M. K. Nielsen

Partial-genome evaluation of postweaning feed intake and efficiency of crossbred beef cattle

The effects of individual SNP and the variation explained by sets of SNP associated with DMI, metabolic midtest BW, BW gain, and feed efficiency, expressed as phenotypic and genetic residual feed intake, were estimated from BW and the individual feed intake of 1,159 steers on dry lot offered a 3.0 Mcal/kg ration for at least 119 d before slaughter. Parents of these F(1) × F(1) (F(1)(2)) steers were AI-sired F(1) progeny of Angus, Charolais, Gelbvieh, Hereford, Limousin, Red Angus, and Simmental bulls mated to US Meat Animal Research Center Angus, Hereford, and MARC III composite females. Steers were genotyped with the BovineSNP50 BeadChip assay (Illumina Inc., San Diego, CA). Effects of 44,163 SNP having minor allele frequencies >0.05 in the F(1)(2) generation were estimated with a mixed model that included genotype, breed composition, heterosis, age of dam, and slaughter date contemporary groups as fixed effects, and a random additive genetic effect with recorded pedigree relationships among animals. Variance in this population attributable to sets of SNP was estimated with models that partitioned the additive genetic effect into a polygenic component attributable to pedigree relationships and a genotypic component attributable to genotypic relationships. The sets of SNP evaluated were the full set of 44,163 SNP and subsets containing 6 to 40,000 SNP selected according to association with phenotype. Ninety SNP were strongly associated (P < 0.0001) with at least 1 efficiency or component trait; these 90 accounted for 28 to 46% of the total additive genetic variance of each trait. Trait-specific sets containing 96 SNP having the strongest associations with each trait explained 50 to 87% of additive variance for that trait. Expected accuracy of steer breeding values predicted with pedigree and genotypic relationships exceeded the accuracy of their sires predicted without genotypic information, although gains in accuracy were not sufficient to encourage that performance testing be replaced by genotyping and genomic evaluations.

Genetic and phenotypic parameter estimates for feed intake and other traits in growing beef cattle, and opportunities for selection.

Growth, feed intake, and temperament indicator data, collected over 5 yr on a total of 1,141 to 1,183 mixed-breed steers, were used to estimate genetic and phenotypic parameters. All steers had a portion of Hereford, Angus, or both as well as varying percentages of Simmental, Charolais, Limousin, Gelbvieh, Red Angus, and MARC III composite. Because the steers were slaughtered on various dates each year and the animals thus varied in days on feed, BW and feed data were adjusted to a 140-d feeding period basis. Adjustment of measures of feed efficiency [G:F or residual feed intake (RFI), intake adjusted for metabolic body size, and BW gain] for body fatness recorded at slaughter had little effect on the results of analyses. Average daily gain was less heritable (0.26) than was midtest BW (MBW; 0.35). Measures of feed intake had greater estimates of heritability, with 140-d DMI at 0.40 and RFI at 0.52; the heritability estimate for G:F was 0.27. Flight speed (FS), as an indicator of temperament, had an estimated heritability of 0.34 and a repeatability of 0.63. As expected, a strong genetic (0.86) correlation was estimated between ADG and MBW; genetic correlations were less strong between DMI and ADG or MBW (0.56 and 0.71). Residual feed intake and DMI had a genetic correlation of 0.66. Indexes for phenotypic RFI and genotypically restricted RFI (no correlation with BW gain) were compared with simple economic indexes incorporating feed intake and growth to elucidate expected selection responses under different criteria. In general, few breed differences were detected across the various measurements. Heterosis contributed to greater DMI, RFI, and MBW, but it did not significantly affect ADG, G:F, or FS. Balancing output (growth) with input costs (feed) is needed in practicing selection, and FS would not be recommended as an indicator trait for selection to change feed efficiency. An index including BW gain and RFI produced the best economic outcome.

Differences in mitochondrial efficiency between lines of mice divergently selected for heat loss

Divergent selection for heat loss was applied to lines of mice for 15 generations (G) in 3 replicates. Selection resumed at G42 and continued through G51 across all replicates. At the end of G51, differences in heat loss and feed intake per unit of BW were approximately 56 and 34%, respectively, between high heat loss (MH) and low heat loss (ML) lines, as a percentage of the control line (MC) mean. Rates of liver mitochondrial respiration states, degree of coupling, and mitochondrial efficiency were measured in G58 using a Clark-type oxygen electrode to investigate possible causes of underlying variation in maintenance requirements. Body composition, BW, liver weight, feed intake, and residual feed intake (RFI) were also measured or calculated. Results reported here represent data from 197 mature male mice from all replicates. There were no differences in BW (P = 0.91) between the selection lines. Selection had an effect on lean percentage (P = 0.02), with MH mice being leaner. Fat percentage differences between the selection lines tended toward significance (P = 0.13). Livers of MH mice were approximately 13% larger than livers of ML mice (P = 0.01). An effect of selection was observed (P < 0.01) in feed intake per unit BW, with MH mice consuming 29% more feed than ML mice in G58. Differences in state 2 and state 4 respiration rates were significant (P = 0.01), whereas state 3 rates approached significance (P = 0.06). Mitochondria of MH mice respired at a greater rate than mitochondria of ML mice in all states of respiration; ML mice had respiratory control ratios that were, on average, 8% greater than MH mice (P = 0.14). Although this difference only tended toward significance, we suspect a greater degree of coupling of mitochondrial processes exists in ML animals. Mice selected for reduced heat loss had ADP:oxygen ratios that were approximately 20% greater than MH mice (P = 0.03). Therefore, greater mitochondrial efficiency was expressed in the ML animals. Within a line-replicate, there was no correlation between ADP:O and feed intake per unit BW (P = 0.71). In addition, no correlation of ADP:O and RFI existed (P = 0.92). Although the selection lines differed in mitochondrial traits, including overall mitochondrial efficiency (ADP:oxygen), these differences were not a significant underlying cause of variation in feed intake per unit BW or in RFI estimates.

Hepatic Mitochondrial Efficiency in Lines of Mice Differing in Feed Intake

Selection was practiced for high (MH) and low (ML) heat loss using direct calorimetry to create lines of mice differing in feed intake per unit average BW (FI/BW). A total of 25 generations of selection was practiced during maintenance of an unselected control (MC). As a percentage of MC, mice of MH and ML lines differed in heat loss and FI/BW by 56 and 34%, respectively, when selection ceased. The purpose of this study was to determine hepatic mitochondrial efficiency in mice at maintenance across the genetic lines. After measurement of feed intakeand BW over a 2-wk period, mice were euthanized, and their livers were extracted. Livers were homogenized and mitochondria were isolated in buffer for measurement of oxygen consumption and, hence, mitochondrial activity using a Clark-type oxygen electrode. Mitochondria of ML mice expressed greater (P < 0.03) respiratory control ratio by 22% than that for MH mice, revealing greater degree of uncoupling during electron transport in MH mice. Difference in FI/BW (P < 0.001) was large with MH mice consuming 34% more than ML mice. The respiratory control ratio differed between mice of MH and ML lines as expected; however, regression analysis revealed that it explained essentially no variation in feed intake per BW.

Lifetime reproductive performance and survival analysis of mice divergently selected for heat loss.

Divergent selection for heat loss was implemented in mice creating maintenance high (MH) and low maintenance (ML) lines and an unselected control (MC) in 3 independent replicates. Mice from the ML line have improved feed efficiency, due to decreased maintenance energy requirement, but there is potential for a correlated decline in reproductive performance and survivability. Number fully formed (NFF), number born alive (NBA), number weaned (NW), litter weaning weight (LWW), pup weaning weight (PWW), fraction alive at birth (FAB), fraction alive at weaning, and birth interval were recorded at every parity on 21 mating pairs from each line × replicate combination cohabitated at 7 wk of age and maintained for up to 1 yr. Traits were summed over parities to evaluate lifetime production. Pairs were culled due to death or illness, no first parity by 42 d cohabitation, 2 consecutive litters with none born alive, 3 consecutive litters with none weaned, 42 d between parities, or average size of most recent 2 litters less than half the average of first 3 litters. Survival probabilities were produced and evaluated for each line and used to calculate mean number of parities using a Markov-chain algorithm assuming a maximum of 4, 6, 8, 10, or 12 parities or 1 yr. Line was insignificant for all litter traits while NFF, NW, and FAB decreased with parity (P < 0.05) and PWW tended to increase (P < 0.07). The MC mice had higher lifetime NW, LWW, and PWW (P < 0.04). Birth interval showed that MH mice had increasingly larger intervals while remaining the same in ML mice (P < 0.01). In the survival analysis, MC mice had the greatest survival rates overall, but ML mice had the greatest rates in the period up to 5 parities while MH mice had the greatest rates in later parities. This resulted in greater mean number of parities for ML mice up to maximum of 8 parities and higher means for MH mice when the maximum number of allowed parities was 10 or higher. Reproductive performance was not substantially affected by changing maintenance energy requirements. The ML animals appear to survive well in early parities and produce more parities when a low number of maximum parities is enforced, but this benefit declines in later parities and MH animals survive better and increase mean number of parities when turnover rates are low. Therefore, selection for low maintenance animals may be beneficial for systems desiring a short generation interval but less so for systems desiring longevity.

Body composition and feed intake of reproducing and growing mice divergently selected for heat loss

Changes in maintenance energy requirements and in feed efficiency have been achieved by divergent selection for heat loss in mice in 3 replicates, creating high heat loss, high maintenance (MH) and low heat loss, low maintenance (ML) lines and an unselected control (MC). However, feed intake has mainly been measured in mature animals and not during growth or reproduction. Additionally, there is evidence that reducing maintenance energy will increase fat content, an undesirable result. To evaluate if selection has altered body composition and lifecycle feed intake, mating pairs were continuously mated and maintained for up to 1 yr unless culled. Offspring pairs were sampled from each line at each parity and maintained from 21 to 49 d of age. Feed intake was recorded for mating pairs throughout the year and on offspring pairs. Body weight was recorded on all animals at culling as well as percent fat, total fat, and total lean, measured by dual X-ray densitometry. Average daily gain was also recorded for offspring. Energy partitioning was achieved using 2 approaches: Approach I regressed energy intake of the pair on sum of daily metabolic weight and total gain to obtain maintenance (bm) and growth (bg) coefficients for each line, replicate, feeding period, and sex (offspring pairs only); Approach II calculated bm for each pair assuming constant energy values for lean and fat gain. Energy coefficients and body composition traits were evaluated for effect of selection (MH vs. ML) and asymmetry of selection ([MH + ML]/2 vs. MC). Both MC mating and offspring pairs tended to have greater BW than the average of the selection lines (P < 0.08). Males of offspring pairs weighed more than females (P < 0.01), while females of mating pairs weighed more than males (P < 0.01). Line was insignificant (P > 0.15) for body composition traits. Using Approach I, MH mice had a greater bm than ML mice for mating pairs (P = 0.03) but not offspring pairs (P = 0.50). For Approach II, MH had a greater bm than ML mice for both mating (P = 0.01) and offspring pairs (P = 0.01). The effect of selection for heat loss on body composition was smaller than previously reported and unlikely to outweigh the benefit of reduced feed intake, which was shown to be maintained throughout an entire lifecycle that included reproducing animals. Additionally, the reduction in energy intake seems primarily due to reduced maintenance energy costs, validating the success of the selection procedure.

Locomotor activity and body temperature in selected mouse lines differing greatly in feed intake

Locomotor activity, body temperature, feed intake, and BW were measured on 382 mature male mice sampled from lines previously selected (25 generations) for either high (MH) or low (ML) heat loss and an unselected control (MC). Animals were from all 3 independent replicates of the 3 lines and across 4 generations (68 through 71). Locomotor activity and body temperatures were obtained using implanted transmitters with data collection over 4 d following a 3-d postsurgery recovery period. Data were collected every minute and then averaged into 30-min periods, thus providing 192 data points for each mouse. Least-squares means for feed intake adjusted for BW (Feed/BW, feed·BW(-1)·d(-1), g/g) were 0.1586, 0.1234, and 0.1125 (±0.0022) for MH, MC, and ML, respectively, with line being a highly significant source of variation (P < 0.0003). Line effects for locomotor activity counts, transformed to the 0.25 power for analysis, were significantly different, with MH mice being 2.1 times more active than ML mice (P < 0.003); MC mice were intermediate. Differences in body temperature were significant for both line (P < 0.03) and day effects (P < 0.001), with a 0.32°C difference between the MH and ML lines. Fourier series analysis used the combined significant periodicities of 24, 18, 12, 9, 6, and 3 h to describe circadian cycles for activity and body temperature. All 3 lines expressed daily peaks in body temperature and locomotor activity ∼3 h into darkness and ∼2 h after lights were turned on. There was a stronger relationship between locomotor activity and Feed/BW (P < 0.0001) than between body temperature and Feed/BW (P < 0.01); differences between lines in locomotor activity and body temperature explained 17% and 3%, respectively, of differences between lines in Feed/BW. Thus, line differences in locomotor activity contribute to line differences in maintenance, but approximately 80% of the differences between the MH and ML selection lines in Feed/BW remains independent of differences in locomotor activity.

Life cycle biological efficiency of mice divergently selected for heat loss

Divergent selection in mice for heat loss was conducted in 3 independent replicates creating a high maintenance, high heat loss (MH) and low maintenance, low heat loss (ML) line and unselected control (MC). Improvement in feed efficiency was observed in ML mice due to a reduced maintenance energy requirement but there was also a slight decline in reproductive performance, survivability, and lean content, particularly when compared to MC animals. The objective of this study was to model a life cycle scenario similar to a livestock production system and calculate total inputs and outputs to estimate overall biological efficiency of these lines and determine if reduced feed intake resulted in improved life cycle efficiency. Feed intake, reproductive performance, growth, and body composition were recorded on 21 mating pairs from each line × replicate combination, cohabitated at 7 wk of age and maintained for up to 1 yr unless culled. Proportion of animals at each parity was calculated from survival rates estimated from previous research when enforcing a maximum of 4, 8, or 12 allowed parities. This parity distribution was then combined with values from previous studies to calculate inputs and outputs of mating pairs and offspring produced in a single cycle at equilibrium. Offspring output was defined as kilograms of lean output of offspring at 49 d. Offspring input was defined as megacalories of energy intake for growing offspring from 21 to 49 d. Parent output was defined as kilograms of lean output of culled parents. Parent input was defined as megacalories of energy intake for mating pairs from weaning of one parity to weaning of the next. Offspring output was greatest in MC mice due to superior BW and numbers weaned, while output was lowest in ML mice due to smaller litter sizes and lean content. Parent output did not differ substantially between lines but was greatest in MH mice due to poorer survival rates resulting in more culled animals. Input was greatest in MH and lowest for ML mice for both offspring and parent pairs, consistent with previous results in these lines. Life cycle efficiency was similar in MC and ML mice, while MH mice were least efficient. Ultimately, superior output in MC mice slightly outweighed the lower inputs in ML animals resulting from decreased maintenance energy requirements. Therefore, selection to reduce maintenance energy requirements may be more useful in terminal crosses or in a selection index to reduce possible negative effects on output, especially reproductive performance.

Selection for postweaning gain in rats: III. Correlated response in maternal performance.

Effects from 34 generations of selection, either up (U) or down (D), for 3- to 9-wk weight gain on genetic direct and postnatal maternal effects, dam size and maternal efficiency were evaluated in 25 cross-fostered sets of rats. Direct genetic effects on 12-d pup weight were 14 and 9% above controls (C) for U and D lines, respectively (P less than .01). Postnatal maternal effects on 12-d pup weight were also 11% higher for the U line (P less than .01), but not different from C for the D line. Females of the U line were heavier (P less than .01) by 13% at mating, 12% at parturition and 15% at 12 d of lactation, relative to controls. They also produced 11% greater total litter weight at 12 d and consumed 9% more feed, but were only nonsignificantly lower in feed per unit of pup 12-d litter weight. Females of D line were -8, -7 and -3% relative to controls in the three weights, but did not differ from controls in 12-d litter weight, feed consumed or in feed efficiency during lactation.