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Biomedical subjects

A S Blix

Publications and source records attributed to A S Blix.

At least 19 recordsLinked to original sources

Water flux and early signs of entrance into phase III of fasting in grey seal pups.

Body water loss and turnover rate were measured in fasting newly weaned grey seal (Halichoerus grypus) pups without access to water for 52 days, by use of bolus injections of deuterated water. Total body water (N) was 41.8 +/- 2.3 (SD)% of total body weight at day 3 and 44.5 +/- 5.9 (SD)% at day 51 of fasting (P greater than 0.05), while the water content of body core decreased from 73 to 61% over the same period. The average water efflux rate in the same period was 362 +/- 17 (SD) ml d-1, the reduction in N over the same period contributing 35% to this end. The biological half-time of deuterated water was 38.2 +/- 3.3 (SD) d. Plasma osmolality increased from a stable level of 329 +/- 11 (SD) mosmol kg-1 to 445 +/- 11 (SD) mosmol kg-1 after day 38 of fasting. Plasma urea was the major contributor to this increase, in spite of a steadily decreasing urine urea concentration (Nordøy et al. 1990) throughout the entire experimental period. It is concluded that grey seal pups endure 52 d of fasting without intake of water with a minor dehydration of body core, while they become hyperosmotic due to increased catabolism of protein after 38 d when sparing of the insulating blubber layer in preparation for life in cold water seems to occur.

Animals

Metabolic rates of minke whales (Balaenoptera acutorostrata) in cold water.

Body temperature, blubber thickness and lung capacity (Vc) were recorded in newly killed minke whales, while respiratory frequency (f) was determined in free-swimming animals. Mean deep (thoracic) body temperature was 34.7 +/- 0.8 (SD) degrees C (n = 14). Weighted mean core/blubber interface temperature in animals caught in 2.5-5.5 degrees C water was 28.8 +/- 1.7 degrees C (n = 8). The minimum average rate of sensible heat loss (HLs) was 3.81 +/- 0.53 (SD) W kgw-0.75 (n = 8) in animals with body masses (w) in the range of 1840 to 5740 kg, HLs being inversely proportional to w (HLs = -2.98 10(-4) w +4.89 W kgw-0.75 (n = 8, r2 = 0.73, P less than 0.01)). The average rate of respiratory heat loss (HLr) was 0.26 +/- 0.04 (SD) W kgw-0.75, regardless of w, in the same 8 animals. Total rates of heat loss (HL = HLr+HLs) in 2.5-5.5 degrees C water ranged between 3.40 and 4.87 W kgw-0.75, with an average of 4.06 +/- 0.52 (SD) W kgw-0.75 (n = 8). Estimates of oxygen consumption based on records of f and Ve, and data on oxygen extraction from other cetaceans, yielded a range of metabolic rates which compared nicely with the calculated HL values.

Adipose Tissue

Glucose and ketone body turnover in fasting grey seal pups.

Concentration and metabolic replacement (turnover) rate of glucose and ketone bodies were determined at intervals during a 52 day postweaning fast in five grey seal (Halichoerus grypus) pups, using bolus injections of radiotracers. Blood glucose was maintained at a high level throughout the fast, while beta-hydroxybutyrate increased 26 times from day 3 to day 37, whereafter it by and large was maintained. Glucose replacement rate decreased to 56% of the day 9 value at day 37 and dropped further to only 32% of the day 9 level at day 52 in two seals, while in another 2 seals it increased at this late stage. The average ketone body replacement rate ranged between 8.6 and 13.8 mumols min-1 kg-1, but did not change significantly (P greater than 0.05) during the fasting period. These results suggest a reduced gluconeogenesis from protein and increased production of ketone bodies, which may in part replace glucose as energy source during fasting.

Animals

Strategies of thermal protection in arctic animals.

Anatomical, physiological and behavioural adaptations to cold and lack of food combined with long periods of darkness are discussed. In large animals such adaptations comprise body size and insulation and controlled peripheral cooling in the legs and heat exchange in the nasal passages, whereby expiratory heat and water loss is minimized. In such animals grave thermal problems are incurred when the animal is forced to run. In some species the heat exchanger in the nose is operated in conjunction with a carotid rete for selective cooling of the brain. Small and poorly insulated mammals evade the brunt of the cold arctic winter in nests under the snow were convective heat loss is eliminated and ground heat creates a relatively comfortable micro-climate. Most newborn mammals are poorly insulated, but some survive birth, wet and miserable, at ambient temperatures down to -30 degrees C. Precocious forms depend heavily on NST in brown fat and skeletal muscle. Altricial forms are born very small and naked and are virtually ecto-thermic at birth. They depend on shelter and/or huddling, but survive a cooling of body core down to +2 degrees C.

Animals

Depressed metabolism and low protein catabolism in fasting grey seal pups.

Grey seal pups (Halichoerus grypus) were collected at the time of weaning (mid-October) and fasted for 52 days at thermoneutrality in separate cages. Body weight decreased exponentially, while metabolic rate dropped 45% from an average of 2.95 +/- 0.15 (SEM) W kg-1 at day 2 of fasting to a stable level of 1.62 +/- 0.06 (SEM) W kg-1 from day 10 to day 47 of fasting. Respiratory quotient was low, indicating extensive catabolism of triglycerides, while plasma cortisol was fairly stable at 110 +/- 8 (SEM) nmol l-1 throughout the fasting period. Daily urinary output decreased from 236 +/- 20 (SEM) ml day-1 at day 2 to a stable value of 87 +/- 6 (SEM) ml day-1 between days 8 and 50 of fasting. The urine was analysed for urea, uric acid, creatinine, ammonia, total nitrogen and osmolality. Urea was always the principal excretory end-product, amounting to between 70 and 80% of the total excreted nitrogen. The urine was moderately concentrated (range 770-1300 mosmol kg-1). Total excreted urinary nitrogen decreased by 68% from 3.7 +/- 0.7 (SEM) g day-1 to 1.2 +/- 0.4 (SEM) g day-1 between days 2 and 50. The urinary nitrogen was used to calculate the daily amount of protein being oxidized and its energy content was compared with the measured basal metabolic rate of individual animals. Approximately 6% of the energy expended by grey seal pups during the post-weaning fast is derived from oxidation of protein. It is concluded that a rapid depression of basal metabolic rate and extensive blubber catabolism enable grey seal pups to endure prolonged periods of fasting without any apparent signs of discomfort or stress.

Animals

Immobilization of Norwegian reindeer (Rangifer tarandus tarandus) and Svalbard Reindeer (R. t. platyrhynchus) with medetomidine and medetomidine-ketamine and reversal of immobilization with atipamezole.

The sedative action of medetomidine (-ketamine) was studied in 12 captive Norwegian semidomesticated reindeer (NR), including 4 newborn calves, and in 7 free-living Svalbard reindeer (SR). Medetomidine, with or without ketamine, caused effective, reliable immobilization in NR. Doses of 50-200 micrograms/kg medetomidine alone or 30-125 micrograms/kg medetomidine combined with greater than or equal to 300 micrograms/kg ketamine induced complete immobilization, good muscle relaxation and persistent, deep sedation with little respiratory depression in NR; SR required higher doses. Atipamezole successfully antagonized medetomidine (-ketamine) resulting in rapid and persistent reversal of immobilization in all cases (NR and SR). Both medetomidine and atipamezole had wide safety margins and no conspicuous lasting side effects after reversal.

Adrenergic alpha-Antagonists

Thermoregulatory control of expired air temperature in diving harp seals.

Expired air temperature (Tex), metabolic rate (MR), and skin (Ts) and body (Tb; rectal) temperatures were recorded in four or five young (1-2 yr) harp seals (Phoca groenlandica) in air [mean air temperature (Ta) = -30, -10, or 10 degrees C] and in water [mean water temperature (Tw) = 2.3 or 24.8 degrees C, with Ta = -30, -10, 0, or 10 degrees C]. Apparent lower critical temperature was below -10 degrees C in air. Above this Ta, mean MR was 5.85 W.kg-0.75 (2.23 W.kg-1), while mean MR was 12.56 W.kg-0.75 (4.69 W.kg-1) at Ta -30 degrees C. When seals were immersed in water of 2.3 degrees C, mean MR was 6.13 W.kg-0.75 (2.31 W.kg-1), regardless of Ta. At Ta -30, -10, and 10 degrees C, mean Tex in air were 9.5, 13.0, and 25.0 degrees C, respectively. The corresponding values for seals in water (Tw = 2.3 degrees C) were 8.0, 9.5, and 15.5 degrees C, respectively. The low Tex recorded at Ta -30 and -10 degrees C in air and at all Ta in water (Tw = 2.3 degrees C) suggests that heat was conserved by nasal heat exchange. At Ta 10 degrees C, mean Tex of seals in air was approximately 10 degrees C higher than mean Tex of seals in water (Tw = 2.3 degrees C). Furthermore, seals subjected to a Tw of 24.8 degrees C at Ta 0 degrees C had a mean Tex 10 degrees C higher than when subjected to Tw 2.3 degrees C at the same Ta. These observations suggest that Tex in seals is under thermoregulatory control. In a series of forced dives of up to 5-min duration Tex was found to be the same before and after the dive regardless of dive duration and Ta.

Animals

Seasonal changes in the cecal microflora of the high-arctic Svalbard reindeer (Rangifer tarandus platyrhynchus).

The dominant cecal bacteria in the high-arctic Svalbard reindeer were characterized, their population densities were estimated, and cecal pH was determined in summer, when food quality and availability is good, and in winter, when it is very poor. In summer the total culturable viable bacterial population was (8.9 +/- 5.3) X 10(8) cells ml-1, whereas in winter it was (1.5 +/- 0.7) X 10(8) cells ml-1, representing a decrease to 17% of the summer population density. Of the dominant species of cultured bacteria, Butyrivibrio fibrisolvens represented 23% in summer and 18% in winter. Streptococcus bovis represented 17% in summer and 5% in winter. Bacteroides ruminicola represented 10% in summer and 26% in winter. In summer and winter, respectively, the proportion of the viable population showing the following activities was as follows: fiber digestion, 36 and 48%; cellulolysis, 10 and 6%; xylanolysis, 33 and 48%; and starch utilization, 77 and 71%. The most abundant cellulolytic species in summer was Butyrivibrio fibrisolvens, representing 62% of the total cellulolytic population, and in winter it was Ruminococcus albus, representing 80% of the total cellulolytic population. The most abundant xylanolytic species in summer was Butyrivibrio fibrisolvens, and in winter it was Bacteroides ruminicola, representing 59 and 54% of the xylanolytic isolates in summer and winter, respectively. The cecal bacterial of the Svalbard reindeer have the ability to digest starch and the major structural carbohydrates of the diet that are not digested in the rumen. The cecum in these animals has the potential to contribute very substantially to the digestion of the available plant material in both summer and winter.

Animals

Selective cooling of the brain in reindeer.

Cineangiographic examination of reindeer exposed to local (hypothalamic) or general heating and cooling revealed that the angular oculi veins are constricted during cold stress but dilated during heat stress. Moreover, during heat stress a segment of the facial vein appeared to be occluded, causing the cold venous return from the nasal mucosa to be routed directly to the cavernous sinus for selective cooling of the brain. Histological examination of the vasoactive segment of the facial vein showed unusually thick longitudinal and circular layers of smooth muscle cells. Obstruction of angular oculi blood flow by clamping of the veins in the heat-stressed animal resulted in an immediate rise in brain temperature. When reindeer under heat stress shift from closed- to open-mouth panting, only the expiratory phase of the respiratory cycle takes place through the mouth, whereas inspiration through the nose is continued. In this way, cooling of the nasal mucosa and, hence, cooling of the brain, is maintained.

Animals

Nasal heat and water exchange in gray seals.

Metabolic rate (MR), expired air temperature (Tex), respiratory frequency (f), respiratory minute volume (V), and skin (Ts) and body (Tb) temperatures were recorded in three gray seals (Halichoerus grypus) at ambient air temperatures (Ta) between -40 and +20 degrees C. At Ta within the thermoneutral zone, MR averaged 3.7 W.kg-0.75, while mean V was 0.26 1.min-1.kg-0.75. At Ta below -11 degrees C [apparent lower critical temperature (Tlc)], both MR and V increased linearly with decreasing Ta. Average maximum MR (9.6 W.kg-0.75) and V (0.57 1.min-1.kg-0.75) were both recorded at Ta of -40 degrees C. Tex decreased with decreasing Ta to an average minimum value of 8 degrees C at Ta of -30 degrees C. The highest Tex recorded was 32 degrees C at Ta of +20 degrees C. At Ta of -20 degrees C, both total respiratory heat loss, with one exception, and respiratory evaporative water loss reached their lowest values. At this Ta, 66% of the heat and 80% of the water added to the inspired air were regained on expiration. We suggest that nasal heat exchange may be of considerable importance for thermal and water balance in many pinnipeds.

Algorithms

Food intake, feeding rhythm, and body mass regulation in Svalbard rock ptarmigan.

Food intake (FI), feeding activity (FA), and body mass (BM) were recorded continuously throughout a 13-mo period in Svalbard rock ptarmigan kept under natural conditions of light and ambient temperature at Svalbard (79 degrees N). FI was persistently high from March until August, including the period when daylight is continuous, whereas it was low from November until January, when it is permanently dark. From August until November, BM doubled, while FI dropped to one-third. BM fell rapidly from mid-November until April despite a doubling of FI from February until March. From August until mid-November and from February until mid-April FA occurred mainly during the light period of the day. From late November until February and from mid-April until August intermittent FA occurred continuously. It is suggested that the seasonal changes in BM are not determined by FI alone but depend heavily on seasonal changes in locomotor activity as reflected in FA.

Animals

Endocrine changes associated with fat deposition and mobilization in svalbard ptarmigan (Lagopus mutus hyperboreus).

Plasma concentrations of Triiodothyronine (T3), thyroxine (T4) and growth hormone (GH) have been measured in blood samples taken from Svalbard ptarmigans (Lagopus mutus hyperboreus), shot throughout 1 whole year at Svalbard (79 degrees N). Plasma T3 levels varied in a monophasic pattern with low levels in winter and a peak in August, whereas plasma T4 levels remained constant throughout the year. High plasma T3 levels coincide with molt and a large food intake while low plasma levels of T3 coincide with molt arrest and a low food intake. Plasma GH levels were highest in winter and lowest in May and September. The low plasma GH levels in early autumn coincide with elevated liver weights and maximum rate of fat deposition. High GH levels in midwinter coincide with low liver weights and the mobilization of fat stores. A possible relationship between molt, food intake, fat deposition/mobilization, and plasma levels of T3 and GH is discussed.

Adipose Tissue

Seasonal changes in T3, FT4, and cortisol in free-ranging Svalbard reindeer (Rangifer tarandus platyrhynchus).

Serum levels of 3,5,3'-triiodothyronine (T3), free thyroxine (FT4), and cortisol were determined for free-ranging Svalbard reindeer during winter (March), summer (June-July), and autumn (October). A total of 48 animals representing adult males, adult barren, pregnant, and lactating females, and calves were shot, and blood sampled from their hearts. T3 and FT4 were found to vary seasonally, the levels being lower in winter than in summer or autumn. Differences due to state of pregnancy, sex, or age were represented by low FT4 levels in lactating females (summer) and rutting males (autumn), and high T3 levels in young calves (summer). Serum cortisol levels were found to change seasonally, summer levels being higher than winter or autumn levels in all groups of animals. This finding suggests that glucocorticoids (cortisol) do not affect the deiodination from T4 to T3 in reindeer. The seasonal changes in T3 and FT4 in wild animals were similar to the changes in captive animals. It is therefore concluded that the changes in thyroidal hormone levels do not indicate changes in basal metabolic rate in this species.

Acclimatization

Orienting and freezing responses in incubating ptarmigan hens.

Behavior studies and telemetric recordings of heart and respiration rates were performed on five wild and two captive, incubating, willow ptarmigan hens (Lagopus lagopus lagopus) and on four wild, incubating, Svalbard ptarmigan hens (Lagopus mutus hyperboreus). Sounds and sight of approaching humans, egg predators, or dogs near the nests elicited behavior in the hens which we have interpreted as an orienting response (OR) followed by freezing behavior. During both types of behavioral responses, heart rate was reduced from 204 +/- 39 (mean +/- SE) to 119 +/- 26 beats per minute and respiration from 25 +/- 2 to 12 +/- 3 breaths per minute. In wild incubating willow ptarmigan, further approach led to tachycardia and increased respiration. Some birds maintained freezing behavior, while others became restless before flying off. Two of the four incubating Svalbard ptarmigan hens showed the OR followed by freezing behavior accompanied by decreased heart and respiration rates. The other two birds showed flight response, restless behavior accompanied by increased heart and respiration rates. Flight behavior was also typical for willow ptarmigan incubating in captivity. Repeated auditory provocation of incubating hens caused progressive decrement in behavioral and cardiac responses that is interpreted as habituation of the OR.

Animals

Seasonal changes in the relative importance of different avenues of heat loss in resting and running reindeer.

Exhaled air temperature (Te), respiratory frequency (f), and rectal temperature (Tb) were measured in two reindeer before, during and after running for 45 min at a constant speed of 9.2 km X hr-1 on a level treadmill at different ambient temperatures (Ta), in summer with fur insulation at a minimum and in winter with fur insulation at its prime. Before and immediately after the 45-min running period, skin temperature (Tsk) and radiative surface temperature (Tr) were also measured. Te, Tsk and Tr decreased at rest with decreasing Ta both summer and winter Tb was elevated during running except at low Ta in summer. During running Tsk, Te and f were higher in winter than in summer, while Tr was higher in summer than in winter. The changes in Tr and Tsk in response to running were more pronounced on the legs than on the trunk. It is suggested that in the running reindeer the body surface is an effective avenue of heat loss during summer, while heat dissipation by way of the respiratory tract attains increased importance in winter, when fur insulation is at its best.

Animals

Seasonal changes in lipogenesis and lipolysis in isolated adipocytes from Svalbard and Norwegian reindeer.

Arctic reindeer exhibit marked seasonal changes in fat deposition and mobilization. At intervals throughout the year, therefore, we have measured feed intake of both Svalbard (SR) and Norwegian reindeer (NR) together with the seasonal changes in size, lipogenic and lipolytic capacity of isolated adipocytes from both sub-species. Feed intake of both NR and SR was maximal in August, but declined thereafter, reaching minimum values in January (NR) and March (SR), 55 and 69% below the August value, respectively. NR and SR adipocyte volume changed in parallel and were reduced to the same extent (69%) from their maximum in August to their minimum in May. Adipocyte lipogenic capacity, measured as acetate incorporation into cellular lipid at saturated acetate concentrations, was lowest in January (NR adipocytes) and March (SR adipocytes), 92 and 90%, respectively, below the maximum values, which were obtained in August. Lipolytic capacity, measured as maximum adrenaline-stimulated glycerol release, was high in SR adipocytes from March through to October and in NR adipocytes from July through to January. Minimum lipolytic capacity, on the other hand, was found in January (SR adipocytes) and March (NR adipocytes). The present findings may be explained by alterations in lipogenic enzyme activity and in the lipolytic activation system.

Adipose Tissue

Some aspects of thermoregulation in newborn reindeer calves (Rangifer tarandus tarandus).

At birth reindeer calves often are exposed to sub-zero ambient temperatures (Ta) sometimes even combined with wind and precipitation. The resting metabolism was measured in three different age groups (1, 7 and 14 days old) at Ta's of -20, -5, 10 and 20 degrees C. Resting metabolism in the thermoneutral zone decreased from 5.1 W X kg-1 at day 1 and 7 to 4.8 W X kg-1 at day 14. At day 1 apparent lower critical temperature (Tlc) was 11 degrees C, while at day 7 it was 7.7 degrees C and at day 14 7.3 degrees C, but total body conductance continued to decrease below apparent Tlc. At Ta's of -5 and -20 degrees C total body conductance was: 0.77 and 0.72 W X degrees C-1 at day 1, 0.98 and 0.92 W X degrees C-1 at day 7, 1.08 and 0.91 W X degrees C-1 at day 14, respectively. Thermal conductance of pelt samples from the trunk was determined in vitro at different combinations of windspeed, Ta and wetness. The conductance of dry fur increased from 5.9 to 11.8 W X m-2 X degrees C-1 at a windspeed of 0 and 10 m X s-1, respectively, as compared to 28.7 W X m-2 X degrees C-1 when wetted without wind. Newborn reindeer calves seem to be heavily dependent on non-shivering thermogenesis in brown adipose tissue for their cold defence since deep body temperature in a calf subjected to propranolol infusion when exposed to a Ta of -25 degrees C in combination with a 10 m X s-1 windspeed increased its cooling rate five times.

Adipose Tissue