Fat in lactation rations: review.

Dietary requirements and functions of α-linolenic acid in animals

Specialized techniques, previously used in surgically prepared goats, which simultaneously measure udder metabolism (arteriovenous difference of milk precursors x udder blood flow) and the whole body turnover of the milk precursors, have been successfully transferred to dairy cows. Methods of obtaining representative samples of arterial and mammary venous blood and of measuring udder blood flow are described.The rates of entry into the circulation, as determined by isotope dilution, of glucose, acetate and plasma free fatty acids were 3·3–4·0, 1·7–2·1 and 0·5 kg/day respectively. Acetate and glucose contributed 32–50 and 4–11% respectively of the total CO2output by the animal.Measurement of the uptake of precursors of milk constituents and their transfer into milk showed that there were substantial arteriovenous differences of glucose, acetate, triglyceride and β-hydroxybutyrate which were not significantly different between breeds or related to milk yield. Isotopic and balance data confirm that glucose is the main precursor of lactose and that the oxidation and transfer of glucose into lactose accounted for 69–98% of the glucose entry rate. As in the goat, plasma triglycerides and blood acetate accounted for 35–80% and 25–50% of the milk triglycerides respectively. Propionate was extracted from plasma but the uptake was only about 8% of the value for acetate.There was no net arteriovenous difference of phospholipids, cholesterol esters or free fatty acids, but the fall in specific radioactivity of free fatty acids across the mammary gland indicated there was an exchange of free fatty acids between plasma and mammary tissue. In agreement with previous findings, acetate contributed to all the milk fatty acids up to a chain length of C14 and part of the C16 fatty acid. Plasma triglycerides contributed to the remainder of the C16 fatty acid and all the milk fatty acids with a chain length of C18 or higher.In contrast to the lactating goat, cow plasma contained very few chylomicrons. The majority of the triglycerides taken up by the udder were derived from the lowdensity lipoprotein fraction.The essential amino acids were extracted from blood in amounts sufficient to account for the essential amino acids secreted into milk protein. Although the plasma level of methionine was low, 52–72% of the material reaching the mammary gland was taken up. The uptake of arginine was far in excess of the requirement for milk protein synthesis.

The metabolism of glucose, acetate, lipids and amino acids in lactating dairy cows

Lipid metabolism in the adipose tissue of ruminant animals

Lipid metabolism in the lactating mammary gland.

1. Measurements were made of milk yield, mammary blood flow and arteriovenous differences of each plasma lipid fraction, and their specific radioactivities, during the infusion of [U-(14)C]stearate, [U-(14)C]oleate, [U-(14)C]palmitate and [1-(14)C]acetate into fed lactating goats. 2. Entry rates of fatty acids into the circulation were 4.2mg./min./kg. body wt. for acetate, and 0.18, 0.28 and 0.42mg./min./kg. for stearate, oleate and palmitate respectively. Acetate accounted for 23% of the total carbon dioxide produced by the whole animal, and contributed to the oxidative metabolism of the mammary gland to about the same extent. Corresponding values for each of the long-chain acids were less than 1%. 3. There were no significant arteriovenous differences of phospholipids, sterols or sterol esters, and their fatty acid composition showed no net changes during passage through the mammary gland. 4. There were large arteriovenous differences of plasma triglycerides, and their fatty acid composition showed marked changes across the gland. The proportions of palmitate and stearate fell, and that of oleate increased. 5. Arteriovenous differences of plasma free fatty acids (FFA) were small and variable, but a large fall in the specific radioactivity of each of the long-chain acids examined indicated substantial uptake of plasma FFA, accompanied by roughly equivalent FFA release from mammary tissue. The uptake of FFA was confirmed by the extensive transfer of radioactivity into milk. The FFA of milk were similar in composition and radioactivity to the milk triglyceride fatty acids, and quite unlike plasma FFA. 6. The formation of large amounts of oleic acid (18-21 mg./min.) from stearic acid was demonstrated. 7. During the terminal stages of the [(14)C]acetate infusion, milk triglyceride fatty acids of chain length C(4)-C(14) showed specific radioactivities that were 75-90% of that of blood acetate, and that of palmitate was roughly one-quarter of this value. Oleate and stearate were unlabelled. 8. The results confirmed that milk fatty acids of chain length C(4)-C(14) arise largely from blood acetate, and palmitate is derived partly from acetate and partly from plasma triglyceride, the latter fraction being almost the sole precursor of oleate and stearate.

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The oxidation and utilization of palmitate, stearate, oleate and acetate by the mammary gland of the fed goat in relation to their overall metabolism, and the role of plasma phospholipids and neutral lipids in milk-fat synthesis.

1. Mammary glands of lactating goats were perfused for 12.5-15hr. with heparinized whole blood and infused with a substrate mixture of glucose, acetate and amino acids (and sometimes chylomicra) containing either [1-(14)C]acetate, d(-)-beta-hydroxy[1-(14)C]butyrate or [U-(14)C]stearate. 2. There was a substantial net uptake of acetate by the glands and transfer of radioactivity into milk fat. Acetate was extensively utilized for the synthesis of milk fatty acids of chain length up to C(14) and to a smaller extent for the synthesis of palmitate. 3. There was a small and variable net uptake of stearate and beta-hydroxybutyrate and negligible oxidation of these substrates. However, tissue uptake was demonstrated by a substantial fall in specific radioactivity across the glands and an extensive transfer of radioactivity into milk fatty acids. 4. With beta-hydroxybutyrate the labelling of milk fat was very similar to that with acetate, but the distribution of radioactivity suggested a cleavage into C(2) fragments of about 40%. 5. Labelled stearate gave rise to highly labelled stearate and oleate in the milk fat. Small amounts of radioactivity were detected in stearate of plasma triglycerides and oleate of plasma free fatty acids. 6. In experiments where there was a decline in milk-fat secretion late in perfusion, the milk fatty acids showed a marked decline in the proportion of stearate and oleate and a rise in the proportion of myristate and palmitate. This did not occur in experiments where milk-fat secretion was maintained at a higher level. 7. The present results confirm that there is a large pool of long-chain fatty acids in mammary tissue that can act as an endogenous source of these substrates.

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E. F. Annison

Papers

The oxidation and utilization of palmitate, stearate, oleate and acetate by the mammary gland of the fed goat in relation to their overall metabolism, and the role of plasma phospholipids and neutral lipids in milk-fat synthesis.

The incorporation of acetate, stearate and D(-)-β-hydroxybutyrate into milk fat by the isolated perfused mammary gland of the goat.