Ovarian follicular dynamics in selected and control (relaxed selection) male- and female-lines of broiler breeders fed ad libitum or on restricted allocations of food

1. The reproductive structures of 4 broiler breeder lines fed ad libitum or restricted were compared at 1st egg. The lines were contemporary male- and female-lines and their respective progenitor lines maintained without selection for 20 years. 2. The selected lines were larger and had more hierarchical yellow follicles than the relaxed lines. The numbers of positions in the hierarchy were greater in the selected lines. The number of atretic follicles and the proportion of multiple follicles were relatively high in the selected male-line. 3. There were more white follicles in the range 1.8 to 5.0 mm diameter in the selected male-line than in the relaxed line, whereas selection did not change white follicle numbers in the female-line. 4. Food restriction decreased the numbers of normal and atretic yellow follicles and the proportion of multiples but had no effect on the number of positions in the hierarchy. Restricted feeding decreased the number of white follicles 1.4 to 2.4 mm and increased follicles 2.4 to 5.0 mm diameter. 5. There was little change in age at 1st egg among lines fed ad libitum. Restricted feeding increased age at 1st egg and the difference was greater in female- than in male-lines and in relaxed compared with selected birds. 6. Abdominal fatness was similar among the lines fed ad libitum and was decreased by restricted feeding. 7. Oviduct weights were greater in selected than in relaxed lines and were decreased by restricted feeding only in the selected male-line. 8. The weight of the largest follicle was higher in the selected lines and in restricted compared with ad libitum-fed birds. Plasma triglyceride concentrations were lower in restricted birds from the selected and relaxed female-lines compared with the other groups. Differences in plasma oestrogen concentrations could not be detected.     

Control of the development of the ovarian follicular hierarchy in broiler breeder pullets by food restriction during rearing

1. The increase in the numbers of yellow and of white ovarian follicles was studied from 10 weeks of age to sexual maturity in broiler breeder pullets fed ad libitum and on a restricted intake. The growth of follicles greater than 1 mm diameter commenced at 14 to 16 weeks of age in ad libitum birds and at 22 to 24 weeks in those which were restricted. 2. In a second experiment, groups of birds restricted from 4 d of age were transferred to ad libitum feeding at 2-week intervals and examined at first egg. The numbers of yellow follicles at first egg were not affected by restriction to 14 weeks of age but, thereafter, declined linearly with time on restriction. 3. In the second experiment there were no comparable responses in the numbers of white follicles or in the rates of atresia. 4. Food restriction to 22 weeks of age produced about half the response in numbers of yellow follicles as restriction continued to point of lay (27 to 32 weeks of age). In practical situations, the results suggest that food intake should be restricted up to the onset of lay and then increased in relation to the rise in egg production. 5. Atresia was greatest in white follicles 3.15 to 6.30 mm diameter and was less outside this range.     

Effects of ad libitum feeding on performance of different strains of broiler breeders

(1) Tolerance to ad libitum feeding was compared in three genotypes of broiler breeder hens: a standard broiler breeder fed ad libitum (SA) or restricted (SR), a slow growing 'label' broiler breeder (L) and an experimental dwarf heavy broiler breeder (E). Two similar experiments were conducted in two distinct research centres. (2) Feed intake and body weight were measured every 3 weeks from hatch to 40 to 49 weeks of age. Egg production and egg abnormalities were recorded. The number of yellow follicles in ovaries was counted at the age of 32 weeks. (3) Body weight was stabilised at 2.2, 3.7 and 5.4 kg after 24 weeks of age in L, E and SA hens, respectively. Growth of the SR hens was similar to that of L up to 20 weeks and stabilised at a similar level to that of E hens after 30 weeks of age. (4) Sexual maturity was delayed by 6 weeks in restricted breeders compared to ad libitum fed hens that started to lay at 20 weeks. SA hens had low egg production and a high proportion of defective eggs, which was largely compensated for by feed restriction. However, productivity of SR hens remained lower than that of L breeders. (5) Compared to the low viability and reproductive fitness observed with SA hens, the E dwarf broiler breeder tolerated ad libitum feeding and had better egg production, fewer egg abnormalities and yellow follicles per ovary and a higher egg production. However, laying rate was still lower than that of the SR and L groups. Energy conversion (kJ/g egg) from 32 to 40 weeks of age was much higher in the SA group than in the other three groups. 6. The feasibility of feeding a dwarf broiler breeder ad libitum calls for further research on implications of specific IGF and GH-receptor expression at the level of the ovary in dw chickens.     

Plasma oestrogen and triglyceride concentrations and reproductive characteristics of broiler chickens after ten generations of selection at seven weeks of age for high or low plasma very low density lipoprotein concentration.

1. Day old female chicks from two lines of broilers, selected for high or low plasma concentrations of very low density lipoproteins, were fed ad libitum or restricted until point of lay. All birds were fed ad libitum after laying an egg, and aspects of growth and reproduction were measured to 40 weeks of age. 2. Compared with lean line birds the growth rate of fat line females fed ad libitum declined from 16 weeks of age. 3. Mortality from apparently stress-related causes in ad libitum fed fat line females increased from 16 weeks of age and was 71% overall, compared with 26% in ad libitum fed lean line females. Mortality in the corresponding restricted birds was 20% and 7%, respectively. 4. Few fat line females fed ad libitum commenced lay and those which were restricted during rearing stopped laying after the initiation of ad libitum feeding. Egg productions in lean line birds fed ad libitum or restricted were similar. 5. Fat line females fed ad libitum had few yellow follicles and a high rate of atresia at first egg and between 37 and 39 weeks, as did restricted fat line birds between 37 and 39 weeks of age after being fed ad libitum. 6. Plasma concentrations of oestradiol-17 beta were 5-fold higher in fat compared with lean line females at 35 weeks of age. There was a similar large difference in concentrations of plasma triglyceride which reached 59 mg/ml in fat line females fed ad libitum.     

Changes in ovarian function and egg production in commercial broiler breeders through 40 weeks of lay

1. Egg production, body weight, feed intake and mortality were recorded in over 78?000 broiler breeder hens in 4 commercial flocks housed in 16 houses from early lay though 40 weeks of lay. A total of 420 hens were sampled at regular intervals throughout the laying period to determine the changes in body weight and the numbers of yellow follicles, paired follicles and hierarchical positions with time in the ovary throughout lay to relate ovarian function to productivity. Average egg weight was recorded weekly from one flock.

2. A quadratic equation fitted the changes in time for the number of yellow follicles, body weight, feed intake and mortality; a linear equation described the decline in the number positions in the hierarchy and there was a linear decline in the logistic scale of the proportions of yellow follicles developing as pairs of similar weight. Egg production was described by a cubic equation and egg weight by a line plus exponential model.

3. The average number of yellow follicles declined from 7·2 to 5·4 and the number of hierarchical positions from 6 to 5 from 4 to 40 weeks after photostimulation. The proportion of follicles developing as pairs of similar weight was over 25% at the onset of lay and declined to less than 10% from 20 weeks after photostimulation, representing a substantial loss of potential productivity.

4. Body weight and egg production were similar to the breeder's targets whereas average egg weight and mortality were higher than expected.

5. The relationship defining ovarian function will facilitate the development of an improved model of egg production in broiler breeder hens.     

Ovarian follicular structure of White Leghorns fed ad libitum and dwarf and normal broiler breeders fed ad libitum or restricted until point of lay

1. Yellow follicle numbers when the first egg was laid were 6.3, 9.0 and 12.4 for ad libitum reared Leghorns, sex-linked Dwarf (AL Dwarf) and Normal broiler breeders respectively. Multiple ovulation resulted in low egg production and a high proportion of defective egg shells in AL Dwarf and Normal broiler breeders during early lay. 2. Restricting the growth of broiler breeders during rearing by restricting food intake (R Dwarfs) reduced the number of yellow follicles to 6.8 and increased the rate of lay. 3. Atretic yellow follicles were common among broiler breeders but not in Leghorns or R Dwarfs when the first egg was laid. 4. Poor egg production in older broiler breeders was caused by birds with few or no developing yellow follicles, atresia in yellow follicles and the continued occurrence of multiple ovulations. 5. There was no relationship between the number of yellow follicles and the number of white follicles less than 5 mm diameter but atresia in white follicles 2 to less than 5 mm was negatively related to the number of yellow follicles. 7. No relationship was observed between abdominal fat weight and yellow follicle number, though birds which ate more had more yellow follicles. 8.The sex-linked dwarfing gene dw was associated with increased atresia among the white follicles and low numbers of yellow follicles compared with the Normal DW broiler breeder genotype.     

Role of body weight and food intake after photostimulation on ovarian function at first egg in broiler breeder females

1. The role of body weight during rearing and food allocation after photostimulation on ovarian function in broiler breeder females was examined in two experiments. In experiment 1 birds were fed ad libitum, or to grow to 0.7 or 0.4 of ad libitum‐fed body weight at 20 weeks of age. After photostimulation at 16, 19 and 22 weeks respectively they were fed ad libitum, 145 g/d or 115 g/d in a factorial design. The ovary of each bird was examined after it had laid an egg. In experiment 2, restricted birds (0.4 of ad libitum‐fed body weight) were fed 125 g/d after laying an egg or ad libitum and a third group were given 125 g/d for 3 to 4 weeks and then fed ad libitum. The ovaries of these birds were examined 6 to 7 weeks after their first egg.

2. Food restriction after photostimulation had little effect on ovarian function in birds fed ad libitum during rearing. These birds had 2 to 3 hierarchies of yellow follicles of similar size. Restricted birds fed ad libitum after photostimulation had a large increase in the numbers of yellow follicles (1.7 to 2.0 hierarchies) compared with birds fed 145 g/d or 115 g/d which had 1.2 to 1.6 hierarchies of yellow follicles.

3. Age at first egg was largely a function of rearing treatment but feeding 115 g/d to the 0.4 treatment birds delayed the onset of lay compared with the other treatments.

4. Ad libitum feeding in restricted birds after the onset of lay resulted in a similar increase in the number of yellow follicles as the response to ad libitum feeding after photostimulation.

5. The prevalence of birds with atretic follicles was relatively high in birds fed ad libitum during rearing and in restricted birds fed ad libitum after photostimulation.

6. Yolk weight increased with age at photostimulation and food restriction decreased yolk weight by about 1 g compared with ad libitum feeding after photostimulation.     

Assessment of the effects of separate sex feeding on the welfare and productivity of broiler breeder females

1. The welfare and productivity of female broiler breeders were studied in pens containing 15 females and 1 male where the sexes were fed together or separately by means of a grid with 40 mm spaces over the female feeders. 2. Average head width did not increase after 25 weeks of age. 3. The rate of lay was lower in separate compared with mixed-sex-fed pens but egg weight was similar. 4. The incidence of swollen heads and beak lesions was higher and feather loss and rate of food intake was lower in separate compared with mixed-sex-fed pens. 5. It was concluded that the grid spaces should be 42 mm wide to avoid adverse effects on welfare and productivity.     

Changes in ovarian function of female turkeys photostimulated at 18, 24 or 30 weeks of age and fed ad libitum or restricted until point of lay.

1. Females from a medium body weight strain of turkeys were fed ad libitum or restricted to 0.6 of ad libitum-fed body weight at sexual maturity and were photostimulated at 18, 24 or 30 weeks of age. Ovarian follicular structure was assessed in 8 birds from both rearing treatments at photostimulation, after photostimulation for 6 or 12 weeks and at 55 weeks of age. 2. The patterns of small white follicles and of larger yellow follicles were similar in restricted and in ad libitum-fed turkeys for birds in lay at the same age, regardless of the time elapsed since photostimulation. 3. Among birds in lay the number of normal yellow follicles and the proportion of multiple yellow follicles declined with age. A relatively low proportion of restricted birds photostimulated at 18 and 24 weeks were in lay after 12 weeks of photostimulation. 4. For birds in lay the number of white follicles 1.4 to less than 1.8 mm in diameter increased and the number greater than 1.8 mm declined with age. There was a corresponding increase in the rates of atresia in follicles 1.8 to less than 5.0 mm in diameter.     

Bone mineral density QTL at sexual maturity and end of lay

1. An F₂ cross of a broiler male line and a White Leghorn layer line was used to identify quantitative trait loci (QTL) for bone density at the onset of lay and at the end of the laying period. A total of 686 measures of humeral bone density were available for analysis.

2. There was no evidence for epistasis.

3. Genome-wide significant QTL for bone density at the onset of lay were identified on chromosomes 1 (311?cM) and 8 (2?cM) and on chromosomes 1 (311?cM), 3 (57?cM) and 8 (2?cM) with a covariate for the number of yellow follicles (a proxy for the concentration of circulating oestrogen).

4. Evidence for only 4 chromosome-wide suggestive QTL were detected at the end of lay (72 weeks).

5. Analysis of the combined data confirmed two genome-wide suggestive QTL on chromosome 1 (137 and 266?cM) and on chromosomes 8 (2?cM) and 9 (10?cM) in analyses with or without the covariate.

6. Positive QTL alleles came from the broiler line with the exception of 2 suggestive QTL at the onset of lay on chromosomes 3 and 5 in an analysis with the covariate.

7. In general, QTL acted additively, except that dominant effects were identified for three suggestive QTL at the onset of lay on chromosomes 3 (57 and 187?cM) and 5 (9?cM).

8. The significant QTL in this study were at similar locations to QTL identified in a range of crosses in other publications, suggesting that they are prime candidates for the search for genes and mutations that could be used as selection criteria to improve bone strength and decrease fractures in commercial laying hens.     

Effects of age and of early lighting on ovarian function in breeding turkeys

1. Yellow follicle numbers and production of defective egg shells were greater in the early part of the laying period in conventionally-reared turkeys photostimulated at 24 rather than 30 weeks of age. 2. The number of yellow follicles declined with age and were similar in both groups at 40 weeks of age. 3. At 55 weeks of age more birds photostimulated at 24 weeks were out of lay and birds in lay had fewer yellow follicles and more atresia compared with turkeys photostimulated at 30 weeks of age. 4. The numbers of white follicles in the 1 to less than 2 mm size were similar in the two groups at each age. The proportion of white follicles which were atretic was negatively related to the number of yellow follicles, particularly in follicles 5 to less than 8 mm diameter.     

Comparison of the effects of the degree of food restriction during rearing on ovarian function at the onset of lay in unselected ducks (anas platyrhynchos) and in a line selected for improved food efficiency

1. Ovarian structure and function at sexual maturity in two lines of ducks (selected or not selected for juvenile food efficiency), fed ad libitum or restricted to achieve 0.50, 0.65 or 0.80 of unrestricted body weight at 24 weeks, were compared in a randomised block factorial experiment.

2. Selected ducks were larger but not fatter or older at sexual maturity, had more yellow follicles, giving a greater potential for multiple ovulation, and a higher proportion of atretic yellow follicles, compared with unselected ducks.

3. There was a linear relationship between the number of normal yellow follicles and body weight at the onset of lay which was similar in the two lines.

4. Several birds on the most severe regimen failed to lay and it was concluded that ducks should be at least 0–6 of unrestricted body weight for the onset of lay.

5. Weight of the ovary with follicles less than 8 mm diameter was greater in selected compared with unselected ducks and in birds fed ad libitum compared with restricted ones.     

Ovarian follicular dynamics in lines of sheep (Finn, Merinos) selected on ovulation rate

Ewes from selected lines of sheep from each of two breeds, Finns (high ovulation rate, low ovulation rate and control lines with respective ovulation rates of 5.4, 2.7 and 3.3) and Merinos (T Merinos selected for increased ovulation rate and control Merinos with respective ovulation rates of 1.9 and 1.2) were used to examine how selection to alter ovulation rate had altered follicle development. Ovarian antral follicles were counted, measured, classified as nonatretic or atretic (more than five pyknotic bodies). The growth of ovulatory follicles in vivo, followed by repeated follicle ink marking, also was compared in the three lines of Finns. Regardless of breed, ewes selected for high ovulation rate had a similar number of antral follicles and a similar extent of atresia compared with their controls. Alterations induced by selection were located in the last stages of folliculogenesis. T Merinos exhibited a lower proportion of atretic follicles among follicles greater than 3 mm and a larger diameter of the largest healthy follicle when preovulatory follicles were excluded. High-line Finn ewes recruited more follicles, which produced smaller preovulatory follicles, each containing a smaller number of granulosa cells compared with either the low- or control-line ewes. Hence, physiological selection for high ovulation rate raised it by different methods in Merino than in Finn ewes.     

Effect of dietary zinc proteinate supplementation on growth performance, and skin and meat quality of male and female broiler chicks

1. An experiment was conducted to evaluate the effect of dietary zinc proteinate (ZP) supplementation on growth performance and on skin and meat quality of male and female broiler chicks.

2. A total of 240 1-d-old male and 240 1-d-old female broiler chicks were randomly distributed into 24 floor pens (12 replicate pens/sex; 20 birds/pen) and were given either 0 (Control diet) or 40?mg/kg ZP (ZP 40), resulting in a 2?×?2 factorial arrangement of treatments.

3. The growth performance of male and female broiler chicks was not affected by the ZP supplementation, but the males showed significantly higher growth performance than did females.

4. ZP supplementation increased the total thickness of skin in both sexes, and males had thicker skin than females. It also increased the collagen content of skin, but not that of meat. Males had higher skin collagen contents than did females, but no sex difference was found in the meat collagen contents.

5. ZP supplementation did not affect the shear force values of skin and meat; however, males had higher shear force values of back skin than females. ZP supplementation increased the zinc contents of thigh meat and plasma in both sexes. Males had higher zinc contents in back skin than females.

6. It is concluded that dietary ZP supplementation could increase the skin quality of broiler chicks in both sexes, particularly in female broilers, without any effect on growth performance. Male broilers have better growth performance and skin quality than females.     

Welfare assessment of modified rearing programmes for broiler breeders

1. Female broiler breeders were fed ad libitum or a restricted quantity of food to achieve either a recommended body weight curve or a modified (linear) growth curve that allowed more generous feeding between 6 and 15 weeks of age. The birds were fed a ration containing either a normal or low concentration of crude protein. The welfare of the birds was assessed using a profile of indices of welfare at 6, 12, 18 and 24 weeks of age. 2. The low-protein ration decreased the body weight of birds fed ad libitum and restricted birds were fed substantially more of the low-protein ration to meet target body weights from 3 to 14 weeks of age. 3. The modified restricted rearing programme did not have an effect on indices of welfare. 4. Water intakes and plasma corticosterone concentrations were lower in restricted birds fed the low-protein ration and they spent more time resting, and less time spot-pecking, than birds fed the high-protein ration. 5. Birds fed ad libitum on both rations spent more time resting and less time foraging, drinking and spot-pecking than food restricted birds and were more fearful than restricted birds at 6, 12 and 18 weeks of age. 6. The heterophil:lymphocyte ratio in restricted birds was higher at 6 weeks and lower at 24 weeks compared with birds fed ad libitum. Humoral immunity was lower at 6 weeks of age in birds fed ad libitum and was similar at other ages. Cell-mediated immunity was similar among all treatments. 7. Plasma corticosterone concentrations were higher in restricted birds compared with birds fed ad libitum. Plasma creatine kinase activity was higher in birds fed ad libitum at 6, 12 and 18 weeks of age. Alkaline phosphatase and lactate dehydrogenase activity were higher whereas AST was lower in restricted birds compared with those fed ad libitum. 8. There was no evidence to support the use of low protein rations or linear growth curves to improve welfare in restricted broiler breeder females.     

Male to female ratios for broiler‐type and egg production‐type breeders 1

Two experiments using commercial broiler‐type breeders and one experiment using commercial egg production‐type breeders were conducted to determine the effect of male to female ratios on fertility and hatchability. Breeding pens were made up with 36 females and 4, 3, 2 or 1 male(s) per pen giving ratios of one male to 9, 12, 18 and 36 females. Five replicate pens were used per treatment.

Within the ratios and genetic strains used, maximum fertility was obtained with male to female ratios of 1 to 12 with broiler‐type breeders and 1 to 18 with egg production‐type breeders. It is concluded that using fewer females per male than in these ratios would be unwarranted and in some cases more females per male would be feasible.

Hatchability of fertile eggs was not affected by male to female ratios.     

Comparative development of the ovary and production,fertility and hatchability of eggs from traditional turkeys and a contemporary male-line fed ad libitum or restricted

1. Ovarian function was assessed at first egg and at 44 and 54 weeks of age in traditional turkeys and in a male-line fed ad libitum or restricted to 0.5 of ad libitum during rearing. Egg production was recorded daily and fertility and hatchability were determined at 2 ages. 2. Age at sexual maturity did not differ between the 3 treatments. 3. The number of follicles at first egg in the male line averaged 16.3 compared with 8.9 in traditional turkeys. Food restriction in the male line decreased the number of yellow follicles to 14.6 at first egg and restricted birds were largely in a non-reproductive state from 44 to 54 weeks of age. 4. The total number of recorded ovipositions to 54 weeks of age in male line fed ad libitum or restricted and traditional turkeys respectively were 40.1, 25.2 and 70.7; comparable productions of normal shelled eggs were 28, 20 and 66.3. 5. There were no differences between treatments in fertility after a single insemination at 2 different ages. Hatch of fertiles after the first inseminations were 71.4, 45.5 and 90.3 respectively for male line fed ad libitum or restricted and traditional turkeys.     

Similar improvements in reproductive performance of male line, female line and parent stock broiler breeders genetically selected in the UK or in South America

1. Field data on the reproductive performance of parent stock and grandparent female line and male line broiler breeders were extracted from a commercial database covering the years of hatch from 1989 to 2002. The data were from the UK, Western Europe, Central and South America and Japan. Quadratic regression curves were fitted to the data to estimate the long-term phenotypic trends in total egg production, hatching egg production, chick numbers, hatchability and female mortality. 2. Changes in performance over time were similar in the UK, Western Europe and Central and South America and are consistent with the suggestion that genetic selection in separate breeding programmes in the UK (supplying the UK and Western Europe) and Brazil (serving Central and South America) was associated with similar improvements in the performance of commercial parent stock and grandparent flocks. Changes in performance in Japan (supplied from the UK) were different but recent performance was similar to those in the other regions. 3. The mean annual improvements over 10 years to 2002 for male line, female line and parent stock, respectively, were 0.4, 1.7 and 1.7 for total egg number; 0.5, 1.8 and 1.6 for hatching egg number; 0.6, 1.8 and 1.3 for total chick production; 0.25, 0.47 and 0.09% for hatchability; and -0.69, -0.05 and -0.41% for female mortality. 4. The results are consistent with claims that significant genetic change in pedigree selection (nucleus) flocks by the primary breeding company are reflected in substantial improvements in reproductive performance in commercial flocks. The presence of similar rates of improvement in different geographical regions as a result of genetic selection in two continents representing tropical and temperate regions suggests that genotype-environment interactions for reproduction traits are not important.     

Cell Proliferation in the Atretic Follicles of Buffalo and Cattle Ovary

The present study was carried out to describe the proliferative activity of granulosa and theca cells in healthy antral and atretic follicles of Philippine buffaloes (BU) and Holstein-Friesian (HF) cows. Paraffin sections of ovary were immunostained with mouse monoclonal antibody to proliferating cell nuclear antigen (PCNA). Then the follicles were classified into healthy and various stages of atretic follicles. The granulosa layer of healthy follicles had a significantly higher frequency of PCNA-positive cells than the early and advanced atretic follicles in both breeds. In the theca interna, significantly reduced populations of the PCNA-positive cells were found in both breeds as atresia progressed. Moreover, HF had significantly higher PCNA-positive cells in the theca interna of healthy, early atretic and advanced atretic follicles than BU. A reduction of PCNA-positive cells during atresia was also noted in the theca externa in both animals although differences were not significant. The results of the present work suggest that the proliferative activity of granulosa and theca cells decreases in association with follicular atresia in the BU similar to HF. Furthermore, a significantly deficient cell proliferative activity of theca interna was found in BU compared with HF.     

Modification of intra‐ovarian follicular distributions in broiler breeder hens by ad libitum or restricted feeding

1. Ovarian follicles in broiler hens from 3 lines fed ad libitum or food restricted were examined at point of lay and from 2 of these lines at 47 weeks of age. The lines were Ross 1 birds and birds selected to be lean or fat on the basis of their plasma very low density lipoprotein concentrations at 7 weeks of age.

2. Follicles of diameter greater than 1.4 mm were classified into size groups, and the numbers in these groups in the anterior and posterior halves of the ovary were compared for the effects of line and feeding régimen.

3. There were more yellow follicles ( > 8 mm diameter) in the anterior half of the ovary, and this difference was greater for ad libitum birds. Ad libitum feeding did not influence the number of yellow follicles in the posterior segment of the ovary.

4. The increase in numbers of yellow follicles in the anterior segment of the ovary differed according to line at point of lay, but not at 47 weeks of age.

5. Birds of 47 weeks of age showed higher numbers of follicles 1.4–6 mm in diameter in the posterior segment of the ovary, and smaller numbers of follicles 6–8 mm in diameter. Thus, although large numbers of white follicles were in the earlier stages of growth, there were fewer ovulable yellow follicles because of higher levels of atresia.