A.L. Lock

Modifying Milk Fat Composition of Dairy Cows to Enhance Fatty Acids Beneficial to Human Health

There is increased consumer awareness that foods contain microcomponents that may have beneficial effects on health maintenance and disease prevention. In milk fat these functional food components include EPA, DHA, and CLA. The opportunity to enhance the content of these FA in milk has improved as a result of recent advances that have better defined the interrelationships between rumen fermentation, lipid metabolism, and milk fat synthesis. Dietary lipids undergo extensive hydrolysis and biohydrogenation in the rumen. Milk fat is predominantly TG, and de novo FA synthesis and the uptake of circulating FA contribute nearly equal amounts (molar basis) to the FA in milk fat. Transfer of dietary EPA and DHA to milk fat is very low (<4%); this is, to a large extent, related to their extensive biohydrogenation in the rumen, and also partly due to the fact that they are not transported in the plasma lipid fractions that serve as major mammary sources of FA uptake (TG and nonesterified FA). Milk contains over 20 isomers of CLA but the predominant one is cis-9,trans-11 (75–90% of total CLA). Biomedical studies with animal models have shown that this isomer has anticarcinogenic and anti-atherogenic activities. cis-9,trans-11-CLA is produced as an intermediate in the rumen biohydrogenation of linoleic acid but not of linolenic acid. However, it is only a transient intermediate, and the major source of milk fat CLA is from endogenous synthesis. Vaccenic acid, produced as a rumen biohydrogenation intermediate from both linoleic acid and linolenic acid, is the substrate, and Δ9-desaturase in the mammary gland and other tissues catalyzes the reaction. Diet can markedly affect milk fat CLA content, and there are also substantial differences among individual cows. Thus, strategies to enhance milk fat CLA involve increasing rumen outflow of vaccenic acid and increasing Δ9-desaturase activity, and through these, several-fold increases in the content of CLA in milk fat can be routinely achieved. Overall, concentrations of CLA, and to a lesser extent EPA and DHA, can be significantly enhanced through the use of diet formulation and nutritional management of dairy cows.

Biosynthesis of conjugated linoleic acid in ruminants and humans.

Publisher Summary This chapter discusses the biosynthesis of conjugated linoleic acid (CLA) in ruminants and humans. Conjugated linoleic acid is a mixture of positional and geometric isomers of linoleic acid with a conjugated double-bond system. Because of its potential to improve human health, there is great interest to increase the amount of CLA in the human food supply. This has caused great effort to be expended toward increasing the concentration of CLA, and more specifically rumenic acid (RA), in the milk and tissues of ruminant foods because these are the predominant source of CLA in human diets. RA is the predominant CLA isomer present in ruminant products, and the major source of its occurrence is endogenous synthesis via desaturation of vaccenic acid (VA) by ∆-9-desaturase. The chapter focuses on improving the understanding of biohydrogenation in the rumen and examining milk and tissue CLA responses to a range of diets. The diversity of various BH intermediates in digesta, milk, and tissues indicates the complexity of the BH processes as a whole and the population dynamics of the ruminal bacteria involved. Predicting the outcome of changes in the diet is complicated by the interactions of the ruminal environment, substrate supply and forms of dietary lipids, all of which influence the BH process simultaneously.

A reappraisal of the impact of dairy foods and milk fat on cardiovascular disease risk.

BackgroundThis review provides a reappraisal of the potential effects of dairy foods, including dairy fats, on cardiovascular disease (CVD)/coronary heart disease (CHD) risk. Commodities and foods containing saturated fats are of particular focus as current public dietary recommendations are directed toward reducing the intake of saturated fats as a means to improve the overall health of the population. A conference of scientists from different perspectives of dietary fat and health was convened in order to consider the scientific basis for these recommendations.AimsThis review and summary of the conference focus on four key areas related to the biology of dairy foods and fats and their potential impact on human health: (a) the effect of dairy foods on CVD in prospective cohort studies; (b) the impact of dairy fat on plasma lipid risk factors for CVD; (c) the effects of dairy fat on non-lipid risk factors for CVD; and (d) the role of dairy products as essential contributors of micronutrients in reference food patterns for the elderly.ConclusionsDespite the contribution of dairy products to the saturated fatty acid composition of the diet, and given the diversity of dairy foods of widely differing composition, there is no clear evidence that dairy food consumption is consistently associated with a higher risk of CVD. Thus, recommendations to reduce dairy food consumption irrespective of the nature of the dairy product should be made with caution.

Effects of Ruminant trans Fatty Acids on Cardiovascular Disease and Cancer: A Comprehensive Review of Epidemiological, Clinical, and Mechanistic Studies

There are 2 predominant sources of dietary trans fatty acids (TFA) in the food supply, those formed during the industrial partial hydrogenation of vegetable oils (iTFA) and those formed by biohydrogenation in ruminants (rTFA), including vaccenic acid (VA) and the naturally occurring isomer of conjugated linoleic acid, cis-9, trans-11 CLA (c9,t11-CLA). The objective of this review is to evaluate the evidence base from epidemiological and clinical studies to determine whether intake of rTFA isomers, specifically VA and c9,t11-CLA, differentially affects risk of cardiovascular disease (CVD) and cancer compared with iTFA. In addition, animal and cell culture studies are reviewed to explore potential pro- and antiatherogenic mechanisms of VA and c9,t11-CLA. Some epidemiological studies suggest that a positive association with coronary heart disease risk exists between only iTFA isomers and not rTFA isomers. Small clinical studies have been conducted to establish cause-and-effect relationships between these different sources of TFA and biomarkers or risk factors of CVD with inconclusive results. The lack of detection of treatment effects reported in some studies may be due to insufficient statistical power. Many studies have used doses of rTFA that are not realistically attainable via diet; thus, further clinical studies are warranted. Associations between iTFA intake and cancer have been inconsistent, and associations between rTFA intake and cancer have not been well studied. Clinical studies have not been conducted investigating the cause-and-effect relationship between iTFA and rTFA intake and risk for cancers. Further research is needed to determine the health effects of VA and c9,t11-CLA in humans.

Nutrigenomics, Rumen-Derived Bioactive Fatty Acids, and the Regulation of Milk Fat Synthesis

Mammary synthesis of milk fat continues to be an active research area, with significant advances in the regulation of lipid synthesis by bioactive fatty acids (FAs). The biohydrogenation theory established that diet-induced milk fat depression (MFD) in the dairy cow is caused by an inhibition of mammary synthesis of milk fat by specific FAs produced during ruminal biohydrogenation. The first such FA shown to affect milk fat synthesis was trans-10, cis-12 conjugated linoleic acid, and its effects have been well characterized, including dose-response relationships. During MFD, lipogenic capacity and transcription of key mammary lipogenic genes are coordinately down-regulated. Results provide strong evidence for sterol response element-binding protein-1 (SREBP1) and Spot 14 as biohydrogenation intermediate responsive lipogenic signaling pathway for ruminants and rodents. The study of MFD and its regulation by specific rumen-derived bioactive FAs represents a successful example of nutrigenomics in present-day animal nutrition research and offers several potential applications in animal agriculture.

Nutrition, Metabolism, and Fertility in Dairy Cows: 1. Dietary Energy Source and Ovarian Function

In previous studies, high plasma insulin was associated with earlier resumption of postpartum estrous cycles in dairy cows. The objective of this experiment was to quantify hormonal and ovarian responses to dietary starch and fat contents. Thirty cows were fed on a standard diet from calving until 40 d in milk (DIM) and then 6 cows were allocated to each of 5 isoenergetic diets containing 231, 183, 159, 135, and 87 g of starch and 39, 42, 43, 45, and 48 g of fat/kg of dry matter (DM) for diets 1 to 5, respectively, until 70 DIM. Estrus was synchronized at 60 DIM. Between 60 and 70 DIM, energy intake, milk yield, and energy balance were similar among diet groups. Plasma insulin-to-glucagon ratio increased with increasing dietary starch and decreasing dietary fat concentrations, reaching a break point at 159 g of starch, 43 g of fat/kg of DM (diets 1 to 5: mean 3.86, 3.78, 3.59, 2.98, 2.06 +/- standard error 0.22). Growth hormone, insulin-like growth factor-I, and leptin did not vary among diets. The greatest dietary starch concentration was associated with elevated plasma urea-N (diets 1 to 5: mean 3.69, 3.01, 2.94, 2.95, 2.75, +/- standard error 0.13 mmol/L, respectively) and delayed postovulatory progesterone increase (progesterone at 3 to 5 d postovulation for diets 1 to 5: mean 2.7, 5.9, 4.2, 5.6, 4.3 +/- standard error 0.9 ng/mL, respectively). The number of small (<5 mm) ovarian follicles was positively related to starch intake (r = 0.381) and plasma insulin concentration (r = 0.402). It is concluded that to maintain adequate insulin-to-glucagon ratio in cows at the start of the breeding period, dietary starch concentration should be above 160 g/kg of DM and dietary fat below 44 g/kg of DM, and this should have a positive effect on ovarian function.

Feeding a C16:0-enriched fat supplement increased the yield of milk fat and improved conversion of feed to milk

Previous work has indicated that dietary palmitic acid (C16:0) may increase milk fat yield. The effect of a dietary C16:0-enriched fat supplement on feed intake, yield of milk and milk components, and feed efficiency was evaluated in an experiment with a crossover arrangement of treatments with 25-d periods. A fermentable starch challenge on the last 4d of each period was utilized as a split-plot within period. Sixteen mid-lactation Holstein cows (249 ± 33 d in milk) were assigned randomly to treatment sequence. Treatments were either a C16:0-enriched (~85% C16:0) fat supplement (fatty acid treatment, FAT, 2% dry matter) or a control diet (CON) containing no supplemental fat. Diets containing dry ground corn grain were fed from d 1 through 21 of each period. On the last 4d of each period, dry ground corn was replaced by high-moisture corn grain on an equivalent dry matter basis to provide a fermentable starch challenge. Response variables were averaged for d 18 to 21 (immediately before the fermentable starch challenge) and d 22 to 25 (during the fermentable starch challenge). We observed no treatment effects on milk yield or milk protein yield. The FAT treatment increased milk fat concentration from 3.88 to 4.16% and fat yield from 1.23 to 1.32 kg/d compared with CON. The FAT treatment decreased dry matter intake by 1.4 kg/d and increased conversion of feed to milk (3.5% fat-corrected milk yield/dry matter intake) by 8.6% compared with CON. The increase in milk fat yield by FAT was entirely accounted for by a 27% increase in 16-carbon fatty acid output into milk. Yields of de novo and preformed fatty acids were not affected by FAT relative to CON. The fermentable starch challenge did not affect milk fat concentration or yield. Results demonstrate the potential for a dietary C16:0-enriched fat supplement to improve milk fat concentration and yield as well as efficiency of conversion of feed to milk. Further studies are required to verify and extend these results and to determine whether responses are similar across different diets and levels of milk production.

Palmitic acid increased yields of milk and milk fat and nutrient digestibility across production level of lactating cows

The effects of palmitic acid supplementation on feed intake, digestibility, and metabolic and production responses were evaluated in dairy cows with a wide range of milk production (34.5 to 66.2 kg/d) in a crossover design experiment with a covariate period. Thirty-two multiparous Holstein cows (151 ± 66 d in milk) were randomly assigned to treatment sequence within level of milk production. Treatments were diets supplemented (2% of diet DM) with palmitic acid (PA; 99% C16:0) or control (SH; soyhulls). Treatment periods were 21 d, with the final 4 d used for data and sample collection. Immediately before the first treatment period, cows were fed the control diet for 21 d and baseline values were obtained for all variables (covariate period). Milk production measured during the covariate period (preliminary milk yield) was used as covariate. In general, no interactions were detected between treatment and preliminary milk yield for the response variables measured. The PA treatment increased milk fat percentage (3.40 vs. 3.29%) and yields of milk (46.0 vs. 44.9 kg/d), milk fat (1.53 vs. 1.45 kg/d), and 3.5% fat-corrected milk (44.6 vs. 42.9 kg/d), compared with SH. Concentrations and yields of protein and lactose were not affected by treatment. The PA treatment did not affect dry matter (DM) intake or body weight, tended to decrease body condition score (2.93 vs. 2.99), and increased feed efficiency (3.5% fat-corrected milk/DM intake; 1.60 vs. 1.54), compared with SH. The PA treatment increased total-tract digestibility of neutral detergent fiber (39.0 vs.35.7%) and organic matter (67.9 vs. 66.2%), but decreased fatty acid (FA) digestibility (61.2 vs. 71.3%). As total FA intake increased, total FA digestibility decreased (R(2) = 0.51) and total FA absorbed increased (quadratic R(2) = 0.82). Fatty acid yield response, calculated as the additional FA yield secreted in milk per unit of additional FA intake, was 11.7% for total FA and 16.5% for C16:0 plus cis-9 C16:1 FA. The PA treatment increased plasma concentration of nonesterified FA (101 vs. 90.0 μEq/L) and cholecystokinin (19.7 vs. 17.6 pmol/L), and tended to increase plasma concentration of insulin (10.7 vs. 9.57 μ IU/mL). Results show that palmitic acid fed at 2% of diet DM has the potential to increase yields of milk and milk fat, independent of production level without increasing body condition score or body weight. However, a small percentage of the supplemented FA was partitioned to milk.

A Conjugated Linoleic Acid Supplement Containing Trans-10, Cis-12 Conjugated Linoleic Acid Reduces Milk Fat Synthesis in Lactating Goats

The effect of conjugated linoleic acid (CLA) supplements containing trans-10, cis-12 for reducing milk fat synthesis has been well described in dairy cows and sheep. Studies on lactating goats, however, remain inconclusive. Therefore, the current study investigated the efficacy of a lipid-encapsulated trans-10, cis-12 CLA supplement (LE-CLA) on milk production and milk fatty acid profile in dairy goats. Thirty multiparous Alpine lactating goats in late lactation were used in a 3 x 3 Latin square design (14-d treatment periods separated by 14-d intervals). Does were fed a total mixed ration of Bermuda grass hay, dehydrated alfalfa pellets, and concentrate. Does were randomly allocated to 3 treatments: A) unsupplemented (control), B) supplemented with 30 g/d of LE-CLA (low dose; CLA-1), and C) supplemented with 60 g/d of LE-CLA (high dose; CLA-2). Milk yield, dry matter intake, and milk protein content and yield were unaffected by treatment. Compared with the control, milk fat yield was reduced 8% by the CLA-1 treatment and 21% by the CLA-2 treatment, with milk fat content reduced 5 and 18% by the CLA-1 and CLA-2 treatments, respectively. The reduction in milk fat yield was due to decreases in both de novo fatty acid synthesis and uptake of preformed fatty acids. Milk fat content of trans-10, cis-12 CLA was 0.03, 0.09, and 0.19 g/100 g of fatty acids for the control, CLA-1, and CLA-2 treatments, respectively. The transfer efficiency of trans-10, cis-12 CLA from the 2 levels of CLA supplement into milk fat was not different between treatments and averaged 1.85%. In conclusion, trans-10, cis-12 CLA reduced milk fat synthesis in lactating dairy goats in a manner similar to that observed for lactating dairy cows and dairy sheep. Dose-response comparisons, however, suggest that the degree of reduction in milk fat synthesis is less in dairy goats compared with dairy cows and dairy sheep.

NUTRITION, METABOLISM, AND FERTILITY IN DAIRY COWS: 2. DIETARY FATTY ACIDS AND OVARIAN FUNCTION

Plasma insulin has important implications for ovarian function in dairy cows. Previous work demonstrated that plasma insulin increased with increasing dietary starch and decreasing dietary fatty acid concentrations. The objective of this experiment was to investigate hormonal and ovarian responses to dietary fatty acid content with no change in other dietary components. Thirty cows were fed a standard diet from calving until 40 d in milk (DIM) and then 6 cows were transferred to each of 5 diets containing 0, 8, 15, 23, and 30 g/kg of dry matter (DM) of calcium salts of palm fatty acids (CaPFA; Megalac) until 70 DIM. Estrus was synchronized at 60 DIM. Between 60 and 70 DIM, energy intake, milk yield, and energy balance were similar among diet groups. Plasma insulin decreased when dietary concentration of CaPFA exceeded 15 g/kg of DM (insulin: 0.46, 0.41, 0.46, 0.33, 0.28 +/- SE 0.034 ng/mL for diets containing 0 to 30 g of CaPFA/kg of DM, respectively). Maximum plasma insulin to glucagon ratio was observed with 15 g of CaPFA/kg of DM (ratios: 3.99, 4.33, 4.67, 3.45, 2.89 +/- SE 0.156 for diets containing 0 to 30 g of CaPFA/kg of DM, respectively). Plasma concentrations of growth hormone, insulin-like growth factor-I and leptin did not vary between diets. The number of small (<5 mm) ovarian follicles was negatively related to plasma insulin concentration (r = -0.328) and was stimulated by CaPFA supplementation at all rates tested compared with cows receiving zero CaPFA (small follicles preovulation: 6.7, 11.2, 11.5, 11.3, 11.9 +/- SE 1.48 for diets containing 0 to 30 g of CaPFA/kg of DM, respectively). The number of medium-sized follicles, and diameters of the ovulatory follicles and corpora lutea, were not affected by CaPFA supplementation. It is concluded that dietary total fat concentration should be below 50 g/kg of DM to avoid depressing plasma insulin concentration in cows at the start of the breeding period.