Great Lakes Dairy Sheep Symposium - the Department of Animal ...

Great Lakes Dairy Sheep Symposium - the Department of Animal ... Great Lakes Dairy Sheep Symposium - the Department of Animal ...

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Stelwagen, K., Davis S.R., Farr V.C., Prosser C.G., and Sherlock R.A. 1994. Mammary epithelial cell tight junction integrity and mammary blood flow during an extended milking interval in goats. J. Dairy Sci., 77:426-432. Swalve H.H. 2000. Theoretical basis and computational methods for different test day genetic evaluation methods. J. Dairy Sci., 83:1115-1124. Swalve, H. H. 1995. Genetic relationship between dairy lactation persistency and yield. J. Anim. Breed. Genet., 112:303-311. Thomas D. L., Berger Y. M. , and McKusick B. C. 2001. Effects of breed, management system, and nutrition on milk yield and milk composition of dairy sheep. J. Anim. Sci., 79(suppl. E):E16-E20. Togashi, K., and C. Y. Lin. 2003. Modifying the lactation curve to improve lactation milk and persistency. J. Dairy Sci. 86:1487-1493. Tonner E., Allan G.J., and Flint D.J. 2000. Hormonal control of plasmin and tissue-type plasminogen activator activity in rat milk during involution of the mammary gland. J. Endocrinol., 167:265-273. Wilde C.J., and Knight C.H. 1990. Milk yield and mammary function in goats during and after once-daily milking. J. Dairy Res., 57:441-447. Wilde C.J., Henderson A.J., Knight C.H., Blatchford D.R., Faulkner A., and Vernon R.G. 1987. Effects of long-term thrice-daily milking on mammary enzyme activity, cell population and milk yield in the goat. J. Anim. Sci., 64:533-539. Wohlt J.E., Foy Jr V.L., Kniffen D.M., Trout J.R. 1984. Milk yield by Dorset ewes as affected by sibling status, sex and age of lamb, and measurement. J.Dairy Sci., 67:802-807. Wood P.D.P. 1967. Algebraic model of the lactation curve in cattle. Nature, 216:164–165. Zamiri M. J., Qotbi A., and Izadifard J. 2001. Effect of daily oxytocin injection on milk yield and lactation length in sheep. Small Rum. Res., 40:179-185. 68

Introduction MILK FAT SYNTHESIS AND ITS REGULATION IN DAIRY SHEEP Adam L. Lock,* Liam A. Sinclair, † and Dale E. Bauman* *Cornell University, New York, USA † Harper Adams University College, Shropshire, UK Milk and dairy products were recognized as important foods as early as 4000 B.C. as evidenced by rock drawings from the Sahara. Today, the important contribution of milk and dairy products in meeting our dietary requirements for energy, high quality protein and several key minerals and vitamins are well documented. With the projected growth in world population and the increased demand for animal-derived food products as living standards improve, dairy products will undoubtedly continue to be an important dietary source of nutrients (Bauman et al., 2005a). Fat is an important component of milk for a number of reasons. Nutritionally, fat is the major energy component of milk and it accounts for many of the physical properties, manufacturing characteristics, and sensory attributes of milk and dairy products. The fat content of milk is highly variable and the environmental and physiological factors affecting milk fat have been extensively characterized. Furthermore, nutritional quality has become increasingly important in food choices because of consumer awareness of the link between diet and health; importantly, milk fat has been shown to contain a number of bioactive components that may provide benefits to human health which has led to an interest in “designing” milk fat to improve its healthfulness and functional properties. Nutrition is the predominant factor affecting milk fat and it represents a practical tool to alter the yield and fatty acid composition of milk fat. The impact of nutrition on fat content and fatty acid composition of milk in the bovine has been extensively reviewed (Palmquist et al., 1993; Lock and Shingfield, 2004; Lock and Bauman, 2004). Therefore, the purpose of this review is to provide an overview of the biology of milk fat synthesis in ruminants, discuss an innovative method for altering the fat content of sheep milk and its possible application, and consider the manipulation of the fatty acid composition of sheep milk fat. Milk Fat Synthesis Fat is the most variable component in the milk of ruminants. The concentration of fat in milk is influenced by animal and environmental factors such as breed, diet, stage of lactation, season of year, ambient temperature, and body condition (Figure 1). Milk fat consists of droplets of triglycerides that are coated with cell membrane (milk fat globular membrane; MFGM). Thus, the majority of the fat content of milk is triglycerides (~95%) with phospholipids, cholesterol, diacylglycerols, monoacylglycerols, and free fatty acids constituting the remainder (Lock and Shingfield, 2004). Ruminants are estimated to have over 400 different fatty acids comprising milk fat triglycerides, but the majority of the fatty acids have chain lengths between four and eighteen carbons. In ruminants, fatty acids in milk fat arise from two sources that contribute equally (molar basis), de novo synthesis within the mammary gland and uptake of preformed fatty acids from circulation (McGuire and Bauman, 2002). An overview of the metabolic pathways and some of the key enzymes involved in milk fat synthesis are shown in Figure 2. 69

Introduction<br />

MILK FAT SYNTHESIS AND ITS REGULATION IN DAIRY SHEEP<br />

Adam L. Lock,* Liam A. Sinclair, † and Dale E. Bauman*<br />

*Cornell University, New York, USA<br />

† Harper Adams University College, Shropshire, UK<br />

Milk and dairy products were recognized as important foods as early as 4000 B.C. as<br />

evidenced by rock drawings from <strong>the</strong> Sahara. Today, <strong>the</strong> important contribution <strong>of</strong> milk and<br />

dairy products in meeting our dietary requirements for energy, high quality protein and several<br />

key minerals and vitamins are well documented. With <strong>the</strong> projected growth in world population<br />

and <strong>the</strong> increased demand for animal-derived food products as living standards improve, dairy<br />

products will undoubtedly continue to be an important dietary source <strong>of</strong> nutrients (Bauman et al.,<br />

2005a). Fat is an important component <strong>of</strong> milk for a number <strong>of</strong> reasons. Nutritionally, fat is <strong>the</strong><br />

major energy component <strong>of</strong> milk and it accounts for many <strong>of</strong> <strong>the</strong> physical properties,<br />

manufacturing characteristics, and sensory attributes <strong>of</strong> milk and dairy products. The fat content<br />

<strong>of</strong> milk is highly variable and <strong>the</strong> environmental and physiological factors affecting milk fat have<br />

been extensively characterized. Fur<strong>the</strong>rmore, nutritional quality has become increasingly<br />

important in food choices because <strong>of</strong> consumer awareness <strong>of</strong> <strong>the</strong> link between diet and health;<br />

importantly, milk fat has been shown to contain a number <strong>of</strong> bioactive components that may<br />

provide benefits to human health which has led to an interest in “designing” milk fat to improve<br />

its healthfulness and functional properties. Nutrition is <strong>the</strong> predominant factor affecting milk fat<br />

and it represents a practical tool to alter <strong>the</strong> yield and fatty acid composition <strong>of</strong> milk fat. The<br />

impact <strong>of</strong> nutrition on fat content and fatty acid composition <strong>of</strong> milk in <strong>the</strong> bovine has been<br />

extensively reviewed (Palmquist et al., 1993; Lock and Shingfield, 2004; Lock and Bauman,<br />

2004). Therefore, <strong>the</strong> purpose <strong>of</strong> this review is to provide an overview <strong>of</strong> <strong>the</strong> biology <strong>of</strong> milk fat<br />

syn<strong>the</strong>sis in ruminants, discuss an innovative method for altering <strong>the</strong> fat content <strong>of</strong> sheep milk<br />

and its possible application, and consider <strong>the</strong> manipulation <strong>of</strong> <strong>the</strong> fatty acid composition <strong>of</strong> sheep<br />

milk fat.<br />

Milk Fat Syn<strong>the</strong>sis<br />

Fat is <strong>the</strong> most variable component in <strong>the</strong> milk <strong>of</strong> ruminants. The concentration <strong>of</strong> fat in milk<br />

is influenced by animal and environmental factors such as breed, diet, stage <strong>of</strong> lactation, season<br />

<strong>of</strong> year, ambient temperature, and body condition (Figure 1). Milk fat consists <strong>of</strong> droplets <strong>of</strong><br />

triglycerides that are coated with cell membrane (milk fat globular membrane; MFGM). Thus,<br />

<strong>the</strong> majority <strong>of</strong> <strong>the</strong> fat content <strong>of</strong> milk is triglycerides (~95%) with phospholipids, cholesterol,<br />

diacylglycerols, monoacylglycerols, and free fatty acids constituting <strong>the</strong> remainder (Lock and<br />

Shingfield, 2004). Ruminants are estimated to have over 400 different fatty acids comprising<br />

milk fat triglycerides, but <strong>the</strong> majority <strong>of</strong> <strong>the</strong> fatty acids have chain lengths between four and<br />

eighteen carbons. In ruminants, fatty acids in milk fat arise from two sources that contribute<br />

equally (molar basis), de novo syn<strong>the</strong>sis within <strong>the</strong> mammary gland and uptake <strong>of</strong> preformed<br />

fatty acids from circulation (McGuire and Bauman, 2002). An overview <strong>of</strong> <strong>the</strong> metabolic<br />

pathways and some <strong>of</strong> <strong>the</strong> key enzymes involved in milk fat syn<strong>the</strong>sis are shown in Figure 2.<br />

69

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