Feeding high lupine based diets for broiler chickens: Effect of soybean meal substitution with yellow lupine meal at various time points of growth cycle

Accepted Manuscript

Feeding high lupine based diets for broiler chickens: Effect of soybean meal substitution with yellow lupine meal at various time points of growth cycle Boguslaw Olkowski PII: DOI: Reference:

S1871-1413(18)30593-6 https://doi.org/10.1016/j.livsci.2018.10.017 LIVSCI 3561

To appear in:

Livestock Science

Received date: Revised date: Accepted date:

18 December 2017 22 September 2018 29 October 2018

Please cite this article as: Boguslaw Olkowski , Feeding high lupine based diets for broiler chickens: Effect of soybean meal substitution with yellow lupine meal at various time points of growth cycle, Livestock Science (2018), doi: https://doi.org/10.1016/j.livsci.2018.10.017

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ACCEPTED MANUSCRIPT Highlights 

Lupines based diets in chickens during the first 3 weeks of growth are not recommended

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Lupines meal can be used as soybean substitute for broilers aged 4 weeks and onwards Chicken fed lupines may have nutritional quality beneficial for consumers

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ACCEPTED MANUSCRIPT Feeding high lupine based diets for broiler chickens: Effect of soybean meal substitution with yellow lupine meal at various time points of growth cycle

Boguslaw Olkowski*

Institute of Bioengineering and Animal Breeding,

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08-110 Siedlce, 14 B. Prusa, Poland

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Siedlce University of Natural Science and Humanities, Faculty of Life sciences,

*Corresponding author: [email protected]

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Bogusław Olkowski

Siedlce University of Natural Science and Humanities, Faculty of Life sciences, Institute of Bioengineering and Animal Breeding, 08-110 Siedlce, 14 B. Prusa, Poland.

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ACCEPTED MANUSCRIPT Abstract

The present work examined the potential benefits and limitations of yellow lupine seeds meal (LM) as a substitute for soybean meal (SBM) in broiler diets. The experimental work was focused on evaluation of effects of dietary lupine introduced in the diet at various time points The nutritional efficacy of LM for broiler chickens was evaluated using

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of the growth.

performance parameters including body mass gain, feed conversion ratio, and carcass analysis. Breast muscle was subjected to color and chemical analyzes. Patho-physiological variables monitored in response to LM included overt livability, morbidity, mortality, and

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morphometry of key organs involved in digestive processes.

Complete replacement of SBM with LM in the broiler diet introduced at days 1, 8, and 15 resulted in significant decline of feed intake and growth rate (both P<0.05). High level of LM in the diet for young chickens caused enlargement of the liver, pancreas, gizzard, and

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intestines (all P<0.05). Chickens fed a LM based diet did not show gross changes in carcass quality or yield of main commercially important parts (breast muscle and leg quarters), but Also, LM slightly

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showed increased (P<0.05) carcass proportion of giblets and fat pad.

decreased breast muscle protein content and increased muscle fat content (P<0.05), but it is

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noteworthy that fat from LM fed broilers had more desirable profile of fatty acids. Although nutritional characteristic of LM appeared to be comparable to SBM, its

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nutritional potential cannot be completely utilized by young chickens. Complete substitution of SBM with LM in diet for broilers during first stages of growth is not realistic, because of it

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would considerably lower the nutritional efficiency of the ration and would have detrimental effects on performance.

In conclusion, from a practical stand point, LM based diet for

broilers during the first 3 weeks of growth is not recommended, but the LM based diet during later stages of growth (4 weeks and onwards) can be used in the broiler ration without risks of compromising production parameters or health.

Key words: Lupine seeds, broiler chicken, soybean replacement, age 3

ACCEPTED MANUSCRIPT 1. Introduction

Lupine seeds contain more proteins than other grain legumes, and therefore LM is more suitable as a dietary protein supplement in poultry. The high cost of feeds for organic poultry production generates increased interest in organically-grown lupines as an alternative feed

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ingredient. Among the main cultivated lupine species (L. albus, L. angustifolius, L. luteus), seeds of L. luteus have highest content of protein and better amino acids profile, making it a preferred feed for poultry (Petterson, 2000).

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Historically, a long-standing issue limiting the use of lupines in animal feeding had been a high level of toxic alkaloids in the so-called bitter varieties, but with the development of “sweet” lupine varieties containing insignificant level of alkaloids (Reinhard et al., 2006), the problem of toxicity has been practically eliminated (Petterson, 2000).

Also the levels other

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significant anti-nutritional factors, typical for grain legume such as phytates, tannins, lectins, saponis and tripsin inhibitors in modern lupine cultivars are negligible (Petterson, 2000).

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However, relatively high levels of non-starch polysaccharides (NSP) still presents a major factor limiting large scale use of lupine seeds in young chickens’ diets (Brenes et al., 2003;

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Hughes et al., 1999; Steenfeldt et al., 2003, Petterson, 2000). Of note, yellow lupins contain considerable lower level of NSP in comparison to other lupine species (Kluge et al., 2003).

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Given the nutritional characteristics, yellow lupines have great potential in the feeding of broilers, especially as alternative to soy bean meal protein concentrate, (Petterson, 2000). In

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addition, lupines, being very efficient in nitrogen binding, are environmentally friendly, and as such are ideally suited for ecological poultry rearing systems (Hammershoj and Steenfeldt, 2005).

Although it is generally accepted that lupine seeds can be used in broiler diets, practical aspect of lupine based ration formulation is somewhat controversial. Some authors (Olver, 1987; Perez-Alba et al., 1990b; Olver and Jonker, 1997; Froidmont et al., 2004; Laudadio 4

ACCEPTED MANUSCRIPT and Tufarell, 2011a) suggest that lupine based diets formulated to totally replace SBM had no negative effects on health or performances. In contrast data from other studies indicate that in fact such high levels of lupine in diet may be detrimental to growth and health in young birds (Naveed et al., 1998; Hughes, 1999; Olkowski et al., 2001, 2005; Steenfeldt et al., 2003; Janiuk et al., 2014) because of anti-nutritional impact. However, it is well established

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that poor performance of broilers fed lupine based diets is not associated with inadequacy of riboflavin (Olkowski 2008) and the detrimental effects can be reduced by enzyme addition (Olkowski et al., 2001; Olkowski, 2011).

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In view of the inconsistencies among published reports, more work is needed to examine chickens responses to high levels of dietary lupine at various growth stages. Accordingly, the present study was focused on responses of broilers to diets formulated to replace SBM with lupine meal, where the lupine based diets introduced at various stages of

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2. Material and Methods

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growth cycle.

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2.1. Birds and feeding regime: A total of 180, Ross 308 day old broilers (mixed sex) were randomly assigned to 5 treatment groups, 36 birds per group. Each group was replicated 6

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times, with 6 chickens per replication. Feeding regime assigned to respective treatment groups was according to the schedule presented in Table 1. The chickens were reared in

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metal battery cages in environmentally controlled room. The feed and water were offered ad libitum. The birds were daily monitored for overt clinical abnormalities. The experiment was conducted in two phases, consisting of a starter phase from day 1 to 21 and a finisher phase from day 22 to 42. The experiment was conducted in compliance with the European Union (2010/63/EU) directives on the care and use of animals for experimental and other scientific purposes.

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ACCEPTED MANUSCRIPT 2.2. Diets: Two basal diets were formulated using either soybean meal (SBM) or lupine meal LM) as main protein supplement. Otherwise, the basal diet contained all required nutrients to fulfil nutritional requirements of broiler chickens (NRC 1994). Maize gluten was used to level balance of protein in diets, whereas the level of metabolic energy was balanced by soya oil. All diets were supplemented with adequate synthetic amino acids (lysine, methionine,

diets (starter and grower) are presented in Table 2.

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tryptophan, and threonine) and a mineral-vitamin mix. Ingredients and nutritional value of

2.3. Data collection: Birds live weights were recorded on day 1, and then on days 7, 14, 21, 42. The feed consumption data were collected accordingly. Performance assessment data

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included feed intake (FI), body weight gain (BWG), and feed conversion ratio (FCR). At the completion of the experiment, all birds were weighted, and routinely processed for post slaughter analysis.

Selected gastrointestinal tract (GI) components were subjected to

morphometric measurements. Morphometric analyses were performed on organs from 12

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chickens per each group, with masses being recorded for liver, pancreas and empty gizzard. Length were recorded for duodenum, jejunum, ileum, and caeca. Carcasses were subjected

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to abbreviated post-slaughter analysis where weight of primary commercial cuts (breasts, legs), edible giblets (gizzard, liver, heart) and abdominal fat data were recorded.

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Representative samples of breast fillets were randomly selected from each group (six

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specimens per one group) for further analysis.

2.4. Chemical analyses

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Samples of experimental feeds and breast muscles were subjected to analyses for proximate nutrient composition according to methods described in the Association of Official Analytical Chemists (AOAC, 2001).

The samples of breast muscle were evaluated for fatty acid

composition in lipid fraction using capillary gas chromatography according to the procedure described by Zapletal et al. (2010). The following parameters were calculated on the basis of the fatty acid content results: saturated fatty acid (SFA), unsaturated fatty acid (UFA), monounsaturated

fatty

acid

(MUFA),

polyunsaturated 6

fatty

acid

(PUFA),

and

ACCEPTED MANUSCRIPT hypercholesterolemic fatty acid (OFA). Breast fillet meat samples were also subjected to color analyses. Color measurement was performed on muscle surface free from unnatural discoloration using a Minolta CR-310 chroma meter; calibration was done on a white standard.

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2.5. Statistical analysis Statistical analyses were carried out by GLM ANOVA from the computer package NCSS (Hintze 1995). The means were compared using Tukey’s test. Statistical significance was

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assumed to exist when the probability was less than 0.05.

3. Results

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In comparison with control (SBM group) and other LM groups, chickens fed lupine diet from day 1 (group LM1) showed the lowest feed intake, weight gain, and feed conversion ratio, The detrimental effects of LM on

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both during the starter and grower period (Table 3).

performance parameters were also apparent, although of lesser magnitude, when SBM diet

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was replaced with LM diet from day 8 (group LM8), from day 15 (group LM15), and from day 22 (group LM22).

However, it is noteworthy that the performance indexes significantly

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improved (P<0.05) as the LM diet was introduced during various stages of growth, with general trend being for group LM1 < group LM8 < group LM15 < group LM22. There were

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no significant differences in FC and BWG between group LM and SBM group. Detailed data on performance parameters for different treatment groups are shown in table 3.

Overall mortality was very low, one chicken died in the SBM group in the 1 st week, one chicken in LM1 and one in LM22 died in the 2 nd week. Post mortem examination revealed no signs of toxicity or pathological lesions. The effect of lupine on hypertrophy of GI organs was noticeable, but the differences were not always statistically significant (Table 5). The LM diet 7

ACCEPTED MANUSCRIPT increased liver mass (P<0.05) and overall increased the length of the small intestine, but not caeca and colon. Introducing LM in diet on day 8 (group LM8) has similar effect, but of lesser magnitude. The LM diet introduced on day 22 (group LM22) has a significant effect (P<0.05) on the length of duodenum, while the remaining GI parameters did not differ in

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comparison with the SBM group.

The replacement of SBM diet with LM diet in the starter period did have any significant effect on slaughter efficiency or share of main commercial elements of carcasses (Table 5). The LM introduced on day 1 (group LM1), on day 8, and on day 15 significantly increased

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(P<0.05) abdominal fat pad.

Substitution of SBM with LM affected meat chemistry (Table 6), with notable (P<0.05) reduction of protein content and increase in intramuscular fat content. Breast muscle's color

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analysis did not show any major changes (Table 6). Brightness (L*) and redness (a*) did not differ markedly between groups, but there was a slight but significant (P<0.05) effect of LM

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on stronger yellowish hue (b*) of breast muscles in group LM1.

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The LM based diet affected composition of lipid fraction (Table 7). Lipid fractions from chicken fed LM diet showed lower levels of palmitic acids (C16:0) and stearic acids (C18:0)

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in comparison to those fed SBM diet (P<0.05). In contrast, the level of linolenic acid (C18:3) tended to be higher in chickens fed LM diet, and this effect was significant (P<0.05) when LM

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diet was introduced in 3rd or 4th week of chicken rearing. Overall, LM based diets changed the content of total saturated fatty acids (SFA) and total unsaturated fatty acids (UFA) in lipid fractions. In contrast to SBM fed chickens, lipids in chickens fed LM diet had markedly lower total SFA (P<0.05), while total UFA level was significantly higher (P<0.05). Generally, the contents of monounsaturated fatty acids (MUFA) did not differ between groups, but the polyunsaturated fatty acids (PUFA) in lipid fraction of LM fed chickens were higher (P<0.01) in relation to lipids of SBM fed chickens. 8

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4. Discussion Based on the analysis of the various stages of SBM replacement with a LM in the diet, it can be inferred that LM substitution in the diet at any point during the first 3 weeks of rearing would have detrimental effects on performance parameters, but It is noteworthy that, neither

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pathological lesions nor signs of lupine toxicity described by Olkowski et al. (2001) were observed in the present study, which suggest that further improvement in quality of lupine seeds was achieved since the 1990’s.

The observed effect of a high level of LM diet introduced at the beginning of growth

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(group LM1) on performance and morphology of the GI concur with the results published previously (Farrell et al., 1999; Brenes et al., 2002; Olkowski et al., 2005). However, it is worth noting that in the present study the magnitude of adverse responses declined when the LM diet was introduced at later stages of the growth period, and the difference between

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control (SBM group) and LM fed group became statistically insignificant after day 21 of age. In the present study, production indices for the entire growth period in group fed LM

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diet introduced on day 22 were comparable to those seen in SBM group, indicating that complete substitution of SBM in broiler grower rations introduced during the 4 th week is a

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realistic approach without any risk of adverse effects on performance. Similar conclusions were derived by other authors (Roth-Maier and Kirchgessner, 1993, 1994; Roth-Maier and

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Paulics, 2003; Suchy et al., 2006). The findings from the present study indicate that the diet containing very high levels

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(400g/kg) of LM introduced from first week had negative effect on feed intake of young broilers and performance. In essence, observations from the present study are consistent with reports of other researchers (Olkowski et al., 2001, 2005, Strakova et al., 2010, Olkowski 2011), and therefore generally, research findings from a wide range of sources indicate that substitution of SBM with high levels of LM in rations for young broilers should be avoided. In order to overcome the problems resulting from complete substitution, some authors 9

ACCEPTED MANUSCRIPT investigated the effects of partial replacement of SBM with lupine. Nalle et al. (2011) reported positive results in terms of weight gain and feed intake with broilers at an inclusion rate of 200 g/kg of white lupine. In contrast, Steenfeldt et al. (2003) reported that incorporation of blue lupine at 200 g/kg in broiler diets did not reduce feed intake, but this did not abrogate negative effects on weight gain and feed conversion efficiency. The latter study indicate that

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depressed feed intake is not the only factor associated with detrimental effects of dietary lupine on performance of broilers.

Interestingly, when the LM diet is supplemented with suitable enzyme preparations, performance parameters are not different from those obtained with SBM (Olkowski 2011).

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Other authors (Steenfeldt et al., 2003 and Brenes et al., 2005) also showed that the enzyme addition can improve the nutritional value of lupine cultivars that were found to depress chick performance. In some trials, only a slight improvement of performance in chickens fed diets with high levels lupine was reported due to enzyme supplementation (Rubio et al., 2003),

2006).

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while at lower levels of lupine inclusion the enzyme effects were insignificant (Orda et al., Although studies with enzymes revealed that some anti-nutritional factors are

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associated with negative effects of lupine on performance, more comprehensive research is required to fully understand detrimental effects of lupine seeds.

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Interestingly, substituting SBM with LM did not affect slaughter efficiency and yield of main culinary elements of carcasses (breasts and legs); however significantly increased the

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share of edible giblets and fat deposits. Results pertaining to slaughter efficiency observed in the present study are in agreement with the findings of Suhy et al. (2006) and Diaz et al.

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(2006). Notably, LM induced beneficial changes in the fatty acid composition of lipid fraction with pro-health attributes of broilers meat. Similar effect on fatty acid profile was reported in studies with white lupine (Strakova et al., 2010). Also, the effects of white lupine in diet for boiler chicken causing decline in protein content and increase in fat content in muscles seen in the present study are similar to the findings reported by Froidmont et al. (2004). In the assessment of breast muscle color there was a significant impact on yellowness. Similar effect of yellowish tinge was apparent on skin of carcasses from chicken 10

ACCEPTED MANUSCRIPT fed LM diets. The effect of LM on meat and skin color corresponded with considerable amount of carotenoids (Wang et al., 2008) and riboflavin (Olkowski, 2008) in lupine seeds. Interestingly, feeding lupine seed meal to hens resulted in a more distinct yellow-red color of egg yolk (Dvorak et al., 2006; Laudadio and Tufarelli, 2011b; Lee et al., 2016). It seems that natural pigments from LM diets was highly bioavailable in chicken, therefore lupines can be

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considered as suitable source of natural pigments in animal product in the context of positive impact for consumers.

5. Conclusion remarks

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Although yellow lupine only slightly affected slaughter efficiency and yield of breast and legs in carcasses, the application of high levels of yellow lupine seed in starter rations is not practical due to its negative effect on performance. Nevertheless, LM can be used as an alternative protein supplement for SBM in grower rations, but the benefits of LM in broilers

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diet must be weighed against negative effects on carcass characteristics which include higher abdominal fad pad and edible giblets, and fat composition, as well as slight decrease

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Acknowledgements

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in the content of protein in breast fillets.

This work was supported by the Ministry of Science and Higher Education (Statutory Theme No.

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46/91/S).

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Conflict of interest: No conflict of interest to declare.

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ACCEPTED MANUSCRIPT Wang, S., Errington, S., Yap, H. H., 2008. Studies on Carotenoids from Lupin Seeds. Lupins for Health and Wealth. Palta, J. A. & Berger, J. B. (eds.). Fremantle, Western Australia ed. Canterbury, New Zealand: International Lupin Association. Vol. 1, pp. 198 –202.

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Tierzucht. 53, 589–599.

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ACCEPTED MANUSCRIPT Table 1. Experimental design

Feeding period with given diet (days: from - to) SBM based diet

LM based diet

SBM

1 – 42

-

LM1

-

1 – 42

LM8

1-7

LM15

1 - 14

LM22

1 – 21

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Treatment

8 – 42

15 – 42 22 – 42

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LM1, LM8, LM15, LM22 - day of introduction LM diet instead SBM diet; SBM = soybean

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meal, LM= lupine meal

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ACCEPTED MANUSCRIPT Table 2. Composition and nutritional value of diets used according to treatment. SBM

Component

Grower

Starter

315 -

400

-

-

40

30

70

72

82

88

40

45

45

45

L-Lysine (g/kg)

1.4

1.4

4.0

1.7

DL-Methionine (g/kg)

2.3

2.3

3.5

2.3

L-Tryptophan (g/kg)

-

-

1

0.6

L-Threonine (g/kg)

0.1

0.2

1

0.3

up to 1000

up to 1000

up to 1000

up to 1000

Soybean meal (g/kg) Lupine meal (g/kg)

365 Starter -

Maize gluten (g/kg) Soybean oil (g/kg) Mineral-vitamin mixture

1ab

(g/kg)

Maize meal (g/kg)

SBM

2

LZ

Grower 350

CR IP T

Starter

LM

Metabolizable Energy(AMEN ; MJ/kg) and nutrient composition (g/kg) 13.0

13.4

13.1

13.4

Dry matter*

919.4

923.2

930.4

922.0

Crude protein*

225.9

201.0

226.5

202.0

Crude fiber*

41.3

37.8

71.1

65.6

Calcium

9.7

9.4

10.8

10.5

Available Phosphorus

5.0

5,0

5,1

5.1

Linoleic acid

25.5

27.3

26.5

28.5

12.1

10.5

12.0

10.2

8,5

8.1

9.0

8.7

7.8

6.8

7.6

6.8

2.3

2.0

2.2

2.0

M

AN US

AMEN

Lysine

ED

Methionine + Cystine Threonine Tryptophan

- mineral-vitamin mixture (75% dicalcium phosphate, salt 7%, 6% limestone and 12%

PT

1

premix for broilers (starter or grower). The minerals and vitamins were supplemented diets at

CE

a rate: a - Starter (mg) Fe 80.0; Zn 70.0; Mn 80.0; Cu 15.0; I 0.12; Se 0.275; Co 0.3; vitamins

AC

(mg): E 80.0; K3 3.0; B1 2.5; B2 8.75; B6 5.0; B12 0.024; Biotyne 0.2; Folic acid 1.5; Calcium pantothenate 27.0; Nicotinic acid 50.0; Choline 500.0 (IU): A 20000; D3 2500. b-Grower (mg): Fe 75.0; Zn 65.0; Mn 70.0; Cu 7.5; I 0.125; Se 0.275; Co 0.25; vitamins (mg): E 40.0; K3 2.5; B1 2.0; B2 7.75; B6 4.0; B12 0.024; Biotyne 0.15; Folic acid 1.0; Calcium pantothenate 13.6; Nicotinic acid 45.0; Choline 500.0 or (IU): vit. A 15000; D3 2500. 2

- AMEN Calculations were based on chemical analysis of feedstuffs using formula and

digestion coefficients according to WPSA (Jansen, 1989). *- Analyzed 19

ACCEPTED MANUSCRIPT Table 3. Feed intake (FI), body weight gain (BWG) and feed conversion ratio (FCR) of broilers according to treatment. Parameter

SBM

LM1 c

LM8 a

LM15 a

LM22 b

bc

SEM

FI (g) BWG (g)

4225.1 2236.1b

3590.6 1859.2a

3645.3 1944.8a

3834.8 1983.9a

3991.4 2182.6b

96.1 72.2

FCR (g/g)

1.89

1.93

1.88

1.91

1.82

0.09

LM1, LM8, LM15, LM22 - day of introduction LM diet instead SBM diet; SEM - standard error

AC

CE

PT

ED

M

AN US

CR IP T

of the mean. Means in rows with different letters differ significantly P<0.05.

20

ACCEPTED MANUSCRIPT Table 4. Morphometric parameters of gastrointestinal tract (in relation to body weight) of breast meat according to treatment. Parameter

units

SBM

LM1

LM8

LM15

LM22

SEM

Gizzard Pancreas

g g

1.83 0.18

2.04 0.20

2.11 0.19

2.02 0.19

1.82 0.17

0.11 0.12

Liver

g

1.95a

2.21b

2.31b

2.00a

1.89a

0.12

1.16

a

1.28

b

1.31

b

ab

1.26

b

0.07

a

2.56

b

2.78

b

2.43

a

0.15

cm

Jejunum

cm

2.27

Ileum

cm

2.70

Ceca Colon

cm

1.42a

cm

a

0.31

2.92

2.50

1.63

b

0.39

b

1.47 0.36

a

ab

1.22

2.60

b

CR IP T

Duodeum

2.81

1.52

0.36

a

ab

2.86

0.20

1.53

a

0.08

0.30

a

0.03

LM1, LM8, LM15, LM22 - day of introduction LM diet instead SBM diet; SEM - standard error

AC

CE

PT

ED

M

AN US

of the mean. Means in rows with different letters differ significantly P<0.05.

21

ACCEPTED MANUSCRIPT Table 5. Dressing efficiency and participation of primary carcass elements(%) according to treatment. Parameter

SBM

LM1

LM8

LM15

LM22

SEM

Dressing efficiency Breast

74.35 22.94

72.26 21.41

72.16 21.93

72.78 22.12

73.37 21.52

1.50 0.80

Legs

26.54

25.57

25.40

25.53

25.69

0.80

3.89

a

4.68

c

4.41

bc

bc

ab

0.08

2.54

a

3.37

c

3.17

bc

b

0.06

Edible giblets

3.02

b

4.18

3.07

CR IP T

Abdominal fat

4.37

LM1, LM8, LM15, LM22 - day of introduction LM diet instead SBM diet; SEM - standard error

AC

CE

PT

ED

M

AN US

of the mean. Means in rows with different letters differ significantly P<0.05.

22

ACCEPTED MANUSCRIPT Table 6. Proximate chemical composition and color parameters of breast muscles according to treatment. Component

SBM

LM1

LM8

LM15

LM22

SEM

Dry matter (%) Crude ash (%)

26.53 1.20

25.92 1.20

25.90 1.19

26.65 1.18

26.40 1.17

0.32 0.01

Crude protein (%)

24.36b

23.32a

23.59a

23.60a

23.81a

0.29

0.77

Lightnes (L*)

46.22

48.82

48.06

Rednes (a*)

4.77

5.41

4.60

3.82

a

5.1 6

b

1.51

b

Crude fat (%)

Yellownes (b*)

1.59

bc

4.48

ab

1.73

c

47.59

b

0.12

47.71

0.84

1.22

CR IP T

a

5.12

4.30

a

4.74

0.43

a

0.24

4.03

LM1, LM8, LM15, LM22 - day of introduction LM diet instead SBM diet; SEM - standard error

AC

CE

PT

ED

M

AN US

of the mean. Means in rows with different letters differ significantly P<0.05.

23

ACCEPTED MANUSCRIPT Table 7. Fatty acids (FA) profile in lipid fraction (% of total FA) according to treatment. Components

SBM

LM1 b

LM8 a

LM15 a

LM22 a

SEM

20.64 6.18b

19.41 4.60a

18.79 4.29a

19.34 4.19a

19.03a 5.07a

0.47 0.25

C16:1

1.42a

1.57a

1.20a

1.56ab

1.77b

0.15

C18:1

32.51

31.64

32.27

32.22

31.99

0.61

C18:2

36.99a

40.61c

41.32c

39.93bc

38.33abc

0.97

C18:3 C 20:4

0.98

a

0.62

c

1.15 0.43

a

ab

1.09

a

0.49

b

CR IP T

C16:0 C18:0

1.33

b

0.35

a

1.37

b

0.11

c

0.07

0.57

Total SFA

27.08b

24.26a

22.41a

23.88a

24.37a

0.61

Total MUFA

34.04

33.32

33.56

33.95

33.88

0.61

Total UFA

38.71 72.75

a

42.07

b

75.39

b

c

43.89

42.07

b

AN US

Total PUFA

a

c

77.45

76.02

bc

41.59

b

1.07

75.47

b

0.55

LM1, LM8, LM15,LM22 = days of introduction LM diet instead SBM diet; SEM - standard error of the mean. Means in rows with different letters differ significantly P<0.05.

M

SFA = saturated fatty acids; UFA - unsaturated fatty acids; MUFA - monounsaturated fatty

AC

CE

PT

ED

acids; PUFA - polyunsaturated fatty acids.

24