Influence of hyperbaric stress on metabolic reactions of the organism of divers, prevention

Research of metabolic rates is conducted in a blood of the subjects whose professional work is connected with the systematic implementation of diving on the large and medium depths. For restoration of working capacity and preservation of health of divers was carried pharmacological prophylaxis with supplementation "Kalifen", which is the total polyphenol complex isolated from viburnum (Viburnum sargentii Koehne). The blood plasma of divers, investigated after a dive, there was hypertriglyceridemia and hypercholesterolemia, the suppression of esterified liver function, an imbalance of fractional content of phospholipids, the strain of antioxidant defense of an organism and increased lipid peroxidation. Preventive administration supplementation "Kalifen" for 2 months before the dive will help remove metabolic disorders caused by hyperbaric factors, and also to store up of anabolic reserves and increase body resistance level in extreme conditions.     

The influence of body size on the diving behaviour and physiology of the bimodally respiring turtle, Elseya albagula

In aquatic vertebrates that acquire oxygen aerially dive duration scales positively with body mass, i.e. larger animals can dive for longer periods, however in bimodally respiring animals the relationship between dive duration and body mass is unclear. In this study we investigated the relationships between body size, aquatic respiration, and dive duration in the bimodally respiring turtle, Elseya albagula. Under normoxic conditions, dive duration was found to be independent of body mass. The dive durations of smaller turtles were equivalent to that of larger individuals despite their relatively smaller oxygen stores and higher mass specific metabolic rates. Smaller turtles were able to increase their dive duration through the use of aquatic respiration. Smaller turtles had a relatively higher cloacal bursae surface area than larger turtles, which allowed them to extract a relatively larger amount of oxygen from the water. By removing the ability to respire aquatically (hypoxic conditions), the dive duration of the smaller turtles significantly decreased restoring the normal positive relationship between body size and dive duration that is seen in other air-breathing vertebrates.     

Urinary C-peptide as a method for monitoring body mass changes in captive bonobos (Pan paniscus)

In recent years methodological improvements have allowed for more precise estimates of nutrient intake in wild primates. However, estimates of energetic condition have remained relatively imprecise due to the difficulties of estimating digestive efficiency and energy expenditure in these animals. In the absence of a reliable intake-expenditure calculation, a method is needed that directly links changes in energetic condition, such as body mass, to physiological changes that can be detected via markers in body excretions such as urine or feces. One promising marker is C-peptide, a metabolic byproduct of insulin synthesis. Here we present the results of a food restriction experiment carried out in a group of captive bonobos (Pan paniscus). We measured changes in food availability and body mass and determined urinary C-peptide levels with the help of a time-resolved fluoroimmunoassay routinely used for measuring C-peptide in human blood. Urinary C-peptide levels decreased during a period of food restriction and increased again when food availability was continuously increased. During this refeeding phase an increase in body mass was significantly correlated with an increase in urinary C-peptide levels. Our results suggest that urinary C-peptide levels are an accurate indicator of individual energy balance. In conclusion, measuring C-peptide in urine is a promising method to quantify the energetic condition of wild apes.     

Physiological and behavioral determinants of the aerobic dive limit in Weddell seal (Leptonychotes weddellii) Pups

The aerobic dive limit, as defined by an increase in plasma lactate levels following dives, has to date only been determined in adult and juvenile Weddell seals (Leptonychotes weddellii). However, theoretical aerobic dive limits based on calculated total body oxygen stores, estimated metabolic rates, and dive duration frequencies have been published for several species. Using data collected over the past 3 years in McMurdo Sound. Antarctica, the aerobic dive limit of Weddell seal pups was determined by both the physiological and modeling methods. Time-depth diving recorders deployed on 36 pups between 2 and 14 weeks of age allowed the aerobic dive limit to be predicted from duration-frequency histograms. The aerobic dive limit was also calculated from estimates of total body oxygen stores and predicted diving metabolic rates. Finally, these two estimates were compared with aerobic dive limits determined from post-dive lactate levels in three pups between 5 and 7 weeks old. The aerobic dive limits of pups increased with age, but pup aerobic dive limits were still significantly shorter than those of yearlings and adults. In addition, the aerobic dive limits determined by the three methods were not equivalent for pups, yearlings, or adults, and indicate that care should be taken when modeling methods are used to estimate the aerobic dive limit in other species. Changes in hematocrit, plasma glucose, and plasma lactate levels during and between rest, diving, and recovery in pups were compared to known values for juveniles and adults. Plasma metabolite levels were more highly regulated in older pups, and together with the increasing aerobic dive limit, suggest that Weddell seal pups are not refined divers until after they are weaned, and that their diving ability continues to develop over several years.     

Blood Oxygen Depletion Is Independent of Dive Function in a Deep Diving Vertebrate,the Northern Elephant Seal

Although energetics is fundamental to animal ecology, traditional methods of determining metabolic rate are neither direct nor instantaneous. Recently, continuous blood oxygen (O₂) measurements were used to assess energy expenditure in diving elephant seals (Mirounga angustirostris), demonstrating that an exceptional hypoxemic tolerance and exquisite management of blood O₂ stores underlie the extraordinary diving capability of this consummate diver. As the detailed relationship of energy expenditure and dive behavior remains unknown, we integrated behavior, ecology, and physiology to characterize the costs of different types of dives of elephant seals. Elephant seal dive profiles were analyzed and O₂ utilization was classified according to dive type (overall function of dive: transit, foraging, food processing/rest). This is the first account linking behavior at this level with in vivo blood O₂ measurements in an animal freely diving at sea, allowing us to assess patterns of O₂ utilization and energy expenditure between various behaviors and activities in an animal in the wild. In routine dives of elephant seals, the blood O₂ store was significantly depleted to a similar range irrespective of dive function, suggesting that all dive types have equal costs in terms of blood O₂ depletion. Here, we present the first physiological evidence that all dive types have similarly high blood O₂ demands, supporting an energy balance strategy achieved by devoting one major task to a given dive, thereby separating dive functions into distinct dive types. This strategy may optimize O₂ store utilization and recovery, consequently maximizing time underwater and allowing these animals to take full advantage of their underwater resources. This approach may be important to optimizing energy expenditure throughout a dive bout or at-sea foraging trip and is well suited to the lifestyle of an elephant seal, which spends > 90% of its time at sea submerged making diving its most “natural” state.     

Body oxygen stores, aerobic dive limits, and the diving abilities of juvenile and adult muskrats (Ondatra zibethicus)

Intraspecific variability in body oxygen reserves, muscle buffering capacity, diving metabolic rate, and diving behavior were examined in recently captured juvenile and adult muskrats. Allometric scaling exponents for lung (b=1.04), blood (b=0.91), and total body oxygen storage capacity (b=1.09) did not differ from unity. The concentration of skeletal muscle myoglobin scaled positively with mass in 254-600-g juveniles (b=1.63) but was mass-independent in larger individuals. Scaling exponents for diving metabolic rate and calculated aerobic dive limit (ADL) were 0.74 and 0.37, respectively. Contrary to allometric predictions, we found no evidence that the diving abilities of muskrats increased with age or body size. Juveniles aged 1-2 mo exhibited similar dive times but dove more frequently than summer-caught adults. Average and cumulative dive times and dive&rcolon;surface ratios were highest for fall- and winter-caught muskrats. Total body oxygen reserves were greatest in winter, mainly due to an increase in blood oxygen storage capacity. The buffering capacity of the hind limb swimming muscles also was highest in winter-caught animals. Several behavioral indicators of dive performance, including average and maximum duration of voluntary dives, varied positively with blood hemoglobin and muscle myoglobin concentration of muskrats. However, none of the behavioral measures were strongly correlated with the total body oxygen reserves or ADLs derived for these same individuals.     

In situ metabolic budget for the calanoid copepod Acartia clausi in a tropical brackish water lagoon (Ebrié Lagoon,Ivory Coast)

Simultaneous measurements of respiration, excretion and production rates were carried out several times over a year period at five representative stations of the Ebrié Lagoon. Assuming a constant assimilation efficiency rate of 69.4%, we derived metabolic budgets for carbon, nitrogen and phosphorus. Daily specific ingestion rates calculated were rather generally high, and ranged between 54 and 159% of body carbon, between 26 and 102% of body nitrogen and between 108 and 307% of body phosphorus. Regional and seasonal variations depended mainly upon variations in trophic conditions. Curvilinear relationships between ingestion production, or net production efficiency K2, and food concentration (as chlorophyll-a + phaeopigments) showed that food could have been a limiting factor. Furthermore, K2 were low when compared with data from the literature (mean of 21% in carbon, 39% in nitrogen and 11% in phosphorus).Complementary laboratory experiments carried out on adults fed with enriched natural particles or algal cultures (Tetraselmis sp. or Dunaliella sp.) showed similar production (egg-production) vs food concentration curvilinear relationships as in the field. However, considerably higher maximal ingestion and production rates were obtained for animals fed algal cultures suggesting that optima for food acquisition and transformation were not reached in field conditions.Consequently, A. clausi, which represents more than 50% of the zooplankton biomass, appears to be rather inefficient in transforming the abundant particulate organic matter produced in the lagoon. This results from its high level of metabolic expenditure through respiration or excretion (about 50% of ingestion in terms of carbon) and from the small size and poor trophic value of food particles (high percentage of detritus).     

Tufted ducks Aythya fuligula do not control buoyancy during diving

Work against buoyancy during submergence is a large component of the energy costs for shallow diving ducks. For penguins, buoyancy is less of a problem, however they still seem to trade‐off levels of oxygen stores against the costs and benefits of buoyant force during descent and ascent. This trade‐off is presumably achieved by increasing air sac volume and hence pre‐dive buoyancy (B_pre) when diving deeper. Tufted ducks, Aythya fuligula, almost always dive with nearly full oxygen stores so these cannot be increased. However, the high natural buoyancy of tufted ducks guarantees a passive ascent, so they might be expected to decrease B_pre before particularly deep, long dives to reduce the energy costs of diving. Body heat lost to the water can also be a cause of substantial energy expenditure during a dive, both through dissipation to the ambient environment and through the heating of ingested food and water. Thus dive depth (d_d), duration and food type can influence how much heat energy is lost during a dive. The present study investigated the relationship between certain physiological and behavioural adjustments by tufted ducks to d_d and food type. Changes in B_pre, deep body temperature (T_b) and dive time budgeting of four ducks were measured when diving to two different depths (1.5 and 5.7 m), and for two types of food (mussels and mealworms). The hypothesis was that in tufted ducks, B_pre decreases as d_d increases. The ducks did not change B_pre in response to different diving depths, and thus the hypothesis was rejected. T_b was largely unaffected by dives to either depth. However, diving behaviour changed at the greater d_d, including an increase in dive duration and vertical descent speed. Behaviour also changed depending on the food type, including an increase in foraging duration and vertical descent speed when mussels were present. Behavioural changes seem to represent the major adjustment made by tufted ducks in response to changes in their diving environment.     

Food shortage affects flight migration of the giant water bug Lethocerus deyrolli in the prewintering season

The endangered giant water bug Lethocerus deyrolli (Vuillefroy) is frequently attracted in large numbers to artificial lights in Japan. To examine factors enhancing flight migration for L. deyrolli, we carried out field work in western Hyogo Prefecture, central Japan, in September during the nonreproductive and prewintering season. The body weight of specimens collected under flight migration (flight bugs) was significantly less than that of those collected in ponds (pond bugs). A field experiment using open cages in a rice paddy field was carried out with two treatments, with and without a food supply. The remaining rate of L. deyrolli for the food present treatment was significantly higher than that for the food absent treatment for the first two days. These results suggest that L. deyrolli would fly in search of food when the food supply of the present habitat becomes unsuitable.     

Solid State 13C-NMR Analysis of Cell Wall Components of Fusarium oxysporum

An abdominal fat accumulation complicated by high blood triglycerides is regarded as a risk factor of metabolic syndrome. Feeding powdered nacre, mother of pearl, from Pinctada maxima, resulted in reduced body weight, visceral fat amount, and blood triglyceride level without influencing the food intake, body length, or amount of muscular tissue, suggesting that nacre powder specifically could decrease visceral fat.     

Behavioural and metabolic responses of the Antarctic sea star Odontaster validus to food stimuli of different concentration

Experiments were carried out to study the behavioural and metabolic responses of the Antarctic sea star, Odontaster validus, to different concentration of food signals (glutamic acid, aspartic acid, glicyne, alanine, tyrosine and a mixture of 11 amino acids). Stimulus concentration increase caused a rise in both the percentage of reacting animals and in the reaction intensity. At low stimulus concentration, O. validus reaction consisted of tube foot waving, and only high concentrations elicited a complicated sequence of several types of behaviour. The metabolic rate of small O. validus was more sensitive to chemical stimuli than of large individuals, with small sea stars reacting even to low signal concentration. It can be speculated that small body size can be a factor restricting sea star capabilities and influencing its reaction to chemical signals.     

Diving behavior in a free‐living,semi‐aquatic herbivore,the Eurasian beaver Castor fiber

Semi‐aquatic mammals have secondarily returned to the aquatic environment, although they spend a major part of their life operating in air. Moving both on land, as well as in, and under water is challenging because such species are considered to be imperfectly adapted to both environments. We deployed accelerometers combined with a depth sensor to study the diving behavior of 12 free‐living Eurasian beavers Castor fiber in southeast Norway between 2009 and 2011 to examine the extent to which beavers conformed with mass‐dependent dive capacities, expecting them to be poorer than wholly aquatic species. Dives were generally shallow (<1 m) and of short duration (<30 s), suggesting that the majority of dives were aerobic. Dive parameters such as maximum diving depth, dive duration, and bottom phase duration were related to the effort during different dive phases and the maximum depth reached. During the descent, mean vectorial dynamic body acceleration (VeDBA—a proxy for movement power) was highest near the surface, probably due to increased upthrust linked to fur‐ and lung‐associated air. Inconsistently though, mean VeDBA underwater was highest during the ascent when this air would be expected to help drive the animals back to the surface. Higher movement costs during ascents may arise from transporting materials up, the air bubbling out of the fur, and/or the animals’ exhaling during the bottom phase of the dive. In a manner similar to other homeotherms, beavers extended both dive and bottom phase durations with diving depth. Deeper dives tended to have a longer bottom phase, although its duration was shortened with increased VeDBA during the bottom phase. Water temperature did not affect diving behavior. Overall, the beavers’ dive profile (depth, duration) was similar to other semi‐aquatic freshwater divers. However, beavers dived for only 2.8% of their active time, presumably because they do not rely on diving for food acquisition.     

Heart rate and plasma lactate responses during submerged swimming and trained diving in California sea lions, Zalophus californianus

California sea lions, Zalophus californianus, were trained to elicit maximum voluntary breath holds during stationary underwater targeting, submerged swimming, and trained diving. Lowest heart rate during rest periods was 57 bpm. The heart rate profiles in all three protocols were dominated by a bradycardia of 20–50 bpm, and demonstrated that otariid diving heart rates were at or below resting heart rate. Venous blood samples were collected after submerged swimming periods of 1–3 min. Plasma lactate began to increase only after 2.3-min submersions. This rise in lactate and our inability to train sea lions to dive or swim submerged for periods longer than 3 min lead us to conclude that an aerobic limit had been reached. Due to the similarity of heart rate responses and swimming velocities recorded during submerged swimming and trained diving, this 2.3-min limit should approximate the aerobic dive limit in these 40-kg sea lions. Total body O₂ stores, based on measurements of blood and muscle O₂ stores in these animals, and prior lung O₂ store analyses, were 37–43 ml O₂ kg⁻¹. The aerobic dive limit, calculated with these O₂ stores and prior measurements of at-sea metabolic rates of sea lions, is 1.8–2 min, similar to that measured by the change in post-submersion lactate concentration. Accepted: 7 July 1996     

BUFFERING CAPACITY OF THE LOCOMOTOR MUSCLE IN CETACEANS: CORRELATES WITH POSTPARTUM DEVELOPMENT, DIVE DURATION, AND SWIM PERFORMANCE

Skeletal muscles of marine mammals must support the metabolic demands of exercise during periods of reduced blood flow associated with the dive response. Enhanced muscle buffering could support anaerobic metabolic processes during apnea, yet this has not been fully investigated in cetaceans. To assess the importance of this adaptation in the diving and swimming performance of cetaceans, muscle buffering capacity due to non-bicarbonate buffers was measured in the longissimus dorsi of ten species of odontocete and one mysticete. Immature specimens from a subset of these species were studied to assess developmental trends. Fetal and neonatal cetaceans have low buffering capacities (range: 34.8–53.9 slykes) that are within the range measured for terrestrial mammals. A lengthy developmental period, independent of muscle myoglobin postnatal development, is required before adult levels are attained. Adult cetacean buffering capacities (range: 63.7–94.5 slykes) are among the highest values recorded for mammals. Cetacean species that demonstrate extremely long dive durations or high burst speed swimming tend to have greater buffering capacities. However, the wide range of body size across cetaceans may complicate these trends. Enhanced muscle buffering capacity may enable small-bodied species to extend breath-hold beyond short aerobic dive limits for foraging or predator evasion when necessary.     

Benefits of thermal acclimation in a tropical aquatic ectotherm,the Arafura filesnake, <Emphasis Type="Italic">Acrochordus arafurae</Emphasis>

The presumption that organisms benefit from thermal acclimation has been widely debated in the literature. The ability to thermally acclimate to offset temperature effects on physiological function is prevalent in ectotherms that are unable to thermoregulate year-round to maintain performance. In this study we examined the physiological and behavioural consequences of long-term exposure to different water temperatures in the aquatic snake Acrochordus arafurae. We hypothesised that long dives would benefit this species by reducing the likelihood of avian predation. To achieve longer dives at high temperatures, we predicted that thermal acclimation of A. arafurae would reduce metabolic rate and increase use of aquatic respiration. Acrochordus arafurae were held at 24 or 32°C for 3 months before dive duration and physiological factors were assessed (at both 24 and 32°C). Although filesnakes demonstrated thermal acclimation of metabolic rate, use of aquatic respiration was thermally independent and did not acclimate. Mean dive duration did not differ between the acclimation groups at either temperature; however, warm-acclimated animals increased maximum and modal dive duration, demonstrating a longer dive duration capacity. Our study established that A. arafurae is capable of thermal acclimation and this confers a benefit to the diving abilities of this snake.     

Ecological consequences of scaling of chew cycle duration and daily feeding time in Primates

Feeding systems and behaviors must evolve to satisfy the metabolic needs of organisms. This includes modifications to feeding systems as body size and metabolic needs change. Using our own data and data from the literature, we examine how size-related changes in metabolic needs are met by size-related changes in daily feeding time, chew cycle duration, volume of food processed per chew, and daily food volume intake in primates. Increases in chew cycle duration with body mass in haplorhine primates are described by a simple power function (cycle time α body mass^0.181). Daily feeding time increases with body mass when analyzed using raw data from the “tips” of the primate phylogenetic tree, but not when using phylogenetically independent contrasts. Whether or not daily feeding time remains constant or increases with body mass, isometry of ingested bite size and the slow rate of increase in chew cycle time with body size combine to allow daily ingested food volume to scale faster than predicted by metabolic rate. This positive allometry of daily ingested food volume may compensate for negative allometry of nutrient concentration in primate foods. Food material properties such as toughness and hardness have little impact on scaling of chew cycle durations, sequence durations, or numbers of chews in a sequence. Size-related changes in food processing abilities appear to accommodate size-related changes in food material properties, and primates may alter ingested bite sizes in order to minimize the impacts of food material properties on temporal variables such as chew cycle duration and chew sequence duration.     

Food Viscosity Influences Caloric Intake Compensation and Body Weight in Rats

Objective: To determine the effects of food viscosity on the ability of rats to compensate for calories in a dietary supplement. Research Methods and Procedures: In a series of four experiments, rats consumed dietary supplements equated for caloric and nutritive content but differing in viscosity. Experiments 1 to 3 examined the ability of the rats to compensate for the calories consumed in low‐ compared with high‐viscosity premeals by reducing intake of a subsequent test meal. Caloric compensation was assessed with a wide range of premeal viscosity levels and with two different non‐nutritive thickening agents. Experiment 4 assessed the effects of consuming daily a low‐viscosity compared with an equicaloric high‐viscosity dietary supplement on longer term body weight gain. Results: Consuming a lower viscosity premeal was followed by significantly more caloric intake (i.e., less caloric compensation) compared with consuming premeals with higher viscosity levels. This effect was not specific to one thickening agent. Furthermore, rats given a low‐viscosity supplement daily gained significantly more weight over a 10‐week period compared with rats given a high‐viscosity supplement. Discussion: The results of these experiments suggest that food viscosity may be an important determinant of short‐term caloric intake and longer term body weight gain.     

Diving through the thermal window: implications for a warming world

Population decline and a shift in the geographical distribution of some ectothermic animals have been attributed to climatic warming. Here, we show that rises in water temperature of a few degrees, while within the thermal window for locomotor performance, may be detrimental to diving behaviour in air-breathing ectotherms (turtles, crocodilians, marine iguanas, amphibians, snakes and lizards). Submergence times and internal and external body temperature were remotely recorded from freshwater crocodiles (Crocodylus johnstoni) while they free-ranged throughout their natural habitat in summer and winter. During summer, the crocodiles'' mean body temperature was 5.2 ± 0.1°C higher than in winter and the largest proportion of total dive time was composed of dive durations approximately 15 min less than in winter. Diving beyond 40 min during summer required the crocodiles to exponentially increase the time they spent on the surface after the dive, presumably to clear anaerobic debt. The relationship was not as significant in winter, even though a greater proportion of dives were of a longer duration, suggesting that diving lactate threshold (DLT) was reduced in summer compared with winter. Additional evidence for a reduced DLT in summer was derived from the stronger influence body mass exerted upon dive duration, compared to winter. The results demonstrate that the higher summer body temperature increased oxygen demand during the dive, implying that thermal acclimatization of the diving metabolic rate was inadequate. If the study findings are common among air-breathing diving ectotherms, then long-term warming of the aquatic environment may be detrimental to behavioural function and survivorship.     

The diving behaviour of the Manx Shearwater Puffinus puffinus

The diving capabilities of the Procellariformes remain the least understood component of avian diving physiology. Due to their relatively small size, shearwaters may have high oxygen consumption rates during diving relative to their available oxygen stores. Dive performance in this group should be strongly limited by the trade‐off between oxygen consumption and oxygen stores, and shearwaters could be a good model group for testing predictions of dive theory. Many earlier measurements of shearwater dive behaviour relied on observations from the surface or potentially biased technology, and it is only recently that diving behaviour has been observed using electronic recorders for many of the clades within the family. The diving behaviour of Manx Shearwaters Puffinus puffinus breeding in Wales, UK, was studied on a large sample of birds using time–depth–temperature recorders deployed on chick‐rearing shearwaters in July and August over 3 years (2009–2011). Light availability apparently limited diving as dives only occurred between 04:00 and 19:00 h GMT. All individuals routinely dived deeper than traditionally assumed, to a mean maximum depth of 31 m and occasionally down to nearly 55 m. We compiled all available data for a comparison of the dive depth across shearwater species. There was a positive allometric relationship between maximum dive depth and body mass across Puffinus and Ardenna shearwater species, as expected, but only if samples of fewer than two individuals were excluded. The large intra‐specific range in maximum dive depth in our study illustrates that apparent diversity in diving performance across species must be interpreted cautiously.     

Seasonal changes in the oxygen storage capacity and aerobic dive limits of the muskrat (Ondatra zibethicus)

Summary The oxygen storage capacity and partitioning of body oxygen reserves were compared in summer-and winter-acclimatized muskrats (Ondatra zibethicus). Blood volume, blood oxygen capacity, and skeletal muscle myoglobin content were higher in December than in July (P<0.02). Total lung capacity increased only slightly in winter (P>0.05). The oxygen storage capacity of a diving muskrat was calculated at 25.2 ml O₂ STPD · kg^-1 in July, compared to 35.7 ml O₂ STPD · kg^-1 in December. Blood comprised the major storage compartment in both seasons, accounting for 57% and 65% of the total oxygen stores in summer and winter, respectively. Based on available oxygen stores and previous estimates of the cost of diving, the aerobic dive limit (ADL) increased from 40.9 s in July to 57.9 s in December. Concurrent behavioral studies suggested that most voluntary diving by muskrats is aerobic. However, the proportion of dives exceeding the calculated ADL of these animals was shown to vary with the context of the dive. Only 3.5% of all dives initiated by muskrats floating in the water exceeded their estimated ADL. Provision of a dry resting site and access to a submerged food source increased this proportion to 18–61%, depending on the underwater distance that foraging muskrats were required to swim. Serial dives exceeding the estimated ADL were not accompanied by extended postdive recovery periods.Abbreviations ADL acrobic dive limit - Hb hemoglobin - Hct hematocrit - Mb myoglobin - PaO₂ arterial O₂ tension - STPD standard temperature and pressure, dry