Effects of dietary eicosapentaenoic acid on growth, survival, pigmentation and fatty acid composition in Senegal sole (Solea senegalensis) larvae during the Artemia feeding period

We examined the effect of dietary eicosapentaenoic acid (20:5n-3, EPA) on growth, survival, pigmentation and fatty acid composition of Senegal sole larvae using a dose-response design. From 3 to 40 days post hatch (dph), larvae were fed live food that had been enriched using one of four experimental emulsions containing graduated concentrations of EPA and constant docosahexaenoic acid (22:6n-3, DHA) and arachidonic acid (20:4n-6, ARA). Proportions of EPA in the enriched Artemia nauplii were described as “nil” (EPA-N, 0.5% total fatty acids, TFA), “low” (EPA-L, 10.7% TFA), “medium” (EPA-M, 20.3% TFA) or “high” (EPA-H, 29.5% TFA). Significant differences among dietary treatments in larval length were observed at 25, 30 and 40 dph, and in dry weight at 30 and 40 dph, although no significant correlation could be found between dietary EPA content and growth. The stage of eye migration at 17 and 25 dph was significantly affected by dietary levels of EPA. Significantly lower survival was observed in fish fed EPA-H enriched nauplii. A significantly lower percentage of fish fed EPA-N (82.7%) and EPA-L (82.9%) diets were normally pigmented compared to the fish fed EPA-M (98.1%) and EPA-H (99.4%) enriched nauplii. Tissue fatty acid concentrations reflected the corresponding dietary composition. Arachidonic and docosahexaenoic acid levels in all the tissues examined were inversely related to dietary EPA. There was an increase in the proportion of docosapentaenoic acid (22:5n-3, DPA) in the tissues relative to the diet, which is indicative of chain elongation of EPA. This work concluded that Senegal sole larvae have a very low EPA requirement during the live feeding period

Principles of resilient coding for plant ecophysiologists

Plant ecophysiology is founded on a rich body of physical and chemical theory, but it is challenging to connect theory with data in unambiguous, analytically rigorous and reproducible ways. Custom scripts written in computer programming languages (coding) enable plant ecophysiologists to model plant processes and fit models to data reproducibly using advanced statistical techniques. Since many ecophysiologists lack formal programming education, we have yet to adopt a unified set of coding principles and standards that could make coding easier to learn, use and modify. We identify eight principles to help in plant ecophysiologists without much programming experience to write resilient code: (i) standardized nomenclature, (ii) consistency in style, (iii) increased modularity/extensibility for easier editing and understanding, (iv) code scalability for application to large data sets, (v) documented contingencies for code maintenance, (vi) documentation to facilitate user understanding; (vii) extensive tutorials and (viii) unit testing and benchmarking. We illustrate these principles using a new R package, {photosynthesis}, which provides a set of analytical and simulation tools for plant ecophysiology. Our goal with these principles is to advance scientific discovery in plant ecophysiology by making it easier to use code for simulation and data analysis, reproduce results and rapidly incorporate new biological understanding and analytical tools

Effects of dietary protein and fat level and rapeseed oil on growth and tissue fatty acid composition and metabolism in Atlantic salmon (Salmo salar L.) reared at low water temperatures

A 12 week feeding trial was conducted to elucidate the interactive effects of dietary fat and protein contents and oil source on growth, fatty acid composition, protein retention efficiency (PRE) and β-oxidation activity of muscle and liver in Atlantic salmon (Salmo salar L.) at low water temperatures (4.2 oC). Triplicate groups of Atlantic salmon (initial weight 1168 g) were fed six isoenergetic diets formulated to provide either 390 g kg-1 protein and 320 g kg-1 fat (high protein (HP) diets) or 340 g kg-1 protein and 360 g kg-1 fat (low protein (LP) diets); within each dietary protein/fat level crude RO comprised 0, 30 or 60% (R0, R30, R60, respectively) of the added oil. After 12 weeks the overall growth and FCR were very good for all treatments (TGC; 4.76 (±0.23), FCR; 0.85 (±0.02)). Significant effects were shown due to oil source on SGR and TGC only. The liver and muscle FA compositions were highly affected by the graded inclusion of RO. The PRE was significantly affected by the dietary protein level, while no significant effects were shown in total β-oxidation capacity of liver and muscle. The results of this study suggest that more sustainable, lower protein diets with moderate RO inclusion can be used in Atlantic salmon culture at low water temperatures with no negative effects on growth and feed conversion, no major detrimental effects on lipid and fatty acid metabolism and a positive effect on protein sparing

Fatty acid metabolism in marine fish: Low activity of fatty acyl Δ5 desaturation in gilthead sea bream ( Sparus aurata ) cells

Marine fish are known to have an absolute dietary requirement for C20 and C22 highly unsaturated fatty acids. Previous studies using cultured cell lines indicated that underlying this requirement in marine fish was either a deficiency in fatty acyl Δ5 desaturase or C18-20 elongase activity. Recently, Ghioni et al. (Biochim. Biophys. Acta, 1437, 170-181, 1999) presented evidence that in turbot cells there was low activity of C18-20 elongase whereas Δ5 desaturase had high activity. In the present study, the fatty acid desaturase/elongase pathway was investigated in a cell line (SAF-1) from another carnivorous marine fish, sea bream. The metabolic conversions of a range of radiolabelled polyunsaturated fatty acids that comprised the direct substrates for Δ6 desaturase ([1-14C]18:2n-6 and [1-14C]18:3n-3), C18-20 elongase ([U-14C]18:4n-3), Δ5 desaturase ([1-14C]20:3n-6 and [U-14C]20:4n-3) and C20-22 elongase ([1-14C]20:4n-6 and [1-14C]20:5n-3) were utilized. The results showed that fatty acyl Δ6 desaturase in SAF-1 cells was highly active and there was substantial C18-20 elongase and C20-22 elongase activities. A deficiency in the desaturation/elongation pathway was clearly identified at the level of the fatty acyl Δ5 desaturase which was very low, particularly with 20:4n-3 as substrate. In comparison, the apparent activities of Δ6 desaturase, C18-20 elongase and C20-22 elongase were approximately 94-fold, 27-fold and 16-fold greater than that for Δ5 desaturase towards their respective n-3 polyunsaturated fatty acid substrates. The evidence obtained in the SAF-1 cell line is consistent with the dietary requirement for C20 and C22 highly unsaturated fatty acids in the marine fish, the sea bream, being primarily due to a deficiency in fatty acid Δ5 desaturase activity

Effects of dietary vegetable oil on atlantic salmon hepatocyte fatty acid desaturation and liver fatty acid compositions

Fatty acyl desaturase activities, involved in the conversion of the C18 EFA, 18:2n-6 and 18:3n-3, to the highly unsaturated fatty acids (HUFA) 20:4n-6, 20:5n-3 and 22:6n-3, are known to be under nutritional regulation. Specifically, the activity of the desaturation/elongation pathway is depressed when animals, including fish, are fed fish oils rich in n-3HUFA compared to animals fed vegetable oils rich in C18 EFA. The primary aims of the present study were a) to establish the relative importance of product inhibition (n-3HUFA) versus increased substrate concentration (C18 EFA) and, b) to determine whether 18:2n-6 and 18:3n-3 differ in their effects, on the hepatic fatty acyl desaturation/elongation pathway in Atlantic salmon (Salmo salar). Smolts were fed ten experimental diets containing blends of two vegetable oils, linseed (LO) and rapeseed oil (RO), and fish oil (FO) in a triangular mixture design for 50 weeks. Fish were sampled after 32 and 50 weeks, lipid and fatty acid composition of liver determined, fatty acyl desaturation/elongation activity estimated in hepatocytes using [1-14C]18:3n-3 as substrate, and the data subjected to regression analyses. Dietary 18:2n-6 was positively correlated, and n-3HUFA negatively correlated, with lipid content of liver. Dietary 20:5n-3 and 22:6n-3 were positively correlated with liver fatty acids with a slope greater than unity suggesting relative retention and deposition of these HUFA. In contrast, dietary 18:2n-6 and 18:3n-3 were positively correlated with liver fatty acids with a slope of less than unity suggesting metabolism via β-oxidation and/or desaturation/elongation. Consistent with this, fatty acyl desaturation/elongation in hepatocytes was significantly increased by feeding diets containing vegetable oils. Dietary 20:5n-3 and 22:6n-3 levels were negatively correlated with hepatocyte fatty acyl desaturation. At 32 weeks, 18:2n-6 but not 18:3n-3, was positively correlated with hepatocyte fatty acyl desaturation activity whereas the reverse was true at 50 weeks. The data indicate that both feedback inhibition through increased n-3HUFA and decreased C18 fatty acyl substrate concentration are probably important in determining hepatocyte fatty acyl desaturation activities, and that 18:2n-6 and 18:3n-3 may differ in their effects on this pathway

Cultured fish cells metabolize octadecapentaenoic acid (all-cis delta3,6,9,12,15–18∶5) to octadecatetraenoic acid (all-cis delta6,9,12,15–18∶4) via its 2-trans intermediate (trans delta2, all-cis delta6,9,12,15–18∶5)

Octadecapentaenoic acid (all-cis Δ3,6,9,12,15-18:5; 18:5n-3) is an unusual fatty acid found in marine dinophytes, haptophytes and prasinophytes. It is not present at higher trophic levels in the marine food web but its metabolism by animals ingesting algae is unknown. Here we studied the metabolism of 18:5n-3 in cell lines derived from turbot (Scophthalmus maximus), gilthead sea bream (Sparus aurata) and Atlantic salmon (Salmo salar). Cells were incubated in the presence of approximately 1 μM [U-14C] 18:5n-3 methyl ester or [U-14C] 18:4n-3 (octadecatetraenoic acid; all-cis Δ6,9,12,15-18:4) methyl ester, both derived from the alga Isochrysis galbana grown in H14CO3, and also with 25 μM unlabelled 18:5n-3 or 18:4n-3. Cells were also incubated with 25 μM trans Δ2, all-cis Δ6,9,12,15-18:5 (2-trans 18:5n-3) produced by alkaline isomerization of 18:5n-3 chemically synthesized from docosahexaenoic acid (all-cis Δ4,7,10,13,16,19-22:6; 22:6n-3). Radio- and mass analyses of total fatty acids extracted from cells incubated with 18:5n-3 were consistent with this fatty acid being rapidly metabolized to 18:4n-3 which was then elongated and further desaturated to eicosatetraenoic acid (all-cis Δ8,11,14,17,19-20:4; 20:4n-3) and eicosapentaenoic acid (all-cis Δ5,8,11,14,17-20:5; 20:5n-3). Similar mass increases of 18:4n-3 and its elongation and further desaturation products occurred in cells incubated with 18:5n-3 or 2-trans 18:5n-3. We conclude that 18:5n-3 is readily converted biochemically to 18:4n-3 via a 2-trans 18:5n-3 intermediate generated by a Δ3,Δ2-enoyl-CoA-isomerase acting on 18:5n-3. Thus, 2-trans 18:5n-3 is implicated as a common intermediate in the β-oxidation of both 18:5n-3 and 18:4n-3

Utilization and metabolism of palmityl and oleoyl fatty acids and alcohols in caecal enterocytes of Atlantic salmon (Salmo salar L.)

The substitution of fish oil with wax ester-rich calanoid copepod-derived oil in diets for carnivorous fish, such as Atlantic salmon, has previously indicated a lower lipid digestibility. This suggests that the fatty alcohols (FAlc) present in wax esters may be a poorer substrate for intestinal enzymes than the fatty acids (FA) in triacylglycerol, the major lipid in fish oil. The hypothesis tested was that the possible lower utilization of dietary FAlc by salmon enterocytes is at the level of uptake and that subsequent intracellular metabolism was identical to that of FA. A dual-labelled FAlc-FA metabolism assay was employed to determine simultaneous FAlc and FA uptake and relative utilisation in enterocytes isolated from pyloric caeca of Atlantic salmon fed either a diet supplemented with fish oil or wax ester-rich Calanus oil. The diets were fed for 10 weeks before caecal enterocytes from each dietary group were isolated and incubated with equimolar mixtures of either [1-14C]16:0 FA and [9,10(n)-3H]16:0 FAlc, or [1-14C]18:1n-9 FA and [9,10(n)-3H]18:1n-9 FAlc. Uptake was measured after 2 h with relative utilization of labelled FAlc and FA calculated as a percentage of uptake. Differences in uptake were observed, with FA showing higher uptake than FAlc, and 18:1 chains a higher uptake than 16:0. A proportion of unesterified FAlc was possibly recovered in the cells, but the majority of FALc was recovered in lipid classes such as triacylglycerol and phospholipids indicating substantial conversion of FAlc to FA followed by esterification. However, incorporation of FA and FAlc into esterified lipids was higher when derived from FA than from FAlc. Twenty-five to fifty percentage of the absorbed 16:0 FA was recovered in TAG fraction of the enterocytes compared with fifteen to seventy-five percentage of 18:1 FA. Twenty to thirty percentage of the absorbed 16:0 FA was recovered in the PC fraction of the enterocytes compared with only five to fifteen percentage of the 18:1 FA. Less than 15% of the fatty chains taken up by the cells was used for energy production, with significantly higher oxidation of 18:1 in enterocytes from fish fed the fish oil diet compared to the Calanus oil diet. However, overall, dietary copepod oil had little effect on FAlc and FA metabolism. Metabolic modification by elongation and/or desaturation was generally low at 1-5% of uptake. We conclude that our hypothesis was generally proved in that the uptake of FAlc by salmon enterocytes was lower than the uptake of FA and that subsequent intracellular metabolism of FAlc was similar to that of FA. However, unesterified FAlc was possibly recovered in the cells suggesting that the conversion to FA may not be concomitant with uptake

Egg quality determinants in cod (Gadus morhua L.): egg performance and lipids in eggs from farmed and wild broodstock

Lipids and essential fatty acids, particularly the highly unsaturated fatty acids, 20:5n-3 (eicosapentaenoic acid; EPA), 22:6n-3 (docosahexaenoic acid; DHA) and 20:4n-6 (arachidonic acid, AA) have been shown to be crucial determinants of marine fish reproduction directly affecting fecundity, egg quality, hatching success, larval malformation and pigmentation. In Atlantic cod (Gadus morhua L.) culture, eggs from farmed broodstock can have much lower fertilisation and hatching rates than eggs from wild broodstock. The present study aimed to test the hypothesis that potential quality and performance differences between eggs from different cod broodstock would be reflected in differences in lipid and fatty acid composition. Thus eggs were obtained from three broodstock, farmed, wild/fed and wild/unfed, and lipid content, lipid class composition, fatty acid composition and pigment content were determined and related to performance parameters including fertilisation rate, symmetry of cell division and survival to hatching. Eggs from farmed broodstock showed significantly lower fertilisation rates, cell symmetry and survival to hatching rates than eggs from wild broodstock. There were no differences in total lipid content or the proportions of the major lipid classes between eggs from the different broodstock. However, eggs from farmed broodstock were characterised by having significantly lower levels of some quantitatively minor phospholipid classes, particularly phosphatidylinositol. There were no differences between eggs from farmed and wild broodstock in the proportions of saturated, monounsaturated and total polyunsaturated fatty acids. The DHA content was also similar. However, eggs from farmed broodstock had significantly lower levels of AA, and consequently significantly higher EPA/AA ratios than eggs from wild broodstock. Total pigment and astaxanthin levels were significantly higher in eggs from wild broodstock. Therefore, the levels of AA and phosphatidylinositol, the predominant AA-containing lipid class, and egg pigment content were positively related to egg quality or performance parameters such as fertilisation and hatching success rates, and cell symmetry

ALMA and Herschel reveal that AGN and main-sequence galaxies have different star formation rate distributions

Using deep Herschel and ALMA observations, we investigate the star formation rate (SFR) distributions of X-ray AGN host galaxies at 0.5<z<1.5 and 1.5<z<4, comparing them to that of normal, star-forming (i.e., "main-sequence", or MS) galaxies. We find 34-55 per cent of AGNs have SFRs at least a factor of two below that of the average MS galaxy, compared to ~15 per cent of all MS galaxies, suggesting significantly different SFR distributions. Indeed, when both are modelled as log-normal distributions, the mass and redshift-normalised SFR distributions of AGNs are roughly twice as broad, and peak ~0.4 dex lower, than that of MS galaxies. However, like MS galaxies, the normalised SFR distribution of AGNs appears not to evolve with redshift. Despite AGNs and MS galaxies having different SFR distributions, the linear-mean SFR of AGNs derived from our distributions is remarkably consistent with that of MS galaxies, and thus with previous results derived from stacked Herschel data. This apparent contradiction is due to the linear-mean SFR being biased by bright outliers, and thus does not necessarily represent a true characterisation of the typical SFR of AGNs

Branch-and-lift algorithm for deterministic global optimization in nonlinear optimal control

This paper presents a branch-and-lift algorithm for solving optimal control problems with smooth nonlinear dynamics and potentially nonconvex objective and constraint functionals to guaranteed global optimality. This algorithm features a direct sequential method and builds upon a generic, spatial branch-and-bound algorithm. A new operation, called lifting, is introduced, which refines the control parameterization via a Gram-Schmidt orthogonalization process, while simultaneously eliminating control subregions that are either infeasible or that provably cannot contain any global optima. Conditions are given under which the image of the control parameterization error in the state space contracts exponentially as the parameterization order is increased, thereby making the lifting operation efficient. A computational technique based on ellipsoidal calculus is also developed that satisfies these conditions. The practical applicability of branch-and-lift is illustrated in a numerical example. © 2013 Springer Science+Business Media New York