Fewer Exposed Lysine Residues May Explain Relative Resistance of Chicken Serum Albumin to In Vitro Protein Glycation in Comparison to Bovine Serum Albumin.

Protein glycation and formation of advanced glycation end products is associated with several diseases resulting from high blood glucose concentrations. Plasma albumin is directly exposed to circulating glucose concentrations and is therefore at greater risk of glycation than hemoglobin. As plasma glucose concentrations in birds are 1.5-2 times higher than mammals of similar mass, avian albumin may be particularly at risk of glycation. Thus, the goal of the present study was to compare the in vitro formation of glycated albumin in chicken serum albumin (CSA) and bovine serum albumin (BSA) exposed to a range of glucose concentrations over a 16-week period. The level of glycation for CSA and BSA was quantified using boronate affinity columns to separate glycated albumin from non-glycated albumin and calculating the difference in protein concentration of each sample. The results indicate that CSA is glycated to a lesser degree than BSA when the albumins are exposed to increasing concentrations of glucose (38.8-500 mM). This was most apparent at week sixteen (500 mM glucose) where BSA expressed a higher degree of glycation (37.8 ± 0.76%) compared to CSA (19.7 ± 1.06%, P < 0.05). Additionally, percent glycation at week sixteen was significantly higher than the glucose-free solutions for both BSA and CSA, indicating that glycation is glucose-dependent. Analyses of the protein structures suggest that the relative resistance of CSA to glycation may be due to fewer lysine residues and variations in protein folding that shield more lysine residues from the plasma. Moreover, comparisons of reconstructed ancestral albumin sequences show that the ancestor of birds had 6-8 fewer lysine residues compared to that of mammals.

Novel role of insulin in the regulation of glucose excretion by mourning doves (Zenaida macroura).

In mammals, insulin primarily lowers plasma glucose (PGlu) by increasing its uptake into tissues. Studies have also shown that insulin lowers PGlu in mammals by modulating glomerular filtration rate (GFR). Birds have naturally high PGlu and, although insulin administration significantly decreases glucose concentrations, birds are resistant to insulin-mediated glucose uptake into tissues. Since prior work has not examined the effects of insulin on GFR in birds, the purpose of the present study was to assess whether insulin can augment renal glucose excretion and thereby lower PGlu. Therefore, the hypothesis of the present study was that insulin lowers PGlu in birds by augmenting GFR, as estimated by inulin clearance (CIn). Adult mourning doves (Zenaida macroura) were used as experimental animals. Doves were anesthetized and the brachial vein was cannulated for administration of [14C]-inulin and insulin and the brachial artery was cannulated for blood collections. Ureteral urine was collected via a catheter inserted into the cloaca. Ten minutes following administration of exogenous insulin (400µg/kg body mass, i.v.) plasma glucose was significantly decreased (p=0.0003). Twenty minutes following insulin administration, increases in GFR (p=0.016) were observed along with decreases in urine glucose concentrations (p=0.008), glucose excretion (p=0.028), and the fractional excretion of glucose (p=0.003). Urine flow rate (p=0.051) also tended to increase after administration of insulin. These data demonstrate a significant role for insulin in modulating GFR in mourning doves, which may in part explain the lower PGlu measured following insulin administration.

Chicken albumin exhibits natural resistance to glycation.

Glycation of proteins and subsequent production of advanced glycation end products (AGEs) is a major contributor to the pathophysiology of diabetes. The objective of the present study was to compare the glycation of avian and human serum albumin to elucidate the mechanisms by which protein glycation in birds is prevented in the presence of naturally high plasma glucose concentrations. Solutions of purified chicken and human serum albumin (CSA and HSA) were prepared with four different glucose concentrations (0, 5.56, 11.1, and 22.2mM) and incubated at three temperatures (37.0, 39.8, and 41.4°C) for seven days. The solutions were sampled on Days 0, 3, and 7 and analyzed by liquid chromatography-electrospray ionization-mass spectrometry for the presence of glycated albumin. Four-way repeated measures ANOVA (p=0.032) indicate that all independent variables (albumin type, glucose concentration, temperature and time) interacted to affect the degree of glycation. With increasing glucose concentration, the glycation of both HSA and CSA increased with time at all temperatures. In addition, HSA was glycated to a greater extent than CSA at the two higher glucose concentrations for all temperature conditions. Glycation was elevated with increasing temperatures for HSA but not CSA. The results suggest an inherent difference between human and chicken albumin that contributes to the observed differences in glycation. Further research is needed to characterize this inherent difference in an effort to elucidate mechanisms by which avian plasma protein is glycated to a lesser degree than that of mammals (humans).

Comparison of metabolic substrates in alligators and several birds of prey.

On average, avian blood glucose concentrations are 1.5-2 times those of mammals of similar mass and high concentrations of insulin are required to lower blood glucose. Whereas considerable data exist for granivorous species, few data are available for plasma metabolic substrate and glucoregulatory hormone concentrations for carnivorous birds and alligators. Birds and mammals with carnivorous diets have higher metabolic rates than animals consuming diets with less protein whereas alligators have low metabolic rates. Therefore, the present study was designed to compare substrate and glucoregulatory hormone concentrations in several birds of prey and a phylogenetically close relative of birds, the alligator. The hypothesis was that the combination of carnivorous diets and high metabolic rates favored the evolution of greater protein and fatty acid utilization leading to insulin resistance and high plasma glucose concentrations in carnivorous birds. In contrast, it was hypothesized that alligators would have low substrate utilization attributable to a low metabolic rate. Fasting plasma substrate and glucoregulatory hormone concentrations were compared for bald eagles (Haliaeetus leucocephalus), great horned owls (Bubo virginianus), red-tailed hawks (Buteo jamaicensis), and American alligators (Alligator mississippiensis). Avian species had high circulating ß-hydroxybutyrate (10-21 mg/dl) compared to alligators (2.81 ± 0.16 mg/dl). In mammals high concentrations of this byproduct of fatty acid utilization are correlated with insulin resistance. Fasting glucose and insulin concentrations were positively correlated in eagles whereas no relationship was found between these variables for owls, hawks or alligators. Additionally, ß-hydroxybutyrate concentrations were low in alligators. Similar to carnivorous mammals, ingestion of a high protein diet may have favored the utilization of fatty acids and protein for energy thereby promoting the development of insulin resistance and gluconeogenesis-induced high plasma glucose concentrations during periods of fasting in birds of prey.

Reciprocal inhibition of in vitro substrate movement into avian skeletal muscle.

Plasma glucose and ketone concentrations are much higher in birds than in humans and birds exhibit resistance to insulin-mediated glucose uptake into muscle. Therefore, birds may offer a model in which to examine the effects of high plasma glucose and free fatty acid (FFA) concentrations on substrate preference. The present study examined the uptake of radiolabeled oleic acid (OA; C18:1) and radiolabeled glucose by skeletal muscle isolated from the forewing of English sparrows (Passer domesticus). In dose-response studies, unlabeled glucose and OA (20 mM each) inhibited the uptake of their respective radiolabeled counterparts. To examine the effects of glucose on OA uptake, muscles were incubated for 60 min in a buffer containing 20 mM glucose with the addition of radiolabeled OA. This level of glucose significantly decreased radiolabeled OA uptake by 36%. Using the same methodology, 20 mM OA significantly decreased radiolabeled glucose transport by 49%. Comparing control values for glucose (0.952 ± 0.04 µM/mg muscle) and OA uptake (2.20 ± 0.29 µM/mg muscle), it is evident that OA is preferentially taken up by avian skeletal muscle. As FFAs provide a greater amount of energy per mole (146 ATP/OA) than carbohydrates (36 ATP/glucose), storing and utilizing fats may be more energy-efficient for birds. As studies in mammals have shown that FFAs may impair glucose uptake pathways, it is suspected that high FFA uptake by avian skeletal muscle may induce their notably lower glucose transport.

Architecture of kangaroo rat inner medulla: segmentation of descending thin limb of Henle's loop.

We hypothesize that the inner medulla of the kangaroo rat Dipodomys merriami, a desert rodent that concentrates its urine to more than 6,000 mosmol/kgH(2)O water, provides unique examples of architectural features necessary for production of highly concentrated urine. To investigate this architecture, inner medullary nephron segments in the initial 3,000 µm below the outer medulla were assessed with digital reconstructions from physical tissue sections. Descending thin limbs of Henle (DTLs), ascending thin limbs of Henle (ATLs), and collecting ducts (CDs) were identified by immunofluorescence using antibodies that label segment-specific proteins associated with transepithelial water flux (aquaporin 1 and 2, AQP1 and AQP2) and chloride flux (the chloride channel ClC-K1); all tubules and vessels were labeled with wheat germ agglutinin. In the outer 3,000 µm of the inner medulla, AQP1-positive DTLs lie at the periphery of groups of CDs. ATLs lie inside and outside the groups of CDs. Immunohistochemistry and reconstructions of loops that form their bends in the outer 3,000 µm of the inner medulla show that, relative to loop length, the AQP1-positive segment of the kangaroo rat is significantly longer than that of the Munich-Wistar rat. The length of ClC-K1 expression in the prebend region at the terminal end of the descending side of the loop in kangaroo rat is about 50% shorter than that of the Munich-Wistar rat. Tubular fluid of the kangaroo rat DTL may approach osmotic equilibrium with interstitial fluid by water reabsorption along a relatively longer tubule length, compared with Munich-Wistar rat. A relatively shorter-length prebend segment may promote a steeper reabsorptive driving force at the loop bend. These structural features predict functionality that is potentially significant in the production of a high urine osmolality in the kangaroo rat.

Naturally high plasma glucose levels in mourning doves (Zenaida macroura) do not lead to high levels of reactive oxygen species in the vasculature.

Plasma glucose (P(Glu)) concentrations in birds are 1.5-2 times higher than those of mammals of similar body mass. In mammals, sustained elevations of P(Glu) lead to oxidative stress and free radical-mediated scavenging of endogenous vasodilators (e.g., nitric oxide), contributing to elevated blood pressure. Despite the relatively high P(Glu) levels in birds, they appear resistant to the development of oxidative stress in tissues such as the heart, brain and kidneys. To our knowledge no information exists on oxidative stress susceptibility in the resistance vasculature of birds. Therefore, we compared endogenous antioxidant mechanisms in the resistance vasculature of mourning doves (MODO; Zenaida macroura) and rats (Rattus norvegicus). Reactive oxygen species (ROS) were assessed with the fluorescent indicator 7'-dichlorodihydrofluorescein diacetate, acetyl ester in mesenteric arteries from rats and wild-caught MODO. Despite having significantly higher P(Glu) than rats, there were no significant differences in ROS levels between mesenteric arteries from rats or doves. Although superoxide dismutase and catalase activities were lower in the plasma, total antioxidant capacity, uric acid, vitamin E (α-tocopherol), and carotenoids (lutein and zeaxanthin) were significantly higher in MODO than in rats. Thus, compared to rats, MODO have multiple circulating antioxidants that may prevent the development of oxidative stress in the vasculature.

Glucose regulation in birds.

Birds maintain higher plasma glucose concentrations (P(Glu)) than other vertebrates of similar body mass and, in most cases, appear to store comparatively very little glucose intracellularly as glycogen. In general, birds are insensitive to the regulation of P(Glu) by insulin. However, there appears to be no phylogenetic or dietary pattern in the avian response to exogenous insulin. Moreover, the high levels of P(Glu) do not appear to lead to significant oxidative stress as birds are longer-lived compared to mammals. Glucose is absorbed by the avian gastrointestinal tract by sodium-glucose co-transporters (SGLTs; apical side of cells) and glucose transport proteins (GLUTs; basolateral side of cells). In the kidney, both types of glucose transporters appear to be upregulated as no glucose appears in the urine. Data also indicate that the avian nervous system utilizes glucose as a metabolic substrate. In this review, we have attempted to bring together information from a variety of sources to portray how glucose serves as a metabolic substrate for birds by considering each organ system involved in glucose homeostasis.

Inhibition of lipolysis does not affect insulin sensitivity to glucose uptake in the mourning dove.

Birds have much higher plasma glucose and fatty acid levels compared to mammals. In addition, they are resistant to insulin-induced decreases in blood glucose. Recent studies have demonstrated that decreasing fatty acid utilization alleviates insulin resistance in mammals, thereby decreasing plasma glucose levels. This has yet to be examined in birds. In the present study, the levels of glucose and beta-hydroxybutyrate (BOHB), a major ketone body and indicator of fatty acid utilization, were measured after the administration of chicken insulin, acipimox (an anti-lipolytic agent), or insulin and acipimox in mourning doves (Zenaidura macroura). Insulin significantly decreased whole blood glucose levels (19%), but had no effect on BOHB concentrations. In contrast, acipimox decreased blood BOHB levels by 41%, but had no effect on whole blood glucose. In addition to changes in blood composition, levels of glucose uptake by various tissues were measured after the individual and combined administration of insulin and acipimox. Under basal conditions, the uptake of glucose appeared to be greatest in the kidney followed by the brain and skeletal muscle with negligible uptake by heart, liver and adipose tissues. Acipimox significantly decreased glucose uptake by brain (58% in cortex and 55% in cerebellum). No significant effect of acipimox was observed in other tissues. In summary, the acute inhibition of lipolysis had no effect on glucose uptake in the presence or absence of insulin. This suggests that free fatty acids alone may not be contributing to insulin resistance in birds.

Relationships between renal morphology and diet in 26 species of new world bats (suborder microchiroptera).

The renal morphology of 24 species of mormoopid and phyllostomid bats feeding on six different diets was examined to test evolutionary changes in several structural traits presumably led by dietary shifts from ancestral insectivorous diets. The kidneys of a fish-eating vespertilionid and an insect-eating emballonurid were also examined but not included in the phylogenetic comparison. The length, width, and breadth of the kidneys were used to calculate relative medullary thickness (RMT). Tissues were processed for stereological analysis, and the volumes of the kidney, nephron components, and vasculature were determined. RMT did not correlate with body mass in either animal-eating or plant-eating phyllostomid and mormoopid bats. The shift from insectivory to frugivory and nectarivory was accompanied by a reduction in RMT, a reduction in the percent of renal medulla, and an increase in the percent of renal cortex. No changes in these traits were observed in bats that shifted to carnivorous, omnivorous or sanguinivorous habits. No changes were observed in renal vasculature, in the percentage of cortical and medullary nephron components or of capillaries surrounding the nephrons in any feeding group. Vespertilionid and emballonurid species had similar values in all traits examined as compared to insectivorous phyllostomids and mormoopids. Our data suggest that diet does not influence a single area of the nephron, but rather the entire nephron such that the relative amounts of renal cortex and medulla are affected.

Glucose transporter expression in English sparrows (Passer domesticus).

Patterns of glucose transporter expression have been well-characterized in mammals. However, data for birds is currently restricted to isolated cells, domestic chickens and chicks, and ducklings. Therefore, in the present study, protein and gene expression of various glucose transporters (GLUTs) in English sparrow extensor digitorum communis, gastrocnemius and pectoralis muscles as well as heart, kidney, and brain tissues were examined. The hypothesis is that the expression pattern of avian GLUTs differs from mammals to maintain the high plasma glucose levels of birds and insulin insensitivity. Our studies failed to identify a GLUT4-like insulin responsive transporter in sparrows. GLUT1 gene expression was identified in all tissues examined and shares 88% homology with chicken and 84% homology with human GLUT1. Compared to the rat control, GLUT1 immunostaining of sparrow extensor digitorum communis muscle was weak and appeared to be localized to blood vessels whereas immunostaining of gastrocnemius muscles was comparable to rat muscle controls. Gene expression of GLUT3 was identified in all tissues examined and shares 90% gene sequence homology with chicken embryonic fibroblast and 75% homology with human GLUT3. Protein expression of GLUT3 was not determined as an avian antibody is not available. Moreover, the C-terminus of the mammalian GLUT3 transporter, against which antibodies are typically designed, differs significantly among species. The predominant difference of chicken and sparrow GLUT expression patterns from that of mammals is the lack of an avian GLUT4. The absence of this insulin responsive GLUT in birds may be a contributing factor to the observed high blood glucose levels and insulin insensitivity.

Oleic acid uptake by in vitro English sparrow skeletal muscle.

Studies of prolonged avian flight have shown it to require large amounts of energy supplied mainly by free fatty acids (FFA). In the present study, the high levels of plasma ketone bodies found in sparrows (2.58 mmol l(-1)) are supportive of the use of fatty acids for flight. To determine the nature of fatty acid (oleic acid, OA) uptake, various pharmacological agents were used. The uptake of OA was examined using the extensor digitorum communis (EDC) muscle of English sparrows incubated in vitro. Initial studies demonstrated that radiolabeled OA uptake decreased in the presence of increasing unlabeled OA, suggesting that uptake occurred by a facilitative transport process. To further characterize OA uptake, EDC muscles were incubated with either: insulin (2 ng ml(-1)), insulin-like growth factor isoform-1 (IGF-I; 48 ng ml(-1)), 5'-Aminoimidazole-4-carboxamide-1-beta-D-ribofuranoside (AICAR; 2 mmol) or caffeine (5 mmol). Insulin, but not IGF-I, significantly increased OA uptake by avian EDC (P < 0.01). Caffeine and AICAR were ineffective at increasing OA uptake. A specific inhibitor of FFA transport by fatty acid transporters (FAT/CD36), sulfo-N-succinimidyl oleate (SSO; 500 micromoles), significantly decreased OA uptake at 2.5 min. The effectiveness of SSO suggests that a FAT/CD36-like protein is expressed in avian tissues. As uptake of OA was not completely blocked by SSO, it is likely that other mechanisms for FFA movement across membranes, such as diffusion, may be present.

Skeletal muscle fiber composition of the English sparrow (Passer domesticus).

Substrate utilization by English sparrow skeletal muscle has been extensively studied in our lab. However, there are few published studies on the muscle fiber composition of English sparrow wing and gastrocnemius muscles. The objective of the present study was to examine the fiber type composition of a variety of muscles in the English sparrow. The classification of a muscle fiber as fast glycolytic, slow oxidative, or fast oxidative glycolytic provides insight into the physiological function of muscles. Therefore, we completed mATPase and NADH stains on four muscles of the sparrow wing, as well as the gastrocnemius muscle, to characterize these muscle fiber types. Results show that the fibers of extensor digitorum communis, extensor metacarpi ulnaris, and extensor metacarpi radialis are homogeneous fast oxidative. The fibers of the supinator are homogeneous fast oxidative in 62.5% of samples, and heterogeneous (45.2% fast oxidative, 54.8% fast nonoxidative) in 37.5% of samples. Whereas the gastrocnemius muscle fibers are heterogeneous (10% fast oxidative, 64% fast nonoxidative, 26% slow oxidative) in all muscles examined.

Transepithelial urate transport by avian renal proximal tubule epithelium in primary culture.

Birds are uricotelic, and because they excrete urate by renal tubular secretion, they provide a convenient model for examination of this process. Primary monolayer cultures of the isolated renal proximal tubule epithelium from the domestic chicken, Gallus gallus L., were mounted in Ussing chambers where several substrates/inhibitors of renal organic anion transporters were tested for the sidedness and specificity of their effects on transepithelial urate transport. Transepithelial electrical resistance, electrical potential and sodium-dependent glucose current were monitored to detect nonspecific effects. Under control short-circuited conditions the ratio of unidirectional fluxes of [(14)C]urate was found to be 3:1. Active net secretion was specifically inhibited by 1 mmol l(-1) probenecid and 10 mmol l(-1) para-aminohippuric acid (PAH). Bromocresol Green, cimetidine, nocodozole, cytochalasin D and ouabain also inhibited secretion but were toxic. Interstitial-side lithium (5 mmol l(-1)) and glutarate (1 mmol l(-1)) specifically blocked transport, but 10-100 micromol l(-1) glutarate had no effect. Interstitial estrone sulfate (ES) stimulated urate secretion at 10 micromol l(-1) but was inhibitory at 500 micromol l(-1). Active PAH secretion (5:1 flux ratio) was inhibited 34% by 330 micromol l(-1) urate. ES (500 micromol l(-1)) blocked the remainder. From the lumen side, glucose-free, Cl(-)-free and high K(+) (30 mmol l(-1)) solutions, or an alkaline pH of 7.7 had no effect on urate transport and neither did several compounds known to be uricosuric. Lumen-side methotrexate (500 micromol l(-1)) and MK571 (20 micromol l(-1)) strongly inhibited urate secretion. MK571 had no effect from the interstitial side. RT-PCR revealed mRNA for OAT1-, OAT3-, MRP2- and MRP4-like organic anion transporters in chicken proximal epithelium.

The functional morphology of the english sparrow cecum.

As birds do not have a urinary bladder, the kidneys and lower gastrointestinal tract must function in concert to maintain fluid and electrolyte homeostasis. In birds, urine is conveyed to the cloaca, and moved by reverse peristalsis into the colon and digestive ceca. Digestive ceca have been well studied for non-passerine birds and have been shown to absorb substrates and water. The ceca of passerine birds have been suggested to be non-functional because of their small size. The present study was undertaken to examine the morphology and cytochemistry of the small ceca of the English sparrow (Passer domesticus). Three-dimensional reconstruction of the ceca from serially sectioned tissue showed these organs to have a central channel with a large number of side channels. Electron micrographs indicated that all of the channels are lined by epithelial cells with a very dense microvillus brush border as well as a region densely packed with mitochondria just below the brush border. Specific staining for Na(+), K(+)-ATPase indicated the enzyme to be localized to the brush border. Quantification of Na(+), K(+)-ATPase activity showed it to be comparable to the coprodeum of domestic fowl. The data suggest that the small ceca of passerine birds may function in fluid and electrolyte homeostasis.

Glucose transport by English sparrow (Passer domesticus) skeletal muscle: have we been chirping up the wrong tree?

Glucose uptake by mammalian skeletal muscle has been extensively covered in the literature, whereas the uptake of glucose by avian skeletal muscle has yet to be examined. As skeletal muscle provides the majority of postprandial glucose uptake in mammals, this study was designed to characterize the glucose transport mechanisms and glycogen content of avian skeletal muscle. In addition, plasma glucose levels were measured. English sparrow extensor digitorum communis (EDC) skeletal muscles were used for this study to quantify in vitro radiolabeled-glucose uptake. Uptake of labeled glucose was shown to decrease in the presence of increasing unlabeled glucose and was maximal by 60 minutes of incubation. Various agents known to increase glucose transport in mammalian tissues, via the insulin and contraction-responsive pathways, were used to manipulate and characterize in vitro transport in birds. The typical effectors of the mammalian insulin pathway, insulin (2 ng/ml) and insulin-like growth factor-1 (48 ng/ml), did not increase skeletal muscle glucose transport. Likewise, inducers of the mammalian contraction-responsive pathway had no effect on glucose transport by in vitro avian skeletal muscle (5 mM caffeine, 2 mM AICAR (5'-aminoimidazole-4-carboxamide-1-b-D-ribofuranoside). Interestingly, 200 microM phloretin, an agent used to block glucose transport proteins, significantly inhibited its uptake (P<0.001). These results suggest that a glucose transporter is responsible for glucose uptake by avian skeletal muscle, albeit at unexpectedly low levels, considering the high plasma glucose concentrations (265.9+/-53.5 mg/dl) and low skeletal muscle glycogen content (9.1+/-4.11 nM glucose/mg) of English sparrows.

Protein location and elemental composition of urine spheres in different avian species.

We examined the internal morphology, location of protein, and identity and location of elements, in avian urate-containing spheres in 9 species of birds. The urine spheres were collected from voided samples. The spheres ranged in size from 0.5-5.0 microm, except in the domestic fowl, where they ranged up to 10 microm in diameter. The internal morphology of the spheres was examined using freeze-fracture microscopy. Protein location within the spheres was identified using fluorescein isothiocyanate (FITC). The urine spheres were analyzed for content and internal location of elements using Energy Dispersal System Analysis (EDS). Internally, the spheres consisted of a central nidus surrounded by 3-4 concentric narrow rings of protein. Elements found within the spheres included nitrogen, potassium, calcium, sodium, phosphorus, chloride and sulfur; however, only nitrogen, potassium and chloride were common in the spheres of all species. Nitrogen comprised the majority of the elemental content of the spheres (77-90%) followed by potassium (8-45%), with all other ions present in trace amounts. Unlike protein, the location of elements was random within the spheres. Protein and urate are both negatively charged and known to associate to form the spheres and as potassium is the only cation common to all spheres, it too may play a role in their formation.

Regulation of renal and lower gastrointestinal function: role in fluid and electrolyte balance.

For the majority of vertebrates, the kidneys are not the sole organs that function to maintain homeostasis of body fluid and electrolytes. Mammals are unusual in this respect, as the kidneys are the organs that fill this role. For non-mammalian vertebrates, other organs such as gills, skin, salt glands, urinary bladders and the gastrointestinal (GI) system function in concert with the kidneys in the control of fluid and ion balance. Birds are of particular interest and unique as they do not possess a urinary bladder and the renal output enters the lower GI tract. The physiology of the interaction of avian kidneys and lower GI tract is an excellent example of integrative physiology and several aspects of it have been examined, for example, the role of the avian antidiuretic hormone (arginine vasotocin, AVT) in controlling renal output. AVT produces both a tubular and glomerular antidiuresis. The glomerular antidiuresis is important, as the fluid from the kidneys that enters the GI should not be highly concentrated. Another hormone, aldosterone, has been shown to play an important role in regulating the transport of sodium by the GI epithelium. In addition, the lower GI tract plays a significant role in recycling a portion of the nitrogen that leaves the kidneys as uric acid. Furthermore, the output of avian kidneys contains large amount of protein that is conserved by the lower GI tract.

Glomerular and medullary architecture in the kidney of Anna's Hummingbird.

Hummingbirds have rates of water turnover that are among the highest of any bird, consuming up to five times their body mass in nectar each day. To determine if the processing of these extraordinary volumes of water is associated with structural specializations in the kidney, we examined the renal morphology of Anna's hummingbird (Calypte anna) using scanning electron microscopy of vascular and tubular casts. The glomerular tufts are simple, containing a single, unbranched capillary that is spiraled or folded back on itself only one or two times. There is no evidence that nectarivory in this species is associated with a relative increase in the size of the glomeruli. The medullary cones are small, containing only a few loops of Henle and collecting ducts. The vasa recta form a complex network of branching and anastomosing capillaries. In this nectarivore, the structures necessary to produce urine that is hyperosmotic to plasma are poorly developed or absent, which is consistent with urine osmolalities that are uniformly low. J. Morphol. 240:95-100, 1999. © 1999 Wiley-Liss, Inc.