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Daily rhythms of lipogenesis in fat and lean white-throated sparrows Zonotrichia albicollis.

Lipogenesis was assayed at 6 times a day in the migratory white-throated sparrow Zonotrichia albicollis by measuring the incorporation of label in liver and body lipid 4 h after injection of [3H] acetate. There were daily rhythms of 3H incorporation by liver lipid both in fat birds in the vernal migratory condition and in lean birds in the summer photorefractory condition. Most of the liver lipid incorporation of 3H was restricted to the late afternoon and early evening in both the fat and lean birds. In addition, the total liver incorporation was similar for both groups. On the other hand, nearly twice as much 3H was incorporated in the body lipid of fat birds as in that of lean birds, and a daily rhythm was present only in fat sparrows. Injections of prolactin given simultaneously with [3H] acetate early during the day (at a time when daily injections of the hormone can cause severe losses in body fat stores) drastically reduces liver and body lipid incorporation of 3H. However, injections of prolactin during the afternoon (when daily injections of the hormone produce large increases in body fat stores) have no apparent influence on the incorporation of label. Our results indicate that a principal difference between lean and fat birds resides in a greater capacity in fat birds to transport triglycerides from the liver, in which they are produced, to the body storage depots. The daily rhythm of prolactin in some temporal relations may cause fattening by increasing the transport capacity or cause loss in fat stores in other temporal relations by directly inhibiting hepatic lipogenesis.

Acetates↗

Lower intestinal modification of ureteral urine in hydrated house sparrows.

The ureters of birds empty into the posterior portion of the lower intestine, thereby providing the possibility for modification of ureteral urine by this latter organ. We have used in vivo perfusion to measure the transport of Na+, K+, and water across the lower intestine (colon and coprodaeum) of anesthetized house sparrows (Passer domesticus). Na+ was reabsorbed from (Vmax = approximately 22 mu eq . cm-2 . h-1, Km = approximately 69 meq/l) and K+ was secreted (at variable rates) into all saline perfusion fluids. The osmotic permeability of the intestinal epithelium was 0.39 microliter . cm-2 . h-1 . mosM-1 in the mucosal-to-serosal direction and 0.43 microliter . cm-2 . h-1 . mosM-1 in the serosal-to-mucosal direction. At isosmotic perfusion, Na+-linked water transport occurred at a rate of 1.7 microliter/mu eq Na+. In hydrated house sparrows the composition of ureteral urine (osmolarity = 351 mosM, Na+ = 86.5 meq/l, K+ = 60.5 meq/l) was significantly modified by transport in the lower intestine (voided fluid osmolarity = 344 mosM, Na+ = 60 meq/l, K+ = 90 meq/l). Interspecific comparisons of lower intestinal resorptive surface area and transport parameters at the level of the tissue, organ, and whole animal reveal no consistent pattern of adaptation related to habitat.

Animals↗

Cholecystokinin octapeptide decreases food intake in white-crowned sparrows.

White-crowned sparrows maintained on short days (9:15-h light-dark cycle) were peripherally injected with 1.0, 4.0, and 16 micrograms/kg ip of cholecystokinin octapeptide (CCK-8). Meal size over the subsequent 30 min was significantly depressed in a dose-dependent fashion. Water intake was not affected. The anorexic effect caused by 4.0 micrograms/kg was attenuated by 100 micrograms/kg of the type-A CCK receptor antagonist MK-329 but not by 300 micrograms/kg of the type-B CCK receptor antagonist L 365,260, suggesting that CCK-induced suppression of food intake in this species is mediated by a CCK-A receptor. Administration of both CCK-A and CCK-B receptor antagonists alone resulted in no change in meal size. These experiments suggest that white-crowned sparrows, when weight stable, respond to CCK-8 in a manner comparable with several mammalian species.

Animals↗

NPY increases food intake in white-crowned sparrows: effect in short and long photoperiods.

To determine if altered sensitivity to neuropeptide Y (NPY) underlies premigratory fattening, white-crowned sparrows held on short day length (9:15-h light-dark) received injections into the third ventricle (ivt) of saline or several doses of NPY. An inverted-U function occurred with food intake increasing 30 and 60 min after doses of 1.0 and 2.0 micrograms NPY. When photostimulated (20:4-h light-dark), birds increased daily caloric intake and gained weight rapidly. Birds maintained on long day lengths significantly increased food intake after 0.25 and 0.5 micrograms of NPY and did not respond to higher doses. The effective dose range for NPY to increase food intake moved to the left, suggesting an increase in sensitivity to the peptide on long day lengths. In summary, white-crowned sparrows consume more food when administered NPY ivt and have increased sensitivity when photostimulated and gaining weight. Hence NPY may be a natural stimulator of food intake in this species.

Animals↗

Regulation of food intake by metabolic fuels in white-crowned sparrows.

Migratory birds rely on increased fat storage and fatty acid utilization to meet seasonal changes of energy expenditure and as a result increase food intake and fat stores before migration. To determine whether their feeding behavior is sensitive to carbohydrate and/or fatty acid utilization, white-crowned sparrows maintained on short daylength (9L15D) were injected intraperitoneally with 2-deoxy-D-glucose (2-DG) or 2,5-anhydro-D-mannitol (2,5-AM). Low doses of 2-DG (25 or 50 mg/kg) had no effect on food intake, and higher doses (100 or 300 mg/kg) significantly suppressed feeding after 1 and 2 h. No dose of 2-DG increased meal size. Similarly, low doses of 2,5-AM (25, 50, or 100 mg/kg) had no effect on food intake, and higher doses (300 and 600 mg/kg) significantly suppressed intake. These data suggest that decreased carbohydrate metabolism does not elicit feeding in this species. Importantly, these drugs, as well as insulin and glucagon, were demonstrated to increase plasma fatty acids as well as to decrease feeding. Injections of tributyrin (100, 300, 600, or 2,000 mg/kg i.p.) or glycerol (300, 450, and 600 mg/kg) also significantly suppressed 60-min and 120-min food intake dose dependently in these birds, and equimolar glucose (1,200 mg/kg) had no effect. We conclude that feeding by the white-crowned sparrow is unresponsive to manipulations of carbohydrate metabolism and is decreased after manipulations that increase plasma lipids.

Animals↗

Circadian rhythm resetting in sparrows: early response to doublet light pulses.

Circadian responses were studied using the perching activity of house sparrows (Passer domesticus). The sparrows were subjected to single or double 4-hr light pulses (the single pulses or the second pulses of the doublets scanned 24 hr) in the first cycle after previous entrainment to a light-dark cycle (LD 12:12). The differences in times at which the birds commenced perch-hopping in LD 12:12 before the pulses and in the five cycles immediately following the pulses were determined (phase shifts). A 24-hr time profile for phase shifts in response to single light pulses replicated our previous study: Early-night pulses delayed the rhythm (-1.7 hr), while late-night pulses advanced the rhythm (+3.8 hr). After pretreatment with a light pulse that advanced the birds +2.7 hr, the resetting curve was advanced. There were no delays; the range of average shifts was +0.1 hr to +6.2 hr. After pretreatment with a light pulse that delayed the birds -1.7 hr, the resetting curve was delayed. Average delays as much as -1.1 hr and advances up to +2.1 hr were measured. The data for double pulses were interpreted from predictions made from single-pulse data.

Animals↗

Effects of N-methyl-D-aspartate on luteinizing hormone release and Fos-like immunoreactivity in the male White-crowned sparrow (Zonotrichia leucophrys gambelii).

Seasonal breeding is terminated in the White-crowned sparrow by the onset of absolute photorefractoriness, a condition in which the reproductive system is switched off indefinitely until it is dissipated by short day lengths. Absolute photorefractoriness is controlled by the central nervous system; however, the mechanisms underlying GnRH quiescence in photorefractory birds have yet to be elucidated. Using the excitatory amino acid glutamate agonist N-methyl-D-aspartate (NMDA), plasma LH levels in White-crowned sparrows were significantly elevated regardless of the reproductive or photoperiodic condition. NMDA also significantly induced Fos-like immunoreactivity (FLI) within the infundibular nucleus and median eminence, regions previously shown to express FLI after a photoperiodically driven LH rise. NMDA did not induce FLI within GnRH I neurons; instead, it significantly activated cells within the organum vasculosum of the lamina terminalis in close proximity to GnRH I perikarya. These findings provide the first evidence that photorefractoriness is not due to depletion of GnRH stores, as LH and presumably GnRH were secreted in response to excitatory amino acid stimulation. NMDA activation of FLI in the region of the organum vasculosum of the lamina terminalis and the basal tuberal hypothalamus suggests that seasonal reproductive neuroendocrine control may be mediated via cells in the region of the GnRH I perikarya and terminals.

Animals↗

The low-affinity glucocorticoid receptor regulates feeding and lipid breakdown in the migratory Gambel's white-crowned sparrow Zonotrichia leucophrys gambelii.

Plasma corticosterone increases during spring migration in a variety of bird species, including the Gambel's white-crowned sparrow Zonotrichia leucophrys gambelii. Corticosterone is elevated specifically in association with migratory flight, suggesting that corticosterone may promote processes such as energy mobilization and/or migratory activity. General effects of glucocorticoids support such a prediction. Because glucocorticoids exert permissive effects on food intake, corticosterone may also participate in the regulation of migratory hyperphagia. To examine the role of corticosterone during migration, we induced Gambel's white-crowned sparrows to enter the migratory condition and compared food intake and locomotor activity between controls and birds injected with RU486--an antagonist to the low-affinity glucocorticoid receptor (GR). In addition, we investigated effects of RU486 in birds that were subjected to a short-term fast. Results indicate that RU486 did not affect locomotor activity. However, consistent with its effects in mammals, RU486 suppressed food intake. Thus, hyperphagia and migratory restlessness, the two behaviors that characterize migration, may be regulated by different mechanisms. Lastly, RU486 antagonized fasting-induced lipid mobilization, as evidenced by decreased plasma free fatty acids. Thus, data on spring migrants suggest that endogenous corticosterone levels act through the GR to support hyperphagia and that the GR promotes availability of lipid fuel substrates in association with periods of energetic demand, e.g. during migratory flight.

Analysis of Variance↗

How the house sparrow Passer domesticus absorbs glucose.

According to the hypothesis that most glucose absorption occurs passively across intestinal tight junctions (paracellular absorption), one would predict fairly similar rates of in vivo absorption of L-glucose, the stereoisomer of D-glucose that is absorbed only passively and is not catabolized, and of 3-O-methyl-D-glucose (3OMD-glucose), the D-glucose analogue that is actively and passively transported and not catabolized. In house sparrows Passer domesticus, we applied a pharmacokinetic method to measure simultaneous in vivo absorption of [14C]L-glucose and [3H]3OMD-glucose in a situation in which intestinal glucose transporters were relatively saturated (gavage solution contained 200 mmol l(-1) 3OMD-glucose). Fractional absorptions (F) were not significantly different between [3H]3OMD- and [14C]L-glucose (0.80 vs 0.79), and the apparent rates of absorption did not differ significantly. When we performed the same experiment on other sparrows in a situation in which intestinal glucose transporters were relatively unsaturated (200 mmol l(-1) mannitol replaced 3OMD-glucose in the gavage solution), the apparent rate of absorption was significantly reduced for [14C]l-glucose by 39% and for [3H]3OMD-glucose by 26%. A simulation model showed that a reduction is not predicted if most of the [3H]3OMD-glucose is actively absorbed, because the absorption rate of the tracer should increase when competitive inhibitor (unlabeled 3OMD-glucose) is removed. The similar extent and rates of absorption of [3H]3OMD- and [14C]L-glucose, and the acceleration of their rates of absorption in the presence of luminal 3OMD-glucose, are most consistent with Pappenheimer's hypothesis that the majority of dietary D-glucose is absorbed passively.

3-O-Methylglucose↗

The relationship of central and peripheral organ masses to aerobic performance variation in house sparrows

We evaluated the relationship between organ mass and the limits to aerobic metabolism in house sparrows Passer domesticus. The results were used to test three models of performance limitation (the central limitation, peripheral limitation and symmorphosis concepts). Basal metabolic rate (BMR) was determined during the rest phase. The maximum rate of oxygen consumption during exercise ( v_dot (O2max)) was measured in an enclosed wheel that allowed limited hovering flight. Neither BMR nor v_dot (O2max) was affected by gender, but adults had significantly higher v_dot (O2max) and lower BMR than juveniles. The masses of most central organs (gut, gizzard, liver, heart, kidney and reproductive organs) differed significantly between ages. There were no gender differences in organ mass among juveniles, but liver mass differed between male and female adults. In the pooled data, BMR was positively correlated with the mass of three central organs (gut, liver and kidney) and with one peripheral effector (breast muscle); together, these explained more than half the variance in BMR (r(2)=0.57). In adults, BMR was positively correlated with the mass of reproductive tissue. The masses of one peripheral effector (breast muscle) and one central organ (the heart) were positively correlated with v_dot (O2max) (r(2)=0.17 for the pooled data set). These results are consistent with a symmorphosis model of aerobic capacity. We found a significant positive relationship between BMR and v_dot (O2max) in juveniles, but not in adults. Taken together, our data indicate that house sparrows can achieve elevated v_dot (O2max) without paying a 'penalty' (fitness trade-off) in the form of an increased BMR.

Journal Article↗

Passive absorption of hydrophilic carbohydrate probes by the house sparrow Passer domesticus.

To evaluate the permeability of the intestine of the house sparrow Passer domesticus to hydrophilic compounds, we applied a pharmacokinetic technique to measure in vivo absorption of two carbohydrate probes, l-arabinose and d-mannitol. Probes were fed or injected, and blood and excreta were subsequently collected and analyzed by gas chromatography/mass spectrometry. Following injection, plasma probe concentration decreased in a log-linear fashion, implying single-compartment, first-order kinetics. Following oral administration, plasma probe concentrations increased, reached a maximum at 10 min and then decreased in log-linear fashion. Mannitol and arabinose absorption were calculated from the areas under the post-absorption plasma curve and the respective distribution spaces and elimination constants. The amounts absorbed increased linearly with the concentration administered (range 1-1000 mmol x l(-1)), implying a passive process. The mouth-to-cloaca retention time of digesta, measured using the non-absorbable compound potassium ferrocyanide, was independent of probe concentration. On average, 69% of the oral dose of probe was absorbed and this was independent of the concentration of probe administered. This paper supports an earlier report of substantial passive glucose absorption in house sparrows and offers a method to study the extent of hydrophilic solute absorption, which has importance for future research in areas as diverse as biomedical, ecological and evolutionary physiology.

Animals↗

Effects of season on kidney morphology in house sparrows.

Seasonal variability in kidney morphology of the house sparrow Passer domesticus was examined using light microscopy. Sparrows were captured from the wild in winter, spring, summer and autumn. The kidneys were perfused with half-strength Karnovsky's fixative and processed for light microscopy by embedding in either paraffin wax or JB4 acrylic resin. Absolute volumes of the kidneys, their components (cortex, medulla and blood vessels), components of the nephron (renal corpuscles, proximal tubules, loops of Henle, distal tubules and collecting ducts) and the capillaries surrounding the nephron were quantified using stereology. Tissue processed in paraffin wax had a mean shrinkage of 17.7 % compared with 10.1 % in JB4 resin. Absolute volumes of the kidneys and nephrons were compared statistically between tissue processing methods and among seasons. Absolute volumes of the structures within the kidneys were not significantly different between treatments or among seasons. Within the nephron, the only measured variables to show significant differences were the absolute volumes of the distal tubules and cortical collecting ducts between tissue processing treatments. Thus, kidney morphology was relatively unaffected by changes in season. In addition, the results show that embedding tissue using acrylic resin causes less shrinkage, and it should therefore be the preferred embedding medium for quantitative morphologists.

Acrylic Resins↗

Altitudinal variation in parental energy expenditure by white-crowned sparrows.

We used the doubly labeled water technique to measure daily energy expenditure (DEE) during the incubation and feeding nestling stages in two populations of white-crowned sparrows (Zonotrichia leucophrys) - one montane and migratory, the other coastal and sedentary - that differ in thermal environment and clutch size. We assessed the birds' thermal environment by continuously monitoring (among other variables) operative temperature and wind speed both in the open and within bushes and willow thickets occupied by sparrows. From these measurements, we derived several estimates of the birds' thermal environment, including standard operative temperature (T(es)). Shade air temperature and T(es) averaged 6.6 and 10.3 degrees C lower, respectively, at the montane study site during DEE measurements. The montane population's DEE averaged 24% higher than that of the sea-level population (103.6+/-12.2 versus 83.7+/-9.6 kJ day(-1); means +/- S.D., N=31 and 22, respectively), reflecting both its larger brood size (3.7 versus 2.9) and the colder environment. The DEE:BMR ratio was lowest in the sea-level population (2.1 versus 2.6), but neither population worked to their physiological capacity to produce young. DEE was significantly correlated with temperature across populations, with T(es) explaining 42% of the variation in DEE. Statistically removing the effect of temperature by adjusting DEE to a common temperature reduced the difference in DEE between populations by 34% to 87.7 and 100.8 kJ day(-1), respectively, for sea-level and montane populations. Basal and resting metabolic rates were similar in both populations, implying that greater activity in the montane population accounted for its higher temperature-adjusted DEE. Our results indicate that the thermal context within which behavior occurs can significantly affect interindividual variation in DEE. Attempts to assess reproductive effort by measuring DEE should therefore account explicitly for the effect of temperature.

Altitude↗

Androgen feedback-dependent and -independent control of photoinduced LH secretion in male tree sparrows (Spizella arborea).

Photoperiodic control of gonadotrophin secretion in male tree sparrows was studied by examining changes in plasma LH in castrated birds retained on short daylengths and in castrated birds transferred to long daylengths. Plasma LH concentrations were markedly higher in photostimulated birds than in non-photostimulated birds throughout the 25-day experiment, and implantation of the antiandrogen cyproterone (free alcohol), which should have blocked the action of castration-resistant androgens, did not increase plasma LH in either group. Such results, obtained from birds in which testosterone feedback was inoperative, indicate that the gonadostimulatory effect of long daylengths in intact males must be mediated, at least in part, by an androgen feedback-independent mechanism. To determine whether changes in testosterone feedback facilitate gonadotrophin secretion during photostimulation, two feedback performance characteristics (i.e. set point (minimum concentration of testosterone that suppresses plasma LH) and sensitivity (change in plasma LH per unit change in testosterone)) were quantified by evaluating plasma LH responses of non-photostimulated castrated birds and of photostimulated castrated birds to replacement testosterone (0-4.16 mumol). The data indicate that, in addition to stimulating LH secretion by an androgen feedback-independent mechanism, long daylengths reduce feedback inhibition of LH secretion by increasing the putative set point and decreasing the sensitivity of the testosterone feedback mechanism. The feedback-independent effect is the predominant effect of photostimulation on LH secretion in male tree sparrows.

Animals↗

An androgen-independent mechanism maintains photorefractoriness in male tree sparrows (Spizella arborea).

A series of experiments was performed to clarify whether photorefractoriness in male tree sparrows is maintained by an androgen-dependent mechanism. Castration did not raise plasma LH in photorefractory males held under a daily photoperiod of 20 h light:4 h darkness (20L:4D). Castrated photorefractory males were implanted with the antiandrogen cyproterone or injected s.c. with the antiandrogen flutamide to determine whether androgens which may be resistant to castration inhibit LH secretion. Neither cyproterone nor flutamide raised plasma LH above values found in castrated control birds. Castrated photorefractory males were treated with testosterone to determine whether plasma LH in photorefractory males is androgen-suppressible. Concentrations of plasma LH were independent of plasma testosterone over a wide range of concentrations. The lack of LH response to castration, to castration coupled with antiandrogen therapy, and to castration coupled with testosterone replacement argues that photorefractoriness in male tree sparrows is maintained by an androgen-independent mechanism.

Animals↗

Testosterone sensitivity of the seminal sacs of tree sparrows (Spizella arborea) in different reproductive states.

Testosterone sensitivity of the seminal sacs of castrated tree sparrows from each of three reproductive states was evaluated by measuring the change in seminal-sac mass per unit change in the logarithm of replacement or plasma testosterone. Birds were exposed to exogenous testosterone for 38 days. Replacement doses less than 0.17 mumol or plasma concentrations less than about 0.7 nmol/l did not induce seminal-sac growth in photosensitive castrated birds held on short days, in photosensitive castrated birds transferred from short to long days, or in photorefractory castrated birds retained on long days. Higher replacement doses or plasma concentrations, however, stimulated log dose-dependent growth of the seminal sacs in castrated birds from all three reproductive states. The change in seminal-sac mass per unit change in the logarithm of the dose of replacement testosterone was less (P = 0.0495) in photosensitive castrated birds held on short days than in photosensitive castrated birds transferred to long days. A more critical test of sensitivity (i.e. the change in seminal-sac mass per unit change in the logarithm of mean plasma testosterone concentration) indicated, however, that sensitivity of the seminal sacs to testosterone is independent of reproductive state. That result, when considered in the context of the plasma testosterone profile of intact males during a simulated reproductive cycle, argues that the seminal sacs of sexually quiescent (photosensitive or photorefractory) tree sparrows are small not because of their insensitivity to androgens, but because of a deficiency of circulating androgens.

Animals↗

Changes in the daily rhythm of plasma corticosterone concentration related to seasonal conditions in the white-throated sparrow, Zonotrichia albicollis.

Circadian variations in concentrations of plasma corticosterone were investigated in the white-throated sparrow maintained on short (10-hr) or long (16-hr) daily photoperiods. In addition, the plasma concentrations of corticosterone were determined throughout a day in birds that were in the reproductively photosensitive spring migratory condition, the reproductively photorefractory post nuptial molt condition, and the fall migratory condition. Distinct unimodal rhythms were found in photosensitive birds. The daily rise occurred 12 hr after the offset of light in birds kept on both the short and the long photoperiodic regimens. There was no discernible daily variation in photorefractory birds kept on a 16 hr daily photoperiod and there was a bimodal rhythm in the birds that were in the fall migratory condition. The results are consistent with an hypothesis that assigns an important role to the circadian rhythm of corticosteroid concentration in the photoperiodic mechanism controlling seasonal reproductive and migratory conditions in the white-throated sparrow.

Animals↗

Diel rhythms of basal and stress-induced corticosterone in a wild, seasonal vertebrate, Gambel's white-crowned sparrow.

Glucocorticoids have a wide array of actions in vertebrates. Daily fluctuations in basal levels of glucocorticoids are thought to regulate homeostatic mechanisms. In contrast, elevated levels secreted in response to stress stimulate changes in physiology and behavior. These changes are thought to aid an animal in avoiding chronic stress or death. Twenty-four-hour rhythms in basal and stress-induced glucocorticoids have been detected in laboratory mammals, but studies in wild, seasonal vertebrates are rare. Identification of plasticity in hormone secretion in wild vertebrates is critical to understanding the effects of hormones on physiology and behavior, and therefore the success of an animal in its natural environment. In the present study, we characterized diel patterns of basal and stress-induced corticosterone (the avian glucocorticoid) under two photoperiods in Gambel's white-crowned sparrow (Zonotrichia leucophrys gambelii). In contrast to previous findings in the white-crowned sparrow, we demonstrated a robust rhythm in basal corticosterone secretion, in which corticosterone reaches peak levels at the end of the inactive period, and has returned to trough levels just after the active period has begun. We also demonstrated a diel rhythm in secretion of corticosterone in response to a stressor, showing the greatest response at the beginning of the active period. Patterns of CORT secretion were similar under both photoperiods. These patterns show interesting similarities and differences to classical mammalian rhythms.

Animals↗