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Department of Dairy and Animal Science, The Pennsylvania State University, University Park 16802
2 Corresponding author: ajh{at}psu.edu
| ABSTRACT |
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Key Words: heifer forage to concentrate ratio growth milk production
| INTRODUCTION |
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Reducing the length of the growing period by decreasing age at first calving below recommendations (22 to 24 mo; Ettema and Santos, 2004) could shorten the time between expenditure and revenue generation and reduce the costs associated with the nonproductive period. This could be accomplished by increasing prepubertal ADG, which would subsequently result in a lower age at first breeding and presumably a lower age at first calving. Although this strategy would ultimately lead to an earlier return on investment, increased prepubertal ADG has been demonstrated to have a negative impact on mammary parenchyma cell numbers (Sejrsen et al., 1982; Meyer et al., 2006) and first-lactation milk yield (Van Amburgh et al., 1998; Lammers et al., 1999; Radcliff et al., 2000). Determining a strategy to accelerate the growth of dairy heifers without subsequently reducing production should continue to be investigated. Even if a methodology to accelerate growth without reducing lactation performance could be implemented, the comparatively low efficiency of heifer rearing remains, albeit for a shorter duration of time.
It has been known for some time that diets containing high proportions of concentrate feedstuffs are utilized with greater efficiency than those containing high proportions of forages (Blaxter and Wainman, 1964; Garrett, 1979). Several experiments have offered dairy heifers high-concentrate (HC) diets for ad libitum consumption, which has led to reduced first-lactation milk production (Swanson, 1960; Radcliff et al., 2000). However, other experiments have demonstrated that if HC diets are restricted so that ADG is comparable to a high-forage (HF) control diet, mammary development and first-lactation milk production do not differ between groups (Hof and Lenaers, 1984; Sejrsen and Foldager, 1992; Carson et al., 2000). Despite these results, dairy heifers commonly receive diets in which the majority of nutrients come from forages instead of concentrates. Therefore, the objective of this experiment was to evaluate growth and first-lactation milk production in dairy heifers fed HF or HC rations for similar levels of prepubertal ADG.
| MATERIALS AND METHODS |
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The treatment rations were formulated on a quantity basis to meet or exceed the NRC (2001) nutrient recommendations for a 150-kg Holstein heifer gaining 0.800 kg/d and for all macronutrients to be equal between the rations (Table 1
). The HF control diet was formulated to contain 75% forage and 25% concentrate, and the HC diet was formulated to contain 25% forage and 75% concentrate. Heifers were individually fed a TMR twice daily at 12-h intervals (0800 and 2000 h). Heifers were fed these rations for 245 d in an attempt to encompass and isolate the prepubertal allometric mammary growth phase (Sinha and Tucker, 1969). An important consideration in this experiment was to maintain similar ADG throughout the prepubertal phase so that effects of differing levels of forage and concentrate could be determined independently from alterations in prepubertal ADG. Although the energy density of the 2 rations differed considerably, ADG was controlled by controlling the amount of each ration offered. Heifers were weighed on 2 consecutive days, 2 h prior to the morning feeding (0600 h), at weekly intervals; from the mean of these weights, the amount of feed offered in the subsequent week was adjusted to maintain the desired ADG. Concurrent with weekly weighing, several structural measurements also were recorded, including withers height, body length (point of the shoulders to the ischium), heart girth, paunch girth, and hip width (Brody, 1945). At the conclusion of the experimental period, heifers were transitioned onto a common HF diet (66% grass silage, 20% corn silage) that was group-fed until animals entered the precalving transition group between 30 and 60 d prior to calving. After the experimental feeding period was complete, heifers from each treatment were managed as a single group in a manner consistent with other heifers on the farm.
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Heifers were considered eligible for first breeding at 350 kg and 13 mo. In an effort to obtain a first calving age within the recommended range of 22 to 24 mo, heifers not inseminated prior to 13.5 mo of age were subjected to an estrus synchronization program including controlled-release intravaginal progesterone (Eazi-Breed CIDR, InterAg, Hamilton, New Zealand) and an injection of prostaglandin (Lutalyse, Pharmacia & Upjohn, Kalamazoo, MI). This protocol also was used for services subsequent to an open pregnancy diagnosis. The number of services required per pregnancy was recorded for comparative purposes, and heifers requiring >4 services were eliminated from the experiment (n = 2/treatment).
Lactation
Cows calved in individual pens, were milked, and were then transferred to a tie-stall barn (stall dimensions 132 x 183 cm) where the animals were transitioned to the lactation ration and monitored for postparturient problems. After approximately 1 mo, cows were transferred to a free-stall, sand-bedded barn with other first-lactation cows. The lactation ration was formulated as a TMR designed to meet nutrient requirements based on level of milk production according to the NRC (2001). Feeding occurred once per day at approximately 0700 h, and cows were allowed ad libitum access to feed and water. Cows were administered bST (Posilac, Monsanto Co., St. Louis, MO) biweekly beginning at 63 DIM.
Cows were milked twice daily at 0500 and 1700 h in a double-10 herringbone milking parlor equipped with the Afifarm system (S.A.E. Afikim, Kibbutz Afikim, Israel; US distributor: Germania Dairy Automation, Waunakee, WI). Milk yield was recorded for individual animals at each milking, and cows were weighed upon exiting the parlor. Monthly, at 2 consecutive milkings, milk samples were collected and analyzed for fat and protein (Dairy One DHIA, Ithaca, NY). Mature equivalent 305-d production values were calculated by DHIA procedures and are reported based on the first 154 DIM.
Calculations and Statistics
Because heifers were individually fed and intake and ADG for each animal were known, the individual heifer was considered the experimental unit. The previous experiment treatment blocking variable was retained in all cases, except where noted below, as a penalty against complete randomization. As mentioned previously, heifers were grouped in pens with other animals receiving the same treatment. For this reason, a fixed covariate effect was included for pen nested within treatment to account for potential pen-to-pen effects.
Any response for which there was one observation per animal (age at puberty, BW at calving, etc.) was analyzed according to a randomized complete block design including previous treatment and pen within treatment as covariates in PROC MIXED (SAS Institute, 2006). Repeated measurements during lactation were analyzed as in the above model, expanded to include the fixed effect of week and the treatment by week interaction. Intercept and week within cow within pen within treatment were included as random effects as variance components, and the correlation between residuals attributable to repeated measures was modeled using the first-order autoregressive covariance structure.
For weight and structural measurement gain, the following linear covariate mixed effects model was fit using PROC MIXED of SAS:
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where Yijkh is a continuous, dependent response variable; Bo is the overall intercept across treatment, previous treatment, pen grouping, and heifer; BoTi is a fixed effect of the ith treatment on the intercept (i = 1, 2); BoPj is a fixed effect of the jth previous treatment on the intercept (j = 1, . . . 5); BoGk(i) is a fixed effect of the kth pen grouping within the ith treatment on the intercept (k = 1, 2, 3); boh[k(i)] is a random effect of the hth heifer within the kth pen within the ith treatment on the intercept (h = 20); B1 is the overall slope (daily gain) across treatment, previous treatment, pen grouping, and heifer; B1Ti is a fixed effect of the ith treatment on the slope (i = 1, 2); B1Pj is a fixed effect of the jth previous treatment on the slope (j = 1, . . . 5); B1Gk(i) is a fixed effect of the kth pen grouping within the ith treatment on the slope (k = 1, 2, 3); b1h[k(i)] is a random effect of the hth heifer within the kth pen within the ith treatment on the slope (h = 20); and t is the continuous effect of time on trial in days after 35 d of adaptation; with
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and eijkh is the residual error.
Error correlation within heifers across days on trial was modeled using the first-order autoregressive covariance structure because of being repeated measures on the same animal across time. This procedure makes use of all measurements taken, does not rely solely on the initial and final measurements, and allows each covariate (treatment, previous treatment, and pen and the random effect of heifer) to be estimated simultaneously. The model was restrained to linear effects, although quadratic or greater effects were significant for some measurements. It must be noted, however, that in the regression equation for BW, the quadratic, nonlinear component was rejected (P > 0.40), indicating that during the time frame of this experiment, heifers grew at an approximately linear rate.
To overcome difficulties associated with the nonlinearity of the structural measurements, allometric equations were used relating the structural response of interest to BW. To assess effects that dietary treatments had on growth of structural characteristics relative to growth of the whole body, an allometric, nonlinear model was fit in the NLMIXED procedure of SAS:
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where Yih is the continuous, dependent response variable; B is a scaling coefficient of the equation across treatment and heifer; BTi is a fixed effect of the ith treatment on the scaling coefficient (i = 1, 2); bh(i) is a random effect of the hth heifer within the ith treatment on the scaling coefficient (h = 20); K is the overall allometric growth coefficient across treatment and heifer; KTi is a fixed effect of the ith treatment on the allometric growth coefficient (i = 1, 2); and kh(i) is a random effect of the hth heifer within the ith treatment on the allometric growth coefficient (h = 20); with
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and eih is the residual error,
N (0,
).
This is the typical expression for the allometric growth equation (Huxley, 1932) extended to account for variation arising from the fixed effect of treatment and the random effect of heifer in a manner similar to that used for the Gompertz growth function by Wang and Zuidhof (2004). This analysis is valuable in determining effects that treatment has on the growth of measured structural parameters relative to growth of the whole body and allows treatment effects and random individual heifer effects to be estimated simultaneously. Being nonlinear, this analysis is also preferred to quadratic or higher order polynomial equations in accounting for nonlinearity of growth curves, which suffer from high levels of multicollinearity between the parameter estimates. The covariate parameters included in previous models were originally included in this analysis as well; however, they were highly nonsignificant (P > 0.90) and made it difficult fitting the complex nonlinear mixed model and were thus excluded from this analysis.
The assumption of normality was assessed using the normal probability plot correlation coefficient test (Filliben, 1975). Homoskedasticity of residuals and the presence of outliers were evaluated by assessing the Studentized residual plot, with outliers defined as Studentized residuals greater than 3 standard deviations. Figures for structural growth measurements present the dependent variable of interest adjusted for random effects according to St-Pierre (2001). This presents graphically only the variation attributable to treatments, while still accounting for random effects in fitting the model. Values presented in Tables 2
and 3
are regression coefficients fit using models 1 and 2; otherwise least squares means are presented. Differences were considered statistically significant where P < 0.05 and tending toward significance where P < 0.10.
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| RESULTS AND DISCUSSION |
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Growth
Heifer growth was monitored weekly to determine potential differences between treatments. Results for initial measurements and daily gains are presented in Table 2
. These results indicate that maintaining the rate of weight gain by controlling access to feed was successful, because ADG between groups were not different (0.828 HF vs. 0.827 HC; SE ± 0.010 kg/d; P > 0.94). Average DMI required to obtain these levels of gain were 5.96 HF and 5.32 HC kg/d (SE ± 0.12; P < 0.001), and the associated FE (kg of ADG/kg of DMI) was 0.142 HF and 0.156 HC (SE ± 0.003; P < 0.002) for the experimental growth period.
Heifers in this trial had, on average, greater withers height and body length per unit of BW than the upper ranges of size recommended by Hoffman (1997). Although only weight gain could be controlled, growth of all structural characteristics, except for paunch girth, was unaffected by dietary regimen, expressed either as daily gains (cm/d) or as allometric expressions (Tables 2
and 3
, and Figure 1
). The near constancy in structural size and gains indicates that, under the conditions of the current experiment, BW was the major factor influencing these variables and that differences in growth attributable to dietary treatment were minimal.
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Although body composition was not measured, this situation could possibly result from a replacement of gut contents and tissue weight with visceral fat tissue weight. Two serial slaughter experiments, however, failed to detect any differences in visceral tissue fat accretion in animals fed HF and HC diets for similar ME intakes, even though ADG was greater for HC-fed animals in both experiments (Coleman et al., 1995; Kim et al., 2003). The precise cause of the greater rate of paunch girth gain for heifers fed HC is unknown from the results of the current experiment; however, knowledge of the composition of the visceral tissues and gut fill would be of great utility in practical feeding systems in which HC diets are fed for controlled growth.
Puberty
Results related to characteristics of heifers at attainment of puberty are presented in Table 4
. Heifers receiving HC were younger and lighter at puberty than were those fed HF, although these results were not significant. Results from this experiment compare closely with other recently published results concerning BW of Holstein heifers at puberty. Capuco et al. (1995) reported BW at puberty lower than the results of this experiment; several other experiments reported higher BW (Radcliff et al., 1997; Lammers et al., 1999), and others were similar (Meyer et al., 2006). The weighted average of BW at puberty of the 221 heifers from these published experiments is 291 kg compared with an average of 290 kg in the current experiment.
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Lifetime and experimental ADG until puberty did not differ between diets. The experimental period began after the calfhood phase, in which nutritional alterations can influence mammary development (Brown et al., 2005), and treatment rations were fed until after puberty had been confirmed. Thus, treatments were delivered during the physiological stage associated with prepubertal allometric mammary growth (Sinha and Tucker, 1969), and ADG were not different between dietary groups during this stage. This result is consistent with our objective of not only keeping ADG similar between groups, but also maintaining it at an optimal level that has been shown to allow maximal levels of production (Zanton and Heinrichs, 2005).
Lactation
After completing the experimental feeding period, all heifers were managed as a group in accordance with protocols of the Penn State dairy. Although the number of animals is too small for inferences about reproduction to be powerful, conception rate was not affected by treatment (P > 0.42) and occurred at a time that allowed heifers to calve at an average age of 23.4 mo (Table 5
; P > 0.50). Average BW at calving (BWC) was 548 kg and was not different between treatments (Table 5
, P > 0.16). These averages are within the currently recommended range; however, average BWC for heifers fed HF equaled the lower limit of the range recommended for BWC (Hoffman, 1997).
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Current recommendations for ADG prior to puberty are related only to LW ADG (Hoffman, 1997; Sejrsen et al., 2000; Zanton and Heinrichs, 2005). How the EBW ADG in this trial fits relative to that which would result in maximum first-lactation milk production is not known. From milk production results, it can be inferred that the level of accelerated prepubertal EBW gains predicted for heifers fed HC in this experiment did not translate into reduced milk production.
The results from the first 150 d of first lactation are shown in Tables 5
and 6
and in Figure 2
. For most production values, heifers fed HC prior to puberty had an advantage, although this advantage was significant only for FCM and fat production (P < 0.02). Covariate analysis of BWC and BW loss to nadir were not signifi-cant for production results (P > 0.30) and are not included in results presented. The only significant covariate determined by regression analysis was total BW change over 150 DIM (P < 0.01 for all variables except protein production and fat concentration: P > 0.10). Including this covariate reduced the significance for FCM to P < 0.086 from 0.020 and for fat production to P < 0.054 from 0.013. In all cases in which covariates were included, heifers fed HC prior to puberty maintained a numerical advantage over heifers fed HF, although with less statistical significance.
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| CONCLUSIONS |
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| ACKNOWLEDGEMENTS |
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| FOOTNOTES |
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Received for publication January 21, 2007. Accepted for publication April 2, 2007.
| REFERENCES |
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