Organic and inorganic selenium: III. Ewe and progeny performance

Published January 20, 2015 Organic and inorganic selenium: III. Ewe and progeny performance W. C. Stewart,* 1 G. Bobe,* G. J. Pirelli,* W. D. Mosher,* J. A. Hall 2,3 *Department of Animal and Rangeland Sciences, College of Agriculture, Oregon State University, Corvallis 97331; Linus Pauling Institute, Oregon State University, Corvallis 97331; Department of Biomedical Sciences, College of Veterinary Medicine, Oregon State University, Corvallis 97331 ABSTRACT: Selenium is an essential micronutrient in sheep, and deficiency can limit lamb growth and survival. To evaluate how different chemical forms of Se administered to mature ewes at comparative dosages affect ewe and progeny performance, 240 ewes were divided into 8 treatment groups (n = 30 each) and drenched weekly with no Se; at the maximum FDAallowed concentration with inorganic Na-selenite or organic Se-yeast (4.9 mg Se/wk); with inorganic Na-selenate (8.95 mg Se/wk); or with inorganic Na-selenite and organic Se-yeast at supranutritional concentrations (14.7 and 24.5 mg Se/wk, respectively). The treatment period started approximately 2 wk before breeding and lasted for 62.5 wk. Ewes of the no-se and Se-yeast groups continued treatments for another 21 to 24 wk through a second lambing season. Chemical form or dosage of Se did not affect ewe reproductive performance based on proportion of ewes lambing in each treatment group, or number of lambs born, nursed, or weaned per ewe (all P > 0.10). Ewes Key words: ewe health, inorganic selenium, lamb growth, lamb vigor, reproductive performance, selenium yeast 2012 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2012.90:4536 4543 doi:10.2527/jas2011-5019 INTRODUCTION Selenium is an essential micronutrient in sheep. Adequate Se transfer from ewes to lambs is important to prevent Se-responsive diseases in lambs such as nutritional myodegeneration and Se-responsive 1 Current Address: Texas AgriLife Research, 7887 U.S. Highway 87 North, San Angelo 76901. 2 Corresponding author: Jean.Hall@oregonstate.edu 3 Funded in part by USDA CSREES 2008-35204-04624, Agricultural Research Foundation, and Animal Health and Disease Project Formula Funds, Oregon State University, Corvallis. Appreciation is also expressed to B. L. Gill, W. C. Minto, K. J. Hooper, M. L. Galbraith, A. M. Harwell, C. O. Walsh, S. C. Walker, and J. M. Van Duzer for their technical assistance during the conduct of these experiments. Received December 12, 2011. Accepted June 21, 2012. 4536 receiving the highest supplementation rate of Se-yeast at 24.5 mg Se/wk had higher BCS (scale 1 to 5) at the end of yr 1 (2.95 vs. 2.66; P = 0.05) than ewes receiving Se-yeast at 4.9 mg Se/wk. Performance was better in lambs from ewes receiving Se-yeast at 24.5 mg Se/wk than in lambs from ewes receiving Se-yeast at 4.9 mg Se/wk or no Se. In yr 1, lambs from ewes receiving Se-yeast at 24.5 vs. 4.9 mg Se/wk were heavier at 120 d of age (37.0 vs. 34.2 kg; P = 0.05). In yr 2, lambs from ewes receiving Se-yeast at 24.5 mg Se/wk were or tended to be heavier at 60 d of age than lambs from ewes receiving no Se (21.2 vs. 19.0 kg; P = 0.04) or lambs from ewes receiving Se-yeast at 4.9 mg Se/wk (19.2 kg; P = 0.09). This effect was more pronounced in ewes raising multiple lambs. We conclude that supranutritional supplementation of ewes with Se-yeast at 24.5 mg Se/wk improves lamb growth and ewe health without negatively affecting reproductive performance. unthriftiness (Muth et al., 1958). Current FDA regulations limit Se supplementation to 0.3 mg Se/kg diet (as fed), which is equivalent to 0.7 mg ewe -1 d -1 (FDA, 2009). The most common inorganic Se sources are Naselenite and Na-selenate, which are usually provided in mineral premixes or are injected. Organic Se sources are seleno-aa [e.g., selenomethionine (SeMet) and selenocysteine (SeCys)], which are found in Se-yeast or in feeds grown on Se-rich soils. Provision of organic Se to the dam is an effective method to meet Se requirements of newborn lambs because Se crosses the placental barrier into fetal tissues and enters mammary secretions with greater transfer efficiency over a wider supplementation range for organic Se vs. inorganic Na-selenite (Taylor et al., 2009; Stewart et al., 2012). Selenium supplementation may improve

Maternal selenium and lamb performance 4537 reproductive efficiency (Balicka-Ramisz et al., 2006; Koyuncu and Yerlikaya, 2007; Muñoz et al., 2009) and lamb survival and growth (Langlands et al., 1991b; Gabryszuk and Klewiec, 2002; Muñoz et al., 2008). There are concerns, however, that Se supplementation around conception reduces embryonic survival (van Niekerk et al., 1996; Sánchez et al., 2008). Little is known about how different chemical sources of Se given at comparative dosages to ewes affect ewe and progeny performance. The objective of this study were to evaluate how different chemical forms of Se (Se-yeast and Na-selenite) administered at comparative dosages (4.9, 14.7, and 24.5 mg/wk) to mature ewes reared in Se-deficient regions affect ewe and progeny performance. We hypothesized that supplementing ewes with organic Se-yeast above current FDAallowed levels improves lamb growth and survival with no detrimental effects on ewe reproduction. MATERIALS AND METHODS Experimental Design and Treatments Experimental procedures used in this study were approved by the Institutional Animal Care and Use Committee of Oregon State University and have been described in detail previously (Hall et al., 2012). Briefly, 240 mature ewes were randomly assigned to 8 treatment groups (n = 30 each) based on Se supplementation rate and source: Na-selenite, Na-selenate (both from RETORTE Ulrich Scharrer GmbH, Röthenbach, Germany), and Se-yeast (Prince Se Yeast 2000, Prince Agri Products Inc., Quincy, IL). One group received no Se (depletion group); 1 group received Na-selenite at 8.95 mg Se/wk. Three groups received Na-selenite at 4.9, 14.7, or 24.5 mg Se/wk, and 3 groups received Se-yeast at 4.9, 14.7, or 24.5 mg Se/wk. The treatment period started approximately 2 wk before breeding and lasted for 62.5 wk for ewes receiving inorganic Se source. Treatments were continued for another 21 to 24 wk in remaining ewes of the no-se and Se-yeast groups until they lambed a second time. All dosages were less than the maximum tolerable level (5 mg/kg as fed) for small ruminants (NRC, 2007). Treatment groups were blocked for age of ewe and footrot (FR) incidence and severity. Ewes were from 3 genotypes (Polypay, Suffolk, and Suffolk Polypay) and ranged in age and BW from 2 to 6 yr and 51 to 93 kg, respectively. The experiments were conducted at the Oregon State University Sheep Center, Corvallis. Treatments were administered individually once weekly by oral drench (with the calculated weekly amount of Se supplement being equal to the summed daily intake). The Se dose (0, 4.9, 14.7, or 24.5 mg Se) was suspended in a reasonable volume of water (5 ml for inorganic Se with more water required for the organic solutions, i.e., 11, 30, and 48 ml for the 4.9, 14.7, or 24.5 mg Se solutions, respectively). Solutions were prepared weekly and administered with a dose syringe. To ensure a homogeneous dosage, the solution was stirred each time before being drawn into the dose syringe. Lambs did not receive any additional Se supplementation after birth. Individual aliquots of inorganic Se solutions were submitted for Se analysis to the Center for Nutrition, Diagnostic Center for Population and Animal Health, Michigan State University, East Lansing. Except for the 4.9 mg weekly dose of Na-selenate (sheep received 8.95 vs. 4.9 mg because we calculated the molecular weight for Na-selenate decahydrate instead of Na-selenate anhydrous), the 4.9, 14.7, and 24.5 mg weekly doses of Na-selenite (4.85, 14.85, and 24.6 mg Se, respectively) were within expected analytical variance of their targeted concentrations. Concentrations of Se in the pasture forage from Sheep Center pastures ranged from 0.12 to 0.14 μg/g DM. Ewes were naturally mated with a ram:ewe ratio of 3:100 starting in wk 3 and 62.5 of the supplementation period. Rams were with ewes the equivalent of 3 estrous cycles. Ewes were fed on pasture, except for a 3-mo period around lambing when ewes were housed in the barn. Ewes on pasture were supplemented with grass hay and later with alfalfa hay when grass was scarce. In the barn, sheep were fed alfalfa hay and shelled corn, except for 2 d in the lambing jug when ewes were fed alfalfa pellets. Ewe feed sources and management details have been previously described (Hall et al., 2012). Sheep were fed to meet or exceed NRC (2007) recommendations. Ewe and Progeny Performance Reproductive performance was assessed by calculating the number of lambs born, nursed, or weaned per ewe (to 120 d of age in yr 1 and to 60 d of age in yr 2). For each treatment group, we also calculated percentage of ewes that lambed (% lambing; ewes that died before lambing were not included), that nursed lambs (% nursing; if insufficient milk production or mastitis was present at lambing, lambs were removed from the ewe), and that raised lambs (% raising; to 120 d of age in yr 1 and to 60 d of age in yr 2). The latter calculations did not account for the number of lambs per ewe. In yr 2, we recorded an additional variable (i.e., % assistance; percentage of ewes requiring assistance at birth for ewes in each treatment group). Ewe health was evaluated for ewes in each treatment group by calculating the number of ewes that completed the trial in yr 1 and ewes that started and completed the trial in yr 2 (% survival). At the end of yr 1, ewes were weighed and BCS were assigned on a 1 to 5 scale (1 =

4538 Stewart et al. emaciated; 5 = severely obese) by two independent evaluators (ASIA, 2002). Ewes with low BCS at the beginning of yr 2 were culled by the flock manager at the time the rams were introduced (wk 62.5). Thus, the numbers of ewes in each group at the beginning of yr 2 were 25, 20, 18, and 27 for the no-se, 4.9, 14.7, and 24.5 Se-yeast groups, respectively. Based on necropsy results of ewes that died, culled ewes most likely also had severe intestinal parasitism with anthelmintic reisistance. To assess lamb performance, lambs were weighed at birth and at 90 and 120 (weaning) d of age in yr 1 and at birth and at 10, 20, and 60 d of age in yr 2. Bodyweights (except for birth weights) were adjusted for lamb age at weighing because not all lambs were weighed at the same age, using the American Sheep Industry Association formulas (ASIA, 2002). To assess lamb health, the percentage of lambs that died before weaning (to 120 d of age in yr 1 and to 60 d of age in yr 2) was calculated for each treatment group. In yr 2, lamb vigor was scored from 0 to 15 min and from 15 to 30 min after lambing on a 1 to 7 scale: 1 = dead; 2 = hypothermic (recorded rectal temperature); 3 = weak and lethargic; does not attempt to hold up head or stand; 4 = holds up head; 5 = holds up head and attempts to stand; 6 = stands for over 20 sec; 7 = vigorous (stands immediately and attempts to nurse). Statistical Analysis Statistical analyses were performed using SAS, Version 9.1 (SAS Inst., Inc., Cary, NC) software. Body weights and vigor scores for lambs from the same ewe were averaged because ewe was the experimental unit. PROC GLIMMIX was used to analyze data for % lambing, % nursing, and % weaning, assuming a binomial distribution. Count data (lambs per ewe born, nursed, and weaned) were analyzed in PROC GLIMMIX assuming a negative binomial distribution. Fisher s exact test was used for assessing % ewe survival and % lamb survival (from nursing to weaning). The remaining data (BW, BCS, and vigor scores) were analyzed with PROC GLM assuming a normal distribution. Fixed effects in the model were treatment (no Se, 8.95 mg/wk Na-selenate, 4.9 mg/wk Na-selenite, 14.7 mg/wk Na-selenite, 24.5 mg/wk Na-selenite, 4.9 mg/wk Se-yeast, 14.7 mg/ wk Se-yeast, and 24.5 mg/wk Se-yeast), ewe FR-status at the beginning of the treatment period (yes or no), ewe breed (Polypay, Suffolk, or Suffolk Polypay; Suffolk and Suffolk Polypay ewes were combined into one group because the animal numbers for both groups were small and phenotypically the crossbreds more closely resembled the Suffolks in size), ewe age at lambing (3 to 6 or 6 yr), sex of lamb (male or female; only available for yr 1; used only for weights and weight gains of lambs), number of lambs born (0, 1, and 2; used only for lamb data), number of lambs reared (0, 1, and 2; used only for lamb growth data). To evaluate whether FR-status, ewe breed, number of lambs born, and number of lambs reared modified the treatment effect, data were checked for interactions and additionally stratified by FR-status, ewe breed, number of lambs born, and number of lambs reared, respectively. To test the effect of chemical form and Se dosage on ewe and progeny performance, orthogonal contrasts were constructed using the ESTIMATE statement in PROC GLIMMIX, PROC GLM, and PROC MIXED. The effect of Se depletion on ewe and progeny performance was evaluated (no-se vs. 7 Se groups). The effect of Se dosage for Na-selenite (only for yr 1) or for Seyeast was evaluated by comparing the 3 dosages through linear (24.5 vs. 4.9 mg Se/wk) and quadratic (14.7 vs. 4.9 and 24.5 mg Se/wk) contrasts, respectively. The no- Se group was not included in testing the effect of Se dosage because the purpose of the no-se group was to measure the effect of Se depletion. During depletion, Se values continued to decrease (Hall et al., 2012); thus, Se values were unstable in the no-se group (unlike what is expected of a true control group). The control groups are those sheep groups receiving the maximum FDAallowed Se amount (4.9 mg Se/wk). The effect of chemical form of Se was evaluated (3 Na-selenite groups vs. 3 Se-yeast groups; only for yr 1). The interaction between Se source and Se dosage was evaluated by constructing orthogonal contrasts between groups of ewes receiving different Se sources and Se dosages of Na-selenite and Se-yeast (only for yr 1). Data are reported as least squares means ± SEM except for % ewe survival and % lamb survival; the latter are means ± SEM. Statistical significance was declared at P 0.05, and a tendency at 0.10 P < 0.05. RESULTS AND DISCUSSION Supplementation of pregnant ewes with Se is an effective method to improve Se status of their lambs; however, producers are primarily interested in how Se supplementation affects ewe and lamb performance. The objective of this study was to evaluate how inorganic (Na-selenite) and organic forms (Se-yeast) of Se administered at FDA-allowed (4.9 mg Se/wk) or supranutritional dosages (14.7 and 24.5 mg Se/wk) to mature ewes affect ewe and lamb performance. Our primary findings are that chemical form or dosage of Se did not significantly affect reproductive performance. However, BCS of ewes and growth of lambs were or tended to be better when ewes received Se-yeast at 24.5 mg Se/wk vs. 4.9 mg Se/wk. We conclude that Se-yeast at 24.5 mg Se/wk may improve lamb performance, compared with FDA-allowed dosages of 4.9 mg Se/wk in regions with Se-deficient soils.

Maternal selenium and lamb performance 4539 Effect of Selenium on Ewe Reproductive Performance and Health Chemical form and dosage of Se, and Se depletion, did not significantly affect the proportion of ewes that lambed or needed assistance at birth, or the number of lambs born, nursed, or weaned per ewe in yr 1 (Table 1) or yr 2 of the study (Table 2). Similar results, albeit with smaller animal numbers and fewer treatment groups, have been reported previously (Rock et al., 2001; Rodinova et al., 2008). Results from other studies show that testing for an effect of Se supplementation on reproductive performance depends upon the Se status of the control group. In Se-deficient ewes, Se supplementation improves reproductive performance (Hemingway, 2003; Muñoz et al., 2009), but not always in Se-replete ewes (Whanger et al., 1977; Langlands et al., 1991a), as shown in our study (Tables 1 and 2). There are some concerns that additional Se supplementation in ewes with adequate Se status may negatively affect embryonic survival shortly after conception through an unknown mechanism (van Niekerk et al., 1996; Sánchez et al., 2008; Palmieri and Szarek, 2011). Davis et al. (2006a,b) addressed these concerns by feeding ewes throughout 2 lambing seasons with up to 20 mg Se/kg of diet as fed of Na-selenite, dietary Se concentrations much greater than in our study, and did not detect detrimental effects on ewe conception rates. Similarly, Glenn et al. (1964) did not observe detrimental effects on fertility or early embryonic development in ewes treated daily with subtoxic to toxic (50 mg Se/d) doses of Na-selenate for extended periods. The previously reported negative effects of Se supplementation on ewe reproduction may be because van Niekerk et al. (1996) used a very high dosage of Ba-selenite (one time injection of 50 mg Se as Ba-selenite; no significant effect of injecting 1 mg Se as Na-selenite) administered 18 to 35 d after mating to achieve the decrease in conception rates. Sánchez et al. (2008) observed that estrus synchronized ewes receiving 0.125 mg Se/d as SeMet in the diet had lower conception and lambing rates than estrus synchronized ewes receiving no Se supplement, although naturally mated and Se-supplemented ewes had similar conception and lambing rates, compared with non-sesupplemented ewes. Therefore, we conclude that there is little evidence to suggest a detrimental effect of supranutritional Se supplementation at dosages up to 24.5 mg Se/wk. However, ours as well as many of the other studies, have the limitation that several hundred animals per treatment group are needed to detect differences of even 10% with binary data. Therefore, an even largerscale study to evaluate the effect of supranutritional Se supplementation with organic Se-yeast on reproductive efficiency is warranted. Ewes receiving Se-yeast at 24.5 vs. 4.9 mg Se/wk were in better body condition at the end of yr 1 (P = 0.05; Table 1). A similar effect was not found with supranutritional dosages of Na-selenite. Muñoz et al. (2009) reported that Se supplementation increased BW and BCS in ewes. Meyer et al. (2010) reported that although dietary Se supply had no effect on ewe BW, it did affect ADG during pregnancy such that ewes fed high Se had a greater BCS increase during midgestation and a lesser BCS loss during late gestation than those fed adequate Se. In most other studies, inorganic or organic dietary Se supplementation above the maximum FDA-allowed level was not associated with changes in BW (Davis et al., 2006b; Neville et al., 2008; Swanson et al., 2008). There are at least 3 groups of selenoproteins: gluthathione peroxidase family of enzymes, responsible for reduction of hydroperoxides; iodothyronine deiodinases, responsible for metabolism of thyroid hormones; and thioredoxin reductases, which reduce thioredoxin and are thus involved in many cell functions including cell growth, control of apoptosis, and maintenance of cellular redox status, often through regulation of transcription factors (Rooke et al, 2004). An understanding of the role of Se in immune function and disease resistance led Finch and Turner (1996) to summarize that Se deficiency could compromise the immune system and result in a decline in production and performance before gross effects became apparent. Selenium supplementation might thereby improve feed efficiency by reducing the incidence of subclinical disease. We conclude that supranutritional Se supplementation with Se-yeast at 24.5 mg Se/wk vs. 4.9 mg Se/wk improves the overall fitness of ewes. Effect of Selenium on Lamb Performance The dosage of Se administered to ewes as Seyeast, but not Na-selenite or Se depletion, affected birth weights of their lambs in yr 1 (Table 1). There were both linear and quadratic effects of Se-yeast dosage (4.9, 14.7, and 24.5 mg Se/wk). Lambs from ewes receiving Se-yeast at 4.9 mg Se/wk had the lightest birth weights, compared with any of the other groups (P 0.05). There was no effect of maternal Se-yeast dosage on lamb birth weights in yr 2 (Table 2). However, lambs from ewes receiving Se-yeast at 24.5 vs. 4.9 mg Se/wk tended to have higher vigor scores 30 min after lambing in yr 2 (P = 0.08). Some studies have shown a significant increase in fetal weight with supranutritional Se supplementation of ewes during pregnancy (Reed et al., 2007); others have not (Neville et al., 2008). Some studies have shown decreased lamb BW at birth (Gabryszuk and Klewiec, 2002); others have shown no effect on birth weight (Muñoz et al., 2008; Swanson et al., 2008) in lambs from

4540 Stewart et al. Table 1. Effect of weekly oral drenching with no Se, inorganic Se (Na-selenate or Na-selenite), or organic Se (Seyeast) at varying supplementation rates (4.9, 14.7, and 24.5 mg Se/wk; Na-selenate only at 8.95 mg Se/wk) for 60 wk (starting 2 wk before mating) on ewe and progeny performance in the first lambing season Oral selenium drench P-value 1 No-Se Na-Selenate Na-Selenite Se-Yeast Se dosage, mg Se/wk 0 8.95 4.9 14.7 24.5 4.9 14.7 24.5 SEM 2 Overall Ewe Reproduction n = 30 n = 30 n = 30 n = 30 n = 30 n = 30 n = 30 n = 30 Lambs/Ewe Born 1.83 1.73 1.56 1.79 1.99 1.60 1.78 1.45 0.27 0.83 Nursed 1.64 1.45 1.49 1.43 1.71 1.40 1.35 1.18 0.25 0.76 Weaned 1.64 1.45 1.49 1.43 1.71 1.40 1.35 1.15 0.25 0.76 Ewes % Lambing 96 97 87 93 100 94 100 85 7 0.75 % Nursing 93 94 87 87 97 94 93 79 8 0.45 % Weaning 93 94 87 87 97 94 93 79 8 0.45 Ewe Health, after 60 wk BW, kg 67.0 65.7 66.6 67.1 65.1 67.3 67.8 70.1 1.7 0.60 BCS 3 2.72 2.57 2.74 2.78 2.64 2.66 2.75 2.95 0.11 0.31 % Survival 90 80 93 100 90 90 90 90 7 0.87 Lamb Growth Individual BW, kg 4 At birth 5 6.30 6.35 6.29 6.32 6.08 5.51 6.36 6.03 0.20 0.01 At 90 d of age 33.2 32.8 32.6 32.4 34.0 31.9 33.0 33.2 1.8 0.75 At 120 d of age Overall 6 36.1 35.1 34.9 34.7 36.7 34.2 34.8 37.0 2.0 0.39 Single lambs 7 n = 7 n = 13 n = 7 n = 8 n = 7 n = 14 n = 15 n = 10 BW, kg 38.0 37.3 36.8 38.8 40.8 36.8 39.1 38.4 3.4 0.92 Multiple lambs 7,8 n = 20 n = 15 n = 19 n = 18 n = 21 n = 13 n = 11 n = 13 BW, kg 32.8 32.3 31.6 30.7 32.7 30.8 29.3 34.3 1.2 0.06 Polypay ewes 9 n = 17 n = 16 n = 14 n = 14 n = 16 n = 13 n = 16 n = 17 BW, kg 34.9 34.9 35.5 33.2 35.9 33.4 33.4 35.4 1.1 0.40 Suffolk ewes 10 n = 10 n = 12 n = 12 n = 12 n = 12 n = 14 n = 10 n = 6 BW, kg 38.7 36.4 33.6 36.7 38.0 34.7 37.5 37.8 2.5 0.44 Sum BW/ewe, kg 11 At 120 d of age 47.4 46.5 45.2 44.7 46.7 44.7 44.4 47.1 3.2 0.79 Lamb health % Survival from onset of 100 100 100 100 98 100 100 98 3 NA nursing to 120 d 1 Overall = any group differences. Preplanned contrasts were performed. The effect of Se depletion on ewe and progeny performance was evaluated (no-se vs. 7 Se groups). The effect of Se dosage for Na-selenite or for Se-yeast was evaluated by comparing the 3 dosages through linear (24.5 vs. 4.9 mg Se/wk) and quadratic (14.7 vs. 4.9 and 24.5 mg Se/wk) contrasts, respectively. The effect of chemical form of Se was evaluated (3 Na-selenite groups vs. 3 Se-yeast groups). The interaction between Se source and Se dosage was evaluated by constructing orthogonal contrasts between groups of ewes receiving different Se sources and Se dosages of Na-selenite and Se-yeast. Significant effects of preplanned contrasts are indicated as separate footnotes. 2 The largest SEM of the 8 treatment groups is shown. 3 BCS were assigned on a 1 to 5 scale (1 = emaciated; 5 = severely obese). Body condition score was greater in ewes receiving 24.5 mg Se/wk as Se-yeast vs. 4.9 mg Se/wk as Se-yeast (P = 0.05). 4 BW values for lambs from the same ewe were averaged then average values for all ewes in a treatment group are shown. 5 Lamb BW at birth was greater in ewes receiving 24.5 mg Se/wk as Se-yeast vs. 4.9 mg Se/wk as Se-yeast (P = 0.05). The quadratic effect of Se dosage was significant for lamb weight at birth (P = 0.008) for Se-yeast. 6 Lamb BW at 120 d of age overall was greater in ewes receiving 24.5 mg Se/wk as Se-yeast vs. 4.9 mg Se/wk as Se-yeast (P = 0.05). 7 Lamb BW at 120 d of age was also stratified by number of lambs nursed (P Interaction Se supplementation number of lambs nursed = 0.38). 8 Lamb BW at 120 d of age was greater in lambs nursed as multiples from ewes receiving 24.5 vs. 4.9 mg Se/wk as Se-yeast (P = 0.03). The quadratic effect of Se dosage was significant for lamb weight at 120 d of age (P = 0.03) for Se-yeast. The effect of the interaction between Se source and Se dosage was significant for lamb weight at 120 d of age (P = 0.009). 9 Lamb BW at 120 d of age was also stratified by ewe breed (P Interaction Se supplementation x ewe breed = 0.43). 10 Suffolk ewes also included Suffolk Polypay ewes. 11 Sum of all lamb BW produced per ewe were averaged and shown for each treatment group.

Maternal selenium and lamb performance 4541 Table 2. Effect of weekly oral drenching with no Se or organic Se (Se-Yeast) at varying supplementation rates (4.9, 14.7, and 24.5 mg Se/wk) for 84 to 87 wk (starting 2 wk before mating in the first lambing season) on ewe and progeny performance in the second lambing season (oral drenching stopped at lambing time) No-Se Se-Yeast Contrast2 Se dosage, mg Se/wk 0 4.9 14.7 24.5 SEM 1 Depletion Linear Quadratic Ewe reproduction n = 25 n = 20 n = 18 n = 27 Lambs/ewe Born 1.59 1.28 1.62 1.50 0.30 0.66 0.53 0.49 Nursed 1.56 1.14 1.41 1.41 0.28 0.40 0.43 0.65 Weaned 1.04 0.86 1.11 1.32 0.25 0.87 0.14 0.89 Ewes % Lambing 95 77 87 84 10 0.19 0.55 0.59 % Assistance 15 9 12 2 9 0.22 0.29 0.28 % Nursing 95 77 87 81 10 0.15 0.74 0.46 % Weaning 81 60 79 81 11 0.58 0.11 0.60 Ewe health % Survival, overall 3 92 100 100 100 6 0.07 1.00 1.00 Lamb growth Individual BW, kg 4 At birth 4.54 4.29 4.66 4.65 0.26 0.98 0.29 0.52 At 10 d of age 7.28 6.92 7.31 7.50 0.47 0.94 0.32 0.84 At 20 d of age 10.22 9.79 10.73 10.38 0.55 0.88 0.39 0.30 At 60 d of age Overall 5 19.04 19.21 20.95 21.18 0.94 0.12 0.09 0.46 Single lambs 6 n = 8 n = 3 n = 5 n = 5 BW, kg 22.17 22.95 20.26 22.40 2.11 0.87 0.84 0.31 Multiple lambs 6,7 n = 10 n = 9 n = 9 n = 17 BW, kg 17.00 17.80 21.43 20.33 1.04 0.01 0.04 0.04 Polypay ewes 8 n = 11 n = 6 n = 10 n = 15 BW, kg 18.08 19.34 19.35 19.81 1.43 0.27 0.78 0.87 Suffolk Ewes 8,9 n = 7 n = 6 n = 4 n = 7 BW, kg 20.44 19.70 24.18 23.21 1.92 0.24 0.05 0.17 Lamb health Vigor score, 15 min 10 4.87 4.70 4.92 4.94 0.26 0.94 0.45 0.74 Vigor score, 30 min 10 6.37 5.97 5.90 6.52 0.25 0.32 0.08 0.21 % Survival from onset of 64 75 75 90 9 0.06 0.16 0.54 nursing to 60 d 11 1 The largest SEM of the 4 treatment groups is shown. 2 Preplanned, orthogonal contrasts were performed to measure the effect of Se Depletion (no-se vs. 3 Se-yeast groups) and the linear (24.5 vs. 4.9 mg Se/wk) and quadratic (14.7 vs. 4.9 and 24.5 mg Se/wk) effect of Se-yeast dose. 3 Survival during dosage period in yr 2. 4 BW values for lambs from the same ewe were averaged then average values for all ewes in a treatment group are shown. 5 Lamb BW at 60 d of age overall was greater in lambs from ewes receiving 24.5 mg Se/wk vs. no Se (P = 0.04). 6 Lamb BW at 60 d of age was also stratified by number of lambs nursed (P Interaction Se supplementation x number of lambs nursed = 0.05). 7 Lamb BW at 60 d of age was greater in lambs nursed as multiples from ewes receiving 24.5 mg Se/wk vs. no Se (P = 0.007). 8 Lamb BW at 60 d of age was analyzed stratified by ewe breed (P Interaction Se supplementation x ewe breed = 0.55). 9 Suffolk ewes also included Suffolk x Polypay ewes. Lamb weight at 60 d of age was greater in lambs nursed from Suffolk or Suffolk x Polypay ewes receiving 24.5 mg Se/wk vs. no Se (P = 0.05); the interaction by ewe breed, however, was not significant (P Interaction Se supplementation x ewe breed = 0.55). 10 Vigor scale: Scale: 1 = dead; 2 = hypothermic (recorded rectal temperature); 3 = weak and lethargic; does not attempt to hold up head or stand; 4 = holds up head; 5 = holds up head and attempts to stand; 6 = stands for over 20 s; 7 = vigorous stands immediately and attempts to nurse. 11 Lamb survival was greater in lambs from ewes receiving 24.5 mg Se/wk vs. no Se (P = 0.01). ewes consuming either organic or inorganic Se sources at various dosages. Ewes receiving Se-yeast at 24.5 vs. 4.9 mg Se/wk had heavier lambs at weaning (120 d of age) in yr 1 (P = 0.05; Table 1) and tended to have heavier lambs in yr 2 (P = 0.09; Table 2). In addition, compared with ewes receiving no Se supplement, lambs from ewes receiving Se-yeast at 24.5 mg Se/wk were heavier (P = 0.04) and had a higher survival rate at 60 d of age in yr 2 (P = 0.01; Table 2). The effect was more pronounced in ewes of heavier breeds (Suffolk or Suffolk Polypay) vs. lighter breeds (Polypay), and in ewes having multiple vs. single

4542 Stewart et al. lambs; however, the sample size got very small and the interaction was not always significant (Tables 1 and 2). The level of selenium supplementation needed for adequate performance may be less under optimum conditions; however, in situations of raising multiple lambs and in breeds with faster growth rates, Se requirements may be increased for optimum performance. Previous studies have reported that Se supplementation of ewes during pregnancy improves lamb growth in the first 2 wk (Abd Elghany et al., 2008), 4 wk (Gabryszuk and Klewiec, 2002), 6 wk (Muñoz et al., 2009), and at 60 d (Koyuncu and Yerlikaya, 2007). Similar to our study, Muñoz et al. (2008) reported reduced perinatal lamb mortality, yet overall mortality from birth to weaning was unaffected when dams were supplemented with Se. Muñoz et al. (2008) also reported heavier weaning weights at 16 wk in lambs from ewes receiving 0.5 mg Se/d as Se-yeast from 14 to 90 d after mating. Kott et al. (1983) reported improved preweaning lamb survival in ewes receiving monthly injections of 4 mg Se as Naselenite during mating and pregnancy. It is well known that Se supplementation of Se-deficient lambs results in significant BW gain responses (Grace and Knowles, 2002; Rooke et al., 2004). We have no explanation why the lower level of Se supplementation had no beneficial effect, compared with Se depletion. One potential reason could be that animals in the Se depletion group were not Se deficient. We did not observe any overt clinical signs of Se deficiency in ewes or their progeny. Another possibility is that the lower level of Se supplementation was not adequate to see beneficial effects. In summary, supranutritional Se supplementation with Se-yeast at 24.5 mg Se/wk compared with the maximum FDA-allowed levels of 4.9 mg Se/wk resulted in increased BW of weaned lambs and improved overall health of ewes and lambs in a region with Se-deficient soils. Supranutritional Se supplementation of pregnant ewes with Se-yeast may be an effective management strategy for meeting the Se requirements of growing lambs, especially when providing Se supplementation to lambs is a complicated endeavor. LITERATURE CITED Abd Elghany, H., A. E. López-Arellano, R. Revilla-Vázquez, E. Ramírez-Bribiesca, and E. J. Tortóra-Pérez. 2008. Effect of preand postpartum selenium supplementation in sheep. J. Anim. Vet. Adv. 7:61 67. American Sheep Industry Association (ASIA). 2002. Sheep Production Handbook. ADS/Nightwing Publishing, Fort Collins, CO. Balicka-Ramisz, A., B. Pilarczyk, A. Ramisz, and M. Wieczorek. 2006. Effects of selenium administration on blood serum Se content and on selected reproductive characteristics of sheep. Arch. Tierz. Dummerstorf 49:176 180. Davis, P. A., L. R. McDowell, N. S. Wilkinson, C. D. Buergelt, R. Van Alstyne, R. N. Weldon, and T. T. Marshall. 2006a. Effects of selenium levels in ewe diets on selenium in milk and the plasma and tissue selenium concentrations of lambs. Small Ruminant Res. 65:14 23. Davis, P. A., L. R. McDowell, N. S. Wilkinson, C. D. Buergelt, R. Van Alstyne, R. N. Weldon, and T. T. Marshall. 2006b. Tolerance of inorganic selenium by range-type ewes during gestation and lactation. J. Anim. Sci. 84:660 668. FDA. 2009. Title 21. Food and Drugs: Food additives permitted in feed and drinking water of animals. Accessed June 3, 2010. http://www.accessdata.fda.gov/scripts/cdrh/cfdocs/cfcfr/ CFRSearch.cfm?fr=573.920 Finch, J. M., and R. J. Turner. 1996. Effects of selenium and vitamin E on the immune responses of domestic animals. Res. Vet. Sci. 60:97 106. Gabryszuk, M., and J. Klewiec. 2002. Effect of injecting 2- and 3-year-old ewes with selenium and selenium-vitamin E on reproduction and rearing of lambs. Small Ruminant Res. 43:127 132. Glenn, M. W., R. Jensen, and L. A. Griner. 1964. Sodium selenate toxicosis: The effects of extended oral administration of sodium selenate on mortality, clinical signs, fertility, and early embryonic development in sheep. Am. J. Vet. Res. 25:1479 1485. Grace, N. D., and S. O. Knowles. 2002. A reference curve using blood selenium concentration to diagnose selenium deficiency and predict growth responses in lambs. N. Z. Vet. J. 50:163 165. Hall, J. A., R. J. Van Saun, G. Bobe, W. C. Stewart, W. R. Voracheck, W. D. Mosher, T. Nichols, N. E. Forsberg, and G. J. Pirelli. 2012. Organic and inorganic selenium: I. Oral bioavailability in ewes. J. Anim. Sci. 90:568 576. Hemingway, R. G. 2003. The influences of dietary intakes and supplementation with selenium and vitamin E on reproduction diseases and reproductive efficiency in cattle and sheep. Vet. Res. Comm. 27:159 174. Kott, R. W., J. L. Ruttle, and G. M. Southward. 1983. Effects of vitamin E and selenium injections on reproduction and preweaning lamb survival in ewes consuming diets marginally deficient in selenium. J. Anim. Sci. 57:553 558. Koyuncu, M., and H. Yerlikaya. 2007. Effect of selenium-vitamin E injections of ewes on reproduction and growth of their lambs. S. Afr. J. Anim. Sci. 37:233 236. Langlands, J. P., G. E. Donald, J. E. Bowles, and A. J. Smith. 1991a. Subclinical selenium insufficiency. 2. The response in reproductive-performance of grazing ewes supplemented with selenium. Aust. J. Exp. Agric. 31:33 35. Langlands, J. P., G. E. Donald, J. E. Bowles, and A. J. Smith. 1991b. Subclinical selenium insufficiency. 3. The selenium status and productivity of lambs born to ewes supplemented with selenium. Aust. J. Exp. Agric. 31:37 43. Meyer A. M., J. J. Reed, T. L. Neville, J. B. Taylor, C. J. Hammer, L. P. Reynolds, D. A. Redmer, K. A. Vonnahme, and J. S. Caton. 2010. Effects of plane of nutrition and selenium supply during gestation on ewe and neonatal offspring performance, body composition, and serum selenium. J. Anim. Sci. 88:1786 1800. Muñoz, C., A. F. Carson, M. A. McCoy, L. E. R. Dawson, D. Irwin, A. W. Gordon, and D. J. Kilpatrick. 2009. Effect of supplementation with barium selenate on the fertility, prolificacy and lambing performance of hill sheep. Vet. Rec. 164:265 272. Muñoz, C., A. F. Carson, M. A. McCoy, L. E. R. Dawson, N. E. O Connell, and A. W. Gordon. 2008. Nutritional status of adult ewes during early and mid-pregnancy. 2. Effects of supplementation with selenised yeast on ewe reproduction and offspring performance to weaning. Animal 2:64 72. Muth, O. H., J. E. Oldfield, L. F. Remmert, and J. R. Schubert. 1958. Effects of selenium and vitamin E on white muscle disease. Sci-

Maternal selenium and lamb performance 4543 ence 128:1090. Neville, T. L., M. A. Ward, J. J. Reed, S. A. Soto-Navarro, S. L. Julius, P. P. Borowicz, J. B. Taylor, D. A. Redmer, L. P. Reynolds, and J. S. Caton. 2008. Effects of level and source of dietary selenium on maternal and fetal body weight, lambs visceral organ mass, cellularity estimates, and jejunal vascularity in pregnant ewe. J. Anim. Sci. 86:890 901. NRC. 2007. Nutrient Requirements of Small Ruminants: Sheep, Goats, Cervids, and New World Camelids. Natl. Acad. Press, Washington, DC. Palmieri, C., and J. Szarek. 2011. Effect of maternal selenium supplementation on pregnancy in humans and livestock. J. Elementol. 16:143 156. Reed, J. J., M. A. Ward, K. A. Vonnahme, T. L. Neville, S. L. Julius, P. P. Borowicz, J. B. Taylor, D. A. Redmer, A. T. Grazul-Bilska, L. P. Reynolds, and J. S. Caton. 2007. Effects of selenium supply and dietary restriction on maternal and fetal body weight, visceral organ mass and cellularity estimates, and jejuna vascularity in pregnant ewe lambs. J. Anim. Sci. 85:2721 2733. Rock, M. J., R. L. Kincaid, and G. E. Carstens. 2001. Effects of prenatal source and level of dietary selenium on passive immunity and thermometabolism of newborn lambs. Small Ruminant Res. 40:129 138. Rodinova, H., V. Kroupova, J. Travnicek, M. Stankova, and L. Pisek. 2008. Dynamics of IgG in the blood serum of sheep with different selenium intake. Veterinarni Medicina 53:260 265. Rooke, J. A., J. J. Robinson, and J. R. Arthur. 2004. Effects of vitamin E and selenium on the performance and immune status of ewes and lambs. J. Agric. Sci. 142:253 262. Sánchez, J., A. Jiménez, S. Regodón, and S. Andrés. 2008. Inhibitory effect of selenium supplementation on the reproductive performance in synchronized Merino sheep at range conditions in a selenium-deficient area. Reprod. Domest. Anim. 43:328 332. Stewart, W. C., G. Bobe, W. R. Voracheck, G. J. Pirelli, W. D. Mosher, T. Nichols, R. J. Van Saun, N. E. Forsberg, and J. A. Hall. 2012. Organic and inorganic selenium: II. Transfer efficiencies from ewes to lambs. J. Anim. Sci. 90:577 584. Swanson, T. J., C. J. Hammer, J. S. Luther, D. B. Carlson, J. B.Taylor, D. A. Redmer, T. L. Neville, J. J. Reed, L. P. Reynolds, J. S. Caton, K. A. Vonnahme. 2008. Effects of gestational plane of nutrition and selenium supplementation on mammary development and colostrum quality in pregnant ewe lambs. J. Anim. Sci. 86:2415 2423. Taylor, J. B., L. P. Reynolds, D. A. Redmer, and J. S. Caton. 2009. Maternal and fetal tissue selenium loads in nulliparous ewes fed supranutritional and excessive selenium during mid- to late pregnancy. J. Anim. Sci. 87:1828 1834. van Niekerk, F. E., S. W. P. Cloete, E. W. P. Heine, G. D. van der Merwe, A. Wellington, S. S. du Plessis, and D. Bekker. 1996. The effect of selenium supplementation during the early postmating period on embryonic survival in sheep. J. S. Afr. Vet. Assoc. 67:209 213. Whanger, P. D., P. H. Weswig, J. A. Schmitz, and J. E. Oldfield. 1977. Effects of selenium and vitamin E deficiencies on reproduction, growth, blood components, and tissue lesions in sheep fed purified diets. J. Nutr. 107:1288 1297.