AN ABSTRACT OF THE THESIS OF Tariq A. AL-Sabbagh for the degree of Masters of Science in Animal Science presented on June 21,1994. Title: The Relationship of Prepartum Body Condition Score To Postpartum Colostrum Ouality And Milk Yield And Composition in Polypay Sheep. Abstract approved. Redacted for Privacy Lloyd Swanson Two experiments were conducted at the OSU Sheep Center to examine the effect of body condition score at lambing (BCSL) on immunoglobulin concentrations (IgG), milk yield and composition and lamb performance. The experiments used Polypay ewes, 4 to 7 years of age, one-half mated to Polypay rams and one-half mated to Columbia rams. adjusted to two lambs shortly after birth. Litters were Milk yield was determined by hand milking, aided by oxytocin. Data were analyzed using least squares analysis of variance. In the first experiment, 101 ewes were divided into three groups at lambing, based on their BCSL (2.5, 3.0, 3.5). Colostrum samples were collected by hand milking from each ewe within 11 hr of lambing and IgG concentrations were determined using radial immunodiffusion. Colostral IgG averaged 57 ± 2.3 mg/ml and decreased significantly with time. The estimated IgG at parturition (time 0) was 76 mg/ml and linear regression analysis predicted that IgG would decline to 0 at 23 hr postpartum. Colostral IgG concentrations were not affected by BCSL, age of ewe, breed of mating sire or birth rank. Lamb mortality through weaning was 17.6% and was not influenced by BCSL, age of ewe, breed of mating sire, sex of lamb, or birth rank. Total weaning weights of lambs was 58.5 ± 1.38 kg and was not affected by BCSL. Total weaning weights were the lowest in 7-yr-old ewes (31.7 ± 3.50 kg) and the heaviest in 6-yrold ewes (53.1 ± 2.56 kg). Total weaning weights were greater in lambs sired by Polypay rams (50.0 ± 2.03 kg) than in lambs sired by Columbia rams (38.0 ± 2.17 kg). The second experiment used 43 ewes, selected from the first experiment, that had BCSL of either 2.5 or 3.5. Body condition scores, milk yield and composition, and lamb weights were obtained at three 28 day intervals beginning at 27 ± 4 days after lambing. Body condition score decreased significantly (24%) in the high BCSL group (3.5) during the first month after lambing but only decreased by 1.6% in the low BCSL group (2.5). Thereafter, the two groups plateaued during the second month and then increased significantly during the third month, yielding a quadratic response. The 4-yr-old ewes were the lowest in BCS each month and ewes giving birth to twins had higher BCS's than ewes having multiple lambs (3). Milk yield declined linearly from 3,260 ± 342 ml at the first month to 1,571 ± 193 ml at the third month. Milk yield was not affected by BCSL or other parameters. Percent milkfat declined curvilinearly from 11.71 ± 0.481% at the first month to 8.67 ± 0.526% at the third month and was not affected by BCSL. Percent milk protein increased in a quadratic fashion from 4.02 ± 0.068% at the first month to 5.25 + 0.082% at the third month. It was higher (P<0.05) in the low BCSL group at the second month and in 7-yr-old ewes during the entire lactation. Percent milk protein was always higher in ewes giving birth to twins, particularly at the third month (P<0.05), and in ewes mated to Polypay rams. Lamb birth weight and growth from birth to weaning were not affected by BCSL. Lamb weights were always the highest at each month for 7-yr-old ewes and purebred lambs were always heavier than crossbred lambs. Twin female lambs were heavier than twin male lambs, although twin male lambs gained weight more rapidly after the second month. It is concluded that BCSL, over the range studied, is not an important factor affecting performance of ewes or their lambs during lactation. The Relationship of Prepartum Body Condition Score to Postpartum Colostrum Quality and Milk Yield and Composition in Polypay Sheep by Tariq Ashour AL Sabbagh A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Completed June 21, 1994 Commencement June 1995 APPROVED: Redacted for Privacy Professor of Animal Science in charge of major Redacted for Privacy Head of Department of Animal Science Redacted for Privacy Dean of Graduate School Date thesis is presented June 21, 1994 Typed by researcher for Tariq Ashour AL Sabbagh TABLE OF CONTENTS CHAPTER 1. INTRODUCTION 1 CHAPTER 2. LITERATURE REVIEW 4 Body condition score as an estimator of nutritional status Effect of body condition score Colostral IgG concentration Factors affecting milk production and composition . . . . . CHAPTER 3. EXPERIMENTAL PROCEDURE Body condition score at lambing Ewe management Lamb management Colostrum collection Milking process Analysis of milk composition . . . Analysis of colostral IgG concentration Data analysis 19 19 20 21 21 23 24 25 RESULTS 31 Colostrum Experiment 31 Colostral IgG Lamb mortality Weaning weight 31 32 32 Lactation Experiment Body condition score Daily milk yield Milk composition (fat & protein) Lamb weight CHAPTER 5. 7 10 12 17 (fat & protein) CHAPTER 4. 4 33 . 33 35 36 37 DISCUSSION 58 Colostrum Experiment 58 Colostrum Lamb Mortality Lactation Experiment Body Condition Score Milk Yield Milk Composition 58 60 62 62 64 69 Table of Content, Continued Lamb Weights Conclusion BIBLIOGRAPHY 72 75 76 LIST OF FIGURES Figure Page 1 Linear regression curve for sheep immunoglobulin (IgG) concentrations using radial immunodiffusion (Exp. 1). 29 2 Linear regression of ewe colostrum immunoglobulin(IgG) concentrations on time of collection after parturition 50 (Exp. 1). 3 Least square means of body condition score during lactation of ewes of different body condition scores at lambing (BCSL) and of different birth rank at lambing (Exp. 2). 4 Least square means of body condition score during lactation of ewes of different ages at parturition (Exp. 2). 52 5 Linear regression of least square means of daily milk yield of ewes of different body condition score at lambing (BCSL)(Exp. 2). 52 6 Linear regression of actual daily milk yield of ewes of different body condition score at lambing (BCSL) (Exp. 2). 7 Least square means of percent fat and protein in milk during lactation of ewes of different body condition score at lambing (BCSL) (Exp. 2). 8 Least square means of total lamb weight during lactation of ewes of different body condition score at lambing (BCSL) (Exp. 2). . . . . . . . . . . . . 50 54 54 56 LIST OF TABLES Table Page 1 Classification of ewes involved in the colostrum (n= 101) and lactation (n= 43) studies. 2 Least square means of colostral immunoglobulin (IgG) concentrations and total weaning weight of lambs (0,1,2) at 95 days of age as affected by body condition score at lambing (BCSL), age of ewe, mating sire and birth rank. 3 Lamb mortality classified by body condition score at lambing (BCSL), age of ewe, mating sire, sex of lambs and birth rank. 4 Least square means of body condition score as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. 5 Least square means of milk yield (ml) as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling . . . . . . . . . . . 28 40 41 42 43 . (twin) lambs. 6 Least square means of percent fat in milk as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. 7 Least square means of percent protein in milk as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. 8 Least square means of total weight born (twin lambs) as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. 46 9 Least square means of total lamb weights' (kg) at 30, 60 and 90 days postpartum and at weaning (95 days) as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. 47 10 Correlation coefficients for ewe and lamb characteristics (Exp. 1). . 44 . . . . . 45 48 List of Tables, Continued 11 Correlation coefficients of some ewe and lamb characteristics during lactation (Exp. 2). . . 49 The Relationship Of Prepartum Body Condition Score To Postpartum Colostrum Quality And Milk Yield And Composition in Polypay Sheep. CHAPTER 1. INTRODUCTION Great Britain and some parts of Europe such as France and Romania are important and famous areas in the sheep industry. These areas are rich in natural resources and enjoy environments conducive to the success of the sheep industry. In the Middle East, however, sheep raising is challenging because of the scarcity of natural resources Yet, the sheep industry remains vital to the Middle East for the following reasons: 1) sheep are sociologically important because they are a source of wealth and respect in this society, thus people prefer meat from sheep as compared to other species, 2) sheep are easier to raise than other grazing animals in an environment which has insufficient natural resources (water and forage), and 3) sheep are important to the religion and customs of the Middle Eastern people. In religious ceremonies such as Hajj and Eid, the demand for sheep increases many times. Millions of sheep are slaughtered at these festivals to share with the poor. The need for sheep for the religious festivals cannot be met by the domestic supply, especially in Saudi Arabia, the Hajj location. As a result, all of the live sheep exported from New Zealand are shipped to Saudi Arabia (Storey & Sri Ramaratnam, 1991). In addition, the milk of ewes is high in 2 fat, which makes it unique and preferred in the Middle East. These people use the milk to produce dairy products, such as cheese and yogurt and use the wool for clothes; even the skin may be used to make musical drums. For these reasons the sheep industry is very important in the Middle East. The socio-economic importance of sheep creates the need to discover the factors which affect their productivity. Scientists have examined the various aspects of sheep husbandry to maximize their benefits while minimizing costs. One area which has been studied closely is the relationship of body condition score to performance of sheep. After a gestation period of about 145 days the pregnant ewe gives birth to single, twins, triplets and occasionally quadruplets and quintuplets. suckled by newborn lambs. Colostrum is the first milk Colostrum is high in immunoglobulin, particularly immunoglobulin G (IgG), which provides passive immunity to the newborn lambs; the level of passive immunity attained is proportional to the concentration and quantity of immunoglobulin and the time interval after birth in which the colostrum is consumed (Esser et al., 1989). During the nursing period, lambs derive most of their nutrients for growth from the ewe's milk. The nursing period the time interval from birth to weaning -varies in length from about 6 to 13 weeks. Generally, the shorter the nursing period the lower the 3 growth rate and the higher the post weaning mortality rate (Fogarty et al., 1992). Thus, milk production is very important to lamb growth. Therefore, it is important to gain more information about factors that affect milk production. Body condition scoring, which reflects the nutritional status of the ewe, may be an important factor affecting milk yield. This study was designed to provide information on the relationship of prepartum ewe body condition score and postpartum colostrum quality and milk yield and composition. 4 CHAPTER 2 LITERATURE REVIEW Sheep are one of many resources used by man. Sheep have the ability to convert low quality food sources from plants into such products as milk, meat, animal oil and wool. The desire for these commodities has created a proliferation of scientific research to develop methods that maximize the ratio of benefits to cost in raising sheep. Numerous studies have investigated ways to measure and enhance sheep performance and productivity. Productivity of sheep can be measured by litter size, number of lambs weaned per ewe, (Sidwell and Miller, 1971; Dickerson and Glimp, 1975; More-O'Ferral, 1976; Parker and Pope, 1983; Lewis and Burfening, 1988) and the nutritional status of ewes and lambs (Jordan and Mayer, 1989). Body Condition Score As an Estimator of Nutritional Status Body condition score (BCS) and live weight are two systems for measuring nutritional status in ewes. Body condition score, the assessment of fat and muscle in the loin region, is a system which estimates the degree of muscling and fat development in an animal (Russel et al., 1969; Ducker and Boyd, 1977). It is considered to be a more subjective, accurate and inexpensive estimator of a ewe's nutritional status than body weight because skeletal size 5 adds to the live weight without being a determinant of the nutritional status. Fat stored in the body becomes a reservoir for energy. Pregnancy, lactation and undernourishment, for example, would be a time when an animal utilizes this fat reserve. Body condition score is a better predictor of fertility and total fat deposits in the animal than body weight (Barth and Neumann, 1991). The regression of BCS on body weight is curvilinear; the change in body weight increases at an increasing rate as BCS increases from 1 to 5 (Teixeira et al., 1989). Body condition scoring was developed by Jefferies (1961), using a scale of one to five, where 1=thin and 5=obese. al. Russel et (1969) refined the system by adding half point gradations. The procedure for scoring the condition of an animal's body is done tactically in the loin area, assessing the degree of muscle and fat that surround the vertebrae. This area is optimal in that it is the first part of the body to be covered with fat and muscle and the last area to lose it. Also, subcutaneous fat correlates more closely with percentage of total body fat than does the intermuscular, mesenteric or perirenal fat (Russel et al., 1971; Greg, 1974). This system has proven to be useful and is reasonably repeatable. Oriordan and Murphy (1987) observed that 370 of the scores were repeatable when they used this system for determining the body condition of 54 pregnant ewes. Ninety- 6 five percent of the remaining scores were within 0.5 units, and no animal had scores differing by more than 1.0 unit, giving credence to the repeatability of this technique. Although BCS has a lower repeatability than live weight, it correlates more closely with percent body fat. Of two ewes with the same skeletal size but with a BCS differing by 1 unit, the ewe with the higher BCS was 10 kg heavier (Russel et al., 1969). Body condition scoring is also used for other species, for example, a scale of 1 to 5 (Meinert et al., 1992; Perkins et al., 1985; Edmonson et al., 1989; Ruegg et al., 1992) or 1 to 10 (Northcutt et al., 1992) is used to score dairy and beef cattle. In dairy cattle, 1 unit of BCS is equivalent to 545.5 kg milk. The goal for commercial ewes, sold for meat purposes, is a body condition score of 3.0, in which fat represents 301; of the weight of a fleece-free carcass. The primary factor affecting BCS is dry matter intake, which is influenced by the age and breed of sheep (Bocquier et al., 1987). However, dry matter intake decreases in ewes with BCS of a 4 (Trabalza-Marinucci et al., 1992). A BCS a 3.0 is important for embryonic development during early pregnancy, resulting in lower prenatal mortality (Alexander, 1964; Nordby et al., 1986). At mid gestation a BCS a 3.0 is necessary for optimal placental growth (Mellor, 1990). The most crucial period is late 7 pregnancy (90 days to parturition) when the greatest fetal growth occurs (Robinson et al., 1977; Mellor, 1990). Effects of Body Condition Score Body condition score has been studied to observe its effect on ewe reproductive performance. Rhind and Schanbacher (1991) studied the effect of BCS on the reproductive performance of crossbred ewes. Ewes with a high BCS had larger follicles than ewes with a low BCS during the luteal and follicular phases of the estrous cycle. However, BCS had no effect on the number of follicles. Generally, reproductive performance improved when BCS ranged from 2.25 to 2.50. After ewes attained a BCS of 2.50, some breeds continued to improve in reproductive performance (Welsh Mountain) while reproductive performance declined in other breeds (Gunn et al., 1991). Coop and Clark (1969), Gunn et al. Doney (1979), Rhind et al. (1969), Gunn and (1984a,b), and Meyer (1985) observed a positive correlation between BCS and mean ovulation rate, although Rhind et al. (1991a) failed to observe this correlation in a small study using Greyface ewes. The BCS had no significant effect on luteinizing hormone (LH) and follicle stimulating hormone (FSH) concentrations (Rhind et al., 1984a) and pulse amplitude of 8 LH during the estrus cycle (Rhind and Schanbacher, 1991). Two days after ovariectomy, ewes with a high BCS had a higher mean LH pulse frequency and higher FSH concentrations (Rhind et al., 1991b; Rhind and Schanbacher, 1991). Body condition score directly affected hypothalamic activity and gonadotropin releasing hormone (GnRH) secretion but not pituitary sensitivity to GnRH (Rhind et al., 1991b). Rhind et al. (1991c) repeated a study using 37 ewes, varying from a high BCS (3.50) to moderate BCS (2.50 to 2.75). No significant differences in pregnancy rate, ova wastage rate and profiles of progesterone and prolactin were observed, although ewes with a lower BCS tended to have lower prolactin levels (Rhind et al., 1991b). Body condition score did not affect progesterone levels following exogenous LH pulses (Rhind et al., 1991b). Unlike the previous findings, the conception rate was slightly higher in ewes with a high BCS (Wallace, 1961; Coop, 1962; Taplin and Everitt, 1964; Coop, 1966; Meyer and Bradford, 1973; Nordby et al., 1986; Gunn et al., 1990; West et al., 1991). While some studies have concluded that embryonic survival was not affected by changes in BCS (Rhind et al., 1984a), most studies have shown a positive correlation between BCS and embryonic survival (Guerra et al., 1971; Gunn et al., 1972; Gunn and Doney, 1975, 1979; Cumming et al., 1975; Gunn and Maxwell, 1989; West et al., 1991). 9 Ewes with a higher BCS tended to have greater lamb survival (Johnson et al., 1982; Sykes, 1982; BerggrenThomas, 1984; Jordan and Feuvre, 1989; Nawaz et al., 1992; Nawaz and Meyer, 1992; King et al., 1990). Jordan and Hanke (1991) failed to observe a positive correlation between ewes with a high BCS and lamb survival but their experiment was conducted on ewes with a high mean BCS which produced a higher quality and quantity of milk for their lambs. Lamb birth weight and lamb survival were positively correlated with a high BCS at late gestation (Alexander, 1964; Robinson, 1983; Beeston, 1984; Scales et al., 1986; Jordan and Mayer, 1989; Hohenboken et al., 1988) due to reduced prenatal growth in ewes with low BCS (Rattray et al., 1974; Mellor and Murray, 1981, 1982). A low plane of nutrition in ewes during the last six weeks of gestation resulted in low birth weight lambs (Treacher, 1971). However, other studies have shown no effect of BCS on birth weight of lambs (Slade, 1980; Jordan and Hanke, 1991; Gunn et al., 1986). The lack of differences in these latter studies, however, may have been due to the small range in BCS's in these ewes. The growth rate of the lambs, which is dependent on milk yield, was not affected by differences in the BCS of their dams in late gestation (Slade, 1980; Gibb and Treacher, 1982; Jordan and Hanke, 1991). However, Treacher (1971), Gibb and Treacher (1980), Rhind et al. Aguilera et al. (1992) and (1992) reported that ewes with a lower BCS 10 during late gestation had decreased milk production when a restricted diet was fed after parturition. Molina et al. (1991), on the other hand, reported that ewes with a higher BCS weaned heavier lambs. The onset of anestrus occurred 51 to 75 days earlier in ewes with low BCS (Forcada et al., 1990). Not only did ewes with a high BCS exhibit a higher incidence of estrus, they also exhibited greater ovarian activity, greater number of corpora lutea (Forcada et al., 1990), and increased litter size (Burditt et al., 1987). Colostral IgG Concentrations Colostrum is the first milk produced by the ewe after parturition. It is rich in immunoglobulins which are important for passive immunity of the lambs. The level of passive immunity is proportional to the concentration of immunoglobulin in colostrum and the quantity of colostrum ingested (Esser et al., 1989). Immunoglobulin G (IgG) is one of the five classes of humoral antibodies produced by the body in response to invasions by bacteria, fungi and viruses. The concentration of IgG is usually determined using the radioimmunodiffusion (RID) method developed by Mancini et al., (1965) and modified by Fahey and McKelvey (1965). Colostrum quality can also be determined by enzyme-linked 11 immunosorbant assay (ELISA) or radioimmunoassay (RIA), that are extremely precise, or less precisely by specific gravity (Toenjes et al., 1991). In general, the RID method is simple and requires a minimal investment in equipment (Check and Piper, 1986). There is no significant relationship between the appearance or color of colostrum and its concentration of immunoglobulins. Toenjes et al. (1991) observed a small but negative relationship between the interval from calving to first milking and immunoglobulin content. The quantity of colostrum was positively related to the colostral IgG production (Esser et al., 1989; Shubber et al., 1979). The age and genotype of the ewe, BCS, litter size, and sex of the lambs had no significant effect on colostral IgG concentration (Esser et al., 1989) or colostrum yield (Thomas et al., 1988). On the other hand, Gilbert et al. (1988) observed higher IgG concentrations in yearling ewes than in older ewes. Colostrum production was lower for single-bearing than for twin-bearing ewes (Gallo and Davies, 1987). Ewes delivering triplets had higher protein and immunoglobulin concentrations in their colostrum than ewes delivering twins (Gallo and Davies, 1987; Gilbert et al.,1988). 12 Factors Affecting Milk Production and Composition Newborn lambs depend on milk for most of their early nutrition. Quality and quantity of milk consumed during the first weeks of life are important determinants of both health and growth of lambs (Brown and Hogue, 1985). Therefore, factors that affect milk yield and composition, such as nutrition, lactation stage, diet, and age and breed become important to lamb survival. Milk yield can be measured by hand milking, machine milking and lamb-suckle-weigh methods. Studies show that the hand milking and machine milking techniques yield similar results (Papachristoforou, 1990). The lamb-suckle- weigh method is not as reliable as the hand or machine milking methods because it depends on the lamb's appetite (Peart, 1967). Doney et al. (1979), however, found that hand or machine milking agreed well with the lamb-suckleweigh method. Milk production begins immediately after parturition, peaks at 2 to 4 weeks postpartum, and then declines (Peart, 1967, 1968, 1970; Wohlt et al., 1984; Brown et al., 1987; Reynolds and Brown, 1991). Peart (1967) observed that milk yield decreased from 2.91 L/day during the third week of lactation to about 0.87 L/day during the tenth week of lactation. During the lactation period, milk composition averaged 5.5% protein and 6.2% fat (Boylan and Sakul, 1988; 13 Banda, 1992). Percentage of fat in milk used to be determined using the Babcock method. The Babcock method involves the use of a strong acid, making it hazardous to Therefore, the infrared method (IR) use. is currently widely used to determine the percentages of fat and protein in milk; it is based on the absorption of IR energy at a specific wave length by carbonyl groups in ester linkages of the fat molecule and by peptide linkages between amino acids of protein molecules. Dorset, Finnsheep, Lincoln, Rambouillet, Ramonov, Suffolk and Targhee ewes were tested for milk production and composition (Sakul and Boylan, 1992). Suffolk's had the highest milk yield while Ramonov's had the lowest milk yield. Milk of the Finn breeds was the lowest in protein content. Bass (1989) failed to detect a difference in milk yield between the Scottish Black Face and East Friesland breeds. Generally, milk production was lower in American breeds than in the "dairy" sheep breeds of Europe and the Middle East (Reynolds and Brown, 1991). Snowder and Glimp (1991) failed to detect differences in milk yield between the Rambouillet, Columbia, Polypay and Suffolk breeds. Cross breeding was an effective means to increase milk yield but not necessarily milk composition (Sakul and Boylan, 1992). Milk fat and protein were unaffected by the number of suckling lambs (Snowder and Glimp, 1991) but increased when low roughage diets (high energy) were 14 introduced (Brown and Hogue, 1985). Milk protein concentrations were higher when ewes were fed on a high plane of nutrition (Barnicoat et al., 1965) or when diets were supplemented with fish meal (Penning et al., 1988). Dorset's and Ramonov's produced milk with the highest protein content while the Finn breed was the lowest in protein content. Milk yield was affected by stage of lactation; it peaked at the third to fourth week and declined gradually during the remainder of the lactation (AL Saiegh and AL Khauzai, 1993). Each 2.3 kg of milk resulted in 0.45 kg of lamb gain (Robinson and Forbes, 1968). Part of the decline in milk production can be attributed to the decreased suckling stimulus as lambs begin supplementing their diets with other feeds (Wohlt et al., 1984; Reynolds and Brown, 1991), resulting in a positive correlation between milk yield and average daily gain in lambs during early lactation (Peart, 1967). Body weight of ewes and lamb birth weight, along with daily lamb weight gain during the different periods of lactation, correlated with milk yield (AL Saiegh and AL Khauzai, 1993) . Nutrition is the primary factor affecting milk yield (Barnicoat et al., 1949). Milk yield was positively correlated with feed consumption at the eighth week of lactation (Barnicoat et al., 1949; 1991). Reynolds and Brown, The quality of the diet affected milk yield as well. 15 When feed was restricted after lambing, ewes with a higher BCS had higher milk yields than ewes with a lower BCS (Peart, 1970; Russel et al., 1977), although this difference disappeared when the ewes were fed to requirement (Peart, 1967, 1968). Low dietary protein during the last six weeks of gestation led to a lower milk yield after parturition (Wallace, 1948a; Slen and Whiting, 1952; Bass, 1989). Accessibility to water also affected milk yield. Ewes watered every 24 hr produced more milk than ewes watered every 48 or 72 hr (Aganga et al., 1988). A decrease in the watering interval also resulted in a significant increase in fat and protein concentrations (Aganga et al., 1988). Milk yield is also influenced by the number of suckling lambs (suckling stimulus). Ewes nursing single lambs had lower milk yield in late lactation than ewes nursing twins (Wallace, 1948a,b; Doney and Munro, 1961; Robinson and Forbes, 1968; Ricketts, 1970; Joseph and Foot, 1980; Torres- Hernandez and Hohenboken, 1980; Hinch et al., 1983; Thomas et al., 1988; McMillan et al., 1988; Bass, 1989; Dove and Milne, 1991; Snowder and Glimp, 1991; Peeters et al., 1992; Rhind et al., 1992). Milk yield was affected by the adjusted number of suckling lambs during the lactation period, regardless of the actual number of lambs born (Rhind and Schanbacher, 1991). Fecht and Boylan (1987) and Reynolds and Brown (1991), however, failed to detect 16 differences in milk yield between ewes nursing single or twin lambs. (n=24) This might be a result of a small sample size . Suckled milk yield was correlated positively with udder width at peak lactation (r=0.66) and with ewe body weight (r=0.75) at breeding (Reynolds and Brown, 1991). Generally, larger udders produce more milk than smaller ones (Linzell, 1966; Davis et al., 1980; Reynolds and Brown, 1991). Milk yield decreased significantly after ewes attained 4.5 years of age. Peeters et al. (1992) observed that two- year-old ewes produced more milk than one-year-old West ewes of the European breed. Ewes with high milk production during early lactation continued high performance of milk yield at 45 days postpartum. 17 CHAPTER 3 EXPERIMENTAL PROCEDURE Two experiments were conducted at the Oregon State University Sheep Research Center during the 1993 lambing season (February March) using 104 Polypay ewes. One-half of these ewes were mated to rams of the Columbia breed and the other half were mated to rams of the Polypay breed. The experiments were conducted to determine the effect of body condition score at lambing (BCSL) to colostral IgG quality (Exp. 1) and the effect of BCSL on body condition score (BCS), milk yield, percentages of fat and protein in the milk and weight gain of lambs at selected intervals during lactation (30, 60 and 90 days postpartum) and at weaning (95 days postpartum)(Exp.2). In Experiment 1 we were only able to collect colostrum samples from 101 of the 104 ewes because a snowstorm prevented access to the Sheep Research Center within 12 hr of lambing for 3 of the ewes. Table 1 illustrates how the 101 ewes were allocated when classified according to BCSL, age of ewe at lambing, breed of sire and number of lambs born (birth rank). All of the ewes were within a BCSL range of 2.5 to 3.5 and varied in age from 4 to 7 yr. The majority of ewes (55%) had twin lambs at parturition, 8% had singles and the remaining 37% had 3 to 5 lambs at parturition (R = 3.4 lambs). Within 2 hr of lambing, the litters of ewes with more than 2 live lambs were reduced to 18 two lambs of similar bodyweight, regardless of sex, to nurse each ewe (hereafter referred to as adjusted lambs). Forty-three ewes were selected from the colostrum experiment for the lactation study (Experiment 2)(Table 1). One ewe was dropped after the first month of lactation when she developed mastitis and two additional ewes were dropped after the second month of lactation; one died of natural causes and one had a severe teat injury which precluded suckling or hand milking. To allow for maximum detection of differences due to BCS, only ewes with the BCS extremes (2.5 or 3.5) were used. Due to a shortage of ewes with a BCS of 3.5, only 16 of the 43 ewes had a BCS of 3.5. The ages of the ewes ranged from 4 to 7 yr; most (46.5%) were 6 years of age and in their fifth season of lambing. Since all ewes were adjusted to nurse two lambs during lactation, ewes lambing with singles were deleted from the lactation study. Forty percent of the ewes gave birth to three to five lambs which were subsequently adjusted to two lambs. Slightly more than one-third (370) of the ewes nursed two lambs of the same sex (an equal number of ewes nursed two males or two females) and the remainder nursed one lamb of each sex (Table 1). 19 Body Condition Score at Lambing Ewes (n=104) were scored weekly for body condition beginning three weeks before lambing and continuing until all ewes had lambed. The last BCS prior to parturition was considered to be the BCSL. The ewes were scored individually by the same person to decrease variability in scoring. The scoring system was based on a 5 point scale with half-point gradations where 1 is very thin and 5 is obese (Russel et al., 1977). This system is based on determining the degree of sharpness of the spinous and transverse processes in the lumbar region by palpation (Jefferies, 1961). The procedure of body condition scoring can be summarized in three steps. The first step is to palpate the degree of prominence of the spinous process in the loin region using the thumb. The second step is to palpate the degree of sharpness of the transverse process in the loin region, using the tips of the fingers for palpation. Finally, the tips of the fingers are used to palpate the depth of muscle and fat in the loin region. Ewe Management During early pregnancy the ewes were grazed on a mixed grass-clover pasture. At two weeks prior to lambing, the 20 ewes were moved to the lambing barn where they were given 0.91 kg rolled corn daily and grass-clover hay to appetite. Immediately after lambing, each ewe and her lambs were moved to a 1.75 m2 enclosure for that and the following day to ensure bonding. Here the ewes received a ration of alfalfa pellets to appetite. On the second day after lambing and after dams were checked for mastitis (using the California Mastitis Test), ewes and lambs were returned to group pens where they again received a ration of 0.91 kg rolled corn daily and grass-clover hay to appetite. After all ewes had lambed, they were returned to a mixed grass-clover pasture. The ewes had been bred in late September and lambed within a three week period in February and lambs were weaned in May. Lamb management Within 2-3 hr of lambing, the lambs were checked for entropion (turned in eyelid) and navels were dipped in a solution of 7% iodine. Within 8-16 hr of lambing, lambs were docked using an elastrator band, weighed, tattooed, ear tagged, vaccinated with IBR\PI3 (Infectious Bovine Rhinotracheitis\Parainfluenza Type 3) and male lambs were castrated with an elastrator band. Lambs were vaccinated for Soremouth and Enterotoxemia (Clostridium perfringens type C & D) twice; at 7-10 d and at 3-4 wk postpartum. 21 Colostrum Collection Colostrum samples (30 ml approximately 15 ml from each one-half of the udder) were obtained from each ewe by hand milking within 12 hr of lambing. The concentration of immunoglobulin is known to sharply diminish after that time (Shubber et al., 1979). sampling was recorded. The time interval from lambing to The colostrum samples were stored at -4 °C for subsequent analysis of colostral IgG concentration. Milking Process Since all ewes had lambed within a three week period of time, the 43 ewes were arranged into three milking groups to overcome the difficulty of hand milking 43 ewes within one day. Each milking group consisted of ewes that had lambed within a 7-day time period. The first milking was centered at 27 ± 4 days postpartum and the second and third milkings at 28-day intervals thereafter (55 ± 4 and 83 ± 4 days. For convenience these will be referred to as 30, 60 and 90 days. On the day of milking, the ewes and lambs were moved to a barn where the lambs were weighed and the ewes were body condition scored by the same individual as previously. The ewes were physically separated from their lambs but within sight and sound of each other. The ewes were individually 22 moved to an elevated platform for hand milking. Oxytocin (5 IU) was administered intramuscularly (im) in the rear leg (McCance, 1959). Although oxytocin is often administered intravenously, Geenty (1980) found that it was just as effective when administered intramuscularly. The ewes were hand milked until all milk had been removed, and a second injection of oxytocin (5 IU, im) was administered. After a 2 min wait the ewes were hand milked again and the milk was discarded. After a separation period of 3-4 hr (McCance, 1959; Thomas et al., 1988, Brown et al.,1987) which corresponds to normal suckling intervals of lambs (Ricordeau et al., 1960) the same hand milking procedure, utilizing two injections of oxytocin (5 IU, im), was followed. Milk from the latter two milkings was combined, the volume recorded and a sample taken for fat, protein and somatic cell determination. Milk volume was later corrected to 24 hr secretion, assuming that milk synthesis was similar during the entire 24 hr as was determined for the 3-4 hr separation time. Following milking, the ewes and their lambs were returned to pasture. The same milking procedure was followed at 60 and 90 days. Hand milking, using the oxytocin-induced method (McCance, 1959; Wohlt et al., 1984; Reynolds and Brown, 1991; Holcombe and Hallford, 1988), was employed rather than the traditional lamb-suckle-weigh method because of greater accuracy for the determination of milk yield. The lamb- 23 suckle-weigh method varies with appetite of the lambs (Peart, 1967) and urination or defecation by the lambs during the suckling period. Analysis of milk composition (fat and protein) Milk samples were filtered to remove debris, preserved with Bronopal, refrigerated overnight and then sent to the DHIA Testing Center in Salem, OR. The samples were analyzed for fat and protein by infrared spectroscopy (Williams, 1984; Snowder and Glimp, 1991) using a Multi Spec spectrometer (Berwind Instrument Group, Florida), and for somatic cell count using a Foss Model 215 (Foss America Inc., New York). Fat determination in sheep milk, using the infrared spectroscopy method, differs slightly from that of cows due to differences in the fat refractive index (Grappin, 1986). Results for sheep milk can be corrected using a regression equation (Reynolds and Brown, 1991). However, because the difference between the two readings would not have exceeded 1.7%, the calculated milk fat percentage was not corrected. Similarly, the percentage of protein in sheep milk as determined by infrared spectroscopy differs from bovine milk due to differences in the ratio of whey protein to true protein (Grappin, 1986). Because this difference did not exceed 10% in any of the samples, the calculated protein percentage was not corrected. 24 Analysis of Colostral IgG Concentration The IgG concentration in colostrum was analyzed using the radioimmunodiffusion (RID) technique (Mancini, 1965; Fahey and McKelvey, 1965). This technique depends on precipitation of the antibody-antigen complex around a well. Plastic plates (10.5 x 5.5 cm) containing a 1 mm layer of gel which incorporated antibody to sheep IgG (The Binding Site Limited, Birmingham, England) were used. Fourteen wells (2 mm diameter) were distributed throughout the gel plate. Diluted colostrum samples or IgG standards (5 Al) were pipetted into each well, the plates covered, and then incubated at room temperature in a humidified atmosphere for 72 hr. As the colostral or standard IgG (antigen) migrated through the gel, it complexed with the antibody. The distance migrated was proportional to the concentration of IgG. When a sufficient concentration of the Ag-Ab complex had accumulated (proportional to the concentration of colostral IgG), it precipitated and formed a visible ring around the well. All colostrum samples were initially diluted 1:40 using 7% BSA in distilled water (Guidry and Pearson, 1979). Any colostral sample that had a precipitate ring diameter greater than that of the high standard (25% of the samples) was further diluted to 1:80 or 1:160. Samples with an indistinct precipitate ring were also reassayed. After the 25 72 hr incubation period, the diameter of the precipitate rings were determined using a jeweller eyepiece (7x) with a micrometer scale inscribed on the objective. If the diameter of the precipitate ring of the high standard was within 0.3 mm of 9 mm, the standard curve provided with the RID kits was used. If not, a standard curve utilizing the three standards was constructed and a regression equation was used to calculate the concentration of the unknowns, using the square of the precipitate ring diameter as the dependent variable (Figure 1). Although the precipitate ring diameter of the high standard (1,000 µg /ml) should have attained 9.0 mm, the 11 plates averaged 8.61 ± 0.06 mm, but were consistent (CV = 2.43%). Of the 11 plates, standards. 5 contained a complete set of The medium (600 µg /m1) and low (100 µg /ml) standards averaged 6.96 ± 0.04 mm and 4.27 ± 0.04 mm (n=5), respectively, and also were consistent between plates (CV 1.75% and 2.65%, respectively). = Theoretically, the precipitate ring diameter of the medium and low standards should have achieved 7.25 and 4.13 mm, respectively. Data Analysis Data from experiments 1 and 2 were analyzed by least squares analysis of variance (ANOVA) using the General Linear Model (GLM) of the Statistical Analysis System (SAS, 26 1988). The Student-Newman-Keuls (SNK) method was used to determine significant differences between least squares means (Steel and Torrie, 1960). All main effects and their interactions were included in the analysis. The main effects examined included treatment (BCSL), age of ewes, birth rank, breed of sire and sex of lambs. Main effects (except treatment) and interaction terms having P>0.50 were deleted from the final model. Classifications with few observations (generally less than 8) were either deleted from the model or were combined with other classifications to eliminate the problem of having empty cells which would not generate least square means and standard errors. Categorical data for lamb mortality were analyzed using Chi-Square (Steel and Torrie, 1960). Since IgG levels were significantly influenced by the interval from parturition to sample collection, time of collection was used as a covariate for the colostral IgG analysis. Adjusted lamb birth weights (total weight of twin lambs born in litters s 3) were used as a covariate in the analysis of the weaning weights of the lambs. Since BCS, milk yield and milk composition were measured repeatedly over time, a repeated measures ANOVA model was used (SAS, 1988). Polynomials were used to test for linearity of the time trends during lactation (SAS, 1988). Date of lambing was used as a covariate for the analysis of BCS, milk yield and milk composition to adjust 27 for variation in parturition date. The same procedure was followed for the analysis of lamb weights during lactation (30, 60, 90 and at weaning) but using adjusted lamb birth weight as a covariate. Pearson correlation coefficient analysis was used to measure the degree of relationships between variables. For Experiment 1 these included BCSL, total weight of lambs born, total weight of adjusted lambs born (multiple births reduced to twins), total weight of lambs weaned, colostral IgG concentration, time of colostral IgG collection, age of ewes, birth rank, number of lambs turned out to pasture, number of lambs that died within 12 hr postpartum and number of lambs that died of natural causes from lambing to weaning. In the second experiment, Pearson correlation analyses included BCSL, BCS at 30, 60 and 90 days, average daily milk yield at 30, 60 and 90 days, percentages of milkfat and milk protein at 30, 60 and 90 days, somatic cell count at 30, 60 and 90 days, age of ewes, birth rank, sex of nursed lambs, total weight of adjusted lambs born and lamb weight at 30, 60 and 90 days and at weaning (95 days postpartum). Table 1. Classification of ewes involved in the colostrum (n= 101) and lactation (n= 43) studies. BCSL Colostrum Age of ewe, yr 2.5 3.0 3.5 4 38 35 28 26 20 34 5 6 Breed of Sire Birth rank Polypay Columbia 1 2 i3 21 52 49 8 58 35 7 Sex of lambs Male/ Male Female female Lactation' 30 d PP 27 16 12 6 20 5 25 18 26 17 8 8 27 60 d PP 27 15 12 5 20 5 25 17 25 17 8 8 26 90 d PP 25 15 12 5 18 5 25 15 24 16 7 7 26 1 All ewes adjusted to twins for lactation study. BCSL = Body condition score at lambing. PP = Postpartum. 29 Figure 1. Linear regression curve for sheep immunoglobulin (IgG) concentrations using radial immunodiffusion (Exp. 1) . Figure 1, continued 80 N E E 60 = Y = 12.1020 + 0.0622X R2 = 0.9961 .._...- L. 'E 40 0 a, 00 0 cy) .g. 20 1 200 400 600 800 IgG Concentration 0.1g/rap 1000 31 CHAPTER 4 RESULTS Colostrum Experiment Colostral IgG Unadjusted colostrum IgG levels averaged 57 ± 2.3 mg/ml and ranged from a low of 14 mg/ml to a maximum of 114 mg/ml. Collection of colostrum samples varied from 0 to 11 hr after lambing and averaged 5.9 hr. Colostrum IgG levels decreased significantly with time (Figure 2, r=-0.46) and linear regression analysis projected colostrum levels at 76 mg/ml at parturition (time 0). Using time of collection as a covariate and adjusting for age of ewe (4-7 yr), breed of mating sire and birth rank (twins vs. multiples), colostrum levels were unaffected by BCSL (Table 2). Sex of lambs and associated interaction terms were deleted from the final model. Birth ranks of 4 and 5 were combined with triplets while singles were deleted from the final least squares model since there were only 9, 2 and 8 ewes with quadruplets, quintuplets and singles, respectively. Ewes which gave birth to twins had higher (P=0.06) levels of colostrum IgG than ewes giving birth to multiples. The interaction of age of ewe with BCSL approached significance (P=0.093). 32 Lamb Mortality Total lamb mortality from birth to weaning (day 95) averaged 17.60 (Table 3). Natural death loss was 10.90, including lambs which died before multiple litters were adjusted to twins. Of the lambs that died, the majority (810) of lambs were dead at birth or died within 2-3 hr of birth. Age of ewe, BCSL, mating sire, sex of lamb, and birth rank had no effect on total death loss, although the effect of birth rank approached significance (P<0.10), as death loss was greater in ewes having larger litters at parturition. Predator kill (mostly coyotes) totaled 6.7% from turnout to weaning and was not affected by BCSL, age of ewe, mating sire, sex of lamb or birth rank. Weaning Weight Neither birth rank and its interactions nor other interaction terms (age of ewe x mating sire, age of ewe x BCSL) had an influence on the model (P>0.50) and were deleted from the final least squares analysis. Body condition score at lambing had no effect on total weight weaned (P=0.28), although ewes with a BCSL of 3.0 tended to wean more total lamb weight than ewes with a BCSL of 3.5 (P=0.13)(Table 2). 33 The oldest ewes (7 yr) weaned the lowest total lamb weight whereas the 4- and 6-yr-old ewes weaned the highest total lamb weight (Table 2). Sire of the lambs was important; purebred lambs (Polypay x Polypay) were weaned at a higher total weight than the crossbred lambs (Columbia x Polypay)(P<0.001; Table 2). Lactation Experiment Body Condition Score Using parturition date as a covariate, the final statistical model included all the explanatory variables (BCSL, sex of lambs, age of ewes, mating sire and birth rank), the interaction of BCSL with birth rank, time (30, 60 and 90 days of lactation), and the interactions of time and the explanatory variables. Body condition score differed significantly between the two treatment groups (BCSL 2.5 vs 3.5) at 30 days but were similar at 60 and 90 days of lactation (Table 4), although the 3.5 BCSL group were consistently higher at each time period. The 3.5 BCSL group decreased significantly in body condition score from lambing to the first month (24%) whereas the 2.5 BCSL group only decreased by 1.60. Thereafter, the two groups plateaued from 30 to 60 days and 34 then increased significantly from 60 to 90 days of lactation (Figure 3), yielding a quadratic response (P=0.0001). Age of ewe also had a significant effect on BCS, with the 4-year-old ewes being the lowest at each time period This is probably because 11 of the 12 4-year-old (Table 4). ewes were in the 2.5 BCSL group at lambing. The interaction of time and age of ewe approached significance (P=0.069), due to the 7-year-old ewes increasing faster in BCS during lactation than the other age groups (Figure 4). Ewes giving birth to twins were consistently higher (P=0.015) in BCS during the lactation than ewes with birth rank of a3 (Table 4), although this difference was only significant at 60 days. This is likely because 75% of the ewes having multiple births were in the 2.5 BCSL group at lambing. The interaction of birth rank and BCSL was significant due to the increase in BCS from 30 to 60 days for the ewes giving birth to twins while the ewes having multiple births decreased slightly during this time period (Figure 3). Breed of mating sire had no effect on BCS although ewes bred to Columbia sires tended to have higher BCS's during the first month of lactation (Table 4). Sex of lambs influenced BCS (P=0.106); ewes nursing twin females tended to have a higher BCS at each month of lactation, particularly at the second month, whereas ewes 35 nursing mixed twins (male, female) tended to have a lower BCS at each month of lactation (Table 4). Daily Milk Yield Using parturition date as a covariate, the final least squares model included all the explanatory variables (BCSL, sex of lambs, age of ewes, mating sire and birth rank), the interactions of BCSL with birth rank and with sex of lambs, the interactions of age of ewe with breed of sire and with time (30, 60 and 90 days of lactation), and the interactions of time and the explanatory variables. Milk yield declined linearly (P<0.001) from 30 to 90 days of lactation; the slopes of the regression lines for the two treatment groups (BCSL = 2.5 and 3.5) were similar (Figure 5). Average daily milk yield was similar between ewes of BCSL = 2.5 and ewes of BCSL = 3.5 (Table 5). Following adjustment for explanatory variables, milk secretion was the same for the two treatment groups at 30 days, the 2.5 BCSL ewes were 8% higher (P=0.75) at 60 days, whereas milk secretion was 33% higher (P=0.24) in the 3.5 BCSL ewes at the third month. The interaction of the independent variables and time was not significant, nor were the interactions significant among the independent variables. Although not significant, ewes at 4 yr of age consistently 36 produced less milk each month than the older ewes (5 to 7 Although not significant, ewes nursing twin female yr). lambs secreted more milk than ewes nursing twin male lambs. Analysis of the simple means indicated that ewes with the higher BCSL tended to produce more milk each month (Figure 6). This difference was significant at the third month. Milk Composition (Fat & Protein) Body condition score at lambing had no effect on percent milkfat during the lactation (Table 6). Percent milkfat decreased significantly during lactation, but primarily from the first to the second month (P=0.002 for quadratic effect). Age of ewe had no effect on percent milkfat although the 4-year-old ewes generally had the highest milkfat percentage each month. Although not significant, ewes having twin lambs at birth consistently had higher percent milkfat over the entire lactation than ewes having multiple lambs at birth. Ewes mated to Columbia rams had higher (P.0.047) percent milkfat at 30 days but not at 60 days (P=0.164) or 90 days of lactation (P=0.999). of lambs had no effect on percent milkfat. Sex The large standard errors of the mean precluded detecting differences between the various main effects. 37 Ewes with a higher BCSL (3.5) had slightly higher percent milk protein at 30 and 90 days of lactation, whereas the percent milk protein of ewes with a lower BCSL (2.5) was significantly higher at 60 days of lactation, yielding a significant interaction of time and treatment (P=0.032)(Table 7). Percent protein in milk increased significantly in a quadratic fashion, primarily from 30 to 60 days of lactation (Figure 7). Older ewes tended to have higher percent milk protein during lactation, but this difference was only significant at 90 days and over the entire lactation when the 7-year-old ewes were higher than the 4- and 5-year-old ewes. Ewes giving birth to twin lambs had higher percent milk protein each month of lactation, but this difference was significant only at 90 days. Ewes mated to Polypay rams had slightly higher percent milk protein at 30 days of lactation and this difference increased with time (P=0.056 at 60 days; P=0.005 at 90 days). Ewes nursing twin male lambs tended to have higher percent milk protein during lactation; at 60 days of lactation ewes nursing twin males were higher (P=0.020) than ewes nursing mixed-sex twins. Lamb Weight Total weight born (TWB), adjusted to twin lambs, was analyzed for the main effects and the interactions of BCSL with sex of lambs and with birth rank and the interaction of 38 age of ewe with breed of mating sire. Birth weight of lambs sired by Columbia rams were heavier (P=0.12) than lambs sired by Polypay rams and birth weight of lambs born as twins were heavier (P=0.006) than the adjusted birth weight of twin lambs born as multiple births (3)(Table 8). The interaction of BCSL with birth rank approached significance (P=0.09); total birth weight of twin lambs born from ewes with a lower BCSL (2.5) were heavier than twin lambs from ewes with a higher BCSL (3.5) whereas lambs from multiple births were heavier than twin births from ewes with a higher BCSL (3.5). Using parturition date as a covariate, the final statistical model for total lamb weights included all the explanatory variables (BCSL, sex of lambs, age of ewes, breed of mating sire and birth rank), the interaction of breed of sire with birth rank, time (30, 60 and 90 days of lactation), and the interactions of time and the explanatory variables. Lamb weights were similar at 30 and 60 days but the lambs of ewes having BCSL = 2.5 gained weight more rapidly thereafter and were heavier at 90 days of lactation (P=0.155) and at weaning (P=0.096)(Table 9). Lamb weight increased over time, exhibiting a cubic effect (P=0.005). Body weights of lambs nursing 4-year-old ewes tended to be lower whereas body weights of lambs nursing 7-year-old ewes tended to be higher at each month of lactation. This 39 difference was significant at 30 days and 60 days of lactation. Lamb weights from multiple birth litters tended to be heavier throughout lactation; this difference only was significant (P=0.054) at 30 days of lactation. Lambs sired by Polypay rams were heavier (P=0.02) throughout lactation. This difference increased during lactation, yielding a significant interaction (P=0.01). The interaction of time and treatment was also significant (P=0.003); lambs of ewes with BCSL = 3.5 gained weight more rapidly prior to 60 days whereas lambs of ewes with BCSL = 2.5 gained weight more rapidly from 60 days to weaning at 95 days (Figure 8). The interaction of age of the ewes with time (P=0.084) was due to lambs of the 4-year-old ewes being heavier than lambs of the 5- and 6-year-old ewes at 60 days but not during the other times of lactation. Twin female lambs were heavier (P=0.02) at weaning than twin mixed-sex lambs. The interaction of sex of lambs with time (P=0.007) was caused by heavier weights of the mixed-sex twin lambs than the twin male lambs at 30 days postpartum whereas the twin male lambs were heavier at 90 days of lactation and at weaning. 40 Table 2. Least square means of colostral immunoglobulin (IgG) concentrations and total weaning weight of lambs (0,1,2) at 95 days of age as affected by body condition score at lambing (BCSL), age of ewe, mating sire and birth rank. IgG (mg/ml) Category Weaning weight No. ewes (kg) n x 2.5 37 60 4.1 38 43.5 2.49 3.0 31 56 4.2 35 47.2 2.48 3.5 25 58 5.7 28 41.4 2.88 4 yr 23 57 6.0 26 47.6be 3.06 5 yr 18 66 6.7 20 43.8b 3.24 6 yr 32 55 3.6 34 53.1' 2.56 7 yr 20 53 7.2 21 31.7a 3.50 Polypay 49 59 3.6 52 50.0a 2.03 Columbia 44 57 4.4 49 38.0b 2.17 2 58 61 2.8 z3 35 55 5.3 SE n x SE BCSL Age of ewe Breed of sire Birth rank 'b'Means in column within a category with different superscripts differ (P<0.05). 41 Table 3 Lamb mortality classified by body condition score at lambing (BCSL), age of ewe, mating sire, sex of lambs and birth rank. Lamb mortality Category Born Killed Natural death n n n Total lambs lost n Lamb mortality rate No. lambs weaned BCSL 2.5 76 7 9 16 21.1 60 3.0 66 4 4 8 12.1 58 3.5 51 2 8 10 19.6 41 193 13 21 34 17.6 159 4 yr 50 4 6 10 20.0 40 5 yr 36 1 5 6 16.7 30 6 yr 69 3 5 8 11.6 61 7 yr 38 5 5 10 26.3 28 100 6 9 15 15.0 85 93 7 12 19 20.4 74 88 6 9 15 17.0 73 105 7 12 19 18.1 86 1 8 1 0 1 12.5 7 2 109 7 8 15 13.8 94 3 51 3 7 10 19.6 41 4 19 1 4 5 26.3 14 5 6 1 2 3 50.0 3 Total Age of ewe Ram breed Polypay Columbia Sex of lambs male female Birth rank Table 4. Least square means of body condition score as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. Days post partum Category 30 60 _ x 90 _ x Overall mean n x 2.5 26 2.46a .056 26 2.56 .062 26 3.10 .067 2.71 .041 3.5 15 2.666 .082 15 2.68 .090 15 3.15 .097 2.83 .059 12 2.92a .094 2.61' .057 3.14'b .132 2.89b .080 3.27b .074 2.796 .045 3.19'6 .131 2.806 .080 SE n SE n SE SE BCSL Age of ewe 4 yr 2.50ab .079 12 2.40a .087 12 5 yr 5 2.75' .110 5 2.776 .121 5 6 yr 19 2.49b .062 19 2.60'b .068 19 2.50'b .109 5 2.70b .120 5 25 2.60 .058 25 2.75a .064 25 3.19 .070 2.85 .042 16 2.52 .073 16 2.50b .080 16 3.07 .087 2.69 .053 25 2.50 .054 25 2.56 .060 25 3.08 .065 2.75 .039 16 2.62 .074 16 2.54 .082 16 3.04 .089 2.79 .054 Males 7 2.60 .086 7 2.62'd .094 7 3.13 .102 2.79 Females .062 7 2.59 .099 7 2.73' .108 7 3.19 .118 2.84 .072 27 2.48 .060 27 2.50d .065 27 3.06 .071 2.68 .043 7 yr Birth rank 2 3 Ram breed Polypay Columbia Sex of 5 lambs Male\Female Overall mean 2.56 2.62 .052 .057 3.13 ''b Means in column within a category with different superscripts differ (P<0.05) cd Means in column within a category with different superscripts differ (P=0.06) .062 I. NJ Table 5. Least square means of milk yield (ml) as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs Days postpartum Category __30 n 60 90 Overall mean x SE n x SE 2385 2209 304.4 25 1347 207.6 465.9 15 1794 317.8 2331 2421 2127 2352 2369 269.6 n x SE SE BCSL 2.5 25 3262 369.1 25 3.5 15 3259 565.0 15 12 2921 515.8 12 1911 425.4 12 1547 5 3610 745.4 5 2159 614.7 5 1287 18 3328 417.0 18 2050 343.9 18 1729 290.1 419.3 234.6 5 3182 639.5 5 3069 527.4 5 1721 359.8 2657 334.3 2 24 3185 397.6 24 2554 1751 223.6 3335 485.4 16 2041 16 1391 273.0 2497 2256 207.8 16 327.9 400.3 24 a3 3239 366.0 25 2110 301.9 25 1557 521.4 15 2485 430.0 15 1584 205.9 293.3 2302 3282 2450 191.3 272.5 2987 547.1 7 2195 451.2 7 1577 307.7 2253 285.9 3662 622.2 7 2014 513.2 7 1602 350.0 2426 325.2 3132 370.6 26 2683 305.7 26 1534 208.5 2450 193.7 3260 342_3 2297 282_3 1571 392 6 Age of ewe 4 yr 5 yr 6 yr 7 yr Birth rank Ram breed Polypay 25 Columbia 15 Sex of lambs Males 7 Females 7 Male\ 26 Female Overall mean 192.9 295.3 389.6 217.9 253.7 Table 6. Least square means of percent fat in milk as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. Days postpartum Category 30 n _ x 60 SE n _ x 90 SE n _ x SE Overall mean SE BCSL 2.5 25 11.50 .617 25 9.24 .707 25 9.08 .676 3.5 9.94 .550 14 11.90 .652 14 9.54 .747 14 8.29 .714 9.91 .581 11 12.07 .641 11 9.94 .734 11 9.18 .702 10.39 5 yr .571 5 12.03 1.075 5 9.44 1.231 5 7.82 1.178 6 yr 7 yr 9.76 .957 18 11.06 .524 18 9.34 .601 18 9.26 .574 9.89 .467 5 11.64 1.105 5 8.84 1.266 5 8.48 1.211 9.66 .984 23 12.06 .498 23 9.80 .570 23 9.21 .545 10.36 .443 16 11.34 .777 16 8.97 .889 16 8.16 .851 9.49 .692 23 10.99' .487 23 8.83 .558 23 8.69 .533 9.50 .434 16 12.41b .674 16 9.95 .772 16 8.69 .738 10.35 .600 7 12.04 .964 7 10.02 1.104 7 9.42 1.056 10.49 .859 7 12.12 .781 7 9.30 .895 7 8.98 10.14 .696 9.14 .505 Age of ewe 4 yr Birth rank' 2 3 Ram breed Polypay Columbia Sex of lambs Males Females .856 Male\Female 25 10.93 .568 25 8.84 .650 25 .622 7.65 Overall 11.71 .481 9.38 .550 8.67 .526 a'b Means in column within a category with different superscripts differ (P<0.05) c (P=0.016) Table 7. Least square means of percent protein in milk as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs Days postpartum Category 30 60 n x SE n BCSL 2.5 25 4.00 .080 25 3.5 14 4.05 .102 14 11 3.92 .104 5 3.98 18 Age of ewe 4 yr 5 yr 6 yr 7 yr Birth rank 2 3 x 90 SE Overall mean SE 5.22 .097 4.71 .068 5.29 .124 4.66 .087 11 5.01' .125 4.53a .088 5 5.02' .172 4.57.1° .120 5.30°b .104 4.74ab .073 5 5.68b .198 4.90b .139 .092 23 5.41° .109 4.78 .076 .095 16 5.10b .113 4.59 .079 .100 4.83' .070 .124 4.54b .087 .160 4.81 .112 .169 4.64 .118 .097 4.61 .068 SE n x 4.918 .081 25 4.64b .104 14 11 4.64 .105 .142 5 4.73 .144 4.09 .086 18 4.83 .087 18 5 4.11 .164 5 4.89 .166 23 4.08 .090 23 4.85 16 3.97 .093 16 4.70 Ram breed Polypay 23 4.10 .083 23 4.90c .084 23 5.49° Columbia 16 3.95 .103 16 4.64d .104 16 5.01b Sex of lambs Males 7 4.12 .133 7 5.00a .135 7 5.32 Females 7 3.97 .140 7 4.71ab .142 7 5.23 Male\ Female 25 3.99 .080 25 4.62b .081 25 5.21 Overall mean 4,02 .068 4.78 .069 5.25 Means in column within a category with different superscripts differ (P<0.05) Means in column within a category with different superscripts differ (P=0.056) .082 46 Table 8. Least square means of total weight born (twin lambs) as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. Category Total weight born, kg n x SE 2.5 27 8.3 .35 3.5 16 9.2 .53 4 yr 12 8.1 .49 5 yr 6 8.8 .62 6 yr 20 8.7 .36 7 yr 5 9.3 .58 2 26 9.5. a3 17 8.0b .45 Polypay 25 8.3 .36 Columbia 18 9.2 .47 Males 8 8.7 .49 Females 7 9.0 .61 28 8.6 .33 BCSL Age of ewe Birth rank .35 Ram breed Sex of lambs Male\ Female a.b Means in column within a category with different superscripts differ (P<0.05) Table 9. Least square means of total lamb weights' (kg) at 30, 60 and 90 days postpartum and at weaning (95 days) as affected by body condition score at lambing (BCSL), age of ewe, birth rank, mating sire and sex of suckling (twin) lambs. n x Days postpartum rn qn Category SE n x 90 SE n x SE n x SE BCSL 2.5 21 21.9 .83 21 36.1 1.25 21 55.2 1.52 21 60.3 1.92 3.5 13 22.4 .76 13 36.6 1.14 13 52.3 1.39 13 55.9 1.76 10 20.7b .78 10 35.68b 1.17 10 52.0 1.44 10 55.5 1.81 5 21.2b 1.06 5 35.08b 1.60 5 54.7 1.95 5 58.6 2.47 17 20.9b .60 17 34.5b .91 17 52.4 1.11 17 58.8 1.41 2 25.88 1.61 2 40.28 2.42 2 55.8 2.96 2 59.6 3.75 21 21.1' .68 21 35.6 1.02 21 53.3 1.25 21 58.6 1.58 13 23.2d .86 13 37.0 1.30 13 54.1 1.58 13 57.6 2.01 20 22.9 .73 20 37.5 1.09 20 55.78 1.34 20 60.9' 1.69 14 55.46 1.70 4 59.7'1' 3.05 6 60.1° 2.25 24 54.56 1.58 Age of ewe 4 yr 5 yr 6 yr 7 yr Birth rank 2 3 Ram breed Polypay Columbia 14 21.5 .73 14 35.2 1.10 14 51.76 1.34 Sex of lambs Males 4 20.5 1.31 4 35.3 1.97 4 54.8 2.41 Females 6 23.4 .97 6 38.3 1.46 6 55.0 1.78 Male\ 24 22.6 .68 24 35.3 1.02 24 51.3 1.25 Female ab Means in column within a category with different superscripts differ (P<0.05) Means in column within a category with different superscripts differ (P=0.054) Only ewes that weaned twin lambs are included in the analysis. ' Table 10. Correlation coefficients for ewe and lamb characteristics (Exp. 1). TWBF BCSL -.25* TWBF TWBA TWBA -.05 .42" WWT HR AGE RANK TRN DL12 -.02 .01 .22* -.25* -.27" .08 -.08 .26" .08 -.09 .07 .44" .13 .02 .32" .09 -.17 .34" -.16 .34" -.21* -.03 -.14 -.07 .00 .08 .64" -.36" -.33 -.02 .08 -.04 .16' -.02 .07 -.07 -.16 .09 -.05 .02 .04 -.11 .29 ** .33" .07 -.37" .01 -.04 WWT IgG -.46" HR AGE .75" -.17a RANK TRN DL12 P<0.01; IgG DED -.07 P<0.05; a P<0.10 BCSL = Body condition score at lambing; TWBF = unadjusted total birth weight of lambs; TWBA adjusted total birth weight of lambs; WWT = total weaning weight of lambs; IgG = immunoglobulin concentration; HR = time from lambing to colostrum collection; AGE = age of the ewes; RANK = birth rank; TRN = number of lambs turned out to pasture; DL12 = number of lambs died before bumming; DED = number of lambs died after turned out to pasture. 49 Table 11. Correlation coefficients of some ewe and lamb characteristics during lactation (Exp. 2). BCSL BCSI BCS2 LMB WTI LMB WT2 LMB WWT .19 35* .36* .24 .16 -.28* .31* .20 .06 -.01 -.29' -.29* -.11 31* 34* .40* .29a .04 .36* -.10 -.12 .25 .42 ** A5** 39* .26 .13 .10 .08 -.06 .06 .20 31' .31" .29" -.29' -.26' .21 -.23 .21 .26 31* .26 28* .28' .22 -.11 -.08 .13 -.14 -.09 .21 .30' 34* .21 .21 .07 .43** .21 33* .25 -.23 -.10 .19 32* 32" .22 -.10 -.15 .12 .12 .12 .03 .21 -.05 -29' -.03 -.12 -.12 .01 .20 .00 .01 .55** .08 .464.4, _Spit. ....23 -.18 35* .37* 32 .28 .16 .55** A3** .18 .18 -.17 -.10 -.27* -.13 .20 -.13 -.15 -.14 -.03 .724* .14 .16 .10 -.04 -.27" -.08 35* .02 -.10 -.12 -.08 .08 .14 .19 .00 -.23 -.11 .26' -.09 -.18 -.16 -.03 .49** .47** .40** -.38* .03 .33* .48** .56** 39* .27 .30" -.02 .02 -.13 -.04 .07 A5** .32' .25 .22 -.09 -.02 .17 .19 .10 -.07 -.20 -.19 -.01 .43** .48** .30` .27 32' -.24 -.63- -.38* -.26 -.32' -.25 .14 .11 -.05 -.07 -.06 .71" .57" .63" .84" .88" 37* BCS1 BCS2 BCS3 MKY1 MKY2 MKY3 PRTI PRT2 PRT3 FATI FAT2 FAT3 SCC1 SCC2 scC3 AGE RANK SIX TWB .44** .42** .40** .24 .09 .41** .28" -.21 .20 -.03 .03 -.02 .14 -.04 .19 .25" -.23 -.21 .62** 37* -.03 .03 .42** .24 -.22 .02 .12 .22 .11 .10 .08 .19 .02 -.21 .61** .10 .06 .38* .43** -.28* .19 .11 .19 -.00 -52** .20 .22 .294 .20 .03 36* .42** -.06 .35* .04 .06 -.02 .51** .22 .25 .15 -.04 .21 -.12 -.06 .01 .22 .17 .03 .18 .22 -.26' -.41** -.17 -.02 .11 -.14 -.10 .02 -.27' -.20 .22 .05 .14 .14 .48** -.15 .13 .48** .09 -.11 BCS3 MKY I MKY2 MKY3 PRT1 PRT2 PRT3 FATI FAT2 FAT3 SCC1 SCC2 SCC3 AGE RANK SEX TWB LMBWT1 LMBWT2 LMBWT3 .91** wT3 .61** .72** .84** ** P<0.01; * P<0.05; a P<0.10 BCSL = Body condition scores at lambing; BCS1 =-first month; BCS2 = second month; BCS3 = third month of lactation; MKY1 = average daily milk yield at first month; MKY2 =-second month; MKY3 = third month of lactation; PRT1 = percentages of protein in milk at first month; PRT2 = second month; PRT3 = third month of lactation (PRT3); FAT' = percentages of fat in milk at first month; FAT2 = second month oflactation; FAT3 = third month; SCC1 = somatic cell count at first month; SCC2 = second month; SCC3 = third month of lactation; AGE = age of ewes; RANK = birth rank; SEX = sex of the nursed (twin) lambs; TWB = total birth weight of the nursed (twin) lambs; LMBVsT1 = total (twin) lamb weight at first month; LMBWT2 = second month; Ll'ABWT3 = and third month of lactation; WWT = total weight of the weaned lambs (twin). 50 Figure 2. Linear regression of ewe colostrum immunoglobulin (IgG) concentrations on time of collection after parturition (Exp. 1). Figure 3. Least square means of body condition score during lactation of ewes of different body condition scores at lambing (BCSL) and of different birth rank at lambing (Exp. 2). 51 Figure 2 120 , E cm E CD cm 1-4 100 80 60 40 20 2 8 6 Time Postpartum (Hr) 4 12 10 Figure 3 3.5 o 2.5 BCSL 3.5 BCSL Birth rank=2 a) L_ 0 (.) v) c 0 Birth rank3 3.0 43 0c 00 >, -0 2.5 0 cc) 2.0 30 60 Days Postpartum 90 52 Figure 4. Least square means of body condition score during lactation of ewes of different ages at parturition (Exp. 2). Figure 5. Linear regression of least square means of daily milk yield of ewes of different body condition score at lambing (BCSL) (Exp. 2). 53 Figure 4 3.5 O 4 years of age A 5 years of age O * 6 years of age o 7 years of age 0 U) 3.0 2.5 2.0 30 60 Days Postpartum 90 Figure 5 + ...--. L 6000 x Y = 4181 3.5 BCSL 27.4X R2 = 0.3273 -0 0 2.5 BCSL Y = 3726 27.7X R2 = 0.3239 4000 _.v --±' >-. ° 2000 a__ oo 30 60 Days Postpartum 90 54 Figure 6. Linear regression of actual daily milk yield of ewes of different body condition score at lambing (BCSL) (Exp. 2). Figure 7. Least square means of percent fat and protein in milk during lactation of ewes of different body condition score at lambing (BCSL)(Exp. 2). 55 Figure 2.5 BCSL E Y = 3726 27.7X R2 = 0.3239 6000 x 3.5 BCSL Y = 4181 - 27.4X R2 = 0.3273 -o a) >- 4000 _y >, (Is 2000 0 30 Figure 12 10 I 60 Days Postpartum 90 O 2.5 BCSL 35 BCSL Milk Fat a) 0 a) CL Milk Protein 30 _a 60 Days Postpartum 90 56 Figure 8. Least square means of total lamb weight during lactation of ewes of different body condition score at lambing (BCSL)(Exp. 2). 57 Figure 8, continued. 60 50 40 30 20 10 30 60 Days Postpartum 90 W 58 CHAPTER 5 DISCUSSION Colostrum Experiment Colostrum The concentration of immunoglobulins in colostrum decreases rapidly with time. Correlation coefficient analysis indicated that colostral IgG concentration was negatively correlated to time of sample collection (r= 0.46; P<0.01; Table 10). Linear regression analysis of the data, collected from 0 to 11 hr postpartum, projected a level of 76 mg/ml at parturition (Figure 2), and with a slope of -3.28, colostrum IgG levels would decrease to 0 at about 23 hr postpartum. Shubber et al. (1979) collected colostrum over a 48 hr period and observed that concentrations of total immunoglobulins (IgG, IgM, IgA) had decreased to low levels by 36 hr postpartum. A study of dairy cattle (Stott et al., 1981) over the first 6 milkings (12 hr intervals) postpartum also indicated a linear decline (semilog plot) in milk IgG concentration with time (slope -0.293). Although the unadjusted concentration of colostral IgG (57 ± 2.3 mg/ml) is not directly comparable since samples were collected up to 11 hr postpartum, they compare 59 favorably with Thomas et al. (1988) and Gilbert et al. (1988) who collected colostrum at parturition and reported concentrations of IgG similar to mine (64 ± 7.6 and 80 ± 2.4 mg/ml, respectively). Hunter et al. (1977) also collected colostrum at parturition but observed IgG concentrations 2- fold higher (115 ± 10.1 mg/ml) than mine and, similar to my 8-fold range between individual ewes (14-114 mg/ml), observed a 6-fold range (43-260 mg/ml) between ewes. Shubber et al. (1979) reported mean levels of total immunoglobulins ranging from 71 ± 11 to 203 ± 28 mg/ml in colostrum collected from 1 to 12 hr after parturition. Different reagents for the radial immunodiffusion assay and the variability between ewes probably account for the major differences between studies. I did not observe differences in colostral IgG concentrations due to BCSL, age of ewe, breed of mating sire or birth rank (Table 2). Similarly, Thomas et al. (1988), in a study involving 3-yr-old Finn-Targhee ewes, failed to detect differences in colostral IgG concentrations of ewes having BCSL of 2.5 or 3.5 (64 ± 7.6 vs 67 ± 7.5 mg/ml). Although Thomas et al. (1988) (twins vs triplets) and Halliday (1974)(singles vs twins) did not observe an effect of birth rank, Gilbert et al. (1988) reported a linear increase in colostral IgG concentrations as birth rank increased (singles, twins and triplets). I observed an 60 opposite trend in our study; colostral IgG concentrations were lower (P =0.06) in ewes having multiple lambs (3) than in ewes having twins. However, our analysis did not include singles and all multiple litters were pooled. In a large study (N =1686) involving several breeds and ages of ewes, Gilbert et al. (1988) found that yearling ewes had concentrations of colostral IgG (100 ± 7.3 mg/ml) significantly higher than older ewes (2-7 yr) but found no difference due to age among the older ewes. Gilbert et al. (1988) also observed higher concentrations of colostral IgG in Polypay ewes (80 ± 2.4 mg/ml) than in Columbia ewes (64 ± 2.1 mg/ml); I did not observe any difference (P>0.10) between Polypay ewes mated to Polypay rams (59 ± 3.6 mg/ml) and Polypay ewes mated to Colombia rams (57 ± 4.4 mg/ml). Lamb Mortality Lamb mortality was not affected by BCSL, age of ewe, breed of mating sire, sex of lamb, or birth rank (Table 3). The majority (81%) of the natural death loss (10.9%) occurred at birth (stillborn) or within 2-3 hr after birth. Predator kill was 6.7% which, when added to the natural death loss, yielded a total death loss of 17.6% from birth to weaning. Gilbert et al. (1988) reported a death loss of 12.3% and found that 52% of the deaths resulted from 61 infectious diseases. They observed differences due to breed (higher in Targhee, lower in Polypay and Finn) and found that death loss was 3-4x greater in lambs having serum IgG levels slO mg/ml. from 7.71%. 1992). Others have reported death losses ranging (Young and Dickerson, 1987) to 15.796 (Otesile, Previous studies at Oregon State University (Khan, 1994; Nawaz and Meyer, 1992; Nawaz et al., 1992), observed differences in lamb survival due to the effect of breed in some, but not all, trials. They also reported a positive relationship between body condition of the ewe and lamb survival. Our narrow range of BCS's (2.5 to 3.5) may have precluded us from observing this effect. Kleemann et al. (1993), however, reported that lamb survival was not influenced by the nutritional status of the ewe during mid and late pregnancy. A negative relationship (r= -0.21; P<0.05) was observed between early death loss (DL12) and adjusted total birth weight (TWBA), indicating that neonatal death loss was lower in lambs having a heavier birthweight (Table 10). Similar findings by others (Alexander, 1964; Mellor, 1990; Jordan and Feuvre, 1989) have been attributed to the failure of smaller birthweight lambs to compete successfully for milk. The positive correlation (r=0.13; P>0.10) between neonatal death loss (DL12) and unadjusted total birth weight (TWBF) was expected since the smaller birthweight lambs were in the 62 larger birth ranks. This is reflected in the large positive correlation (r=0.33; P<0.01) between early death loss (DL12) and birth rank (RANK). The negative correlation (r= -0.33; P<0.01) between death loss in pasture (DED)(largely predator kill) and total weaning weight (WWT) can be attributed to the higher weaning weight of surviving lambs. The negative correlation (r= -0.25; P<0.05) between BCSL and unadjusted total birth weight (TWBF) can be attributed to the negative relationship between BCSL and birth rank (RANK)(r. -0.25; P<0.05); ewes with higher BCSL had a lower birth rank, yielding a lower TWBF (Table 10). The correlations between the number of lambs turned out to pasture at 2-3 weeks postpartum (TRN) and birth weight (TWBF, TWBA) and weaning weight (WWT) were positive and of the opposite sign of early death loss (DL12) as expected, since TRN and DL12 are opposites. Heavier birth weight lambs have a lower death loss. Lactation Experiment Body Condition Score Body condition score of ewes with a higher BCSL (3.5) decreased significantly (240) from lambing to first milking (30 days postpartum) whereas the BCS of ewes with a lower 63 BCSL (2.5) only decreased by 1.6% during this interval (Table 4). Peart (1970) observed even a greater loss in condition score (390) in ewes with a high BCSL (3.1) whereas ewes with a low BCSL (1.1) did not decline in BCS. Similarly, Gibb and Treacher (1980, 1982) observed that ewes with a higher BCSL (3.2) lost condition more rapidly early in lactation (first 10 weeks) than ewes with a lower BCSL (2.8), but that by 16 weeks postpartum, the condition loss was the same between the two groups. Since we failed to detect differences in milk yield at 30 days of lactation between the 2 treatment groups (Table 5), the greater loss of BCS in ewes with the higher BCSL (3.5) was likely due to decreased DMI, since Cowan et al. (1980) has shown that DMI decreases as body fatness increases. After the initial decrease in BCS from parturition to 30 days, the BCS increased in both treatment groups from 30 to 90 days postpartum (Figure 3), although ewes with the lower BCSL (2.5) gained body condition more rapidly during lactation and were only 1.6% lower than the high BCSL group by 90 days of lactation. Although not significant, we found that ewes with a higher birth rank had a lower BCSL (r= -0.23; Table 11), indicating that ewes with a higher birth rank had a larger demand for nutrients by the fetuses during pregnancy, resulting in a lower BCSL. This is similar to the 64 significant correlation (r= -0.25; P<0.01) we observed in Experiment 1 (Table 10) between these two factors. Similarly, Khan (1994) observed a significant decrease in birth rank in ewes having a higher BCSL in 1 of 7 trials he conducted. The positive correlations between BCSL and the BCS's at 30, 60 and 90 days and between the BCS's at those times (Table 11) indicates that the scoring was repeatable when performed each time by the same individual. Milk Yield The lactation period in sheep varies in length, depending on the environment and management. Lambs may be weaned as early as 40 days or as late as 130 days postpartum. In this study, since adequate forage was available, we chose to wean the lambs at 95 days postpartum. Milk production normally peaks at 3-4 weeks postpartum and then exhibits a linear decline during the remainder of the lactation (Peart, 1967). example, observed Reynolds and Brown (1991), for peak daily milk yield (2,459 ml/day) at 2-4 weeks of lactation and then a decline to 1,142 ml/day at 106 days of lactation. (Since many of the studies report milk yield in units of grams, we have converted these data to milliliters using a factor of 1 ml = 1.035 g (Torres- 65 Hernandez and Hohenboken, 1979). Ramsey et al. (1994) reported peak milk yield at 14 days of lactation. In their study, ewes nursing twins produced more milk (3,108 and 4,176 ml/day) than ewes nursing singles (2,136 and 2,664 ml/day) on days 4 and 14, but not on days 50 (2,352 vs 2,112 ml/day), 89 (1,260 vs 1,584 ml/day) or 106 (1,056 vs 1,056 ml/day, respectively) of lactation. The objective of this study was not to determine peak milk production but to begin the milking study after the peak of production (day 30) so that we could study the rate of decline in milk yield during the lactation. Daily milk yield at 30 days of lactation (3,260 ± 342 ml)(Table 5) was higher than that reported by Reynolds and Brown (1991) at 30 days (2,376 ± 130 ml), Shabbir et al. (1992) at 30 days (2,204 ml), Torres-Hernandez and Hohenboken (1979) at 21 days (1,795 ml), Brown and Hogue (1985) at 28 days (1,414 ml) and Langlands (1973) at 31 days (1,728 ml). Others, however, have reported milk yields slightly higher (Snowder and Glimp, 1991 3,434 ml at 28 days; Ramsey et al., 1994 3,336 at 14 days) than ours. Our estimate of total milk yield (231 1) for the 95-day lactation (2.43 l/d) is higher than the estimates of 189 1 in 89 d (2.12 l/d) by Gardner and Hogue (1964), 142 1 in 70 d (2.03 1/d) by Snowder and Glimp (1991), 129 1 in 70 days (1.84 l/d) by Peart (1970), and 137 1 in 70 d (1.96 l/d) by 66 Peart (1967). Our estimate is based on a linear regression equation (Y = 4035 -30.1607X) calculated from the overall least square means (Table 5). An examination of the milk yield data (measured over a comparable time period) from other experiments revealed that they observed a similar rate of decline (negative regression slope) in milk yield (Langlands, 1973, b= -18.9; Snowder and Glimp, 1991, b= -36.7; Reynolds and Brown, 1991, b= -16.6; Torres-Hernandez and Hohenboken, 1979, b= -15.1) as we found (b= -30.2). Simultaneous with the decline in milk yield, Langlands (1973) reported a linear increase in grass consumption with age of the lambs. Geenty (1985) observed that milk production is a good predictor of lamb growth up to 4 weeks of age, after which grass consumption becomes an important variable. We did not observe differences in milk yield due to BCSL, age of ewe, birth rank, breed of mating sire or sex of lambs (Table 5). The adjusted milk yields were the same the first month but the ewes having the higher BCSL (3.5) produced 33% more milk (P=0.24) at 90 days of lactation. An analysis of the simple means indicated that the ewes having the higher BCSL produced more milk each month, although the difference was only significant (P<0.05) at 90 days of lactation (Figure 6). Gibb and Treacher (1980), in agreement with our data, also observed a slight advantage in 67 milk yield in ewes having a higher BCSL (3.2 vs 2.4), and this difference was only significant later in lactation (weeks 9-12). Others have failed to observe a difference in milk yield between ewes of different BCSL (Jordan and Hanke, 1991; Peart, 1968; Gibb and Treacher, 1982). Peart (1967) reported that ewes, severely undernourished during pregnancy, produced less milk the first week of lactation, but produced similar amounts of milk at the peak of lactation and for the entire lactation as ewes that were well nourished during pregnancy. In a later study, Peart (1970) found that milk yield was influenced by BCSL but only when the forage supply was restricted. Since our ewes were adjusted to twin lambs shortly after parturition, we failed to note the increased milk yield in ewes nursing twins compared to ewes nursing singles as reported by Gardner and Hogue (1964), Ramsey et al. (1994), Slen et al. (1963), Snowder and Glimp (1991), Torres-Hernandez and Hohenboken (1979), and Gibb and Treacher (1982). Torres-Hernandez and Hohenboken (1979) observed that age of ewe (3- vs 4-yr-old) had no effect on milk yield and, in a later study (Torres-Hernandez and Hohenboken, 1980), they found that sex of lamb was not a factor. Birth rank was not a factor influencing milk yield in a study by Knight and Bencini (1993). Snowder and Glimp (1991) compared the 68 Rambouillet, Columbia, Polypay, and Suffolk breeds and concluded that milk yield was not affected by breed of ewe. Torres-Hernandez and Hohenboken (1979) reported a negative correlation (r= -0.25; P>0.10) between BCS at the end of lactation and total milk production. Our correlations were positive between milk yield and BCS at 90 days of lactation (r=0.36; P<0.05) and between BCSL and milk yield at 30 (r=0.24; P>0.10), 60 (r=0.09; P>0.10), and 90 days (r=0.41; P<0.01) of lactation (Table 11). The significant correlation at 90 days of lactation is in agreement with our analysis of simple means for milk yield in which ewes with the higher BCSL (3.5) produced more milk each month but the difference was only significant at 90 days of lactation (Figure 6). We failed to detect significant correlations between milk yield and lamb growth. The correlations ranged from r=0.20 at day 30 to r=0.26 at day 60 and r=0.21 at day 90 (P>0.10) of lactation (Table 11). Others have noted significant correlations between these two factors but primarily early in lactation (Geenty, 1980; Geenty et al., 1985; Reynolds and Brown, 1991; Snowder and Glimp, 1991) and primarily in ewes nursing single lambs (Torres-Hernandez and Hohenboken, 1980; Ramsey et al., 1994). Brown et al. (1987) observed that milk yield was only a good predictor of lamb growth until the third or fourth week of lactation. Gardner 69 and Hogue (1964), however, reported a correlation coefficient of 0.83 between total milk yield and 90-day lamb weights. A decline in these correlations during lactation was not observed in my experiment because milk yield and lamb weights were not measured until 30 days of lactation, when the lambs had already begun supplementing their diets with forage. Ramsey et al. (1994) stated that milk yield is a more limiting factor for twins whereas singles have ad lib access to milk which can express itself more directly in growth of lambs. Langlands (1973) reported that lambs increase forage intake to compensate for reduced milk supply, thus maintaining an equivalent growth rate. Milk Composition Milkfat percentage decreased from 11.71 ± 0.481% at 30 days to 8.67 ± 0.526% at 90 days of lactation (Table 6), averaging 9.92% during the entire lactation. These levels are considerably higher than 6.48% reported by Brown and Hogue (1985), 6.99% reported by Brown et al. (1987), and 6.65% reported by Torres-Hernandez and Hohenboken (1979). Our milkfat percentages would change only slightly (Corrected fat% = -1.09 + 1.11(IRMA %fat)) if corrected for infrared analysis as suggested by Reynolds and Brown (1991). Percent milkfat was not significantly correlated with milk 70 yield; some correlations were positive and some negative (Table 11). In contrast to the decrease in percent milkfat that we found, Torres-Hernandez and Hohenboken (1979) observed an increase in percent milkfat during the lactation, but they only reported values at 8 (7.13%) weeks of lactation. Shabbir et al. (6.17%) and 12 (1992), in a study involving various levels of dietary calcium, reported milkfat percentages (11.6%) as high as ours in some of their dietary groups, and a decrease in milkfat percentage in some dietary groups from the first to the second month of lactation. Milk protein, however, increased during lactation from 4.02 ± 0.068% at 30 days to 5.25 ± 0.082% at 90 days (Table 7). 1985 Others have reported similar levels (Brown and Hogue, 4.49%; Brown et al., 1987 Brown, 1991 4.81%; Reynolds and 4.51%; Torres-Hernandez and Hohenboken, 1979 4.90 %) during lactation. Our results agree with Torres- Hernandez and Hohenboken (1979) who reported an increase in percent milk protein during the lactation. Snowder and Glimp (1991) found that milk protein percentage was high immediately after parturition, decreased to a low at 8 weeks, and then increased again at 12 and 14 weeks of lactation. In our study, ewes mated to Polypay rams had a higher milk protein percentage (4.83 ± 0.070%) than ewes mated to Columbia rams (4.54 ± 0.087%); Snowder and Glimp 71 (1991) did not detect differences due to breed of the ewe's sire. Torres-Hernandez and Hohenboken (1979) observed a significant effect of the breed of the ewe's sire on percentage milk protein but not on the percentage milkfat. Also in our study, milk protein percentage was higher in 7yr-old ewes than in 4-yr-old ewes at 90 days and over the entire lactation; Torres-Hernandez and Hohenboken (1979) found no difference in milk composition between 3- and 4-yrold ewes. There was a slight positive relationship between percentage milkfat and milk yield each month, whereas percentage milk protein was negatively correlated with milk yield at 60 (r= -0.41, P<0.01) and 90 days (r= -0.20, P>0.10) of lactation (Table 11). Somatic cell count (SCC) may be an indication of an inflammatory response in the udder, but it is not a direct indicator of mastitis. In dairy cattle, there is a negative regression of SCC on milk yield. We observed a negative correlation (r= -0.29; P<0.10) between milk yield and SCC at 60 days, but not at 30 or 90 days of lactation (Table 11). The positive correlations between the SCC at 30 days with the SCC at 60 days (r=0.49, P<0.01) and 90 days (r=0.47, P<0.01) of lactation indicates that ewes with a high SCC at 30 days tended to have high SCC's at 60 and 90 days of lactation. Older ewes had higher SCC at 30 days (r=0.40; P<0.01) and 90 days (r=0.22; P>0.10) but not at 60 days (r= 72 -0.02; P>0.10) of lactation. Torres-Hernandez and Hohenboken (1979) observed decreased milk yield (11.5%) in ewes with mastitis in half of the udder and a larger decrease (58.3%) in milk yield in ewes with mastitis in both halves of the udder. They reported an overall mastitis incidence of 11.5% in their study. Lamb Weights There was no effect of BCSL on birth weight of lambs born (adjusted to twins), although birth weights were slightly heavier (P =0.15) for ewes with a higher BCSL (Table 8). Similarly, Gibb and Treacher (1980, 1982) found no difference in lamb birth weights in their studies, which compared ewes of high (3.2-3.3) and low (2.4-2.6) BCSL. Peart (1968) and Khan (1994) found the same relationships whereas Russel et al. (1977) observed that birth weights declined when ewes were undernourished during gestation. Kleemann et al. (1993), however, found that birth weights were unaltered by the level of nutrition during mid and late pregnancy. Thomas et al. (1988) reported that lamb birth weight was greater (P<0.10) for ewes with a higher BCSL (3.5) than for ewes with a lower BCSL (2.5). But ewes in that experiment had similar birth ranks in both treatment groups whereas, in our study, ewes with a lower BCSL had a 73 higher birth rank (r= -0.23; P>0.10; Table 11). This relationship was significant in the first experiment (r= 0.25; Table 10). Older ewes had fewer (r= -0.19; P>0.10) but heavier lambs at birth (r=0.43; P<0.01; Table 11). High birth ranks consisted of lambs with lower birth weights (r= -0.63; P<0.01), similar to the results reported by Kleemann et al. (1993). Nawaz and Meyer (1992a) observed a curvilinear effect of ewe age on lamb birth weight; birth weights were the lowest in yearling ewes. In my experiment, lambs sired by Columbia rams tended to be heavier (P>0.10) at birth than those from ewes mated to Polypay rams (Table 8), but, during lactation and at weaning, lambs sired by Polypay rams were heavier. This difference was significant at 90 days and at weaning. Nawaz and Meyer (1992) observed that lambs were smaller at birth from Polypay- and Coopworth-sired ewes than lambs from Suffolk-sired ewes. Ewes with multiple births (3) had lighter lambs at birth than ewes with twins (Table 8). Since all litters were adjusted at birth to twins, litters of twins born as twins would be heavier than twins selected from multiple-birth litters. Neither growth of lambs nor weaning weights were affected by BCSL (Table 9). This is in agreement with Gibb and Treacher (1982) who reported that the nutritional status 74 of ewes during pregnancy had no affect on the growth rate of lambs. Khan (1994), however, who had a wider range of BCSL (2.5-4.0), reported that ewes with a higher BCSL weaned heavier lambs. Our lack of differences due to BCSL is not surprising, given that milk yield did not differ between treatment goups. Although only significant at 30 and 60 days of lactation, lambs of 7-yr-old ewes were heavier at birth (Table 8) and at each period of lactation and at weaning (Table 9). Nawaz and Meyer (1992) reported that yearling ewes weaned the lightest lambs and that weaning weight increased with parity. Similar to birth weights, they also found that weaning weights from Polypay-sired ewes were intermediate between Suffolk- and Coopworth-sired ewes. We observed that twin female lambs were heavier than twin male lambs at 30, 60, and 90 days of lactation, whereas twin males were heavier at birth (Table 8) and at weaning. After 60 days, twin males gained weight faster than twin female lambs. It is interesting to note that, although lambs from twin-bearing litters were heavier at birth than lambs of multiple litters (Table 8), lambs of multiple litters were heavier at 30 days (P=0.054) and slightly heavier at 60 and 90 days of lactation (Table 9). 75 Conclusion Under these conditions (multiparous ewes and spring lambing) I conclude that, in the colostrum experiment, a BCSL ranging from 2.5 and 3.5 had no affect on colostral IgG concentration, mortality rate or weaning weights of lambs. In the lactation experiment, BCS of the ewes after the first month of lactation was similar for ewes with 2.5 and 3.5 BCSL. 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