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Biomedical subjects

L P Folkow

Publications and source records attributed to L P Folkow.

At least 19 recordsLinked to original sources

Cutaneous heat flux models do not reliably predict metabolic rates of marine mammals.

Heat flux models have been used to predict metabolic rates of marine mammals, generally by estimating conductive heat transfer through their blubber layer. Recently, Kvadsheim et al. (1997) found that such models tend to overestimate metabolic rates, and that such errors probably result from the asymmetrical distribution of blubber. This problem may be avoided if reliable estimates of heat flux through the skin of the animals are obtained by using models that combine calculations of conductive heat flux through the skin and fur, and convective heat flux from the surface of the animal to the environment. We evaluated this approach based on simultaneous measurements of metabolic rates and of input parameters necessary for heat flux calculations, as obtained from four harp seals (Phoca groenlandica) resting in cold water. Heat flux estimates were made using two free convection models (double-flat-plate and cylindrical geometry) and one forced convection model (single-flat-plate geometry). We found that heat flux estimates generally underestimated metabolic rates, on average by 26-58%, and that small variations in input parameters caused large variations in these estimates. We conclude that cutaneous heat flux models are too inaccurate and sensitive to small errors in input parameters to provide reliable estimates of metabolic rates of marine mammals.

Animals↗

On the direction and velocity of blood flow in the extradural intravertebral vein of harp seals (Phoca groenlandica) during simulated diving.

Ronald et al. (1977) suggested that blood flow in the caudal/lumbar sections of the extradural intravertebral vein (EIV) of seals changes direction from running towards the head before diving, to the opposite during diving. The possible advantage would be that the oxygen-depleted venous effluent from the brain is routed via the EIV to the posterior parts of the hepatic sinuses and the inferior caval vein and, hence, is prevented from mixing with the more oxygen-rich venous blood in their anterior parts. We have re-examined this hypothesis by use of Doppler flowmetry. A catheter-tip flow probe was introduced into the EIV of two similar-sized juvenile harp seals, and flow direction and rate determined before, during and after simulated dives lasting for 5 min, at three positions (caudal, lumbar and thoracic) along the EIV. Regardless of probe position, blood was mainly flowing towards the head in 11 of 13 experiments prior to diving, in 8 of 13 experiments during diving and in 11 of 13 experiments during recovery after diving (and away from the head in the remaining experiments). Flow direction was most variable in the caudal position. Mean blood velocity in the EIV was substantially lower during diving (0.10 +/- 0.22 cm s-1 (n=5) in thoracic position) than in the pre-dive (3.98 +/- 3.32 cm s-1 [n=5]) and post-dive (5.75 +/- 4.07 cm s-1 [n=5]) situations. Thus, the direction and rate of flow in the EIV was variable, particularly during diving, as is to be expected in a system of anastomosing, valveless veins. We conclude that the hypothesis of Ronald et al. (1977) most likely is false.

Animals↗

Panting in reindeer (Rangifer tarandus).

Two winter-insulated Norwegian reindeer (Rangifer tarandus tarandus) were exposed to air temperatures of 10, 20, 30, and 38 degrees C while standing at rest in a climatic chamber. The direction of airflow through nose and mouth, and the total and the nasal minute volumes, respectively, were determined during both closed- and open-mouth panting. The animals alternated between closed- and open-mouth panting, but the proportion of open-mouth panting increased with increasing heat load. The shifts from closed- to open-mouth panting were abrupt and always associated with a rise in respiratory frequency and respiratory minute volume. During open-mouth panting, the direction of airflow was bidirectional in both nose and mouth, but only 2.4 +/- (SD) 1.1% of the air was routed through the nose. Estimates suggest that the potential for selective brain cooling is markedly reduced during open-mouth panting in reindeer as a consequence of this airflow pattern.

Animals↗

Simulations of the effect of currently used grenade harpoons for the killing of whales using a pig-model.

Physical model experiments, as well as simulations of the effects of grenade harpooning on anaesthetized pigs fully immersed in water suggest that the shock effect of the blast from the currently used grenades is relatively minor. Also the animals are not stunned to death, but loose consciousness and subsequently die from hemorrhage. Survival time is therefore very short if the animals are hit in the thorax, and is likely to be further reduced if the charge which is currently used is increased, or, even better, if shrapnel (fragment scattering) grenades are used instead of blast grenades.

Animals↗

Effects of adrenergic and cholinergic drugs on splenic arteries and veins from hooded seals (Cystophora cristata).

Isolated ring preparations of arteries and veins from hooded seal spleens were subjected in vitro to adrenaline (A), noradrenaline (NA), isoprenaline (Iso), and acetylcholine (ACh), alone or in combination with the blockers phentolamine (Phe), propranolol (Pro), and atropine (Atr). Both arteries and veins constricted in response to A (the estimated effective dose required for half-maximal response (ED50) was 3.3 and 0.2 microM, for arteries and veins, respectively) and NA (estimated ED50 was 1.5 and 0.6 microM, for arteries and veins, respectively), but these effects were abolished when the drugs were given in combination with the alpha-adrenoceptor blocker Phe. The responses of arteries and veins to ACh and the beta-adrenoceptor agonist Iso were minor and inconsistent, and were completely abolished when combined with their respective blockers (Atr and Pro, respectively). The ED50 for both A and NA are quite high in relation to normal plasma levels of A and NA in seals. This implies that these vessels (and, hence, the supply of blood to the spleen) primarily are subjected to neurogenic, rather than humoral physiological control.

Adrenergic Agents↗

Changes in fibrinolytic activity in diving grey seals.

In order to test the hypothesis that enhanced fibrinolytic activity is a factor which prevents the blood of diving seals from clotting, we instrumented two female grey seals (Halichoerus grypus) with subcutaneous electrodes for measurements of heart rate (HR) and an extradural intravertebral venous catheter for collection of blood samples before, during and after simulated dives of 10 min duration. Blood samples were used for in vitro determination of clot lysis time (CLT), which is a measure of the level of fibrinolytic activity, and for analyses of plasma levels of cortisol, noradrenaline and adrenaline (A). The seals displayed profound diving bradycardia indicative of a substantial reduction in blood flow rates (pre-dive HR: 78 (63-98) bpm; dive HR: 8 (7-10) bpm; (median (range); n = 2)) and elevated catecholamine levels (pre-dive A: 121 (98-184) pg.ml-1; peak dive/post-dive A: 3510 (447-6181) pg.ml-1), both of which are factors which promote blood coagulation. Nevertheless, we found that CLT always increased in connection with diving (pre-dive CLT: 436 (356-568) min; peak CLT during diving: 1380 (640-1800) min), which implies a reduced, rather than enhanced, fibrinolytic activity in this situation. These results show that enhanced fibrinolytic activity is not part of the defence system which prevents fatal clotting from occurring in diving grey seals.

Animals↗

Blubber and flipper heat transfer in harp seals.

The trunk of marine mammals is encased in a blubber layer which provides thermal insulation that can be changed by circulatory adjustments. The extremities, on the other hand, are poorly insulated but have vascular arrangements constructed for prevention or promotion of heat loss depending on the thermal state of the animal. We have studied the importance of different body parts as sites for heat dissipation and also assessed the effect of circulatory adjustments on heat transfer through blubber, by combining direct measurements of heat flux from the flippers and trunk with simultaneous recordings of temperature gradients through the blubber and metabolic rates of harp seals (Phoca groenlandica) subjected to water temperatures between 1 and 24 degrees C. We also determined the thermal conductivity of blubber samples from the same animals after death, and compared this with the insulative properties of live blubber. At the lowest water temperatures, the insulative properties of live blubber were similar to those of dead blubber, and heat loss from the flippers only accounted for 2-6% of the metabolic heat production. As heat load increased with increasing water temperatures, the fraction of heat lost from the flippers increased, to 19-48% at 24 degrees C, while the fraction lost from the trunk decreased, despite an increase in the convective (circulatory) heat transfer through the blubber layer.

Adipose Tissue↗

Volume capacity and contraction control of the seal spleen.

Volume changes in the spleens of hooded seals (Cystophora cristata) and harp seals (Phoca groenlandica) were measured plethysmographically in vitro in response to epinephrine, norepinephrine, isoprenaline, phentolamine, and acetylcholine. Dilated spleens contracted forcefully within 1-3 min of alpha-adrenoceptor activation with 1.0-5.0 micrograms epinephrine/kg body mass, whereas stimulation of beta-adrenoceptors and cholinergic receptors had little effect. The mass of dilated hooded seal spleens corresponded to 2-4% (n = 7) of body mass, with volume (V; ml) relating to body mass (M; kg) as follows: V = 12.0M + 910 (r2 = 0.96, n = 4). Thus the spleen of a 250-kg hooded seal maximally expels 3.9 liters, or 13%, of its estimated total blood volume. Average hematocrit in splenic venous outflow from dilated spleens was 90 +/- 3% (n = 3) in hooded seals and 85% (n = 2) in harp seals. From these data we have estimated that the aerobic diving limit of a 250-kg hooded seal increases only 105 s, at the most, if complete emptying of the spleen occurs during diving, while the corresponding estimate for a 112-kg harp seal is 80 s.

Animals↗

Pineal and thyroid functions in newborn seals.

Daily variations of pineal and plasma melatonin and plasma thyroid hormones were measured in harp seals (Phoca groenlandica), grey seals (Halichoerus grypus), and hooded seals (Cystophora cristata), ranging in age from newborn to 14 days. In newborn harp seals the mean mass of the pineal gland was 273 mg (+/- 45 SEM, n = 11), containing 49 ng (median) melatonin. In newborn, 4- and 10-day-old grey seals, the pineal mass was similar, weighing on average 337 mg (+/- 74, n = 6) and containing 90 ng melatonin. Two newborn hooded seal pups had pineals weighing 520 and 1289 mg, with 254 and 7600 ng melatonin, respectively. There were no day-night differences in the pineal contents of melatonin or in the number of pineal beta-adrenergic receptors measured in newborn harp seals, and, in newborn, 4- and 10-day-old grey seals, there were no day-night or age differences in pineal melatonin content. Plasma melatonin levels were 10 times higher in newborn seals than in two 10-day-old grey seals and one 14-day-old harp seal pup. In all seal pups, the levels exhibited a 24-hr rhythmicity, with increasing night- and decreasing daytime concentrations. Plasma levels of thyroxine (T4) and triiodothyronine (T3) were generally higher in newborn seals than in 10- and 14-day-old seals or in adult females. There was no apparent 24-hr rhythmicity, but the thyroid hormone levels generally declined throughout each sampling sequence. High pineal and thyroid activities may play a thermoregulatory role in newborn seals, but the results do not indicate a stimulatory action of melatonin in the peripheral conversion of T4 to T3. It is speculated that the large and active pineal gland, particularly in newborn seals, may be related to aspects of their diving habit.

Aging↗

Daily energy expenditure in free living minke whales.

Six minke whales (Balaenoptera acutorostrata) were instrumented with VHF-radio transmitters and four with sonic speed-depth transmitters off the west coast of northern Norway and Svalbard and followed within view for up to 24 h. During such periods their respiratory rate was continuously recorded and their energy expenditure estimated according to Folkow & Blix (1992) at different swimming speeds and types of activity. We found that cost of swimming is remarkably low in these large animals and that their estimated daily energy expenditure on average only amounts to 80 kJ kg-1 day-1.

Animals↗

Nasal heat and water exchange is not an effector mechanism for water balance regulation in grey seals.

Phocid seals may effectively restrict respiratory heat and water loss by nasal heat and water exchange (NHE), and respiratory heat loss is, in fact, subject to thermoregulatory control. We have investigated whether phocid seals also control NHE and respiratory water loss to regulate water balance. Three resting juvenile female grey seals (Halichoerus grypus) were subjected to: (i) 5 days of food and water deprivation, (ii) intravenous infusion of 1000 ml of a hypersomotic (930 mM) solution of the diuretic mannitol, and (iii) oral injection of 1500 ml distilled water. During these experiments in air of 0 degree C, expired air temperature (T(ex)) and respiratory frequency (f) were recorded, and urine and blood samples collected. The results were compared with results from control experiments. Five days of food and water deprivation caused an average 10.5% and 20.8% increase in plasma (PO) and urine (UO) osmolality, respectively. Mannitol infusion induced excessive diuresis and caused an average 2.45% reduction of the estimated body water pool. Water loading caused an average 4.5% and 60% reduction in PO and UO, respectively, while urine production increased by 365%, on average. However, in no case did either T(ex) or f change significantly from mean control levels of 22.4 (range: 20.7-25.2) degrees C and 7.3 (range: 6.6-8.4) breaths min-1, respectively. Thus, water balance disturbances that initiate renal compensatory mechanisms fail to affect NHE in grey seals. This suggests that control of NHE is not an effector mechanism for regulation of water balance in grey seals.

Administration, Oral↗

Distribution and vasomotor effects of neuropeptides in angular oculi and facial veins of reindeer.

1. The vasomotor responses to neuropeptides of the angular oculi and facial veins of reindeer were examined in vitro and correlated with the neuropeptide distribution in the perivascular nerves, as demonstrated by immunohistochemistry. 2. Nerves displaying calcitonin gene related peptide (CGRP)- or neuropeptide Y (NPY)-like immunoreactivity (-LI) were observed in the media of both veins, while very few fibers were immunoreactive to vasoactive intestinal polypeptide (VIP) or substance P (SP) in either vein. 3. The staining pattern for NPY-LI was largely identical to that of dopamine-beta-hydroxylase, a marker for noradrenaline (NA) producing fibers, indicating coexistence of NPY and NA. 4. Administration of NPY in vitro elicited contractions in both veins in the presence of propranolol, though more conspicuously in the angular oculi vein. 5. The peptide was without any modulating effect on NA-stimulated contractions in the angular oculi vein, whereas a small enhancement of the NA-induced tone was seen in the facial vein. 6. CGRP caused partial relaxation of both veins, whereas atrial natriuretic polypeptide caused relaxation only in the facial vein. VIP and SP had no effect on either vein. 7. The results suggest that in reindeer the sympathetic nerve fibres to both facial and angular oculi veins contain the vasoconstrictor neuropeptide NPY besides NA, even though these fibres exert a vasodilator action on the myogenically active facial vein. 8. The vasodilator neuropeptide CGRP, which is present in other more sparse perivascular nerve fibres mainly in angular oculi vein, is perhaps of afferent nature in which case CGRP might subserve axon reflex functions. 9. If, however, also the CGRP fibres are truly efferent in nature, chances for a central reciprocal control of flow through angular oculi vein might be at hand.

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↗

Adrenergic vasomotor responses in nasal mucosa of hooded seals.

In seals respiratory heat and water losses are restricted through nasal heat exchange. The heat exchange efficiency is apparently controlled through adjustments in the nasal mucosal blood flow rate and/or pattern. In this study the adrenergic mechanisms involved in regulation of mucosal blood flow were investigated. The nasal mucosal vasculature of 14 newly killed hooded seal (Cystophora cristata) pups was perfused by a constant-flow peristaltic pump with 37 degrees C oxygenated modified Krebs solution via the sphenopalatine arteries. The effects of single-dose injections of various drugs on resistance to flow were monitored with a pressure transducer. Epinephrine, norepinephrine, alpha 1-adrenoceptor agonist phenylephrine, alpha 2-agonist clonidine, beta 1-agonist dobutamine, and beta 2-agonist terbutaline caused transient pressure increases that were blocked by alpha-adrenoceptor antagonists. Papaverine and vasoactive intestinal polypeptide induced vasodilatation, showing that some basal vascular tone was present. Nevertheless, the beta 1- and beta 2-agonist isoproterenol had no effect on resistance, and none of the beta-agonists attenuated the pressor responses to alpha-agonists. In conclusion, adrenergic control of nasal mucosal blood flow in seals is essentially exerted through alpha-adrenoceptor-mediated arteriolar constriction, whereas beta-adrenoceptor-mediated dilatation seems to be of little importance. It is suggested that such sympathoadrenergic vascular mechanisms contribute to control nasal heat exchange efficiency in seals.

Animals↗

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↗

Vascular control of brain cooling in reindeer.

The cold venous effluent returning from the nose of the heat-stressed reindeer may be distributed through angular oculi veins for selective cooling of the brain and through facial veins for general body cooling. In vitro experiments indicate that adrenergic receptors of the angular oculi vein are exclusively of the alpha-adrenergic type, whereas facial veins contain mainly beta-adrenergic receptors. We suggest that the antagonistic adrenergic neuroeffector organization of these veins plays a major role in control of brain cooling. Thus simultaneous sympathetic stimulation results in constriction of angular oculi veins and release of a preexisting stretch-induced tone in facial veins. In this situation cold venous blood is directed via facial veins and used for general body cooling. Reduction of sympathetic activity, on the other hand, results in dilatation of angular oculi veins and constriction of facial veins, due to development of myogenic tone in the latter vessels. In this situation cold venous blood is distributed via angular oculi veins and used for selective cooling of the brain.

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↗