title: Adaptations of the reed frog Hyperolius viridiflavus (Amphibia, Anura, Hyperoliidae) to its arid environment : III. Aspects of nitrogen metabolism and osmoregulation in the reed frog, Hyperolius viridiflavus taeniatus, with special reference to the role of iridophores.
abstract: Reed frogs of the superspecies Hyperolius viridiflavus occur throughout the seasonally very dry and hot African savannas. Despite their small size (300-700 mg), estivating reed frogs do not avoid stressful conditions above ground by burrowing into the soil, but endure the inhospitable climate relatively unprotected, clinging to mostly dry grass stems. They must have efficient mechanisms to enable them to survive e.g. very high temperatures, low relative humidities, and high solar radiation loads. Mechanisms must also have developed to prevent poisoning by the nitrogenous wastes that inevitably result from protein and nucleotide turnover. In contrast to fossorial amphibians, estivating reed frogs do not become torpid. Reduction in metabolism is therefore rather limited so that nitrogenous wastes accumulate faster in these frogs than in fossorial amphibians. This severely aggravates the osmotic problems caused by dehydration. During dry periods total plasma osmolarity greatly increases, mainly due to urea accumulation. Of the total urea accumulated over 42 days of experimental water deprivation, 30% was produced during the first 7 days. In the next 7 days rise in plasma urea content was negligible. This strong initial increase of urea is seen as a byproduct of elevated amino acid catabolism following the onset of dry conditions. The rise in total plasma osmolarity due to urea accumulation, however, is not totally disadvantageous, but enables fast rehydration when water is available for very short periods only. Voiding of urine and feces ceases once evaporative water loss exceeds 10% of body weight. Therefore, during continuous water deprivation, nitrogenous end products are not excreted. After 42 days of water deprivation, bladder fluid was substantially depleted, and urea concentration in the remaining urine (up to 447 mM) was never greater than in plasma fluid. Feces voided at the end of the dry period after water uptake contained only small amounts of nitrogenous end products. DSF (dry season frogs) seemed no to be uricotelic. Instead, up to 35% of the total nitrogenous wastes produced over 42 days of water deprivation were deposited in an osmotically inert and nontoxic form in iridophore crystals. The increase in skin purine content averaged 150 μg/mg dry weight. If urea had been the only nitrogenous waste product during an estimation period of 42 days, lethal limits of total osmolarity (about 700 mOsm) would have been reached 10-14 days earlier. Thus iridophores are not only involved in colour change and in reducing heat load by radiation remission, but are also important in osmoregulation during dry periods. The selective advantages of deposition of guanine rather than uric acid are discussed.
authors.affiliation: Zoologisches Institut der Universität, Röntgenring 10, D-8700, Würzburg, Germany.
authors.forename: R
authors.initials: R
authors.lastname: Schmuck
authors.affiliation: Zoologisches Institut der Universität, Röntgenring 10, D-8700, Würzburg, Germany.
authors.forename: K E
authors.initials: KE
authors.lastname: Linsenmair
keywords: Estivation; Hyperolius viridiflavus; Iridophores; Nitrogen metabolism; Osmoregulation