ANIMAL FEED SCIENCE AND TECHNOLOGY

ELSEXIER

Animal Feed Science Technology 63 (1996) 149- 162

Effects of a probiotic yeast in lactating ruminants: interaction with dietary nitrogen level
S. Giger-Reverdin a* *, N. Bezault a, D. Sauvant a, G. Bertin b
aLaboratoire de Nutrition et Alimentation (INRA) de lINA-PG, 16 rue Claude Bernard, 7.5231 Paris Cedex
05, France b SANTEL

Sante' Animale, Direction R & D, 85 rue Anatole France, 92300 Leuallois-Perret, France Accepted5 March 1996

Abstract
This study was designed to test the effects of a probiotic yeast (Saccharomyces cereuisiae at two levels of nitrogen intake in early lactation, expressed as PDI (Protein truly Digestible in the small Intestine) at normal levels of 108gPDIkg- DM (dry matter) vs. low levels of 78 g PDI kg- DM, on dry matter intake, milk production and composition and on blood parameters. Twenty-eight dairy goats received a complete diet including alfalfa hay (25% of dry matter), pressed sugar beet pulp silage (50%) and a compound feed (25%). Supplying yeast decreased energy and nitrogen balances during the first 6 weeks after parturition. Yeast seemed to facilitate increased mobilization of body reserves and to increase milk fatty acid production. Thus, fat-corrected milk yield increased during the period when animals are very susceptible to nutritional stress.
I-1077) Keywords:

Probiotic yeast; Dairy ruminants; Metabolism; Milk yield

1. Introduction The ability of probiotic yeasts to increase dry matter (DM) intake, modify rumen stoichiometry, and increase milk yield and milk protein content in lactating ruminants has been established for early lactating ruminants (Giinther, 1989; Harris and Webb, 1990; Williams et al., 1991). However, not all studies have found these effects (Arambel and Kent, 1990; Wohlt et al., 1991). Nevertheless, researchers agree that yeast limits the decrease in pH when large quantities of easily fermentable carbohydrates are ingested,
Corresponding author.

0377-8401/96/$1_5.00Copyright 0 1996 Elsevier Science B.V. All rights reserved.

PII SO377-8401(96)01011-5

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Feed Science Technology 63 (1996) 149-162

and seems to stimulate microbial activity, especially that of the cellulolytic microflora (Arambel and Kent, 1990; Williams and Newbold, 1990; Williams et al., 1991). The aim of this study was to clarify the effects of a probiotic yeast (Succharomyces cereuisiae I- 1077) whose positive influence on ruminal production of volatile fatty acids has already been shown (Jouany et al., 1991) in ruminants at the onset of lactation, when animals are affected by negative nutritive balances, and are thus more susceptible to benefit from the use of such products (Chase, 1989). This study used two nitrogen (N) levels to test potential interactions (yeast X N supply), and especially to investigate the possible beneficial effects of yeast with nitrogen-deficient diets. In addition to the different effects of yeast on fundamental zootechnical parameters (e.g. intake, milk yield), nutritional balances, blood parameters and fatty acids secretion in milk were measured in the same animals.

2. Materials and methods 2.1. Experimental

design and treatments

The study was conducted on 28 dairy goats in the experimental herd of the Nutrition and Feeding Station (INRA) of the Institut National Agronomique Paris-Grignon. Alpine or Saanen goats were assigned to seven blocks of four animals each as a function of parity (two blocks for the first lactation), prior milk production for multiparous goats and estimated production for primiparous, type of aS1 casein, and liveweight. Animals within each block were randomized among the following four groups: CLw -control, low nitrogen: CNl-control, normal nitrogen; YLw-yeast, low nitrogen; YNI-yeast, normal nitrogen. The four groups made up a 2 X 2 factorial design, within which each of the three effects (yeast, nitrogen, parity) were tested in 14 pairs. 2.2. Diets All goats were fed on alfalfa hay (25% of DM), pressed sugar beet pulp silage (50%) and a compound feed (25%), in which the composition was varied according to the

Table 1 Composition

of compound

feeds Diet Low in nitrogen (Lw) Normal in nitrogen (NI) 18.5 92 185 463 45 30 Zn 7500ppm. Mn

glcg-
Barley Maize grain Oats Soyabean meal Sodium bicarbonate Mineral and vitamin premix a

370 370 185 0 45 30

Premix contained P 20.0%. Ca O.O%, Mg 5.0%, S 2.0%, Cu ISOOppm, Fe 3000ppm, 4500ppm, Vitamin A 45OoOOIU, Vitamin D, 15OOOOIU, Vitamin E 3001U.

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desired N level, in a total mixed ration offered twice daily (Table 1). The energy and N contents of the feeds were estimated, with the French feed unit systems (Vermorel, 1988; V&it6 and Peyraud, 1988): net energy value was 6.82MJ kg- DM for the whole diet. The PDI (Proteins truly Digestible in the small Intestine) levels were respectively 108gkg- for the normal (Nl) N diet and 78 g kg- for the low (Lw) N diet. Yeast was provided by SANTEL Sante Animale and was from Succharomyces cereuisiue strain (I-1077). It was given once daily to animals in the yeast groups (YLw and YNl) with a dosing gun at the beginning of the afternoon. The daily dose was 200 mg of yeast at a concentration of 11 X 109 g - * CFU (colony forming units) in 20 ml of tap water. Each goat thus received 2.2 X lo9 CFUday- from the day of parturition until the end of the sixth week of lactation. Assuming that the rumen of a goat at the onset of lactation contains approximately 91 of liquid, the concentration of the yeast suspension was about 2.45 X 10 cellsml- of rumen fluid. 2.3. Sampling and measurements Feed intake levels were calculated from daily individual measurements of the quantities of feed offered and refused, averaged weekly. The DM contents of the components of the diet were determined weekly. Classical feed analysis (crude fibre, crude protein and ash) allowed definition of the nutritional value of the feeds (MJ of net energy, g of PDI) and thus of the diet. Animals were weighed weekly. The milk yield of each goat was recorded on three consecutive days during each of the experimental weeks and a weekly average was calculated. Milk samples were taken from two consecutive milkings to composite the weekly sample which was analysed for fat, protein and lactose. Additional milk samples collected during the first, second, fourth and sixth weeks of lactation were used to determine DM, ash and urea contents. Milk fatty acids were extracted by a modification of the Rose-Gottlieb method (1926) as described by Bas (19851, and measured by gas-liquid chromatography. Blood samples were collected weekly in heparinized tubes from the jugular vein of the animals, before offering the morning meal, during the first, second, fourth and sixth

Table 2 Effects of main factors on zootechnical Diet YLW MW (kg) DMI (gkg- MW) NE1 (k.lkg- MW) PDI (gkg- MW) 20.8 86.0 589 6.69 a a a a CLW 20.5 a 87.4a 598 6.82 p

parameters Primiparous YNI 19.7 a 88.5 a 606 a 9.54 b CNl 20.4 a 85.1= 581a 9.24 b 18.3 = 83.7 574 7.77 22.4 a 89.8 614 8.37 1.3 6.0 42 0.57 Multiparous SEM Significance N NS NS NS *** P ** NS NS NS

YLw, Yeast and low in nitrogen; CLw, control and low in nitrogen; YNI, yeast and normal in nitrogen; CNl, control and normal in nitrogen. N, Nitrogen; P, parity. MW, Metabolic weight; DMI, dry matter intake; NEI, net energy intake; PDI, protein truly digestible in the small intestine. NS, Not significant; * P < 0.05; ** P <0.01;*** P < 0.00 1. Yeast effect, Y X N, Y X P, N X P and Y X N X P interactions are not significant. Means not bearing the same superscript letters within rows am significantly different (P < 0.05).

Table 3 Milk production Primiparous YNl CNI Y N YXN P YXP Multiparous SEM Significance

characteristics

of goats

Diet

YLW

CLW

YXNXP

B E & 3 e

2
NS 5 NS NS NS NS NS
l

42.6 axb 33.2 a 49.8 a 7.6 a 134.2 a 0.31 = 178 a 202 a 8.33 b 5.77 a 8.84 a 159 175 6.87 5.35 8.06 166 a 178 a.b 6.62 a.b 5.71 a 8.22 a 170 188 7.4s 5.58 8.21 15 18 0.77 0.52 0.73 NS * NS NS NS NS NS NS NS NS NS NS NS NS

47.6 ab 32.9 a 49.6 7.7 a 137.1 a 0.48 b 44.0 34.4 50.7 u 7.8 137.4 0.36 1.7 0.7 0.7 0.2 2.7 0.02 ** NS NS NS NS NS

40.9 b 35.1 a 49.2 a 8.0 134.8 a 0.43 b

44.0 33.2 48.3 u 7.8 133.9 0.37

NS NS NS NS NS **

NS NS ** NS NS NS NS NS NS NS NS

NS * NS NS NS NS NS NS NS NS NS

*
NS NS NS NS NS NS NS NS NS

g.
e $ B @ % !I 2 % g F 2 z!J 2 g 2 % 5

Milk composition (g kg- ) 45.1 a.b Fat Protein 34.0 a 49.5 a Lactose Ash 7.8 a Dry matter 136.3 a 0.25 a Urea Yield per day (gkg- MW) Milk 177 a FCM 199= Fat 7.99 a.b Protein 5.95 a Lactose 8.68 a

137 a 147 b 5.72 a 4.43 a 6.79 a

YLw, Yeast and low in nitrogen; CLw, control and low in nitrogen; YNI, yeast and normal in nitrogen; CNI, control and normal in nitrogen. Y, Yeast; N, nitrogen; P, parity. MW, Metabolic weight; FCM, 35 g kg - fat-corrected milk. Means not bearing the same superscript letters within rows are signiticantly different (P < 0.05). NS, Not significant; *P < 0.05; * * P < 0.01; * * P < 0.001. N XP interaction is not significant.
l

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weeks of lactation. Plasma was assayed for glucose by a modified hexokinase-glucose6-phosphate dehydrogenase procedure (Cooper, 19731, for non-esterified fatty acids (NEFA) by an adaptation of the enzymatic procedure described by Bas (19841, for beta-hydroxybutyrate (BHB) by the procedure of Bamouin et al. (1986) and for urea by a modification of the method of Talke and Schubert (1965). All methods were adapted for a Coulter semi-automatic instrument (P. Bas and A. Rouzeau, unpublished data, 1992). 2.4. Statistical analysis Data were analysed using the general linear model (GLM) procedure of SAS Institute, Inc. (1987).

where Yijk is dependent variable, p is overall mean, cx is effect of yeast (Y), p is effect of N level (N), y is effect of parity of animals (PI, OL X p is Y-N interaction (Y X N), (YX y is Y-P interaction (Y X P), (3 X y is N-P interaction (N X P), cxX p X y is Y-N-P (triple interaction: Y X N X P) and eijk is experimental error. The repeated data version of GLM was used because all the weeks were considered in the same analysis to test the mean effects of different factors. Results are expressed as least-square (LS) means. 3. Results One multiparous goat in Group CM died during the fifth week of lactation and was removed from the statistical analysis.

e-c-

Yeaalandbwinnilropen

Fig. 1. Relationships between 35 g kg- fat corrected milk yield and time post-p&m.

Table 4 Effect of main factors on the percentage Primiparous CNI N YXN P NS NS NS NS NS NS

NS NS

of fatty acids in milk Multiparous SEM Significance YXNXP

Diet

YLW NS NS NS NS NS NS NS * NS

CLW

YNI

a a a a a

* *t
NS
I *

a a = b b a = a.b a a a a

NS NS NS * NS NS NS NS NS NS NS

c4:o C6:O C8:O c1o:o c12:o c14:o c15:o Cl6:O Cl6:l c17:o C17:I Cl&O Cl8:I Cl&2 a a a a.b b a a a.b a a a. a a = 2.05 2.51 2.93 10.55 5.01 11.13 1.07 28.47 1.42 0.98 0.54 6.15 20.3 I 2.55

1.99* 2.47 a 2.96 a 11 12 a.b 5:9 1 ?xb ll.14a 1.11 a 27.41 a 1.44 a 0.96 a 0.55 a 5.83 a 19.65 a 2.80 a

2.08 2.64 3.18 12.10 6.29 ll.96a 1.13 31.19 1.33 0.79 0.37 5.00 15.16 2.29

a b p b b a b b

2.06 2.58 3.03 10.90 4.83 11.22 0.95 30.04 1.42 0.92 0.52 5.37 19.37 2.59

a
a 37.67 2.60 31.09 27.36 37.64 2.36 32.53 26.19

2.10 2.62 3.10 11.31 5.50 11.12 1.08 28.57 1.32 u 1.01 u 0.52 6.27 u 18.50 2.59 2.00 2.48 2.95 11.01 5.52 11.61 1.06 29.99 1.48 p 0.82 p 0.48 4.91 fi 18.74 2.54 * NS * NS 1.27 0.13 1.10 1.69 * NS NS NS NS * * * 0.13 0.12 0.1 1 0.45 0.41 0.35 0.09 1.13 0.08 0.05 0.04 0.48 1.23 0.15 NS NS NS NS NS ** NS NS NS NS NS NS NS NS

NS NS
NS NS NS NS

NS NS NS
NS NS NS NS

z c4-Cl4 37.71 a.b I: Cl5 and 2.63 a Cl7 Z Cl6 29.95 a z Cl8 28.29 a 31.08 a.b 29.01 a

40.42 a 2.30 a

36.38 b 2.40 a

36.10 b 2.60 a

33.63 b 22.45 b

32.56 a.b 27.34 a.b

YLw, Yeast and low in nitrogen; CLw, control and low in nitrogen; YNl, yeast and normal in nitrogen; CNl, control and normal in nitrogen. Y, Yeast; N, nitrogen; P, parity. NS, Not significant; P < 0.05; * * P < 0.01; * * * P < 0.001. Yeast effect and Y X P and N X P interactions are not significant. Means not bearing the same superscript letters within rows are significantly different (P < 0.05).

Table 5 Effects of main factors on the production of milk fatty acids (gday- kg- metabolic weight) Primiparous SEM Y * YNI CNI Multiparous Significance

t&s-

Diet

YLW

CLW

c4:o

c6:o

t * .
NS * NS *

*
I

0.15 0.18 0.22 0.81 0.41 0.86 0.08 2.22 0.11 0.06 0.37 I .42 0.19 2.78 0.18 2.41 1.99 0.02 0.02 0.02 0.09 0.06 0.09 0.01 0.22 0.01 0.0 1 0.01 0.06 0.17 0.02 0.30 0.02 0.24 0.25

** NS * **
NS * * **

C8:O Cl@0 Cl20 Cl40 c15:o Cl6:O Cl6:l c17:o Cl7:l Cl&O Cl8:l C18:2 3.02 B 0.20 a.b 2.69 a,b 2.30 a 2.38 a 0.17 a.b 2.05 a.b 1.93 a.b 2.59 0.18 2.12 1.90

0.16 a,b 0.19 a*b 0.23 a 0.88 a 0.47 a 0.88 a 0.09 a 2.21 a*b 0.12 p 0.07 a 0.04 a 0.46 a 1.58 a 0.22 a 0.17 b 0.21 b 0.25 a 0.90 a 0.40 a 0.93 a 0.08 a 2.48 b 0.12 a 0.08 a 0.04& 0.45 a 1.63 a 0.22 a.b 0.13 a.b 0.16 a.b 0.19a 0.70 a 0.33 a 0.73 a 0.07 a I .88 a.b 0.09 a.b 0.07 a.b 0.04 a 0.40 a 1.35 a.b 0.17 a.b 0.14 0.18 0.21 0.78 0.38 0.76 0.07 1.95 0.09 0.07 0.04 0.43 1.28 0.18

0.12 a 0.15 a 0.18 a 0.70 a 0.36 a 0.69 a 0.07 a 1.76 a 0.07 b 0.04 b 0.02 b 0.30 a 0.85 b 0.13 b

z Z Z Z

c4-Cl4 Cl5 and Cl7 Cl6 Cl8

2.99 a 0.21 a 2.41 a 2.26 a

2.34 a 0.13 b 1.90 b 1.28 b

YLw, Yeast and low in nitrogen; CLw, control and low in nitrogen; YNI, yeast and normal in nitrogen; CNI, control and normal in nitrogen. Y, Yeast. NS, Not significant; * P < 0.05; * P < 0.01; * * * P < 0.001. N (nitrogen) and P (parity) effects, such as Y X N, Y X P, N X P and Y X N X P interactions are not significant. Means not bearing the same superscript letters within rows are significantly different (P < 0.05).

Table 6 Effects of main factors on nutritive balances and on plasma parameters Primiparous YNI - 268.4 a -2.21 b 0.523 a 49.2 b 712 b 0.535 b - 227.0 - 2.71 0.577 OL 44.2 687 a 0.409 -214.9 - 2.69 0.517 p 39.1 436 p 0.411 37.2 0.56 0.022 4.5 94 0.028 * ** NS * * NS - 235.8 a - 1.35 b 0.555 B 42.9 a.b 550 a 0.472 NS ** NS NS NS *** CNI Y N Multiparous SEM Significance YXN NS NS NS NS NS

Diet

YLW

CLW

P NS NS * NS * NS

YXP NS NS NS NS NS

EB &Jkg- 1 PDIB (gkg- 1 Glycaemia (g l- 1 BHB (mgl-) NEFA (mmol I- Uraemia (aI_ )

- 273.0 a -4.91 a 0.548 a 42.5 a.b 585 a.b 0.291 a

- 106.6 b - 2.33 b 0.564 a 32.1 a 399 0.342 a

YLw, Yeast and low in nitrogen; CLw, control and low in nitrogen; YNl, yeast and normal in nitrogen; CNl, control and normal in nitrogen. Y, Yeast; N, nitrogen; P, parity. EB, Energy balance; PDIB, protein tNly digestible in the small intestine balance. NS, Not significant; * P < 0.05; * * P < 0.01; * * * P < 0.001. N XP and Y X N X P interactions are not significant. Means not bearing the same superscript letters within rows are significantly different (P < 0.05).

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3.1. Zootechnical parameters

For all weeks considered, only parity of the animals had a significant effect on metabolic weight (MW; liveweight0.75) which was 18.3 kgo.75 for primiparous goats and 22.4 kg o.75 for multiparous (Table 2). Thus intake, nutritive balances and yields are expressed on the basis of MW. Intake of DM in the four groups was equivalent. As the feeds were isoenergetic, net energy intake was also equivalent in the four groups. Obviously, the level of PDI intake was significantly linked to the N level of the diet. 3.2. Milk yield Milk yield did not differ among the four groups, although there was a trend for it to be higher in groups receiving yeast (Table 3). However, provision of yeast increased milk fat content (mean effect: +4.5 fatpointsg kg-), and had no effect on protein, lactose, ash, dry matter or urea concentrations in milk. This positive effect of yeast on milk fat content also occurred for milk fat yield, and consequently production of fat corrected milk (35 gfatkg-) was increased (Fig. 1). Group CLw tended to differ from the other groups in terms of protein and lactose yields, but only at the P < 0.10 level. The urea content of milk was significantly higher in groups receiving more nitrogen (YNl and CNl), although the extent of the increase was greater with yeast supplemented diets, whereas the nitrogen factor had no effect on milk protein levels. Dietary factors had relatively little effect on the fatty acid composition of milk lipids, expressed in per cent (Table 4). However, Group CLw differed from the others, with lower percentages of long-chain fatty acids (especially (C,s:,)), which result from mobilization, and a trend for a higher percentage of C ibEo, and of short- and medium-chain fatty acids (sum of C,--C ,4) synthesized in the mammary gland. Quantities of fatty acids (expressed as g day - kg- MW) tended to increase with the supply of yeast for all fatty

weeksafterparturition 0 1

-100

-200

-300

-400 500 Fig. 2. Relationship between energy balance and time post-parturn.

158

S. Giger-Reuerdin et al./Animal Feed Science Technology 63 (1996) 149-162

Fig. 3. Relationship

between plasma NEFA concentration

and time post-partum.

acids considered (Table 5). This increase was nearly always significant at the P < 0.05 threshold. 3.3. Nutritional balances and blood parameters Energy balances were consistently negative, which is normal when considering the physiological state of early lactating goats because of the mobilization of lipids. However, provision of yeast caused the energy balance to be more negative (Table 6 and Fig. 2). This decrease tended to be greater in animals receiving the low-nitrogen feed (Y X N interaction, P < 0.09). These effects can be explained by the positive effect of yeast on fat corrected milk production ( + 23%).

weeks after parturition 0 1

-2.5

-5 -

-7.5

-10

--

Yeast and Ncfmal ccolrol

nivogeo
1

g kg-l metabolic weight

and Normal

nllrOQen

Fig. 4.

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The among-group hierarchy of NEFA is a reflection of energy balance or milk production (Fig. 3). At equivalent energy balances, however, the NEFA content of Group YLw tended to be lower than that of Group YNl. The BHB levels were higher when animals received yeast and with the higher nitrogen intake. The among-group hierarchy for BHB levels corresponded to those for NEFA, as well as to lactose production. Glycaemia was not significantly affected by treatments. Groups that received yeast, however, tended to have lower glycaemia than the others. This trend was increased by increasing the nitrogen supply in the feed. The PDI balances were estimated from calculated PDI intakes and PDI requirements of the animals (Morand-Fehr and Sauvant, 1988). They were significantly more negative in groups receiving low-nitrogen diets. Yeast also had a negative effect on these balances (Fig. 4). Uraemia increased significantly with nitrogen supply, although the extent of the increase was greater within goats provided with yeast.

4. Discussion In the present study, yeast had no effect on DM intake. This agrees with results of Arambel and Kent (1990), Kellems et al. (1990) and Wohlt et al. (1991) at the onset of lactation, and those of Glinther (19891, Erdman and Sharma (19891, Harris and Webb (lQQO),Erasmus et al. (1992) and Smith et al. (1993) in mid-lactation. Only Williams et al. (1991) reported a positive effect of yeast on DM intake for cows in mid-lactation. In the present experiment, the animals were starting lactation, and were therefore in a period when they do not meet their nutritional needs, or, in other terms, in a period of physical regulation of appetite. Yeast apparently has an effect on rumen stoichiometry, as several workers have shown that its inclusion in the diet increases the total quantity of volatile fatty acids produced in vivo and in vitro, but not always significantly, as well as the individual quantities of C, , C,, C, and isoacids (expressed in mol l- of rumen fluid) (Arambel et al., 1987; Wiedmeier et al., 1987; Harrison et al., 1988; Gray and Ryan, 1989a, Gray and Ryan, 1989b; Martin et al., 1989; Dawson, 1990; Jouany et al., 1991; J.P. Jouany et al., unpublished data, 1993; Windschitl, 1992; Mutsvangwa et al., 1992). The possibility of increased quantities of C, and C, produced in the rumen, as previously observed with the same strain of yeast by Jouany et al. (1991, and unpublished data, 1993), could explain the increase in plasma BHB levels in groups receiving yeast and thus increased mammary synthesis of short- and medium-chain fatty acids, as acetic and butyric acids are precursors of plasma BHB. This increased synthesis also requires larger quantities of glucose and this is consistent with the numerically lower blood glucose levels (although they are non-significant) in yeast groups. With dairy calves, Quigley et al. (1992) also reported an increase in molar proportions of acetate and butyrate and in plasma ketone bodies (BHB and aceto-acetate) with the inclusion of yeast in the diet. Increased production of propionate in the rumen, as previously observed (Jouany et al., 19911, is consistent with the increase in the production of fatty acids containing 15 or 17 carbon atoms by the udder. Yeast clearly caused more extensive mobilization of reserves, and increased use of NEFA and

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endogenous BHB by the udder would result in higher levels of non-esterified fatty acids and greater production of long-chain fatty acids in milk. Yeast could provide a source of nutrients such as nucleotides, amino acids and vitamins for micro-organisms in the rumen which could promote bacterial growth through autolysis in the forestomach (Guerzoni and Succi, 1972). This phenomenon would be expected to be more significant in a nitrogen-deficient medium. It could explain differences in response to a supply of yeast between groups receiving the low-N diets and those receiving the feed in which N was not limiting. Through this potential increase in bacterial growth, ammonia N would be used better and its concentration in the rumen decreased (Erasmus et al., 1992; J.P. Jouany et al., unpublished data, 1993). This increased incorporation of ammonia into microbial protein could increase the duodenal flux of microbial N (Erasmus, 1991), and even modify the amino acid profile of microbial protein (Erasmus et al., 1992). As energy intake was the same for the four groups, the possible increased protein flow to the intestine may explain the enhanced overall performance of goats fed the low-N diet when yeast was added. Thus, in the case of yeast groups the PDI values of diets need to be reconsidered. The positive effect of yeast on the milk fat content, on milk fatty acid production and on fat corrected milk yield is directly related to the increased secretion of the different fatty acids by the udder. This agrees with the results of Giinther (19891, Harris and Webb (1990) and Williams et al. (1991) obtained in early lactation, and those of Kellems et al. (1990) during a whole lactation. However, Arambel and Kent (1990) and Wohlt et al. (1991) in early lactation, and Harris and Lobe (19881, Quinonez et al. (19881, Erdman and Sharma (19891, Huber et al. (1989), Harris et al. (19921, Piva et al. (1993) and Smith et al. (1993) in mid-lactation could show no positive effect of yeast addition to the diet. The lack of effect of yeast on protein and lactose contents confirms results of Arambel and Kent (19901, Kellems et al. (1990) and Wohlt et al. (19911, who also worked with early lactating ruminants, at a time when energy deficiency is an important factor in changes in milk protein content. These results are also in agreement with those of Piva et al. (1993), who worked in mid-lactation. Nevertheless, the increase in lactose production, only significant at the P < 0.10 level, is related to that of increased plasma BHB.

5. Conclusions

The main effect of yeast in early lactation was to induce the goats to mobilize more energy reserves, as seen by higher NEFA levels, leading to a higher production of fat corrected milk. It would be of interest to continue this study and provide the yeast on the basis of DM intake throughout the experiment (e.g. 1.5 X lo9 CFU kg- I DMI), to clarify whether responses are due to a flash effect of the yeast or to a dose effect. To substantiate our ruminal explanations of the results based on experiments done in males or in vitro, changes in aspects of rumen fermentation and in forestomach and total digestibility should also be studied with early lactating animals using different N levels, as the yeast effect depends on the nitrogen level of feed. This would lead to a better assessment of the value of yeast in ruminant feeding.

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Acknowledgements The authors thank J. Hervieu and N. Lemaire for taking care of the experimental goats and for technical assistance, F. Ternois for skilful laboratory analyses, and Dr. A.A. Ponter and Dr. C. Duvaux-Ponter for correcting the English.

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