M. Doreau

Effect of different types of forages, animal fat or marine oils in cow’s diet on milk fat secretion and composition, especially conjugated linoleic acid (CLA) and polyunsaturated fatty acids

This review summarises the known effects of forages, animal fats or marine oils on bovine milk fat secretion and composition. Special attention is given to fatty acids that could play a positive role for human health, such as butyric acid, oleic acid, C18 to C22 polyunsaturated fatty acids and conjugated linoleic acid (CLA). The efficiency of the transfer of n-3 polyunsaturated fatty acids from diet to milk is reviewed. Milk fat from pasture fed cows seems to be higher in linolenic acid than milk fat from cows receiving preserved grass or maize, but the magnitude of this difference is limited. Indirect comparisons show that milk fat from maize silage diets is richer in short-chain FA and linoleic acid when compared to grass silage diets. Compared to fresh grass, grass silage favours myristic and palmitic acids at the expense of mono- and polyunsaturated FA, including CLA. Protected tallow allows for a large increase in milk fat yield, and in the percentage of milk stearic and oleic acids, at the expense of medium chain FA. Non-protected tallow has a similar effect on medium chain FA without increasing so much C18 FA yield, which explains that it does not increase milk fat yield. Dose–response curves of milk CLA are reviewed for marine oil supplements, as well as the relationship between milk CLA and trans-C18:1 contents. The potential of marine oil supplementation to increase the mean CLA content in cow milk fat is large (more than 300% above basal values). A specific role for dietary C20:5 n-3 in the sharp decrease in milk fat secretion after fish oil supplementation is suggested. However, there is a need to evaluate how the different feeding strategies could change the other aspects of milk fat quality, such as taste, oxidative stability or manufacturing value.

Methane mitigation in ruminants: from microbe to the farm scale

Decreasing enteric methane (CH4) emissions from ruminants without altering animal production is desirable both as a strategy to reduce global greenhouse gas (GHG) emissions and as a means of improving feed conversion efficiency. The aim of this paper is to provide an update on a selection of proved and potential strategies to mitigate enteric CH4 production by ruminants. Various biotechnologies are currently being explored with mixed results. Approaches to control methanogens through vaccination or the use of bacteriocins highlight the difficulty to modulate the rumen microbial ecosystem durably. The use of probiotics, i.e. acetogens and live yeasts, remains a potentially interesting approach, but results have been either unsatisfactory, not conclusive, or have yet to be confirmed in vivo. Elimination of the rumen protozoa to mitigate methanogenesis is promising, but this option should be carefully evaluated in terms of livestock performances. In addition, on-farm defaunation techniques are not available up to now. Several feed additives such as ionophores, organic acids and plant extracts have also been assayed. The potential use of plant extracts to reduce CH4 is receiving a renewed interest as they are seen as a natural alternative to chemical additives and are well perceived by consumers. The response to tannin- and saponin-containing plant extracts is highly variable and more research is needed to assess the effectiveness and eventual presence of undesirable residues in animal products. Nutritional strategies to mitigate CH4 emissions from ruminants are, without doubt, the most developed and ready to be applied in the field. Approaches presented in this paper involve interventions on the nature and amount of energy-based concentrates and forages, which constitute the main component of diets as well as the use of lipid supplements. The possible selection of animals based on low CH4 production and more likely on their high efficiency of digestive processes is also addressed. Whatever the approach proposed, however, before practical solutions are applied in the field, the sustainability of CH4 suppressing strategies is an important issue that has to be considered. The evaluation of different strategies, in terms of total GHG emissions for a given production system, is discussed.

Digestion and utilisation of fatty acids by ruminants

Abstract This review deals with the quantitative and qualitative changes in fatty acids (FA) throughout the total digestive tract of ruminants. Special attention is paid to the causes of variation in the extent to which different mechanisms contribute to the ruminal metabolism and intestinal digestion of FA. Most results obtained with diets not supplemented with lipids show that the FA flow leaving the rumen is higher than FA intake. This is due to bacterial synthesis of FA in the rumen. With diets supplemented with lipids, the FA balance at the end of the rumen is often negative. The cause of this apparent disappearance of FA is not known. In the rumen, lipids are first hydrolysed to a very large extent; then unsaturated FA are hydrogenated. Hydrogenation is almost complete for linolenic acid, and amounts to between 60 and 95% for linoleic acid. This proportion decreases when the level of concentrates increases in the diet. Digestibility of FA in the small intestine ranges from 70 to 90% and is not related to the level of FA intake. Contrary to the situation in monogastric animals, there are only moderate differences in the digestibility of individual FA. It appears to be higher for palmitic and stearic acids than for other saturated FA, and for oleic and linoleic acids than for stearic and linolenic acids. In the large intestine, there is synthesis of FA which are probably not absorbed.

Digestion and metabolism of dietary fat in farm animals

Fat digestion and metabolism differ widely between animal species. In ruminants, dietary fats are hydrogenated in the rumen before intestinal absorption so that absorbed fatty acids (FA) are more saturated than dietary FA. In non-ruminants, intestinal FA digestibility depends on the level of saturation of dietary FA. Fat supplementation of the diet of cows decreases milk protein and has a variable effect on milk fat, depending on the source of dietary lipids. When encapsulated lipids are used, the linoleic acid content of milk is increased, but the organoleptic quality of milk may be altered. Supplementary lipids are incorporated into non-ruminant body fat, whereas de novo lipogenesis is reduced. There is a close relationship between the nature of dietary FA and nonruminant body FA.

Methane output and diet digestibility in response to feeding dairy cows crude linseed, extruded linseed, or linseed oil1

This experiment studied the effect of 3 forms of presentation of linseed fatty acids (FA) on methane output using the sulfur hexafluoride tracer technique, total tract digestibility, and performance of dairy cows. Eight multiparous lactating Holstein cows (initial milk yield 23.4 +/- 2.2 kg/d) were assigned to 4 dietary treatments in a replicated 4 x 4 Latin square design: a control diet (C) consisting of corn silage (59%), grass hay (6%), and concentrate (35%) and the same diet with crude linseed (CLS), extruded linseed (ELS), or linseed oil (LSO) at the same FA level (5.7% of dietary DM). Each experimental period lasted 4 wk. All the forms of linseed FA significantly decreased daily CH(4) emissions (P < 0.001) but to different extents (-12% with CLS, -38% with ELS, -64% with LSO) compared with C. The same ranking among diets was observed for CH(4) output expressed as a percentage of energy intake (P < 0.001) or in grams per kilogram of OM intake (P < 0.001). Methane production per unit of digested NDF was similar for C, CLS, and ELS but was less for LSO (138 vs. 68 g/kg of digested NDF, respectively; P < 0.001). Measured as grams per kilogram of milk or fat-corrected milk yield, methane emission was similar for C and CLS and was less for ELS and LSO (P < 0.001), LSO being less than ELS (P < 0.01). Total tract NDF digestibility was significantly less (P < 0.001) for the 3 supplemented diets than for C (-6.8% on average; P < 0.001). Starch digestibility was similar for all diets (mean 93.5%). Compared with C, DMI was not modified with CLS (P > 0.05) but was decreased with ELS and LSO (-3.1 and -5.1 kg/d, respectively; P < 0.001). Milk yield and milk fat content were similar for LSO and ELS but less than for C and CLS (19.9 vs. 22.3 kg/d and 33.8 vs. 43.2 g/kg, on average, respectively; P < 0.01 and P < 0.001). Linseed FA offer a promising dietary means to depress ruminal methanogenesis. The form of presentation of linseed FA greatly influences methane output from dairy cows. The negative effects of linseed on milk production will need to be overcome if it is to be considered as a methane mitigation agent. Optimal conditions for the utilization of linseed FA in ruminant diets need to be determined before recommending its use for the dairy industry.

TARGETS AND PROCEDURES FOR ALTERING RUMINANT MEAT AND MILK LIPIDS

Beef and dairy products suffer from a negative health image, related to the nature of their lipid fraction. Rumen lipid metabolism involves the presence of saturated lipids in ruminant tissues. Lipolysis, fatty acid biohydrogenation and formation of microbial fatty acids in the rumen and their effects on rumen outflow of fatty acids are discussed. Special emphasis is given to the formation of trans-fatty acids and the possibilities of decreasing biohydrogenation. Small differences in intestinal digestibilities of fatty acids are mentioned, followed by a discussion on transfer of absorbed fatty acids into milk and adipose tissue lipids. The preferential retention of polyunsaturated fatty acids as well as the balance between synthesis and incorporation of fatty acids in tissues is described. Dietary means for the modification of milk fat are listed, with special emphasis on the possibilities for enrichment in polyunsaturated fatty acids and the presence of conjugated linoleic acids. A description of the nature and development of fat depots in beef cattle is followed by a discussion of breed, conformation and feed effects on adipose tissue distribution and fatty acid composition. Special emphasis is given to the very lean Belgian Blue double-muscled breed. The review ends with a consideration of the limits to the modification of ruminant fats, involving considerations of consumer acceptance as well as animal welfare and environmental effects.

Milk fatty acids in dairy cows fed whole crude linseed, extruded linseed, or linseed oil, and their relationship with methane output.

This experiment studied the effect of 3 different physical forms of linseed fatty acids (FA) on cow dairy performance, milk FA secretion and composition, and their relationship with methane output. Eight multiparous, lactating Holstein cows were assigned to 1 of 4 dietary treatments in a replicated 4 x 4 Latin square design: a control diet (C) based on corn silage (59%) and concentrate (35%), and the same diet supplemented with whole crude linseed (CLS), extruded linseed (ELS), or linseed oil (LSO) at the same FA level (5% of dietary dry matter). Each experimental period lasted 4 wk. Dry matter intake was not modified with CLS but was lowered with both ELS and LSO (-3.1 and -5.1 kg/d, respectively) compared with C. Milk yield and milk fat content were similar for LSO and ELS but lower than for C and CLS (19.9 vs. 22.3 kg/d and 33.8 vs. 43.2 g/kg, on average, respectively). Compared with diet C, CLS changed the concentrations of a small number of FA; the main effects were decreases in 8:0 to 16:0 and increases in 18:0 and cis-9 18:1. Compared with diet C (and CLS in most cases), LSO appreciably changed the concentrations of almost all the FA measured; the main effects were decreases in FA from 4:0 to 16:0 and increases in 18:0, trans-11 16:1, all cis and trans 18:1 (except trans-11 18:1), and nonconjugated trans 18:2 isomers. The effect of ELS was either intermediate between those of CLS and LSO or similar to LSO with a few significant exceptions: increases in 17:0 iso; 18:3n-3; trans-11 18:1; cis-9, trans-11 conjugated linoleic acid; and trans-11, trans-13 conjugated linoleic acid and a smaller increase in cis-9 18:1. The most positive correlations (r = 0.87 to 0.91) between milk FA concentrations and methane output were observed for saturated FA from 6:0 to 16:0 and for 10:1, and the most negative correlations (r = -0.86 to -0.90) were observed for trans-16+cis-14 18:1; cis-9, trans-13 18:2; trans-11 16:1; and trans-12 18:1. Thus, milk FA profile can be considered a potential indicator of in vivo methane output in ruminants.

Effect of dietary lipids on nitrogen metabolism in the rumen: a review

This paper deals with the influence of supplementary lipids in the diet on protein degradation and microbial synthesis in the rumen, through the effect of lipids on the microbial ecosystem. Global effects of lipids can be summarized by a trend to decrease ammonia ruminal concentration but duodenal non-ammonia nitrogen flow is not modified. The two components of this flow, non-microbial and microbial flows, generally are not modified so that it can be deduced that modifications of degradation and microbial synthesis due to lipid addition are of low extent. The efficiency of microbial synthesis depends on the nature of fatty acids and is increased especially when organic matter ruminal digestibility is depressed; this can be related to the decrease in protozoa number in the rumen.

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.

Lipid metabolism of liquid-associated and solid-adherent bacteria in rumen contents of dairy cows offered lipid-supplemented diets

The lipid distribution and fatty acid (FA) composition of total lipids, polar lipids and free fatty acids (FFA) were determined in liquid-associated bacteria (LAB) and solid-adherent bacteria (SAB) isolated from the rumen contents of seven dairy cows fitted with rumen fistulas. Two experiments, arranged according to a 4 x 4 and 3 x 3 Latin Square design, were performed using two basal diets consisting of one part hay and one part barley-based concentrate, and five lipid-supplemented diets consisting of the basal diet plus (g/kg dry matter):53 or 94 rapeseed oil, 98 tallow, 87 soya-bean oil or 94 palmitostearin. For all diets used, total lipids were 1.7-2.2 times higher in SAB than in LAB (P less than 0.05); this probably resulted from a preferential incorporation of dietary FA absorbed onto food particles. Addition of oil or fat to the diets did not modify the polar lipid content but increased the FFA content of SAB and LAB by 150%. Lipid droplets were observed in the cytoplasm in SAB and LAB using transmission electron microscopy, which suggested that part of the additional FFA was really incorporated into the intracellular FFA rather than associated with the cell envelope by physical adsorption. Linoleic acid was specifically incorporated into the FFA of SAB, which emphasized the specific role of this bacterial compartment in the protection of this acid against rumen biohydrogenation.