Recent advances in ruminal lipid metabolism have focused primarily on manipulation of physicochemical events in the rumen aimed at two practical outcomes: 1) control of antimicrobial effects of fatty acids so that additional fat can be fed to ruminants without disruption of ruminal fermentation and digestion and 2) regulation of microbial biohydrogenation to alter the absorption of selected fatty acids that might enhance performance or reduce saturation of meat and milk. Properties of lipids that determine their antimicrobial effects in the rumen include type of functional group, degree of unsaturation, formation of carboxylate salts, and physical association of lipids with surfaces of feed particles and microbes. The mechanism of how lipids interfere with ruminal fermentation is a complex model involving partitioning of lipid into the microbial cell membrane, potency of the lipid to disrupt membrane and cellular function, physical attachment of microbial cells to plant surfaces, and expression and activity of microbial hydrolytic enzymes. Lipolytic and hydrogenation rates vary with forage quality (stage of maturity and N content), surface area of feed particles in the rumen, and structural modifications of the lipid molecule that inhibit attack by bacterial isomerases.
Recent advances in chromatographic identification of CLA isomers, combined with interest in their possible properties in promoting human health (e.g., cancer prevention, decreased atherosclerosis, improved immune response) and animal performance (e.g., body composition, regulation of milk fat synthesis, milk production), has renewed interest in biohydrogenation and its regulation in the rumen. Conventional pathways of biohydrogenation traditionally ignored minor fatty acid intermediates, which led to the persistence of oversimplified pathways over the decades. Recent work is now being directed toward accounting for all possible trans-18:1 and CLA products formed, including the discovery of novel bioactive intermediates. Modern microbial genetics and molecular phylogenetic techniques for identifying and classifying microorganisms by their small-subunit rRNA gene sequences have advanced knowledge of the role and contribution of specific microbial species in the process of biohydrogenation. With new insights into the pathways of biohydrogenation now available, several attempts have been made at modeling the pathway to predict ruminal flows of unsaturated fatty acids and biohydrogenation intermediates across a range of ruminal conditions. After a brief historical account of major past accomplishments documenting biohydrogenation, this review summarizes recent advances in 4 major areas of biohydrogenation: the microorganisms involved, identification of intermediates, the biochemistry of key enzymes, and the development and testing of mathematical models to predict biohydrogenation outcomes.
Recent research has demonstrated the effectiveness of added fat in diets to maintain milk production and fat percent. Much of the earlier work which indicated that fat affects digestion negatively may not be applicable because of great differences in the nature of diets and fats fed and especially in total feed intake. Nevertheless, much remains to be learned about interactions of fat, fiber, calcium, and rumen microorganisms if feeding of fat is to be maximized. The uniquely high acidity in the duodenum combined with detergent action of bile acids, lysolecithin, and fatty acids causes saturated fatty acids to be more digestible in ruminants than in nonruminants. Large quantities of added dietary fat increase concentrations in plasma of very low density lipoprotein triglyceride which increases their uptake by the mammary gland with inhibition of short chain fatty acid synthesis and consequent changes in milk fatty acid composition. In some cases, secretion of milk fat is increased. Current research and practice demonstrate that 3 to 5% fat may be added to diets for lactation to increase energy intake of high-producing cows and/or to reduce starch feeding, thereby increasing the ratio of forage to concentrate to prevent depression of milk fat.
A number of major scientific advances have been realized in the last 25 yr in determining the opportunities and limitations of altering milk composition through nutritional manipulation. Because of the greater sensitivity of milk fat to dietary manipulation than either protein or lactose, nutritional control of milk fat content and fatty acid composition received a great deal of attention. New information emerged linking ruminal production of trans fatty acid isomers with milk fat depression. As a result, research on fatty acid biohydrogenation intensified yielding new insight on the origin of specific trans fatty acid isomers originating from ruminal biohydrogenation and how these isomers were modified by the action of mammary enzymes. The discovery of conjugated linoleic acid (CLA) as a potent anticarcinogen also led to extensive work on enhancing its concentration in milk through nutritional manipulation and discovering the physiological effects of specific CLA isomers. New protected fats were developed in recent years that were designed to resist biohydrogenation and enhance the concentration of unsaturated fatty acids in milk. The nutritional factors receiving the most attention during the last 25 yr for their influence on milk protein content were forage-to-concentrate ratio, the amount and source of dietary protein, and the amount and source of dietary fat. New insights were tested on modes of action whereby fat supplements caused a decline in protein concentration. Changes in milk lactose concentration occur only in extreme and unusual feeding situations, but the basic biology of lactose synthesis and regulation are still being explored using modern molecular techniques. This paper highlights the major advances in controlling milk composition by dietary manipulation and how it influences the entire animal system from practical feeding studies to basic cellular work on mammary tissue metabolism.
Holstein (n = 19) and Jersey (n = 18) cows were used to study effects of two feeding systems on fatty acid composition of milk. Confinement cows were fed a total mixed ration with corn silage and alfalfa silage and pastured cows grazed a crabgrass (90%) and clover (10%) pasture and were allowed 5.5 kg of grain per head daily. Two milk samples were collected from each cow at morning and afternoon milkings 1 d each week for four consecutive weeks in June and July 1998. One set of milk samples was analyzed to determine fatty acid composition, and the second set was used for crude protein and total fat analyses. Data were analyzed by the general linear models procedure of SAS, using a split-plot model with breed, treatment, and breed x treatment as main effects and time of sampling and week as subplot effects along with appropriate interactions. Milk from pastured cows was higher than milk from confinement cows for the cis-9, trans-11 octadecadienoic acid isomer of conjugated linoleic acid (CLA). Also, milk from Holsteins was higher than milk from Jerseys for C16:1, C18:1, and CLA and lower than Jerseys for C6:0, C8:0, C10:0, C12:0, and C14:0. Several treatment x week interactions existed, but main effects were still important; for example, proportions of CLA in milk of grazed cows were relatively constant across weeks (0.66, 0.64, 0.64, and 0.69% +/- 0.02%, respectively), but the CLA in milk of confinement cows increased in wk 4 (0.35, 0.31, 0.31, and 0.48% +/- 0.02% for wk 1 to 4, respectively). There are potentially important differences in fatty acid composition of milk from cows consuming a warm season pasture species compared with milk from cows consuming a total mixed ration, as well as differences between Holstein and Jersey breeds.
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