The effects of selection for different combinations of weights at two ages on the growth curve of mice

Summary The weights of mice in lines selected for different combinations of high and low body weights at 5 and at 10 weeks of age were recorded from 3 to 21 weeks of age. The average growth curve for each line was computed using the Gompertz function. The growth curves of lines selected for high or low weight at a single age (ST lines) showed large differences in estimates of mature size and small differences in estimates of maturing rate, i.e. of the relative rate of growth to maturity. The growth curves of lines selected by independent culling for divergent combinations of deviations of opposite sign in 5- and 10-week weights (ICL lines) showed little difference in estimates of mature size and a large difference in estimates of maturing rate. The growth curves of lines selected by index for divergence in 5-week weight with no change in 10-week-weight or for divergence in 10-week-weight with no change in 5-week weight showed large differences in estimates of mature size and large differences in estimates of the maturing rate. The relationship between mature size and maturing rate was affected in different ways by the three types of selection.     

Using quantile regression methodology to evaluate changes in the shape of growth curves in pigs selected for increased feed efficiency based on residual feed intake

Growth rate is a major component of feed efficiency when estimating residual feed intake (RFI). Quantile regression (QR) methodology can be used to identify animals with different growth trajectories. The objective of this study was to evaluate the use of QR to identify phenotypic and genetic differences in pigs selected for low RFI. Using performance data on 750 Yorkshire pigs selected for low RFI, individual average daily gain (ADG), average daily feed intake (ADFI), RFI and Gompertz growth curve parameters (asymptotic weight (a), inflection point (b) and decay parameter (c)) were estimated for each pig. Using QR methodology, three Gompertz growth curves were estimated for the whole population for three quantiles (0.1, 0.5 and 0.9) of the BW data. Each animal was classified into one of the quantile regression groups (QRG) based on their overall Euclidian distance between each observed and estimated BW from the quantile growth curves. These three curves were also estimated using only part of the data (generations −1 to 3, and −1 to 4) in order to evaluate the agreement classification rate of animals from later generations into QRGs. We evaluated the effect of QRG on growth parameters and performance traits. Genetic parameters were estimated for these traits, as well as for QRG. In addition, genetic trends for each QRG were estimated. Three distinct growth curves were observed for animals classified into either quantiles 0.1 (QRG_0.1), 0.5 (QRG_0.5) or 0.9 (QRG_0.9). When only part of the data was used to estimate quantile growth curves, all animals from QRG_0.1 were correctly classified in their group. Animals in QRG_0.1 had significantly lower ADFI, ADG and RFI, and greater a, b and c than animals in the other groups. Quantile regression groups analysed as a trait was highly heritable (0.41) and had high (0.8) and moderate (0.46) genetic correlations with ADG and RFI, respectively. Selection for reduced RFI increased the number of animals classified as QRG_0.1 in the population. Overall, downward genetic trends were observed for all traits as a function of selection for reduced RFI. However, QRG_0.1 was the only group that had a positive genetic trend for ADG. Altogether, these results indicate that selection for reduced RFI changes the shape of growth curves in Yorkshire in pigs, and that QR methodology was able to identify animals having different genetic potential for feed efficiency, bringing a new opportunity to improve selection for reduced RFI.     

Decreased frequency of the rat growth hormone transgene in mouse populations with or without selection for increased adult body weight

Summary Frequencies of mice with the rat growth hormone (rGH) transgene were examined in lines derived from two genetic bases (P/W and P/C). The genetic bases were developed from males (P) with the rGH transgene, mated with non-transgenic females of different origin: a line previously selected for large body size (W) and a corresponding unselected control line (C). They were maintained for six generations under random mating with or without selection for increased 42-day body weight. The frequencies of P/W and P/C males with the rGH transgene wer 0.075 and 0.300, respectively at generation 0 of the genetic bases. They were significantly (P<0.05) lower than the expected frequency (about 0.5). At generation 6, the frequencies had decreased further both in selected and unselected lines (ranging from 0.025 to 0.125). Decreased frequencies of mice with the transgene were confirmed in a separate experiment testing segregation of the transgene. The reasons for these decreases are not clear. The results suggest that transgenes need to be monitored when transgenic animals are mated with animals of different origin.Animal Research Centre Contribution No. 1697     

Divergent selection on 63-day body weight in the rabbit: response on growth,carcass and muscle traits

The effects of selection for growth rate on weights and qualitative carcass and muscle traits were assessed by comparing two lines selected for live body weight at 63 days of age and a cryopreserved control population raised contemporaneously with generation 5 selected rabbits. The animals were divergently selected for five generations for either a high (H line) or a low (L line) body weight, based on their BLUP breeding value. Heritability (h²) was 0.22 for 63-d body weight (N = 4754). Growth performance and quantitative carcass traits in the C group were intermediate between the H and L lines (N = 390). Perirenal fat proportion (h² = 0.64) and dressing out percentage (h² = 0.55) ranked in the order L < H = C (from high to low). The weight and cross-sectional area of the Semitendinosus muscle, and the mean diameter of the constitutive myofibres were reduced in the L line only (N = 140). In the Longissimus muscle (N = 180), the ultimate pH (h² = 0.16) and the maximum shear force reached in the Warner-Braztler test (h² = 0.57) were slightly modified by selection.     

Genetic architecture of growth curve parameters in chickens

Summary Genetic improvement in growth of poultry has traditionally proceeded via selection for body weight at a fixed age. Due to increased maintenance costs and reproductive problems of adult broiler breeders, the potential for genetic manipulation of the growth curve has been receiving increased interest. Research of both male and female progeny of a three-way diallel cross was used to investigate the inheritance of growth curve parameters. The Laird form of the Gompertz equation was used to determine growth curve parameters, and was suited to the juvenile growth data frequently collected from meat-type chickens. Growth rate exhibited significant heterosis due to both autosomes and the sex chromosomes. Age at inflection point also exhibited significant average heterosis, though only among females. Growth rate was also influenced by average line effects, as was age at inflection point. Maternal effects had no influence on growth curve parameters, while additive sex linkage was observed for growth rate. Phenotypic and genetic correlations were calculated among the growth curve parameters and suggest that specific breeding programs could alter the growth trajectory of the contemporary broiler chicken. Moderate heritabilities were observed for the growth curve parameters and support the hypothesis that the growth curve could be altered via genetic manipulation of early postnatal growth, especially during the first 14 days post-hatch.     

Responses to divergent selection for plasma concentrations of insulin-like growth factor-1 in mice

A divergent selection experiment with mice, using plasma concentrations of insulin-like growth factor-1 (IGF-1) at 42 days of age as the selection criterion, was undertaken for 7 generations. Lines were not replicated. To obtain sufficient plasma for the IGF-1 assay, blood from four individuals was volumetrically bulked to obtain a litter mean IGF-1 concentration. This necessitated the use of between family selection. Although inbreeding accumulated in a linear fashion in each of the high, control and low lines, the rates were different for each line (3.6, 1.6 and 5.3% per generation for the high, control and low lines, respectively). As a consequence, the effects of selection and inbreeding are confounded in this experiment. Divergence between the high and low lines in plasma concentrations of IGF-1 continued steadily until generation 5. In generations 6 and 7, there was a reduced degree of divergence and this contributed towards the low realized heritability value of 0.15 +/- 0.12. Six-week liveweight showed a steady positive correlated response to selection for or against plasma concentrations of IGF-1 until generation 4 (high-low difference = 1.7 g = 12%). In generation 5, a substantial drop in 6-week liveweight in the low line relative to both the high and control lines occurred (high-low difference, 3.9; g, 25%). This difference was maintained until generation 7.(ABSTRACT TRUNCATED AT 250 WORDS)     

Replicated selection for insulin-like growth factor-1 and body weight in mice

Summary Five generations of divergent selection for plasma concentration of insulin-like growth factor-1 (IGF-1) and for 12-week body weight were carried out in mice, including randomly selected control lines for each trait. All lines were replicated once (12 lines in total). Each replicate line consisted of eight male and eight female parents per generation. Litter size was standardized to eight pups at birth. Mass selection was applied in the selected lines and within-family random selection in the control lines. Blood was taken from the orbital sinus of individual mice at 12 weeks of age for IGF-1 assay. Realized heritabilities were 0.10±0.01 for IGF-1 and 0.41 ± 0.02 for 12-week weight. The realized genetic correlation between IGF-1 and 12-week weight was 0.58 ± 0.01, with a phenotypic correlation of 0.38. Although the genetic correlation between IGF-1 and body weight in mice is moderately positive, 12-week weight responded 3.5 times as fast to weight selection as to selection for IGF-1.     

Different resource allocation strategies result from selection for litter size at weaning in rabbit does

This study examined the effect of long-term selection of a maternal rabbit line, solely for a reproductive criterion, on the ability of female rabbits to deal with constrained environmental conditions. Female rabbits from generations 16 and 36 (n=72 and 79, respectively) of a line founded and selected to increase litter size at weaning were compared simultaneously. Female rabbits were subjected to normal (NC), nutritional (NF) or heat (HC) challenging conditions from 1st to 3rd parturition. Animals in NC and NF were housed at normal room temperatures (18°C to 25°C) and respectively fed with control (11.6 MJ digestible energy (DE)/kg dry matter (DM), 126 g digestible protein (DP)/kg DM, and 168 g of ADF/kg DM) or low-energy fibrous diets (9.1 MJ DE/kg DM, 104 g DP/kg DM and 266 g ADF/kg DM), whereas those housed in HC were subjected to high room temperatures (25°C to 35°C) and the control diet. The litter size was lower for female rabbits housed in both NF and HC environments, but the extent and timing where this reduction took place differed between generations. In challenging conditions (NF and HC), the average reduction in the reproductive performance of female rabbits from generation 16, compared with NC, was −2.26 (P<0.05) and −0.51 kits born alive at 2nd and 3rd parturition, respectively. However, under these challenging conditions, the reproductive performance of female rabbits from generation 36 was less affected at 2nd parturition (−1.25 kits born alive), but showed a greater reduction at the 3rd parturition (−3.53 kits born alive; P<0.05) compared with NC. The results also showed differences between generations in digestible energy intake, milk yield and accretion, and use of body reserves throughout lactation in NC, HC and NF, which together indicate that there were different resource allocation strategies in the animals from the different generations. Selection to increase litter size at weaning led to increased reproductive robustness at the onset of an environmental constraint, but failure to sustain the reproductive liability when the challenge was maintained in the long term. This response could be directly related to the short-term environmental fluctuations (less severe) that frequently occur in the environment where this line has been selected.     

Changes in the body composition of mice selected for high and low eight week weight

Summary Body composition was studied in three lines of mice, one selected for high (H) and one for low (L) 8 week weight, and one maintained as an unselected control (C). After 25 generations 8 week weights were 41.2g, 30.6 g and 20.5g for the H, C and L lines. Mice were sampled from the lines and analysed for fat, protein, ash and water at generations 14 and 25. Apart from fat in the H line, there was little alteration due to selection in the relationships between individual body components and total body weight. In the H line, the contribution of fat to body weight gain was considerably increased. Although leaner than the C and L mice at low body weights, H line mice rapidly became fatter with increasing body weight. Selection appeared to reduce the body weight at which fat was deposited at its maximum rate in the H line. The H and C lines were equally fat at body weights of 29.0 g and 21.6 g at generations 14 and 25 respectively. Body weights at points of inflection of the growth curves of the H, C and L lines at generation 25 were 18.3 g, 14.3 g and 12.8 g. The implications of these findings for meat species slaughtered at set weights are discussed.     

Prediction of genetic contributions and generation intervals in populations with overlapping generations under selection

A method to predict long-term genetic contributions of ancestors to future generations is studied in detail for a population with overlapping generations under mass or sib index selection. An existing method provides insight into the mechanisms determining the flow of genes through selected populations, and takes account of selection by modeling the long-term genetic contribution as a linear regression on breeding value. Total genetic contributions of age classes are modeled using a modified gene flow approach and long-term predictions are obtained assuming equilibrium genetic parameters. Generation interval was defined as the time in which genetic contributions sum to unity, which is equal to the turnover time of genes. Accurate predictions of long-term genetic contributions of individual animals, as well as total contributions of age classes were obtained. Due to selection, offspring of young parents had an above-average breeding value. Long-term genetic contributions of youngest age classes were therefore higher than expected from the age class distribution of parents, and generation interval was shorter than the average age of parents at birth of their offspring. Due to an increased selective advantage of offspring of young parents, generation interval decreased with increasing heritability and selection intensity. The method was compared to conventional gene flow and showed more accurate predictions of long-term genetic contributions.     

Genetic Analysis of a Strain of Mice Plateaued for Litter Size

A strain of mice (S1) was successfully selected for large litter size for 31 generations, increasing the mean by 4.2 young per litter. After generation 31, there was no further progress and it was concluded that a selection plateau had been reached. Realized heritability decreased during the course of the experiment from 0.16+/-0.06 for the first 15 generations to 0.00+/-0.03 for generations 30 through 45.--In order to explore the nature of the selection plateau, the following groups were derived from line S1 at generation 34 or 35: Upward selection with inbreeding (SF), random (relaxed) selection (SO), and downward selection (SR). Selections were carried out for 10 to 11 generations. The means of SO and SF were similar to those of S1, ruling out any major effect of natural selection or overdominance. SR decreased, the mean averaging 2.3 young per litter below that of S1 during the last three generations. The fact that SR responded to selection indicates that genetic variance was still present in the plateaued population. The SF sublines were crossed when the inbreeding was 95% and a new line, SX, was formed. SX was maintained for three generations and the difference of +0.7 young per litter above the contemporary generations of S1 was significant. The results from this experiment suggest that the selection plateau in line S1 was caused by reduction of additive genetic variance to a very low level. Some nonadditive genetic variance remained, however, and was attributed to recessive alleles at low frequency. In agreement with results reported by Falconer (1971), inbreeding with selection followed by crossing of the inbred sublines proved to be effective in overcoming a selection plateau in litter size.     

Modeling bacterial growth responses

Summary The main steps of modeling bacterial growth responses are summarized and a new model for growth curves is shown. Its advantages are analyzed from some theoretical and practical points of view. The new model fits better and has more advantageous statistical properties than the Gompertz curve.     

Evolution of reduced pre-adult viability and larval growth rate in laboratory populations of Drosophila melanogaster selected for shorter development time

Four large (n > 1000) populations of Drosophila melanogaster, derived from control populations maintained on a 3 week discrete generation cycle, were subjected to selection for fast development and early reproduction. Egg to eclosion survivorship and development time and dry weight at eclosion were monitored every 10 generations. Over 70 generations of selection, development time in the selected populations decreased by approximately 36 h relative to controls, a 20% decline. The difference in male and female development time was also reduced in the selected populations. Flies from the selected populations were increasingly lighter at eclosion than controls, with the reduction in dry weight at eclosion over 70 generations of selection being approximately 45% in males and 39% in females. Larval growth rate (dry weight at eclosion/development time) was also reduced in the selected lines over 70 generations, relative to controls, by approximately 32% in males and 24% in females. However, part of this relative reduction was due to an increase in growth rate of the controls populations, presumably an expression of adaptation to conditions in our laboratory. After 50 generations of selection had elapsed, a considerable and increasing pre-adult viability cost to faster development became apparent, with viability in the selected populations being about 22% less than that of controls at generation 70 of selection.     

Artificial selection on larval growth curves in Tribolium: correlated responses and constraints

Body size is often constrained from evolving. Although artificial selection on body size in insects frequently results in a sizable response, these responses usually bear fitness costs. Further, these experiments tend to select only on size at one landmark age, rather than selecting for patterns of growth over the whole larval life stage. To address whether constraints may be caused by larval growth patterns rather than final size, we implemented a function‐valued (FV) trait method of selection, in which entire larval growth curves from Tribolium were artificially selected. The selection gradient function used was previously predicted to give the maximal response and was implemented using a novel selection index in the FV framework. Results indicated a significant response after one generation of selection, but no response in subsequent generations. Correlated responses included increased mortality, increased critical weight, and decreased development time (DT). The lack of response in size and development time after the first generation was likely caused by increased mortality suffered in selected lines; we demonstrated that the selection criterion caused both increased body size and increased mortality. We conclude that artificial selection on continuous traits using FV methods is very efficient and that the constraint of body size evolution is likely caused by a suite of trade‐offs with other traits.     

Pair housing of rabbits reduces variances in growth rates and serum alkaline phosphatase levels

New Council of Europe regulations mandate housing of two rabbits in the same cage space currently used to house one, provided the animals are socially compatible. This study was designed to assess changes in growth and selected serum chemistry parameters due to pair housing or single housing of rabbits. Six sets of four female siblings of Crl:KBL(NZW)BR rabbits were used. The animals were seven weeks old on arrival. Two siblings of each set were allocated to pair housing, two to single housing. The animals were housed in stainless steel cages (120 cm x 60 cm x 60 cm) with a perforated floor, including a shelf (60 cm x 30 cm) at 30 cm height from the floor. The rabbits were provided with an aspen cube (5 cm x 5 cm x 5 cm), one item per animal. The rabbits were weighed and blood samples were taken from the auricular central artery at four different times during the study. Blood sera were assayed for a set of routinely assayed clinical chemistry parameters: alanine aminotransferase (ALT), aspartate aminotransferase (AST), alkaline phosphatase (APHOS), blood urea nitrogen (BUN), cholesterol (CHOL) and protein (PROT). Mean and variance profiles over the study period were statistically analysed by multivariate analysis of variance. No differences in mean profiles were detected; however, weight (P = 0.0002) and APHOS (P = 0.017) variances were significantly lower in pair-housed animals. The reduction in variance on growth and APHOS attributable to pair housing appears to be rather large. During the 21-week study, occasional fighting was seen between the pair-housed rabbits. After sexual maturity, further major fighting bouts resulted in significant trauma that necessitated the cessation of the study. In conclusion, pair housing appears to have a decreasing effect on growth and APHOS variance, but antisocial behaviour such as fighting remains a serious problem.     

Genetic-statistical analysis of growth in selected and unselected mouse lines

The growth of males sampled from two mouse lines long-term selected for over 86 generations on body weight (DU6) or on protein amount (DU6P) was analysed from birth till 120 days of age and compared to the growth of an unselected control line (DUKs). Animals from the selected lines are already approximately 40 to 50% heavier at birth than the controls. This divergence increases to about 210 to 240% at the 120 day of age. With birth weights of 2.2 and 2.4 g and weights of 78 and 89 g at the 120 day these selection lines are the heaviest known mouse lines.

The fit of three modified non-linear growth functions (Gompertz function, Logistic function, Richards function) was compared and the effect of three different data inputs elucidated. The modification was undertaken to use parameters having a direct biological meaning, for example: A: theoretical final body weight, B: maximum weight gain, C: age at maximum weight gain, D (only Richards function): determines the position of the inflection point in relation to the final weight. All three models fit the observed data very well (r² = 0.949–0.998), with a slight advantage for the Richards function. There were no substantial effects of the data input (averages, single values, fitting a curve for every animal with subsequent averaging the parameters).

The high growth of the selected mice is connected with very substantial changes in the final weight and in the maximum weight gain, whereas the changes of the age at the point of inflection were, although partially significant, relatively small and dependent on the model used.     

Reproductive robustness differs between generalist and specialist maternal rabbit lines: the role of acquisition and allocation of resources

Background

Farm animals are normally selected under highly controlled, non-limiting conditions to favour the expression of their genetic potential. Selection strategies can also focus on a single trait to favour the most ‘specialized’ animals. Theoretically, if the environment provides enough resources, the selection strategy should not lead to changes in the interactions between life functions such as reproduction and survival. However, highly ‘specialized’ farm animals can be required for breeding under conditions that differ largely from selection conditions. The consequence is a degraded ability of ‘specialized’ animals to sustain reproduction, production and health, which leads to a reduced lifespan. This study was designed to address this issue using maternal rabbit lines. A highly specialized line with respect to numerical productivity at weaning (called V) and a generalist line that originated from females with a long reproductive life (called LP) were used to study the strategies that these lines develop to acquire and use the available resources when housed in different environments. In addition, two generations of line V, generations 16 and 36, were available simultaneously, which contributed to better understand how selection criteria applied in a specific environment changed the interplay between functions related to reproduction and survival.

Results

We show that, under constrained conditions, line LP has a greater capacity for resource acquisition than line V, which prevents excessive mobilization of body reserves. However, 20 generations of selection for litter size at weaning did not lead to an increased capacity of nutrient (or resource) acquisition. For the two generations of line V, the partitioning of resources between milk production, body reserves preservation or repletion or foetal growth differed.

Conclusions

Combining foundational and selection criteria with a specific selection environment resulted in female rabbits that had a different capacity to deal with environmental constraints. An increased robustness was considered as an emergent property of combining a multiple trait foundational criterion with a wide range of environmental conditions. Since such a strategy was successful to increase the robustness of female rabbits without impairing their productivity, there is no reason that it should not be applied in other livestock species.