Effect of individual and mixed live yeast culture feeding on growth performance, nutrient utilization and microbial crude protein synthesis in lambs

Animal Feed Science and Technology 155 (2010) 163–171

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Effect of individual and mixed live yeast culture feeding on growth performance, nutrient utilization and microbial crude protein synthesis in lambs M.K. Tripathi ∗ , S.A. Karim Division of Animal Nutrition, Central Sheep and Wool Research Institute, Via Jaipur, Avikanagar 304501, Rajasthan, India

a r t i c l e

i n f o

Article history: Received 10 March 2009 Received in revised form 30 November 2009 Accepted 30 November 2009 Keywords: Probiotic Yeast culture Growth Intake Lamb performance

a b s t r a c t This study investigated effects of feeding three individual, and a mixed, yeast culture (Kluyveromyces marximanus NRRL3234, Saccharomyces cerevisiae NCDC42, Saccharomyces uvarum ATCC9080 all in a 1:1:1, ratio) on growth performance, nutrient utilization and microbial crude protein (CP) synthesis in feedlot lambs during the post-weaning phase of growth. Sixty weaner lambs (90 ± 3.5 d old and 15.9 ± 0.50 kg BW) were fed for 91 d in five equal groups. The control group of lambs received sterilized culture medium while the treatment groups were fed a yeast culture in addition to a ad libitum total mixed ration (TMR). The yeast culture, dosed at 1 ml/kg body weight (BW) had 1.5–2.0 × 109 live cells/ml. Yeast culture supplementation did not influence intake and digestibility of organic matter (OM), CP, neutral detergent fiber (NDF), acid detergent fiber (ADF) and hemicellulose and the metabolizable energy (ME) level of the diets were similar between control and yeast supplemented lambs. Lambs in all groups were in positive N balance, but N intake and N voided in feces and urine, as well as N balance, did not change due to yeast culture supplementation. Urinary allantoin excretion was similar, but purine derivatives absorbed (mM/d) were higher (P<0.05) in yeast culture supplemented lambs. Yeast culture supplementation improved (P<0.05) microbial CP synthesis. Supplementation of SC and mixed yeast improved (P=0.002) BW gain of lambs by 21% and 16% respectively. All yeast culture supplemented lambs had higher feed efficiency in comparison to control lambs. Among the three yeast cultures used, S. cerevisiae had the most potential as a growth promoting feed additive in feedlot lamb production, and it may serve as an alternate to antibiotics and ionophores as a growth promoter of weaner lambs. © 2009 Elsevier B.V. All rights reserved.

1. Introduction Weaning of lambs may alter digestive tract microflora because of the shift from a liquid to a solid diet. This diet transition may result in establishment of less desirable intestinal microflora leading to poor performance. A common method of repressing undesirable intestinal microbes has been use of antibacterials, but growing interest had focused on non-antibiotic feed additives because of public concern over drug residues in animal products and development of

Abbreviations: ADF, acid detergent fiber; ADG, average daily BW gain; BW, body weight; CP, crude protein; DM, dry matter; DOMI, digestible OM intake; ME, metabolizable energy; MP, microbial CP; NDF, neutral detergent fiber; OM, organic matter; TMR, total mixed ration. ∗ Corresponding author at: Division of Nutrition, Feed Resource and Product Technology, Central Institute for Research on Goats, Makhdoom, Farah, Mathura 281122, UP, India. Tel.: +91 565 2763380; fax: +91 565 2763246. E-mail addresses: [email protected], [email protected] (M.K. Tripathi). 0377-8401/$ – see front matter © 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.anifeedsci.2009.11.007

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antibiotic-resistant pathogenic organisms (NRC, 1980). Probiotics are viable edible microorganisms, mainly lactic and nonlactic acid bacteria, yeast (dairy strains) and fungi which are capable of producing beneficial effects on the host when fed (15 × 109 live yeast cells/g of powder; AAFCO, 1994 or 2.6 × 109 spores/animal/d; Erasmus et al., 2005) in sufficient numbers. Probiotic microorganisms, by altering the microflora of the digestive tract of the host (Rook and Brunet, 2005), produce beneficial effects on gut ecology. Possible mechanisms of probiotic benefit in the host, as suggested by McNaught and MacFie (2001) and Bontempo et al. (2006), are competition for nutrients, colonization and adhesion of gut mucosa (i.e. competition for receptors), stimulation of mucosal and systemic immunity and/or production of antimicrobial substances. However, the exact mechanism of the beneficial role of probiotics within the animal, and against harmful microorganisms, remains to be delineated. The function of probiotics fed to monogastric animals has been studied to a large extent relative to protection against enteric diseases, control of pre- and post-weaning diarrhea, mortality, shedding of Escherichia coli and growth promotion (Kyriakis et al., 1999; Kritas and Morrison, 2005). However, effects of probiotics in ruminants remain to be investigated intensively. Limited studies have shown that probiotic feeding to cows during pre- and post-parturition increased milk production and proportions of milk fat, lactose and protein in a dose dependent manner (Kim et al., 2006; Kritas et al., 2006; Stein et al., 2006). Calves fed milk replacer supplemented with an antibiotic or a probiotic from birth to 5 weeks of age had similar levels of diarrhea reduction and growth performance (Donovan et al., 2002; Morrill et al., 1977). Responses to probiotic feeding were increased feacal shedding of E. coli, reduced incidences of diarrhea/mortality, and improved growth in lambs and feedlot cattle (Lema et al., 2001; Brashears et al., 2003; Kritas et al., 2006). Probiotic supplementation controls haemolytic E. coli induced diarrhea, which improved growth and reduced mortality of young ruminants (Kritas et al., 2006). Active yeast supplementation had positive effects on performance of young ruminants through increased dry matter (DM) intake and body weight (BW) gain, changes in hip height and width (Lesmeister et al., 2004). The performance promoting effects of live yeast additives could be due to an improvement in rumen development parameters, such as papillae length and width, and/or rumen wall thickness (Lesmeister et al., 2004), as well as early establishment and stabilization of rumen microbial communities (Chaucheyras-Durand and Fonty, 2001, 2002; Newbold et al., 1996) leading to reduced number of days of diarrhea (Galvano et al., 2005). Establishment of a complex rumen microbial ecosystem subsequently improved rumen function that promoted digestion and absorption (Hopper et al., 2001), thereby improving gut health. Yeast probiotics (Saccharomyces cerevisiae strains) were more potent in producing beneficial effects in comparison to monensin (Castillo et al., 2006a). Yeast culture supplementation also helped to maintain a higher rumen pH (Bach et al., 2007) in high grain fed ruminants. Yeast probiotics produce one or more extracellular compounds, due to metabolic activity in the host system, which exert inhibitory action on some pathogenic bacterial strains (Comitini et al., 2005). The bacteriostatic or bactericidal activity of S. cerevisiae is due to production of proteases (e.g. serine proteases, etc.) (Castagliulo et al., 1996; Comitini et al., 2005). Other metabolites of yeast, such as low-molecular weight fatty acids, may also inhibit bacterial growth, indicating that a complex mixture of yeast metabolites is involved in the inhibitory activity (Lemaresquier, 1987; Dick et al., 1992; Comitini et al., 2005). Performance responses of ruminants, including lambs fed yeast and yeast cultures, have been variable. Feeding yeast cultures, Williams and Newbold (1990), Lesmeister et al. (2004) and Stella et al. (2007) reported improved feed consumption, BW gain and feed efficiency for gain, while Erasmus et al. (2005), Agarwal et al. (2002), Mahender et al. (2005), Kim et al. (2006), Kawas et al. (2007) and Tripathi et al. (2008) found similar or reduced growth rate and efficiency of gain compared to the control. Therefore, beneficial responses to yeast cultures as probiotic supplements in ruminant diets are highly variable, and also may be diet and dose dependent (Wallace, 1994). Most positive animal responses were due to modification of the rumen bacterial population (Newbold et al., 1995), and regulation of rumen pH has been ascribed to high efficiency of lactate utilization (Chaucheyras-Durand et al., 1995; Simova et al., 2004) that benefits ruminants in terms of BW gain and milk production by increasing feed intake rather than feed efficiency. The purpose of this study was to evaluate the influence of feeding three yeast strains, and a mixed liquid culture, on growth performance, nutrient utilization and microbial CP synthesis of lambs in order to select the most promising yeast culture for lamb feeding. 2. Materials and methods The experiment was conducted at the Central Sheep and Wool Research Institute, Avikanagar (Rajasthan, India) located at 26◦ 17 N latitude and 75◦ 28 E longitude and 320 m above sea level. The climate is hot and semi-arid. The study initiated in March and ended in June and, during the experiment, minimum and maximum ambient temperature ranged from 15 to 31 ◦ C and 29 to 48 ◦ C, respectively while relative humidity varied from 24 to 70%. 2.1. Animals, housing and feeding Sixty weaner lambs (90 ± 3.5 d old and 15.9 ± 0.50 kg BW) were divided randomly into five equal groups with six males and six females in each. Lambs were penned in well-ventilated enclosures for the experiment and fed individually for 91 d. Lams were fed ad libitum a composite feed (TMR) having a forage:concentrate (F:C) ratio of 250:750 (Table 1). The control group was supplemented with a sterilized culture medium while the other four groups received one of the three, or a mixed, yeast culture. Fresh feed was offered daily at 9:00 h after discarding the residue in excess of 0.10 of previous day’s intake, while the daily dose of yeast cultures was drenched at 1 ml/kg BW. The lyophilised cultures were procured from the

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Table 1 Ingredient and chemical composition of the diet fed to weaner lambs. g/kg as mixed Ingredient composition Forages sources Khejri (Prosopis cenraria) leaves Pala (Zuzuphus numularia) leaves Concentrate source Maize grain Barley grain Groundnut oil meal Mustard oil meal Supplements Mineral premixa Common salt Vitamin premix (Vitablend)b Chemical composition (g/kg DM)c Dry matter (g/kg) Organic matter Crude protein Neutral detergent fiberd Acid detergent fiberd Cellulosed Lignin(sa) Hemicellulosed a b c d

125 125 280 280 80 80 18 10 2 Total mixed ration

Khejri

Pala

977.0 902.3 166.2 603.7 324.9 126.1 62.0 278.8

962.8 840.5 105.0 481.8 261.7 141.1 75.1 220.1

958.2 842.1 96.2 624.9 527.3 164.3 116.4 97.6

Mineral mixture contained (g/kg): Ca 320, P 62, Mn 2.7, Zn 2.6, Fe 1, F 0.9, I 0.1, Cu 0.1 (Ranmix, Ranbaxy Fine Chemicals Limited, New Delhi, India). Vitamin premix contained (IU/g): vitamin A 50,000, vitamin D3 5000 (Agrivet Farm Care, GlaxoSmithKline Pharmaceuticals Limited, Mumbai, India). Mean of three observations. Determined using a sequential procedure.

National Collection of Dairy Cultures, National Dairy Research Institute in Karnal, India. These cultures were activated on skimmed milk and used for bulk production by subculturing every 30 d. The yeast cultures were bulk produced on a sterilized (autoclaved at 1.05 kg/cm2 and 121 ◦ C for 15 min) liquid medium containing glucose 20 g, peptone 20 g and yeast extract 10 g in each liter of distilled water with pH 6.5 ± 0.2 at 28 ◦ C. Sterilized liquid medium was inoculated aseptically with 10 ml liquid culture and incubated at 28 ◦ C for 36 hrs, with shaking for 60 cycles/min. Three stains were cultivated separately having 1.5–2.0 × 109 live cells/ml. In mixed culture drenching, the live yeast cultures of Kluyveromyces marximanus NRRL3234 (KM), S. cerevisiae NCDC42 (SC) and Saccharomyces uvarum ATCC9080 (SU) were mixed in ratio of 1:1:1 just before dosing. The TMR contained essential constituents recommended for native growing lambs (ICAR, 1998) and the composition of the diet, forage source and TMR is in Table 1. Feed samples were collected at weekly intervals for DM determination and 3- or 4-week samples were pooled for chemical analysis. Free choice water was available from 10:00 to 16:00 h. 2.2. Growth and metabolism experiment The growth experiment lasted for 91 d in a randomized design during which lamb BW were recorded for 2 consecutive days every 7 d and these values were used to determine BW gain. Pattern of growth was calculated based on these 7 d periods. The change in BW was calculated as: Y = a + b1 X + b2 X2 , where Y is change in BW (kg), a is constant of growth (i.e. BW at start), b1 BW on X d, b2 rate of change in BW on X d, X d is days of experiment. A metabolism experiment was conducted near the end of experimental feeding (i.e. 80th d), on six randomly selected lambs from each treatment, for 10 d (i.e. 3 d adaptation followed by 7 d of sample collection) during which daily feed intake and output of feces and urine were collected and recorded. Samples of feed, orts, feces and urine were collected every morning. Feces and urine were collected into acidified containers containing 100 ml H2 SO4 (100 ml concentrated H2 SO4 diluted to 1000 ml with distilled water) using a total collection method. Urine pH was maintained below 3 by addition of diluted H2 SO4 (10 ml/100 ml distilled water) and collected urine was diluted with distilled water to a volume of 2000 ml and a representative sample was frozen and preserved for purine derivative analysis. For chemical analysis samples of feeds, feces and orts, samples were dried to a constant weight in a forced air oven at 70 ◦ C. Dried samples of the 7 d collection were pooled and ground to pass a 1 mm screen and preserved for chemical analysis. 2.3. Chemical analysis The DM of feed, orts and feces were analyzed by drying at 100 ◦ C for 24 h or to constant weight. The AOAC (1995) analytical procedures were used for the OM determination (no. 968.08) by ashing at 550 ◦ C for 4 h and N estimation (no. 988.05)

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by a Kjeldahl technique. Neutral detergent fiber (NDF) and acid detergent fiber (ADF) were determined by a sequential procedure using the same sample. For NDF determination, the procedure of Van Soest et al. (1991) was used without sodium sulfite or ␣-amylase, whereas the procedure described by Robertson and Van Soest (1981) was used for ADF and lignin(sa) determination. The NDF and ADF values are expressed with residual ash. Metabolizable energy (ME) was calculated according to AAC (1990) as: ME (MJ/kg DM) = [(digestible OM, g/kg DM)/1000] × 18.5 × 0.81. Allantoin in urine was estimated according to the method described in IAEA (1997). Since allantoin excretion constitutes 0.7 of the total purine derivatives excreted in urine, hence a factor of 0.7 was used to compute excretion of purine derivatives (Wina et al., 2005). Therefore, urinary excretion of allantoin (Y, mmol/d) was used to calculate microbial purines absorbed (X, mmol/d) from the equation: Y = 0.84X + (0.150 × BW0.75 × e−0.25X ), where BW was animal BW (kg). Microbial N supply (g/d) was calculated from the relationship: 70X/(0.83 × 0.116 × 1000) where 70 is the N content of purines (mg N/mmol), X is as defined above, 0.83 is the assumed digestibility of microbial purines, 0.116 is the ratio of purine N/total N in mixed rumen microbes and 1000 converts mg to g (Chen and Ørskov, 2004). 2.4. Statistical analysis Data on feed intake, growth performance, feed efficiencies, nutrient utilization and microbial CP synthesis were subjected to analysis of variance for statistical significance using a general linear mathematical model as: Yijk =  + Ti + eij where Yijk = observation mean; ␮ = general mean, Ti = effect of ith treatment (i = 1, 5), eij = random error. Significant differences among the dietary treatments were separated using Duncan’s multiple range test. Relationships to describe the influence of different yeast cultures on growth were determined by regression analysis using SPSS Base 14.0 (SPSS, 2005). Weekly BW change of individual lambs was determined by fitting polynomial curves. Treatment differences among growth constants were analyzed using an analysis of variance procedure as above. 3. Results 3.1. Dry matter intake, nutrient digestibility and nutritive value of diets The DM intakes in terms of g/d or relative to BW were similar in lambs fed TMR with or without yeast culture (Table 2). Intake and digestibility of OM, CP, NDF, ADF and hemicellulose were also similar among groups. Yeast culture feeding did not affect the ME contents or N:ME ratio of the TMR (Table 3). Table 2 Dry matter intake, nutrient intake and digestibility coefficients of diets fed to weaner lambs with different live yeast cultures. Dietary treatments (type of yeast culturea ) Control Body weight (kg) Dry matter intake (g/kg BW)

KM

SC

SEM SU

Mixed yeastb

27.9 29.8

27.8 34.5

32.0 31.3

27.3 34.0

Intake and digestibility Dry matter Intake (g/d) Digestibility

833 0.66

937 0.65

988 0.62

929 0.67

1025 0.64

44.6 0.012

Organic matter Intake (g/d) Digestibility

752 0.69

845 0.67

891 0.64

838 0.69

925 0.66

40.3 0.011

Crude protein Intake (g/d) Digestibility

139 0.61

156 0.61

164 0.59

154 0.62

170 0.60

7.4 0.015

Neutral detergent fiber Intake (g/d) Digestibility

503 0.60

565 0.60

596 0.56

561 0.61

619 0.58

26.9 0.012

Acid detergent fiber Intake (g/d) Digestibility

271 0.44

304 0.43

321 0.39

302 0.42

333 0.41

14.5 0.017

Hemicellulose Intake (g/d) Digestibility

232 0.78

261 0.78

275 0.76

259 0.83

286 0.77

12.4 0.012

a b

KM: Kluyveromyces marximanus; SC: Saccharomyces cerevisiae; SU: Saccharomyces uvarum. Mixed culture of the three yeasts in equal proportions, supplemented at 1 ml/kg BW.

29.2 34.9

0.97 1.31

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Table 3 Nutritive value of the diets digestible OM and ME intake of weaner lambs fed diets with different live yeast cultures. Dietary treatments (type of yeast culturea ) Control Nutritive value of the diets Metabolizable energy (MJ/kg DM) N:ME ratio (g N/MJ ME) Digestible OM and ME intake Digestible OM intake (g/d) ME intake (MJ/d) a b

KM

SEM

SC

b

SU

Mixed yeast

9.34 2.85

9.04 2.98

8.66 3.11

9.29 2.87

8.90 3.01

0.149 0.056

516 7.73

561 8.41

565 8.47

571 8.55

610 9.14

25.4 0.380

KM: Kluyveromyces marximanus; SC: Saccharomyces cerevisiae; SU: Saccharomyces uvarum. Mixed culture of the three yeasts in equal proportions, supplemented at 1 ml/kg BW.

Table 4 Dietary N utilization and rumen microbial CP synthesis of weaner lambs fed diets with different live yeast cultures. Dietary treatments (type of yeast culturea ) Control

KM

SC

SU

Mixed yeast

22.2

24.9

26.3

24.7

27.3

1.19

8.6 4.2

10.0 5.0

11.1 4.0

10.0 4.5

11.1 4.4

0.68 0.18

Total

12.9

14.9

15.1

14.0

15.5

0.76

N balance g/d g 100/kg BW

9.3 33.4

10.0 35.8

11.2 35.0

10.7 39.2

11.8 40.4

0.73 0.26

51.0 5.57a 27.8a 56

69.1 7.46ab 39.2ab 71

76.8 9.48b 50.5b 88

59.2 6.07a 30.7a 59

65.5 7.52ab 39.2ab 73

3.30 0.440 2.54 4.9

N utilization (g/d) N intake N voided Feces Urine

Microbial crude protein (MP) synthesis Allantoin excreted (mg/kg BW0.75 ) Purine derivatives excreted (mM/d)c Microbial CP synthesis (g/d)c Efficiency of microbial CP synthesis (g MP/kg DOMI) a b c

SEM b

KM: Kluyveromyces marximanus; SC: Saccharomyces cerevisiae; SU: Saccharomyces uvarum. Mixed culture of three yeasts in equal proportions, supplemented at 1 ml/kg BW. Values bearing different letter in a row differ (P<0.05).

3.2. Nitrogen (N) utilization and microbial protein synthesis All lambs were in positive N balance (Table 4), but N intake, voided in feces and urine and balance did not change due to yeast culture supplementation. Urinary allantoin excretion was similar, but purine derivatives absorbed were higher (P<0.05) in SC yeast culture supplemented lambs. SC yeast culture supplementation increased (P<0.05) microbial CP synthesis, but efficiency of microbial CP synthesis relative to digestible OM intake (DOMI) was not influenced. 3.3. Feed intake, feed efficiency, growth performance and growth constants Weaning and finishing BW of the lambs did not differ, but SC and Mixed yeast culture improved (P=0.002) BW gain (Table 5). All yeast culture supplemented lambs had higher feed efficiency, sometimes due primarily to higher growth rate (i.e. SC and mixed culture) and sometimes due to a combination of higher growth rate and lower feed intake. The SC culture supplemented lambs had a consistently higher growth rate and that the difference with other group of lambs increased with progressive of the experimental period (Table 6). The growth constants were similar among groups, but R2 was lower in KM (P<0.05) indicating that variation in growth was relatively higher in KM lambs in comparison to other groups. 4. Discussion Active yeast products have beneficial effects in ruminant livestock production as feed additives to improve feed efficiency and growth performance (Kmet et al., 1993). Effects include improvement of rumen function by favoring microbial establishment, stabilization of rumen pH in animals fed high cereal diets, interaction with lactate metabolizing bacteria and yeasts, and increased fiber degradation due to more cell wall degrading microorganisms (Chaucheyras-Durand et al., 2008; Yang et al., 2004). The primary objective of this study was to evaluate three yeast strains for their suitability as probiotics for lamb growth promotion. It was expected that the yeast strains would have varying degree

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Table 5 Growth performance of Malpura weaner lambs fed diets with different live yeast cultures. Dietary treatments (type of yeast culturea ) Control

KM

Performance Weaning weight (kg) Finishing weight (kg) Average daily gain (g)c

16.0 29.2 145b

15.5 27.9 136b

Feed intake kg (91 d)c g/d

89.1a 979a

70.2b 77b

94.7a 1040a

6.8b 15.1b

5.7a 17.7a

5.6a 17.9a

Feed efficiency kg feed/kg gainc kg gain/100 kg feedc a b c

SC 16.5 33.2 184a

SEM b

SU

Mixed yeast

15.2 30.0 162ab

16.2 32.0 173a

84.2ab 925ab

87.3a 955a

2.58 28.4

5.6a 18.0a

0.12 0.31

5.7a 17.7a

0.50 0.76 4.5

KM: Kluyveromyces marximanus; SC: Saccharomyces cerevisiae; SU: Saccharomyces uvarum. Mixed culture of the three yeasts in equal proportions, supplemented at1 ml/kg BW. Values bearing different letter in a row differ (P<0.05).

of performance promotion and the most promising strain could be considered for further evaluation and development. 4.1. Dry matter intake and nutrient utilization That yeast culture supplementation did not affect DM intake and nutrient utilization in growing lambs is consistent with Robinson and Erasmus (2009) who concluded that a consistent small improvement in DM intake (2.0–3.7%) occurred in response to yeast product supplementation. The increased digestibility of poor quality forage often reported in response to live yeast supplementation (Guedes et al., 2008; El-Ghani, 2004; Yang et al., 2004) may be due to stimulated growth of cellulolytic bacteria (Plata et al., 1994; Girard and Dawson, 1995; Callaway and Martin, 1997). However, as our lambs were fed a high concentrate diet (0.75 concentrate) beneficial effects of yeast supplementation on digestibility improvements are consistent with Pinos-Rodrigues et al. (2008), even though increased polysaccharide degrading activities in the rumen of sheep fed high concentrate diets in the presence of an active yeast culture were observed, but the impact on digestibility enhancement was small (Jouany et al., 1998; Chaucheyras-Durand and Fonty, 2006). Castillo et al. (2006a) did not observe the expected response of yeast supplementation in steers fed a high grain diet with a high CP level and indicated that the high CP content of the ration could have been responsible for the lack of response to yeast supplementation, possibly because the CP may have acted as a systemic buffer as ammonia (Castillo et al., 2006b). Ruminants fed high concentrate diets have increased acid absorption from the rumen, and the additional ammonia produced from amino acid catabolism may enhance this systemic buffer effect to counteract the high acid loads (Swartz et al., 1994). Positive effects of yeast feeding in ruminants can be substantial in diets containing low levels of CP (Ando et al., 2004) and rations containing dietary CP below 125 g/kg DM have shown improvements in feed degradation and animal performance (Castillo et al., 2006b). As the TMR fed to our lambs had a CP level of 166 g/kg DM, the better performance from these yeasts is consistent with previous findings. 4.2. Nitrogen (N) utilization and microbial protein synthesis Yeast culture supplementation has been reported to enhance microbial growth and decrease N loss by incorporating more digestible carbohydrates into microbial mass (Sniffen et al., 2004). Change in rumen N metabolism in the presence of yeast is due to inhibitory activities of proteolytic rumen bacteria that limit degradation of protein and peptides. The Table 6 Rate of change in lamb body weight (kg) with feeding period (d) on diets fed with different live yeast cultures. Dietary treatments (type of yeast culturea )

Constant (BW kg, 0 d) b1 c b2 d R2 e

SEM b

Control

KM

SC

SU

Mixed yeast

15.5 0.88 0.02 0.99ba

15 0.73 0.02 0.98b

16 0.93 0.03 0.99a

14.8 0.79 0.03 1.00ba

15.8 0.85 0.03 0.99a

Values bearing different letter in a row differ significantly (P<0.05). a KM: Kluyveromyces marximanus; SC: Saccharomyces cerevisiae; SU: Saccharomyces uvarum. b Mixed culture of the three yeasts in equal proportions, supplemented at 1 ml/kg BW. c Lamb BW on X d of feeding. d Rate change in lamb BW with X d feeding. e Coefficient of growth.

0.50 0.058 0.004 0.001

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mechanism of yeast action may be due to competition between yeast cells and bacteria for energy supply and/or by direct inhibitory effects of yeast on small peptides and bacterial peptidases (Chaucheyras-Durand et al., 2008). With a desirable dietary balance between soluble N and carbohydrate supply, the dietary level of soluble N is a key parameter in a production response to probiotic yeast because stimulation of rumen bacterial numbers provide more microbial CP to the host (Kamel et al., 2004). However, some yeast products did not have a positive response on the amount and composition of microbial CP reaching the duodenum (Erasmus et al., 2005; Putnam et al., 1997). In the present study, we also did not observe an effect of supplementation of live yeast preparations on rumen N metabolism in lambs. Although, microbial CP supply was improved (2.9–22.7 g) in yeast culture supplemented lambs, the efficiency of microbial CP supply did not differ. The variable response of yeast culture feeding in our study supports the views of Wallace (1994) and Beauchemin et al. (2003) that positive effects of yeast probiotics in ruminants are variable, and may be strain dependent. 4.3. Feed and nutrient efficiency, and lamb performance Feed intake, growth performance and feed efficiency were in reported ranges for feedlot lambs (Tripathi et al., 2004, 2007a,b; Raghuvansi et al., 2007). Supplementation of SC and mixed culture yeast improved growth performance and feed efficiency for growth. Maximum improvement in daily growth (39 g) and feed intake (52.5 g) was with SC, but feed efficiency was improved in all yeast supplemented lambs. The improvement in growth was a cumulative effect of enhanced feed intake, better feed efficiency and possibly higher microbial CP supply in the SC and mixed yeast supplemented lambs, supporting the hypothesis that growth promotion by yeast probiotics is due to improved feed intake and enhanced feed efficiency (Williams and Newbold, 1990). Most of the positive animal responses to yeast feeding are thought to be due to modifications of the rumen bacterial population (Newbold et al., 1995) and regulation of rumen pH due to higher efficiency of lactate utilization. Yeast strains differ in their ability to produce such responses (Chaucheyras-Durand et al., 1995; Simova et al., 2004), which has been ascribed to the varying potential of yeast strains to produce of one or more extracellular compounds related to its metabolic activity (Castillo et al., 2006a; Bach et al., 2007; Comitini et al., 2005; Lemaresquier, 1987; Dick et al., 1992). SC feeding is known to reduce heat stress and improve performance (Bruno et al., 2009) and, as the present experiment was during hot semi-arid conditions, the SC culture possibly reduced the impact of heat stress on lambs leading to improved growth performance. 5. Conclusions Yeast culture supplementation did not influence DM intake, nutrient digestibility, N utilization or ME intake of weaner lambs. S. cerevisiae and mixed culture (i.e. S. cerevisiae, K. marximanus and S. uvarum; 1:1:1 ratio) supplementation improved feed intake, daily gain, feed efficiency and microbial CP synthesis. The K. marximanus culture improved feed efficiency, but did not improve performance, while growth of the S. uvarum fed group was improved by 10.5%, with a similar level of improvement in feed efficiency over control lambs. 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