F. Glasser

Oilseed Lipid Supplements and Fatty Acid Composition of Cow Milk: A Meta-Analysis

Numerous experiments have studied the use of oilseed supplements in cow diets to alter milk fatty acid (FA) composition, but no quantitative synthesis of these studies is currently available. This article reports a meta-analysis of the response of cow milk FA composition to oilseed lipid supplements from linseed, rapeseed, soybeans, and sunflower seed. First, from a database of 145 oilseed supplementation experiments, we collected the mean FA percentages observed with unsupplemented diets and diets supplemented with the 4 oilseeds given as seeds (after various types of processing), as oils (including Ca salts and amides), or in protected forms. Second, we studied the response of the major milk FA percentages to increasing amounts of supplemental lipids from the 4 oilseeds. Responses were nonsignificant, linear, or quadratic, depending on the FA studied and the supplement. Effects of interfering factors, such as supplement form, forage component of the diet, or lactation stage, were difficult to assess from the available data. Third, we studied the response of the major milk FA percentages to increasing dietary intakes of linoleic or linolenic acids, taken separately. Overall, these results confirm the high plasticity of milk FA composition, with the widest variations being observed in the percentages of medium-chain versus C18 FA, and among the C18 in 18:0, cis-18:1, and trans-18:1. The percentages of the polyunsaturated FA cis-9 cis-12-18:2 and 18:3 were less variable, except when protected lipids (mostly formaldehyde treated) were supplied. However, trans-18:1 and polyunsaturated FA (including conjugated linoleic acid) exhibited the greatest variations when expressed relative to their respective basal values (for unsupplemented diets). Oils, compared with seeds, induced greater percentages of trans-18:1 and tended to decrease C6 to C12 FA more. Intakes of 18:2- and 18:3-rich lipid sources did not differ greatly in their effects on short- and medium-chain FA and trans-18:1 percentages, although the profiles of individual 18:1 and 18:2 isomers in milk differed. This meta-analysis provides quantitative estimates, obtained from the extensive literature produced over more than 40 yr, of the impact of oilseed supplements on milk FA composition.

Digestion of fatty acids in ruminants: a meta-analysis of flows and variation factors: 2. C18 fatty acids.

In ruminants, dietary lipids are extensively hydrogenated by rumen micro-organisms, and the extent of this biohydrogenation is a major determinant of long-chain fatty acid profiles of animal products (milk, meat). This paper reports on the duodenal flows of C18 fatty acids and their absorption in the small intestine, using a meta-analysis of a database of 77 experiments (294 treatments). We established equations for the prediction of duodenal flows of various 18-carbon (C18) fatty acids as a function of the intakes of their precursors and other dietary factors (source and/or technological treatment of dietary lipids). We also quantified the influence of several factors modifying rumen metabolism (pH, forage : concentrate ratio, level of intake, fish oil supplementation). We established equations for the apparent absorption of these fatty acids in the small intestine as a function of their duodenal flows. For all C18 unsaturated fatty acids, apparent absorption was a linear function of duodenal flow. For 18:0, apparent absorption levelled off for high duodenal flows. From this database, with fatty acid flows expressed in g/kg dry matter intake, we could not find any significant differences between animal categories (lactating cows, other cattle or sheep) in terms of rumen metabolism or intestinal absorption of C18 fatty acids.

Long-Chain Fatty Acid Metabolism in Dairy Cows: A Meta-Analysis of Milk Fatty Acid Yield in Relation to Duodenal Flows and De Novo Synthesis

This study is a meta-analysis of the response of milk long-chain fatty acid (FA) yield and composition to lipid supply, based on published experiments reporting duodenal FA flows or duodenal lipid infusions and milk FA composition (i.e., 39 experiments reporting 139 experimental treatments). Analysis of these data underlined the interdependence between milk yields of C18 and short- and medium-chain (C4 to C16) FA. Lipid supplementation (producing an increase in duodenal C18 flow) decreased linearly milk C4 to C16 yield (-0.26 g of C4 to C16 produced per gram of duodenal C18 flow increase) and increased quadratically milk C18 yield. When these 2 effects increased the percentage of C18 in milk FA up to a threshold value (around 52% of total FA), then milk C18 yield was limited by C4 to C16 yield, decreasing the C18 transfer efficiency from duodenum to milk with high-lipid diets. Moreover, for a given duodenal C18 flow, a decrease in milk C4 to C16 yield induced a decrease in milk C18 yield. Despite high variations in C18 transfer efficiency between duodenum and milk, for a given experimental condition, the percentages of C18 FA in milk total C18 could be predicted from their percentages in duodenal C18, and the percentages at the duodenum and in milk were very similar when mammary desaturation was taken into account (i.e., considering the sums of substrates and products of mammary desaturase). The estimated amounts of 18:0, trans-11-, and trans-13-18:1 desaturated by the mammary gland were a linear function of their mammary uptake, and mammary desaturation was responsible for 80, 95, and 81%, respectively, of the yield of their products (i.e., cis-9-18:1; cis-9, trans-11-, and cis-9, trans-13-18:2). Thus, mammary FA desaturation capacity did not seem to be a limiting factor in the experimental conditions published so far.

Effect of linseed fed as rolled seeds, extruded seeds or oil on fatty acid rumen metabolism and intestinal digestibility in cows.

Linseed, a source of linolenic acid, is used in ruminant diets to increase polyunsaturated fatty acids (FA) in animal products. Seed processing is known to have an impact on FA rumen metabolism, but few data are available for linseed. We studied the effect of linseed lipid on ruminal metabolism and intestinal digestibility in cows. Three modes of linseed processing: rolled linseed (RL), extruded linseed (EL) and linseed oil plus linseed meal (LO), supplemented at 7.5% of DM intake, were compared to a control diet (C). Duodenal flows, intestinal digestibility and plasma composition were determined. The duodenal flow of linolenic acid was similar among diets. The sum of t10 and t11-18:1, which were coeluted, was increased with lipid-supplemented diets and represented more than 60% of trans 18:1 for EL and LO diets. The main 18:2 isomers were c9, c12 and t11, c15 among the non-conjugated isomers, and t11, t13 among CLA. Linseed supplementation increased the duodenal flow of unsaturated intermediates of biohydrogenation, and this effect was more pronounced for extruded seeds and oil than for rolled seeds. For most 18-carbon FA, intestinal digestibility was slightly higher for C and LO diets than for RL and EL. Plasma concentrations of non-conjugated 18:2 and linolenic acid were similar among the lipid-supplemented diets. Within diet, profiles of 18:1 isomers (except c9) remained very similar between duodenal and plasma FA.

Prediction of bulk milk fatty acid composition based on farming practices collected through on-farm surveys.

The aim of this study was to predict the fatty acid (FA) composition of bulk milk using data describing farming practices collected via on-farm surveys. The FA composition of 1,248 bulk cow milk samples and the related farming practices were collected from 20 experiments led in 10 different European countries at 44°N to 60°N latitude and sea level to 2,000 m altitude. Farming practice-based FA predictions [coefficient of determination (R(2)) >0.50] were good for C16:0, C17:0, saturated FA, polyunsaturated FA, and odd-chain FA, and very good (R(2) ≥0.60) for trans-11 C18:1, trans-10 + trans-11 C18:1, cis-9,trans-11 conjugated linoleic acid, total trans FA, C18:3n-3, n-6:n-3 ratio, and branched-chain FA. Fatty acids were predicted by cow diet composition and by the altitude at which milk was produced, whereas animal-related factors (i.e., lactation stage, breed, milk yield, and proportion of primiparous cows in the herd) were not significant in any of the models. Proportion of fresh herbage in the cow diet was the main predictor, with the highest effect in almost all FA models. However, models built solely on conserved forage-derived samples gave good predictions for odd-chain FA, branched-chain FA, trans-10 C18:1 and C18:3n-3 (R(2) ≥0.46, 0.54, 0.52, and 0.70, respectively). These prediction models could offer farmers a valuable tool to help improve the nutritional quality of the milk they produce.

Effects of fish oil and starch added to a diet containing sunflower-seed oil on dairy goat performance, milk fatty acid composition and in vivo Δ9-desaturation of [13C]vaccenic acid.

The potential benefits on human health have prompted an interest in developing nutritional strategies for specifically increasing rumenic acid (RA) in ruminant milk. The aims of the present study were to (i) compare two dietary treatments with lipid supplements on milk yield and composition, (ii) measure the in vivo delta9-desaturation of vaccenic acid (VA) to RA using 13C-labelled VA and (iii) determine the effect of the dietary treatments on this variable. Treatments were 90 g sunflower-seed oil (SO) per d or 60 g sunflower-seed oil and 30 g fish oil per d plus additional starch (SFO), in a grassland hay-based diet given to eight Alpine goats in a 2 x 2 cross-over design with 21 d experimental periods. Milk yield and composition were similar between treatments. Goats fed SFO had higher milk 6 : 0-16 : 0 concentration, lower milk sigmaC18 concentrations and showed no effect on milk VA and RA, compared with SO. At the end of the experiment, intravenous injection of 1.5 g [13C]VA followed by measurements of milk lipid 13C enrichment showed that in vivo 31.7 and 31.6 % of VA was delta9-desaturated into milk RA in the caprine with the SO and SFO treatments, respectively. The expression of genes encoding for delta9-desaturase (or stearoyl-CoA desaturase; SCD1, SCD5) in mammary tissues and four milk delta9-desaturation ratios were similar between treatments. In conclusion, the present study provides the first estimates of in vivo endogenous synthesis of RA (63-73 % of milk RA) from VA in goats, and shows no difference between the two lipid supplements compared.

Combined effects of trans-10,cis-12 conjugated linoleic acid, propionate, and acetate on milk fat yield and composition in dairy cows

Diets inducing milk fat depression (MFD) are known to alter ruminal lipid metabolism, leading to the formation of specific isomers [such as trans-10,cis-12 conjugated linoleic acid (CLA)] that inhibit milk fat synthesis in lactating dairy cows. However, ruminal outflow of these isomers does not fully account for the decreases in milk fat synthesis observed during diet-induced MFD. The high-concentrate diets inducing MFD also induce a greater production of propionate, suggesting a possible inhibition of milk fat by propionate associated with trans-10,cis-12-CLA during MFD. The present experiment aimed to study the combined effects of propionate and trans-10,cis-12-CLA (both inhibitors of milk fat synthesis) on milk fat secretion and the effects of the combination of 2 nutrients with opposite effects (acetate and propionate). Six Holstein cows were used in a 6×6 Latin square design with 21-d periods (14 d of nutrient infusion). The treatments were control; ruminal infusion of 1,500 g/d of acetate (A); ruminal infusion of 800 g/d of propionate (P); duodenal infusion of 1.60 g/d of trans-10,cis-12-CLA (CLA); ruminal infusion of 750 g/d of acetate+400 g/d of propionate (A+P); and duodenal infusion of 1.60 g/d of trans-10,cis-12-CLA+ruminal infusion of 800 g/d of propionate (CLA+P). The amounts of nutrients infused were chosen to induce a similar variation in milk fat content. Treatments A and P decreased dry matter intake. Compared with the control, P and CLA treatments decreased milk fat content and yield by 9% and 15% on average. Treatment A increased milk fat content by 6.5% but did not modify milk fat yield (because of a decrease in milk yield). The effects of A and P, and CLA and P on milk fat and fatty acid percentages and yield were additive (A+P and CLA+P treatments). With a same dose of trans-10,cis-12-CLA, the additional supply of propionate induced a decrease in milk fat 40% higher than that induced by trans-10,cis-12-CLA alone. The milk fatty acid profile obtained with CLA+P was similar to those observed with high-concentrate diets inducing MFD. In conclusion, under our experimental conditions, the effects of the 3 nutrients were additive on mammary lipogenesis, regardless of their separate effects. We also show that propionate could contribute to the milk fat reductions unaccounted for by trans-10,cis-12-CLA during MFD induced by high-concentrate diets.

In vivo production and molar percentages of volatile fatty acids in the rumen: a quantitative review by an empirical approach

Despite their major contribution to the energy supply of ruminants, the production of volatile fatty acids (VFA) in the rumen is still poorly predicted by rumen models. We have developed an empirical approach, based on the interpretation of large bibliographic databases gathering published in vivo measurements of ruminal VFA production rate (PR), rates of duodenal and faecal digestion and molar percentages of VFA in the rumen. These databases, covering a wide range of intake levels and dietary composition, were studied by meta-analysis using within-experiment models. We established models to quantify response laws of total VFA-PR and individual VFA molar percentages in the rumen to variations in intake level and dietary composition. The rumen fermentable organic matter (RfOM) intake, estimated from detailed knowledge of the chemical composition of diets according to INRA Feed Tables, appears as an accurate explanatory variable of measured total VFA-PR, with an average increment of 8.03 ± 0.64 mol total VFA/kg RfOM intake. Similar results were obtained when total VFA-PR was estimated from measured apparent RfOM (total VFA-PR/RfOM averaging 8.3 ± 1.2 mol/kg). The VFA molar percentages were related to dry matter intake and measured digestible organic matter (OM), digestible NDF and rumen starch digestibility, with root mean square error of 1.23, 1.45, 0.88 and 0.41 mol/100 mol total VFA for acetate, propionate, butyrate and minor VFA, respectively, with no effect of pH on the residuals. Stoichiometry coefficients were calculated from the slopes of the relationships between individual VFA production (estimated from measured apparent RfOM and individual VFA molar percentages) and measured fermented fractions. Coefficients averaged, respectively, 66, 17, 14 and 3 mol/100 mol for NDF; 41, 44, 12 and 4 mol/100 mol for starch; and 46, 35, 13 and 6 mol/100 mol for crude protein. Their use to predict VFA molar percentages appear relevant for most dietary conditions, that is, when the digested NDF/digested OM ratio exceeded 0.12. This study provides a quantitative review on VFA yield in the rumen. It contributes to the development of feed evaluation systems based on nutrient fluxes.

Short communication: Diet-induced variations in milk fatty acid composition have minor effects on the estimated melting point of milk fat in cows, goats, and ewes: Insights from a meta-analysis

In ruminants, the ability to maintain milk fat melting point within physiological values could play a role in the regulation of milk fat secretion when milk fatty acid (FA) composition varies, such as in response to feeding factors. However, the relationship between milk fat fluidity and changes in milk FA composition is difficult to study experimentally. A meta-analysis was therefore conducted to compare the magnitude of diet-induced variations in milk FA composition and the calculated melting point of milk FA (used as a proxy to estimate the variations in the melting point of milk fat) in 3 dairy ruminant species (cow, goat, and sheep). The coefficient of variation (CV), a scale-free measure of statistical dispersion, was used to compare the variability of criteria differing in their order of magnitude. The analysis of a database of milk FA profiles from cows, goats, and sheep fed different dietary treatments (unsupplemented diets and diets supplemented with lipids rich in oleic acid, linoleic acid, linolenic acid, or C20-22 polyunsaturated FA) revealed that the variability of the calculated melting point of milk FA was narrow (CV of 5%) compared with the variability of milk FA percentages (CV of 18 to 72%). The regulation of the melting point of milk fat is thus probably involved in the control of diet-induced variations in milk fat secretion. The calculated melting point of ewe milk FA was approximately 3°C lower than that of goats or cows across all types of diets, which might be linked to differences in milk fat content (higher in sheep) or the structure of milk triacylglycerides among these species. Lipid supplementation increased the calculated melting point of C18 FA in milk, whereas that of total FA was significantly reduced by supplements rich in oleic, linoleic, and linolenic acids but not C20-22 polyunsaturated FA. However, the slight effects of dietary treatments on the calculated melting point of milk FA did not differ between cows, goats, and ewes.

Response of milk fat concentration and yield to nutrient supply in dairy cows.

Dietary changes alter dairy cow milk fat concentration (MFC) and yield (MFY) through modifications in the supply of nutrients, which act as precursors or inhibitors of mammary fat synthesis. The current models used to formulate dairy cow diets cannot predict changes in milk fat. The knowledge of the effects of the nutrients on milk fat would help to progress toward this prediction. To this end, we quantified and compared the milk fat responses to variations in the supply of seven nutrients derived from digestion: volatile fatty acids, glucose, proteins, long-chain fatty acids (LCFA) and t10,c12-conjugated linoleic acid (CLA). A database was compiled from studies involving digestive infusions of these nutrients in dairy cows. It included 147 comparisons between a nutrient infusion and a control treatment. The nutrient infusions were limited to the range of physiological variations to mimic nutrient changes after dietary modifications. We established models for the response of MFC, MFY and milk fatty acid (FA) composition to the supply of each nutrient. MFC and MFY responses to the nutrients were significant and linear, except for the MFC response to glucose that was curvilinear. The nutrients differed in their effects on MFC and MFY: acetate, butyrate and LCFA increased MFC and MFY, whereas propionate, glucose and t10,c12-CLA decreased them. Protein infusions increased MFY and decreased MFC because of an increase in milk yield. The effects of numerous interfering factors related to animals, diets or experimental conditions were tested on the residuals of the response models. The responses of milk FA percentages are also provided. When adjusted to the in vivo variations in the nutrients observed after dietary changes, the effects of the different nutrients were moderate. Finally, this study showed that several of these nutrients could contribute to the changes in milk fat production and composition observed after dietary changes. This is a first step toward predicting milk fat response to changes in nutrient supply.