Growth and body composition changes in mice selected for high post-weaning weight gain on two levels of feeding

Summary Two lines of mice were selected for high post-weaning weight gain (3 to 6 weeks) adjusted for 3 week weight. One line (F) was grown on freely available food and the other (S) on a feeding scale set at the same level for all mice. Food intake of the S line averaged 80% of the F line. The realised heritabilities after 6 generations of selection were 0.38±0.06 and 0.33±0.07 for the F and S lines, respectively. In generation 7, mice from the F and S lines and from an unselected control line (C) were compared on both free and set levels of feeding from 3 weeks to 9 weeks of age. Measurements taken were growth rate, appetite, food conversion efficiency (weight gain/food intake) and body composition (fat, protein, ash, water). The F and S lines grew more rapidly and efficiently than the C line on both levels of feeding, each line performing best on the level of feeding on which it was selected. The average genetic correlation between growth rates of the same line on the two feeding levels was 0.54±0.10. The F line grew 19% faster and was 9% more efficient than the S line on free feeding but the S line grew 15% faster and was 15% more efficient than the F line on set feeding. Relative to the C line, food intake per day on free feeding was 4% higher in the F line and 6% lower in the S line. There was no difference between the lines in food intake/g body weight. The rate of deposition of all body components increased in both selection lines. In the F, S and C lines respectively, efficiencies of gains in body components (10²x gain/food) were 1.79, 1.31 and 1.06 for fat, 1.53, 1.63 and 1.22 for protein and 5.88, 6.45 and 4.98 for protein + water. Apparently energy lost as heat was reduced in both the F and S lines. The partitioning of energy retained was altered in favour of more fat in the F line and more protein in the S line.     

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.     

Changes in growth,appetite, food conversion efficiency and body composition in mice selected for high post-weaning weight gain on restricted feeding

Summary This study aimed to test the hypothesis that if animals were fed the same amount over the same time period, selection of the fastest growers would result in a change in the partitioning of metabolisable energy toward more protein and less fat deposition. Two mouse lines (S1 and S2) were selected for high 5 to 9 week weight gain corrected to mean 5 week weight. Appetite variation between mice was eliminated by feeding a fixed amount to each mouse daily. After 6 generations of selection, the lines were compared with an unselected control (C) on restricted and ad libitum levels of feeding for growth rate, appetite, food conversion efficiency and chemical body composition.Realised heritabilities of 5 to 9 week gain were 0.36+ 0.05 and 0.19±0.04 for S1 and S2 respectively. Nine week weights were increased by an average of 13% on both feeding levels. Most of this increase, particularly in S2, occurred before 5 weeks and was therefore outside the period of measurement used in selection. On ad libitum feeding, selection increased food intake per unit time by 6% but there was no increase per unit body weight. Food conversion efficiency (gain/food) increased by 12%. Compared with controls at 9 weeks, 3% more of the body weights of selected mice was fat and 1% less was protein. These differences were reduced but were still in the same direction when comparisons were made at the same body weight. Thus the expected change in energy partitioning toward greater protein and less fat deposition in the S lines did not occur.It was concluded that the increased growth and energy retention in the S lines was brought about by a reduction in maintenance requirement. To achieve the desired change in energy partitioning using a similar selection scheme, higher levels of dietary protein should be fed, and some measure of protein deposition rather than growth rate used as the selection criterion.     

Agouti-related protein-deficient mice display an age-related lean phenotype

Endogenous modulators of the central melanocortin system, such as the agouti-related protein (AgRP), should hold a pivotal position in the regulation of energy intake and expenditure. Despite this, AgRP-deficient mice were recently reported to exhibit normal food intake, body weight gain, and energy expenditure. Here we demonstrate that 2- to 3-month-old Agrp null mice do in fact exhibit subtle changes in response to feeding challenges (fasting and MCR agonists) but, of more significance and magnitude, exhibit reduced body weight and adiposity after 6 months of age. This age-dependent lean phenotype is correlated with increased metabolic rate, body temperature, and locomotor activity and increased circulating thyroid hormone (T4 and T3) and BAT UCP-1 expression. These results provide further proof of the importance of the AgRP neuronal system in the regulation of energy homeostasis.     

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.     

A strong response to selection on mass-independent maximal metabolic rate without a correlated response in basal metabolic rate

Metabolic rates are correlated with many aspects of ecology, but how selection on different aspects of metabolic rates affects their mutual evolution is poorly understood. Using laboratory mice, we artificially selected for high maximal mass-independent metabolic rate (MMR) without direct selection on mass-independent basal metabolic rate (BMR). Then we tested for responses to selection in MMR and correlated responses to selection in BMR. In other lines, we antagonistically selected for mice with a combination of high mass-independent MMR and low mass-independent BMR. All selection protocols and data analyses included body mass as a covariate, so effects of selection on the metabolic rates are mass adjusted (that is, independent of effects of body mass). The selection lasted eight generations. Compared with controls, MMR was significantly higher (11.2%) in lines selected for increased MMR, and BMR was slightly, but not significantly, higher (2.5%). Compared with controls, MMR was significantly higher (5.3%) in antagonistically selected lines, and BMR was slightly, but not significantly, lower (4.2%). Analysis of breeding values revealed no positive genetic trend for elevated BMR in high-MMR lines. A weak positive genetic correlation was detected between MMR and BMR. That weak positive genetic correlation supports the aerobic capacity model for the evolution of endothermy in the sense that it fails to falsify a key model assumption. Overall, the results suggest that at least in these mice there is significant capacity for independent evolution of metabolic traits. Whether that is true in the ancestral animals that evolved endothermy remains an important but unanswered question.     

Selection for components related to body composition in mice: correlated responses

Summary Correlated responses were estimated in each of two replicate lines of mice selected within full-sib families for high (HF) or low (LF) 12-week epididymal fat pad weight as a percentage of body weight (epididymal fat pad percentage), or high (HL) or low (LL) 12-week hind carcass weight as a percentage of body weight (hind carcass percentage). Two replicate control lines (RC) were maintained. Correlated traits were measured in each of the 10 generations of selection. Realized (r_{G R}) and offspring-sire genetic correlations generally were in agreement. In HF and LF, 3–6 week postweaning gain (r_{G R} = 0.36±0.04) and feed intake (r_{G R} = 0.50±0.13) had positive correlated responses, but feed efficiency and feed intake/metabolic body size did not change. Positive correlated responses were found for subcutaneous fat pad percentage, body weight-adjusted subcutaneous fat pad weight and fat percentage in the hind carcass (r_{G R}'s were 1.04±0.13, 0.93±0.13 and 0.90±0.08). In the hind carcass, fat-free dry (protein + ash) percentage showed a small negative correlated response, and fat-free dry weight did not change. In HL and LL, the correlated responses for the above traits were generally opposite to those observed in HF and LF. Litter size, percentage of infertile matings, and preweaning mortality showed no consistent correlated responses in any of the lines, but an index of fitness combining the three traits showed a decrease in all four selection treatments.Paper no. 11057 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, 27695-7601. The use of trade names in this publication does not imply endorsement by the North Carolina Agricultural Research Service of the products named, nor criticisms of similar ones not mentioned     

Reduced serotonin reuptake transporter (SERT) function causes insulin resistance and hepatic steatosis independent of food intake

Serotonin reuptake transporter (SERT) is a key regulator of serotonin neurotransmission and a major target of antidepressants. Antidepressants, such as selectively serotonin reuptake inhibitors (SSRIs), that block SERT function are known to affect food intake and body weight. Here, we provide genetic evidence that food intake and metabolism are regulated by separable mechanisms of SERT function. SERT-deficient mice ate less during both normal diet and high fat diet feeding. The reduced food intake was accompanied with markedly elevated plasma leptin levels. Despite reduced food intake, SERT-deficient mice exhibited glucose intolerance and insulin resistance, and progressively developed obesity and hepatic steatosis. Several lines of evidence indicate that the metabolic deficits of SERT-deficient mice are attributable to reduced insulin-sensitivity in peripheral tissues. First, SERT-deficient mice exhibited beta-cell hyperplasia and islet-mass expansion. Second, biochemical analyses revealed constitutively elevated JNK activity and diminished insulin-induced AKT activation in the liver of SERT-deficient mice. SERT-deficient mice exhibited hyper-JNK activity and hyperinsulinemia prior to the development of obesity. Third, enhancing AKT signaling by PTEN deficiency corrected glucose tolerance in SERT-deficient mice. These findings have potential implications for designing selective SERT drugs for weight control and the treatment of metabolic syndromes.     

Thermoregulatory responses of two mouse Mus musculus strains selectively bred for high and low food intake

We examined the thermoregulatory responses of male and female mice Mus musculus that had been divergently selected on voluntary food intake, corrected for body mass, to produce a high-intake and a low-intake strain. Resting metabolic rate was determined by indirect calorimetry (at 30 degrees C, 25 degrees C, 15 degrees C and 5 degrees C). Body temperature responses were measured in a separate group of mice in a parallel protocol. High-intake mice had significantly elevated body masses compared to low-intake mice in both sexes. Lower critical temperature in both strains appeared to be around 28 degrees C. At 30 degrees C there was a significant strain effect on resting metabolic rate, with high strain mice having greater metabolism than low strain mice. Sex and body mass were not significant main effects on resting metabolic rate and there were no significant interactions. Body temperature measured at 30 degrees C, 25 degrees C, 15 degrees C and 5 degrees C differed significantly between sexes (females higher) and there was a significant sexxbody mass interaction effect, but there was no difference between strains. Thermal conductance was significantly related to strain and sex, mice from the high strain and males having greater thermal conductances than mice from the low strain and females. Artificial selection has resulted in high-intake mice having greater body masses and greater thermal conductances, which together account for up to 45% of the elevated daily energy demands that underpin the increase in food intake. The greater levels of food intake were also associated with higher resting metabolic rates at 30 degrees C.     

Basal metabolic rate of aged mice is affected by random genetic drift but not by selective breeding for high early-age locomotor activity or chronic wheel access

The study of correlated evolution can lead to new insights about the inheritance patterns of complex traits. In order to better understand the evolution of metabolic rate, we tested whether voluntary activity levels and basal metabolic rate are genetically correlated in 90-wk-old mice (Mus domesticus) from replicated lines of the sixteenth generation of an artificial selection experiment for high early-age wheel-running activity. We measured basal rates of oxygen consumption and carbon dioxide production and also computed the respiratory exchange ratio. Half of the individuals from both selected and control lines had been allowed free access to running wheels since 4 wk of age, while the other half were in standard cages. This design allowed testing of hypotheses about (1) genetic correlations between voluntary activity and metabolic rate and (2) lifetime training effects on metabolic traits. Selection group did not have a significant effect on metabolic traits; therefore, this study does not support some of the implicit assumptions of the aerobic capacity model for the evolution of vertebrate energetics. Activity group also did not affect metabolic rate, indicating that lifetime training does not alter basal metabolism in these mice. However, strong replicate line-within-selection-group differences were detected, indicating the occurrence of random genetic drift. In females, this divergence in metabolic traits attributable to drift was independent of body mass, but in males it was probably caused by a correlated response to selection involving body mass. This study is the first to show such effects of random genetic drift on metabolic traits.     

Environmental maternal influences on body composition in mice selected for body weight

Summary The effect of the postnatal maternal environment, simulated by rearing mice in litters of three, six or nine, on body weight and body composition was investigated in three lines of mice differing widely in growth rate. The lines were selected for high (H₆) and low (L₆) 6-week body weight while the control line was maintained by random selection. Body weight and weights and percentages of ether extract, water, ash and protein at 21, 42, 63 and 84 days were recorded. With few exceptions, there were positive correlated responses to selection in body weight and in weights of body components. At 21 and 42 days the correlated responses were larger in L₆ mice than in H₆ mice. Body weight and weights of body components were larger for mice reared in litters of three than for those reared in litters of nine. Also, mice reared in litters of six were intermediate in body weight and weights of some of the body components between those reared in litters of three and nine. Differences in body weight and weights of body components due to postnatal maternal environment were small by comparison with differences due to genetic line. There were significant line by maternal environment interactions in body weight at 21 days and in ether extract weight at 21 and 63 days. Line and maternal environment differences in percentages of body components did not follow any consistent trend. The results for percentages of body components were further complicated by line x maternal environment interactions. In general, both line and postnatal maternal environmental differences in percentages of body components diminished with age.Paper No. 5670 of the Journal Series of the North Carolina Agricultural Experiment Station, Raleigh, North Carolina 27650. The use of trade names in this publication does not imply endorsement by the North Carolina Agricultural Experiment Station of the products named, nor criticism of similar ones not mentioned     

LH receptor induction and ovulation rate in mice selected for litter size and body weight

Female mice from lines which had undergone long-term single trait and antagonistic index selection for litter size and body weight were analysed for ovulation rate and LH receptor induction. Compared to randomly selected controls, selection for large litter size increased ovulation rate (60%; P less than 0.001) and decreased LH receptor induction per microgram ovarian DNA (87%; P less than 0.01). Selection for large body weight increased ovulation rate (18%; P less than 0.001), but did not lead to a significant correlated response in LH receptor induction. Index selection for large litter size and small body weight increased ovulation rate (14%; P less than 0.01) and decreased LH receptor induction (72%; P less than 0.01), while index selection for small litter size and large body weight did not significantly alter either ovulation rate or LH receptor induction. LH receptor quantities in testes of males from the 5 lines did not exhibit the among-line profile which was observed in ovaries of females. These results confirm the role of ovulation rate in mediation of the positive genetic correlation between litter size and body weight in mice. Increased ovulation rate in mice selected for large litter size may be due to mechanisms associated with LH receptors as well as factors related to growth. In contrast, increased ovulation rate in mice selected for large body weight may be due exclusively to factors related to growth.     

Effect of selection for development rate on reproductive onset in female mice.

This research reports analyses of correlated response in reproductive onset in ICR mice after 23 generations of restricted index selection for divergent body weight gain, early (birth-10 days) or later (28-56 days) in life. Long-term selection altered growth trajectories and 56 day body weight of individuals under different selection regimes in this study. Mice in lines under early selection have the same percentage mature weight at vaginal opening as controls (63%). Vaginal opening is delayed in mice selected for slow early growth, which take longer to reach what appears to be a weight threshold. In contrast, individuals in lines selected for later slow growth undergo vaginal opening at the same age as controls, but at a lower weight and increased percentage mature weight. Pre-compensation or 'counter-balance growth' is observed in these lines, with mice selected for late enhanced growth reaching 52% of mature weight at vaginal opening while mice with late slow growth attain 71% of mature weight prior to vaginal opening. Only 42% of mice with late slow growth attain first oestrus by 56 days. We speculate this is a function of growth rate and fat/lean ratio. Mice with early slow growth show compensatory growth, reaching first oestrus at a similar time to controls. We conclude that selection for growth rate has asymmetrically affected reproductive onset, with lines selected for suppressed gains experiencing delays in the reproductive onset traits measured.     

Restricted selection index in mice designed to change body fat without changing body weight: correlated responses

Summary Correlated responses were studied in lines of mice selected for eight generations based on the criterion of a restricted selection index. Two replicate lines were selected in each treatment as follows: HE, high epididymal fat pad weight (EF) with zero change in body weight (BW) at 12 weeks of age; LE; low EF with zero change in BW; and RS, randomly. Correlated responses showed considerable variation between replicates, suggesting that genetic drift was important. Further, correlated responses for most traits were relatively small, probably because of low selection intensity. The HE line responded as expected in component traits of the restricted index. Associated compositional traits in HE responded as predicted since traits correlated with adiposity increased and hind carcass weight did not change significantly. Feed intake increased and feed efficiency (weight gain/feed intake) decreased in HE, as predicted. In contrast, the LE line did not respond in component traits as predicted since EF did not decrease and BW increased. Consequently, LE exhibited little change in traits associated with adiposity, but hind carcass weight, feed intake and feed efficiency increased. Of the correlated responses scored for fitness traits (littering rate, number of days from pairing of mate to littering, litter size and preweaning pup survival rate), significant effects were found for decreased littering rate in LE and increased prenatal survival rate in HE. In summary, correlated responses to restricted index selection generally agreed with expectation when responses in component traits of the index were considered.The research reported in this publication was funded by the North Carolina Agricultural Research Service (NCARS), Raleigh, NC 27695-7643, USA. The use of trade names in this publication does not imply endorsement by the NCARS, nor criticism of similar ones not mentioned     

Food consumption and body composition in mice selected for high wheel-running activity

The effects of genetic selection for high wheel running (13th generation) and prolonged access (8 weeks) to running wheels on food consumption and body composition were studied in house mice (Mus domesticus). Mice from four replicate lines selected for high wheel-running activity ran over twice as many revolutions per day on activity wheels as did mice from four replicate control lines. At approximately 49 days of age, all mice were placed individually in cages with access to wheels and monitored for 6 days, after which wheels were prevented from rotating for the "sedentary" individuals. During the experiment, five feeding trials were conducted and body mass was measured weekly. After 8 weeks, body composition was measured by hydrogen isotope dilution. Across the five feeding trials, mice in the "active" group (wheels free to rotate) consumed 22.4% more food than mice in the "sedentary" group (wheels locked); mice from the selected lines consumed 8.4% more food than mice from the control lines (average of all trials; body mass-corrected values). In females, but not males, we found a significant interaction between selection and wheel access treatments: within the "active" group the difference in food consumption between selected and control animals was greater than in the "sedentary" group. At the end of the study, mice from the "active" and "sedentary" groups did not differ significantly in body mass; however, mice from the selected lines were approximately 6% smaller in body mass. Estimated lean body mass did not differ significantly either between selected and control lines or between wheel-access groups (P>0.3). Mice from selected lines had lower total body fat compared to mice from control lines (P=0.05; 24.5% reduction; LSMEANS) as did mice from the "active" compared to "sedentary" group (P= 0.03; 29.2% reduction; LSMEANS). Under these conditions, a sufficient explanation for the difference in body mass between the selected and control lines was the difference in fat content.     

Heterosis among lines of mice selected for body weight

Summary To examine the effect of selection on levels of heterosis, crosses were made between three groups of six lines of mice, one group unselected (controls) and the other two selected for high (large lines) and low (small lines) 6-week body weight, respectively. The coefficient of inbreeding of each line was about 0.60. Each line was crossed reciprocally to one line from each of the parental groups, as well as producing purebred progeny. Heterosis for 3-week weight, 6-week weight and 3–6 week gain averaged 0.0%, 2.4% and 4.2%, respectively, and was higher for males than for females. Heterosis was more extensive in crosses involving large or control lines than in crosses with small lines. There was no detectable heterosis in several measures of developmental rate, such as age at vaginal opening. Food conversion efficiency and carcass composition were measured on a sample of the animals. Food consumption, gonadal fat pad weight, and hindquarters weight, each expressed as a proportion of body weight, exhibited –4.0%, 5.6%, and 2.3% heterosis, respectively.     

Locomotor activity of mice divergently selected for basal metabolic rate: a test of hypotheses on the evolution of endothermy

The aerobic capacity model postulates that high basal metabolic rates (BMR) underlying endothermy evolved as a correlated response to the selection on maximal levels of oxygen consumption () associated with locomotor activity. The recent assimilation capacity model specifically assumes that high BMR evolved as a by‐product of the selection for effective parental care, which required long‐term locomotor activity fuelled by energy assimilated from food. To test both models, we compared metabolic and behavioural correlates in males of laboratory mice divergently selected on body mass‐corrected BMR. elicited by running on the treadmill did not differ between selection lines, which points to the lack of genetic correlation between BMR and . In contrast, there was a positive, genetic correlation between spontaneous long‐term locomotor activity, food intake and BMR. Our results therefore corroborate predictions of the assimilation capacity model of endothermy evolution.     

THE RESPONSES OF DROSOPHILA MELANOGASTER TO ARTIFICIAL SELECTION ON BODY WEIGHT AND ITS PHENOTYPIC PLASTICITY IN TWO LARVAL FOOD ENVIRONMENTS

To investigate the potential response to natural selection of reaction norms for age and size at maturity, fresh body weight at eclosion was mass selected under rich and poor larval food conditions in Drosophila melanogaster. The sensitivity of dry weight at eclosion to the difference between rich and poor larval food was selected using differences in sensitivities among families. For both experiments, the correlated response to selection of age at eclosion was examined. The flies were derived from wild populations and had been mass cultured in the lab for more than six months before the experiments started. These flies responded to selection on body weight upwards and downwards on both rich and poor larval food. Selection on increased or decreased sensitivity of body weight was also successful in at least one direction. Sensitivity was reduced by selection upwards in a poor environment and downwards in a rich environment.     

Correlated Responses to Selection for Postweaning Gain in the Rat

Evidence for correlated responses to selection was investigated in lines of rats selected for 13 generations for high (U line) and low (D line) 3-9-week gain in comparison with random-bred control lines (R and C lines). The increase in 3-9-week gain in the U lines was shown to be due largely to an increase in 9-week weight, although 3-week weight also increased in these lines. In the D lines, where a marked decrease in 3-9-week gain was observed, this was found to be due to a large decrease in 9-week weight and no detectable change in 3-week weight. The average 2-week litter weight, a measure of the lactational performance of the dam, was significanly greater in the U lines than in the D lines. Selection for 3-9-week gain in these lines of rats led to changes of litter size at birth in the same direction as that of selection. This resulted in a significantly higher litter size in the U lines than in the D lines. The number of rats alive 2 and 9 weeks of age and the percentage of mated females pupping were similar in the U and D lines but lower in these lines than the random bred C lines, providing evidence for a reduction of "fitness" in the selected lines. Carcass composition was studied for all lines at the 11th generation of selection. Carcass composition, in terms of water, fat, ash and protein, was similar in the R and C lines. The U lines had more water and lesss fat than the R or C line. The D lines had similar carcass composition to the R and C lines. It is suggested that these selected and random-bred lines of rats are potentially useful animals to investigate further the developmental and physiological mechanisms which control growth.     

Direct responses in males and correlated responses for reproduction in females to selection for testicular size adjusted for body weight in young male lambs

Selection based upon testicular diameter adjusted for body weight at 6, 10 and 14 weeks of age was used to produce two lines of sheep, with either high or low testicular size. Ten generations of selection were carried out and the estimate of the realized heritability of the selection criterion was 0.53 +/- 0.01. There were significant positive correlated responses to selection for testicular diameter at 6, 10 and 14 weeks of age, but the correlated responses in body weight at these ages were negative. In mature females, there were significant negative correlated responses to selection in premating body weight in the 1st, 2nd and 3rd breeding season and in the day of the first oestrus in the 2nd breeding season. Litter size per ewe mated had a small positive correlated response to selection in the second breeding season. This latter response appeared to be due to a positive correlated response in fertility, ewes from the High-line having a significantly higher probability of conceiving to a single mating than those from the Low-line. There was no significant correlated response in ovulation rate or litter size per ewe lambing and the genetic correlation between these traits and the selection criterion is likely to be close to zero. This may be due to the adjustment for body weight used, but it is possible that, in any event, body weight in young rams may be a better predictor of female ovulation rate than testicular diameter. These results do not rule out the possibility that testicular size in rams older than those selected would provide a good predictor of genetic merit for female ovulation rate.