Udder health and milk composition, with special reference to beef cows

A literature review
Miguel Velazquez

____________________________________________________________________________ SLU Specialarbete 11 Institutionen fr husdjurens milj och hlsa Swedish University of Agricultural SciencesSkara 2000 Faculty of Veterinary Medicine ISSN 1402-3342 Department of Animal Environment and Health ISBN 91-576-6004-2 ___________________________________________________________________________

FOREWORD

This literature review was written by Miguel Velazquez, a recently qualified veterinarian from the University of Merida in Mexico. He was invited to participate in two projects within FOOD21, and stayed in Sweden from April to November 2000. FOOD21 is a large research programme mainly working in the area of a sustainable environment and agriculture. The programme has been funded by MISTRA during 1997-2000 and is expected to be funded for a second period during 2001-2004. As a part of the synthesis work, a group called Tema Ko-kalv (theme: Cow-calf) has been organising meetings, seminars, research projects and collaboration with developing countries, as for example Mexico and Colombia. The major interest of this group is to increase the knowledge about systems where dairy cows and calves are kept together, and to study the effects of these systems on behaviour, health and production. One of the projects Miguel Velazquez was working with during his stay in Sweden was on the nursing behaviour, milk production, udder health and milk composition of beef heifers during their first lactation. There was a need in the project to have a good literature review on the udder health and milk composition related to beef cattle. This review has been checked by us and approved for publication after some revision. It is published as a joint publication between the Department of Animal Environment and Health and FOOD21. It is our hope that this review will be of value both for students, and for other researchers going into this field. The visit of Miguel Velazquez was funded by the International Foundation for Science (IFS).

Skara 2000-11-27

Lena Lidfors

Charlotte Berg

CONTENTS

SUMMARY RESUMEN (ESPAOL) INTRODUCTION THE EFFECT OF MASTITIS ON PRODUCTION IN BEEF CATTLE FACTORS INVOLVED IN INFECTION AND MASTITIS COMMON TYPES OF INFECTIOUS MASTITIS Staphylococcal mastitis Streptococcal mastitis Coliform mastitis THE EPIDEMIOLOGY OF MASTITIS IN BEEF CATTLE SOMATIC CELL COUNT FACTORS AFFECTING SOMATIC CELL COUNT Infection status Season Age Stage of lactation Diurnal variation Day to day variation Stress Technical aspects Effects of quarter Milk fraction BOVINE MASTITIS BACTERIOLOGY MILK CONSTITUENTS NUTRITIONAL FACTORS AFFECTING MILK CONSTITUENTS Fat content Protein content Lactose content NON-NUTRITIONAL FACTORS AFFECTING MILK CONSTITUENTS Breed Stage of lactation

7 8 10 11 12 16 16 17 17 18 20 20 20 21 22 22 23 23 24 24 25 25 26 27 28 29 30 30 31 31 31

Mastitis and somatic cell count Milk fraction Other factors affecting milk constituents ARE TEAT TRAITS IMPORTANT IN UDDER HEALTH AND MILK COMPOSITION ASSESSMENTS? FACTORS AFFECTING TEAT LENGTH AND ITS IMPORTANCE ON MILK YIELD AND UDDER HEALTH TEAT INJURY AND TEAT PAPILLOMATOSIS MILK UREA CONCENTRATION NUTRITIONAL FACTORS AFFECTING MILK UREA CONCENTRATION Effect of protein Importance of balance between protein and energy Effect of rumen undegradable protein (RUP) NON-NUTRITIONAL FACTORS AFFECTING MILK UREA CONCENTRATION Diurnal variation Season Stage of lactation Age Mastitis and somatic cell count (SCC) Body weight Milk yield Technical aspects Other factors affecting milk urea concentration EFFECTS OF NURSING ON UDDER HEALTH STATUS RESTRICTED SUCKLING VERSUS ARTIFICIAL REARING CONCLUSION CONCLUSION (ESPAOL) ACKNOWLEDGEMENTS REFERENCES

33 33 33

35 35 35 37 38 38 38 39 40 40 41 41 42 42 43 43 43 44 45 45 49 49 50 52

SUMMARY

Mastitis is a significant disease in beef cattle production. However the prevalence of major pathogens in beef cows is lower compared to dairy cows, S. aureus being the main major pathogen isolated. The reason for the lower prevalence may be the absence of opportunity for infection at machine milking time, which is considered an important factor for mastitis in dairy cows. Immunoglobulins, cell-mediated immunity and anatomic teat traits play an important role in the protection against mastitis. The most important factor affecting somatic cell count (SCC) is the infection status of the quarter, and the threshold for dividing quarters into uninfected and infected (with a major pathogen) is usually set at 200 000 cells/ml or 300 000 cells/ml. However several samplings, at least five during a lactation, must be done to achieve reliable results. Although several factors have a significant effect on SCC, some of them could be explained by the dilution effect, related to the milk yield of the cow. Interpretation of bacteriology and SCC in milk together is not straighforward, and possible reasons for unexpected results must be taken into account. Milk fat percentage is the most variable component in milk. Despite of some contradictory works, most of the literature have reported that in general high amounts of energy in the feed result in milk fat depression, and that the feed protein level has little or no effect on the milk fat content. Increased dietary energy will increase milk protein percentage. Increased protein levels in the diet generally have no effect on milk protein content. However fat supplements may decrease milk protein percentage. Lactose is the most constant component in milk. Milk constituents are also affected by non-nutritional factors and mastitis is the most important such factor. Some nonnutritional factors that have influence on milk composition could be explained by dilution effects as well. The most important nutritional factor affecting milk urea concentration is the balance between energy and protein. Milk urea concentration must be measured only in samples from healthy quarters and preferably in fresh samples or after a short period of conservation. Also, due to the diurnal variation, time sampling and feeding time must be considered. Teat traits have an important role in the development of intramammary infections, therefore this must be taken into consideration in mastitis risk factor assessment. Suckling has a beneficial effect on the incidence of mastitis. This could be due to the mechanical effect of suckling, the better emptying of the udder and to the cleaning effect of the calfs saliva.

RESUMEN

Mastitis es una enfermedad significativa en la produccin de ganado de carne. Sin embargo la prevalencia de patgenos mayores en vacas de carne es menor comparado con vacas de leche, siendo S. aureus el principal patgeno mayor aislado. La razn para esta menor prevalencia puede ser la falta de oportunidad de infeccin al momento de la ordea, lo cual es considerado un factor importante en mastitis en vacas lecheras. Inmunoglobulinas, inmunidad mediada por celulas y caractersticas anatmicas de la teta juegan un papel muy importante en la proteccin contra mastitis. El factor ms importante afectando el conteo de celulas somticas es el estado de infeccin del cuarto, y el lmite para dividir cuartos en infectados y no infectados es usualmente fijado en 200 000 cels/ml o 300 000 cels/ml. Sin embargo varios muestreos, por lo menos cinco durante la lactation, deben ser hechos para conseguir reultados fiables. Aunque varios factores tienen un efecto significante sobre el conteo de celulas somticas, algunos de ellos podran ser explicados por el efecto de dilucin, relacionado con la produccin de leche de la vaca. La interpretacin de examenes bacteriolgicos y conteo de celulas somticas en leche, cuando se utilizan juntos, no es sencillo, y posibles razones para resultados inesperados deben ser tomados en cuenta. El porcentage de grasa en la leche es el componente ms variable en la leche. A pesar de algunos trabajos contradictorios, la mayora de la literatura ha reportado que en general altas cantidades de energa en el alimento resulta en una disminucin de la grasa en leche, y que los niveles de proteina en el alimento tienen poco o ningn efecto sobre el contenido de grasa en leche. Energa diettica incrementada incrementar el porcentage de proteina en leche. Niveles de proteina incrementados en la dieta generalmente no tienen efecto sobre el contenido de proteina en leche. Sin embargo los suplementos grasos pueden disminuir el porcentage de proteina en leche. Lactosa es el componente ms constante en la leche. Los constituyentes de la leche tambin son afectados por factores no nutricionales y mastitis es el ms importante de ellos. Algunos factores no nutricionales que tienen influencia sobre la composicin de la leche podran ser explicados tambin por el efecto de dilucin. El factor nutricional ms importante afectando la concentracin de urea es el balance entre energa y proteina. La concentracin de urea en leche debe ser medida solo en muestras provenientes de cuartos sanos y de preferencia en muestras frescas o despus de un perodo corto de conservacin. Tambin, debido a las variaciones diurnas, tiempo de muestreo y tiempo de alimentacin deben ser considerados.

Las caractersticas de las tetas tienen un papel importante en el desarrollo de infecciones intramamarias, por lo tanto esto debe ser tomado en consideracin en evaluaciones de factores de riesgo de mastitis. El amamantamiento tiene un efecto benfico sobre la incidencia de mastitis. Esto puede ser debido a el efecto mecnico del amamantamiento, el mejor vaciado de la ubre y al efecto de limpieza proporcionado por la saliva del becerro.

INTRODUCTION

The udder is one of the most important physiological and conformational characteristics of the cow (White and Vinson, 1975) due to its importance for milk production. It is often believed that milk production is only important in dairy cattle. However milk production in beef cows is the greatest single factor influencing preweaning weight gains (Beal, et al., 1990). High correlations between milk yield and calf weight have been found by several researchers (Neville, 1962; Totusek et al., 1973; Stobbs and Brett, 1976; Boggs et al., 1980; Doornbos et al., 1981; Beal et al., 1990). Studies done by Neville (1962) and Jeffery et al. (1971) determined that milk yield accounted for more than 60% of variation in calf preweaning weight gain. Rutledge et al. (1971) suggested that approximately 60% of the variance in weaning weight could be attributed to the direct influence of the dams milk yield. Jeffery and Berg (1971) and Boggs et al. (1980) reported that each additional kg of milk per day increased calf preweaning weights by 7 to 14 kg. Therefore factors affecting milk production in beef cattle are certainly important. However, research about factors that can influence the quality and quantity of bovine milk has hitherto mainly been carried out in dairy cows. The effects of feeding and separation in cow-calf pairs on milk yield, cow body weight, calf growth, udder health, milk composition and behaviour have been studied in beef cows in a experimental studies carried out in Sweden recently. The investigation was done with 7 Hereford, 3 Angus and 4 Angus x Hereford primiparous cows with an age of 23-26 months and their calves. The study consisted of two phases, with the first one in the indoor season (5 days after calving) and the second one in the grazing season (3 months after calving). In the first phase cows were fed with silage ad libitum and 1kg of concentrate which was a mixture of barley (35%) and oats (65%). In the second phase 6 kg of concentrate were given. Milk yield was estimated by the calf weighing method using a scale. The cows body weight was recorded by scale (phase 1) and by tape measure (phase 2). Three milk samples were taken per day at a number of occasions, two (in the morning and in the afternoon) for analysis of fat, protein, lactose and somatic cell count and one for bacteriology (only in the morning). The suckling behaviour was recorded using a portable video camera. For each cow separate teat traits were recorded in both phases. Analysis of the results is currently being undertaken. More information is needed in order to interpret, in the best way possible, the results of this investigation and also for future investigations on the subject. The aim of this review is to present some basic knowledge about factors that influence udder health and milk composition in beef cows, mainly by comparing data from dairy and beef cattle reported in the literature.

10

THE EFFECT OF MASTITIS ON PRODUCTION IN BEEF CATTLE

The relationship between cow-calf performance and udder health status has been investigated in beef cattle. Watts et al. (1986) found that intramammary infections with pathogenic organisms were associated with increased somatic cell count (SCC) levels and decreased weaning weights, reporting that calves from cows with one, two, three and four quarters infected weighed 10.4, 22.7, 27.2 and 27.2 kg less (P<0.01), respectively, than calves from uninfected animals. These authors concluded that as the number of infected quarters per cow increased, regardless of pathogen, milk production of the cow was apparently reduced which resulted in lower calf weaning weights. Haggard et al. (1983) reported that mean 205-d weights of calves with uninfected dams were 13.0 kg. greater than those of calves with S. aureus-infected dams. Newman et al. (1991) reported that beef cows with intramammary infections weaned calves that were 9.6 kg lighter (P<0.05) than calves from contemporary non-infected cows. Nickerson et al. (2000) observed that dams with coagulase-negative staphylococci mastitis weaned calves that weighed 15 lb less than uninfected controls, and dams with S. aureus mastitis weaned calves that weighed 23 lb less, concluding that producers may be losing over 5% of sales values due to this disease. However, Simpson et al. (1994) reported that cows with intramammary infections had higher SCC, but that did not affect milk production or weaning weight. Simpson et al. (1995) found similar results in Simmental cows, reporting that higher SCC were associated with lower milk production in cows, but cows infected with intramammary pathogens weaned calves with similar weight to cows negative for intramammary pathogens. These researchers suggested that maybe infections in their study were not as severe or persistent as in other studies. Hence, factors that limit the dams milk production can be expected to reduce calf weaning weights. Mastitis is a disease that reduces milk production in cow (Blosser, 1979).

11

FACTORS INVOLVED IN INFECTION AND MASTITIS

Mastitis (from the Greek word mastos meaning breast and the suffix itis meaning inflammation) classically is defined as inflammation of the mammary gland (Kehrli and Shuster, 1994). Similarly, inflammation is defined simply as a reaction to injury. Hence, injury of any type to mammary tissue may be expected to induce an inflammatory response or mastitis (Jain, 1979). However, the udder disease usually arises as a result of microbial infection. The signs of mastitis vary according to factors in the host and the invading pathogen (Leigh, 1999). The characteristic features of an inflammatory response are swelling, heat, redness, pain and disturbed function. Peracute mastitis exhibits all the signs of inflammation along with systemic signs of fever, depression, shivering, loss of appetite, rapid weight loss and in some cases development of bacteraemia, septicaemia and death of the animal. Acute mastitis also is characterized by all gross signs of inflammation and some signs of systemic disturbance such as fever and mild depression. In Subacute mastitis, the cardinal signs of mastitis are less pronounced and there are no systemic signs. The existence of inflammation in the absence of gross signs is referred to as subclinical mastitis. Mastitis is chronic when the inflammatory process persists for months. It may remain subclinical indefinitely or, in some cases, may have temporary exacerbations of subacute or acute nature. The existence of a pathogen within the mammary gland without any evidence of mastitis is referred to as latent infection (Jain, 1979) The internal environment of a normal mammary gland is ideally sterile, but saprophytic bacteria may be found as commensals in some "normal mammary glands. However, to induce mastitis, a pathogen must first gain entrance into the mammary gland, survive the intramammary bacteriostatic and bactericidal agents, and then multiply in significant numbers. Mastitis begins with penetration of pathogenic bacteria through the teat canal (streak canal) into the interior of the gland. If the internal environment of the gland is favorable to survival and multiplication of the invading bacteria, the products of bacterial growth and metabolism may irritate the delicate mammary tissue and induce an inflammatory reaction. The clinical signs of mastitis are, in reality, an expression of the host defense intended to destroy the invader and to make way for repair to regain normality (Jain, 1979). The normal bovine mammary gland is divided by the medial susphensory ligament and supplied by separate arteries, veins and nerves. The front and rear quarters are separated from each other with regard to secretory tissue but share a common blood and nerve supply. Each quarter has a gland cistern, which serves as a collecting duct for that quarter. The gland cistern continues distally to the teat cistern and is demarcated by a distinc fibrous tissue, which is called the annular ring. At the most distal aspect of the teat cistern is the teat canal canal (papillary duct), which communicates with the outside of the teat (fig. 1) ( Weber, 1977; Trostle and OBrien, 1998).

12

Figure 1. Diagram of the main structures of the bovine udder (adapted from Sisson, 1975 and Willian, 1985).

The immune system of the mammary gland must serve a dual function: to provide passive immunity for the neonate and to protect the organ itself against pathogenic insult (Opdebeeck, 1982). The first line of defence the organisms encounter is the epidermis of the teat (McDonald, 1979; Fox and Gay, 1993; Sordillo et al., 1997). Resistence to bacterial invasion of a mammary quarter is determined, for the most part, by the structure and function of the teat canal. The normal teat canal has several anatomic features that act as barriers to penetration of bacteria (Smith, 1983; Hibbitt, 1983b). These features are most effective in the first lactation and tend to decrease with increasing lactational age. The lining of the teat canal consists of a stratified squamous

13

epithelium, like the skin of the teat, and its surface continually undergoes keratinization to form sebum-like material which fills the lumen of the canal (Smith, 1983; Hibbitt, 1983b). Teat canal keratin act as barrier to microorganisms involved in mastitis (Jack et al., 1992).This material is rich in long-chain fatty acids having a bacteriostatic affect on certain bacteria (Mosdoel, 1978). Partial removal of keratin from the teat canal has been reported to compromise the ability of the teat to prevent passage of bacterial pathogens from the external environment into the mammary gland (Bramley and Dodd, 1984; Capuco et al., 1992). The teat canal is surrounded by a true sphincter of smooth muscle fibers which function in maintaining a tight closure of the canal (fig. 1). Quarters having patent teat canals (lack of tight closure) have a greater incidence of infection (Jain, 1979; Sordillo et al., 1997). McDonald (1968) reported that the teat canals lengthen and dilate with increase in lactational age of cows. Motomura et al. (1994) reported that cows with larger canal diameters could make them more liable to contract mastitis. Other authors have found similar results (Grega and Szarek, 1985; Jorstad et al., 1989; Hamana et al., 1994). However Kartashova et al. (1992) reported no significant correlation between teat diameter and incidence of mastitis. The neutrophile polymorphonuclear (PMN) leukocytes is the second line of defense against mammary gland infection (McDonald, 1979). The PMN leukocytes make up most of the leukocytes in circulating blood of many animal species. However, in the bovine, PMN leukocytes make up only 25% of the total leukocyte count (Paape et al., 1979). These neutrophils are capable of phagocytosing a wide variety of particles. Phagocytosis is the process of recognition, ingestion and digestion of foreign particles. In the bovine species there is a difference between the phagocytic ability of PMN leukocytes of milk and PMN leukocytes of blood (Paape et al., 1979). It has been reported that the number of staphylococci killed by PMN leukocytes isolated from milk was significantly less than the number killed by PMN leukocytes isolated from blood (Russell and Reiter, 1975). This inhibition of function in the milk has been attributed to poor glycogen reserves within the cell and diversion of phagocytic effort into the ingestion of milk fat globules and casein (Opdebeeck, 1982). Immunoglobulins also have an important role in the pathogenesis of mastitis. These in mammary secretation are either humoral origin or are made locally by cells of the lymphocyte plasma cell series located near the glandular epithelium. The most of the IgG in mammary secretation has a humoral derivation whereas IgA and IgM are synthesized locally (Lascelles, 1979; Hibbitt, 1983a). Gram-positive cocci account for more than 90% of infections in the bovine mammary gland. The opsonization of bacteria and neutralization of toxins constitute important effector functions in the protection of the gland and these functions are mainly ascribed to the IgG immunoglobulin (Opdebeeck, 1982).

14

Some studies have investigated if there is a relationship between oestrus and incidence of mastitis (Reece and Murphy, 1943; Berger and Francis, 1951; Frank and Pounden 1961) as well as the influence on mastitis of forages containing legumes which have estrogenic properties (Frank et al., 1959; Pounden et al., 1960; Pounden and Frank, 1961; Frank and Pounden; 1961). The main results of these investigations were that an excess of oestrogens might predispose to mastitis, mainly at oestrus when estrogenic activity is considered to be highest. Also it was found that the incidence of mastitis was related positively to feeding legume-grass forages in the fresh state or as silage.

15

COMMON TYPES OF INFECTIOUS MASTITIS

A total of 137 microbial species, subspecies and serovars have been isolated from the bovine mammary gland (Watts, 1988). However, the most common causes of udder disease include staphylococci (Staphylococcus aureus and Staphylococcus epidermidis), streptococci (Streptococcus agalactiae, Streptococcus dysgalactiae, Streptococcus uberis and Streptococcus bovis) and coliforms (mainly Escherichia coli and Klebsiella pneumoniae). Other etiological agents less frequently encountered include pseudomonads, nocardia, mycoplasmas and yeast (McDonald, 1979). Organisms colonizing the mammary gland can be divided as well into one group, referred to either as minor pathogens or commensals (e.g. corynebacterium bovis or coagulase negative staphylococci) and a second group containing the major pathogens (streptococci spp., S. aureus and coliforms) (Dohoo and Meek 1982).

Staphylococcal mastitis This type of mastitis is usually associated with udder infection by S. aureus. This bacteria mainly produces subclinical and chronic mastitis, but it also may cause peracute mastitis and lead to gangrene of the quarters. Bacterial toxins and toxic products are thought to be involved in the causation of mastitis and gangrene. The alfa toxin is potentially the most damaging, because it causes vasoconstriction leading to ischemic necrosis of affected tissues and gangrene (Schalm, 1977). Gangrenous mastitis is most often found in young cows and after calving. Coagulase and other bacterial products are thought to enhance infection, allowing bacterial growth in the face of host defense mechanisms like phagocytosis (Jain, 1979). Delayed type hypersensitivity is considered an important part of the pathogenesis of staphylococcal mastitis and is related to peptidoglycan fraction of the cell wall (Woolcock, 1979). The principal reservoirs of S. aureus are the udder and teat skin and the milk of infected glands (McDonald, 1977). As few as 10 colony forming units (CFU) of S. aureus can infect the udder. The organisms have the capacity to penetrate tissues producing deep seated foci; hence, intramammary antibiotic therapy is not completely successful in eradication of staphylococcal mastitis. Leukocytosis in milk is not successful in disposing of virulent S. aureus because a) certain bacterial products like alfa toxin and leucocidin are damaging to neutrophils, b) protein A is antiphagocytic, and c) certain bacterial products protect them from intracellular killing. Intramammary infections with S. epidermidis are being increasingly recognized, but it is still not considered a serious mammary pathogen since most infections are eliminated spontaneously (Jain, 1979).

16

Streptococcal mastitis In this case species frequently reported to cause mastitis are Str. agalactiae, Str. dysgalactiae and Str. uberis, with the first one being most prevalent (McDonald, 1977). Str. agalactiae multiplies in the milk and on the mammary epithelial surfaces, generally causing a subacute or chronic inflammatory reaction with periodic acute flareups. The affected tissue eventually is destroyed resulting in reduced milk production or agalactia. Str. agalactiae is an obligate parasite of the udder and, unlike coliform organisms, it does not survive in the environment of the cow; hence, it is relatively easy to eradicate. Str. uberis and Str. dysgalactiae are not obligate udder pathogens. They can survive for long periods in the environment of the cow and may be cultured from skin and other parts of the cow; hence these organisms are difficult to eradicate (Jain, 1979). These infections are not contagious as is Str. agalactiae. These organisms invade the udder when conditions become favorable and may cause acute or chronic mastitis but more commonly a subclinical reaction. Although udder and teat surfaces seem to be the most common reservoirs of Str. uberis, mastitis due to this organism is less frequent (Jain, 1979).

Coliform mastitis The coliforms organisms commonly involved in mastitis are Escherichia coli, Klebsiella sp., and Enterobacter aerogenes, the first one being most prevalent (Eberhart, 1977). Their pathogenic effect is an attribute of endotoxin contained in bacterial cell wall. Coliform bacteria are ubiquitous in the environment of the cow. Coliform mastitis is typically acute or peracute, but chronic and subclinical infections accompanied with periodic acute flareups also occur. Cows suffering from peracute mastitis may die within a few days due to endotoxemia but usually overcome acute mastitis. Coliform mastitis is generally self-limiting and does not cause extensive damage to the mammary parenchyma; hence milk production does not decrease significantly following recovery (Jain 1979). Infections usually are acquired from the environment via the teat canal and are not transmitted directly from cow to cow, as is the case with most streptococcal and staphylococcal infections. Although coliforms are widespread in the environment of the cow and frequently are isolated from teat skin, coliform mastitis is relatively uncommon. This is perhaps due to relative susceptibility of most coliform bacteria to humoral and cellular factors in milk. Natural antibodies to coliform organisms in milk and leukocytosis in the udder generally prevent establishment of udder infection with most of these organisms. Hence, glands infected with more common udder pathogens and experiencing mild leukocytosis generally remain free of coliform infection. Therefore, coliform mastitis is considered to be a disease of the gland uninfected with other pathogens (Jain, 1979).
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THE EPIDEMIOLOGY OF MASTITIS IN BEEF CATTLE

Mastitis can be an important factor in preventing beef cows from achieving their true genetic potential for milk production (Kasari and Gleason, 1996). Two epidemiological tools used in order to quantify udder health are prevalence and incidence. Prevalence measures the amount of mastitis prevailing at a given point in time, and incidence measures the rate of new cases of mastitis over a period of time (Thurmond, 1993). Newman et al. (1991), working with purebred and crossbred beef cows reported that the numbers of new intramammary infections (IMI) with S. aureus at each sampling time (early, mid and late lactation) tended to be about equal to the number of infections lost (infected quarter recovered), so their prevalence did not greatly increase during lactation. Incidence measures are of particular interest because they provide estimates of risk (Thurmond, 1993). Figures on prevalence of mastitis has been reported in a number of studies in beef cattle. In a two-year study made in two beef suckler herds; in which the cow not only raised her own calf but several other calves were frequently purchased and were raised to weaning on the same cow, 2400 quarter milk samples were examined. The researchers found that 18% of all quarters were infected and 67% of all infections were due to staphylococci and 20% to streptococci (Hunter and Jeffrey, 1975). Sobari et al. (1976) reported a prevalence of 42.3% in beef cows from various parts of Northern Queensland. Kirkbride (1977) reported that of 20 beef cows, 8 were infected (40%) with coagulase-positive staphylococci. These cows were 2 to 6 years old and had been lactating 1 to 8 months. Haggard et al. (1983) reported in two different beef herds, a prevalence of 13% (12 of 92) in confined Angus cows and heifers; and 10.7% (8 of 75) in Hereford and Herefordcross-bred females that were in a range-pasture operation. The total prevalence (both groups) was 11.9%. In the confined group the agents identified were S. aureus (n=9), streptococcus spp other than agalactiae (n=1), and klebsiella spp (n=2). For the rangepasture herd S. aureus was the only agent identified. This test was done 30 days after calving. In 1600 Charolais cows 24 (1.5) cases of clinical mastitis were detected, most appearing during the calving period, also infected quarters rarely recovered and frequently dried up, sometimes temporarily but often permanently (Boucomont, 1985). Watts et al. (1986) identified IMI in 37% of Hereford, Hereford x Brahman, and Hereford x Brown Swiss cows and in 18% of the quarters. Coagulase-positive staphylococci and coagulase-negative staphylococci were isolated in 17.9% and 16.1% of the cows respectively. For coagulase-positive staphylococci, S. aureus accounted for 39.9% of these infections. Samples from quarters in this study were collected at parturition and at weaning.

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Timms et al. (1989) working with 80 beef cattle reported high levels of infection at calving (41% of cows, 17.5% of quarters), throughout lactation (33%, 13%) and at weaning (28%, 12%). Predominant organisms in this study were coagulase-negative staphylococci (67% of infections), S. aureus (28%) and environmental streptococci (5%). In 213 quarter milk samples taken at random from 7 beef herds, it was found that 36% of cows and 14% of quarters had subclinical mastitis, mostly associated with coagulase-negative staphylococci (Hoyer et al., 1991). Newman et al. (1991) reported 25.8, 29.2 and 54.4% of the cows being infected with mastitis pathogens in early (2 to 4 weeks postcalving), mid (100 days postcalving) ans late lactation (200 days postcalving) respectively. The corresponding prevalence of infection in quarters in that study was 13.1, 14.9 and 27.5% respectively. S. aureus was isolated from 2.9, 2.7 and 3.2% of quarters in early, mid and late lactation. Simpson et al. (1995) found that in 25 primiparous Simmental cows, 32% of the cows and 18% of the quarters were infected with mastitis organisms at least once during their initial lactation. The sampling on these cows was done on day 34, 49, 80, 108, 147 and 189+/3 postpartum. Nickerson et al. (2000) working with beef heifers observed that the prevalence of infection in quaters was 10% in early lactation and the main microorganisms isolated were coagulase-negative staphylococci, followed by S. ureus. Ridgway et al. (1999) also reported have isolated these two microorganisms from Angus x Hereford cows. As we can see the prevalence of IMI in beef cows in these works ranged from 10% to 54% and the prevalence of infected quarters ranged from 10% to 27%. These data also show that the predominant microorganisms isolated were coagulase-negative staphylococci and S. aureus and that most of the IMI occurred soon after calving (early lactation). In dairy cows it has been found that most of the IMI that ultimately develop into clinical mastitis do so during the first few weeks of lactation, which correlates well with immunosuppression observed during this period (Kehrli and Shuster, 1994) Compared to data from dairy cows the prevalence of infection by major pathogens in beef cows is low. Prevalence of major pathogens within a dairy herd is influenced by mastitis control practices, especially teat dipping and dry cow therapy, and therefore, varies considerably from herd to herd. The usual method of transmitting major pathogens in a dairy herd is from cow to cow during milking. In beef cows, with the calf as a vector, the organism can be spread from one quarter to another in the same cow, but spread from cow to cow seems unlikely because cross-suckling is believed to occur only rarely in beef cows (Hoyer et al., 1991; Newman et al., 1991). Hence it is possible that in beef cows, the prevalence of major pathogens remains relatively low because of the absence of opportunity for infection at milking time (Newman et al., 1991).

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SOMATIC CELL COUNT

The alveolar epithelial cells of the mammary gland are highly active secretory cells that are normally subjected to continuous high turnover and therefore must be replaced continually with new cells. These displaced cells are discharged into the milk as a normal process. In response to irritation, whether physical or microbial, white blood cells enter the milk from the blood and perialveolar interstitium. The displaced alveolar epithelial cells, phagocytes and white blood cells comprise the somatic cells of the milk (Kirk, 1984; Concha, 1986; Sordillo et al., 1997). The measurement of the number of somatic cells in milk is known as a somatic cell count (ORourke, 1999). Somatic cell count (SCC) of milk samples, is one of the techniques used to monitor the level or occurrence of subclinical mastitis in herds or individual cows or quarters (Dohoo and Meek, 1982). The prevalence of subclinical infection is difficult to ascertain, as this can only be determined by the isolation of bacteria from the gland; however, SCC is often used as an indirect estimation (Leigh, 1999).

FACTORS AFFECTING SOMATIC CELL COUNT

The ability to correctly interpret SCC depends on an understanding of the factors which may affect them. These factors may exert their influence at the quarter, cow or herd level (Dohoo and Meek, 1982). The following factors have been identified; infection status, season, age, stage of lactation, diurnal variation, day to day variation, stress, technical aspects, effects of quarter and milk fraction.

Infection status The most important factor affecting the SCC of the milk from an individual quarter, and consequently the cow and the herd, is the infection status of the quarter (Dohoo and Meek, 1982; Kirk, 1984; Sender et al., 1987; Schepers et al., 1997). The normal somatic cell count in milk in various parts of the same udder varies widely from near zero in uninfected areas to something in the order of three hundred million cells per litre in the worst infected areas (ORourke, 1999). It is difficult to nominate appropiate cell count thresholds above which the udder is termed diseased (Francis, 1993). The main problem has been determining a correlation between a given somatic cell count and the corresponding probability that infection might be present (Kirk, 1984). In dairy cattle both 400 000 and 500 000 cells/ml have been evaluated as possible thresholds, for classifying a quarter as being infected, but both resulted in a high false negative rate (i.e. too many cases of subclinical mastitis were missed) (Dohoo and Meek, 1982). On a quarter basis, Poutrel and Rainard (1982) found that 93% of samples from uninfected quarters had cell counts of less than 500 000 cells/ml, but 45% of quarters infected with major pathogens also had counts of less than 500 000 cells/ml. However in beef cattle it has been reported
20

that staphylococcal infections were associated with cell counts over 500 000 cells/ml (Hunter and Jeffrey, 1975; Nickerson et al., 2000). It has been recommended that the threshold be set at 200 000 cells/ml for firts lactation (Sender et al., 1987; Dohoo and Leslie, 1991; Schepers et al., 1997) at 300 000 cells/ml for all lactations (Dohoo and Meek, 1982; Sender et al., 1987). aureus However Neave (1975) found that 11% of quarters with long lasting Staphylococcus aureus had, at one or more samplings, cell counts of less than 100 000 cells/ml, and 19 % of these quarters had at some time a count of less than 300 000 cells/ml. Also secretory disturbances including declining milk yield, have been reported to start once cell counts exceed 100-150 000 cells/ml, and the probability of isolating a major pathogen is increased with counts above 200 000 cells/ml (Dohoo and Meek, 1982). Hence no matter what threshold is used, a proportion of quarters or udders will be incorrectly classified (Francis, 1993). Therefore the evaluation of several successive counts is preferable to the interpretation of an individual count (Dohoo and Meek, 1982). Sampling at least five times during a lactation has been recommended (Clarkson, 1975).

Season The season appears to affect the SCC of dairy herds in various locations and is not entirely related to ambient temperature or stressful seasonal conditions (Dohoo and Meek, 1982). In general, SCC are lowest during winter and highest during the summer. Data from Wisconsin shows a peak SCC from July through August and a low count in March; marked differences also existed between months during different years (Bodoh et al., 1976). Data from Quebec, Canada, indicates a low count in May, with a steady increase beginning in June and registering the highest count in December. The increase, which began when the cows went to the pasture and peaked when they returned to housing is possibly attributable to environmental stress and renewed challenges from bacteria (Kennedy et al., 1982). During the summer in Scandinavia, cows on pasture (cooler temperatures) have had higher cell counts than cows confined to the barn (warmer temperatures), and the increase in cows on pasture was primarily seen in noninfected quarters (Simensen, 1976). In a recent study Pomies et al. (2000) concluded that the increase in SCC observed in summer is not due to the environmental change when cows are turned out to pasture, and alternative explanations were suggested, such as physiological, health or climatic factors which may be exacerbated by the period at pasture. No studies on seasonal

21

effects on SCC in beef cattle were found. In temperate climates, such as in Scandinavia, beef calves are generally born in the spring or early summer. It is therefore difficult to analyse any seasonal diference in SCC, as any seasonal change will always be confounded by the lactational stage.

Age An increase in the cellular content of milk with increasing age has been reported in beef cattle (Duenas et al., 1994) and in dairy cattle (Daniel et al., 1966; Schultz, 1977; Gill and Holmes, 1978; Syrstad et al., 1979; Taralik, 1998; Choi et al., 1999). However, Wilson et al. (1971), working with Angus-Holstein F1 cows (dual-purpose cattle), with an age from 2 to 7 years found that number of lactations did not significantly affect SCC. Laevens et al. (1997) found similar results in dairy cattle.The increase in SCC with the age is primarily due to an increased prevalence of infection in older cows and is not due to any large increase due to the age per se, being a pathological rather than a physiological change (Kirkbride, 1977; Dohoo and Meek, 1982). It has been found that older cows have a greater cellular response to both minor and major pathogens, and therefore older cows usually have high cellular readings (Dohoo and Meek, 1982; Kirk, 1984). This has been attributed to a number of factors, including more quarters being infected, more extensive tissue damage in long lasting infections, and a greater cellular response in quarters that have been previously infected (Dohoo and Meek, 1982).

Stage of lactation In dairy cattle SCC has been found to be high in the first week after calving, with the lowest count occurring at the peak level of production, and tend to rise slightly in the last few weeks of lactation (Kennedy et al., 1982; Kirk, 1984; ORourke, 1999). stensson (1993b), working with dairy cattle, reported that SCC did not change significantly but tended to increase as lactation proceeded. Wilson et al. (1971) in dualpurpose cows found the SCC was significantly influenced by lactation stage, reporting, as in dairy cattle, that SCC tended to be the highest early in lactation, declined, and then increased appreciably at the end of lactation. Another study in beef cows also found that early and late lactation animals were more frequently associated with high cell counts than animals in mid-lactation (Hunter and Jeffrey, 1975). However Laevens et al. (1997) reported no effect of stage of lactation on SCC in dairy cows. A partial explanation for this variation could be the milk yield of the cow during lactation, as Emanuelson and Funke (1991) and Miller et al. (1993) found a dilution effect due to an inverse relationship between milk yield and milk SCC. In these works it was assumed that a dilution effect caused the regression of milk yield on milk SCC. Miller et al. (1993) suggested that the observed negative relationship between milk yield and SCC may partly reflect both the true biological effects of udder inflamation and a dilution effect. stensson (1993b) mentioned that the milk somatic cell counts
22

increases towards the end of lactation, because of the higher prevalence of mastitis, normal involution of the udder and decreased milk production which causes less dilution of the milk leucocytes.

Diurnal variation Diurnal variation in SCC has been reported in dairy cattle; Smith and Schultze (1967) reported that cell counts are lowest just prior to milking and are highest immediately after milking with these levels persisting for up to four hours, then decreasing progressively to a minimum value near the end of the intermilking period. These authors suggested that most normal quarters have a rather high leucocyte concentration for as long as 8 hours after each milking. Syrstad and Ron (1978) found that samples from morning milkings had about 20% lower cell counts than samples from afternoon milkings. White and Rattray (1965) reported similar results. These differences have been attributed to a dilution effect, where SCC decrease as volume of milk increase (Emanuelson and Funke, 1991; Miller et al., 1993). The dilution effect probably result from the cyclic pressure changes in the alveoli, where for the first few hours after milking, the pressure within the alveolus is lowest; this tends to cause a release of polymorphonuclear leucocytes and squamous epithelial cells into the milk. Therefore, the samples taken at this time have the highest daily SCC, whereas samples taken just before milking have the lowest SCC, because increase in alveolar pressure prevents cells from being released into milk while milk synthesis and release of milk into the lumen continues, thereby diluting concentration of somatic cells in milk (Donovan et al., 1992). Newman et al. (1991) working with beef cows suggested that the low levels of SCC found in their study could be a result of all samples being collected about 16 hours after the cows had last been nursed. Thus, cells present may have been diluted by relatively high milk volume.

Day to day variation In dairy cattle it has been reported that cell counts of cows also vary from day-to-day by up to 25% of the baseline count. The variation is small in uninfected cows but may be much greater in cows with active infections (Kirk, 1984). It has been reported that fluctuations in individual quarter samples from uninfected cows have run in parallel, suggesting physiological factors acting at the cow level (Dohoo and Meek, 1982). Donovan et al. (1992) mentioned that day-to-day variation in milk SCC, could be due to other factors affecting SCC like age, stage of lactation, environmental temperature and stress. However, Simpson et al. (1995) reported that day of sampling did not influence SCC in beef cattle. Stress

23

It has been reported in dairy cattle, that stress can affect the SCC. In a study in which the cows were stressed by corticotropin injection, confinement in a heat-humidity chamber, or environmental heat stress by exposure during the hot summer months of June through November in southern Arizona; a modest increase in SCC was found in cows treated with corticotropin injection and exposed to environmental heat stress, but no significant changes occurred during the chamber heat stress period (Wegner et al., 1976). However Convey et al. (1971) reported that corticoid-induced leucocytosis of blood is not reflected by increases in somatic cell numbers in milk. In another trial with a heat-humidity chamber, Roussel et al. (1969) reported that there was no evidence that increased thermal stress significantly caused an elevation in the SCC. Nelson et al. (1969) working in environmental heat stress, reported that samples having somatic cell counts in excess of 500 000/ml were greatest during periods of maximum temperature, and that individual quarters responded independently during the period of high atmospheric temperatures. This indicates that, while stress of the entire animal undoubtedly is a factor, manifestation of the stress condition is determined by factors operative in the individual quarters of the animal. It has been reported that there is no increase in SCC associated with cows being in oestrous. In a study involving 14 cows; it was found that the oestrous did not affect SCC, milk production or occurrence of clinical mastitis (Guidry et al., 1975). stensson (1993b) neither found differences in SCC between oestrous and dioestrous in various milk fractions.

Technical aspects The method of transportation and storage of milk samples, as well as the method used to count the somatic cells, all have an influence on the resultant counts (Dohoo and Meek, 1982; Coleman and Moss,1989). Fresh, nonpreserved samples become unacceptable for use 16 hours following collection. Refrigerated, nonpreserved samples remain acceptable for up to three days (Kirk, 1984). Differences among preservatives were not found in studies comparing bronopol, potassium dichromatic, sodium azide, boric acid and milkofix ( Hamann, et al., 1991, Hanus et al., 1992; Bertrand, 1996). However some differences can be found, which will depend of the amount of preservative used in the milk sample, temperature and time of storage (Heeschen et al., 1993; Gencurova et al., 1994). SCC are lower after freezing than before freezing and the decrease is greater when the freezing period is longer , but when thresholds of 200 000, 250 000 and 500 000 cells/ ml are used to predict infection, freezing appear to have a little impact on the sensitive and specifity of diagnostic parameters (Barkema et al., 1997). Effects of quarter Wilson et al. (1971) reported that in dual-purpose cows means for log SCC were lower for front than for rear quarters, although differences between left front and left rear
24

quarters were not significant. These authors suggested that a partial explanation for this, could be the sucking preferences of the calves, because in their trial they found that there was a preference for calves to nurse the fore quarters more frequently than the rear quarters . Failure to partially or completely empty a quarter would tend to increase the cell count of the milk (Wilson et al., 1971). Natzke et al. (1965) reported increased milk cell content from the omission of a milking. Milk fraction In dairy cattle SCC may vary considerably among different fractions of milk obtained at a single milking (Paape and Tucker, 1966, stensson and strm, 1994). Berning et al. (1986) reported that SCC was higher in residual milk than in foremilk. Berning et al. (1987) found that SCC (log thousand cells/ml) in foremilk was 2.52 and in residual milk 5.53. stensson (1993a) reported that in healthy udder quarters there were significant differences in SCC between foremilk and residual milk, with the higher values for residual milk. Similar results have been found by other authors (Woolford et al., 1998; Hamann and Gyodi 1999; Waldmann et al., 1999). stensson (1993a) also reported that during the course of inflammation there were no consistently significant differences between foremilk and residual milk in the SCC . The higher values for SCC in residual milk, compared with foremilk may suggested that cell counts in residual milk is a more sensitive measure of the inflammatory condition in the udder tissues. Differences between foremilk and residual milk may be attributable to that cells, because of their close contact with the epithelium, are retained in the alveoli and ducts until residual milk is collected (stensson, 1993b). stensson et al. (1988) suggested that for research purposes it may be adequate to analyse 2 milk samples, for instance foremilk and residual milk, in order to get a more precise characterization of the condition of the udder. Although there are no data in beef cattle on some factors reviewed above (that are affecting SCC), it is important to consider them during udder health assessment in beef cows.

25

BOVINE MASTITIS BACTERIOLOGY

The prevalence of subclinical infection is difficult to ascertain, as this can only be determined by isolation of bacteria from the gland (Leigh, 1999). Many organisms present in the cows environment can cause mastitis. When one of these organisms is isolated from mastitic milk the question arises as to whether it is the cause of the mastitis, or a contaminant (Jones, 1994). Bacteriology and cell counting are sometimes used together to determine the infection status of the mammary gland (Barkema et al., 1997). However the interpretation of these combined results is not straightforward (Francis 1993; Shoshani et al., 2000). Francis (1993) present some of the possible reasons for contradictory results (Tab. 1).
Table 1-Possible reasons for inconsistent results using bacteriology and somatic cell counts Together (Francis, 1993). High cell count and NEGATIVE Bacteriology Small numbers of bacteria not being detected in the 10 ul aliquot cultured Culture medium unsuitable for the causal agent Milk sample is bactericidal (death in the post) Physical or chemical damage to the udder Teat canal infections producing endotoxin which raises the cell count Low cell count and POSITIVE Bacteriology Teat canal infections

Sample contaminated during collection

Long-standing subclinical infection

Contaminated sample bottle

26

MILK CONSTITUENTS

Milk is a biological fluid containing a large number of different constituents (Davies et al., 1983; Kennelly, 1996). However most of the research carried out on milk composition has been done investigating the major constituents, which are fat, protein and lactose. The composition of milk in dairy cows generally ranges between 3.0-5.3% fat, 2.84.5% protein and 4.2-5.1% lactose. Some works have reported either only fat (Morgan, 1991; Griinary et al., 1997), or fat and protein (Oldenbroek, 1984; Higginbotham et al., 1988; Jones-Ensley et al., 1997; Pirlo et al., 1997, Wu et al., 1997; Bremer et al., 1997). However other studies have reported the all three constituents (Tab. 2).
Table 2. Percentage of fat, protein and lactose in milk from different dairy breeds. Breed type Milk fat (%) 3.5 3.3 3.9 3.6 3.8 4.9 4.3 5.0 4.3 3.2 4.6 4.0 3.8 Milk protein (%) 3.0 3.7 2.8 3.0 2.9 3.6 4.5 3.7 4.5 3.2 3.5 3.3 3.2 Milk lactose (%) 4.6 4.5 4.9 4.9 5.1 4.7 4.2 5.0 4.2 3.8 4.8 4.6 4.8 Reference

Holstein Holstein Holstein Holstein Holstein Jersey Jersey Jersey Jersey Dairy* Guernsey Ayrshire Brown Swiss

Jennes 1985 Sharaby 1988 Nagel and Broderick 1992 Rodriguez et al. 1997a Rodriguez et al. 1997b Jenness 1985 Sharaby 1988 Rodriguez et al. 1997a Rodriguez et al. 1997b Mondragon et al. 1983 Jenness 1985 Jenness 1985 Jenness 1985

*=Brown Swiss and Holstein and their crosses

In beef cows a variation of 2.5-6.5%, 3.0-4.0% and 3.8-5.3% for fat, protein and lactose respectively has been found. Some researchers have reported 3 major constituents of milk from beef cows (Tab. 3). However some investigations have worked only with fat (Danilevskaya et al., 1972; Carwright et al., 1976; Belcher and Frahm, 1979; Franke and Martin, 1983; Morgan, 1991; Kovacks, 1997), fat and protein (Rahnefeld, et al., 1990; Lalman et al., 2000), or protein and lactose (Mollet et al., 1989).
Table 3. Percentage of fat, protein and lactose in milk from different beef breeds. 27

Breed type

Milk fat (%) 4.1 4.9 6.3 4.3 5.8 4.9 3.4 5.8 5.7 5.9 5.7 6.5 5.7 2.8 3.1 3.6 3.1 3.1 3.9 3.3 4.9 4.8 4.8

Milk protein (%) 3.3 3.5 3.3 3.8 3.0 3.5 3.5 3.0 3.1 3.2 3.1 3.3 3.2 3.3 3.2 3.3 3.5 3.3 3.5 3.5 3.5 3.5 3.5

Milk lactose (%) 4.7 5.5 5.0 5.1 5.2 5.1 5.2 5.0 5.1 5.1 5.1 5.1 5.0 5.3 5.1 5.2 5.2 5.0 5.2 5.1 5.5 5.2 5.3

Reference Beal et al. 1990 Butson and Berg 1984 Daley et al. 1986 Meregalli et al. 1983 Daley et al. 1986 Kovacs et al. 1999 Mondragon et al. 1983 Daley et al. 1986 Daley et al. 1986 Daley et al. 1986 Daley et al. 1986 Daley et al. 1986 Daley et al. 1986 Mondragon et al. 1983 Mondragon et al. 1983 Mondragon et al. 1983 Mondragon et al. 1983 Mondragon et al. 1983 Mondragon et al. 1983 Mondragon et al. 1983 Butson and Berg 1984 Butson and Berg 1984 Butson and Berg 1984

Angus Hereford Hereford Chianina Red Poll Hungarian Grey Charolais Angus x Hereford Hereford x Red Poll Red Poll x Hereford Angus x Charolais Brahman x Hereford Brahman x Angus British1 Dairy x British Charolais x British Burwash x British Dairy x Burwash Jersey x British Exotic x Dairy Beef-Synthetic Dairy-Beef
5 4 3 2

Dairy-Synthetic6

1=Angus, Hereford, Shorthorn and their crosses 2=Brown Swiss, Holstein and their crosses 3= Chianina, Limousin, Charolais and Maine Anjou 4=crosses of Charolais, Angus and Galloway 5=crosses of dairy sires with Hereford or beef-synthetic dams 6=crosses of Holstein or Brown Swiss with beef breeds

NUTRITIONAL FACTORS AFFECTING MILK CONSTITUENTS

When discussing milk constituents, one must be careful to distinguish between yield (quantity) and content (percentage) (DePeters and Cant, 1992). The chemical composition of milk depends on factors related to the animal and to its environment (Hoden and Coulon, 1991). Fat content
28

The percentage of milk fat is influenced by a number of interacting dietary factors (Ashes et al., 1997). It has been proposed that the percentage of milk fat is affected by plasma insulin concentrations. The glucogenic-insulin theory proposes that increased insulin release which occurs when high starchy concentrate diets are fed, preferentially channels nutrients to adipose tissue, resulting in a shortage of nutrients at the mammary gland and thus milk fat depression (Brockman and Laarveld, 1986; Sutton et al., 1988; Griinari et al., 1997a). This theory has been questioned because injections of insulin in some cases did not result in a lower milk fat content (Mcguire et al., 1995; Griinari et al., 1997a). It is therefore possible that other hormones are also involved in the regulation of milk fat synthesis (Samuelsson, 1996). Samuelsson et al. (1998) found a positive relation between somatostatin and milk fat content. They hypothesised that one way for somatostatin to stimulate milk fat synthesis was through a modulation of the action of insulin in which high levels of somatostatin promotes deposition of fat in the milk through an inhibition of lipogenesis in adipose tissue, i.e. the milk synthesis in the udder is given priority before the deposition of fat in the body. Samuelsson (1996) mentioned that cows on a high plane of nutrition generally have a reduced fat content in the milk whereas cows with low energy intake have an increased fat content. It has been reported a depresion in milk fat content in diets contain high amounts of grain (fermentable starch)(Palmquist et al., 1993; Jenkins, 1993). In primiparous beef heifers Lalman et al. (2000) comparing four treatments with low (L), maintenance (M), maintenance high (MH) and high dietary energy concentration (H), found that increasing dietary energy it was increased milk fat percentage, however this effect apparently was seen only in the L, M and MH treatments, as the lowest milk fat percentage was found in the H treatment (3.5, 3.8, 3.7% vs. 3.3%). Lowman et al. (1979) working with beef cows reported that increasing the energy allowance significantly increased milk yield but did not affect milk composition. The effect of dietary energy on milk composition vary depending of the source (ingredients) of energy used in the cows diet and other dietary factors like effective fiber (chewing time), particle size and additives (buffers) (Schultz, 1974; Emery, 1988; Jelec, 1990; Hoden and Coulon, 1991). The role of protein in milk fat was investigated by Jones-Enslay et al. (1997) who reported that the percentage of milk fat produced by grazing dairy cows was unaffected by crude protein in the supplement or by the amount of supplement offered. Emery (1988) mentioned that amount of protein have a little or no effect on content of milk fat. Emery (1991) mentioned that increasing crude protein from 10% to about 18% of the diet in lactating dairy cows, milk fat percentage may decrease due to dilution by increased milk yield. In studies investigating the effect of ruminally undegradable protein (RUP) milk fat percentage was reported to be affected by RUP (Christense et al., 1992a; Christensen et al., 1992b; Wiley et al., 1991) whereas others reported no effect (Wu et al., 1997). Also is important to take into consideration the intake of water. In a study done in Israel, it was reported that dairy herds given insufficient drinking water produced milk with significantly increased water content and significantly reduced fat content. It was
29

suggested that these changes are physiological adaptations to enable sucking young to survive (Yagil et al., 1986).

Protein content Protein percentage in milk is also influenced by nutritional factors. The amino acids which are taken up by the mammary gland for milk protein synthesis arise from dietary undegraded protein which reaches the duodenum and rumen microbial protein (Murphy and OMara, 1993). According to this higher values of percentage of milk protein have been observed in abomasal infusion of casein (Griinari et al., 1997b) and in diets with high RUP (Komaragiri and Erdman, 1992; Bremmer et al., 1997; Wu et al., 1997). However Rodriguez et al. (1997a) reported that milk protein content was reduced in diets with high amounts of RUP. It has been found that increasing dietary energy concentration resulted in an increase in milk protein percentage (Murphy and OMara, 1993; Lalman et al., 2000) and vice versa (Wiley et al., 1991). However Pirlo et al. (1997) reported that milk protein percentage was positively influenced by diets with low energy. In some investigations it has been found that fat supplements often have decreased milk protein percentage (Coppock and Wilks, 1991; DePeters and Cant, 1992; Wu and Huber, 1994; Rodriguez et al, 1997a). The mechanism for this decrease in milk protein is not understood. Current theories include ruminal effects; such as toxic effects of fat on ruminal microbes, decreased microbial protein synthesis when fat is substituted for ruminally available carbohydrates, or absorption of certain fatty acids that may directly or indirectly alter uptake of amino acids by the mammary gland (Chow et al., 1990). Grummer (1991) suggested that protein percentage decrease maybe due to a dilution effect when milk yield increases as a result of fat supplementation. It has been reported that increasing the protein level in the diet had no effect on the milk protein concentration (Emery, 1991; Christensen et al., 1992a; Murphy and OMara, 1993; Jones-Endsley et al., 1997). However Pirlo et al. (1997) reported higher milk protein percentages in high protein diets.

Lactose content Milk lactose is the main osmotic component in milk and therefore the lactose content is rather constant (Schultz, 1974; Samuelsson, 1996). Wiley et al. (1991) evaluated the effect of diets made to induce or prevent weight loss before parturition and diets after calving with ruminally undegradable or degradable protein supplement. They found no

30

effect of either of these four treatments on lactose content of milk from primiparous beef cows. However Rodriguez et al. (1997a) found that the lactose content of milk increased 1.4% with added fat and increased 1.6% with diets having high RUP content in dairy cows.

NON-NUTRITIONAL FACTORS AFFECTING MILK CONSTITUENTS

There are several factors which are non-nutriotional and can have an affect on the constituents in milk, they are mainly breed, stage of lactation, udder health, milk fraction and some other factors.

Breed The effect of breed has been reported in some studies with beef cattle. Cartwright et al. (1976) compared Aberdeen-Angus x Jersey cows with Hereford cows and reported differences in milk fat percentages. Belcher et al. (1979) found no significant differences for milk fat percentage, but milk protein percentages had differences between breeds. Daley et al. (1986) found that milk fat percentage differed significantly among breed types. Breed effects for other milk traits were reported nonsignificant in that study. Rahnefeld et al. (1990) reported breed effects on milk fat percentage and on milk protein percentage. Sharaby (1988) found breed differences for fat, protein and lactose content in milk in Jersey and Holstein cows. Butson and Berg (1984) working with beef and dairy-beef cows reported that all milk constituents percentages were higher than those reported for commercial dairy cattle.

Stage of lactation The stage of lactation has been reported to have an effect on milk composition as well. In dairy cows in general tha fat (Dash et al., 1978; Yadav and Sharma, 1984; Chaudhary et al., 1998), protein (Ettala, 1976; Taralik, 1998) and lactose (Sharma et al., 1990; Holdaway et al., 1996) are high in early lactation, increase in mid lactation (peak milk yield) and decrease during mid- to late-lactation. However in some studies it has been reported that lactation stage had no significant effect on milk constituent percentages (Sharaby, 1988; Ibeawuchi and Dangut, 1996). In beef cattle milk fat percentage was found to be highest in early lactation, while milk protein percentage increased during lactation, and milk lactose percentage remained constant (Mondragon et al., 1983). Morgan (1991) reported the lowest values for milk fat percentage during the peak milk yield (middle of lactation) for Hereford cows. Butson and Berg (1984) working with beef and dairy-beef cows observed that all recorded milk constituent percentages increased significantly from June to September at about 130 days of lactation. Other authors have found similar results (Yanagita et al., 1978; Daley et al., 1986).
31

These data suggest that maybe this effect can partly be explained by the lactation curve of the cow, resulting in a dilution effect (fig.2). Dilution effect is the inverse relationship between milk yield and percentage of milk constituents. This is supported by Pitcher et al. (1992) who reported that the dilution effect was responsible for the changes in milk composition in dairy cows and by Ibeawuchi and Dangut (1996) who found a significant inverse correlation between milk yield and milk fat in Zebu cattle. Rahnefeld et al. (1990) found that the percentage of fat and protein decreased as milk yield increased in beef cows. Also Butson and Berg (1984) found that in beef cows, correlations between milk yields and constituent percentages were significant and negative. According to the previously mentioned Ettala (1976) observed that when the effect of milk yield was eliminated in the analysis, stage of lactation had only a slight effect on milk constituents. Owing to that changes in milk yield result in a dilution effect, the changes in percentage of milk constituents will automatically be more significant in dairy cattle than in beef cattle, as dairy cows have a higher milk production. This is belived to be a result of dairy cows having a greater number of secretory cells and greater activity per cell (Keys et al., 1989).

Fig. 2. Lactation curves 0 to 25 weeks for mature crossbred cows ( Angus x Hereford or Hereford x Angus; Charolais x Angus or Hereford; Jersey x Angus or Hereford; Simmental x Angus or Hereford). (From Jenkins and Ferrel, 1984).

32

Mastitis and somatic cell count (SCC) In a study done with Bos taurus and Bos indicus x Bos taurus breed types, Daley et al. (1986) found that mastitis had an effect on milk constituent percentages, reporting that cows that had mastitis in the tested quarter had a higher percentage of fat and protein in their milk, but a decreased percentage of lactose. Watts et al. (1986) working with beef cows reported reductions in fat and protein in milk from udders with intramammary infection caused by Staphylococcus aureus. Simpson et al. (1995) in Simmental cows reported an increase in fat and protein in milk with high SCC. These authors suggested that maybe inflammatory proteins and antibodies present in response to intramammary infections could have contributed to the higher percentage of protein in high SCC. However casein is the primary compound detected in the milk protein analysis. In dairy cattle, high cell count milk has lower fat and lactose levels than low cell count milk (Dohoo and Meek, 1982). Miller et al. (1983) reported low percentage of lactose in high SCC, but high percentages of fat and protein. Roussel et al. (1969) found a significant positive correlation between milk fat and SCC in dairy cattle. However Eicher et al. (1999b) found that SCC did not influence protein in milk from dairy cows..

Milk fraction In dairy cattle an effect of milk fraction on some milk constituents has been found. Moore et al. (1981) reported that milk fat percentage was significantly higher at evening than at morning milking for foremilk, but significantly higher at morning than at evening milking for residual milk. Protein content was significantly higher at evening than at morning milking for machine milk and residual milk. Carlsson and Bergstrm (1994) reported that fat percentage was higher in residual milk than in foremilk. Ugarte (1977; 1991) comparing normal milk (machine milking) and residual milk, reported that fat percentage was higher in residual milk, but protein percentage was lower in this fraction of milk. Waldmann et al. (1999) reported similars results.

Other factors affecting milk constituents In beef cattle has been found that parity had no effect on milk composition (Mondragon et al., 1983; Meregalli et al., 1983). In beef cattle it has been observed that dams with bull calves produced more fat and less protein than cows with heifer calves, and the calf age had a positive effect upon percentage of lactose, and as calf age increased the percentage of protein increased significantly (Daley et al., 1986). However Meregalli et al. (1983) have not found effect sex of calf on milk composition in beef cows. It has been found that estimates for all milk traits decrease as the length of time of separation (from their calves) increased in beef cows (Chenette and Frahm, 1981). However Belcher et al. (1979) reported that milk fat percentage decreased as time of
33

cow-calf separation increased in beef cattle but milk protein percentage was not affected. In dairy cows it was reported that udder shape had significant effects on milk fat and on milk protein, but not on lactose (Barbary et al., 1999). However Baruah et al. (1991) found that milk composition of dairy cows was not significantly correlated with any udder measurements. The effect of photoperiod and heat load on milk composition has been studied in dairy cows, and protein as found to be affected by photoperiod, fat was affected by head load and lactose was affected by both environmental factors (Aharoni, et al., 1999).

34

ARE TEAT TRAITS IMPORTANT IN UDDER HEALTH AND MILK COMPOSITION ASSESSMENTS?

FACTORS AFFECTING TEAT LENGTH AND ITS IMPORTANCE ON MILK YIELD AND UDDER HEALTH

The importance of teat length has been studied in beef cattle, and it has been found that calves born from cows with large teats had higher pre-weaning growth rates than those born from cows with smaller teats (Makarechian and Paputungan, 1998). In dairy cattle correlation values of 0.15 to 0.88 between teat length and mastitis have been reported (Janicki, 1977; Soldatov and Kholodkov, 1990). Lund et al. (1994) reported a correlation between clinical mastitis and teat length in Danish Holsteins cows; and concluded that their results indicated predisposition to mastitis with long teats. Similar results were found by Roy et al. (1993) in crossbred cows. On the contrary Shukla et al. (1997) reported that small teats were a risk factor for mastitis in dairy cattle and Murrah buffaloes. An increase in teat length has been found in beef cows with increasing age (Logan and Gibson, 1975; Olson et al., 1989; Prajapati et al., 1995). In dairy cattle fore teats have been reported to be longer than rear teats (Naidu, 1972; Oldenbroek, 1984; Gupta et al., 1991; Singh and Gupta, 1995). In dairy cows it has also been observed that the length of all teats increased with lactation number (Tomar, 1973, Sharma and Sidhu, 1980; Baruah et al., 1991). It has been reported that teat dimensions (including teat length) does not affect milk yield in dairy cattle (Naidu, 1972; Pander and Chopra, 1986; Ghosh and Prasad, 1998; Ozbeyaz et al., 1998) or in beef cattle (Prajapati et al., 1998). However, Tomar (1973) reported that milk yield in dairy cattle was significantly correlated with the length of both fore and rear teats. Baruah et al. (1991) working with Jersey cows reported a significant correlation between milk yield and rear teat length in the 1st and 3rd lactation of 0.36 and 0.65 respectively. Similar results were found in Gir cows by others (Tripathi et al. 1982; Qureshi et al. 1984) In Zebu, European and crossbreed cattle teat length has been reported to be affected by genetic group and lactation number, but not by stage of lactation (Sharma and Sidhu, 1980). However, Baruah et al. (1991) reported that udder measurements (including teat length) were greater in early-lactation than in mid-lactation and late lactation in dairy cows.

TEAT INJURY AND TEAT PAPILLOMATOSIS

The highly vascular and vulnerable mammary tissue is protected by the skin of the udder. In young animals it is thin and closely attached to the udder. Specialised flexible and elastic skin is covering the teats, the epidermis of which is thicker than in other
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areas of the body and is tightly attached to the underlying dermis. The teat is normally capable of withstanding minor trauma including suckling by the calf and milking by hand or machine (Jackson, 1996). Any injury or infection involving the skin of the teat is a potential threat to its efficient function and to the welfare of the animal. The sources of injury may be barbed wire, sharp objects and damaged or inadequate fencing (Jackson, 1996). In dairy cows tramped teats are a common source of teat injury (Oltenacu et al., 1990). However, beef cows generally have smaller udders and shorter teats, which decreases the risk of teat trampling. Multiple lacerations of varying depth and length may be seen in the skin of the udder and teats. Deep horizontal cracks may occur as a result of excessive sucking or biting by older calves or even older animals (Jackson, 1996). The likelihood of cows developing mastitis is 50% higher in injured than in non-injured cows (Modransky and Welker, 1993). Similar results were found by Pyorala et al. (1992) and Geishauser et al. (1999). Teat injury has been associated with high SCC in dairy cattle (Jorstad, et al., 1989), and numerous studies have indicated a positive relationship between milk SCC and udder disease (Coffey et al., 1986a; Coffey et al., 1986b; Emanuelson, 1988; Emanuelson et al., 1988; Shook, 1989; Weller et al., 1992). Single or multiple papillomas caused by bovine papilloma viruses can be found on the teats of heifers and cows (Meischke, 1979; Olson et al., 1982). Lindholm et al. (1984) found that Hereford cows and their crosses had a high incidence of papillomas and showed the highest numbers of warts, but it was not possible to make a realistic comparison with other breeds because of the small numbers of animals involved in the study. This study was done with seven breeds and their crosses. Pobric et al. (1990) working with cows of various breeds found a prevalence of papillomatosis of 8% (91 of 1124). William et al. (1992) reported that of 171 animals examined, 43 (25%) had papillomatosis, of which 56% were cows, 28% heifers and 12% calves, and the highest incidence of papillomatosis was on the udder and navel. Wadhwa et al. (1996) reported a prevalence of 83% (19 of 23). Nooruddin et al. (1997) reported a prevalence of 16% (73 of 459) for crossbred and pure bred dairy cows; they also found a significant higher prevalence of papillomatosis in small herds compared to large herds. These data show that teat traits have an influence on mastitis, but the effect on milk yield is apparently not as significant as for intramammary infections.

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MILK UREA CONCENTRATION

Milk urea reflects the balance between dietary carbohydrate and protein and therefore serve as a feed efficiency indicator (Kirchgessner et al., 1986; Spohr and Wiesner, 1991; Roth et al., 1996; Lyatuu and Eastridge, 1998). The main theory behind this concept is that urea concentrations in blood and milk can provide information on nitrogen losses following absorption of ammonia from the gut, particularly the rumen (Oltner et al., 1985). The urea in milk arises primarily from massive transfer of urea from the blood (Roseler et al., 1993). Several authors have reported in dairy cattle a close correlation (0.770.98) between concentrations of urea in blood and milk (Bakanov et al., 1976; Oltner and Wiktorsson, 1983; Refsdal, 1983; Gajdosik and Szaboova, 1984; DePeters and Ferguson, 1992; Sato et al., 1992). The equilibration between blood urea and milk urea is relatively rapid; urea is equilibrated with serum with a time lag of 1 to 2 hours (Gustafsson and Palmquist, 1993). Due to its rapid and passive diffusion through cell membranes (DePeterson and Cant, 1992), equilibration may be explained by diffusion of urea along the mammary ducts and through the mucosa in the alveoli (Gustafsson and Palmquist, 1993). Milk has the advantage of being easy to obtain from the lactating cow and the stress of blood sampling is avoided (Oltner and Wiktorsson, 1983). In experimental studies a variation of 2.1 to 9.1 mmol/l in urea concentration has been reported in dairy cattle (Oltner et al., 1985; Refstal et al., 1985; Gustafsson and Carlsson 1993; Carlsson and Pehrson, 1994; Eicher et al., 1999a;). In beef cattle it has been reported a variation of 0.8 to 10.1 mmol/l (Sinclair, et al., 1994; Ponter et al., 1997; Knaus et al., 1999; Ponsart, et al., 1999). Carlsson and Perhson (1994) suggested that a milk urea concentration of 4.0-5.5 mmol/l could be considered as normal. This is supported by Gustafsson and Carlsson (1993) who observed a range from 4.5 to 5.0 mmol/l in milk urea concentration from cows with optimal reproductive efficiency. It has been reported that either low or high milk urea concentrations are associated with low reproductive efficiency (Ropstad and Refsdal, 1987; Ferguson et al., 1988; Canfield et al., 1990; Pehrson et al, 1992). Other authors have reported milk urea values in mg/dl and they have found a variation of 5.6 to 34 mg/dl in dairy cattle (Kirchgessner et al., 1988; Pestevsek et al., 1990; Roseler et al., 1993; Krober et al., 1999; Ikuta et al., 2000b) and a variation of 3.7 to 31.3 mg/dl in beef cows (Mathison et al., 1981; Kushibiki et al., 1991; Hess et al., 1998; Huston, et al., 1999; Lima et al., 1999). A range from 18 to 23 mg/dl has been reported as normal (Kirchgessner et al., 1988). Paulicks (1992) suggested that when protein and energy supply of dairy cows is balanced with requirements, milk urea concentrations are between 15 and 25 mg/dl. This variation in milk urea concentration is due to the influence of nutritional and nonnutritional factors (Carlsson, 1994). NUTRITIONAL FACTORS AFFECTING MILK UREA CONCENTRATION
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Effect of protein Voluntary dry-matter intake and composition of the forages of the ration can be considered responsible for the base level of milk urea concentration. In dairy cattle the concentrates are usually distributed according to the daily milk yield and are mostly responsible for the individual variations of milk urea (Eicher et al., 1999b). Urea concentration in blood and milk are positively correlated with the intake of dietary protein; when protein intake exceeds requirements, the levels of ammonia will increase in the rumen and therefore in blood and milk, whereas a deficiency in protein should result in low urea concentration (Carlsson, 1994). This relationship has been found in beef cattle (Fishwick et al., 1974; Weisenburger and Mathison, 1977; Hennessy et al., 1995; Lima et al., 1999) and in dairy cows ( Kreuzer et al., 1991; Homolka and Vencl, 1993; Mackle et al., 1996; Sutton et al., 1997; Ikuta et al., 2000b).

Importance of balance between protein and energy Although protein intake affect urea level in blood and milk; urea concentrations are particularly affected by the balance between energy and protein in the diet (Ide et al., 1966; Payne et al., 1970; Oltner and Wiktorsson, 1983; Klein et al., 1987; Hof et al., 1997). The relation between protein and energy in the diet has a greater influence on milk urea concentration than has the absolute amount of feed of a certain composition that is consumed by the cow (Oltner and Wiktorsson, 1983; Oltner et al., 1985; Kirchgessner and Windisch, 1989; Ropstad et al., 1989). Some recent studies have shown that if protein is according to or over the requeriments of the cow and the energy is low, the urea concentrations are high, but if the protein is according to the requirements and there is a surplus of energy, decreases in urea concentration will be found (Carlsson and Pehrson, 1994; Kampl and Martincic, 1995; Carlsson et al., 1995; Steinwidder et al., 1998; Riha and Hanus, 1999). The reason for these changes is that the rumen microorganisms require energy to synthesise protein from ammonia; too little energy allows more ammonia to be converted into urea and excess energy allows the microorganisms to synthesise more protein and deplete the rumen of ammonia (Farries, 1982; Carlsson, 1994) In support of this a positive correlation between rumen ammonia and blood and milk urea concentrations have been reported (Foldager and Huber, 1979; Folman et al., 1981; Hammond , 1983; Ha and Kennelly, 1984; Kaim et al., 1987). Hoffman and Steinhfel (1990) found that dairy cows being fed more than their requirement of both energy and protein had urea concentrations lower than cows being fed according to their requirements of both energy and protein, they attributed this decrease to the more efficient utilisation of the dietary protein for the synthesis of microbial protein. However Carlsson and Pershon (1994) found conflicting results, reporting that when cows fed with a diet balanced according to their requeriments for
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energy and protein were fed with extra energy and protein, urea concentrations were increased. Carlsson and Pershon (1994) suggested that 1 possible reason for this discrepancy might be the much higher milk yields of the cows used in their experiment, as it is possible that ruminal flora of intensively fed cows are less able to use aditional energy for the synthesis of microbial protein.

Effect of rumen undegradable protein (RUP) Carlsson (1994) suggested that a diet containing a high proportion of ruminally undegradable protein (RUP) should result in lower concentrations of ammonia in the rumen and, as a result, lower concentrations of urea in blood and milk and vice versa. The same author mentioned that in his opinion, Ropstad et al. (1989) are in accordance with this supposition, because they found a strong positive correlation between the protein balance in rumen (the balance in the rumen between degraded feed protein and nitrogen used in the microbial protein synthesis) and milk urea concentration. However, Ropstad et al. (1989) in their study used two mixes (low and high protein) that were made to provide similar amounts of RUP. Therefore, with the same levels of RUP in the diet, it is not possible to measure the effect of RUP on urea concentration. Studies done in beef cattle (Hess et al., 1998) and in dairy cattle (Higginbotham et al., 1989) have reported that urea concentrations were increased in cows receiving additional protein, but no differences due to degradability were detected. Roseler et al. (1993) found that the intake of undegradable protein elevated plasma and milk urea concentrations to a similar extend as intake of degradable protein. Similar results have been found also in dairy cows by other authors (Roseler et al., 1990; Rodriguez et al., 1997b; Wu et al., 1997). In this aspect Roseler et al. (1993) suggested that the ability to detect subtle imbalances in degradable intake protein or undegradable intake protein by milk urea probably is masked by the intake of dietary crude protein and the balance between protein and energy, and that ruminal and postruminal excesses of nitrogen are eliminated from the body in the same process of hepatic urea synthesis; therefore excess protein, whether degradable or undegradable will elevate urea concentrations. However it has been reported that milk urea concentration was increased by effect of RUP in the diet in beef (Wiley et al., 1991) and in dairy cows (Figueroa et al., 1992; Rodriguez et al., 1997a).

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NON-NUTRITIONAL FACTORS AFFECTING MILK UREA CONCENTRATION

Non-nutriotional factors sometimes can affect significantly the urea concentration, therefore is important to consider them in milk urea assessments. Diurnal variation, season, stage of lactation, age, udder health, body weight, milk yield, technical aspects and others factors play an important role in the milk urea concentration.

Diurnal variation Diurnal variations of milk urea are influenced by feeding strategy (Miettinen and Juvonen, 1990; Gustafsson and Palmquist, 1993, Hess et al., 1998; Eicher et al., 1999a), time of sampling (Miettinen and Juvonen, 1990; Gustafsson and Palmquist, 1993; Eicher et al., 1999a) and the quality of protein (Miettinen and Juvonen, 1990). Carlsson and Bergstrm (1994) reported that there was a significant diurnal variation in milk urea and the highest values were found 3-5 hours after the beginning of morning feeding and the lowest values during late night in dairy cattle.. These authors observed that within 1 hour after the start of morning feeding the urea values had increased significantly. Ikuta et al. (2000a) reported also that milk urea concentrations increased after feed intake and reached a peak from about 3 to 5 hours after feeding. Ponsart et al. (1999) working with beef cattle observed the highest values of urea concentration 3-4 hours after the morning feeding. Similar results have been reported in other investigations in dairy cows (Tomas and Kelly, 1976; Gustafsson and Palmquist, 1993). Some researchers have found that milk urea concentrations were highest in the morning (Gustafsson et al., 1993; Eicher et al., 1997a; Eicher et al., 1999a) while others have reported that milk urea is higher in the evening samplig (Miettinen and Juvonen, 1990; Broderick and Clayton, 1997). Miettinen and Juvonen (1990) attributed their results to the time sampling and feeding, because milk were taken 1-3 hours after the onset of feeding when urea concentrations are still increasing and the feeding seem to influence the urea level in the morning but not in the evening; this may due to the long fasting period (12 hours) during the preceding night. Coggins and Fileld (1976) working with beef cows also found diurnal variations in urea concentration. They suggested that this variation was due to feeding time. However Oltner and Wiktorsson (1983) found no significant differences in milk urea levels between morning and afternoon sampling and the day-to-day variations were small. Since urea is mostly excreted by the kidneys, the blood urea and consequently milk urea, is increased in renal failure. However in healthy animals the urea levels is primarily affected by their feeding regime (Miettinen and Juvonen, 1990). Eicher et al. (1999a) suggested that the difference of the diurnal patterns could be the result of the
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different feeding practices, possibly the concentrate feeding frequency. Miettinen and Juvonen (1990) observed that the circadian rhythm of milk urea was closely related to the rhythmic feeding and fasting. According to that they mentioned a peak of urea appearing 2-4 hours after feeding in twice daily feeding (Tomas and Kelly, 1976; Gustafsson and Palmquist, 1993), whereas at higher feeding frequencies, a diurnal pattern is absent (Tomas and Kelly, 1976; Roth et al., 1996). In some studies it has been concluded that under comparable feeding practices, a milk sample taken 2 to 4 hours after first concentrate feeding, adequately reflects the maximum urea levels; also in some cases it could be useful to analyze both morning and afternoon milk to get an indication of the dynamic changes of circulating urea due to inadequate concentrate feeding (Eicher et al., 1999a) and if urea is to be used as an indicator of nutritional status, diurnal variations of milk urea should be considered; time sampling versus time of feeding is crucial (Gustafsson and Palmquist, 1993).

Season Investigations of dairy cattle have found that milk urea concentrations were significantly higher during the grazing season than during the indoor season (Hoffman and Steinhfel, 1990; Carlsson and Pehrson, 1993; Carlsson et al., 1995; Roth et al., 1996). Carlsson et al (1995) suggested that the difference was probably due to the high content of easily digestible crude protein in grass, relative to its energy content. Grazing animals often have a high milk urea concentrations indicating an excess formation of ammonia in the gut (Oltner et al., 1985). According to this, Kushibiki et al (1991) reported in beef cows that urea concentrations increased rapidly at the onset of grazing and remained high (25-30 mg/dl). Carlsson et al. (1995) reported that the changes in the concentration of urea in milk were much more regular throughout lactation when the cows where housed than when they were at grass. These authors suggested that it was probably because any changes in the nutritional content of the cows diet were more regular and controlled during the housing period.

Stage of lactation The effect of stage of lactation has been investigated in some studies. Milk urea concentration varies largely according to the stage of lactation (Faverdin and Verite, 1998). Some researchers have found that urea concentration is lower during the first month of lactation than later in beef cattle (Hess et al., 1998) and in dairy cows (Bruckendal et al., 1980; Ng-Kwai-Hang et al., 1985; DePeters and Cant, 1992; Emanuelson et al., 1993; Carlsson et al., 1995). Carlsson et al. (1995) suggested that the low urea concentration in this period is because in very early lactation the output of nutrients in milk generally exceeds the feed intake ability of the cow, which might induce suboptimal function of the ruminal flora and that another factor may be the high risk of metabolic disturbances during the first month of lactation. Ikuta et al. (2000b) observed the opposite situation and reported lower levels of milk urea in late-lactation than in mid- and early lactation in dairy cows. However in some studies it has been
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reported that stage of lactation did not significantly influence milk urea concentration (Hoffman and Steinhfel, 1990; Schepers and Meijer, 1998).

Age In dairy cattle it has been reported that age did not contribute significantly to the variation of urea levels in milk (Ropstad et al., 1989; Canfield et al., 1990; Schepers and Meijer, 1998). Carlsson et al. (1995) observed in their study that a preliminary analysis suggested that the multiparous cows had slightly higher milk urea concentrations than the primiparous cows, but when other factors where taken into account, this difference disappeared. The influence of age probably reflects differences in management and feeding practices for primiparous cows. This indicates the necessity to differentiate between first calf heifers and pluriparous cows in milk urea concentration evaluation (Eicher et al., 1999b). However, Oltner et al. (1985) reported that primiparous cows had a lower milk urea concentration than older animals. These authors suggested that this could be due to the fact that primiparous cows have a pronounced drive to grow and may therefore utilise amino acids more effectively and consequently this will reduce deamination and urea formation in the liver. Carlsson et al. (1995) suggested that effect of age found by Oltner and co-workers (1985) could be due to the small numbers of animals used in the experiment (49 cows of the Swedish Red and White breed), and concluded that it seems reasonable that milk urea concentration can be evaluated adequately without considering the age of the cows.

Mastitis and somatic cell count (SCC) Some studies have shown that SCC did not affect milk urea concentrations (Eicher et al., 1997a; Eicher et al., 1999a, Eicher et al., 1999b). However, a high SCC has been associated with high (Ng-Kwai-Hang et al., 1985) and low (Licata, 1985) milk urea values. Mert et al. (1992) comparing milk samples from healthy cows and from cows with subclinical mastitis reported significant differences in urea concentrations. Gutjahr et al. (1997) comparing milk from healthy quarters and from quarters with acute mastitis, chronic mastitis and subclinical mastitis observed that urea concentration in the milk from udder quarters with symptoms of mastitis was significantly lower than in the milk from healthy quarters of the corresponding udders. Eicher et al. (1999a) concluded that a milk sample from one clinically healthy quarter can be considered representative of the momentary urea concentration, but in the case of clinical mastitis, the more severely impaired permeability of the epithelium could produce a noticeable alteration. Therefore the urea concentration should be measure only in milk samples from healthy quarters.

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Body weight An inverse relation between body weight and milk urea has been reported by some authors (Oltner et al., 1983; Oltner et al., 1985), whereas others have reported that the body weight did not contribute significantly to the variation of urea levels in milk (Ropstad et al., 1989). The negative relation between milk urea concentration and live weight might be explained partly on the basis of a simple dilution. In a big cow the compartment for urea distribution is naturally larger than in a small cow, and if the same amount of urea is formed in the liver, concentrations in blood and milk will obviously be lower. With simple dilution the effect of weight on milk urea concentration will also depend on the absolute amount of urea to be distributed in the body. On diets producing much urea and consequently high milk urea concentrations , the effect of live weight on milk urea will be greater on a concentration basis (Oltner et al., 1985).

Milk yield It has been reported higher milk urea concentrations with increasing milk yield (Oltner et al., 1985; Carlsson et al., 1995; Eicher et al., 1999b), but this effect has been suggested to be due to feeding (Eicher et al., 1999b). As previously discussed milk urea concentration increase along with an increase in the protein/energy ratio in the diet (Oltner and Wiktorsson, 1983). According to standard feeding recommendations, more protein per unit of energy is needed for milk production than for maintenance, which will lead to an increase in the protein/energy ratio of the diet as milk production increases. However, Ropstad et al. (1989) reported that milk yield did not contribute significantly to the variation of urea levels in milk.

Technical aspects Sample conservation has influence on milk urea concentration. It has been reported that milk samples conserved in refrigeration at 4 oC with or without preservative (Bronopol or Sodium azide) remain unchanged for up to 1 week on their urea levels (Miettinen and Juvonen, 1990; Carlsson and Bergstrm, 1994; Eicher et al., 1997a; Eicher et al., 1999a). Samples in refrigeration with preservative could remain unchanged on its urea levels for up to 11 (Sodium azide) (Miettinen and Juvonen, 1990) or 17 days (Bronopol) (Carlsson and Bergstrm, 1994) and preservation by freezing at 20 oC increase storage ability without affecting the result of the analysis of milk urea for up to 1 month (Carlsson and Bergstrm, 1994; Eicher et al., 1997a; Eicher et al., 1999a). Storage at room temperature (20 oC) should be avoided (Eicher et al., 1999a) because in samples stored at room temperature milk urea decreased significantly at 24 hours after sampling (Ikuta et al., 2000a) and about 50% in two days (Miettinen and Juvonen, 1990).. Eicher et al. (1999a) reported that the agreement with the urea levels were fairly

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good between analysis in fresh and refrigerated samples but poor between analysis in fresh and frozen samples. They concluded that analyses of milk urea should be performed preferably in fresh samples or after only a short period of conservation. Some works have studied the precision of the method to measure urea concentrations. Ikuta et al. (2000a) found significant correlations between milk urea concentration and DRICHEM (dry chemical autoanalyzer), absorbance, differential pH and infrared spectroscopy, but no significant correlation was observed with the quick dipstick method. Other authors have reported that the flow injection analysis (FIA) technique (Carlsson and Bergstrm, 1994) and the kinetic enzymatic method(Eicher et al., 1999a) for the determination of milk urea concentrations had a high capacity.

Other factors affecting milk urea concentration There have not been found any breed effects on milk urea between dairy breeds in some studies (Carlsson et al., 1995; Chaiyabutr et al., 1999). However Rodriguez et al. (1997a) observed that milk urea was higher in Holstein than in Jersey cows. Wolfschoon-Pombo (1981) reported that the milk urea concentrations of two German breeds of cows were different, although it was accepted that the difference might have been due to the different feeding systems used. Hunter and Siebert (1985) comparing Bos taurus (Hereford) and Bos indicus (Brahman) cattle reported that urea concentrations were significantly (P<0.005) higher in Brahman than in Hereford cows. In dairy cows no significant differences in the milk urea concentration of different udder quarters have been found (Gustafsson and Palmquist, 1993; Carlsson and Bergstrm, 1994; Eicher et al., 1997a; Eicher et al., 1999a). However Ikuta et al. (2000a) reported that in the right front quarter of the udder, a significant difference was observed in milk urea concentration between pre-milking and post-milking. A physiological explanation for this, was not found in this review.

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EFFECTS OF NURSING ON UDDER HEALTH STATUS

It has been found that suckling has an effect on udder health status. Joshi et al. (1998) working with cows (n=18) and buffaloes (n=56) found a high incidence of clinical mastitis (69.0%) in animals without a nursing calf. Krohn et al. (1990) studying the effect of cow-calf separation on milk production and udder health reported that dairy cows that were separated immediately from their calves at calving had higher percentage (50%) of clinical mastitis compared to cows that were allowed to suckled their calves for five days after calving (13%), attributing this to the presence of the nursing calf. In an experimental group of 140 multi- and primiparous cows that suckled their calves for 10 days after birth and in a control group of 130 cows that were separated from their calves immediately after birth, it was reported that the incidence of subclinical and latent mastitis in the experimental group was 1.3-2.5% lower than in the control group (Twardon, 1988). In another studyl where the cows were allowed to nurse their calves for one hour or during 2, 4, 6 and 10 days, the frequency and duration of mastitis during two months after calving was reported be lower in cows that were allowed to suckle their calves for 6 or 10 days (Tsolov et al., 1989).

RESTRICTED SUCKLING VERSUS ARTIFICIAL REARING

Most of the research where the effect of suckling on udder health has been investigated has been done comparing restricted suckling (RS) with artificial (AR). Although Costa et al. (1997) found a higher incidence in cows where the calves were allowed to have access to the udder after completion of machine milking (RS) than in cows milked in the absence of calves (AR), most of the studies have found opposite results. In a study done in Vietnam by Mai Van et al. (1997) a mastitis incidence of 25% for AR and 0% for RS was reprted in F1 (Holstein x Local) cows. Alvarez et al. (1980) in crossbred cows (Holstein x Zebu and Brown Swiss x Zebu) observed that the incidence of mastitis was reduced by stimulus of the calf (RS). Ugarte and Preston (1972) working with Holstein and F1 (Holstein x Brahman) cows reported that the incidence of clinical and subclinical mastitis was lower in cows with RS than with AR. Rigby et al. (1976) in Holstein x Zebu heifers inoculated with S. aureus, reported that heifers nursing their calves were free of S. aureus after 6 days of the inoculation. Magana et al. (1996) working with Holstein cows in the Mexican tropics found that the health status of the mammary gland was better in cows that were suckling their calves with residual milk (RS); reporting that the incidence of mastitis was significantly lower in this group compared with cows with AR, i.e. machine milking only (40% vs 63%).

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Walsh (1974) studying the effect of RS and AR through lactation on production and health of dairy cows, found that the incidence of infected quarters was higher in AR (29.2%) than in RS (2.1%) in early lactation. Ugarte and Preston (1975) reported that the lower incidence of mastitis in RS compared with AR, was observed during the first ten weeks after calving and then, from the 11th week until the end of lactation there were no significant differences between treatments (RS and AR). Other authors have also reported lower mastitis incidence in RS than in AR (Veitia and Simon 1972; Rayner et al., 1977; Knowles and Edwards, 1983; Preston 1983; Mejia et al., 1998). It has been suggested that the machine milking is important in the genesis of new infections (OShea and Meaney, 1979). This is supported by Hamann and Stanitzke (1990) who found that the teat tissue reactions (teat shape and temperature) were greater after machine milking than after calf nursing. These authors assumed that the suckling allowed a better blood flow and by contrast machine milking may cause accumulation of fluid (blood and lymph) in the region of the teat which can be diagnosed by palpation as oedema. The risk of new mastitis infection is closely related to the magnitude of these changes (Hamann, 1991). However Rasmussen and Larsen (1998) reported that teat skin condition became worse after suckling when compared with machine milking. Still, these authors also reported that the number of bacteria isolated from the teat skin was lower in suckling than in machine milking. In studies resulting in reports on the beneficial effects on mastitis incidence using RS rather than AR, the time the calves are allowed to be nursed by their dams has been around 15 to 60 minutes, in general approximately 30 minutes. Chaverri (1981) found in Brahman cross cows that the percentage of quarters infected with mastitis was lower in cows that were allowed to nurse their calves for 1 hour (0.8%) compared to cows that were nursing their calves for 6 hours (2.2%) after milking. Skei and Waage (1998) testing nursing as a method of treatment of clinical mastitis as an alternative to drug therapy, reported that 10 (63%) of 16 cows with acute or subacute mastitis were cured by allowing them to nursed foster calves for one week. The lower clinical and subclinical incidence of mastitis found in RS has been attributed to the mechanical effect of suckling (Rigby et al., 1976; Ugarte, 1989), the better emptying of the udder (Rigby et al., 1976; Ugarte, 1989; Mejia et al., 1997) and also to the cleaning (bacteriostatic) effect of the saliva (Rigby et al., 1976; Ugarte 1989; Mejia et al., 1997). Twomey and Arnold (1975) suggested that the effect of the saliva could be due to the high level of the cationic polypeptide ubiquitin found in the calves saliva. Simpson et al. (1995) suggested that cows nursing their calves frequently may decrease the impact of potential mastitis infections via removal of bacteria from the mammary glands at relatively frequent intervals.

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In contrast other researchers have not observed any differences on the incidence of mastitis between these management systems (Fulkerson et al., 1978; Thomas et al., 1981; Bar-Peled et al., 1995). Ryle and Orskov (1990) suggested that this could be due to the generally low incidence of mastitis found by the researchers. In fact Fulkerson et al. (1978) attributed their results to a good hygiene of the milking operation kept during the study.

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CONCLUSION

In scientific studies it is important to take into consideration all factors that may have a substantial effect on the milk trait studied e.g. milk yield through lactation. Hence when milk traits are being analyzed, it is important to make comparisons with studies including all relevant factors listed in this review. It is also important to remember that some of the contradictory results found in the literature may be due to the circumstances under which such works were carried out, e.g. small sample sizes. The main contribution of this review is the discussion of some factors that have so far not been thoroughly investigated in beef cows (e.g. milk fraction), which may be used in future investigations in order to get more precise results.

CONCLUSION En estudios cientficos es importante tomar en consideracin todos los factores que pueden tener un considerable efecto sobre la caracterstica de la leche estudiada e.g. produccin de leche a travz de la lactacin. Por lo tanto cuando caractersticas de la leche estan siendo analizadas, es importante hacer comparaciones con estudios que incluyen todos los factores relevantes enlistados en esta revisin. Tambin es importante recordar que algunos de los resultados contradictorios encontrados en la literatura pueden ser debidos a las circunstancias bajo las cuales tales trabajos fueron llevados a cabo, e.g. pequeno tamano de muestra. La principal contribucin de esta revisin es la discusin de algunos factores que hasta ahora no han sido minuciosamente investigados en vacas de carne (e.g. fraccin de la leche), los cuales pueden ser usados en futuras investigaciones para obtener resultados ms precisos.

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ACKNOWLEDGEMENTS

Tack s vldigt mycket fr den otroligt trevliga tiden hr i Sverige. Jag har inte bara lrt mig mer om forskning utan ocks mycket om socialt beteende hos svenskar. Ni r ett vldigt trevligt och vnligt folk. Jag kommer att sakna era kaffe raster, era punsch luncher och ert lttl. Im very grateful to all the people and staff at the deparment, but I would like to express my sincere thanks to: Dr. Lotta Berg and Dr. Lena Lidfors for supervision and for their constructive comments on the writing of this manuscript, and for their frienship. PhD student Jens Jung and Dr. Lotta Berg for giving me the opportunity to come to Sweden. PhD student Karin Schtz for the wonderful time in midsummer at Vstervik and for introducing me to the innebandys world. Assistant researcher Anette Wichman for gaving me skating classes and for teaching me that life is better with nemas problemas. PhD student Margret Wlbers-Mindermann and Engineer Matthias Brandner for the nice barbecues in summer and for the wonderful time at Germany. PhD student Jenny Loberg for teaching me some Swedish and for her frienship. Mexico is waiting for you!!. PhD student Jenny Yngvesson for her delicious mushrooms and for being so nice with me and also Kalle for showing me the strangest crawling system that I have ever seen in my life. PhD student Karin Pettersson for the nice walk at mountain (not a hill!!) Kinnekulle. The diploma will be very useful in my curriculum vitae!! . MSc. Eva Persson for showing me beautiful places in Sweden and for her frienship. Technician Gunilla Jacobsson for her great help in computer questions. My girlfriend Dr. Tina McAdie for checking my spanienglish , for being herself, for making my stay in Sweden the best time of my life and for the most important, for love me. Staff at the library of the Faculty of Veterinary Medicine for helping me to get the literature used on this review.
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British Society of Animal Science for gave permission to publish the graph on page 30. International Fundation for Science (IFS) for funding my travel and subsistence in Sweden, as a part of the cooperation with the FOOD21 research programme.

Tack allihopa, jag hoppas att vi ses igen. Ni har en vn i Mexiko, glm aldrig det!!!!!!.

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REFERENCES

Aharoni, Y., Brosh, A. and Ezra, E., 1999. Effects of head load and photoperiod on milk yield and composition in three dairy herds in Israel. Anim. Sci., 69: 37-47. Alvarez, F.J., Saucedo, G., Arriaga, A. and Preston, T.R., 1980. Effect on milk production and calf performance of milking cross bred European/Zebu cattle in the absence or presence of the calf, and of rearing their calves artificially. Trop. Anim. Prod., 5: 25-37. Ashes, J.R., Gulati, S.K. and Scott, T.W., 1997. Potential to alter the content and composition of milk fat through nutrition. J. Dairy Sci., 80: 2204-2212. Bakanov, B.N., Ovsisher, B.R., Bondareva, N.I., Mamaev, V.A. and Filippov, VF., 1976. Nonprotein nitrogen and urea in blood and milk of cows in relation to ration composition (in Russian, English summary). Izvestiya Timiryazevskoi Selskokhozyaistvennoi Akademii., 4: 177-182. Barbary, E.A.S.A., Mahdy, A.E., Shafie, E.O.M. and Aly, H.M., 1999. Some factors affecting milk production and milk constituents and their relation to udder measurements in pure Friesian cows. Alexandria J. Agric. Res., 44: 17-35. Barkema, H.W., Van Der Schans, J., Schukken, Y.H., De Gee, A.L.W., Lam, T.J.G.M. and Benedictus, G., 1997. Effect of freezing on somatic cell count of quarter milk samples as determined by a fossomatic electronic cell counter. J. Dairy Sci., 80: 422-426. Bar-Peled, U., Maltz, E., Bruckental, I., Folman, Y., Kali, Y., Gacitua, H. and Lehrer, A.R., 1995. Relationship between frequent milking or suckling in early lactation and milk production of high producing dairy cows. J. Dairy Sci., 78: 2726-2736. Baruah, H.B., Sarker, A.B. and Roy, T.C., 1991. Studies on some measurements of udder and teat and their relation with milk yield and milk composition in Jersey local (F1) cows of Assam. Livest. Adv., 16: 26-34. Beal, W.E., Notter, D.R. and Akers, R.M., 1990. Techniques for estimation of milk yield in beef cows and relationships of milk yield to calf weight gain and postpartum reproduction. J. Anim. Sci., 68: 937-943. Belcher, C.G. and Frahm, R.R., 1979. Productivity of two-year-old crossbred cows producing three-breed cross calves. J. Anim. Sci., 49: 1195-1206. Belcher, C.G., Frahm, R.R., Belcher, D.R. and Bennett, E.N., 1979. Comparisons of lactational performance among varios four-year-old crossbred cow groups. Okla. Agric. Exp. Sta. Misc. Pub., MP-104: 132-136. Berger, J. and Francis, J., 1951. Mastitis in practice. 1. Epizootiology and bacteriology. Vet. Rec., 63: 283-293. Berning, L.M., Paape, M.J., Miller, R.H. and Hoffner, R.A., 1986. Variation in N-acetyl-beta-Dglucosaminidase (NAGase) activity and somatic cell count among various milk fractions. J. Dairy Sci. 69 (Suppl. 1): 246 (Abstr.). Berning, L.M., Paape, M.J., Miller, R.H. and LeDane, R.A., 1987. Variation in N-acetyl-beta-Dglucosaminidase activity and somatic cell count among various milk fractions. J. Dairy Sci., 70: 1054-1060. Bertrand, J.S., 1996. Influence of shipping container, preservative and breed on analysis of milk components of shipped samples. J. Dairy Sci., 79: 145-148. Blosser, T.H., 1979. Economic losses from and the national research program on mastitis in the United States. J. Dairy Sci., 62: 119-127. Bodoh, G.W., Battista, W.J., Schultze, L.H. and Johnston, R.P., 1976. Variation in somatic cell counts in dairy herd improvement milk samples. J. Dairy Sci., 59: 1119-1123. Boggs, D.L., Smith, E.F., Schalles, R.R., Brent, B.E., Corah, L.R. and Pruitt R.J., 1980. Effects of milk and forage intake on calf performance. J. Anim. Sci., 51: 550-553. Boucomont, D., 1985. Mastitis in nursing cows (in French, English summary). Bull. des Groupements Techniques Veterinaires., No. 3: 5-9. Bramley, A.J. and Dodd, F.H., 1984. Mastitis control: progress and prospects. J. Dairy Sci., 51: 481-512. Bremmer, D.R., Overton, T.R. and Clark, J.H., 1997. Production and composition of milk from Jersey cows administered bovine somatotropin and fed ruminally protected amino acids. J. Dairy Sci., 80: 1374-1380. 52

Brockman, R.P. and Laarveld, B., 1986. Hormonal regulation of metabolism in ruminants: a review. Livest. Prod. Sci., 14: 313-334. Broderick, G. A. and Clayton, M.K., 1997. A statistical evaluation of animal and nutritional factors influencing concentrations of milk urea nitrogen. J. Dairy Sci., 80: 2964-2971. Bruckental, I., Oldham, J.D. and Sutton, J.D., 1980. Glucose and urea kinetics in cows in early lactation. Br. J. Nutr., 44: 33-45. Butson, S. and Berg, R.T., 1984. Lactation performance of range beef and dairy-beef cows. Can. J. Anim. Sci., 64: 253-265. Canfield, R.W., Sniffen, C.J. and Butler, W.R., 1990. Effects of excess degradable protein on postpartum reproduction and energy balance in dairy cattle. J. Dairy Sci., 73: 2342-2349. Capuco, A.V., Bright, S.A., Pankey, J.W., Wood, D.L., Miller, R.H. and Bitman, J., 1992. Increased susceptibility to intrammary infection following removal of teat canal keratin. J. Dairy Sci., 75: 2126-2130. Carlsson, J., 1994. The value of the concentration of urea in milk as an indicator of the nutritional value of diets for dairy cows, and its relationships with milk production and fertility. Thesis, Swe. Univ. Agric. Sci., Skara, Sweden. Carlsson, J. and Bergstrm, J., 1994. The diurnal variation of urea in cows milk and how milk fat content, storage and preservation affects analysis by a flow injection technique. Acta Vet. Scand., 35: 67-77. Carlsson, J. and Pehrson, B., 1994. The influence of the dietary balance between energy and protein on milk urea concentration. Experimental trials assessed by two different protein evaluation systems. Acta Vet. Scand., 35: 193-205. Carlsson, J., Bergstrm, J. and Pehrson, B., 1995. Variations and breed, age, season, yield, stage of lactation and herd in the concentration of urea in bulk milk and individual cows milk. Acta Vet. Scand., 36: 245-254. Cartwright, T.C., Ellison, D.R., Thomas, R.C. and Fitzhugh, H.A., 1976. Production of F1 Angus-Jersey cows compared with Hereford cows. In: Beef cattle research in Texas.Tex. Agric. Exp. Sta., 2440. Chaiyabutr, N., Preuksagorn, S., Komolvanich, S. and Chanpongsang, S., 1999. Urea and allantoin in milk of crossbred Holstein cattle feeding on different types of roughage. Int. J. Anim. Sci., 14: 916. Chaudhary, A.P., Parmar, O.S. and Singh, K.P., 1998. Evaluation of milk constituents and quality under machine milking cows. Proc. 6th Wld. Cong. Genet. Appl. Livest. Prod., 25: 11-14. Chaverri, R.A.A., 1981. Effect of 3 methods of suckling on the growth of calves and the production and reproduction of their dams (in Spanish, English summary). Asoc. Latinoamericana Prod. Anim., 16: 30-31. Chenette, C.G. and Frahm, R.R., 1981. Yield and composition of milk from various two-breed cross cows. J. Anim. Sci., 52: 483-492. Choi, Y.L., Ahn, B.S., Seo, K.H., Kim, J.S. and Kim, N.S., 1999. Estimation of environmental effects and genetics parameters for SCS (somatic cell score) in Holstein cows (in Korean, English summary). Korean J. Anim. Sci., 41: 141-146. Chow, J.M., DePeters, E.J. and Baldwin, R.L., 1990. Effect of rumen-protected methionine and lysine on casein in milk when diets high in fat or concentrate are fed. J. Dairy Sci., 73: 1051-1061. Christensen, R.A., Lynch, G.L. and Clark, J.H., 1992a. Effect of protein degradability and intake on performance of lactating dairy cows. J. Dairy Sci., 75 (Suppl. 1): 201 (Abstr.). Christensen, R.A., Cameron, M.R., Klusmeyer, T.H. and Clark, J. H., 1992b. Effect of protein degradability and intake on nutrient utilization by dairy cows. J. Dairy Sci., 75 (Suppl. 1): 201 (Abstr.). Clarkson, A.R., 1975. Milk leucocyte counts and their significance in mastitis control. N. Z. Vet. J., 23: 284-286. Claypool, D.W., Pangborn, MC. and Adams, H.P., 1980. Effect of dietary protein on high-producing dairy cows in early lactation. J. Dairy Sci., 63: 833-837. Coffey, E.M., Vinson, W.E. and Pearson, R.E., 1986a. Somatic cell counts and infection rates for cows of varying somatic cell count in initial test of first lactation. J. Dairy Sci., 69: 552-555.

53

Coffey, E.M., Vinson, W.E. and Pearson, R.E., 1986b. Potential of somatic cell concentration in milk as a sire selection criterion to reduce mastitis in dairy cattle. J. Dairy Sci., 69: 2163-2172. Coggins, C.R.E. and Fileld, A.C., 1976. Diurnal variation in the chemical composition of plasma from lactating beef cows on three dietary energy intakes. J. Agric. Sci. (Camb.), 86: 595-602. Coleman, D.A. and Moss, B.R., 1989. Effects of several factors on quantification of fat, protein and somatic cells in milk. J. Dairy Sci., 72: 3295-3303. Concha, C., 1986. Cell types and their immunological functions in bovine mammary tissues and secretions-A review of the literature. Nord. Vet.-Med., 38: 257-272. Convey, E.M., Miller, L.S. and Tucker, H.A., 1971. Acute effect of 9 fluoroprednisolone acetate (9 FFA) on blood leucocytes and somatic cell content of milk. J. Dairy Sci., 54: 360-363. Coppock, C.E. and Wilks, D.L., 1991. Supplemental fat in high-energy rations for lactating cows: effects on intake, digestion, milk yield and composition. J. Anim. Sci., 69: 3826-3837. Costa, E.O. De, Carciofi, A.C., Melville, P.A., Prada, M.S., Pantano, T. and Ribeiro, A.R., 1997. Influence of the presence of calves during machine milking on the occurrence of mastitis (in Portuguese, English summary). Rev. Brasileira Med. Vet., 19: 19-22. Daley, D.R., McCuskey, A. and Bailey, C.M., 1986. Caracterization of milk constituents of Bos taurus and Bos indicus x Bos taurus breed types. Proc. 3rd Wld. Cong. Genet. Appl. Livest. Prod: 308313. Daniel, R.C.W., Barnum, D.A. and Rennie, J.C., 1966. Variation in modified California mastitis test scores in dairy cattle. J. Dairy Sci., 49: 1226-1229. Danilevskaya, N.T., Timchenko, A.G. and Shevchenko, D.I., 1972. Milk production and composition of Aberdeen-Angus cows (in Russian, English summary). Nauchnye Trudy Ukrainskaya Selskokhozyaistvennaya Akademiya., 73: 114-117, 135. Dash, R.C., Singh, B.P. and Misra, B.S., 1978. Inheritance studies on some of the milk constituents in Hariana cattle. Indian Vet. Med. J., 2: 141-145. Davies, D.T., Holt, C. and Christie, W.W., 1983. The composition of milk. In: Biochemistry of lactation. ed. Mephan, T.B., Elsevier Science Publisher B.V. .71-117. DePeters E.J. and Cant J.P., 1992. Nutritional factors influencing the nitrogen composition of bovine milk: a review. J. Dairy Sci., 75: 2043-2070. DePeters E.J. and Ferguson J.D., 1992. Nonprotein nitrogen and protein distribution in the milk of cows. J. Dairy Sci., 75: 3192-3209. Dohoo, I.R. and Leslie, K.E., 1991. Evaluation of changes in somatic cell counts as indicators of new intramammary infections. Prev. Vet. Med., 10: 225-237. Dohoo, I.R. and Meek A.H., 1982. Somatic cell counts in bovine milk. Can. Vet. J., 23: 119-125. Donovan, G.A., Risco, C.A. and Shearer, J.K., 1992. Assessment of the mammary system. Vet. Clin. North Am. Food Anim. Pract., 8: 361-373. Doornbos, D., Anderson, D. and Kress, D., 1981. Selecting for milk production in beef cattle. Large udders dont always mean more milk. Mont. Agric. Exp. Sta., Res. Rep. No. 180: 34-46. Duenas, M.I., Wettemann, R., Paape, M.J. and Cifrian, E., 1994. Effect of parity on intramammary infection and milk somatic cell counts (MSCC) in beef cows. J. Anim. Sci., 72 (Suppl.1): 254 (Abstr.). Eberhart, R.J., 1977. Coliform mastitis. J. Am. Vet. Med. Assoc., 170: 1160-1163. Eicher, R., Bouchard, E. and Bigras-Poulin, M., 1999b. Factors affecting milk urea nitrogen and protein concentrations in Quebec dairy cows. Prev. Vet. Med., 39: 53-63. Eicher, R., Bouchard, E. and Tremblay, A., 1997a. Factors that influence the results and interpretation of urea analysis in milk of dairy cows (in French, English). Epidemiologie et Sante Animale., 31-32: 05.B.12. Eicher, R., Bouchard, E. and Tremblay, A., 1999a. Cow level sampling factors affecting analysis and interpretation of milk urea concentrations in 2 dairy herds. Can. Vet. J., 40: 487-492. Emanuelson, M., Ahlin, K.- and Wiktorsson, H., 1993. Long-term feeding of rapeseed meal and full-fat rapeseed of double low cultivars to dairy cows. Livest. Prod. Sci., 33: 199-214. Emanuelson, U., 1988. Recording of production disseases in cattle and possibilities for genetic improvements: a review. Livest. Prod. Sci., 20: 89-106 Emanuelson, U., Danell, B. and Philipsson J., 1988. Genetic parameters for clinical mastitis, somatic cell counts, and milk production estimated by multipletrait restricted maximum likelihood. J. Dairy. Sci., 71: 467-476. 54

Emanuelson, U. and Funke, H., 1991. Effect of milk yield on relationship between bulk milk somatic cell count and prevalence of mastitis. J. Dairy Sci., 74: 2479-2483. Emery, R.S., 1988. Milk fat depression and the influence of diet on milk composition. Vet. Clin. North Am. Food Anim. Pract., 4: 289-305. Emery, R.S., 1991. Feeding for milk components. Large Anim. Vet., 46: 30-32. Ettala, E., 1976. Factors affecting the composition of milk. II. Effect of stage and number of lactation and some other external factors, and significance of choice of test days in estimation of protein production ability (in Finnish, English summary). Ann. Agric. Fenniae., 15: 196-213. Farries, E., 1982. Metabolic disorders and their effect on milk composition (in German, English summary). Proc. XIIth Wld. Cong. Dis. Cattle, Neth., 1: 539-546. Faverdin, P. and Verite, R., 1998. Use of milk urea concentration as an indicator of protein nutrition and nitrogen losses in dairy cows (in French, English summary). Rencontres Recherches Ruminants., 5: 209-212. Ferguson, J.D., Blanchard, T.L., Galligan, P.F., Hoshell, D.S. and Chalupa, W., 1988. Infertility of dairy cattle fed a high percentage of protein degradable in the rumen. J. Am. Vet. Med. Assoc., 192: 659662. Figueroa, M.R., Dawson, D.P., Batallas, C.E., Kim, DY., Arambel, M.J. and Walters, J.L., 1992. Nutritional and physiological effects of feeding two levels of rumen degradable and undegradable protein. J. Dairy Sci., 75 (Suppl. 1): 202 (Abstr.). Fishwick, G., Fraser, J., Hemingway, R.G., Parkins, J.J. and Ritchie, N.S., 1974. The voluntary intake and digestibility of oat straw by pregnant beef cows as influenced by urea and phosphorus supplementation contained in molassed sugar-beet pulp. J. Agric. Sci. (Camb.), 82: 427-432. Foldager, J. and Huber, J.T., 1979. Influence of protein percent and source on cows in early lactation. J. Dairy Sci., 62: 954-964. Folman, Y., Neumark, H., Kaim, M. and Kaufmann, W., 1981. Performance, rumen and blood metabolites in high-yielding cows fed varying protein percents and protected soybean. J. Dairy Sci., 64: 759-768. Fox, L.K. and Gay, J.M., 1993. Contagious mastitis. Vet. Clin. North Am. Food Anim. Pract., 9: 475-487. Francis, P., 1993. Origins, interpretations and uses of high milk somatic cell counts. In Pract. 15: 288290. Frank, N.A. and Pounden, W.D., 1961. Prevalence of bovine mastitis during various stages of lactation. J. Am. Vet. Med. Assoc., 138: 184-187. Frank, N.A., Pounden, W.D., Weaver, C.R. and Gilmore, L.O., 1959. Activity in milk of a mastitisproducing staphylococcus, a coliform organism and Streptococcus agalactiea. Am. J. Vet. Res., 20: 736-741. Franke, D.E. and Martin, S.E., 1983. Yield and composition of milk compared in beef cattle. Louisiana Agric., 26: 16-18. Fulkerson, W.J., Hooley, R.D. and Findlay, J.K., 1978. Improvement in milk production of first calf heifers by multiple suckling. Aust. J. Agric. Res., 29:351-357. Gajdosik, D. And Szaboova, E., 1984. Applicability of milk urea and acidity determinations in the assessment of protein nutrition of cows (in Slovakian, English summary). Veterinarni Medicina., 29: 329-336. Geishauser, T., Querengasser, K., Nitschke, M. and Sorbiraj, A., 1999. Milk yield, somatic cell counts, and risk of removal from the herd for dairy cows after covered teat canal injury. J. Dairy Sci., 82: 1482-1488. Gencurova, V., Hanus, O., Matous, E., Gabriel, B. and Kopecky, J., 1994. Should bronopol replace potassium dichromate in the preservation of milk samples in the Czech Republic? (in Czech, English summary). Vyzkum v Chovu Skotu., 36: 7-14. Ghosh, B. and Prasad, J., 1998. Milk yield and composition as influenced by udder measurements in Jersey x Red Sindhi crosses. Indian J. Anim. Prod. Man., 14: 23-25. Gill, M.S. and Holmes, C. W., 1978. Somatic cell counts, mastitis and milk production in dairy herds. N. Z. Dairy Sci. and Tech., 13: 157-161. Grega, T. and Szarek, J., 1985. Relationship of milking rate and teat dimensions to cell count in milk (in Polish, English summary). Zeszyty Naukowe Akademii Rolniczej w Krakowie., Zootechnika 23: 13-22. 55

Griinari, J.M., McGuire, M.A., Dwyer, D.A., Bauman, D.E. and Palmquist, D.L., 1997a. Role of insulin in the regulation of milk fat synthesis in dairy cows. J. Dairy Sci., 80: 1076-1084. Griinari, J.M., McGuire, M.A., Dwyer, D.A., Bauman, D.E., Barbano, D.M. and House, W.A.,1997b. The role of insulin in the regulation of milk protein synthesis in dairy cows. J. Dairy Sci., 80: 2361-2371. Grummer, R.R., 1991. Potential for modifying milk composition through feeding practices. Proc., CDR Cheese Res. Tech. Conf., 69-77. Guidry, A. J., Paape, M. J. and Pearson, R.E., 1975. Effects of estrus and exogenous estrogen on circulating neutrophils and milk somatic cell concentration, neutrophil phagocytosis, and occurrence of clinical mastitis in cows. Am. J. Vet. Res., 36: 1555-1560. Gupta, R., Singh, R.P. and Tomar, S.S., 1991. Udder and teat measurements and their association with milk production in Karan-Fries cows. Indian J. Anim. Res., 25: 23-28. Gustafsson, A.H. and Carlsson, J., 1993. Effects of silage quality, protein evaluation systems and milk urea content on milk yield and reproduction in dairy cows. Livest. Prod. Sci., 37: 91-105. Gustafsson, A.H. and Palmquist, D.L., 1993. Diurnal variation of rumen ammonia, serum urea and milk urea in dairy cows at high and low yields. J. Dairy Sci., 76: 475-484. Gutjahr, S., Schulz, J., Muniem, A. and Beck, K., 1997. Influence of udder health on urea content of cow milk samples (in German, English summary). Praktische Tierarzt., 78: 573-580. Ha J.K. and Kennelly, J.J., 1984. Effect of protein on nutrient digestion and milk production by Holstein cows. J. Dairy Sci., 67: 2302-2307. Haggard, D.L., Farnsworth, R. J. and Springer, J. A., 1983. Subclinical mastitis of beef cows. J. Am. Vet. Med. Assoc., 182: 604-606. Hamana, K., Motomura, Y., Yasuda, N., Kamimura, S. and Trenti, F., 1994. Bovine teat morphology and ultrasonic tomography related to milk quality and bacteria. Proc. 18th Wld. Buiat. Cong., 1: 377380. Hamann, J., 1991. Milking related teat tissue changes as a predisposing factor for mastitis. In: New insights into the pathogenesis of mastitis., ed. Burvenich, C., Van Messom, G.V. and Hill, A.W., Institute for hygiene, Dairy Research Center, Germany. 57-68. Hamann, J. and Gyodi, P., 1999. Electrical conductivity in fraction-collected quarter milk samples with low somatic cell counts (in German, English summary). Milchwissenschaft., 54: 487-491. Hamann, J., Gyodi, P. and Heeschen, W., 1991. The influence of varying preserving agents and storage times on somatic cell counting in milk using the Fossomatic-360 and Fossomatic-180 (in German, English summary). Kieler Milchwirtschaftliche Forschungsberichte., 43: 273-295. Hamann, J. and Stanitzke, U., 1990. Studies on pathogenesis of bovine mastitis by comparison of milking conditions as calf suckling, hand milking and machine milking: reactions of the teat tissue. Milchwissenschaft., 45: 632-637. Hammond, A.C., 1983. Effect of dietary protein level, ruminal protein solubility and time after feeding on plasma urea nitrogen to other ruminal ans plasma parameters. J. Anim. Sci., 57 (Suppl. 1): 435 (Abstr.). Hanus, O., Gencurova, V., Gabriel, B. and Zvackova, I., 1992. A comparison of the efficiency of a Milkofix preservative substance with traditional preservatives used to determine somatic cell counts in milk samples by a fluoro-opto-electronic method (in Czech, English summary). Veterinarni Medicina., 37: 91-99. Heeschen, W.H., Ubben, E.H., Gyodi, P. and Beer, P., 1993. Counting somatic cells in milk: comparative studies using fluorescence optical (Fossomatic 360) and flow cytometric (Somascope) measuring (in German, English summary). Kieler Milchwirtschaftliche Forschungsberichte., 45: 109-136.

56

Hennessy, D.W., Kohun, P.J., Williamson, P.J, Brown, D.A. and Nolan, J.V., 1995. The effect of nitrogen and protein supplementation on feed intake, growth and digestive function of steers with different Bos indicus, Bos taurus genotypes when fed a low quality grass hay. Aust. J. Agric. Res., 46: 1121-1136. Hess, B. W., Scholljegerdes, E.J., Coleman, S.A. and Williams, J. E., 1998. Supplemental protein plus ruminally protected methionine and lysine for primiparous beef cattle consuming annual rye hay. J. Anim. Sci., 76: 1767-1777. Hibbitt, K.G., 1983a. Immunity factors in mastitis. Proc. Br, Cattle Vet. Assoc.: 61-65. Hibbitt, K.G., 1983b. Defence mechanism of the bovine teat. Proc. Br. Cattle Vet. Assoc.: 235-240. Higginbotham, G.E., huber, J. T., Wallentine, M.V., Johnston, N.P. and Andrus, D., 1989. Influence of protein percentage and degradability on performance of lactating cows during moderate temperature. J. Dairy Sci., 72: 1818-1823. Hoden, A. and Coulon, J.B., 1991. Regulating the composition of milk: influence of nutritional factors on the quantity and the fat and protein content of milk (in French, English summary). Prod. Anim., 4: 361-367. Hof, G., Vervoorn, M.D., Lenaers, P.J. and Tamminga, S., 1997. Milk urea nitrogen as a tool to monitor the protein nutrition of dairy cows. J. Dairy Sci., 80: 3333-3340. Hoffmann, M. and Seinhfel, O., 1990. Possibilities and limitations for appraisal of energy and protein supply through monitoring of milk urea level (in German, English summary). Mh. Vet. Med., 45:223-227. Holdaway, R.J., Holmes, C.W. and Steffert, I.J., 1996. A comparison of indirect methods for diagnosis of subclinical intramammary infection in lactating dairy cows. Part 1. The effects of bacterial infection, stage of lactation and age of cow on eight parameters in foremilk from individual quarters, with an initial study of differences between milk fractions. Aust. J. Dairy Tech., 51: 6471. Homolka, P. and Vencl, B., 1993. Urea concentration in milk in relation to dietary crude protein to energy ratio (in Czech, English summary). Zivocisna Vyroba., 38: 529-535. Hoyer, M.J., Codd, R., Bishi, A.S., Pawandiwa, A. and Usenik, E.A., 1991. The prevalence of subclinical mastitis in beef herds in Zimbabwe. Zimbabwe Vet. J., 22: 1-11. Hunter, A.C. and Jeffrey, D.C., 1975. Subclinical mastitis in suckler cows. Vet. Rec., 96: 442-447. Hunter, R.A. and Siebert, B.D., 1985. Utilization of low-quality roughage by Bos taurus and Bos indicus cattle. 2. The effect of rumen-degradable nitrogen and sulphur on voluntary food intake and rumen characteristics. Br. J. Nutr., 53: 649-656. Huston, J.E., Lippke, H., Forbes, T.D.A., Holloway, J.W. and Machen, R.V., 1999. Effects of supplemental feeding interval on adult cows in western Texas. J Anim. Sci., 77: 3057-3067. Ibeawuchi, J.A. and Dangut, A.J., 1996. Influence of stage of lactation on milk constituents of Bunaji (Zebu) cattle in a hot humid tropical environment. Disc. Inn., 8: 249-256. Ide, Y., Shimbayashi, K. and Yonemura, T., 1966. Effect of dietary conditions upon serum- and milkurea nitrogen in cows. 1. Serum- and milk-urea nitrogen as affected by protein intake. Jnp. J. Vet. Sci., 28: 321-327. Ikuta, K., Kokamo, M., Sasakura, K. and Hakogi, E., 2000a. Methods of analysis for milk urea nitrogen and influence of sampling and storage on its concentrations (in Japanese, English summary). J. Jpn. Vet. Med. Assoc., 53: 285-288. Ikuta, K., Kokamo, M., Sasakura, K. and Hakogi, E., 2000b. Evaluation of nutritional condition on the basis of milk urea nitrogen and milk protein concentrations in dairy cows (in Japanese, English summary). J. Jpn. Vet. Med. Assoc., 53: 289-292. Jack, L.J.W., Capuco, A.V., Wood, D.L., Aschenbrenner, R.A., Bitman, J., Bright, S.A. and Miller, R.H., 1992. Protein composition of teat canal keratin collected from lactating cows before and after milking. J. Dairy Sci., 75 (Suppl. 1): 195 (Abstr.). Jackson, P., 1996. Skin diseases of the bovine udder and teat. In Pract., 18: 76-80. Jain, N.C., 1979. Common mammary pathogens and factors in infection and mastitis. J. Dairy Sci., 62: 128-134. Janicki, C., 1977. The effect of some factors on udder disease (in Polish, English summary). Roczniki Akademii Rolniczej w Poznaniu., 94: 109-118. Jeffery, H.B. and Berg, R.T., 1971. Evaluation of milk variables as measures of milk effect on preweaning performance of beef cattle. Can. J. Anim. Sci., 51: 21-30. 57

Jeffery, H.B., Berg, R.T. and Hardin, R.T., 1971. Factors affecting preweaning performance in beef cattle. Can. J. Anim. Sci., 51: 561-577. Jelec, S., 1990. Nutritional factors and milk composition with especial reference to milk fat (in SerboCroatian, English summary). Stocarstvo., 44: 199-208. Jenkins, T.G. and Ferrel, C.L., 1984. A note on lactation curves of crossbred cows. Anim. Prod., 39: 479482. Jenkins, T.C., 1993. Lipid metabolism in the rumen. J. Dairy Sci., 76: 3851-3863. Jenness, R., 1985. Biochemical and nutritional aspects of milk and colostrum. In: Lactation.Ed. Larson, B.L., Iowa State University Press, Ames, Iowa. 164-197. Jones, T.O., 1994. Bovine mastitis bacteriology: Problems and pitfalls. Cattle Pract., 2: 55-57. Jones-Ensley, J.M., Cecava, M.J. and Johnson, T.R., 1997. Effects of dietary supplementation on nutrient digestion and the milk yield of intensively grazed lactating dairy cows. J. Dairy Sci., 80: 32833292. Jorstad, A., Farver, T.B. and Riemann, H., 1989. Teat canal diameter and other cow factors with possible influence on somatic cell counts in cow milk. Acta Vet. Scand., 30: 239-245. Joshi, H.D., Joshi, B.R. and Shrestha, H.K., 1998. Epidemiological investigation on clinical mastitis in cattle and buffaloes in the western hills of Nepal. Vet. Rev. Kathmandu., 13: 12-15. Kaim, M., Neumark, H., Folman, Y. and Kaufmann, W., 1987. The Effect of two concentrations of dietary protein and of formaldehyde-treated soya-bean meal on the performance of high yielding dairy cows. Anim. Prod., 44: 333-345. Kampl, B. and Martincic, T., 1995. Relationship between milk urea and liver cytosolic phosphoenolpyruvate carboxykinase activity in cows (in Croation, English summary). Veterinarski Arhiv., 65: 57-62. Kartashova, V., Kasyanchuk, V. and Skorokhodov, Y., 1992. Factors influencing mastitis infection in cows (in Russian, English summary). Molochnoe i Myasnoe Skotovodstvo., No.1: 34-35. Kasari, T. and Gleason, D., 1996. Herd management practices that influence total beef calf production . Part II. Compend. Contin. Educ. Pract. Vet., 18: 1358-1366. Kehrli, M.E. and Shuster, D.E., 1994. Factors affecting milk somatic cells and their role in health of the bovine mammary gland. J. Dairy Sci., 77: 619-627. Kennedy, B.W., Sethar, M.S., Tong, A.K.W., Moxley, J.E. and Downey, B.R., 1982. Environmental factors influencing test-day somatic cell counts in Holsteins. J. Dairy Sci., 65: 275-280. Kennelly, J.J., 1996. The fatty acid composition of milk fat as influenced by feeding oilseeds. Anim. Feed Sci. Tech., 60: 137-152. Keys, J.E., Capuco, A.V., Akers, R.M. and Djiane, J., 1989. Comparative study of mammary gland development and diferentiation between beef and dairy heifers. Dom. Anim. Endoc., 6: 311-319. Kirchgessner, M., Kreuzer, M., and Roth-Maier, D.A., 1986. Milk urea and protein content to diagnose energy and protein malnutrition of dairy cows. Arch. Anim. Nutr., 36: 192-197. Kirchgessner, M., Paulicks, B.R. and Schwarz, F.J., 1988. Changes in urea contents of milk of cows on diets low or excessive in protein (in German, English summary). J. Anim. Physiol. Anim. Nutr., 59: 79-84. Kirchgessner, M. and Windisch, W., 1989. Milk urea content and allantoin excretion of cows during and after energy and protein depletion. 4. The influence of energy and protein depletion and its residual effects (in German, English summary). J. Anim. Physiol. Anim. Nutr., 62: 113-118. Kirk, J.H., 1984. Somatic cells in milk: Current concepts. Compend. Contin. Educ. Pract. Vet., 6: S237S242, S244. Kirkbride, C.A., 1977. Mastitis in beef cows. J. Am. Vet. Med. Assoc., 170: 1141-1142. Klein, B., Schmidt, B., Zucker, H., 1987. Serum urea determinations in dairy herds for evaluating protein and energy supply (in German, English summary). Tierarztliche Umschau., 42: 532, 535-539. Knaus, W., Luger, K., Zollitsch, W., Gufler, H., Gruber, L., Murauer, C. and Lettner, F., 1999. Effects of grass clover-pellets and whole plant maize-pellets on the feed intake and performance of dairy cows. Anim. Feed Sci. Tech., 81: 265-277. Knowles, R.T. and Edwards, M.D., 1983. A comparison of the effects of restricted suckling and artificial calf rearing systems on dam and calf performance. Malaysian Agric. J., 54: 1-9. Komaragiri, M. and Erdman, R., 1992. Effect of two levels of dietary protein on milk production and composition in early lactating dairy cows. J. Dairy Sci., 75 (Suppl. 1): 201 (Abstr.).

58

Kovacs, A.Z., 1997. Milk composition of beef cattle breeds in Hungary (in Hungarian, English summary). Allattenyesztes es Takarmanyozas., 46: 175-187. Kovacs, A. Z., Szirtes, S. and Szakaly, S., 1999. The ancient Hungarian Grey cattle. 2. Composition of milk of Hungarian Grey cattle (in Hungarian, English summary). Tejgazdasag., 59: 27-32. Kreuzer, M., Kirchgessner, M. and Blum, J.W., 1991. Concentrations of hormones and metabolites in blood plasma of cows during and subsequent to different crude protein supply (in German, English summary). J. Anim. Physiol. Anim. Nutr., 65: 11-20. Krober, T.F., Steingass, H., Funk, R. and Drochner, W., 1999. Effects of reduced crude protein supply on dry matter intake, digestibility, N-excretion and performance of lactating dairy cows (German, English summary). Zuchtungskunde., 71: 182-195. Krohn, C.C., Jonasen, B. and Munksgaard, L., 1990. Investigations on cow-calf relations. 2. The effect of 0 versus 5 days suckling on behaviour, milk production and udder health of cows in different stabling (in Danish, English summary). Beretning fra Statens Husdyrbrugsforsog., No. 678: 1-20. Kushibiki, S., Umemura, K. and Hayashi, T., 1991. Effect of fertilizer application on plasma mineral levels of grazing beef cows. 1. Changes in plasma mineral levels in relation to components of fertilizer (in Japanese, English summary). Bull. Nat. Grass. Res. Inst., No. 44: 63-70. Lalman, D.L., Williams, J.E., Hess, B.W., Thomas, M.G. and Keisler, D.H., 2000. Effect of dietary energy on milk production and metabolic hormones in thin, primiparous beef heifers. J. Anim. Sci., 78: 530-538. Laevens, H., Deluyker, H., Schukken, Y.H., De Meulemeester, L., Vandermeersch, R., De Muelenaere, E. and De Kruif, A., 1997. Influence of parity and stage of lactation on the somatic cell count in bacteriologically negative dairy cows. J. Dairy Sci., 80: 3219-3226. Lascelles, A.K., 1979. The immune system of the ruminant mammary gland and its role in the control of mastitis. J. Dairy Sci., 62: 154-160. Leigh, J.A., 1999. Streptococcus uberis: A permanent barrier to the control of bovine mastitis?. Vet. J., 157: 225-238. Licata, E., 1985. Subclinical mastitis and urea determination in cows milk (in Italian, English summary). Obiettivi e Documenti Veterinari., 6: 65-67. Lima, G.F. da C., Sollenberger, L.E., Kunkle, W.E., Moore, J.E. and Hammond, A.C., 1999. Nitrogen fertilization and supplementation effects on performance of beef heifers grazing limpograss. Crop Sci., 39: 1853-1858. Lindholm, I., Murphy, J., ONeil, B.W., Campo, M.S. and Jarret, W.F.H., 1984. Papillomas of the teats and udder of cattle and their causal viruses. Vet. Rec., 115: 574-577. Logan, E.F. and Gibson, T., 1975. Serum immunoglobulin levels in suckled beef calves. Vet. Rec., 97: 229-230. Lowman, B.G., Edwards, R.A., Somerville, S.H. and Jolly, G.M., 1979. The Effect of plane of nutrition in early lactation on the performance of beef cows. Anim. Prod., 29: 293-303. Lund, T., Miglior, F., Dekkers, J.C.M. and Burnside E.B., 1994. Genetic relationships between clinical mastitis, somatic cell count, and udder conformation in Danish Holsteins. Livest. Prod. Sci., 39: 243-251. Lyatuu, E.T. and Eastridge, M.L., 1998. Nutritional factors affecting milk production, milk composition, milk urea nitrogen and plasma urea nitrogen. Ohio Agric. Res. Develop. Cent. Spec. Circ., 163: 65-71. Mackle, T.R., Parr, C.R. and Bryant, A.M., 1996. Nitrogen fertiliser effects on milk yield and composition, pasture intake, nitrogen and energy partitioning, and rumen fermentation parameters of dairy cows in early lactation. N. Z. J. Agric. Res., 39: 341-356. Magana, J.M., Valencia, H.E. and Delgado, L.R., 1996. Effect of restricted suckling and artificial rearing on the performance of Holtein cows and their calves in the Subhumid tropic region of Mexico (in Spanish, English summary). Vet. Mx., 24: 271-277. Mai Van, S., Preston, T.R. and Le Viet, L., 1997. Effects of restricted suckling versus artificial rearing on performance and fertility of crossbred F1 (Holstein Friesian x Local) cows and calves in Vietnam. Livest. Res. Rural Dev., 9: 1-7 Makarechian, M.H. and Paputungan, U., 1998. Effects of dams weight, body condition and udder scores on calf birth weight and preweaning growth rate in beef cattle. J. Dairy Sci., 81 (Suppl. 1): 271 (Abstr.).

59

Mathison, G.W., Hardin, R.T. and Beck, B.E., 1981. Supplemental protein, magnesium and selenium plus vitamin E for beef cows fed straw diets in winter. Can. J. Anim. Sci., 61: 375-392. McDonald, J.S., 1968. Radiographic method for anatomic study of the teat canal. Changes with lactation age. Am. J. Vet. Res., 29: 1207-1210 McDonald, J.S., 1977. Streptococcal and Staphylococcal mastitis. J. Am. Vet. Med. Assoc., 170: 11571159. McDonald, J.S., 1979. Bovine mastitis: Introductory remarks. J Dairy Sci., 62: 117-118. Mcguire, M.A., Griinari, J.M., Dwyer, D.A. and Bauman, D.E., 1995. Role of insulin in the regulation of mammary synthesis of fat and protein. J. Dairy Sci., 78: 816-824. Meischke, H.R.C., 1979. A survey of bovine teat papillomatosis. Vet. Rec., 104: 28-31. Mejia, C.E., Preston, T.R. and Pernilla, F., 1998. Effects of restricted suckling versus artificial rearing on milk production, calf performance and reproductive efficiency of dual purpose Mpwapwa cattle in a semi-arid climate. Livest. Res. Rural Dev., 10: 1-11. Meregalli, A., Aiutolo, D., Montaghi, A. and Acciaioli, A., 1983. Milk production in Chianina cows and growth of their calves (in Italian, English summary). Rivista di Zootecnia e Veterinaria., 11: 380389. Mert, N., Tayar, M., Ogan, M., Muftuoglu, A. and Yavuz, M., 1992. Biochemical changes in the milk of cows with subclinical mastitis (in Turkish, English summary). Veteriner Fakultesi Dergisi Uludag Universitesi., 11: 13-18. Miettinen, P.V.A. and Juvonen, R.O., 1990. Diurnal variations of serum and milk urea levels in dairy cows. Acta Agric. Scand., 40: 289-296. Miller, R.H., Emanuelson, U., Persson, E., Brolund, L., Philipsson, J. and Funke, H., 1983. Relationships of milk somatic counts to daily milk yield and composition. Acta Agric. Scand., 33: 209-223. Miller, R.H., Paape, M.J., Fulton, L.A. and Schutz, M.M., 1993. The relationship of milk somatic cell count to milk yields for Holstein heifers after first calving. J. Dairy Sci., 76: 728-733. Modransky, P. and Welker, B., 1993. Management of teat lacerations and fistulae in cows. Vet. Med., 88: 995-1000. Mollet, T.A., Brooks, C.B. and Leighton, E.A., 1989. Effect of total CMT score and milk quality of the dam upon calf weaning weight in beef cattle. J. Dairy Sci., 72 (Suppl. 1): 467 (Abstr.). Mondragon, I., Wilton, J.W., Allen, O.B. and Sang H., 1983. Stage of lactation effects, repeatabilities and influences on weaning weights of yield and composition of milk in beef cattle. Can. J. Anim. Sci., 63: 751-761. Moore, R.K., Ng-Kwai-Hang, K.F., Sabler, J. and Moxley, J.E., 1981. Fat, protein and somatic cell content of foremilk, normal milk and after milk. Res. Rep. Dep. Anim. Sci. McGill Univ.: 35-38. Morgan, J.H.L., 1991. Estimates of the yield and composition of milk from two-year-old Hereford and Friesian cows and inter-relationships with liveweight gains of calves and calves. Wld. Rev. Anim. Prod., 26: 59-64. Mosdoel, G., 1978. Mastitis pathology in cows, goats and sheep. A literature review (in Norwegian, English summary). Nor. Vet.-Med., 30: 489-497. Motomura, Y., Yugi, H., Kamimura, S. and Hamana, K., 1994. Teat measurements and milk quality in dairy cows (in Japanese, English summary). J. Jpn. Vet. Med. Assoc., 47: 387-389. Murphy, J.J. and OMara, F., 1993. Nutritional manipulation of milk protein concentration and its impact on the dairy industry. Livest. Prod. Sci., 35: 117-134. Naidu, K.N., 1972. Genetic and phenotypic variation in shape and size of udder in four breeds of indian dairy cattle. Agra Univ. J. Res. Sci., 21: 75-77. Natzke, R.P., Schultz, L.H., Barr, G.R. and Holtmann, W.B., 1965. Variation in mastitis screening test and milk composition of udder quarters under normal conditions and following omission of a milking. J. Dairy Sci., 48: 1295-1299. Neave, F.K., 1975. Diagnosis of mastitis by bacteriological methods alone. Ann. Bull. Int. Dairy Fed., No.85: 19-36. Nelson, F.E., Tranmal, H., Schuh, J.D., Wegner, T.N. and Stott G.H., 1969. Criteria of abnormal milk from individual quarters during a period of high temperatures. J. Dairy Sci., 52:912 (Abstr.). Neville, W.E., 1962. Influence of dams milk production and other factors on 120- and 240-day weight of Hereford calves. J. Anim. Sci., 21: 315-320. Newman, M.A., Wilson, L.L., Cash, E.H., Eberhart, R.J. and Drake, T.R., 1991. Mastitis in beef cows and its effects on calf weight gain. J. Anim. Sci. 69: 4259-4272. 60

Ng-Kwai-Hang, K.F., Hayes, J.F., Moxley, J.E. and Monardes, H.G., 1985. Percentages of protein and nonprotein nitrogen with varying fat and somatic cells in bovine milk. J. Dairy Sci., 68: 12571262. Nickerson, S.C., Owens, W.E. and DeRouen, S.M., 2000. Mastitis prevalence in first calf beef heifers and effect on calf weaning weight. Large Anim. Pract., 21: 20-23. Nooruddin, M., Rahman, M.S. and Rahman, M.M., 1997. Prevalence and distribution of teat papillomatosis in crossbred and exotic dairy cows. Bangladesh Vet., 14: 1-2, 5-7. Oldenbroek, J.K., 1984. A comparison of Holstein Friesians, Dutch Friesians and Dutch Red and Whites. I. Production characteristics. Livest. Prod. Sci., 11: 69-81. Olson, D.P., Duren, I.P., Bramwell, K.A., Henson, S.R., Panting, R.R., Ritter, T.W. and Sharp, D.W., 1989. Clinical observations and laboratory analysis of factors that affect the survival and performance of beef cattle. Bovine Pract., No.24: 4-11. Olson, R.O., Olson, C. and Easterday, B.C., 1982. Papillomatosis of the bovine teat (mammary papilla). Am. J. Vet. Res., 43: 2250-2252. Oltenacu, P.A., Bendixen, P.H., Vilson, B and Ekesbo, I., 1990. Tramped teats-Clinical mastitis disease complex in tied cows. Environmental risk factors and interrelationships with other diseases. Acta Vet. Scand., 31: 471-478. Oltner, R., Emanuelson, M. and Wiktorsson, H., 1983. Factors affecting urea concentration in cows milk. Proc. 5th Int. Conf. Prod. Dis. Farm Anim., Swed. Univ. Agric. Sci., Uppsala, Swede.,: 195198. Oltner, R., Emanuelson, M. and Wiktorsson, H., 1985. Urea concentrations in milk in relation to milk yield, live weight, lactation number and amount and composition of feed given to dairy cows. Livest. Prod. Sci., 12: 47-57. Oltner, R. and Wiktorsson, H., 1983. Urea concentrations in milk and blood as influenced by feeding varying amounts of protein and energy to dairy cows. Livest. Prod. Sci., 10: 457-467. Opdebeeck, J.P., 1982. Mammary gland immunity. J. Am. Vet. Med. Assoc., 181: 1061-1065. ORourke, D.J., 1999. Cell counts and their uses. Cattle Pract., 7: 11-12. OShea, J. and Meaney, W.J., 1979. New intramammary infections and teat-orifice infections in suckled cows and in teat-dipped and non-teat-dipped machine-milked cows. Ir. J. Agric. Sci., 18: 37-43. stensson, K., 1993a. Total and diferencial leukocyte counts, N-acetyl-B-D-glucosaminidase activity, and serum albumin content in foremilk and residual milk during endotoxin-induced mastitis in cows. Am. J. Vet. Res., 54: 231-238. stensson, K., 1993b. Variations during lactation in total and differential leukocyte counts, N-acetyl-BD-glucosaminidase, antitrypsin and serum albumin in foremilk and residual milk from noninfected quarters in the bovine. Acta Vet. Scand., 34: 83-93. stensson, K and strm, G., 1994. Differential cell count in varios milk fractions-A sensitive method to evaluate the inflammatory status of the udder. Proc. Int. Symp.- Prosp. Fut. Dairy.: Chall. Sci. Ind., ed. Lind, O. And Svennersten, K. Alfa Laval Agri. AB, Tumba, Sweden and Swe. Uni. Agric. Sci., Uppsala, Sweden: 80-85. stensson, K., Hageltorn, M. and strm, G., 1988. Differential cell counting in fraction-collected milk from dairy cows. Acta Vet. Scand., 29: 493-500. Ozbeyaz, C., Unal, N. and Colakoglu, N., 1998. Effects of udder and teat measurements and teat shape on milk yield and milkability in Brown Swiss cows. I. Udder and teat measurements (in Turkish, English summary). Lalahan Hayvancilik Arastirma Enstitusu Dergisi., 38: 1-23. Paape, M.J. and Tucker, H.A., 1966. Somatic cell content variation in fraction-collected milk. J. Dairy Sci., 49: 265-267. Paape, M.J., Wergin, W.P., Guidry, A.J. and Pearson, R.E., 1979. Leukocytes-second line of defense against invading mastitis pathogens. J. Dairy Sci., 62: 135-153. Palmquist, D.L., Beaulieu, A.D. and Barbano, D.M., 1993. Feeds and animal factors influencing milk fat composition. J. Dairy Sci., 76: 1753-1771. Pander, B.L. and Chopra, S.C., 1986. Effect of udder and teat conformation on milk flow rate and milk production in crosbred dairy cattle. Asian J. Dairy Res., 5: 181-185. Paulicks, B., 1992. When is the urea test useful? (German, English summary). Tierzuchter., 44: 36-38. Payne, J.M., Dew, S.M., Manston, R and Faulks, M., 1970. The use of metabolic test in dairy herds. Vet. Rec., 87: 150-157.

61

Pehrson, B., Forshell, K.P. and Carlsson, J., 1992. The effect of additional feeding on the fertility of high-yielding dairy cows. J. Vet. Med., 39: 187-192. Pestevsek, U., Likosar, D., Zust, J. and Vengust, A., 1990. Use of the milk urea test to detect malnutrition in dairy cows (in Slovenian, English summary). Zbornik Veterinarske Fakultete Univerza Ljubljana., 27: 139-148. Pirlo, G., Capelletti, M. and Marchetto, G., 1997. Effects of energy and protein allowances in the diets of prepubertal heifers on growth and milk production. J. Dairy Sci., 80: 730-739. Pitcher, P.M., Ferguson, J.D., Galligan, D.T. and Chalupa, W., 1992. Importance of differentiating true and dilution effects on milk composition. J. Dairy Sci., 75 (Suppl. 1): 291 (Abstrc.). Pobric, H., Huskic, E., Filipovic, P.M. and Podzo, M., 1990. Frequency of udder abnormalities in cows on private farms (in Slovenian, English summary). Veterinaria Sarajevo., 39: 67-71. Pomies, D., Gasqui, P., Bony, J., Coulon, J.B. and Barnouin, J., 2000. Effect of turning out dairy cows to pasture on milk somatic cell count. Ann. Zoot., 49: 39-44. Ponsart, C., Ponter, A.A., Khireddine, B., Humblot, P., Sauvant, D., Mialot, J.P. and Grimard, B., 1999. A period of energy supplementation but not the type of supplement influences the insulin response to exogenous glucose in food-restricted post-partum suckler beef cows. Anim. Sci., 68: 749-761. Ponter, A.A., Grimard, B., Humblot, P., Novak, N., Khireddine, B., Sauvant, D., Thibier, M. and Mialot, J.P., 1997. Parity influences the utilization of exogenous glucose in suckler anoestrous Charolais beef cows. Anim. Sci., 65: 183-192. Pounden, W.D. and Frank, N.A., 1961. Influence of forages on mastitis. J. Am. Vet. Med. Assoc., 138: 146-150. Pounden, W.D., Frank, N.A. and Vandersall, J.H., 1960. Further observations on mastitis in cows fed legume-grass forage as soilage and as silage. J. Am. Vet. Med. Assoc., 137: 53-57. Poutrel, B. and Rainard, P., 1982. Predicting the probability of quarter infection (by a major pathogen) from somatic cell counts. Am. J. Vet. Res., 43: 1296-1299. Prajapati, K.B., Ashwar, B.K., Patel, J.P., Patel, J.B. and Singh, D.V., 1995. Size and shape of udder and teats in Kankrej cows. Indian J. Anim. Prod. Man., 11: 43-38. Prajapati, K.B., Patel, J.P., Singh, D.V., Ashwar, B.K. and Patel, J.B., 1998. Association of udder and teat measurements with milk yield in kankrej cows. J. Dairy. Food Home Sci., 17: 107-109. Preston, T.R., 1983. Restricted suckling: effects on cow and calf performance. In: Maximun Livestock Production for Minimun Land., ed. Davis, C.H., Preston, T.R., Hague, M. and Saadullah, M., Bangladesh Agricultural University, Mymensingh., 54-66. Pyorala, S., Somer, H.J. and Mero, M., 1992. Clinical, bacteriological and therapeutic aspects of bovine mastitis caused by aerobic and anaerobic pathogens. Br. Vet. J., 148: 54-62. Qureshi, M.I., Taylor, C.M. and Singh, B.N., 1984. A note on teat measurements and shape of udder and teat and its correlation with milk yield in Gir cows. Indian Vet. J., 61: 255-258. Rahnefeld, G.W., Weiss, G.M. and Fredeen, H.T., 1990. Milk yield and composition in beef cows and their effect on cow and calf performance in two environments. Can. J. Anim. Sci., 70: 409-423. Rasmussen, M.D. and Larsen, H.D., 1998. The effect of post milking teat dip and suckling on teat skin condition, bacterial colonisation, and udder health. Acta Vet. Scand., 39: 443-452. Rayner, I.H., Edmunds, J. and Stokoe, J., 1977. Comparison of dairy and multiple suckled calf production with calf access restricted. Aust. J. Exp. Agric. Anim. Husb., 17: 728-734. Reece, R.P. and Murphy, J.M., 1943. The influence of diethylsbestrol dipropionate on the lactating mammary gland of the cow. J. Dairy Sci., 26: 748-749. Refsdal, A.O., 1983. Urea in bulk milk as compared to the herd mean of urea in blood. Acta Vet. Scand. 24: 518-520. Refsdal, A.O., Baevre, L. and Bruflot, R., 1985. Urea concentration in bulk milk as indicator of the protein supply at the herd level. Acta Vet. Scand., 26: 153-163. Ridgway, T.D., Wettemann, R.P., Lents, C.A., Spicer, L.J. and Paape, M.J., 1999. Influence of mastitis, weaning and intramammary antibiotic treatment on somatic cells and tumor necrosis factor-alpha in milk of beef cows. Anim. Sci. Res. Rep., Okla. Agric. Exp. Sta., No.P-973: 296-300. Rigby, C., Ugarte, J. and Boucourt, R., 1976. Rearing dairy calves by restricted suckling. VII. Effect on mastitis development caused by Staphylococcus aureus. Cuban J. Agric. Sci., 10: 35-40. Riha, J. and Hanus, O., 1999. Relationships between reproductive parameters and selected metabolic parameters in bulk milk samples of dairy cows (in Czech, English summary). Vyzkum v Chovu Skotu., 41: 1-6. 62

Rodriguez, L.A., Stallings, C.C., Herbein, J.H. and McGilliard, M.L, 1997a. Effect of degradability of dietary protein and fat on ruminal, blood and milk components of Jersey and Holstein cows. J. Dairy Sci., 80: 353-363. Rodriguez, L.A., Stallings, C.C., Herbein, J.H. and McGilliard, M.L., 1997b. Diurnal variation in milk and plasma urea nitrogen in Holstein and Jersey cows in response to degradable dietary protein and added fat. J. Dairy Sci., 80: 3368-3376. Ropstad, E. and Refsdal, A.O., 1987. Herd reproductive performance related to urea concentration in bulk milk. Acta Vet. Scand., 28: 55-63. Ropstad, E., Vik-Mo, L. and Refsdal, A.O., 1989. Levels of milk urea, plasma constituents and rumen liquid ammonia in relation to the feeding of dairy cows during early lactation. Acta Vet. Scand., 30: 199-208. Roseler, D.K., Ferguson, J.D. and Sniffen, C.J., 1990. The effects of dietary protein degradability/undegradability on milk urea, milk NPN and blood urea in lactating dairy cows. J. Dairy Sci., 73 (Suppl. 1): 168 (Abstr.). Roseler, D.K., Ferguson, J.D., Sniffen, C.J. and Herrema, J., 1993. Dietary protein degradability effects on plasma and milk urea nitrogen and milk nonprotein nitrogen in Holstein cows. J. Dairy Sci., 76: 525-534. Roth, A.C., Gustaffson, A.H., Emanuelson, M. and Bertilsson, J., 1996. The urea concentration in milkan aid to milk production (in Swedish, English summary). Fakta Husdjur., No.14: 4. Roussel, J.D., Ortego, J.D., Gholson, J.H. and Frye, J.B., 1969. Effect of thermal stress on the incidence of anormal milk. J. Dairy Sci., 52: 912 (Abstr.). Roy, S.K., Pyne, A.K. and Maitra, D.N., 1993. Studies on teat size and lactation number in relation to incidence of subclinical mastitis in some herds of crossbred cows. Indian Vet. J., 70: 677-678. Russell, M.W. and Reiter, B., 1975. Phagocytic deficiency of bovine milk leucocytes: an effect of casein. J. Reticuloendothelial Soc., 18: 1-13. Rutledge, J.J., Robison, O.W., Ahlschwede, W.T. and Legates J.E., 1971. Milk yield and its influence on 205-day weight of beef calves. J. Anim. Sci., 33: 563-567. Ryle, M. and Orskov, E.R., 1990. On milk yields and calf rearing. Livest. Res. Rural Dev., 2: 1-6. Samuelsson, B., 1996. The influence of management routines on endocrine systems involved in the control of lactation in dairy cattle. Thesis. Swed. Univ. Agric. Sci., Uppsala, Sweden. Samuelsson, B., Emanuelson, M., Olsson, G., Moberg, K.U. and Sjaunja, K.S., 1998. Hormonal profiles, rumen volatile fatty acids and milk trans-C18:1 fatty acids in relation to milk fat content in restricted or ad libitum-fed dairy cows. Acta Vet. Scand., 48: 76-85. Sato, H., Hanasaka, S. and Matsumoto, M., 1992. Relationships among plasma metabolite levels, nutrient intakes, milk urea, fat and protein levels in dairy cattle (in Japanese, English summary). Anim. Sci. Tech., 63: 1075-1080. Schalm, O.W., 1977. Pathologic changes in the milk and udder of cows with mastitis. J. Am. Vet. Med. Assoc., 170: 1137-1140. Schepers, A.J., Lam, T.J.G.M., Schukken, Y.H., Wilmink, J.B.M. and Hanekamp, W.J.A., 1997. Estimation of variance components for somatic cell counts to determine thresholds for uninfected quarters. J. Dairy Sci., 80: 1833-1840. Schepers, A.J. and Meijer, R.G.M., 1998. Evaluation of the utilization of dietary nitrogen by dairy cows based on urea concentration in milk. J. Dairy Sci., 81: 579-584. Schultz, L.H., 1974. Changes in nutritional value mediated by the cow. J. Dairy Sci., 57: 729-737. Schultz, L.H., 1977. Somatic cell counting of milk in production testing programs a a mastitis control technique. J. Am. Vet. Med. Assoc., 170: 1244-1246. Sender, G., Glabowna, M., Lukaszewicz, M., Chabielska, L.J.B., 1987. Environmental variability and threshold value of somatic cell count in milk (in German, English summary). Milchwissenschaft., 42: 17-19. Sharaby, M.A., 1988. Factors influencing the concentrations and yields of milk constituents and their interrelationships. J. Dairy Res., 55: 171-177. Sharma, H.K. and Sidhu, N.S., 1980. A study of genetic group differences in teat and udder characteristics of Zebu, European and crossbred cattle. Indian J. Heredity., 12: 21-32. Sharma, A.K., Wilcox, C.J., Martin, F.G. and Thatcher, W.W., 1990. Effects of stage of lactation and pregnancy and their interactions on milk yield and constituents. J. Dairy Sci., 73: 1586-1592. Shook, G.E., 1989. Selection for disease resistance. J. Dairy Sci., 72: 1349-1362. 63

Shoshani, E., Leitner, G., Hanochi, B., Saran, A., Shpigel, N.Y. and Berman, A., 2000. Mammary infection with Staphylococcus aureus in cows: progress from inoculation to chronic infection and its detection. J. Dairy Res., 67: 155-169. Shukla, S.K., Dixit, V.P., Thapliyal, D.C., Garg, S.K., Kumar, A., 1997. A note on the incidence of bovine mastitis in relation to teat shape size and quarters affected. Indian Vet. J., 74: 989-990. Simensen, E., 1976. Milk somatic cells in dairy cows kept on pasture or confined indoors during the summer. Nord. Vet.-Med., 28: 603-609. Simpson, R.B., Nickerson, S.C., Phillips, J.M. and Shockey, J.D., 1994. Incidence and effect of mastitisrelated bacteria on milk production and composition in beef cows with and without selenium supplementation. J. Anim. Sci., 72 (Suppl. 1): 85 (Abstr.). Simpson, R.B., Wesen, D.P., Anderson, K.L., Armstrong, J.D. and Harvey, R.W., 1995. Subclinical mastitis and milk production in primiparous Simmental cows. J. Anim. Sci., 73: 1552-1558. Sinclair, K.D., Broadbent, P.J. and Hutchinson, J.S.M., 1994. The effect of pre- and post-partum energy and protein supply on the blood metabolites and reproductive performance of single- and twinsuckling beef cows. Anim. Prod., 59: 391-400. Singh, R.P. and Gupta., 1995. Udder and teat size measures in relation to milk production of Karan Swiss cows. J. Dairy. Fd. Home Sci., 14: 131-136. Sisson, S., 1975. Ruminant urogenital system. In: Sisson and Grossmans the Anatomy of the Domestic Animals. ed. Getty, R. Philadelphia, W.B. Saunders Company. 951. Skei, O. and Waage, S., 1998. Use of suckling calves in the treatment of clinical mastitis in cows (in Norwegian, English summary). Norsk Veterinaertidsskrift., 110: 789-797. Smith, R.N., 1983. Anatomy and physiology of the bovine teat. Proc. Br. Cattle Vet. Assoc.: 225-234. Smith, J. W. and Schultze, W.D., 1967. Variation in cell content of milk associated with time of sample collection. I. Diurnal variation. J. Dairy Sci., 50: 1083-1087. Sobari, S., Ladds, P.W., Flanagan, M. and Lee, C.G., 1976. A pathological and bacteriological study of the mammary glands of beef cows in North Queensland. Aust. Vet. J., 52: 458-461. Soldatov, A.P. and Kholodkov, S.A., 1990. Genetic resistance of cattle to diseases (in Russian, English summary). Zootekhniya., No. 6: 21-24. Sordillo, L.M., Shafer-Weaver, K. and DeRosa, D., 1997. Immunobiology of the mammary gland. J. Dairy Sci., 80: 1851-1865. Spohr, M. and Wiesner, H.U., 1991. Monitoring herd health and milk production by means of the extended milk production performance (in German, English summary). Milchpraxis., 29: 231-236. Steinwidder, A., Schweiger, P., Gruber, L., Lettner, F. and Schmid, W., 1998. Influence of feeding time and protein and energy intake on milk urea content (in German, English summary). Agrob. Res., 51: 341-355. Stobbs, T.H. and Brett, D.J., 1976. Short-term effects of level of dry matter intake upon milk yield, fatty acid composition of milk fat, and some blood constituents of Hereford cows. Aust. J. Agric. Res., 27: 175-182. Sutton, J.D., Hart, I.C., Morant, S.V., Schuller, E. and Simmonds, A.D., 1988. Feeding frequency for lactating cows: diurnal patterns of hormones and metabolites in peripheral blood in relation to milk-fat concentration. Br. J. Nutr., 60: 265-274. Sutton, J.D., Abdalla, A.L., Phipps, R.H., Cammell, S.B. and Humphries, D.J., 1997. The effect of the replacement of grass silage by increasing proportions of urea-treated whole-crop wheat on food intake and apparent digestibility and milk production by dairy cows. Anim. Sci., 65: 343-351. Syrstad, O. and Ron I., 1978. Day-to-day variation in cell counts in milk (in Norwegian, English summary). Nord. Vet.-Med., 30: 192-198. Syrstad, O., Ron, I. and Wiggen, J., 1979. Factors affecting cell counts in milk from individual cows (in Norwegian, English summary). Nord. Vet.-Med., 31: 114-121. Taralik, K., 1998. Correlation between changes in milk yield and composition and genetic and environmental factors (in Hungarian, English summary). Allattenyesztes es Takarmanyozas., 47: 153-164. Thomas, P.C. and Kelly, M.E., 1976. The effect of frecuency of feeding on milk secretion in the Ayrshire cow. J. Dairy Res., 43: 1-7. Thomas, G.W., Spiker, S.A. and Mickan, F.J., 1981. Influence of suckling by Friesian cows on milk production and anoestrus. Aust. J. Exp. Agric. Anim. Husb., 21: 5-11.

64

Thurmond, M.C., 1993. Epidemiologic methods in mastitis treatment and control. Vet. Clin. North Am. Food. Anim. Pract., 9: 435-444. Timms, L.L., Morrical, D. and Mumin, I., 1989. Dynamics of subclinical intramammary infections across lactation in beef cattle, sheep and pigs. J. Dairy Sci., 72 (Suppl. 1): 15 (Abstr.). Tomar, S.S., 1973. Udder and teat measurements and their relation with milk production in Hariana cattle. Indian J. Dairy Sci., 26: 25-28. Totusek, R., Arnett, D.W., Holland, G.L. and Whiteman, J.V., 1973. Relation of estimation method, sampling interval and milk composition to milk yield of beef cows and calf gain. J. Anim. Sci., 37: 153-158. Tripathi, G.S., Koul, G.L. and Katpatal, B.G., 1982. Biometrical studies on shape and size of udder and teats and their relation with milk yield in Gir cattle. Indian J. Dairy Sci., 35: 539-543. Trostle, S.S. and OBrien, R.T., 1998. Ultrasonography of the bovine mammary gland. Compend. Contin. Educ. Pract. Vet., 20 (Suppl ): s64-s71. Tsolov, S., Dimitrov, M., Koleva, M. and Burzilov, G., 1989. Effect of suckling a calf on the frequency of mastitis (in Bulgarian, English, summary). Veterinarna Sbirka., 87: 6-11. Twardon, J., 1988. Effects of suckling and absence of suckling on uterine involution and fertility in cows (in Polish, English summary). Zeszyty Naukowe Akademii Rolniczej we Wroclawiu, Weterynaria., 44: 25-44. Twomey, A. and Arnold, M., 1975. Bactericidal properties of ubiquitin. N. Z. J. Dairy Sci. Tech., 10: 78. Ugarte, J., 1977. Rearing dairy calves by restricted suckling. 10. Residual milk in cows suckling or not their calves after milking. Cuban J. Agric. Sci., 11:253-262. Ugarte, J., 1991. Restricted suckling in dual purpose systems. FAO Anim. Prod. Health Paper., No. 86: 199-207. Ugarte, J. and Preston, T.R., 1972. Rearing dairy calves by restricted suckling. 1. Effect of suckling once or twice daily on milk production and calf growth. Rev. Cubana Cienc. Agric. (Eng. ed.)., 6: 173182. Ugarte, J. and Preston, T.R., 1975. Restricted suckling. VI. Effects on milk production, reproductive performance and incidence of clinical mastitis throughout the lactation. Cuban J. Agric. Sci., 9: 15-26. Veitia, J.L. and Simon, L., 1972. Effect of two restricted suckling systems of calf rearing on milk production and calf growth. Rev. Cubana Cienc. Agric. (Eng. ed.)., 6: 189-193. Wadhwa, D.R., Prasad, B., Rao, V.N. and Dhaliwal, A.S., 1996. Clinico-therapeutic and histopathologic studies on bovine cutaneous papillomatosis. Indian J. Dairy Sci., 49: 206-208. Waldmann, A., Ropstad, E., Landsverk, K., Sorensen, K., Solverod, L. and Dahl, E., 1999. Level and distribution of progesterone in bovine milk in relation to storage in the mammary gland. Anim. Reprod. Sci., 56: 79-91. Walsh, J.P., 1974. Milk secretion in machine-milked and suckled cows. Ir. J. Agric. Res., 13: 77-89. Watts, J.L., 1988. Etiological agents of bovine mastitis. Vet. Microbiol., 16: 41-66. Watts, J.L., Pankey, J.W., Oliver, W.M., Nickerson, S.C. and Lazarus, A.W., 1986. Prevalence and effects of intramammary infection in beef cows. J. Anim. Sci., 62: 16-20. Weber, A.F., 1977. The bovine mammary gland: Structure and function. J. Am. Vet. Med. Assoc., 170: 1133-1136. Wegner, T.N., Schuh, J.D., Nelson, F.E. and Stott, G.H., 1976. Effect of stress on blood leucocyte and milk somatic cell counts in dairy cows. J. Dairy Sci., 59: 949-956. Weisenburger, R.D. and Mathison, G.W., 1977. Protein requirements of beef cows fed pelleted, ground or chopped barley straw in the winter. Can. J. Anim. Sci., 57: 719-725. Weller, J.I., Saran, A. and Zeliger, Y., 1992. Genetic and environmental relationships among somatic cell count, bacterial infection, and clinical mastitis. J. Dairy Sci., 75: 2532-2540. White, F. and Rattray, E.A.S., 1965. Diurnal variation in the cell content of cows milk. J. Comp. Path., 75: 253-261. White, J.M. and Vinson, W.E., 1975. Relationships among udder characteristics, milk yield and nonyield traits. J. Dairy Sci., 58: 729-738. Wiley, J.S., Petersen, M.K., Ansotegui, R.P. and Bellows, R.A., 1991. Production from first-calf beef heifers fed a maintenance or low level of prepartum nutrition and ruminally undegradable or degradable protein postpartum. J. Anim. Sci., 69: 4279-4293.

65

William, H.C., 1985. Milk collection. In: Lactation.Ed. Larson, B.L., Iowa State University Press, Ames, Iowa. 199. William, B.J., Kirubaharan, J.J., Uthuman, K.M., Kumanan, K. and Balachandran, S., 1992. Survey of incidence and complications of bovine cutaneous papillomatosis. Indian Vet. J., 69: 843-844. Wilson, L.L., Eberhart, R.J., Simpson, M.J., Varela-Alvarez, H., Rugh, M.C. and Bair, L.G., 1971. Incidence of intramammary infections and effects of number of lactations, lactation stage, quarter and calf sex on somatic cell content of milk from Angus-Holstein F1 cows. J. Anim. Sci., 33: 433437. Wolfschoon-Pombo, A., Klostermeyer, H., Buchberger, J. and Graml R., 1981. Urea in the NPN-fraction of cows milk determination, content and influences (in German, English summary). Milchwissenschaft., 36: 462-466. Woolcock, J.B., 1979. Staphylococcus aureus- A bacterial model for defense against host reactions. In: Bacterial infection and immunity in domestic animals.Elsevier scientific publishing company, Amsterdam., 13-24. Woolford, M.W., Williamson, J.H. and Henderson, H.V., 1998. Changes in electrical conductivity and somatic cell count between milk fractions from quarters subclinically infected with particular mastitis pathogens. J. Dairy Res., 65: 187-198. Wu, Z., Fisher, R.J., Polan, C.E. and Schwab, C.G., 1997. Lactational performance of cows fed low or high ruminally undegradable protein prepartum and supplemental methionine and lysine postpartum. J. Dairy Sci., 80: 722-729. Wu, Z. and Huber, J.T., 1994. Relationship between dietary fat supplementation and milk protein concentration in lactating cows-a review. Livest. Prod. Sci., 39: 141-155. Yadav, S.B.S. and Sharma, J.S., 1984. Trends of milk constituents across various stages of lactation in cross bred cows. Asian J. Dairy Res., 3: 51-54. Yagil, R., Amir, H., Rabiya, Y.A. and Etzion, Z., 1986. Dilution of milk. A physiological adaptation of mammals to water stress?. J. Arid Env., 11: 243-247. Yanagita, K., Oyamada, T., Nakanishi, Y., Tojo, H. and Ogawa, K., 1978. Variation in milk yield and milk quality of Japanese Black cattle on pasture all year round in relation to season and calving season (in Japanese, English summary). Bull. Fac. Agric. Kagoshima Univ., 28: 19-24.

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I denna serie publiceras resultat frn mindre forskningsarbeten genomfrda inom ramen fr den akademiska utbildningen och som handletts av forskare vid institutionen fr husdjurens milj och hlsa. Arbetena omfattar normalt 10 pong eller mindre. Fr publikationernas innehll ansvarar frfattarna tillsammans med handledarna. ven andra mindre forskningsarbeten frn institutionen publiceras i denna serie. Mer omfattande forskningsarbeten publiceras i institutionens rapportserie. _____________________________________________________________________ Specialarbete nr 1 Tillgren, T., Pehrson, B., 1997 Konsumtionsmnster, nringsbalans, produktions- och djurhlsoparametrar vid olika utfodringsintensitet kring kalvning hos mjlkkor. Sow response to honest begging during post-massage. Calves (Bos taurus) preference for high versus low milk replacer concentration and fat content in cows milk in a semi-natural situation. Utvrdering av Vencomatic inhysningssystem fr vrphns. I enlighet med SJV:s program fr frprvning av ny teknik.

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Augustsson, H., 1997 Alavi Moghaddam, S. L., 1997

Ekstrand, C., Odn, K., Gunnarsson, S., Onila, M., Algers, B., Svedberg, J., 1997 Kasanen, S. 1998

Additional sow gruntings - can they affect piglets suckling behaviour? Domestication effects on foraging behaviour in poultry. Effect of milkflow rate and presence of a floating nipple on abnormal sucking between dairy calves. Components of the human - farm animal relationship. Gummispaltgolv fr bundna kor och ungdjur. Provning av ny teknik enligt Djurskyddsfrordningen. Identifying factors influencing the risk of horse-related accidents. Udder health and milk composition, with special reference to beef cows. A literature review.

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Nordin. E., 1998 hrberg, J., 1999

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Lund, V., 1999 Hultgren, J., 1999

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Blomberg, A., 2000

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Velazquez, M., 2000

DISTRIBUTION: SLU Institutionen fr husdjurens milj och hlsa Box 234 532 23 Skara, Sweden, Tel 0511 - 67000

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