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PubMed1994

Oxytocin antagonist blocks the vasodepressor but not the vasopressor effect of neurohypophysial peptides in chickens.

Cockerels with permanent cannulas in the brachial artery and vein were put into isolated slings. Arterial pressure and heart rate were continuously recorded. Following habituation, tests were initiated. In each cockerel 2 nmol/kg of the tested neurohypophysial peptide (NPs) or analogue was IV injected six times at 6-min intervals. Arginine vasotocin (AVT) caused an immediate vasodepressor (VDP) effect and tachycardia. These subsided within 20-30 s and were followed by a vasopressor (VP) response and bradycardia. On repeated injections of AVT, the VDP response declined and bradycardia intensified. Arginine vasopressin (AVP), oxytocin (OT), and mesotocin (MT) had short-lasting VDP effect in the following order of potency: OT = MT > AVT > AVP. Only AVT and, more effectively, AVP, caused a VP response. The VDP effect of MT and OT declined on repeated injections. When AVT was injected after three injections of MT, it had mostly an immediate VP effect. Although the V1 agonist is VP in chickens, at the dose used the V1 antagonist, [d(CH2)5,O-Me-Tyr2]AVP, had no effect on cardiovascular responses to AVT. Pretreatment with OT antagonist, [d(CH2)5-O-Me-Tyr2-Thr4.Tyr9.Orn8]VT, abolished the VDP effect of all NPs. Thus, MT had no effect on blood pressure, whereas AVP and, more effectively, AVT, had a marked immediate VP action. In chickens the VDP effect of NPs is probably mediated by an OT/MT-like receptor, wherein the peptide's ring structure, shared by AVT, OT, and MT, is important.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1986

The regulation of the corticomelanotropic cell activity in Aves--II. Effect of various peptides on the release of ACTH from dispersed, perfused duck pituitary cells.

We have estimated the corticotropin-releasing activity (CRA) of different neurohypophyseal peptides and synthetic corticotropin-releasing factor (CRF) in the duck, using perfused dispersed pituitary cells and an ACTH radioimmunoassay adapted to duck material. Log dose-response curves were obtained for different doses of arginine-vasopressin (AVP), arginine-vasotocin (AVT), mesotocin (MT), oxitocin (OT) and ovine CRF (oCRF) and compared to the response obtained with dilutions of duck median eminence extracts (DME). All peptides tested behaved as partial agonists compared to DME. AVT and MT were the most potent of all peptides tested, with a capacity of 60% relative to DME. CRF was a weak agonist together with AVP and OT. AVT and CRF perfused together at equal doses significantly potentiated the effect of each other, yielding a dose-response line whose slope approximated that of DME. A similar design was used to test the CRA of the same substances in the rat. The main difference in the pattern of response between the two species was the low potency displayed by all the neurohypophyseal peptides in the rat, compared with CRF which, in contrast with what occurred with the duck system, was the most potent secretagogue of all peptides tested. It is concluded that in birds, as in mammals, the control of ACTH secretion may be exerted by neurohypophyseal peptides and a CRF-like peptide acting synergistically upon the corticomelanotropic cell.

Adrenocorticotropic Hormone↗

PubMed1988

The pneumostome rhythm in slugs: a response to dehydration controlled by hemolymph osmolality and peptide hormones.

1. One response of the terrestrial slug, Limax maximus to dehydration is the initiation and modulation of the pneumostome rhythm. When a slug has lost 15-20% of its initial body weight by evaporation, the frequency of pheumostome closures, which is less than 0.5 closures/min in fully hydrated slugs, begins to increase. 2. The frequency increases with further dehydration, but the average duration of each closure remains constant. Thus, the proportion of time during which the pneumostome is closed increases. Simultaneously, the area of the pneumostome opening decreases. 3. This behavior appears to be controlled in part by both the osmolality of the slug's hemolymph and by a peptide closely related to arginine vasotocin (AVT) and arginine vasopressin (AVP). Injecting intact slugs with mannitol, which increases the osmolality of the hemolymph, or with AVT or AVP, can initiate the pneumostome rhythm. 4. Mannitol injections, however, do not provoke the decrease in the area of the pneumostome opening which is induced by natural dehydration or by AVT or AVP injection. This suggests that at least two systems may be involved in the overall control of the pneumostome.

Animals↗

PubMed1995

Antidiuretic hormone decreases the intracellular pH in distal nephron epithelium (A6) by inhibiting Na+/H+ exchange.

1. To clarify the effect of arginine vasotocin (AVT) on intracellular pH (pHi) of A6 cells (an amphibian renal cell line), we measured pHi with a single-cell fluorescent imaging system in an HCO3(-)-nominally-free medium. 2. AVT (40 mU/ml) significantly decreased pHi from 7.44 +/- 0.11 to 7.19 +/- 0.22 (mean +/- SE) by inhibiting the Na+/H+ exchanger. 3. A membrane-permeable analogue of cAMP, dibutyryl cAMP (1 mM), significantly decreased pHi from 7.40 +/- 0.02 to 7.17 +/- 0.02, a response similar to that of AVT. 4. These data indicate that, in A6 cells, AVT suppresses the Na+/H+ exchanger via a cAMP-dependent pathway and the Na+/H+ exchanger maintains pHi under HCO3(-)-nominally-free conditions.

Amiloride↗

PubMed1981

REM sleep suppression in cats by melatonin.

The pineal indole melatonin, injected into the third ventricle of unanesthetized cats, induced NREM sleep within 11-15 min and suppressed REM sleep for about 3 hr. A dose-related effect could be established between 1, 10 and 100 ng melatonin. The reappearance of REM sleep at about 3 hr following melatonin administration occurred with a marked rebound. The present results further support the hypothesis that melatonin represents the releasing hormone for pineal arginine vasotocin.

Animals↗

PubMed1988

The regulation of the corticomelanotropic cell activity in aves. III--Effect of various peptides on the release of MSH from dispersed, perfused duck pituitary cells. Cosecretion of ACTH with MSH.

1. The melanotropin-releasing activity of arginine-vasopressin (AVP), arginine-vasotocin (AVT), oxitocin (OT), mesotocin (MT) and corticotropin-releasing factor (CRF) was studied in the duck using dispersed, perfused pituitary cells and a specific alpha-MSH RIA. 2. Log dose-response curves were obtained for all the peptides ranging from 5 to 100 ng/ml. All peptides behaved as partial agonists compared to duck median eminence extracts (DME). 3. AVT and MT displayed an alpha-MSH releasing capacity of 60% relative to DME whereas all other peptides behaved as weak agonists with less than 15% capacity relative to DME. 4. AVT and CRF when perfused together acted synergistically on alpha-MSH release yielding a dose response line whose slope approximated that of DME. 5. ACTH was cosecreted together with alpha-MSH in all situations studied with an ACTH to alpha-MSH molar ratio of about 10. 6. It is concluded that CRF and neurohypophyseal peptides may be physiological stimulators of both alpha-MSH and ACTH release in aves.

Adrenocorticotropic Hormone↗

PubMed1992

Cardiac activity of some peptide hormones in the frog, Rana tigrina.

1. The cardiac responses of isolated frog (Rana tigrina) atria to peptide hormones were studied. 2. Calcitonin gene-related peptide (CGRP), arginine vasotocin (AVT), bovine parathyroid hormone fragment (bPTH-(1-34)) and oxytocin (OXY) produced dose-related positive chronotropic and inotropic responses; atrial natriuretic peptide (ANP) was negative chronotropic and inotropic; cholecystokinin (CCK), vasoactive intestinal peptide (VIP) were without effects. 3. The dose-related responses under bPTH-(1-34) stimulation but not CGRP or AVT were attenuated in the presence of ANP (300 ng/ml, approximately 0.98 x 10(-7) M). As expected ANP decreased the basal AR and AT responses of the isolated atria and the inhibitory effects were dose-dependent. 4. As shown previously, propranolol blocked the atrial tension stimulated by bPTH (1-34) but did not alter the cardiac responses to CGRP and AVT. 5. In the presence of beta-adrenergic blocker (propranolol 10(-7) M) or ANP (10(-7) M), the AR and AT changes under ISO stimulation in the frog were also decreased. 6. These cardiac changes suggest the cardiac inhibitory effects of ANP are related to beta-adrenoceptor activity and ANP might be a beta antagonist.

Animals↗

PubMed1995

Ontogeny of vasotocinergic and mesotocinergic systems in the brain of the South African clawed frog Xenopus laevis.

For a better understanding of the development of neurotransmitter systems and of their putative functional significance during ontogenesis, the development of the vasotocin (AVT) and mesotocin (MST) systems in the brain of Xenopus laevis was studied by means of immunohistochemical techniques. Weakly immunoreactive fibers were already present at late embryonic stage 38 in the caudoventral part of the telencephalon and in the ventral part of the diencephalon. The earliest immunodetectable AVT and MST immunoreactive cell bodies were found in the developing preoptic area at late embryonic stage 43. At the end of the embryonic period (stage 45), AVT immunoreactive fibers have reached the future medial amygdala, the midbrain tegmentum, the median eminence and the neural lobe of the pituitary. When compared with AVT immunoreactive fibers, the development of MST fibers shows some temporal delay. During the premetamorphosis (stages 45-52), AVT immunoreactive cell bodies appear in the medial part of the suprachiasmatic nucleus, the dorsal infundibular region, and the midbrain tegmentum, whereas fibers can now be traced to the nucleus accumbens, the septum and the medial amygdala in the forebrain, to the midbrain tegmentum, the reticular formation, the raphe nuclei, and the solitary tract nucleus in the brainstem, and to the spinal cord. Further maturation of the AVT system during prometamorphosis (stages 53-58) includes the appearance of immunoreactive cell bodies in the lateral part of the suprachiasmatic nucleus, the ventral preoptic area, and the dorsal infundibular region. By the end of the metamorphosis (stage 65), the maturation of the AVT/MST systems reaches an almost adult-like pattern. It should be noted that in amphibians, in contrast to mammals, the early appearance of the AVT/MST systems, including their extensive extrahypothalamic component, suggests that the two neuropeptidergic systems may play a significant role during development.

Animals↗

PubMed2005

The role of protein kinase A and cyclin-dependent (CDC2) kinase in the control of basal and IGF-II-induced proliferation and secretory activity of chicken ovarian cells.

The aim of these experiments was to study the role of protein kinase A (PKA), cyclin-dependent kinase 2 (CDC2) and insulin-like growth factor II (IGF-II) in the control of ovarian function in domestic fowl, as well as the role of PKA and CDC2 in mediating the effects of IGF-II on the ovary. For this purpose, we studied the influence of an inhibitor of PKA (KT5720; 50 ng/ml), a CDC2 blocker (olomoucine; 1 microg/ml), IGF-II (0, 1, 10 or 100 ng/ml) and their combinations on cultured fragments of chicken ovarian follicular wall. Accumulation of PKA and CDC2 and secretion of progesterone (P4), testosterone (T), estradiol (E2) and arginine-vasotocin (AVT) were evaluated by using SDS-PAGE-Western blotting and RIA/EIA. IGF-II addition to culture medium stimulated T, E2 and AVT secretion and inhibited P4 secretion. These changes were associated with an increase in PKA and a decrease in CDC2 accumulation. The PKA blocker KT5720, when given alone, increased accumulation of PKA and secretion of T and E2, but not AVT and inhibited P4 secretion. The PKA blocker also prevented and even reversed the effects of IGF-II on PKA and steroid hormones secretion, but enhanced the action of IGF-II on AVT. The inhibitor of CDC2, olomoucine, when given alone, suppressed the expression of CDC2 and the secretion of P4 and AVT (but not T and E2). When given together with IGF-II, it augmented IGF-II-induced suppression of CDC2 and reversed the effects of IGF-II on P4 (but not on T, E2 or AVT). These observations demonstrate the involvement of PKA, CDC2 and IGF-II in regulating the secretory activity of avian ovarian cells. Our data also suggest the involvement of PKA in the mediation of IGF-II effects on P4, T, E2 and AVT secretion. CDC2 can mediate the effects of IGF-II on ovarian P4 secretion but not on other hormones.

Animals↗

PubMed2004

Ion transport mechanisms in the mesonephric collecting duct system of the toad Bufo bufo: microelectrode recordings from isolated and perfused tubules.

It is not clear how and whether terrestrial amphibians handle NaCl transport in the distal nephron. Therefore, we studied ion transport in isolated perfused collecting tubules and ducts from toad, Bufo bufo, by means of microelectrodes. No qualitative difference in basolateral cell membrane potential (Vbl) was observed between tubules and ducts in response to ion substitutions, inhibitor and agonist applications. Cl- substitution experiments indicated a small Cl- conductance in the basolateral membrane. The apical membrane did not have a significant Cl- conductance. Luminal [Na+] steps and amiloride application showed a small apical Na+ conductance. Arginine vasotocin depolarized Vbl. The small apical Na+ conductance indicates that the collecting duct system contributes little to NaCl reabsorption when compared to aquatic amphibians. In contrast, Vbl rapidly depolarized upon lowering of [Na+] in the bath, demonstrating the presence of a Na+-coupled anion transporter. [HCO3-] steps revealed that this transporter is not a Na+-HCO3- cotransporter. Together, our results indicate that a major task of the collecting duct system in B. bufo is not conductive NaCl transport but rather K+ secretion, as shown by our previous studies. Moreover, our results indicate the presence of a novel basolateral Na+-coupled anion transporter, the identity of which remains to be elucidated.

Animals↗

PubMed2005

The cloned avian neurohypophysial hormone receptors.

Arginine vasotocin (AVT), a neurohypophysial hormone, has many essential functions in birds including the regulation of salt and fluid balance, blood pressure, the stress response and a variety of behaviors. In addition, AVT controls reproductive functions in birds that are served by oxytocin in mammals. In the following review, we examine the functions of AVT in birds with an emphasis on the present state of knowledge concerning the cloned receptors for this important hormone. Receptor and gene structure, signal transduction mechanisms and expression pattern are all discussed. Finally, we explore the phylogenetic relationships between the cloned avian receptors and other vertebrate and invertebrate neurohypophysial hormone receptors.

Amino Acid Sequence↗

PubMed2005

Molecular cloning of an oxytocin-like receptor expressed in the chicken shell gland.

The avian homologs of arginine vasopressin (AVP) and oxytocin (OT) are arginine vasotocin (AVT) and mesotocin (MT), respectively. In birds, AVT shares many of the functions of AVP including regulation of fluid balance, blood pressure regulation and the stress response. AVT also plays an oxytocin-like reproductive role in birds by stimulating uterine (shell gland) contraction during oviposition. The role of MT in avian reproduction is not clear. Here, we report the cloning of a third neuropeptide receptor in the chicken (Gallus gallus). Parsimony analysis reveals that the new receptor has highest homology to mammalian OT receptors and the MT receptors of non-mammalian vertebrates. Moreover, the receptor bears far less homology to the two avian VT receptors that have been cloned. Reverse transcription-polymerase chain reaction and in in situ hybridization analyses reveal the receptor is expressed in both the endometrium and myometrium of the shell gland. The expression pattern and high homology to OT receptors suggest that the receptor may stimulate myometrial contraction and therefore play a critical role in oviposition.

Amino Acid Sequence↗

PubMed2006

Effects of ghrelin and its analogues on chicken ovarian granulosa cells.

The aim of these in vitro experiments was (1) to examine the effects of ghrelin on the basic functions of ovarian cells (proliferation, apoptosis, secretory activity); (2) to determine the possible involvement of the GHS-R1a receptor and PKA- and MAPK-dependent post-receptor intracellular signalling cascades; (3) to identify the active part of the 28-amino acid molecule responsible for the effects of ghrelin on ovarian cells. We compared the effect of full-length ghrelin 1-28, a synthetic activator of GHS-R1a, GHRP6, and ghrelin molecular fragments 1-18 and 1-5 on cultured chicken ovarian cells. Indices of cell apoptosis (expression of the apoptotic peptide bax and the anti-apoptotic peptide bcl-2), proliferation (expression of proliferation-associated peptide PCNA), and expression of protein kinases (PKA and MAPK) within ovarian granulosa cells were analysed by immunocytochemistry. The secretion of progesterone (P(4)), testosterone (T), estradiol (E(2)) and arginine-vasotocin (AVT) by isolated ovarian follicular fragments was evaluated by RIA/EIA. It was observed that accumulation of bax was increased by ghrelin 1-28, GHRP6 and ghrelin 1-18, but not by ghrelin 1-5. Expression of bcl-2 was suppressed by addition of ghrelin 1-28, GHRP6 and ghrelin 1-5, but promoted by ghrelin 1-18. The occurrence of PCNA was reduced by ghrelin 1-28, GHRP6, ghrelin 1-18 and ghrelin 1-5. An increase in the expression of MAPK/ERK1, 2 was observed after addition of ghrelin 1-28, GHRP6 and ghrelin 1-18, but not ghrelin 1-5. The accumulation of PKA decreased after treatment with ghrelin 1-28 and increased after treatment with GHRP6 and ghrelin 1-18 but not ghrelin 1-5. Secretion of P(4) by ovarian follicular fragments was decreased after addition of ghrelin 1-28 or ghrelin 1-5 but stimulated by GHRP6 and ghrelin 1-18. Testosterone secretion was inhibited by ghrelins 1-28 and 1-18, but not by GHRP6 or ghrelin 1-5. Estradiol secretion was reduced after treatment with ghrelin 1-28 but stimulated by ghrelins 1-18 and 1-5; GHRP6 had no effect. AVT secretion was stimulated by ghrelin 1-28, GHRP6 and ghrelin 1-18, but inhibited by ghrelin 1-5. The comparison of the effects of the four ghrelin analogues on nine parameters of ovarian cells suggest (1) a direct effect of ghrelin on basic ovarian functions-apoptosis, proliferation, steroid and peptide hormone secretion; (2) that the majority of these effects can be mediated through GHS-R1a receptors; (3) an effect of ghrelin on MAPK- and PKA-dependent intracellular mechanisms, which can potentially mediate the action of ghrelin at the post-receptor level; (4) that ghrelin residues 5-18 may be responsible for the major effects of ghrelin on the avian ovary.

Animals↗

PubMed2006

Novel expression and functional role of ghrelin in chicken ovary.

Ghrelin has recently emerged as pleiotropic regulator of a wide array of endocrine and non-endocrine functions. The former likely includes the control of gonadal function, as expression of ghrelin and its putative receptor, the GH secretagogue receptor type 1a (GHS-R1a), has been described in mammalian gonads, and direct effects of ghrelin in the control of testicular secretion and cell proliferation have been reported. Yet, the expression and/or functional role of ghrelin in gonads from non-mammalian species remain to be analyzed. The present study aimed to evaluate the expression of ghrelin and GHS-R genes in the chicken ovary, and to assess the potential involvement of ghrelin in the direct control of chick ovarian function. To this end, RT-PCR assays for ghrelin and GHS-R1a mRNAs were performed in ovarian tissue, and cultures of chicken ovarian cells were conducted in the presence of increasing doses (1, 10 or 100 ng/ml) of the ghrelin analog, ghrelin 1-18. Our results demonstrate that both ghrelin and GHS-R1a mRNAs are expressed in chick ovarian tissue. Moreover, challenge of ovarian granulosa cells with ghrelin 1-18 was able to induce markers of proliferation (i.e. expression of both PCNA and cyclin), and to modulate markers of apoptosis (i.e. decreased expression of caspase-3, bax, bcl-2 and TUNEL-positive cells). Moreover, ghrelin 1-18 increased the expression of PCNA, cyclin, bax and p53 in cultures of ovarian follicular fragments, where it also stimulated the release of progesterone, estradiol, arginine-vasotocin (AVT) and IGF-I, but not of testosterone. In conclusion, our study provides novel evidence for the gonadal expression of the genes encoding ghrelin and its cognate receptor in a non-mammalian species, i.e. the chicken ovary, and unravels the potential involvement of this newly discovered molecule in the control of key gonadal functions in the chick, such as proliferation, apoptosis, and hormone release.

Animals↗

PubMed2005

Central oxytocinergic and dopaminergic mechanisms regulating penile erection in conscious rats.

A series of in vivo studies in a conscious rat model was conducted to investigate the role of oxytocinergic and dopaminergic neurotransmission in the central regulation of penile erection. Oxytocin, when administrated either intracerebroventricularly (i.c.v.) or intrathecally (i.t.) at the spinal levels of L4-L6, produced dose-related erectogenic effects with a maximum at 0.1 microg/rat i.c.v. or 0.03 microg/rat i.t. Oxytocin-evoked penile activity was attenuated by the inhibitory effect of the selective oxytocin antagonist vasotocin analog [Pmp-Tyr(Me)-Ile-Thr-Asn-Cys]-Pro-Orn-Tyr-NH2 (0.1-1 microg, i.c.v. or i.t.). Penile erection induced by oxytocin was blocked by the dopaminergic receptor antagonist clozapine (1-10 micromol/kg i.p.) in a dose-dependent manner. Conversely, oxytocin antagonist microinjected locally (i.c.v. or i.t.) significantly attenuated the pro-erectile effects of systemic (s.c.) apomorphine, a centrally acting erectogenic agent through dopaminergic receptors. Together, these data indicate a possible concomitant role between dopamine and oxytocin in mediating penile erection at both the spinal and supraspinal sites.

Animals↗

PubMed2004

Evolutionary insights into the regulation of courtship behavior in male amphibians and reptiles.

Comparative studies of species differences and similarities in the regulation of courtship behavior afford an understanding of evolutionary pressures and constraints shaping reproductive processes and the relative contributions of hormonal, genetic, and ecological factors. Here, we review species differences and similarities in the control of courtship and copulatory behaviors in male amphibians and reptiles, focusing on the role of sex steroid hormones, the neurohormone arginine vasotocin (AVT), and catecholamines. We discuss species differences in the sensory modalities used during courtship and in the neural correlates of these differences, as well as the value of particular model systems for neural evolution studies with regard to reproductive processes. For example, in some genera of amphibians (e.g., Ambystoma) and reptiles (e.g., Cnemidophorus), interspecific hybridizations occur, making it possible to compare the ancestral with the descendant species, and these systems provide a window into the process of behavioral and neural evolution as well as the effect of genome size. Though our understanding of the hormonal and neural correlates of mating behavior in a variety of amphibian and reptilian species has advanced substantially, more studies that manipulate hormone or neurotransmitter systems are required to assess the functions of these systems.

Amphibians↗

PubMed2004

Changes in expression of AVT and AVT receptor (VT1) gene in hypothalamus and shell gland in relation to egg laying in white leghorn hen.

Oviposition is a complex phenomenon involving various regulatory mechanisms at the neuroendocrine levels. Present study was designed to access the changes in arginine vasotocin (AVT) and its receptor (VT1) gene expression in relation to the time of egg laying of white leghorn hen. The expression of AVT gene (Northern blot analysis and in situ hybridization) in the hypothalamus and localization of ir-AVT in the magnocellular neurons of paraventricular nuclei was studied 2 h before (-2 h), immediately after (0 h) and 2 h after (+2 h) egg laying. Simultaneous changes in the AVT and VT1 receptor gene in the shell gland, which finally responds to AVT for smooth muscle contraction and expulsion of egg, were also determined by semi-quantitative reverse transcriptase-polymerase chain reaction. The findings indicated increased hypothalamic AVT gene expression immediately after egg laying (0 h) when compared to 2 h before and 2 h after egg laying. AVT receptor gene expression in the shell gland also followed the same pattern. However, AVT gene expression in the shell gland, unlike that of hypothalamus was higher at -2 h compared to 0 and +2 h of oviposition. While highly significant increase was noted in plasma AVT concentration at the time of egg laying, other parameters such as plasma osmolality and ionic concentration (Na(+), K(+), Ca(2+), and Cl(-)) did not show any change. It is suggested that in addition to increased hypothalamic AVT transcript and peripheral release, local synthesis of AVT in the shell gland (paracrine release) may contribute to the contraction of shell gland smooth muscles during egg laying. Moreover, these findings clearly indicate temporal correlation of AVT and its receptor gene expression in different tissues during oviposition.

Animals↗

PubMed2006

Hormonal regulation of ion and water transport in anuran amphibians.

Amphibians occupy a wide variety of ecological habitats, and their adaptation is made possible through the specialization of the epithelia of their osmoregulatory organs, such as the skin, kidney, and urinary bladder, which control the hydromineral and acid-base balance of their internal medium. Amphibians can change drastically plasma Na+, Cl-, and urea levels and excretion rates in response to environmental stimuli such as acute desiccation and changes in external salinity. Several hormones and the autonomic nervous system act to control osmoregulation. Several ion channels including an epithelial sodium channel (ENaC), a urea transporter (UT), and water channels (AQPs) are found in epithelial tissues of their osmoregulatory organs. This mini review examines the currents status of our knowledge about hormone receptors for arginine vasotocin, angiotensin II and aldosterone, and membrane ion channels and transporters, such as ENaC, UT, and AQPs in amphibians.

Aldosterone↗