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PRODUCTION, MODELING, AND EDUCATION |
* Department of Agricultural, Food and Nutritional Science, University of Alberta, Edmonton, Alberta Canada, T6H 5T6; and Agriculture Research Division, Alberta Agriculture and Food, Edmonton, Alberta, Canada, T6H 2P5
1 Corresponding author: frank.robinson{at}ualberta.ca
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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Key Words: broiler breeder • genetic strain • growth profile • fleshing • frame size
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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There are benefits to delaying photostimulation (PS) to 22 or 23 wk of age (Robinson et al., 1996; Renema et al., 2001). When pullets are reared on a conventional BW target, there is no advantage to photostimulating the flock before 22 wk of age (Robinson et al., 1996). Sexual maturation of low-BW pullets can lag behind heavier birds (Robinson and Robinson, 1991; Renema et al., 1999a). Katanbaf et al. (1989b) suggested that the attainment of sexual maturity depends more on age in ad libitum-fed broiler breeders and on BW and carcass composition in feed-restricted hens. A concern with delayed PS is that this may affect the length of the productive egg-laying period. This potential inefficiency is thought to be compensated for by increasing the rate of lay during the production period. It has been proposed that the attainment of BW, body composition, and hypothalamic maturity thresholds required for pubertal development can be accelerated by rearing the pullets on a heavier BW profile to allow a greater BW at PS.
Further research is needed to determine if relative increases in BW in the prelay period (16 to 24 wk) or absolute BW is more critical in triggering puberty and follicular recruitment. Precision feeding of broiler breeder pullets and hens is best attained when the breeder strain is well defined in terms of sexual maturation and nutrient allocation. In commercial broiler strains, continuous improvement in yield and efficiency traits makes characterizing the reproductive implications of genetic selection a challenge.
The current paper is the first component of a comprehensive project that examined the effects of strain, BW profile, and age at PS on pullet rearing and breeding characteristics. This paper is focused on the effects of strain, BW profile, and age at PS on pullet growth and development of selected metabolic and reproductive organs to 24 wk of age. This study will provide a better understanding of the physiological mechanisms driving the interaction among nutrient allocation, selection for growth rate, and age at PS on the development of pullets as they enter production.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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This project was a 3 x 4 x 2 factorial design to determine the effect of 3 broiler breeder strains, 4 target BW profiles, and 2 PS ages on pullet growth and development. Pullets were randomly preassigned at hatch to an experimental fate. Some birds were killed for collection of carcass data at specific ages from 0 to 24 wk of age. Other birds were individually caged and killed on the day following the first oviposition (Renema et al., 2007). The final group of birds was individually caged at 17 wk of age to characterize the egg production period to 58 wk of age (Zuidhof et al., 2007). The current paper deals with the first group of birds to 24 wk of age. The research project, which is reported here, was carried out in compliance with the Guide to the Care and Use of Experimental Animals (Canadian Council on Animal Care, 1984) and was approved by the University of Alberta’s Faculty of Agriculture, Forestry and Home Economics Animal Policy and Welfare Committee.
Stocks and Management
A total of 560 day-old Hubbard Hi-Y (Hubbard ISA, Walpole NH), Ross 508, and Ross 708 (Aviagen Inc., Huntsville, AL) pullets (total = 1,680) were hatched commercially and housed in 24 floor pens of 70 chicks per pen (10 birds/m2) in a light-controlled facility. All chicks were individually identified by neck tags (Swift tack, Heartland Animal Health, Fair Play, MO). The photoperiod was 24L:0D for the first 3 d and 8L:16D to 18 wk of age. Feeder space and floor space exceeded industry recommendations.
Four BW profiles were used, varying in BW from 0 to 32 wk of age as shown in Figure 1
. The profiles were designed to diverge at 4 wk of age and converge at 32 wk. The control profile (standard) was calculated as the mean target BW profile of the strains used. A low profile based on an early reduction in feed allocation was followed by a period of large feed increases during sexual maturation. Birds of this low profile were 25% lower in BW than the standard pullets at 12 wk of age. A moderate profile was based on birds being 150% of the BW of standard birds at 12 wk followed by lower rate of gain to 32 wk. A high profile was based on birds being 200% of 12-wk BW of the standard birds. Pullets in this profile had minimal BW gains during the period of sexual maturation. Birds were group-weighed twice per week from 0 to 18 wk of age to facilitate precise feed allocations for close adherence to the target BW curves. All birds were fed a common diet: starter from 0 to 5 wk, grower from 5 to 22 wk, and a breeder diet after 22 wk (Table 1). Birds were fed ad libitum for the first 4 wk of age, after which quantitative feed restriction was applied. Feed was provided daily on a pen basis based on BW, BW gain during the previous period, and predicted BW gain for the immediate week. The actual feed allocations for each BW profile are shown in Figure 2.
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Data Collection
From 0 to 18 wk, at 14-d intervals, 4 birds from each interaction (total of 48 birds per time) were killed before feeding for collection of carcass morphometric data. The number of treatments doubled following PS of half of the birds at 18 wk of age, and 96 birds (4 per interaction) were dissected at 20, 22, and 24 wk.
Birds were euthanatized by cervical dislocation and weighed. The length of the right shank (tibiotarsus measured from the hock to the footpad) and the keel (distance from the hypocleidoclavical joint to the caudal end of the sternum) were measured using a Vernier caliper (Griffin et al., 2005). The pectoralis major and pectoralis minor, abdominal fat pad, ovary, and oviduct, were dissected from the carcass and weighed. Total breast weight was calculated as the sum of the weights of the pectoralis major and minor. The weights of the breast muscle and abdominal fat pad were calculated as a percentage of BW.
Statistical Analysis
Data were analyzed using the MIXED procedure of SAS (SAS 9.1, SAS Institute Inc., Cary, NC). Strain, BW profile, and PS age were analyzed as fixed effects, and age was analyzed as a random effect. Because variance across age was heterogeneous, separate variances were estimated for each age. No covariance across age was estimated, because measurements were not repeated on individual birds. Differences between least squares means were determined using pairwise differences and were reported as significant at the P < 0.05 level. In some instances, in which convergence was not possible with the full model, a systematic process was engaged in which parameters were removed to achieve convergence and maximize the number of significant parameters in a reduced model.
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RESULTS AND DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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By design, mean BW was similar among strains throughout rearing (data not shown). There were occasional differences among the smaller groups of birds dissected at 2-wk intervals due to random selection of individuals for dissection at each period (Table 3). Close adherence to the target BW was attributed to biweekly BW determinations, weighing all birds, and precision in estimating growth in feed allocation calculations. The high and moderate BW profiles were consistently heavier than the standard and low birds after 6 wk (Table 3). The low treatment was more similar to the standard treatment than the other heavier treatments by design. The intent was to slow the early growth in the low group without substantially stunting their growth to feed them more aggressively than standard birds during the sexual maturation period. After 18 wk of age, PS had little effect on BW (Table 3). Within the strain x PS age and BW profile x PS interactions, there were some unexplainable significant differences, due to what appeared to be random effects of bird selection (Table 3). To reduce sampling bias, birds were assigned to a processing age at hatch allowing the full range of possible phenotypic expressions of growth potential to be expressed.
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. The low treatment was characterized by very low BW gains early (approximately 6 g per d from 6 to 8 wk of age), followed by larger BW gains later in rearing. The converse was true for the moderate and high treatment groups. The peak gain was seen in the high treatment group pullets (approximately 40 g per d) from 7 to 8 wk of age. The week-to-week variability in BW was greater after 18 wk of age. Due to changes in nutrient partitioning priorities at the time of sexual maturity, BW gains typically dropped dramatically around the time of first oviposition (data not shown). Some of the variation in BW gain after 18 wk is related to variation in sexual maturation status, because some birds commenced lay during this time.
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vs. Figure 4). The implications of feed provision (relative to maintenance) were more readily apparent from BW gain data than from BW data. In a commercial setting, breeder flocks with automatic hen scales could be programmed to calculate the gain on a daily or 2-d basis, providing timely data on which to base feed allocation decisions. The effects of feed allocation on the metabolic state of pullets are much more difficult to determine from BW data than from BW gain data.
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), with the Ross 708 chicks having longer shanks than the other 2 (lower breast yielding) strains. It was interesting to note that by wk 2 and 4, the Ross 708 chicks had shorter shanks than the other 2 strains. At 4 wk, Ross 708 shank length was 52.6 mm vs. 56.5 and 59.3 mm in Ross 508 and Hubbard Hi-Y, respectively. There were no differences in shank length among strains from 6 to 22 wk of age. At 22 and 24 wk of age, the differences in shank length among strains were small and were not consistent, suggesting that this may be due to factors specific to the birds randomly selected for dissection. Birds of all strains were limited to a common frame size for most of the rearing period once the feed restriction programs were imposed.
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). There were also differences between the high and moderate birds during some weeks. The standard profile was designed to be representative of the industry standard target BW. Results for this group indicate that current commercial feed restriction practices limit frame size. Shank length was not influenced by PS age overall (Table 5).
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) and again at 10 wk, although some period differences may be artifacts of different birds being represented each week. However, beginning at 8 wk and continuing throughout the rearing program, the feed restriction programs limited keel length in all strains. Body weight profiles affected keel length in a similar fashion to shank length; standard feed restriction practices limited both indicators of frame size. The feed allocations used to achieve the BW targets in this study were very diverse from 6 to 18 wk of age (Figure 2). Clearly, the range in feed allocation was enough to influence frame development already at a young age.
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). By 24 wk, birds photostimulated early (18WK) had longer keels than the 22WK birds (Table 5). This was true for all 3 strains and for the 3 heaviest BW profiles (Table 6). This observation was not consistent with previous research, which has shown that mature frame size increases when birds were photostimulated later due to delayed cessation of growth (Yuan et al., 1994; Robinson et al., 1996; Joseph et al., 2002). The relationship between hormones released at sexual maturity and growth of the keel requires further investigation. Breast Muscle Weight
Ross 708 birds had the greatest breast muscle development (expressed as a % of BW) early in rearing (Table 7). From 12 to 18 wk, the results were more often similar among strains. By 20 wk of age, the greater degree of fleshing of the Ross 708 birds was again statistically clear and remained this way to the end of the 24-wk study period.
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). It is interesting to note that PS did not influence breast muscle fleshing in terms of the main effect (Table 5 and Table 7), whereas there were some spurious differences in 1 or more BW profiles at 20, 22, and 24 wk (Table 7). This is the period in which breast muscle size in the 18WK birds may have been limited compared with the 22WK birds due to diversion of nutrients into reproductive development. These data indicate that at 18 wk of age, variability in percentage of breast muscle resulting from the diverse feed allocations needed to achieve the 4 BW profiles was greater than the genetic variability among the 3 lines. However, at 22 wk, there was more variability among strains than there was between BW profiles.
Abdominal Fat Pad Weight
At wk 2 and 4, during the period of ad libitum consumption, Hubbard HI-Y pullets had the greatest relative fat pad weights (Table 8). At wk 16, Hubbard Hi-Y and Ross 508 birds had greater fat pad weights than the Ross 708 pullets. Presumably this observation was related to the increased propensity for breast muscle deposition in the Ross 708 birds, rather than diverting excess nutrients to storage as fat. The Ross 708 birds consistently had the lowest abdominal fat levels from 20 wk of age. The efficiency and yield traits this strain had been selected for appear to have limited fat deposition. Consequently, the lower fat stores in Ross 708 compared with the other strains may be linked to their inability to maintain early egg production traits under conditions of under-nutrition (Zuidhof et al., 2007). As heavier hens come into production earlier and with greater lipid content (Summers and Leeson, 1983), age at sexual maturity has been considered to relate to BW and composition within the context of hormonal balance (Leeson et al., 1988). With early PS, the result can be a nonuniform flock, with variation in the timing of their prime sequence and corresponding nutrient requirements. With a later PS age, the flock will begin egg production in a more uniform manner (Robinson et al., 1996).
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). After wk 16, the high birds had heavier fat pads than the standard birds (data not shown). The low treatment pullets had similar fat pad weights to the standard birds from 2 to 18 wk of age. It appeared that in most weeks, neither of these 2 treatments deposited abdominal fat reserves; the abdominal fat pad weight recorded was mostly connective tissue that supported fat pad development in more generously fed birds. There was a 4-fold increase in relative fat pad weight in the low birds from 18 to 24 wk of age. This change was only 2-fold in standard birds and nonexistent in high birds (Table 8). These changes demonstrate the effect of the large increase in feed allocation provided to the low birds during this period. It also shows that decreasing feed allocations in this period (high birds) did not result in a decrease in abdominal fat.
Photostimulation at 18 or 22 wk of age had no significant effect on absolute or relative fat pad weight (Table 5). However, in the Ross 708 strain and on the standard, moderate, and high BW profiles, there were instances in which PS age influenced fat pad weight (Table 8). In most of these cases, earlier PS resulted in smaller fat pads, presumably due to a redirection of nutrients from carcass lipid deposition to the production of yolk precursors in the liver and ultimately in the production of yolky ovarian follicles. Ovarian estrogen, produced in response to PS, will increase very low density lipoprotein production that is ultimately represented in recruited follicles (Griffin et al., 1982; Walzem, 1996).
Oviduct Weight
The oviduct weights for the 0 to 16-wk period are not shown, because the values were very small, with several small inconsistent, though statistically significant, differences between strains and BW profiles. At 18 wk of age, absolute oviduct weight was not different among strains, but it was heavier in high BW profile birds compared with the other 3 lower BW profiles (Table 9). This was similar to the previous findings of the largest birds in a random population of pullets also having the greatest oviduct and ovary weights (Renema et al., 1999b).
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and Table 9). These differences persisted to 24 wk. In 18WK birds, there were no strain effects on oviduct weight from 20 to 24 wk of age. However, unphotostimulated Ross 708 birds had smaller oviducts than the other 2 strains at 20 wk of age. Although age at sexual maturity for companion birds was not different among strains, there was a numerical difference, with the Ross 708 birds commencing lay 4 d later than the Hubbard Hi-Y birds and 2 d later than the Ross 508 birds (Renema et al., 2007). The BW profile data show clearly that oviduct weights are higher in birds that are on heavier growth curves throughout sexual maturation.
From 22 to 24 wk of age, the oviduct weight of 18WK-low birds had increased in size 4-fold but was still approximately half the weight of oviducts from birds on the 3 heavier BW profiles. The age-related change in oviduct weight in the heavier bird profiles suggested that the 18WK-low birds were several weeks behind the high birds and less than a week behind the 18WK-standard birds. Sexual maturation of the 18WK-low birds was ultimately delayed by 17 and 5 d, respectively, compared with the high and standard birds (Renema et al., 2007). Oviduct weights were similar across all 4 BW profiles in the 22WK birds. The potential BW effect in the moderate and high birds was likely tempered by these groups being allocated less feed than the smaller standard and low birds in the period immediately following PS (Figure 2). Oviduct weight can be very responsive to feed allocation. During sexual maturation, the growth and development of the oviduct is typically seen before growth of the ovary (Melnychuk et al., 1997), making it a good variable to determine the short-term response to PS during this period.
Ovary Weight
By 24 wk of age, strain influenced ovary weight, indicating that initiation of the sexual maturation process differed among strains (Table 10). In the 18WK birds, the Ross 708 and Ross 508 pullets had a smaller ovary than the Hubbard Hi-Y birds. Ovary weight was influenced by BW profile and possibly level of fatness in a manner similar to oviduct weight. Also in 18WK birds, moderate and high birds demonstrated earlier ovary growth than standard and low birds (Table 10). Varying feed intake before, during, and immediately after sexual maturation can result in a difference of 1 extra large yellow follicle, with a concomitant 10-egg reduction (Robinson et al., 1998a,b). Even small degrees of over- or under-feeding are thought to negatively affect egg and chick production (Katanbaf et al., 1989a,b).
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). Overall, age at PS had the greatest effect of any treatment on ovary development. Current commercial BW profiles limit frame size, breast muscle fleshing, abdominal fat pad weight, and the onset of the pubertal growth of the reproductive organs (liver, ovary, and oviduct; Renema et al., 1999a,b). Analyses of BW profiles need to consider the relative allocation of nutrients early in life, because they affect the establishment of carcass frame and fleshing (maintenance) and later in rearing during the development of the reproductive system. There are significant strain differences in response to aggressive early feeding and to subsequent feed restriction. Some strain differences are obscured by feed restriction. The predisposition of high-yielding pullets to deposit breast muscle may influence the concomitant deposition of carcass fat, other organs and tissues, or both. Photostimulation at 18 wk of age alters the nutrient-partitioning priorities and hence carcass and organ morphology. Organs in which growth is stimulated in response to early PS include the liver, ovary, and oviduct.
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ACKNOWLEDGMENTS |
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Received for publication January 24, 2007. Accepted for publication June 15, 2007.
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REFERENCES |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONREFERENCES |
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