Effects of increasing amounts of
Lupinus albus
seeds without or with whole egg powder in
the diet of growing pigs on performance
C. Van Nevel
a,c,*, M. Seynaeve
b, G. Van De Voorde
c,
S. De Smet
c, E. Van Driessche
a, R. De Wilde
baLaboratory of Protein Chemistry, Institute for Molecular Biology and Biotechnology, Vrije Universiteit Brussel, Paardenstraat, 65, B-1640 Sint-Genesius-Rode, Belgium bLaboratory of Animal Nutrition, Faculty of Veterinary Medicine, University of Ghent,
Heidestraat, 19, B-9820 Merelbeke, Belgium
cDepartment of Animal Production, Faculty of Agricultural and Applied Biological Sciences, University of Ghent, Proefhoevestraat, 10, B-9090 Melle, Belgium
Received 4 November 1998; received in revised form 30 March 1999; accepted 19 October 1999
Abstract
In a growth trial with pigs, the effect of the inclusion of 150 and 300 g ofLupinus albusseeds/kg of feed in the diet was investigated. Parameters studied were: growth, feed utilization, digestibility of nutrients, slaughter and carcass characteristics. Spray dried whole egg powder, a speci®c inhibitor of lectins inL. albusseeds was also added (50 g/kg of feed), with the aim of verifying whether the unfavourable effects of high levels of lupin seeds could be neutralized. Feeding the diet containing 300 g of lupin seeds/kg lowered the average daily gain from 727 to 674 g and feed intake from 2.32 to 2.05 kg, while feed conversion ratio remained unaltered. The presence of whole egg powder in the lupin seed diets did not abolish the negative effects. Apparent faecal digestibility of most nutrients in the diets was not in¯uenced by addition of lupin seeds or egg powder, except for the crude fat fraction, whereas the digestibility coef®cient increased from 0.51 to 0.61. Crude ®bre digestibility also increased, but only at the lowest lupin seed level. Carcass weight and dressing percentage were lower in the groups fed the highest lupin seed level. Fatty acid pro®le of backfat was determined and slightly higher proportions of C18:1 were observed when lupin seeds were fed. Possible reasons accounting for the lower performance of animals receiving lupin seeds are discussed, but the exact reason could not be derived from this experiment.#2000 Elsevier Science B.V. All rights reserved.
Keywords:Pig growth;L. albusseeds; Antinutritional compounds; Egg powder 83 (2000) 89±101
*Corresponding author. Tel.:32-9-2649014; fax:32-9-2649099.
1. Introduction
In the search for alternative protein sources replacing the more traditional ingredients, such as soyabean meal, meat meal or ®sh meal in pig diets, much attention has been paid during the last decade to the use of lupin seeds, e.g.Lupinus angustifolius,L. albusand
L. luteus. Their potential use was made possible by genetic selection for cultivars with low levels of alkaloids, which were thought to be the major antinutritional compounds in lupin seeds (Petterson et al., 1986). Data concerning lectin content are very scarce.The lack of detection of lectins in lupin seeds in earlier works was probably due to the fact that, for the detection of lectins, the hemagglutination method was applied using non-protease treated erythrocytes (Hill, 1977), and the test may not have been adequate. Consequently, during the genetic selection process no attention was paid to the aspect of lectin content. However, in earlier works using rabbit erythrocytes pretreated with Pronase, hemagglutination activities of 41±82 units/mg of seed meal ofL. albusand 5±20 units/mg of seed meal of L. angustifolius (Van Nevel et al., 1998) were determined. Lectins inL. angustifoliusseeds have also been found by Kim and Madhusudhan (1988) and Rahman et al. (1997) using trypsinized red blood cells. A brief literature survey showed that growth and feed intake of pigs decreased when the diet contained 150±430 g of L. albus seeds/kg (Pearson and Carr, 1977; Batterham, 1979; Roth-Maier and Kirchgessner, 1994; Zettl et al., 1995). Although it was generally thought that alkaloids were the inhibitory compounds, it was put forward that other, yet unknown factors could have been responsible for the negative effects on animal performance (Pearson and Carr, 1977). It is not inconceivable that the presence of lectins also played a role in the unfavourable growth response when lupin seeds were fed to piglets or pigs. On the other hand, in previous in vitro works studying the use of receptor-analogues, saturating the sugar-binding site of lectins and thus inhibiting their activity in extracts of different legume seeds, it was observed that whole egg powder was a very ef®cient inhibitor of lectin activity inL. albus seeds (Van Nevel et al., 1998). Therefore, it was decided to investigate in vivo, whether addition of whole egg powder (50 g/kg) was able to neutralize the negative effect of lupin seed containing diets (0, 150 or 300 g/kg) on performance of growing pigs. The following measurements were undertaken: daily gain, feed intake, feed conversion ratio, apparent faecal digestibility, slaughter data and carcass quality including fatty acid pro®le of backfat.
2. Materials and methods
2.1. Animals
2.2. Diets
The experiment was divided into two periods and, for each period, diets with different crude protein content were fed: 186 g/kg during a ®rst period (growth phase from 24 to 43 kg LW) and 170 g/kg during a second growth phase (43±102 kg LW). Seven diets were formulated. Two diets acted as control without lupin seeds: a ®rst diet (G) without egg powder, and a second diet (GE) containing 50 g of egg powder per kg of feed. Four diets with lupin seeds were formulated containing, respectively, 150 and 300 g of lupin seeds/kg of feed without (L15, respectively, L30) or with addition of 50 g of egg powder/ kg (respectively, L15E and L30E). A seventh diet (L30Ly) with extra supplementation of lysine was included, because Batterham et al. (1986) found that when lupin seed meal was fed, the ef®ciency of utilization of absorbed lysine was only 0.53. In this manner, it could be demonstrated whether eventual negative effects of a diet containing 300 g of lupin seeds/kg was due to lysine being limiting for growth. The composition of the different diets is presented in Tables 1 and 2. Within each growth phase, diets were balanced for total protein content, net energy value (9.43 and 9.18 MJ/kg for growth phases 1 and 2, respectively ), Ca, P and some essential amino acids (based on small intestinal digestibility). All ingredients were ground (mesh, 2.0 mm) and diets fed in the dry form. Whole egg powder was provided by VEOS nv., Zwevezele, Belgium (VEPRO
Table 1
Composition (g/kg) of the diets (growth phase 24±43 kg) Ingredients Diets
1G 1GE 1L15 1L15E 1L30 1L30E 1L30Ly Manioc 237.5 274.1 191.0 225.0 144.6 177.5 142.9 Wheat 400.0 400.0 400.0 400.0 400.0 400.0 400.0 Soyabean meala 316.3 245.0 210.0 138.7 100.0 300.0 100.0 Lupin seeds 0.0 0.0 150.0 150.0 300.0 300.0 300.0 Egg powder 0.0 50.0 0.0 50.0 0.0 50.0 0.0 Soyabean oil 14.0 0.0 15.0 1.5 16.5 2.5 16.5 Vitamins and mineralsb 10.0 10.0 10.0 10.0 10.0 10.0 10.0 Limestone 6.0 6.5 6.0 6.5 6.0 6.5 6.0 Dicalcium phosphate 11.0 9.5 11.0 9.5 11.0 9.5 11.0
Salt 3.0 3.0 3.0 3.0 3.0 3.0 3.0
L-Lysine 0.6 0.8 1.2 11.5 2.2 2.3 3.9 DL-Methionine 0.6 0.1 0.8 0.3 1.1 0.6 1.1 L-Threonine 0.0 0.0 0.0 0.0 0.4 0.4 0.4 L-Tryptophan 0.0 0.0 0.0 0.0 0.2 0.2 0.2
Stafoc 1.0 1.0 1.0 1.0 1.0 1.0 1.0
KHCO3 0.0 0.0 1.0 3.0 4.0 6.5 4.0 aSolvent-extracted meal, containing 412 g of crude protein/kg.
bProviding per kg of feed: vitamins: A, 12 000 IU; D
3, 2000 IU; E, 50 mg; K, 1 mg; B1, 0.6 mg; B2, 4 mg;
B3, 12.5 mg; B6, 1 mg; B12, 0.02 mg; niacin, 20 mg; folic acid, 1.5 mg; biotin, 0.10 mg; choline, 350 mg;
Minerals: iron, 150 mg; copper, 25 mg; manganese, 50 mg; cobalt, 1 mg; zinc, 100 mg; iodine, 2 mg; selenium, 0.4 mg; calcium, 870 mg; sodium, 0.2 mg; magnesium, 136 mg; Ethoxyquin, 1 mg; BHT, 1 mg.
55EP). Lupin seeds (L. albus, cv. Lublanc) were imported from France (Groupe CANA, 44150 Ancenis).
2.3. Digestibility measurements
Faecal digestibility of nutrients was determined with three pigs per diet (barrows), using 15 g of Celite 545 (VEL, Leuven, Belgium) as marker per kg of feed. Collection of faecal material was undertaken during 5 days, after an adaptation period of at least 1 week during which the Celite containing diet was fed. Faecal material was stored at ÿ228C until further processing. At the time of digestibility measurements, pigs weighed 70±80 kg, except for diet 2L30 where LW was67 kg.
2.4. Slaughter and carcass characteristics
At an LW of 100±105 kg, animals were transported to the experimental slaughterhouse of the Department of Animal Production, University of Ghent at Melle (Belgium). The feed withdrawal period before slaughter was20 h. The transportation time was 15 min and the pigs did not receive sedatives. The following determinations were carried out: dressing percentage, backfat thickness (measured at 3 locations: at the 1st, 7th and last rib) and percentage of lean in carcass. The latter parameter was calculated by the following regression model with the weight percentages of backfat (X1), loin (X2) and leg (X3) as parameters, determined by a wholesale cut: percentage of lean 7.326ÿ1.334X11.511X20.794X3; R20.966, RSD1.35 (Van De Voorde, to be published). Backfat was sampled for determination of fatty acid pattern.
Table 2
Composition (g/kg) of the diets (growth phase 43±102 kg)a
Ingredients Diets
2G 2GE 2L15 2L15E 2L30 2L30E 2L30Ly Manioc 300.0 300.0 300.0 300.0 300.0 300.0 300.0 Wheat 387.1 343.0 338.5 296.0 287.0 251.0 285.3 Soyabean meal 270.0 185.0 177.5 93.5 85.0 0.0 85.0 Lupin seeds 0.0 0.0 150.0 150.0 300.0 300.0 300.0 Egg powder 0.0 50.0 0.0 50.0 0.0 50.0 0.0 Wheat bran 20.0 100.0 10.0 85.0 0.0 70.0 0.0 Vitamins and minerals 10.0 10.0 10.0 10.0 10.0 10.0 10.0 Limestone 3.0 4.5 3.0 4.5 3.0 4.5 3.0 Dicalcium phosphate 6.0 3.5 6.0 3.5 6.0 3.5 6.0
Salt 3.0 3.0 3.0 3.0 3.0 3.0 3.0
L-Lysine 0.2 0.5 1.0 1.0 1.1 1.5 2.8
DL-Methionine 0.2 0.0 0.5 0.0 0.7 0.2 0.7
L-Threonine 0.0 0.0 0.0 0.0 0.1 0.2 0.1
L-Tryptophan 0.0 0.0 0.0 0.0 0.1 0.1 0.1
Stafo 0.5 0.5 0.5 0.5 0.5 0.5 0.5 KHCO3 0.0 0.0 0.0 3.0 3.5 5.5 3.5
2.5. Chemical analysis
Approximate composition (Weende analysis) on feed and faecal material (homo-genized and lyophilized) was undertaken following standard European Union procedures. Celite was determined as 4 N HCl insoluble ash. Hemagglutinating activity (units/mg of feed) as a measure of lectin presence in extracts of L. albus seed meal and diets was determined using protease (Pronase, Boehringer, Mannheim, Germany) treated rabbit erythrocytes as described previously (Van Nevel et al., 1998). Trypsin-inhibiting activity was determined by Masterlab (Trouw Nutrition Nederland bv., Putten, The Netherlands) using benzoyl-l-arginine-p-nitro-anilidide as substrate (Nederlands Normalisatie Insti-tuut, 1993). Total and individual alkaloids in L. albus seeds were analyzed by Dr. M. Muzquiz using a capillary GLC-MS procedure (Muzquiz et al., 1994). Fatty acid pattern in diets and lupin seeds was determined as described by Sukhija and Palmquist (1988), but C17:0 (heptadecanoic acid) was used as an internal standard. As seeds ofL. albus
have a rather high lipid content (80±140 g/kg), containing 500±550 g/kg of oleic acid (C18:1), 170±200 g/kg of linoleic acid (C18:2) and 90±100 g/kg of a-linolenic acid (C18:3), fatty acid pattern in samples of backfat was also determined (Hill, 1977; Green and Oram, 1983; Zettl et al., 1995). Lipids in backfat samples were extracted as described by Folch et al. (1957). Transesteri®cation and extraction of fatty acid methyl esters was done as outlined previously (Demeyer et al., 1978). C17:0 was used as internal standard and individual fatty acids were determined by GC as reported by Van Nevel and Demeyer (1996).
2.6. Statistics
The effect of lupin seeds and egg powder was tested in a 23 factorial model using the data of diet one-to-six (General Linear Models procedure of SPSS, 1997). The least squares means are given in the tables. Linear and quadratic polynomials were evaluated for the effect of increasing lupin content of the diets. The effect of lysine supplementation was separately tested by comparison of diet 7 (L30Ly) with diet 5 (L30) using a simplet -test.
3. Results
3.1. Diet composition
lupin seeds were not incorporated. The reason for this was an unexpected very high HA in extracted soyabean meal. In a further experiment, HA in extracts of soyabean meal, wheat and manioc was measured using untreated erythrocytes and all results were negative. During a last trial using Pronase-treated erythrocytes, HA was again determined in
Table 3
Chemical composition (Weende analysis) of diets, lupin seeds and whole egg powder Diets Nutrients (g/kg)a
DM OM CP CL CF NFE
1G 875 810 172 35 61 542
1GE 878 813 180 34 54 545
1L15 879 818 199 48 72 499
1L15E 880 818 187 45 69 517
1L30 878 822 186 59 90 487
1L30E 878 821 194 59 79 489
1L30Ly 878 818 192 56 84 486
2G 865 805 171 22 61 551
2GE 869 809 170 34 59 546
2L15 867 807 169 30 72 536
2L15E 870 808 173 46 69 520
2L30 865 804 166 39 87 512
2L30E 867 807 167 53 83 504
2L30Ly 865 804 172 40 87 505
Lupin seeds 870 838 299 93 165 281
Egg powder 965 899 566 272 21 40
aDM, OM, CP, CL, CF, NFE, respectively, dry matter, organic matter, crude protein, crude fat, crude ®bre,
nitrogen-free extractive.
Table 4
Trypsin inhibiting activity (TIA) and hemagglutination activity (HA) of diets and some ingredients
Dieta TIAb,c HAd
1G 0 41
1GE 2.0 20±41
1L15 0.09 20±41
1L15E 2.09 20±41
1L30 0.18 20±41
1L30E 2.18 10±41
1L30Ly 0.18 20±41
Lupin seeds 0.59 20±41
Egg powder 40.0 nd
Soyabean meal n.d.e 1024±2048
aGrowth phase 1 only.
bTrypsin inhibited, in mg/g of feed.
cCalculated from TIA in lupin seeds and egg powder, assuming that other ingredients contained no trypsin
inhibitors.
extracts of the same ingredients and again high values were observed for soyabean meal (1024 to 2048 units/mg), very low values for wheat (2±4 units) and no HA in manioc extracts. It thus seems that use of Pronase treated erythrocytes increased the sensitivity of the HA method to a very considerable extent.
3.2. Animal performance
3.2.1. General
Statistical treatment indicated that for all parameters studied, no signi®cant differences could be observed between the group receiving the diet containing 300 g of lupin seeds (L30) and the group fed the same diet but with extra supplementation of lysine (L30Ly). Therefore, data concerning the latter group were not mentioned in the following tables. Furthermore, statistical analysis also showed that between both the treatments, namely addition of egg powder and lupin seed inclusion, no signi®cant interaction could be calculated, except for one parameter, i.e. faecal digestibility of crude fat (Table 6). Consequently, results for both treatments could be discussed separately in the text, while the aspect of crude fat digestibility was brie¯y treated in Section 3.2.3. For statistical treatment of the data concerning daily gain and feed intake during Growth phase 2 and the entire trial, ®nal body weight was included as a covariate. The same procedure was undertaken for carcass weight (Table 7).
3.2.2. Growth, feed intake and feeding ef®ciency
Addition of egg powder to the diet had no effect on the parameters investigated (Table 5). A trend to a lower average daily gain was observed for the groups fed 300 g of lupin seeds/kg of feed. Feed conversion ratio was not in¯uenced.
3.2.3. Digestibility of nutrients
Digestibility of the crude fat fraction was higher in diets containing egg powder and lupin seeds (Table 6). Statistical treatment revealed interaction between both treatments due to the fact that digestibility coef®cients increased with increasing amounts of lupin seed in the diet, but only in the presence of egg powder. Compared to the control group, crude ®bre digestibility tended to be higher for lupin seed-containing diets. Digestibility of the other nutrients was not altered by both the treatments.
3.2.4. Slaughtering performance and carcass characteristics
Table 5
Reponse of growing pigs to diets containingL.albusseeds fed without or with egg powder
Parametera Treatment SEMc
egg powder(g/kg) lupin seeds (g/kg)
0(18)b 50(18) 0(12) 150(12) 300(12) Phase 1
LW1 (kg) 23.9 23.7 23.9 23.8 23.5 0.37 LW2 (kg) 43.1 42.8 43.5 42.5 42.9 0.29
ADG (g)d 534 552 598 528 504 15
Feed intake (kg/d)e 1.31 1.38 1.47 1.35 1.21 0.03
FCR 2.49 2.53 2.54 2.57 2.42 0.04
Phase 2
LW1 43.1 42.8 43.5 42.5 42.9 0.29
LW2 102.1 101.8 102.8 102.6 100.4 0.53
ADG 767 791 792 800 744 13
Feed intakef 2.62 2.66 2.71 2.78 2.44 0.05
FCR 3.45 3.38 3.44 3.48 3.34 0.08
Total trial
ADGf 689 715 727 706 674 10
Feed intakef 2.20 2.25 2.32 230 2.05 0.04
FCR 3.21 3.17 3.20 3.27 3.10 0.07
aLW1, LW2, respectively, initial body weight and weight at end of growth phase; ADG. average daily gain;
FCR, feed conversion ratio (kg of feed/kg of gain).
bNumber of observations in parentheses.
cStandard of the mean, calculated from pooled values (n36). dTrend for linear effect of lupin seeds (p< 0.1).
eLinear effect of lupin seeds (p< 0.01). fLinear effect of lupin seeds (p< 0.05).
Table 6
Apparent faecal digestibility coef®cients of nutrients
Nutrient Treatment SEMb
egg powder(g/kg) lupin seeds (g/kg)
0(9)a 50(9) 0(6) 150(6) 300(6)
Dry matter 0.79 0.81 0.80 0.81 0.79 0.005 Organic matter 0.84 0.85 0.84 0.85 0.83 0.005 Crude protein 0.73 0.80 0.77 0.80 0.78 0.010 Crude fatc 0.46 0.67 0.51 0.57 0.61 0.011 Crude ®bred 0.50 0.51 0.44 0.57 0.51 0.016 NFEe 0.93 0.92 0.92 0.93 0.93 0.003
aNumber of observations in parentheses.
bStandard error of the mean, calculated from pooled values (n18).
cLinear and quadratic effect of lupin seeds (p< 0.01). An interaction effect between the treatments egg
powder and lupin seeds was noted.
Table 7
Slaughter and carcass characteristics
Parameter Treatment SEMf
egg powder(g/kg) lupin seeds (g/kg)
0(18)a 50(18) 0(12) 150(12) 300(12)
LW (kg)b 101.1 101.1 102.3 101.3 99.6 0.5 Carcass weight (kg)c 84.9 84.9 86.1 85.1 83.5 0.3 Dressing %d 84.0 84.0 85.0 84.1 82.8 0.3 % Lean 60.2 60.3 59.1 60.2 61.6 0.9 Backfat thicknesse(mm) 80.7 77.8 83.7 83.3 70.7 2.9
aNumber of observations between parentheses. bBody weight at slaughter.
cLinear effect of lupin seeds (p< 0.01). dLinear effect of lupin seeds (p< 0.05).
eSum of measurements at three locations: 1st, 7th and last rib. fStandard error of the mean, calculated from pooled values (n36).
Table 8
Fatty acid pro®le (g/kg) of lipid in diets, egg powder, lupin seeds and backfat Diet Fatty acid
C16:0 C16:1 C18:0 C18:1 C18:2 C18:3 C20:0 C20:1
2G 223 0 67 331 308 26 0 0
2GE 211 12 57 297 310 25 0 0
2L15 195 17 44 314 319 38 0 0
2L15E 197 20 57 346 245 30 0 13
2L30 172 26 27 369 289 42 0 0
2L30E 137 21 32 455 223 52 0 18
2L30Ly 191 0 53 371 312 46 0 0
55EPa 265 28 98 463 125 6 0 0
L. albusseeds 92 10 20 550 140 67 11 41
Backfat
Fatty acid Treatment
egg powder (g/kg) lupin seeds (g/kg)
0(18)b 50(18) 0(12) 150(12) 300(12) SEMc
C16:0 250 248 252 254 242 2
C16:1 7 13 11 11 8 2
C18:0 157 149 158 154 146 3 C18:1d 518 522 505 517 538 5
C18:2 68 68 75 64 66 3
aEgg powder (from Cotterill et al., 1978). bNumber of observations in parentheses.
4. Discussion
4.1. Diet composition and digestibility of nutrients
In agreement with literature data (Hill, 1977), TIA ofL. albusseeds was very low, as opposed to whole egg powder. Ovomucoid (100 g/kg egg protein) is one of the most important trypsin-inhibiting substances (Kato and Matsuda, 1997). Hartmann (1995) also found very high concentrations of trypsin and cathepsin D inhibitors in egg white and yolk. The present experiment showed that TIA in the diets fed was too low to have caused growth inhibition of the animals. The fact that in extracts of solvent extracted soyabean meal considerable HA as a measure of lectin activity was found, was due to the use of protease-treated erythrocytes making the method much more sensitive. Very recently, small but substantial amounts of carbohydrate-binding lectin could be determined in several samples of conventionally processed soyabean meal (Maenz et al., 1999). It is certainly not known to what extent these residual lectins can affect performance of pigs or other animals. However, this phenomenon cannot account for the lower performance of the groups fed lupin seed diets in this experiment, as the proportion of solvent-extracted soyabean meal in these diets was much lower than in the control diet (Tables 1 and 2).
Apparent faecal digestibility coef®cients of the different nutrients were within the normal range observed with similar diets (Zettl et al., 1995; Flis et al., 1996). Higher digestibility of crude fat in diets containing egg powder are related to somewhat higher lipid content of these rations (Table 3, Growth phase 2) and this fraction must have been highly digestible. Digestibility of the crude-®bre fraction was higher in the present experiment, but in the literature this value was found to be rather variable and also dependent on the other ingredients in the ration. Additionally, Aguilera et al. (1985) determined a digestibility coef®cient of 0.83 for crude ®bre inL. albus, cv. Multolupa.
4.2. Growth, feed intake and feeding ef®ciency
Daily gain and feed conversion ratios observed were comparable with results of other studies feeding similar diets (Pearson and Carr, 1977; Batterham, 1979; Zettl et al., 1995). Supplementing the L30 diet with lysine (L30Ly) had no effect on performance, which agrees with Batterham et al. (1986), who investigated the effect of mixing seeds of L. angustifolius in pig diets. The possibility of the presence of factors in the seeds also decreasing utilization of the lysine supplement, a phenomenon thought to be speci®c to pigs, was also considered. In experiments with rats receiving L. angustifolius seeds, Rahman et al. (1997) concluded that a low utilization of absorbed N was caused by an increased excretion in urine, indicating disturbances in the systemic metabolism. N-utilization was also negatively in¯uenced by the presence of important amounts of non-starch polysaccharides (NSP).
showed that feeding diets containing >200 g of lupin seeds/kg of feed caused a lowered daily gain and feed intake, especially during a ®rst growth phase, although differences were not always statistically signi®cant (Pearson and Carr, 1977; Batterham, 1979; Zettl et al., 1995). Feed conversion ratio was not in¯uenced. Several mechanisms explaining these observations have been proposed. Seeds ofL. albuscan contain very high amounts of Mn (467±3750 ppm) and it has been suggested that such high concentrations can possibly result in depressed growth in pigs (National Research Council, 1998). On the other hand, the presence of quinolizidine alkaloids decrease palatability of the diet causing lower feed intake. Analysis of alkaloids in the lupin seeds fed in the present experiment showed a total amount of 0.0123 mg/100 mg of seeds with the following pattern: 0.0011 mg of angustifoline, 0.0090 mg of lupanine and 0.0022 mg of 13-OH lupanine. Taking into account the available literature data, it can be concluded that alkaloids in the current diets were not responsible for the lowered feed intake or growth performance because their concentration was very low (Godfrey et al., 1985; Buraczewska et al., 1993; Wasilewko et al., 1997). It was also clearly established by Pearson and Carr (1977) that an eventual high amount of manganese in lupin seeds was not responsible for unfavourable effects; the authors suggested that besides an eventual high content of alkaloids, other factors were probably responsible for lower feed intake of lupin seed-containing diets. As it has been shown that lysine in lupin seeds and also lysine supplemented to diets containing lupin seed was only utilized with an abnormally low ef®ciency, it cannot be completely excluded that poorer performances of the animals observed in the current experiment were due to an inadequate provision of this essential amino acid (Batterham et al., 1986).
Addition of whole egg powder (50 g/kg) to the different diets had no effect on growth performance and feed intake, which was completely different from the effect of egg powder in piglet and chicks diets, where growth inhibition has been observed (Peo et al., 1969; Watkins and Veum, 1986; Kratzer et al., 1988). It also ruled out the possibility of biotin de®ciency caused by the presence of avidin in the egg powder in the current trial (Kratzer et al., 1988).
4.3. Carcass characteristics and fatty acid pro®le of backfat
The lower carcass weight and dressing percentage of animals receiving the highest amount of lupin seeds is probably due to a larger gut ®ll caused by the higher ®bre content of this diet (Batterham, 1979), while the tendency towards more lean and less fat in their carcass is probably the result of slower growth rates related to lower fat deposit.
5. Conclusions
It can be stated with certainty that the presence of lectins did not play a role in the problems observed when lupin seeds were fed, as their concentration was very low, and the fact that adding whole egg powder had no effect on parameters studied supports this statement. On the other hand, the exact reasons for the negative effects on feeding lupin seeds remain unknown and could not be determined from this experiment.
Acknowledgements
The research was ®nancially supported by the Ministry of Agriculture, DG6, Brussels and VEOS nv. Zwevezele, Belgium. Technical help from S. Galle, E. Maes, Daisy Bayens, H. De Rycke, N. Vanrolleghem and Jenny Martens is gratefully acknowledged.
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