FEEDING DAIRY COWS: 1.FIBRE IMPORTANCE ON ANIMAL WELFARE, MILK PRODUCTION AND COMPOSITION
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1 FEEDING DAIRY COWS: 1.FIBRE IMPORTANCE ON ANIMAL WELFARE, MILK PRODUCTION AND COMPOSITION A.B. Rodríguez 1-2, P. Llorente 3, S. Andrés 1, F.J. Giráldez 1 1 Instituto de Ganadería de Montaña, CSIC-ULE 2 Pania Animal S.L. 3 INATEGA S.L. s: A.B. Rodríguez (rganabel@gmail.com) P. Llorente (pablollorente@inatega.com) S. Andrés (sonia.andres@eae.csic.es) F.J. Giráldez (j.giraldez@eae.csic.es) INTRODUCTION Forage is an essential ingredient in dairy cattle feed. Its inclusion in the diet is necessary to ensure an adequate supply of fibre, proper rumen function, animal welfare and productive performance, i.e. milk production and composition. To understand the importance of forage in the dairy cow's diet, it is necessary to know what fibre is, the role it plays in ruminant nutrition and the relationship between its effect and chopped forage size. This first article will address the following questions: What is fibre? Are there different types of fibre? What role does insoluble fibre play in the dairy cow's diet? What is rumen acidosis? Does rumen acidosis affect milk yield and composition? What percentage of fibre should be included in the dairy cow's ration?
2 WHAT IS FIBRE? ARE THERE DIFFERENT TYPES OF FIBRE? The term forage refers to fibrous or voluminous foods, i.e. feeds that are high in fibre and low in energy. Fibre is only found in feeds of plant origin, and in general terms it consists of the plant cell wall components. The intracellular content basically includes non-structural polysaccharides such as starch and fructans. The importance of the content and type of fibre included in rations renders it necessary to conduct a detailed analysis of the fibre fractions necessary to achieve a correct formulation; these are crude fibre (CF), neutral detergent fibre (NDF) and acid detergent fibre (ADF). These various fractions must be included in the calculation because there is no single laboratory method for determining the total fibrous fraction, and not everyone possesses the technical capacity to routinely determine the different types.
3 From an analytical perspective, determination of crude fibre, the most basic type of fibre, is the first method to separate cell wall components from the remaining plant cell components. However, this procedure does not achieve complete separation, and underestimates the content of structural polysaccharides, i.e. those that comprise the cell wall and are less digestible than polysaccharides found within the cell, as shown in Figure 1. Hence, a new method has been developed that optimises separation of structural cell wall polysaccharides and polyphenols from the other components (Figure 2). This new method makes it possible to determine neutral detergent fibre (NDF), also known as insoluble fibre, which includes all the cellulose, hemicellulose and lignin present in feed. Acid detergent fibre (ADF) is the fraction of NDF that consists solely of cellulose and lignin. The NDF content of forage is related to the forage dry matter intake and ADF content with the animal's capacity to digest forage. Thus, an increase in NDF content will decrease intake, while an increase in ADF will decrease digestibility of the feed and the energy available for milk production. Polysaccharides that do not form part of NDF basically include starch, pectins and fructans. These polysaccharides are highly digestible but present some differences in the fermentation
4 process that occurs in the rumen, and therefore the concept of soluble fibre was developed, which includes fructans and pectins but not starch. The fermentation of fructans and pectins produces volatile fatty acids (e.g. acetic, propionic and butyric acid), whereas the fermentation of starch not only produces volatile fatty acids but also lactic acid (Figure 3). This aspect is very important due to its effect on rumen ph, as will be discussed below. HOW DOES ONE COMPARE FORAGE QUALITY? An animal's capacity to ingest and digest forage depends on the latter's quality, and types of forage are classified according to relative feed value (RFV) (Linn & Martin, 1989) and relative forage quality (RFQ) (Moore & Undersander, 2002).
5 The RFV index classifies forage quality. The RFV is an index to classify forage quality, combining the ingestibilidad and digestibility of forage. The RFV is expressed as a percentage of the value that would have an alfalfa of reference (RFV = 1. 29), whose content of NDF and FAD would be 53 and 41%, respectively. It is calculated according to the following equation: calculating forage intake potential and digestibility according to the following equation: RFV = DMI x DDM/1.29 Where: Dry matter intake (DMI) = 120 / %NDF Digestible dry matter (DDM) = (0.779 x %ADF) According to RFV, 5 categories of alfalfa are considered: Supreme, Premium, Good, Fair and Utility (see Figure 4). The bigger the content of NDF and FAD the lower intake and digestibility and worse quality of fodder. NDF and ADF content primarily depends on the phenological stage of the plant (development of buds, leaves and lateral buds, elongation of stems and rosette, development of vegetative parts, emergence of floral organ, bloom, fruit formation, and ripening of fruits and
6 seeds) at the time of harvest (Bosworth & Stingler, 1992), although it may also be influenced by other factors such as genetic variety, season and number of cutting, environmental conditions (temperature, water regime) and harvest and drying conditions. Note that alfalfa is an important source of protein (CP) in the dairy cow's diet, and that protein content is usually inversely related to NDF content; thus, an increase in NDF is associated with a reduction CP content. Relative forage quality (RFQ) is estimated using the following equation: RFQ = DMI x TDN/ 1.23 Where: DMI: Dry matter intake as a percentage of live weight TDN: Total digestible nutrients (DM%) DMI is calculated: DMI =[(120/%NDF)+(In vitro digestibility at 48 h of NDF-45)*0.374]/(135*100) TDN=(0.98*NFC)+(0.93*CP)+(2.25*0.97*CF)+(Dvitro*NDF)-7 Where: NFC: non-fibrous carbohydrates (%DM); CP: crude protein (%DM); CF: crude fat (%DM); Dvitro: In vitro digestibility at 48 h of NDF; NDF: neutro detergent fibre (%DM). This concept is basically the same as RFV and even generates similar values; however, the method uses total digestible nutrients (TDN) rather than ADF to calculate dry matter digestibility. In addition, it also uses different prediction equations to estimate DMI. However, the values of RFV and RFQ are similar only when the value of the NDF digestibility presents an average value. If the value of alfalfa that we want to assess moves away from the mean value, the RFQ would be a more appropriate index. This is so because the RFV system assume that the digestibility is inversely proportional to the content of FAD and, this relationship, however is not fully linear, and as a result, either the relationship between RFV and % of NDF.
7 WHAT ROLE DOES INSOLUBLE FIBRE PLAY IN THE DAIRY COW'S DIET? The rumen is where fermentation takes place, and it contains various microorganisms (bacteria, protozoa and fungi) which digest the feed ingested by the animal. The feed is basically fermented in three stages: colonisation by rumen microorganisms, dissociation of cell wall polysaccharides, and hydrolysis and fermentation of intracellular components (Russell & Hespell, 1981). Fermentation of structural and non-structural carbohydrates (NDF components and starch, respectively) basically produces volatile fatty acids (e.g. acetic, propionic and butyric acid), which are absorbed by the rumen mucosa and used as precursors in the synthesis of glucose, amino acids, long-chain fatty acids, etc., for maintenance of the animal and milk production. Structural carbohydrates are less digestible by rumen microorganisms, especially if lignin content is high, and favour the development of populations of cellulolytic bacteria, which mainly produce acetic acid as a product of metabolism. With an appropriate feed particle size, the higher the NDF content, the longer the time required for mastication during intake and rumination and the greater the production of saliva (Welch & Smith, 1970). Saliva has various functions, one of which is to act as a buffer for variations in rumen fluid ph caused by the production of acids during fermentation. However, non-structural polysaccharides such as starch generally ferment very rapidly in the rumen and produce a higher amount of propionic and lactic acid. In contrast to forage, cereals have a high starch content and do not favour rumination due to their NDF lower content and smaller particle size, thus also reducing saliva production and ph buffer capacity. Therefore, as the percentage of grain in the ration is increased and the percentage of forage is decreased, the production of propionic acid increases and ph decreases. In broad terms, forage and therefore NDF content in the ration is directly proportional to the acetic acid content and ph value of rumen contents, and inversely proportional to the propionic acid content (Figure 5). High production cows consuming starch-rich rations with a very low proportion of forage, or containing forage which has been chopped too finely, may develop a condition known as rumen acidosis.
8 WHAT IS RUMEN ACIDOSIS? Acute rumen acidosis is a condition characterised by a prolonged period of time during which rumen ph is below 5, and is generally and primarily associated with a sharp rise in intake of easily fermentable carbohydrates (starch), a substrate that induces increased bacterial growth, produces high amounts of volatile fatty acids and decreases rumen fluid ph. In turn, this decrease in ph favours the growth of lactic acid-producing bacteria (Streptococcus bovis and Selenomonas ruminantium first and Lactobacilli second), while reducing the growth rate of bacteria that consume this acid (mainly Megasphera elsdenii). The result is a greater accumulation of lactic acid and volatile fatty acids, with a consequent decrease in rumen ph (Figure 6).
9 A very steep and prolonged drop in ph will cause metabolic acidosis (Owens et al., 1998). Due to the high osmotic pressure of the rumen, water from the blood capillaries is released into the gastrointestinal tract, causing diarrhoea. The clinical signs of acute acidosis include anorexia, abdominal pain, tachycardia, lethargy and even death. Intermittent periods of reduced ph will lead to subclinical acidosis. This causes the rumen papillae to atrophy, triggering ruminitis and fibrosis characteristic of acidosis. These changes in the rumen induce a reduction in feed intake, partially mitigating episodes of acidosis. However, these will recur once the animal again consumes high amounts of feed (Nocek, 1997; Enemark et al., 2002; Oetzel, 2003).
10 DOES RUMEN ACIDOSIS AFFECT MILK YIELD AND COMPOSITION? The most obvious and visible symptoms of subclinical acidosis include cyclical variations in feed consumption and a reduction in digestibility, in particular of the fibrous component of the diet. If prolonged over time, a reduction in intake can cause a reduction in milk production. Acidosis is also associated with other conditions that affect production. For example, laminitis, an inflammation of connective tissue in the hoof, has been associated with a drop in systemic ph and subsequent tissue inflammation during episodes of acidosis (Nocek, 1997). More recently, Gozho et al. (2005) associated this inflammation with toxins from bacteria that enter the rumen through wall surfaces damaged by the effect of acidosis and subsequently enter and spread through the bloodstream. Animals with laminitis spend more time lying down, resulting in a lower intake and lower milk production.
11 It is also common for cows consuming starch-rich rations to produce milk with a lower fat content, and milk composition can even be altered. This phenomenon is known as low milk fat syndrome or milk fat depression, and occurs in nutritionally healthy animals in optimal physical condition with a positive energy balance. It has a multifactorial origin, but one of the main causes is a reduction in digestion of cellulose and consequently, in the proportion of acetic acid, the precursor of short-chain fatty acids (4-16 carbon atoms). In addition, an increase in rumen propionic acid due to high starch diets promotes gluconeogenesis in the liver and insulin release, hindering an adequate supply of precursors for fat synthesis in the mammary gland. Recently, milk fat depression in dairy cows has also been linked with some of the fatty acids present in milk, intermediaries in the biohydrogenation phenomena that occur in the rumen, especially those with double bonds at the 10 carbon position in conjugated linoleic acid (trans-10 cis-12 CLA and cis-10 trans-12 CLA), which have an inhibitory effect on fat synthesis in the mammary gland (Shingfield et al., 2010).
12 WHAT PERCENTAGE OF FIBRE SHOULD BE INCLUDED IN THE DAIRY COW'S RATION? Given the differences between forage and concentrates as regards fibre content, the National Research Council (NRC, 2001) introduced recommendations for dairy cattle rations that establish fibre requirements for both types of feed (Figure 9). These recommendations are based on NDF and non-forage carbohydrate (NFC) content in the diet. Thus, the higher the forage NDF content, the lower the percentage of NDF required in the ration as a whole and the higher the percentage of carbohydrates provided by the concentrate, and vice versa. In the case of dairy cattle, acidosis is often the consequence not only of a high content of highly fermentable carbohydrates, but also of the administration of a diet with a deficient particle size, which renders the fibre ineffective. It is therefore necessary to pay attention not only to fibre content but also to particle size since this determines whether the fibre provided fulfils its function or not. This essential aspect merits detailed analysis, and will therefore be discussed in subsequent articles.
13 BIBLIOGRAPHY Bosworth, C. & Stringer, W.C. (1992). Cutting Management of alfalfa, red clover, and birdsfoot trefoil. Agronomy Facts, 7, Penn State Extension. Enemark, J.M.D, Jorgensen, R.J, Enemark, P.S. (2002). Rumen acidosis with special emphasis on diagnosis aspects of subclinical rumen acidosis: A review. Veterinarijair Zootechnika, 42, Lechartier, C., Peyraud, J.L. (2010). The effects of forage proportion and rapidly degradable dry matter from concentrate on ruminal digestion in dairy cows fed corn silage-based diets with fixed neutral detergent fiber and starch contents. Journal of Dairy Science, 93: Linn, J. & Martin, N. (1989). Minnesota Dairy Conference, p. 9.
14 Moore, J. E. & D. J. Undersander, Relative Forage Quality: An alternative to relative feed value and quality index. p En: Proc. Florida Ruminant Nutrition Symposium, University of Florida, Gainesville. Nocek, J.E. (1997). Bovine acidosis: implications on laminitis. Journal of Dairy Science, 80: NRC (2001). Nutrient Requirements of Dairy Cattle. 7th revision. Ed. National Academy Press, Washington, DC. Oetzel GR: Subacute ruminal acidosis in dairy cattle. In: The Merck Veterinary Manual,10th edition, Merial Ltd, Duluth, GA, Owens. F. N., Secrist, D. S., Hill, W. J., Gill, D. R. (1998). Acidosis in cattle: a review. Journal of Animal Science, 76: Russell, J.B, Hespell, R.B Microbial rumen fermentation. Journal of Dairy Science, 64: Shingfield, K.J., Bernard, L., Leroux, C., Chilliard, Y. (2010). Role of trans fatty acids in the nutritional regulation of mammary lipogenesis in ruminants. Animal, 4:7: Welch, B & Smith, A.M. (1970). Forage quality and rumination time in cattle. Journal of Dairy Science, 53:
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