Effect of Selenium Source on Production, Reproduction and Immunity of Lactating Dairy Cows in...

7/31/2019 Effect of Selenium Source on Production, Reproduction and Immunity of Lactating Dairy Cows in Florida and Calif

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Effect of selenium source on production,reproduction and immunity of lactating dairy cows inFlorida and CaliforniaPublished on: 08/27/2007

Rating:Author : FLAVIO T. SILVESTRE, HELOISA M. RUTIGLIANO, WILLIAM W. THATCHER, JOS E.P.SANTOS and CHARLES R. STAPLES (Courtesy of Alltech Inc.)

A nutraceutical is defined as a product isolated or purified from feeds that is demonstrated to have aphysiological benefit or provide protection against chronic disease. Indeed selenium (Se) is an essentialnutrient in that selenium deficiency is associated with increased incidences of retained fetal membranes,clinical mastitis, calf mortality and increased milk somatic cell counts (SCC).

Selenium supplementation in Se-deficient diets reduces the incidence of these clinical problems.Consequently, selenium should be considered a nutraceutical that has an array of biological responses.

Cows fed Se-deficient diets have reduced blood glutathione peroxidase (GSH-Px) activity compared withthose fed Se-supplemented diets (Grasso et al., 1990; Gunter et al., 2003). Likewise, cows receiving

injections containing sodium selenite have greater concentrations of selenium in plasma and greater GSH-Px activity than cows not injected with sodium selenite (Hogan et al., 1990). Glutathione peroxidase is oneof the selenoenzymes (Se-GSH-Px) capable of protecting the cell against oxidative injury.

The Se-GSH-Px catalyzes the reduction of hydrogen peroxide (H2O2) to water and organichydroperoxides to alcohols while utilizing the peptide glutathione as a cofactor (Mezzetti et al., 1990). It isimportant, for example, that neutrophils provide a high oxidizing intracellular environment to ki llphagocytized bacteria, but it is essential that neutrophils regulate the balance between reactive oxygenmetabolites, superoxide (O2 -) and H2O2, in order not to damage cells, leading to cell death.

Iodothyronine deiodinase enzyme is a selenoenzyme that catalyzes the activation and inactivation ofthyroid hormones that regulate metabolic processes and may contribute to production responses.

Basal feed ingredients and selenium yeast provide mostly selenoamino acids (i.e., selenomethionine

(SeMet) and selenocysteine (SeCys)), whereas inorganic selenium supplements provide selenate andselenite. The basic pathways for selenium metabolism have been summarized by Weiss (2003).

Inorganic selenium sources undergo reduction to form selenide, which leads to the formation of SeCys(i.e., the hydroxyl group of a serine molecule linked to a specific tRNA [UGA codon] is replaced with aselenol moiety to form SeCys-tRNA, which is inserted into selenoproteins). Thus, various seleniumsources (both inorganic and organic) must first be converted to inorganic selenide before synthesis ofSeCys, which contributes to the bioactive components of selenoproteins. After absorption of SeMet fromthe intestinal tract, SeMet can be found in blood proteins and in the plasma methionine pool as it istransported to body tissues.

For example, the mammary gland extracts large quantities of methionine to synthesize milk proteins. Thiswould account for the large amounts of selenium found in milk, which may benefit the neonate or serve asa selenium source for human consumption.

During the immediate postpartum period, the cows immune system is challenged severely (Goff, 2006)and the innate and humoral defense systems are reduced. The incidence of diseases and disorders canbe high during this time period and have a negative impact on reproductive performance. For example, therisk of pregnancy (odds ratio) is reduced if cows have retained fetal membranes (RFM) or lose one bodycondition score (BCS) (Loeffler et al., 1999). Reduction in adaptive and innate immunity at parturitionincreases the risk of health disorders such as RFM, metritis, and mastitis.

Selenium has long been associated with immunity. In some studies, cattle supplemented with seleniumyeast had an 18% increase in plasma selenium compared with those fed sodium selenite (Weiss, 2003).

We have conducted joint experiments between the laboratories of W.W. Thatcher at the University ofFlorida (Silvestre et al., 2006a,b) and J.E.P. Santos at the University of California, Davis (Rutigliano et al.,2006) to evaluate a supplemental source of organic selenium on immune, health, reproductive, andlactation responses by dairy cows.

Objectives were to evaluate effects of organic selenium on pregnancy rate (PR) at the first and secondpostpartum artificial insemination (AI) services, uterine health, milk yield and periparturient immune


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responses during the summerheat stressperiod. The concept of replicating the experiment at two sites wasto have sufficient numbers of lactating dairy cows to test the effects of selenium supplements onpregnancy rates to first and second services. The state of Florida is a selenium-deficient area, andlactating dairy cows are exposed to a seasonal period of heat stress that impairs reproductive performanceand health.

Materials and methods

EXPERIMENTAL DESIGN

Florida. Cows were assigned (23 8 days prepartum) to diets containing organic selenium (Sel-Plex,Alltech Inc.; n = 289) or inorganic selenium (sodium selenite [SS; n = 285]) fed at 0.3 mg/kg (dry matter(DM) basis) for more than 81 days postpartum (dpp).

Rectal temperature was recorded each morning for 10 days postpartum. Vaginoscopies were performed at5 and 10 days postpartum. Cows within diet were assigned randomly to two reproductive managementprograms (Presynch-Ovsynch vs CIDR-Ovsynch).

Cows enrolled in the Presynch-Ovsynch received two injections of 25 mg of PGF2 (dinoprosttromethamine; Lutalyse sterile solution, Pfizer Animal Health, New York, NY) on study days 45 + 3 and 59.

Cows in the CIDR-Ovsynch received an intravaginal insert containing 1.38 g of progesterone (EAZI-BREED CIDR; Pfizer Animal Health, New York, NY) from study days 53 to 60. At removal of theintravaginal insert, cows received an injection of PGF2.

All cows were artificially inseminated (AI) at fixed time on study day 81 (i.e., 81 dpp) following the Ovsynchprotocol starting at 12 or 3 days after Presynch- and CIDR-presynchronization protocols, respectively. Forthe Ovsynch protocol, cows received an injection of 100 g of GnRH (gonadorelin diacetate tetrahydrate,Cystorelin; Merial, Ltd, Iselin, NJ), followed 7 days later by an injection of 25 mg of PGF2, and a secondinjection of 100 g of GnRH 48 hrs later, and AI 12 hrs after the final injection of GnRH.

All cows were resynchronized for a second service with Ovsynch 20 to 23 days after first service. Anultrasound pregnancy diagnosis was conducted 27 to 30 days after first timed AI (TAI) and re-confirmed 55days after TAI. Cows in estrus following Presynchs were AI up to the second TAI service. The pregnancyrate at second service was determined by rectal palpation ~42 days after AI. Strategic blood samplingdetermined anovulatory status at Ovsynch and ovulatory response after TAI to first service. Lactationperformance was monitored daily through 81 days of lactation and monthly via DHIA testing throughoutlactation.

California. Cows were assigned to treatments as a randomized block design in a 2 x 2 factorialarrangement of treatments at approximately 25 3 days before the expected date of calving (study day 0= day of calving). Animals were blocked by parity (201 primiparous, and 365 multiparous) and expectedcalving date and, within each block, randomly assigned to one of the four treatments. Treatments were thetwo sources of dietary selenium and the two methods of presynchronization (see Florida, above).

Selenium was supplemented in the pre- and postpartum diets at 0.3 mg/kg of diet DM either as sodiumselenite or Sel-Plex. Cows were also assigned to one of the two presynchronization methods. Cowsenrolled in the Presynch-Ovsynch received two injections of 25 mg of PGF2 on study days 37 and 51.

Cows in the CIDR-Ovsynch received the intravaginal progesterone insert from study days 53 to 60. At

removal of the intravaginal insert, cows received an injection of PGF2. All cows were artificiallyinseminated at fixed time on study day 72 following the Ovsynch protocol starting at 12 or 3 days after thepresynchronization with Presynch and CIDR-PS protocols, respectively. Body condition was scored atstudy enrollment. Lactation performance was followed for the first 80 days postpartum. Cows werediagnosed for pregnancy at 28, 42 and 56 days after AI.

DIETS

Florida and California: Diets were formulated to meet or exceed nutrient requirements for pre-partum andlactating cows (NRC, 2001). Table 1 depicts the respective diets for pre-partum and lactating experimentalcows in Florida and California.

Results and discussionPLASMA SELENIUM AND IMMUNE RESPONSES
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Florida. Blood was sampled on (n=20 cows/diet) -25, 0, 7, 14, 21, and 37 days postpartum to determineselenium and immune status. Plasma selenium increased in Sel-Plex -fed cows (0.087 vs 0.069 0.004g/mL; P


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Anti-IgG to Ovalb did not differ between dietary groups at -60 and -22 days postpartum (0.18 0.01 and0.97 0.04 optical density [OD]). Concentrations of anti-IgG to Ovalb were higher (P


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percentage of phagocytized bacteria killed, number of phagocytized bacteria, and percentage ofneutrophils killing at least one bacterium compared with cows at 42 days postpartum.

Multiparous cows tended (P=0.08) to kill a greater percentage of intracellular bacteria compared withprimiparous cows (46.9 vs 40.5%).

Oxidative burst of neutrophils was evaluated in vitro by reduction of nitroblue tetrazolium (NBT). Source of

selenium and parity did not affect oxidative burst in nonstimulated and stimulated neutrophils. A greater(P=0.04) proportion of nonstimulated neutrophils collected from cows at study day 42 reduced NBTcompared with those from cows at calving, but days postpartum did not affect oxidative burst in stimulatedneutrophils.

Table 2. Effect of selenium source on neutrophil function (LS means) in experimental cows ofCalifornia.

1Study day 0 = calving.

2Se = source of Se; Study day; and Se DPP = interaction between Se and DPP.

3

Percentage of neutrophils containing at least one intracellular bacterium (live or dead)4Number of bacteria phagocytized / number of neutrophils phagocytizing at least one bacterium.

5Number of dead, phagocytized bacteria / (number of live + dead phagocytized bacteria) 100.

6Number of neutrophils containing at least one dead bacterium / number of neutrophils phagocytizing at

least one bacterium.

Source of selenium did not influence serum titers for anti-ovalbumin IgG of cows in the first 42 dayspostpartum. Concentrations of anti-ovalbumin IgG in serum increased (P=0.002) over the study period asexpected, but no interaction was observed between source of selenium and study day on serum anti-ovalbumin IgG titers. Primiparous and multiparous cows had similar responses to immunization withovalbumin, and no treatment parity interaction on serum titers for anti-ovalbumin IgG was observed.

UTERINE HEALTH AND PREGNANCY RATES

Florida and California. Both in Florida and California, frequencies of retained placenta, mastitis, ketosis,displacement of abomasum, and subclinical endometritis were not affected by diets (Table 3). Likewise,frequencies of cows cycling were not affected by diets or reproductive program (Table 3).

Sel-Plex reduced the frequency of multiparous cows detected with more than one event of fever inFlorida (rectal temperature >39.5 C; Sel-Plex, 13.3% [25/188] vs selenite, 25.5% [46/181]; P39.5 C. As in Florida, multiparous cows were lesslikely (P


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1Selenite (n=285 FL, n=285 CA; Sel-Plex, n=289 FL, n=281 CA).

In Florida cervical discharge scores measured at 5 and 10 days postpartum were better for the Sel-Plexgroup (Table 4). The frequency of cows with a purulent-fetid discharge was reduced and proportion ofcows with a clean discharge was increased. These findings lend additional support to the conclusion thatfeeding Sel-Plex in the Florida experiment improved immunocompetence and was associated withimproved uterine health measured at 5 and 10 days postpartum.

In Florida, diet failed to alter first service pregnancy rate at ~30 days post AI or pregnancy losses between~30 and ~55 days post AI (Table 5). These low pregnancy rates and high embryonic losses are typical ofcows managed during the summer heat stress period of Florida. Diet did indeed alter second service

pregnancy rate (Table 5).

The benefit of Sel-Plex on second service pregnancy rate is very interesting. We hypothesize that cowsof the Sel-Plex group were better able to reestablish an embryotrophic environment at second serviceafter either early or late embryonic losses. For example, cows presented for second service may not havebeen pregnant to the first service by 30 days at the ultrasound diagnosis or were pregnant and underwentembryonic loss and required a second service. Indeed, pregnancy rate to the second service for cows thathad lost an embryo was 22.7% for the Sel-Plex group vs 4.2% in the selenite group.

Table 4. Overall frequencies of cervical discharge scores measured at 5 and 10 days postpartum.

Parity (P


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The product Mu-Se (Schering-Plough Animal Health Corp, NJ), which contains inorganic selenium (50mg) andvitaminE (500 mg), increased pregnancy rate to second service (i.e., no effect at first service),reduced services per conception, and decreased the interval from calving to conception (Arechiga et al.,1998).

These results are somewhat similar to the Florida results (Table 5) in which Sel-Plex increasedpregnancy rate to the second service in cows that did not become pregnant at first service. In California,dietary selenium supplements failed to alter conception rates at day 28 and day 56, pregnancy loss from28 to 56 days of gestation, or conception rate at the second timed insemination (Table 5).

MILK PRODUCTION AND SOMATIC CELLS

Florida. Mean milk yield and milk composition determined from monthly samples for cows fedsupplemental inorganic and organic selenium are presented in Table 6. Both monthly milk yield and fat-corrected milk (FCM) were greater for cows fed Sel-Plex.

An additional series of analyses were conducted to determine when the differences in milk productionoccurred. Milk production was monitored on a daily basis for the first 81 days of lactation and mean dailymilk yield did not differ between treatments (35.6 kg/d).

Additional analyses were conducted with the monthly estimates of milk production from monthly DHIAmeasurements and are presented in Figure 4. Monthly milk yields were elevated in the Sel-Plex group inlater stages of lactation between 6-8 months of lactation. The increase in milk production in later stages oflactation occurred primarily in primiparous cows but not in multiparous cows (Figure 4).

Table 6. Effect of selenium source on performance of lactating dairy cows in Florida and Californiaexperiments.
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1SCS = somatic cell score.

2SCC = somatic cell count.

Figure 4. Mean monthly milk yield (kg/day) for primiparous and multiparous cows fed Sel-Plex orsodium selenite. Diet parity month interaction, P


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Figure 5. Mean monthly concentrations of milk somatic cells, Somatic Cell Score (SCS), for

lactating cows fed Sel-Plex or sodium selenite. Parity month interaction P


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increased pregnancy losses compared with those in California (Table 5).

Analyses of the composite feed samples for both experiments identified clear differences in dietaryselenium concentrations. Selenium concentrations in the prepartum diets differed substantially betweenthe Florida (selenite of 0.44 and Sel-Plex of 0.49 mg/kg of DM) and California (selenite of 0.60 and Sel-Plex of 0.74 mg/kg of DM) experiments.

Differences in selenium concentrations (mg/kg of DM) for the postpartum diets were even greater; that is,Florida with 0.36 for control and 0.36 for Sel-Plex and California with 0.71 for control and 0.60 for Sel-Plex. Based upon the higher overall selenium concentrations (i.e., supplemental + background Se levels)in the California experiment, it was not surprising that clinical and immuno-responses did not differ.

In contrast, Florida was essentially a Se-deficient environment in which the virtually all dietary seleniumwas derived via the two supplements. Under these conditions, Sel-Plex (0.33 mg Se/kg) fed in the rationbeginning at 26 days prepartum, elevated plasma selenium concentrations, increased neutrophil functionat the time of parturition, improved immuno-responsiveness in multiparous cows, improved uterine health,and increased second service pregnancy rate during summer in an environment that is selenium deficient.

Such effects were not observed in the California study. It is noteworthy that in the California study, Sel-Plex supplementation during early lactation did result in a substantial increase in milk production. Incontrast, this effect on milk production in early lactation was not evident in the Florida experiment although

Sel-Plex had numerous effects on immunological, uterine health, and pregnancy rate to second serviceoutcomes with a population of cows managed in a selenium deficient environment.

References

Arechiga, C.F., S. Vazquez-Flores, O. Ortz, J. Hernandez-Ceron, A. Porras, L.R. McDowell and P. J.Hansen. 1998. Effect of injection of -carotene or vitamin E and selenium on fertility of lactating dairycows. Theriogenology 50:65-76.

Goff, J.P. 2006. Major advances in our understanding of nutritional influences on bovine health. J. DairySci. 89: 12921301.

Grasso, P.J., R.W. Scholz, R.J. Erskine, and R.J. Eberhart. 1990. Phagocytosis, bactericidal activity, and

oxidative metabolism of mammary neutrophils from dairy cows fed selenium-adequate and selenium-deficient diets. Am. J. Vet. Res. 51:269 274.

Gunter, S.A., P.A. Beck and J.M. Phillips. 2003. Effects of supplementary selenium source on theperformance and blood measurements in beef cows and their calves. J. Anim. Sci. 81: 856864.

Hogan, J.S., K.L. Smith, W.P. Weiss, D.A. Todhunter and W.L. Shockey. 1990. Relationships amongvitamin E, selenium, and bovine blood neutrophils. J. Dairy Sci. 73: 23722378.

Loeffler, S.H., M.J. de Vries and Y.H. Schukken. 1999. The effects of time of disease occurrence, milkyield, and body condition on fertility of dairy cows. J. Dairy Sci.82: 2589-2604.

Mezzetti, A., C. Di Ilio, A.M. Calafiore, A. Aceto, L. Marzio, G. Frederici, and F. Cuccurullo. 1990.Glutathione peroxidase, glutathione reductase and glutathione transferase activities in the human artery,vein and heart. J. Mol. Cardiol. 22: 935- 938.

National Research Council. 2001. Nutrient Requirements of Dairy Cattle. Natl. Acad. Press, WashingtonDC.

Rutigliano, H.M., F.S. Lima, R.L.A. Cerri, L.F. Greco, J.M. Vilela, V. Magalhes, J. Hillegass, W.W.Thatcher and J.E.P. Santos. 2006. Effects of source of supplemental selenium and method ofpresynchronization onreproductionand lactation of dairy cows. J. Dairy Sci. 89 (Suppl. 1):207 (Abstr.).

Silvestre, F.T., D.T. Silvestre, C. Crawford, J.E.P. Santos, C.R. Staples and W.W. Thatcher. 2006a. Effectof selenium (Se) source on innate and adaptive immunity of periparturient dairy cows. Biol. Repro. 39thAnnual Meeting. Special Issue. p. 132.

Silvestre, F.T., D.T. Silvestre, J.E.P. Santos, C. Risco, C.R. Staples and W.W. Thatcher. 2006b. Effects ofselenium (Se) sources on dairy cows. J. Anim. Sci. 89: (Suppl 1.) 52.

Weiss, W.P. 2003. Selenium nutrition of dairy cows: comparing responses to organic and inorganicselenium forms. In: Nutritional Biotechnology in the Feed and Food Industries: Proceedings of Alltechs
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19th Annual Symposium (T.P. Lyons and K.A. Jacques, eds). Nottingham University Press, UK, pp. 333-343.

Authors: FLAVIO T. SILVESTRE1, HELOISA M. RUTIGLIANO2, WILLIAM W. THATCHER1, JOS E.P.SANTOS2 and CHARLES R. STAPLES11 University of Florida, Gainesville, Florida, USA2 University of California-Davis, Tulare, California, USA

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Effect of Selenium Source on Production, Reproduction and Immunity of Lactating Dairy Cows in...

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