approached this question by analysis of cytochalasin B binding to the human erythrocyte sugar export site in the presence of sugars that bind to the sugar import site. Extracellular maltose does not enter human erythrocytes. High concentrations of maltose (1-100 mM) inhibit cytochalasin B binding to human red cells. Low concentrations (25-500 microM) increase the level of erythrocyte cytochalasin B binding. Maltose modulation of cytochalasin B binding is mediated by altered affinity of sugar export sites for cytochalasin B. Similar results are obtained with other cell-impermeant inhibitors of sugar uptake. Extracellular D-glucose (a transported sugar) stimulates cytochalasin B binding at low D-glucose concentrations (10-250 microM), but this effect is lost at higher concentrations. Intracellular D-glucose inhibits cytochalasin B binding. Low concentrations of extracellular maltose and other nontransported inhibitors stimulate 3-O-methylglucose uptake in erythrocytes. Higher sugar concentrations (1-100 mM) inhibit transport. These data support the hypothesis that the erythrocyte sugar transporter presents two sugar import sites and at least one sugar export site. This conclusion is consistent with the proposed oligomeric structure of the sugar transporter, a complex of four GluT1 proteins in which each subunit presents a translocation pathway.     

Sugar preferences of nectar feeding birds – a comparison of experimental techniques

Experiments to determine sugar preferences of nectarivorous animals have been conducted using a wide variety of experimental procedures, all of which aim at ensuring that the solutions offered in choices are “equivalent”. Each method used historically has controlled for a particular variable, such as number of molecules in solution, weight of sugar in solution, or amount of energy in solution, depending on what question the researchers have tried to answer. Biologists interpreting these results in terms of bird sugar preference have seldom taken these differences into account. The consequences of using different experimental procedures for sugar preferences exhibited by a nectarivorous bird, the malachite sunbird Nectarinia famosa, were examined using paired sucrose and hexose sugar solutions made up to be either equimolar, equiweight or equicaloric. We found the effect of methodology on bird sugar preference to be quite distinct, especially at low concentrations, where malachite sunbirds showed either sucrose preference, no preference, or hexose preference, depending on the method used. This study highlights the need for researchers to consider methodology when interpreting, or comparing among, results from previous studies.     

Functional asymmetry of the human Na+/glucose transporter (hSGLT1) in bacterial membrane vesicles

The functional characteristics of the forward and reverse transport modes of the human Na(+)/glucose transporter (hSGLT1) were investigated using plasma membrane vesicles of E. coli expressing the recombinant transporter. Correctly and inverse-oriented vesicles were employed to measure the initial rates of methyl-alpha-D-glucose uptake, under zero-trans conditions, as a function of Na(+), sugar, and phlorizin concentrations and membrane potential. This approach enabled the analysis of the two faces of hSGLT1 in parallel, revealing the reversibility of Na(+)/sugar cotransport. While the key characteristics of secondary active sugar transport were maintained in both modes, namely, Na(+) and voltage dependence, the kinetic properties of the two sides indicated a functional asymmetry of the transporter. That is, the apparent affinity for sugar and driver cation Na(+) exhibited a difference of more than 1 order of magnitude between the two modes. Furthermore, the selectivity pattern of ligands and the interaction of the transporter with the competitive inhibitor phlorizin were different. Whereas the high-affinity substrates, D-glucose and D-galactose, inhibited uptake of radioactive sugar tracer at their physiological concentrations (10 mM) in the forward reaction, they were poor inhibitors even at high concentrations in the reverse transport mode. Taken together, these results confirm the successful employment of E. coli to express and characterize a human membrane protein (hSGLT1), elucidating the functional asymmetry of this cotransporter.