Attraction and Electrophysiological Response to Identified Rectal Gland Volatiles in Bactrocera frauenfeldi (Schiner).

Bactrocera frauenfeldi (Schiner) (Diptera: Tephritidae) is a polyphagous fruit fly pest species that is endemic to Papua New Guinea and has become established in several Pacific Islands and Australia. Despite its economic importance for many crops and the key role of chemical-mediated sexual communication in the reproductive biology of tephritid fruit flies, as well as the potential application of pheromones as attractants, there have been no studies investigating the identity or activity of rectal gland secretions or emission profiles of this species. The present study (1) identifies the chemical profile of volatile compounds produced in rectal glands and released by B. frauenfeldi, (2) investigates which of the volatile compounds elicit an electroantennographic or electropalpographic response, and (3) investigates the potential function of glandular emissions as mate-attracting sex pheromones. Rectal gland extracts and headspace collections from sexually mature males and females of B. frauenfeldi were analysed by gas chromatography-mass spectrometry. Male rectal glands contained (E,E)-2-ethyl-8-methyl-1,7-dioxaspiro [5.5]undecane as a major component and (E,E)-2,8-dimethyl-1,7-dioxaspiro[5.5]undecane as a moderate component. Minor components included palmitoleic acid, palmitic acid, and ethyl oleate. In contrast, female rectal glands contained (E,E)-2,8-dimethyl-1,7-dioxaspiro[5.5]undecane and ethyl laurate as major components, ethyl myristate and ethyl palmitoleate as moderate components, and 18 minor compounds including amides, esters, and spiroacetals. Although fewer compounds were detected from the headspace collections of both males and females than from the gland extractions, most of the abundant chemicals in the rectal gland extracts were also detected in the headspace collections. Gas chromatography coupled electroantennographic detection found responses to (E,E)-2,8-dimethyl-1,7-dioxaspiro[5.5]undecane from the antennae of both male and female B. frauenfeldi. Responses to (E,E)-2-ethyl-8-methyl-1,7-dioxaspiro[5.5]undecane were elicited from the antennae of females but not males. The two spiroacetals also elicited electropalpographic responses from both male and female B. frauenfeldi. Ethyl caprate and methyl laurate, found in female rectal glands, elicited responses in female antennae and palps, respectively. Y-maze bioassays showed that females were attracted to the volatiles from male rectal glands but males were not. Neither males nor females were attracted to the volatiles from female rectal glands. Our findings suggest (E,E)-2,8-dimethyl-1,7-dioxaspiro[5.5]undecane and (E,E)-2-ethyl-8-methyl-1,7-dioxaspiro[5.5]undecane as components of a sex-attracting pheromone in B. frauenfeldi.

Are the glandular trichomes in Jacquinia armillaris (Theophrastoideae-Primulaceae) salt glands?

Salt stress is harmful to plants, especially for those that live under conditions of intense salt aport. For this reason, several species present alternatives to prevent or diminish the damages that high salt concentrations may cause to the cells. Salt glands are one of these alternatives once they are specialized structures that secrete salt. Here, we aimed to investigate if the glandular trichomes in the leaves of Jacquinia armillaris are salt glands. Anatomical and ultrastructural observations showed that the glandular trichomes in J. armillaris resemble the salt glands from other recretohalophytes Primulaceae, such as, their occurrence in sunken regions in the leaf epidermis, the presence of a large basal cell that acts as a collecting cell, the detachment of the cuticle from the outer periclinal walls forming a cuticular chamber, the thickness of the cuticle in the stalk portion of the trichome, and the presence of sodium and chloride ions in the secretion and in the xylem. Altogether, the gathered results support the hypothesis that the glandular trichomes in J. armillaris are adapted to salt secretion, thus characterizing as salt glands.

Effects of short-term hyper- and hypo-osmotic exposure on the osmoregulatory strategy of unfed North Pacific spiny dogfish (Squalus suckleyi).

The North Pacific spiny dogfish (Squalus suckleyi) is a partially euryhaline species of elasmobranch that often enter estuaries where they experience relatively large fluctuations in environmental salinity that can affect plasma osmolality. Previous studies have investigated the effects of altered salinity on elasmobranchs over the long term, but fewer studies have conducted time courses to investigate how rapidly they can adapt to such changes. In this study, we exposed unfed (no exogenous source of nitrogen or TMAO) spiny dogfish to hyper- and hypo-osmotic conditions and measured plasma and tissue osmolytes, nitrogen excretion, and changes in enzyme activity and mRNA levels in the rectal gland over 24h. It was shown that plasma osmolality changes to approximately match the ambient seawater within 18-24h. In the hypersaline environment, significant increases in urea, sodium, and chloride were observed, whereas in the hyposaline environment, only significant decreases in TMAO and sodium were observed. Both urea and ammonia excretion increased at low salinities suggesting a reduction in urea retention and possibly urea production. qPCR and enzyme activity data for Na(+)/K(+)-ATPase did not support the idea of rectal gland activation following exposure to increased salinities. Therefore, we suggest that the rectal gland may not be a quantitatively important aspect of the dogfish osmoregulatory strategy during changes in environmental salinity, or it may be active only in the very early stages (i.e., less than 6h) of responses to altered salinity.

THE SHARK RECTAL GLAND MODEL: A CHAMPION OF RECEPTOR MEDIATED CHLORIDE SECRETION THROUGH CFTR.

The dogfish shark salt gland was predicted by Smith and discovered by Burger at the Mount Desert Island Biological Laboratory in Salisbury Cove, Maine. It is an epithelial organ in the intestine composed of tubules that serve a single function: the secretion of hypertonic NaCl. Many G protein receptors are present on the basolateral surface of these tubules, including stimulatory receptors for vasoactive intestinal peptide, adenosine A2, growth hormone releasing hormone, and inhibitory receptors for somatostatin and adenosine A1. An entirely different class of stimulatory receptors is present as C-type natriuretic peptide receptors. Each stimulatory receptor evokes powerful NaCl secretion. G protein receptors bind to Gαs to activate the catalytic unit of adenylate cyclase to form cyclic adenosine monophosphate (cAMP) and protein kinase A that phosphorylates the regulatory domain of cystic fibrosis transmembrane conductance regulator, opening the channel. The C-type natriuretic peptide receptor stimulates by activating guanylate cyclase and endogenous cyclic guanosine monophosphate which inhibits type 3 phosphodiesterase, the enzyme that breaks down cAMP, thereby elevating cAMP and activating the protein kinase A pathway.

Osmoregulatory and metabolic costs of salt excretion in the Rufous-collared sparrow Zonotrichia capensis.

Recent experiments on shorebirds have demonstrated that maintaining an active osmoregulatory machinery is energetically expensive. This may, in part, explain diet and habitat selection in birds with salt glands. However little is known about the osmoregulatory costs in birds lacking functional salt glands. In these birds, osmotic work is done almost exclusively by the kidneys. We investigated the osmoregulatory cost in a bird species lacking functional salt glands, the passerine Zonotrichia capensis. After 20 days of acclimation to fresh water (FW) and salt water (200 mM NaCl, SW), SW birds tended to be heavier than FW birds. However, this difference was not statistically significant. Total basal metabolic rate was higher in SW birds as compared with FW birds. Renal and heart masses were also higher in the SW group. We also found greater medullary development and an increase in urine osmolality in the SW group. In spite of Z. capensis' ability to tolerate a moderate salt load in the laboratory, we hypothesize that increased cost of maintenance produced by salt consumption may significantly affect energy budget, dietary, and habitat choices in the field.

Perspectives on the convergent evolution of tetrapod salt glands.

Since their discovery in 1958, the function of specialized salt-secreting glands in tetrapods has been studied in great detail, and such studies continue to contribute to a general understanding of transport mechanisms of epithelial water and ions. Interestingly, during that same time period, there have been only few attempts to understand the convergent evolution of this tissue, likely as a result of the paucity of taxonomic, embryological, and molecular data available. In this review, we synthesize the available data regarding the distribution of salt glands across extant and extinct tetrapod lineages and the anatomical position of the salt gland in each taxon. Further, we use these data to develop hypotheses about the various factors that have influenced the convergent evolution of salt glands across taxa with special focus on the variation in the anatomical position of the glands and on the molecular mechanisms that may have facilitated the development of a salt gland by co-option of a nonsalt-secreting ancestral gland. It is our hope that this review will stimulate renewed interest in the topic of the convergent evolution of salt glands and inspire future empirical studies aimed at evaluating the hypotheses we lay out herein.

Functional and morphological plasticity of crocodile (Crocodylus porosus) salt glands.

The estuarine crocodile, Crocodylus porosus, inhabits both freshwater and hypersaline waterways and maintains ionic homeostasis by excreting excess sodium and chloride ions via lingual salt glands. In the present study, we sought to investigate the phenotypic plasticity, both morphological and functional, in the lingual salt glands of the estuarine crocodile associated with chronic exposure to freshwater (FW) and saltwater (SW) environments. Examination of haematological parameters indicated that there were no long-term disruptions to ionic homeostasis with prolonged exposure to SW. Maximal secretory rates from the salt glands of SW-acclimated animals (100.8+/-14.7 micromol 100 g(-0.7) body mass h(-1)) were almost three times greater than those of FW-acclimated animals (31.6+/-6.2 micromol 100 g(-0.7) body mass h(-1)). There were no differences in the mass-specific metabolic rate of salt gland tissue slices from FW- and SW-acclimated animals (558.9+/-49.6 and 527.3+/-142.8 microl O(2) g(-1) h(-1), respectively). Stimulation of the tissue slices from SW-acclimated animals by methacholine resulted in a 33% increase in oxygen consumption rate. There was no significant increase in the metabolic rate of tissues from FW-acclimated animals in response to methacholine. Morphologically, the secretory cells from the salt glands of SW-acclimated animals were larger than those of FW-acclimated animals. In addition, there were significantly more mitochondria per unit volume in secretory tissue from SW-acclimated animals. The results from this study demonstrate that the salt glands of C. porosus are phenotypically plastic, both morphologically and functionally and acclimate to changes in environmental salinity.

Adaptive strategies for post-renal handling of urine in birds.

Birds are a diverse vertebrate class in terms of diet and habitat, but they share several common physiological features, including the use of uric acid as the major nitrogenous waste product and the lack of a urinary bladder. Instead, ureteral urine refluxes from the urodeum into the more proximal coprodeum and portions of the hindgut (colon or rectum and ceca). This presents a potential problem in that hyperosmotic ureteral urine in contact with the permeable epithelia of these tissues would counteract renal osmotic work. This review describes and provides examples of different strategies used by avian species to balance renal and post-renal changes in urine composition. The strategies described include: 1. a "reptilian" mode, with moderate renal concentrating ability, but high rates of post-renal salt and water resorption; 2. the "mammalian" strategy, in which the coprodeum effectively functions like a mammalian urinary bladder, preserving the osmotic concentrating work of the kidney; 3. an interaction strategy, in which post-renal transport processes are hormonally regulated in order to optimize renal function under varying conditions of salt or water stress; 4. the salt gland strategy seen in marine or estuarine birds with functional salt glands, in which post-renal transport mechanisms are used to conserve urinary water and to recycle excess NaCl to the nasal salt glands. Finally, we also describe some features of an as-yet unstudied group of birds, the birds of prey. At least some species in this group are relatively good renal concentrators, and would be predicted to have post-renal mechanisms to preserve this work. This new synthesis illustrates the marked diversity of adaptive mechanisms used by avian species to maintain osmotic homeostasis.

Osmoregulation in elephant fish Callorhinchus milii (Holocephali), with special reference to the rectal gland.

Osmoregulatory mechanisms in holocephalan fishes are poorly understood except that these fish are known to conduct urea-based osmoregulation as in elasmobranchs. We, therefore, examined changes in plasma parameters of elephant fish Callorhinchus milii, after gradual transfer to concentrated (120%) or diluted (80%) seawater (SW). In control fish, plasma Na and urea concentrations were about 300 mmol l(-1) and 450 mmol l(-1), respectively. These values were equivalent to those of sharks and rays, but the plasma urea concentration of elephant fish was considerably higher than that reported for chimaeras, another holocephalan. After transfer to 120% SW, plasma osmolality, urea and ion concentrations were increased, whereas transfer to 80% SW resulted in a fall in these parameters. The rises in ion concentrations were notable after transfer to 120% SW, whereas urea concentration decreased predominantly following transfer to 80% SW. In elephant fish, we could not find a discrete rectal gland. Instead, approximately 10 tubular structures were located in the wall of post-valvular intestine. Each tubular structure was composed of a putative salt-secreting component consisting of a single-layered columnar epithelium, which was stained with an anti-Na(+),K(+)-ATPase serum. Furthermore, Na(+),K(+)-ATPase activity in the tubular structures was significantly increased after acute transfer of fish to concentrated SW (115%). These results suggest that the tubular structures are a rectal gland equivalent, functioning as a salt-secreting organ. Since the rectal gland of elephant fish is well developed compared to that of Southern chimaera, the salt-secreting ability may be higher in elephant fish than chimaeras, which may account for the lower plasma NaCl concentration in elephant fish compared to other chimaeras. Since elephant fish have also attracted attention from a viewpoint of genome science, the availability of fish for physiological studies will make this species an excellent model in holocephalan fish group.

Effect of melatonin on salt gland and kidney function of gulls, Larus glaucescens.

This study examined effects of exogenous melatonin on osmoregulatory hormones and water and sodium secretion by salt glands and excretion via the kidneys of Glaucous-winged gulls (Larus glaucescens). Six saline acclimated gulls were injected with inulin and paraminohippuric acid and then infused with 500 mM NaCl to stimulate salt gland secretion. Each bird was given infusions of NaCl alone and NaCl plus melatonin. Experiments were made one week apart in a randomized order. A large blood sample (to measure osmoregulatory hormones) was taken before infusion, at secretion, and at the end of infusion. A small blood sample was taken at the midpoint of each of six 10 min sequential collections of salt gland secretion and urine. Melatonin tended to increase plasma sodium concentration, did decrease plasma osmolality, but did not affect potassium concentration. Melatonin did not affect salt gland secretion rate or concentration nor renal plasma flow or glomerular filtration. Melatonin increased urine flow rate, tended to increase urine sodium concentration, and did decrease urine potassium concentration. Combined renal and extrarenal sodium excretion was greater during MT treatment. During NaCl infusion, angiotensin II increased, aldosterone decreased, and arginine vasotocin remained unchanged. Melatonin did not affect these responses. These data suggest an osmoregulatory role for melatonin in birds with salt glands.

Control of rectal gland secretion by blood acid-base status in the intact dogfish shark (Squalus acanthias).

In order to address the possible role of blood acid-base status in controlling the rectal gland, dogfish were fitted with indwelling arterial catheters for blood sampling and rectal gland catheters for secretion collection. In intact, unanaesthetized animals, isosmotic volume loading with 500 mmol L-1 NaCl at a rate of 15 mL kg-1 h-1 produced a brisk, stable rectal gland secretion flow of about 4 mL kg-1 h-1. Secretion composition (500 mmol L-1 Na+ and Cl-; 5 mmol L-1 K+; <1 mmol L-1 Ca2+, Mg2+, SO(4)2-, or phosphate) was almost identical to that of the infusate with a pH of about 7.2, HCO3- mmol L-1<1 mmol L-1 and a PCO2 (1 Torr) close to PaCO2. Experimental treatments superimposed on the infusion caused the expected disturbances in systemic acid-base status: respiratory acidosis by exposure to high environmental PCO2, metabolic acidosis by infusion of HCl, and metabolic alkalosis by infusion of NaHCO3. Secretion flow decreased markedly with acidosis and increased with alkalosis, in a linear relationship with extracellular pH. Secretion composition did not change, apart from alterations in its acid-base status, and made negligible contribution to overall acid-base balance. An adaptive control of rectal gland secretion by systemic acid-base status is postulated-stimulation by the "alkaline tide" accompanying the volume load of feeding and inhibition by the metabolic acidosis accompanying the volume contraction of exercise.

Shark rectal gland vasoactive intestinal peptide receptor: cloning, functional expression, and regulation of CFTR chloride channels.

Vasoactive intestinal peptide (VIP) is a secretagogue that mediates chloride secretion in intestinal epithelia. We determined the relative potency of VIP and related peptides in the rectal gland of the elasmobranch dogfish shark and cloned and expressed the VIP receptor (sVIP-R) from this species. In the perfused rectal gland, VIP (5 nM) stimulated chloride secretion from 250 +/- 66 to 2,604 +/- 286 microeq x h(-1) x g(-1); the relative potency of peptide agonists was VIP > PHI = GHRH > PACAP > secretin, where PHI is peptide histidine isoleucine amide, GHRH is growth hormone-releasing hormone, and PACAP is pituitary adenylate cylase activating peptide. The cloned sVIP-R from shark rectal gland (SRG) is only 61% identical to the human VIP-R1. It maintains a long, extracellular NH2 terminus with seven cysteine residues, and has three N-glycosylation sites and eight other residues implicated in VIP binding. Two amino acids considered important for peptide binding in mammals are not present in the shark orthologue. When sVIP-R and the CFTR chloride channel were coexpressed in Xenopus oocytes, VIP increased chloride conductance from 11.3 +/- 2 to 127 +/- 34 microS. The agonist affinity for activating chloride conductance by the cloned receptor was VIP > GHRH = PHI > PACAP > secretin, a profile mirroring that in the perfused gland. The receptor differs from previously cloned VIP-Rs in having a low affinity for PACAP. Expression of both sVIP-R and CFTR mRNA was detected by quantitative PCR in shark rectal gland, intestine, and brain. These studies characterize a unique G protein-coupled receptor from the shark rectal gland that is the oldest cloned VIP-R.

Selective permeability barrier to urea in shark rectal gland.

Elasmobranchs such as the dogfish shark Squalus acanthius achieve osmotic homeostasis by maintaining urea concentrations in the 300- to 400-mM range, thus offsetting to some degree ambient marine osmolalities of 900-1,000 mosmol/kgH(2)O. These creatures also maintain salt balance without losing urea by secreting a NaCl-rich (500 mM) and urea-poor (18 mM) fluid from the rectal gland that is isotonic with the plasma. The composition of the rectal gland fluid suggests that its epithelial cells are permeable to water and not to urea. Because previous work showed that lipid bilayers that permit water flux do not block flux of urea, we reasoned that the plasma membranes of rectal gland epithelial cells must either have aquaporin water channels or must have some selective barrier to urea flux. We therefore isolated apical and basolateral membranes from shark rectal glands and determined their permeabilities to water and urea. Apical membrane fractions were markedly enriched for Na-K-2Cl cotransporter, whereas basolateral membrane fractions were enriched for Na-K-ATPase. Basolateral membrane osmotic water permeability (P(f)) averaged 4.3 +/- 1.3 x 10(-3) cm/s, whereas urea permeability averaged 4.2 +/- 0.8 x 10(-7) cm/s. The activation energy for water flow averaged 16.4 kcal/mol. Apical membrane P(f) averaged 7.5 +/- 1.6 x 10(-4) cm/s, and urea permeability averaged 2.2 +/- 0.4 x 10(-7) cm/s, with an average activation energy for water flow of 18.6 kcal/mol. The relatively low water permeabilities and high activation energies argue strongly against water flux via aquaporins. Comparison of membrane water and urea permeabilities with those of artificial liposomes and other isolated biological membranes indicates that the basolateral membrane urea permeability is fivefold lower than would be anticipated for its water permeability. These results indicate that the rectal gland maintains a selective barrier to urea in its basolateral membranes.

Plasma osmolyte concentrations and rectal gland mass of bull sharks Carcharhinus leucas, captured along a salinity gradient.

Bull sharks (Carcharhinus leucas) were captured across a salinity gradient from freshwater (FW) to seawater (SW). Across all salinities, C. leucas were hyperosmotic to the environment. Plasma osmolarity in FW-captured animals (642 +/- 7 mosM) was significantly reduced compared to SW-captured animals (1067 +/- 21 mosM). In FW animals, sodium, chloride and urea were 208 +/- 3, 203 +/- 3 and 192 +/- 2 mmol l(-1), respectively. Plasma sodium, chloride and urea in SW-captured C. leucas were 289 +/- 3, 296 +/- 6 and 370 +/- 10 mmol l(-1), respectively. The increase in plasma osmolarity between FW and SW was not linear. Between FW (3 mosM) and 24 per thousand SW (676 mosM), plasma osmolarity increased by 22% or 0.92% per 1 per thousand rise in salinity. Between 24 per thousand and 33 per thousand, plasma osmolarity increased by 33% or 4.7% per 1 per thousand rise in salinity, largely due to a sharp increase in plasma urea between 28 per thousand and 33 per thousand. C. leucas moving between FW and SW appear to be faced with three major osmoregulatory challenges, these occur between 0-10 per thousand, 11-20 per thousand and 21-33 per thousand. A comparison between C. leucas captured in FW and estuarine environments (20-28 per thousand ) in the Brisbane River revealed no difference in the mass of rectal glands between these animals. However, a comparison of rectal gland mass between FW animals captured in the Brisbane River and Rio San Juan/Lake Nicaragua showed that animals in the latter system had a significantly smaller rectal gland mass at a given length than animals in the Brisbane River. The physiological challenges and mechanisms required for C. leucas moving between FW and SW, as well as the ecological implications of these data are discussed.

Cross-talk of phosphoinositide- and cyclic nucleotide-dependent signaling pathways in differentiating avian nasal gland cells.

In many bird species, the nasal glands secrete excess salt ingested with drinking water or food. In ducks ( Anas platyrhynchos), osmotic stress results in adaptive cell proliferation and differentiation in the gland. Using 'naive' nasal gland cells isolated from animals that had never ingested excess salt or 'differentiated' cells from animals fed with a 1% NaCl solution for 48 h, we investigated the allocation of metabolic energy to salt excretory processes and to other cellular activities. Activation of muscarinic acetylcholine receptors (carbachol) or beta-adrenergic receptors (isoproterenol) in nasal gland cells resulted in a transient peak in metabolic rate followed by an elevated plateau level that was maintained throughout the activation period. Activation of cells using vasoactive intestinal peptide, however, had only marginal effects on metabolic rate. In differentiated cells, sequential stimulation with carbachol and isoproterenol resulted in additive changes in metabolic rate during the plateau phase. Naive cells, however, developed supra-additive plateau levels in metabolic rates indicating cross-talk of both signaling pathways. Using bumetanide, TEA or barium ions to block different components of the ion transport machinery necessary for salt secretion, the relative proportion of energy needed for processes related to ion transport or other cellular processes was determined. While differentiated cells in the activated state allocated virtually all metabolic energy to processes related to salt secretion, naive cells reserved a significant amount of energy for other processes, possibly sustaining cellular signaling and regulating biosynthetic mechanisms related to adaptive growth and differentiation.

Regulation of salt gland, gut and kidney interactions.

Marine birds can drink seawater because their cephalic 'salt' glands secrete a sodium chloride (NaCl) solution more concentrated than seawater. Salt gland secretion generates osmotically free water that sustains their other physiological processes. Acclimation to saline induces interstitial water and Na move into cells. When the bird drinks seawater, Na enters the plasma from the gut and plasma osmolality (Osm(pl)) increases. This induces water to move out cells expanding the extracellular fluid volume (ECFV). Both increases in Osm(pl) and ECFV stimulate salt gland secretion. The augmented intracellular fluid content should allow more rapid expansion of ECFV in response to elevated Osm(pl) and facilitate activation of salt gland secretion. To fully utilize the potential of the salt glands, intestinally absorbed NaCl must be reabsorbed by the kidneys. Thus, Na uptake at gut and renal levels may constrain extrarenal NaCl secretion. High NaCl intake elevates plasma aldosterone concentration of Pekin ducks and aldosterone stimulates intestinal and renal water and sodium uptake. High NaCl intake induces lengthening of the small intestine of adult Mallards, especially males. High NaCl intake has little effect on glomerular filtration rate or tubular sodium Na uptake of birds with competent salt glands. Relative to body mass, kidney mass and glomerular filtration rate (GFR) are greater in birds with salt glands than in birds that do not have them. Birds with salt glands do not change GFR, when they drink saline. Thus, their renal filtrate contains excess Na that is, in some species, almost completely renally reabsorbed and excreted in a more concentrated salt gland secretion. Na reabsorption by kidneys of other species, like mallards is less complete and their salt glands make less concentrated secretion. Such species may reflux urine into the hindgut, where additional Na may also be reabsorbed for extrarenal secretion. During exposure to saline, marine birds maintain elevated aldosterone levels despite high Na intake. Marine birds are excellent examples of physiological plasticity.

Comparison of renal and salt gland function in three species of wild ducks.

Three processes central to osmoregulation of marine birds were compared in three species of ducks that differ in habitat affinity, diet and saline tolerance. These processes are filtration of Na+ and water from the plasma by the kidneys, their reabsorption along the renal tubules, and secretion by the salt glands. Barrow's goldeneyes Bucephala islandica, the most marine species, have the highest rates for all three processes and only this species can secrete all the infused salt via the salt glands. Rates of all three processes are lower in mallards Anas platyrhynchos, the most freshwater species. Following saline acclimation, mallards could excrete all the infused Na+ by a combined Na+ excretion of the kidneys and salt glands. Canvasbacks Aythya valisineria, despite being more saline tolerant than mallards, are unable to excrete all the infused Na+. They produce a large volume of urine (like mallards) that has a low [Na+] (like goldeneyes). Salt gland secretion Na+ concentration did not differ among the three species, but only goldeneyes secrete at a rate sufficient to eliminate all infused Na+ via the salt glands. Differences in saline tolerance of these ducks species cannot be fully explained by differences in their filtration, reabsorption and secretion of Na+ and water, suggesting that the intestinal tract plays an important role.

Effects of cadmium ingestion on plasma and osmoregulatory hormone concentrations in male and female Pekin ducks.

Effects of ingested cadmium (Cd) on body mass and plasma, urine, salt-gland secretion, and osmoregulatory hormone concentrations were assessed in male and female Pekin ducks, Anas platyrhynchos, acclimated to 450 mM NaCl over 6 wk and then held an additional 13 wk on 300 mM NaCl (prolonged saline exposure). Groups of six birds ate diets containing 0 (control), 50 (low-Cd diet), or 300 (high Cd diet) micrograms Cd/g food. Ducks that ingested Cd, especially females, lost body mass. Cadmium ingestion did not affect salt-gland secretion concentration. Control males had higher plasma osmolality and lower relative plasma volume. These increased in both sexes during saline acclimation. The high-Cd diet suppressed the increase in plasma osmolality in both sexes, but a rise in relative plasma volume occurred only in females. Following prolonged exposure to saline, plasma osmolality and relative plasma volume were reduced in control ducks but further elevated in ducks on the high-Cd diet. Cadmium ingestion suppressed the increase in urine osmolality that occurred in control ducks during saline acclimation. Neither saline acclimation nor Cd ingestion affected plasma concentrations of arginine vasotocin or prolactin. Arginine vasotocin was not correlated with plasma osmolality; prolactin was negatively correlated with plasma osmolality, but only in males. Cadmium suppressed the increase in angiotensin II that occurred at higher salinities in control and low-Cd males. This study examined the effects of gradually increasing body cadmium content on osmotic homeostasis. Cadmium affected plasma and urine, but not salt-gland secretion, concentrations and some of these responses were sexually disparate. Cadmium did not affect osmoregulatory hormones (arginine vasotocin and prolactin) by which observed changes in plasma concentration might have been influenced.

Effect of saline intake on water flux and osmotic homeostasis in Pekin ducks (Anas platyrhynchos).

The physiological regulation of body water volume and concentration was evaluated in Pekin ducks, Anas platyrhynchos, slowly acclimated to increasingly saline drinking water (six equal 75 mM NaCl increments). Body mass, total body water (TBW), water flux, plasma osmolality (Osm(pl)), and ionic and osmoregulatory hormone concentrations were measured at the end of each increment. The salinity at which each variable deviates from its homeostatic set point was calculated by continuous two-phase linear regression. We hypothesized that, as drinking water salinity increases: (1) body water increases in concentration before it decreases in volume and (2) that regulating variables that help determine homeostatically set values (plasma hormone concentrations and water flux) deviate from values of freshwater ducks at lower drinking water salinities than the variables they regulate (Osm(pl), hematocrit, TBW). Osm(pl) was the first variable for which we could calculate a deviation from its homeostatically controlled value. It increases at much lower drinking water salinity than that at which TBW decreases, supporting our first hypothesis, but not our second hypothesis. We further hypothesized that, because the concentration of Pekin duck salt gland secretion is only slightly higher than that of their drinking water, they increase water flux (drinking) as salinity of drinking water increases, until the latter exceeds the secretion concentration and then they drink less. There was no change in water flux until it decreases when TBW decreases, 329 mM NaCl and 335 mM NaCl, respectively. The results do not support our hypothesis that Pekin ducks increase drinking as the salinity of their drinking water increases, but do indicate that, at tolerable salinities, Pekin ducks maintain body water volume while allowing body water osmolality to increase. At higher salinities, ducks decrease drinking and use body water to get rid of the excess salt.