Chapter 4 Physiology of Milk Synthesis De novo synthesis 30 Preformed W eig ht % 25 20 15 10 5 0 4:0 6:0 10:0 12:0 14:0 16:0 F a tty ac id 16:1 18:0 18:1 18:2 Milk Fat Synthesis Circulation Synthesis Translocation Secretion UFA TAG LPL FABP NEFA + Glycerol SFA (C16 - C18) TAG MFGM synthesis Glycerol Glycerol-P Glucose Glucose Acetate ßHBA Basal membrane de novo FA synthesis (C4 - C16) ACC FAS ER membrane Luminal membrane Epithelial Cell Lumen of alveolus Fat Droplet Fat Droplet migrating Golgi Body Lysosome Nucleus Mitochondria Mitochondria E.R . Blood Vessel Epithelial Cell Lumen of alveolus Fat Droplet Fat Droplet migrating Golgi Body Lysosome Nucleus Mitochondria Mitochondria E.R . Blood Vessel Epithelial Cell Lumen of alveolus Cell pinches off Fat Droplet Fat Droplet migrating Golgi Body Lysosome Nucleus Mitochondria Mitochondria E.R . Blood Vessel Epithelial Cell Lumen of alveolus Cell pinches off Fat Droplet Fat Droplet migrating Golgi Body Lysosome Nucleus Mitochondria Mitochondria E.R . Blood Vessel Major Fatty Acids (wt%) of Bovine Milk Saturated Fatty Acids 4:0 6:0 8:0 10:0 12:0 14:0 16:0 18:0 Butyric acid Hexanoic acid Octanoic acid Decanoic acid Lauric acid Myristic acid Palmitic acid Stearic acid 4.5 2.3 1.3 2.7 3.0 10.6 28.2 12.6 Major Fatty Acids (wt%) of Bovine Milk Monounsaturated Fatty Acids 16:1 18:1 Palmitoleic acid Oleic acid 1.6 21.4 Major Fatty Acids (wt%) of Bovine Milk Polyunsaturated Fatty Acids 18:2 Linoleic acid 18:3 Linolenic acid 2.9 0.3 Effect of hydrogenation by rumen microorganisms on dietary lipids Wt% of Fatty Acids Fatty acid 16:0 18:0 18:2 18:3 Diet Abomasal digesta 26 3 17 31 29 45 4 6 Changes in rumen conditions that lead to milk fat depression linoleic acid (cis-9, cis-12 C18:2 ) conjugated linoleic acid (cis-9, transtrans-11 C18:2) vaccenic acid (trans-11 C18:1) stearic acid (C18:0 ) conjugated linoleic acid trans-10, ciscis-12 C18:2 trans-10 C18:1 MFD induced by diet MFD induced by Ca-salts of CLA 5 5 2 y = 1.19x 2 - 3.37x + 4.47 y = 1.04x - 2.76x + 3.93 R2 = 0.63 3 3 2 2 1 1 0.0 0.5 1.0 1.5 trans -10, cis -12 CLA, mg/g milk fat R2 = 0.66 4 Milk fat% Milk fat, % 4 2.0 0.0 0.5 1.0 1.5 trans -10, cis -12 CLA, mg/g milk fat 2.0 Milk Fat Concentration Most variable component of milk Increased during periods of energy deficit Decreased when fed diets containing unsaturated fatty acids (most grains) and low effective fiber (change rumen conditions) – Milk fat depression (<2.8% fat) Fatty acid changes also occur Milk Proteins Caseins (80%) beta-lactoglobulin (10%) Non-casein (Whey) (20%) alpha-lactalbumin (4%) Serum albumin, Immunoglobulins (5%) Milk Protein Concentration Not too variable Changes related to dietary energy intake, not dietary protein – little effect of dietary fat on milk protein concentration unless perturb normal rumen function – fermentable carbohydrate intake is the key! Maximize VFA and microbial protein production 1.5 60 50 40 30 20 10 0 1.0 0.5 0 0 10 20 30 40 hours since last milking Secretion Rate (kg/hr) (____) Udder Pressure (mm Hg) (----) Udder Pressure & Secretion Rate Secretion Rate • Influence of udder pressure • Milking intervals – 12 hour vs. other – 2X vs. 3X – young vs. old cows Yield and Milking Frequency (Erdman & Varner, 1995 JDS) Item Milk yield Fat% Protein % # studies 40 18 4 2X 42.9 3.65 3.10 3X 50.6 3.51 3.05 4 3 3 2X 54.6 4.71 3.49 4X 64.5 4.53 3.44 # studies Milk Let Down • Refractory period to oxytocin? • Adrenaline interferes Milk Ejection brain nerves P.P. Stimulus Milk Ejection nerves P.P. Milk Ejection P.P. Milk Ejection Abdominal aorta Oxytocin Jugular vein heart Milk Ejection Oxytocin Milk Ejection Oxytocin Milk Ejection Milk Ejection Milk Ejection Contraction of an Alveolus arteriole venuole Lumen Myoepithelial Cell Oxytocin Duct Contraction of an Alveolus Oxytocin Myoepithelial Cell contracts Contraction of an Alveolus Releases milk into duct system Contraction of an Alveolus Myoepithelial cell relaxes Contraction of an Alveolus Contraction of an Alveolus 5 4.5 4 3.5 3 2.5 2 1.5 1 0.5 0 Milk Fat % as Milking Progresses foremilk early middle late leftover Milk Fat % & Exercise 3.8 3.7 3.6 3.5 3.4 3.3 3.2 limted moderate considerable Biosynthesis of Milk components Precursors from blood One liter of milk requires 500 L of blood Fat biosynthesis Protein biosynthesis Lactose biosynthesis Milk Fat biosynthesis Requires fatty acids and glycerol Two sources of fatty acids: 1- blood lipids (dietary lipids & adipose tissue) 2- De novo synthesis (within the MG) Major Fatty Acids (wt%) of Bovine Milk Polyunsaturated Fatty Acids 18:2 Linoleic acid 18:3 Linolenic acid 2.9 0.3