Amino acids utilization comprises two phases, cellular uptake and intracellular metabolism. The paper reviews reports on amino acid uptake in different species and at different stages of lactation and presents new data on changes at lactogenesis. The relative importance of changes in factors which determine uptake, mammary blood flow, arterial concentration, and mammary extraction, are discussed. In addition to direct incorporation into milk protein, some absorbed amino acids are an energy source and carbon precursors of other amino acids synthesized in mammary cells. Branched-chain amino acids and ornithine act as nitrogen precursors in these syntheses. Ornithine, which is derived partly from absorbed arginine, is also a precursor of spermidine, a postulated intracellular mediator of hormone action. The question of whether milk protein yield is limited by supply to the mammary gland of certain amino acids is discussed. Much evidence indicates the importance of certain essential amino acids that are transferred stoichiometrically from blood to milk. Results of experiments involving administration of exogenous growth hormone to goats are presented. Increases in milk yield and protein yield are associated with increased rates of mammary blood flow, emphasizing the role of the latter in determining uptake.
1. The following techniques, which have been applied successfully to goats, were used to study mammary metabolism in lactating sows:(I) measurements of mammary arteriovenous (A-V) differences in milk precursors in the conscious undisturbed animal (five sows); (2) continuous intravenous infusion of [U-14C]glucose with concomitant arterial and mammary venous blood sampling for measurement of mammary blood flow and specific radioactivity of glucose and CO, (one sow); (3) perfusion of the isolated gland in vitro (eight glands from four sows), with the inclusion of [U-14C]glucose (two glands) and [U-14C]acetate(two glands) in the substrate mixture.2. Sow mammary tissue was similar to that goats in its milk yield, blood flow, and glucose uptake per unit weight of tissue. As in goats, mammary uptake of glucose was many times that of the rest of the body and the total mammary tissue was utilizing about half of the total glucose entering the circulation. Glucose was a major source of milk lactose and glycerol and of mammary CO2.3. Of the plasma lipid components, only the triglyceride fraction was consistently and significantly removed by the gland. In contrast to the results obtained for the goat, both [U-14C[acetate and [U-14C]glucose carbon were used for milk fatty acid synthesis, and although the pattern of labelling of fatty acid from each precursor was similar, the formation of fatty acids from glucose was at least five times greater than that from acetate. Quantitative evaluation of the contribution of these precursors was not possible, but the RQ (1.09–1.63) suggest that in some instances it may have been considerable.4. The substantial A-V differences of most plasma essential amino acids suggest that these are the sole precursors of the corresponding residues in the mammary synthesized protein. The low A-V differences for several non-essential amino acids suggest that these are synthe- sized in the gland; this suggestion is supported by the incorporation of glucose carbon into non-essential amino acid residues of casein observed in one experiment. However, in contrast to results with the goat, mammary absorption of serine was consistently large.
SUMMARY1. The time course of changes in specific activities of citrate, lactose and fatty acids in milk during frequent milking, following the ixv. administration of labelled glucose, acetate and chylomicrons in goats has been studied. Peak specific activities of lactose and citrate in milk were reached at 2-3 hr, while peak specific activities of fatty acids were reached at 5-7 hr.2. Following short I.A. infusions of 24Na, 3"Cl and 42K, peak specific activities in milk were reached in 1 hr or less.3. The mammary epithelium of lactating goats was found to be virtually impermeable to labelled citrate in both directions. 4. Labelled citrate had an apparent volume of distribution in lactating guinea-pigs mammary slices in vitro similar to that of extracellular space markers.5. Treatment of goats with large doses of oxytocin markedly increased the permeability of the secretary epithelium to labelled citrate.6. In the goat mammary gland, citrate, protein and calcium failed to enter milk which had been diluted with isosmotic lactose by intraductal injection, whereas Na, K and Cl did enter, thus tending to restore the concentrations of these ions to normal.7. It is suggested that citrate, which is formed within the secretary cell, enters milk not by passage across the apical cell membrane but, in common with lactose and milk protein, by exocytosis of Golgi vesicles. It appears that citrate is held at high concentrations in milk by virtue of the impermeability of the mammary epithelium to the forms in which it occurs in milk.
1. Arteriovenous differences of plasma free amino acids across the lactating mammary glands of six goats have been measured. 2. In four experiments, measurements of blood flow, amino acid arteriovenous differences, milk yield and milk nitrogen showed that the uptake of nitrogen in the form of amino acids was sufficient to provide all the nitrogen of the milk proteins synthesized in the mammary gland. 3. In the same four experiments the uptake from the plasma and output into the milk of individual amino acids per unit time were compared. The uptakes of essential amino acids and glutamic acid were approximately equal to the corresponding output figures. The uptake of serine was consistently less than the output, and the uptake of other non-essential amino acids was very variable, in some experiments being approximately equal to the output figures and in others being considerably less. 4. As in cows, there was an uptake of ornithine in all experiments, though ornithine is absent from milk. In goats, though not in cows, the uptake of arginine was consistently greatly in excess of the requirement for arginine residues in milk protein. 5. The possible significance of the uptakes of arginine and ornithine for the synthesis of serine and other non-essential amino acids in the mammary gland is discussed. 6. The importance of clamping the external pudic vein, when sampling mammary venous blood from the caudal superficial epigastric vein, is indicated.
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