Heart rate as a predictor of energy expenditure in undisturbed fasting and incubating penguins.

Heart rate (f(H)) measurement offers the possibility to monitor energy expenditure (EE) in wild animals if the EE/f(H) relationship for the species, physiological stages and activities of interest is known. This relationship has been extensively studied using oxygen consumption rate ( ) measurement in captive, repeatedly handled king penguins (Aptenodytes patagonicus). Unfortunately, the potential effects of stress on the observed relationships resulting from handling and confinement were not considered. This study is the first involving undisturbed animals, and determines the EE/f(H) relationship in naturally fasting and freely incubating or captivity-acclimatized male and female king penguins. EE determination was based on (1) the measurement of body mass loss during periods of phase II fasting, and (2) the calculation of its energy equivalent from changes in body composition, i.e. 23.9 kJ g(-1). f(H) levels in freely incubating and captivity-acclimatized birds were found to be 50-70% lower than those previously reported for resting king penguins during measurements. Significant EE/f(H) relationships were found in freely incubating and captive males and females (R(2)=0.59 to 0.84), with no difference observed between genders. The best overall relationship was obtained by including fasting duration (t, days) in the model: EE=818+43.7xf(H)+36.3t-1.4txf(H) (R(2)=0.91). This equation yielded EE estimates approximately 26% higher than the previously reported 'best' predictive equation in king penguins, and even more so when f(H) was low. This result suggests that stress induces a disproportionate increase of f(H) vs O(2) consumption, and that the use of EE/f(H) relationships obtained in stressed birds could lead to underestimated EE values.

Assessing a relationship between bone microstructure and growth rate: a fluorescent labelling study in the king penguin chick (Aptenodytes patagonicus).

Microstructure-function relationships remain poorly understood in primary bone tissues. The relationship between bone growth rate and bone tissue type, although documented in some species by previous works, remains somewhat unclear and controversial. We assessed this relationship in a species with extreme adaptations, the king penguin (Aptenodytes patagonicus). These birds have a peculiar growth, interrupted 3 months after hatching by the austral winter. Before this interruption, chicks undergo extremely rapid statural and ponderal growth. We recorded experimentally (by means of fluorescent labelling) the growth rate of bone tissue in four long bones (humerus, radius, femur and tibiotarsus) of four king penguin chicks during their fastest phase of growth (3-5 weeks after hatching) and identified the associated bone tissue types ('laminar', 'longitudinal', 'reticular' or 'radial' fibro-lamellar bone tissue). We found the highest bone tissue growth rate known to date, up to 171 microm day(-1) (mean 55 microm day(-1)). There was a highly significant relationship between bone tissue type and growth rate (P<10(-6)). Highest rates were obtained with the radial microarchitecture of fibro-lamellar bone, where cavities in the woven network are aligned radially. This result supports the heuristic value of a relationship between growth rate and bone primary microstructure. However, we also found that growth rates of bone tissue types vary according to the long bone considered (P<10(-5)) (e.g. growth rates were 38% lower in the radius than in the other long bones), a result that puts some restriction on the applicability of absolute growth rate values (e.g. to fossil species). The biomechanical disadvantages of accelerated bone growth are discussed in relation to the locomotor behaviour of the chicks during their first month of life.

Changes in tissue fatty acid composition during the first month of growth of the king penguin chick.

The switch from yolk to food (myctophid fishes) as the nutrient source for the newly hatched chick of the king penguin ( Aptenodytes patagonicus) results in a profound change in the pattern of fatty acid provision. This is characterized by major increases in the proportionate intake of n-3 polyunsaturates (20:5n-3 and 22:6n-3) and long chain (C(20-24)) monounsaturates, accompanied by relatively lower levels of n-6 polyunsaturates (18:2n-6 and 20:4n-6). The effects of this change on the fatty acid composition of tissue lipids during the first month of growth, a period of tissue maturation leading to thermal emancipation, were determined. The composition of adipose tissue triacylglycerol responded rapidly to the switch in nutrient source, the proportion of long chain monounsaturates (mainly 20:1n-9 and 22:1n-11) increasing five-fold between hatch and emancipation while the relative levels of 20:5n-3 and 22:6n-3 also increased significantly, by 3- and 1.2-fold, respectively. At emancipation, the fatty acid profile of adipose tissue triacylglycerol was essentially identical to that of the diet. At hatch, the main polyunsaturates of muscle phospholipid were 20:4n-6, 20:5n-3, and 22:6n-3, respectively, forming (w/w of fatty acids) 13.2%, 5.0%, and 12.0%. By emancipation, 20:4n-6 had decreased to 4.8%, 20:5n-3 increased to 10.9%, and 22:6n-3 at 11.4% showed little change. The main polyunsaturate in brain phospholipid at hatch was 22:6n-3 (19.3%): this remained almost constant until day 15 but then increased significantly to 23.6% by emancipation. Significant but minor changes in the proportions of 20:4n-6 (from 5.2% at hatch to 3.5% at emancipation) and 20:5n-3 (from 3.0% to 3.9%) were also observed in brain phospholipid. The data do not allow us to completely distinguish changes that are solely diet driven from those which are a consequence of tissue differentiation. Nevertheless, it is evident that, whereas the fatty acid composition of adipose tissue responds faithfully to the change in nutrient source, the phospholipids of muscle and, especially, of brain are much more refractory to the effects of diet during this period of tissue maturation.

Glycerol and NEFA kinetics in long-term fasting king penguins: phase II versus phase III.

In spontaneously fasting birds such as penguins, below a body mass threshold corresponding to the phase II-phase III transition, a metabolic and hormonal shift occurs and feeding behaviour is stimulated ('refeeding signal'). The major aim of this study was to determine whether a decrease in non-esterified fatty acid (NEFA) release from adipose tissue could be a component of this signal. Lipolytic fluxes and primary triacylglycerol:fatty acid (TAG:FA) cycling were determined in vivo in breeding, fasting king penguins (Aptenodytes patagonicus) using continuous infusions of 2-[3H]glycerol and 1-[14C]palmitate under field conditions. In phase II (after approximately 8 days of fasting, large fat stores, body protein spared, N=8), the rate of appearance (R(a)) of glycerol and of NEFA were 5.7+/-0.8 and 10.5+/-0.4 micromol kg(-1) min(-1), respectively, and the percentage of primary TAG:FA cycling was 41+/-7%. In phase III (after approximately 25 days of fasting, fat stores reduced by fourfold, increased body protein catabolism, N=9), R(a) glycerol kg(-1) body mass remained unchanged, whereas R(a) glycerol kg(-1) fat mass and R(a) NEFA kg(-1) body mass were increased by 2.8-fold and 1.5-fold, respectively. Increased R(a) glycerol kg(-1) fat mass was possibly the result of a 3.5-fold increase in circulating glucagon, the increased R(a) NEFA kg(-1) body mass being attributable to decreased primary TAG:FA cycling. Thus, triggering of the refeeding signal that redirects the behavior of fasting, incubating penguins from incubation towards the search for food after entrance into phase III cannot be ascribed to a reduction in lipolytic fluxes and NEFA availability.

Transfer of Vitamins E and A from yolk to embryo during development of the king penguin (Aptenodytes patagonicus).

Since the yolk lipids of the king penguin (Aptenodytes patagonicus) are rich in n-3 fatty acids, which are potentially susceptible to peroxidative damage, the yolk contents and yolk-to-embryo transfer of antioxidants and lipid-soluble vitamins were investigated under conditions of natural incubation in the wild. The concentration of vitamin E in the unincubated egg was 155 microg/g wet yolk, of which 88% was alpha-tocopherol and the rest was gamma-tocopherol. Vitamin A (2.9 microg/g) was present in the yolk entirely as retinol; no retinyl esters were detected. Throughout the latter half of the incubation period, vitamins E and A were taken up from the yolk into the yolk sac membrane (YSM) and later accumulated in the liver, with vitamin A being transferred in advance of vitamin E. In the YSM, vitamin A was present almost entirely as retinyl ester, indicating that the free retinol of the yolk is rapidly esterified following uptake. Retinyl esters were also the predominant form in the liver. The retinyl esters of the liver and YSM displayed different fatty acid profiles. At hatching, the brain contained relatively little vitamin E (4.7 microg/g) compared to the much higher concentration in the liver (482.9 microg/g) at this stage. Ascorbic acid was not detected in the yolk but was present at a high concentration in the brain at day 27 (404.6 microg/g), decreasing to less than half this value by the time of hatching. This report is the first to delineate the yolk-to-embryo transfer of lipid-soluble vitamins for a free-living avian species. The yolk fatty acids of the king penguin provide an extreme example of potential oxidative susceptibility, forming a basis for comparative studies on embryonic antioxidant requirements among species of birds whose yolk lipids differ in their degree of unsaturation.

Milking strategy in subantarctic fur seals Arctocephalus tropicalis breeding on Amsterdam Island: evidence from changes in milk composition.

Milk composition was investigated throughout the 10-mo pup-rearing period in subantarctic fur seals (Arctocephalus tropicalis) breeding on Amsterdam Island. The mean milk composition was 42.8% +/- 5.7% lipid, 12.1% +/- 1.5% protein, and 42.6% +/- 7.3% water. Subantarctic fur seals breeding on Amsterdam Island produced one of the richest milks ever reported in otariids (20.4 +/- 2.9 kJ/g), with lipid content contributing 85% of total gross energy. The high lipid levels measured in the milk of subantarctic fur seals breeding on Amsterdam Island is consistent (i) with the relatively long time lactating females spend at sea, due to the relatively poor local trophic conditions near the colony that necessitate that they travel long distances to reach the foraging grounds, and (ii) with the consequently short time mothers spend with their pups ashore. Milk composition changed according to the time mothers were fasting ashore: milk produced during the first 2 d spent ashore, when more than 80% of milk transfer occurred, had higher levels of lipids, proteins, and gross energy than milk produced later during the visit ashore, suggesting that the pups were fed with two types of milk during a suckling period. Throughout the year, mothers in good condition produced milk of higher lipid content than others, suggesting that individual foraging skills contribute to enhance milk quality. Milk lipid and gross energy content varied with pup age, according to quadratic relationships, increasing during the earlier stages of lactation before reaching asymptotic values when pups were 180 d old. The stage of lactation appears to be a better predictor of milk lipid content than the duration of the preceding foraging trip, suggesting that either changes in the nutritional requirements of the pup and/or seasonal changes in trophic conditions act on milk composition. These changes in milk quality may also be related to changes in maternal care; lactating subantarctic fur seals apparently reallocate their body reserves toward gestation rather than lactation at the end of the pup-rearing period.

Long-term fasting and re-feeding in penguins.

Spontaneous fasting during reproduction (sometimes with a full stomach) and moult is a major characteristic of the annual cycle of penguins. Long-term fasting (up to four months in male emperor penguins) is anticipated by the accumulation of fat (incubation fast) and of fat and protein (moult fast). During most of the incubation fast, birds rely almost entirely on lipids as an energy source, body proteins being spared. However, below a critical (but non-total) fat store depletion, marked behavioural, metabolic, and endocrine changes occur. Spontaneous locomotor activity increases and the egg is transitorily left unincubated for increasingly long periods, until its definitive abandon and the bird departs to re-feed at sea. These changes are thought to be activated by an endogenous re-feeding signal triggered before lethal energy depletion. An increase in body protein catabolism in the face of a reduction in lipid availability and utilisation, and an increase in circulating corticosterone vs. a decrease in plasma prolactin, are likely to be major metabolic and hormonal components of this signal. The survival and rapid restoration of energy stores in birds having departed to re-feed at a stage of near total lipid depletion demonstrates the effectiveness of the re-feeding signal. Penguins, and possibly other seabirds, are therefore appropriate animal models for understanding the long-term interactions between body energy reserves and fasting, breeding and feeding physiology and behaviour.

Transfer of n-3 and n-6 polyunsaturated fatty acids from yolk to embryo during development of the king penguin.

This study examines the transfer of lipids from the yolk to the embryo of the king penguin, a seabird with a high dietary intake of n-3 fatty acids. The concentrations of total lipid, triacylglycerol (TAG), and phospholipid (PL) in the yolk decreased by ~80% between days 33 and 55 of development, indicating intensive lipid transfer, whereas the concentration of cholesteryl ester (CE) increased threefold, possibly due to recycling. Total lipid concentration in plasma and liver of the embryo increased by twofold from day 40 to hatching due to the accumulation of CE. Yolk lipids contained high amounts of C(20-22) n-3 fatty acids with 22:6(n-3) forming 4 and 10% of the fatty acid mass in TAG and PL, respectively. Both TAG and PL of plasma and liver contained high proportions of 22:6(n-3) ( approximately 15% in plasma and >20% in liver at day 33); liver PL also contained a high proportion of 20:4(n-6) (14%). Thus both 22:6(n-3) and 20:4(n-6), which are, respectively, abundant and deficient in the yolk, undergo biomagnification during transfer to the embryo.

Refeeding signal in fasting-incubating king penguins: changes in behavior and egg temperature.

This study is directed toward understanding the process of feeding stimulation ("refeeding signal") that has been suggested to operate below a body mass threshold or critical metabolic status in spontaneously fasting birds. Behavior and egg temperature (T(egg)) were continuously monitored by video monitoring and biotelemetry, respectively, in fasting-incubating king penguins kept in a pen to prevent relief by the partner until spontaneous egg abandonment. Penned birds fasted 10 days more and lost 1.2 kg more than birds relieved normally by their partner, abandoning the egg about 1 wk after reaching a critical body mass. Definitive egg abandonment was preceded by transitory abandonments of progressively increasing duration during which time the birds went further and further away from their egg. There were marked interindividual differences but on average transitory abandonments began 36 +/- 5 h before the definitive abandonment and were paralleled by resumption of display songs signaling the readiness of the bird to depart for feeding. T(egg) was maintained at around 35.7 degrees C during normal incubation but significantly decreased the last 2 days before egg abandonment. These changes are interpreted as reflecting a stimulation to refeed at a threshold body mass corresponding to a critical fat store depletion. Thus the fasting-incubating king penguin appears to be an interesting animal model for understanding the long-term metabolic control of feeding behavior in relation to energy status.

Lipid composition of erythrocytes and thrombocytes of a subantarctic seabird, the king penguin.

Phospholipid (PL) compositions and fatty acid (FA) patterns of PL were determined in the erythrocytes and blood thrombocytes of a seabird, the king penguin, living in the subantarctic area and feeding on prey rich in n-3 polyunsaturated FA. Results were compared between birds in three different physiological states (breeding and molting adults, chicks) to those reported for other birds. In erythrocytes, the ratios of cholesterol to PL and of sphingomyelin to phosphatidylcholine (PC) were lower than in other birds. The PL distribution was similar to those previously reported in the hen and pigeon. In contrast to other birds, cardiolipin levels were unexpectedly high (4%). Very long chain n-3 FA were abundant (13-27%) in phosphatidylethanolamine (PE), phosphatidylserine and PC, probably in relation to the natural diet of these birds. Among n-3 FA, 22:6n-3 was the most abundant in all PL (2-20%), whereas the highest levels of arachidonic acid were observed in PE (14%). In thrombocytes, the PL distribution and FA composition of the main PL (PC, PE) differed from those of erythrocytes, and in particular, levels of n-3 FA (9-12%) were 1.5-2 times lower. The highest levels of arachidonic acid were found in phosphatidylinositol (24%). The lipid profile of penguin erythrocytes could contribute to the efficiency of blood circulation and oxygen delivery in microvascular beds, thus favoring diving capacity of these animals. Our observations do not support the hypothesis of a common origin of avian thrombocytes and erythrocytes.

Lipid composition of blood platelets and erythrocytes of southern elephant seal (Mirounga leonina) and antarctic fur seal (Arctocephalus gazella).

Erythrocyte and blood platelet phospholipid compositions were studied in three elephant seals and two fur seals, two species of marine mammals living in the Subantarctic region feeding on preys rich in (n-3) polyunsaturated fatty acids. Results were compared with those reported for related species and humans. In erythrocytes, the phospholipid (PL) and cholesterol (CHOL) contents were lower in pinnipeds than in humans. Phosphatidylcholine (PC) levels were higher in elephant seals than in fur seals, with a reverse trend for phosphatidylethanolamine (PE) and phosphatidylserine (PS). Both species had lower SM/PC ratios and PE plasmalogen concentrations than human. Erythrocytes were richer in (n-3) fatty acids (FA) in pinnipeds than in humans. In platelets, the PL content was lower and the CHOL content higher in elephant seals than in humans or in other phocid seal species studied to date. The SM/PC ratio was much higher than in other seal species or in man. In both species, the proportion of PE plasmalogens was higher in platelets than in erythrocytes. PL were more saturated in elephant seals than in fur seals. These results suggest that the erythrocytes and platelets of wild marine mammals may prove useful models to study the influence of dietary lipids on the structure and hemostatic function of these cells.

Fatty acid composition of the adipose tissue and yolk lipids of a bird with a marine-based diet, the emperor penguin (Aptenodytes forsteri).

The emperor penguin (Aptenodytes forsteri) is an Antarctic seabird feeding mainly on fish and therefore has a high dietary intake of n-3 polyunsaturated fatty acids. The yolk is accumulated in the developing oocyte while the females are fasting, and a large proportion of the fatty acid components of the yolk lipids are derived by mobilization from the female's adipose tissue. The fatty acid composition of the total lipid of the yolk was characterized by high levels of n-3 polyunsaturated fatty acids. However, it differed in several respects from that of the maternal adipose tissue. For example, the proportions of 14:0, 16:1n-7, 20:1n-9, 22:1n-9, 20:5n-3, and 22:6n-3 were significantly greater in adipose tissue than in yolk. Thus adipose tissue lipids contained 7.6+/-0.3% and 8.0+/-0.3% (wt% of total fatty acids; mean +/- SE; n = 5) of 20:5n-3 and 22:6n-3, respectively, whereas the yolk total lipid contained 1.6+/-0.1 and 5.5+/-0.3% of these respective fatty acids. The proportions of 16:0, 18:0, 18:1n-9, 18:2n-6, and 20:4n-6 were significantly lower in the adipose tissue than in the yolk lipids. The proportions of triacylglycerol, phospholipid, free cholesterol, and cholesteryl ester in the yolk lipid were, respectively, 67.0+/-0.2, 25.4+/-0.3, 5.3+/-0.2, and 1.8+/-0.2% (wt% of total yolk lipid). The proportions of 20:4n-6, 20:5n-3, 22:5n-3, and 22:6n-3 were, respectively, 5.7+/-0.3, 2.8+/-0.2, 1.4+/-0.1, and 11.7+/-0.5% in phospholipid and 0.4+/-0.0, 1.2+/-0.1, 0.8+/-0.1 and 3.6+/-0.3% in triacylglycerol. About 95% of the total vitamin E in the yolks was in the form of alpha-tocopherol with gamma-tocopherol forming the remainder. Two species of carotenoids, one identified as lutein, were present.

Behavioral changes in fasting emperor penguins: evidence for a "refeeding signal" linked to a metabolic shift.

This study examines the relationships between metabolic status and behavior in spontaneously fasting birds in the context of long-term regulation of body mass and feeding. Locomotor activity, escape behavior, display songs, body mass, and metabolic and endocrine status of captive male emperor penguins were recorded during a breeding fast. We also examined whether body mass at the end of the fast affected further survival. The major part of the fast (phase II) was characterized by the maintenance of a very low level of locomotor activity, with almost no attempt to escape, by an almost constant rate of body mass loss, and by steady plasma levels of uric acid, beta-hydroxybutyrate, and corticosterone. This indicates behavioral and metabolic adjustments directed toward sparing energy and body protein. Below a body mass of approximately 24 kg (phase III), spontaneous locomotor activity and attempts to escape increased by up to 8- and 15-fold, respectively, and display songs were resumed. This probably reflected an increase in the drive to refeed. Simultaneously, daily body mass loss and plasma levels of uric acid and corticosterone increased, whereas plasma levels of beta-hydroxybutyrate decreased. Some experimental birds were seen again in following years. These findings suggest that at a threshold of body mass, a metabolic and endocrine shift, possibly related to a limited availability of fat stores, acts as a "refeeding signal" that improves the survival of penguins to fasting.

Site-specific regulation of gene expression by n-3 polyunsaturated fatty acids in rat white adipose tissues.

Dietary n-3 polyunsaturated fatty acids (n-3 PUFAs) limit abdominal fat depot hypertrophy. This could be due to regulation of the expression of proteins involved in adipose tissue metabolism. We investigated in vivo whether fatty acid synthase (FAS), hormone-sensitive lipase (HSL), lipoprotein lipase (LPL), phosphoenolpyruvate carboxykinase (PEPCK), CCAAT/enhancer binding protein alpha (C/EBP alpha), and leptin mRNA levels are affected in retroperitoneal (RP) and subcutaneous adipose tissues (SC) of rats fed n-3 PUFAs. For 4 weeks rats were fed high fat diets (20% fat) containing n-3 PUFAs given as eicosapentaenoic acid (EPA group), docosahexaenoic acid (DHA group), a mixture of these two fatty acids (MIX group), or native fish oil (FO group). A control group was fed with lard plus olive oil (LOO group). Final mean fat cell weight in RP ranged according to: LOO > or = EPA > or = DHA = FO = MIX. There was no difference in fat cell size of SC when comparing the LOO and MIX groups. The fatty acid compositions of RP and SC were similar and resemble that of dietary fat within each experimental group. In RP and compared to the LOO group, FAS, HSL, PEPCK, LPL, C/EBP alpha, and leptin mRNA levels decreased although not significantly in the EPA group, and were 40-75% lower in the DHA and MIX groups. mRNA levels were positively correlated to fat cell size in RP. In contrast, n-3 PUFAs had no effect on gene expression in SC. We conclude that n-3 PUFAs and mainly 22:6n-3 affect gene expression in a site-dependent manner in white adipose tissues via possible antiadipogenic effects.

Brown fat thermogenesis in rats fed high-fat diets enriched with n-3 polyunsaturated fatty acids.

OBJECTIVE: To examine the possible involvement of an increase in diet-induced thermogenesis from brown adipose tissue (BAT) in the n-3 polyunsaturated fatty acids (n-3 PUFA) induced limitation of the development of white fat pads during high-fat feeding. DESIGN: Rats fed for four weeks on a low-fat/high-carbohydrate diet (C group) or high-fat diet without n-3 PUFA (REF group), with eicosapentaenoic acid (EPA group), with docosahexaenoic acid (DHA group) or with a mixture of these two fatty acids (MIX group). MEASUREMENTS: Epididymal and retroperitoneal fat pad mass, BAT composition, Guanosine 5'-diphosphate (GDP) binding and uncoupling protein (UCP) content were measured in the five groups of rats. RESULTS: The masses of retroperitoneal and epididymal white fat pads were lower in the groups fed n-3 PUFA than in the C and REF groups. The total BAT GDP binding was 1.6 times higher in the MIX and EPA groups than in the REF group. The BAT from the EPA group presented an enrichment in mitochondria compared to the C and REF groups whereas the BAT from the DHA and REF groups presented a hyperplasia and an increase in thermogenic activity of the mitochondria compared to the C group. The higher thermogenic activity of BAT was observed in the MIX group and is due to hyperplasia and to an increase in thermogenic activity of mitochondria. CONCLUSIONS: n-3 PUFA induce a marked stimulation of BAT thermogenic activity without changes in the UCP content compared to a high-fat diet without n-3 PUFA. The mixture of EPA and DHA has the more pronounced effect while EPA and DHA seem to act in synergy on BAT thermogenesis via different mechanisms.

Selective release of human adipocyte fatty acids according to molecular structure.

The objective of the present study was to investigate the mobilization of individual fatty acids from human white fat cells. Mammary adipose tissue from eight healthy non-obese women in their normal dietary state was collected, and isolated adipocytes were incubated with lipolytic agents. The mobilization of 34 individual fatty acids was measured by comparing the composition of non-esterified fatty acids (NEFA) with that of the triacylglycerols (TAG) from which they originated through lipolysis. Compared with TAG, NEFA were enriched in some polyunsaturated fatty acids with 18-20 carbon atoms. Conversely, the percentage of very-long-chain (20-22 carbon atoms) saturated and monounsaturated fatty acids was approx. 2 times lower in NEFA than in TAG. The relative mobilization (% in NEFA/% in TAG) of the most readily mobilized fatty acid (C20:5, n-3; 2.25) was more than 6-fold higher than that of the least readily mobilized (C22:1,n-11; 0.37). Relationships were found between the molecular structure of fatty acids and their mobilization rate. For a given chain length, the relative mobilization rate increased with increasing unsaturation, whereas for a given unsaturation, it decreased with increasing chain length. The relative mobilization rate for essential fatty acids decreased in the following order: C20:5,n-3>C20:4,n-6>C18:3,n-3>C18:2, n-6>C22:6,n-3. Interestingly, C20:5,n-3 and C20:4,n-6, which are respectively precursors of the 3- and 2-series of prostaglandins, were preferentially mobilized. It is concluded that fatty acids are selectively mobilized from human fat cells according to molecular structure, in full agreement with animal studies. By modulating the qualitative fatty acid supply to organs and by remodelling the fatty acid composition of adipose tissue, this selectivity would be relevant for consideration in physiology, health and epidemiology.

Fasting-induced selective mobilization of brown adipose tissue fatty acids.

This study was conducted to determine whether there is selective mobilization of fatty acids from brown adipose tissue. Rats were fed a fish-oil diet and then fasted for 0, 7, or 10 days followed by analysis of the fatty acid content and composition of triacylglycerols (TAG) and phospholipid (PL) in interscapular brown adipose tissue (IBAT). Fatty acids were selectively lost from IBAT triacylglycerols, the mobilization following the same structural rules as those previously demonstrated for white adipose tissue. Fractional mobilization increased with unsaturation at a given chain length and tended to decrease with chain length at a given unsaturation. However, linoleic acid (18:2 n-6) was mobilized significantly less than predicted by these structural rules. In IBAT phospholipid, fatty acids were also selectively lost but there was no such relationship between the fractional mobilization of a fatty acid and its structure. The fatty acids of the n-6 series were exceptional in their behavior because they displayed below average fractional mobilization. In fact, linoleic and arachidonic acids actually increased their mass in IBAT phospholipid during the fast. It is concluded that, in IBAT, fatty acids are selectively mobilized during a fast, and that fasting-induced remodeling of the fatty acid composition leads to the selective retention of linoleate in the case of TAG and all the n-6 fatty acids in the case of PL.

Selectivity of fatty acid mobilization: a general metabolic feature of adipose tissue.

This study extends our earlier work (T. Raclot and R. Groscolas. J. Lipid Res. 34: 1515-1526, 1993), which showed that, under norepinephrine-stimulated lipolysis, fatty acids of rat retroperitoneal fat cells are selectively mobilized. The present study examines whether this selective mobilization of fatty acids 1) is based on their proportions in adipose tissue, 2) is a metabolic feature common to all adipose tissues, and/or 3) depends on the lipolysis-stimulating agent. Rat fat cells with two markedly different fatty acid compositions were isolated from four white adipose tissues and treated with three lipolytic agents. Fatty acid composition of in vitro released free fatty acids was compared with that of fat cell triacylglycerols, the ratio of percent in free fatty acid to percent in triacylglycerol being defined as the relative mobilization rate (RMR). The RMR of individual fatty acids was related to their molecular structure. It increased exponentially with unsaturation for a given chain length and decreased with increasing chain length for a given unsaturation. The selectivity of fatty acid mobilization was similar regardless of the fatty acid composition of adipose tissue, the tissue location, and the lipolytic agent used. Under conditions of stimulated lipolysis, the selectivity of fatty acid mobilization is therefore a general metabolic feature of adipose tissue. Fatty acids with 16-20 carbon atoms and 4 or 5 double bonds had the highest RMR (from 1.4 to > 5), whereas fatty acids with 20-22 carbon atoms and 0 or 1 double bond had the lowest RMR (from 0.3 to 0.7). For the other fatty acids, RMR was close to unity.(ABSTRACT TRUNCATED AT 250 WORDS)

The selective mobilization of fatty acids is not based on their positional distribution in white-fat-cell triacylglycerols.

Fatty acids have been shown to be selectively mobilized from rat white fat-cells, whatever the dietary manipulations. For convenience, fatty acids have been classified as being highly, weakly and moderately mobilizable. The aim of this study was to examine whether the selective mobilization of fatty acids can be explained, even partly, by their positional distribution in adipose-tissue triacylglycerols (TAG) via the known specificity of hormone-sensitive lipase for the sn-1 and sn-3 positions. Adipose tissue was dietarily manipulated in order to obtain a wide spectrum of fatty acids, including large amounts of either very-long-chain polyunsaturated fatty acids (VLC-PUFA) or very-long-chain monounsaturated fatty acids (VLC-MUFA). The determination of fatty acid distribution in adipose tissue TAG was based on random formation of 1,2-diacyl-rac-glycerols by Grignard degradation, followed by synthesis of phosphatidic acids and hydrolysis in the sn-2 position by phospholipase A2. Regardless of the fatty acid composition and location of fat depots, highly (e.g. 18:4n-3 and some of the VLC-PUFA) and weakly (e.g. VLC-MUFA) mobilizable fatty acids were located mainly in the outer (sn-1 and sn-3) positions of the glycerol moiety (79.5% and 92.5% on average, respectively). Other fatty acids, which are rather moderately mobilizable, were more randomly distributed. We conclude that the selective mobilization of white-fat-cell fatty acids is not based on their positional distribution in TAG.