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Feeding and Excretion in the Scorpion Paruroctonus Mesaensis: Water and Material Balance

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Feeding and Excretion in the Scorpion Paruroctonus Mesaensis: Water and Material Balance J. exp. Biol. 110, 253-265 (1984) 253 gutted in Great Britain © The Company of Biologists Limited 1984 FEEDING AND EXCRETION IN THE SCORPION PARUROCTONUS MESAENSIS: WATER AND MATERIAL BALANCE BY STANLEY D.YOKOTA Department of Biology, University of California at Riverside, Riverside, California 92521, U.SA. Accepted 2 December 1983 SUMMARY Scorpions feed by a process involving the external grinding and digestion of prey, with the ingestion of only the soluble fraction. The water obtained from the prey represents the most important source of water intake for scorpions inhabiting arid regions, placing great importance on the animals' ability to utilize prey water effectively. The scorpionParuroctonus mesaen- sis (Stahnke) was found to ingest a mean of 88 % of the body water of selected prey. However, the scorpion loses 0-37 ml of its own body water per ml of prey water extracted, resulting in a net water gain of 0-51 ml water for every ml of prey water. Fluid uptake by the scorpion has been ascribed to a pharyngeal pumping mechanism. Direct measurements of the suction generated by the pharynx yielded a minimal estimate of its pumping capabil- ity of 130mmHg. The uptake and excretion of nitrogen and electrolytes by Paruroctonus mesaensis on a diet of Tenebrio molitor adults were also analysed. Almost all the potassium ingested was excreted, whereas most of the sodium and chloride were retained, possibly serving to expand haemolymph volume. Assuming a steady state for nitrogen, it was estimated that the net utilizable water obtained from prey, that is the water intake minus the excretory water necessitated by nitrogen excretion, was equivalent to about 35 % of the initial prey water or 69 % of the water ingested. INTRODUCTION The maintenance of water balance plays a prominent role in the lives of many terrestrial animals. Studies of the water relations of many diverse animal groups have focused largely on adaptations for decreasing water loss. The arthropods excel at two approaches to water conservation, having the lowest transpiratory loss rates and excreta with the lowest water activities recorded (Edney, 1977). In general, the means of water intake have received much less attention. Important sources of water for terrestrial arthropods include free standing water, preformed water in food, and water produced by metabolism, with a few species being able to obtain net water from Present address: Department of Physiology, College of Medicine, University of Arizona, Tucson, Arizona 14, U.S.A. * ey words: Arachnid physiology, water balance, feeding. 254 S. D. YOKOTA subsaturated atmospheres (Edney, 1977). In arid environments, in the absence^ drinking water, dietary and metabolic sources become increasingly important. Scor- pions, which are common inhabitants of such regions and which lack the ability to utilize water vapour, must rely on the water gained from their prey and their metabol- ism for most of the year (Hadley, 1974). Scorpions feed by a process involving external grinding with the ingestion of only a portion of their prey. Thus, not all the water and nutrients contained in the captured prey become available for utilization by this predator. There is also the potential for considerable evaporative losses during feeding, a situation exacerbated by the high temperatures, low humidities and strong winds frequent in desert regions. And, in addition to the water and energy acquired from each meal, the scorpion incurs a concomitant electrolyte load. In this study, the water, electrolyte and nitrogen intake through feeding and output by excretion have been evaluated in the scorpion Paruroctonus mesaensis. The ingestion of prey fluids by spiders and scorpions is thought to be due to the actions of the muscular pharynx (Savory, 1964; Snodgrass, 1965). Consequently, the ability of the scorpion to assimilate prey water may be limited by the maximal suction pressure developed by this pump. This same mechanism is likely to account for the ability to take up soil water demonstrated for several spiders (Parry, 1954) and scor- pions (Crawford & Wooten, 1973; Riddle, Crawford & Zeitone, 1976). Direct measurements of the suction pressures produced in the preoral cavity were made to assess the function of the pharyngeal pump. MATERIALS AND METHODS Adult specimens of Paruroctonus mesaensis were collected from sand dunes in the Coachella Valley of the Mojave Desert in Riverside County, California. The scorpions (2-5-3-5 g) were maintained in plastic vials at room temperature (21-24°C) and humidities (approximately 20-40 %) and fed Tenebrio molitor larvae and Periplaneta americana nymphs. Water uptake during feeding A flow-through system was constructed to determine the water fluxes during feed- ing. Air, dried by passage through a dry ice vapour trap and anhydrous calcium sulphate (Drierite), was passed through a plastic feeding chamber at a flow rate of 0-5 1 min"1, creating an average air velocity of 087 cm s"1. The water vapour gained during passage through the feeding chamber was then condensed in a collection tube immersed in an acetone-dry ice bath. The feeding chamber was maintained at 24—25 °C in an incubator. Adult crickets (Acheta domesticus) or nymphal cockroaches {Periplaneta americana), from 0-25-068 g fresh mass, to be used as prey were injected with about 70nCi of tritiated water in 5^1 water. After a 3h equilibration period, a 5/il haemolymph sample was taken for the determination of the specific activity of tritium. Each prey item was then weighed and offered to a preweighed scorpion which had been allowed to acclimate to the feeding chamber for at least 2 h. During feeding the scorpion ingests the soluble portion of the prey and compacts the residue into small masses termed feeding pellets. Within 15min of the scorpion discarding the ^ Scorpion feeding and excretion 255 J let, the air flow was stopped, the water vapour collection tube capped, and the scorpion and feeding pellets were weighed to the nearest 0-1 mg. A sample of the scorpion haemolymph was taken 2 h later for the determination of the specific activity of tritium. The feeding pellets were dried at 100°C for the measurement of dry mass. The water vapour collection tube was rinsed twice with 10 ml of scintillation fluid for the measurement of the tritiated water collected. Tritium activity was measured by liquid scintillation counting (Beckman CPM-100) with a dioxane-naphthalene-PPO scintillation fluid using the external standard method for quench correction. The initial prey body water was calculated from the live mass of the prey using the mean water content, determined by drying to constant mass, for crickets, 72-0±l-4% (±S.E., AT =12), and cockroaches, 65-9±0-9% (iV=5). The prey water ingested by the scorpion during feeding was calculated from the specific activity of tritium in the haemolymph of the prey before feeding and the specific activity in the scorpion haemolymph after feeding, assuming the scorpion body water was equal to 70% of the fresh mass (the mean for laboratory-kept animals), by the formula: Wpi = 0-70M.f(S./Sp), (1) where Wpi is the prey water ingested, M^ is the live mass of the scorpion after feeding, S, and Sp are the specific activities of tritium in the haemolymph of the scorpion and the prey, respectively, and the factor 0-70 has the units of mlg"1. The prey water which is not obtained by the scorpion is the difference between the initial prey body water and the volume ingested by the scorpion. However, the scorpion invests part of its own body water in the feeding process, by exuding digestive fluids into the prey mass being ground. Thus, the net water uptake during feeding is the difference between the prey water ingested and the net scorpion body water which is either evaporated during feeding or left with the pellets, and can be calculated by the following equation: Wnet = (M,f - M.i) - (Mpd - Mfd), (2) where Msf and MB are the final and initial live mass of the scorpion respectively, is the calculated dry mass of the prey, and Mfd is the dry mass of the feeding pellets produced. The water which is not ingested by the scorpion is therefore composed of water both from the prey and the scorpion and has two components: the fraction evaporated during feeding and the liquid fraction left with the feeding pellets. Because several pellets may be produced sequentially from a single prey, the water left with the first pellets gradually evaporates to be collected as water vapour. Consequently, the precise measurement of the water content of the feeding pellets when discarded was not made. Although these considerations influence the relative proportion of loss in the two fractions, they should not affect the measurement of the prey water ingested or net water uptake. Electrolyte and nitrogen uptake and excretion The uptake and balance of electrolytes (Na, K and Cl) and nitrogen were evaluated scorpions on a diet of adult Tenebrio molitor beetles. Scorpions were offered a weighed beetle every second day for 2 months. The feeding pellets, faeces and 256 S. D. YOKOTA urine were collected daily, dried and weighed. Uneaten beetles were discarded Faeces and urine were distinguished visually by colour, white excrement being con- sidered urine and dark excrement being considered faeces (Said, 1961; Yokota & Shoemaker, 1981). Electrolytes were extracted from samples with 0-1 N nitric acid. Sodium and potassium were analysed by flame photometry and chloride was deter- mined with a chloridometer (Buchler-Cotlove). The nitrogen content of the feeding pellets was determined by the Dumas method with a nitrogen analyser (Coleman). The nitrogen content of the faeces and the urine was determined with a modified Kjeldahl method (Jaenicke, 1974) after dissolving the samples in 0-1 N sodium hydroxide.
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