Biomedical subjects

P M Hocking

Publications and source records attributed to P M Hocking.

At least 37 records · Page 2Linked to original sources

PubMed2003

Quantitative trait locus detection in commercial broiler lines using candidate regions.

A QTL that explained a large proportion of the phenotypic difference between broiler and layer chickens in an experimental cross was evaluated in a commercial broiler line. A three-generation design, consisting of 15 grandsires, 608 half-sib hens, and more than 50,000 third-generation offspring, was implemented within the existing breeding scheme of a broiler breeding company. Four markers from a candidate region on chicken chromosome 4 were selected for their informativeness in the grandsires and used to genotype the first two generations. Using half-sib analyses, linkage was studied between these markers and 13 growth and carcass traits. The QTL analyses confirmed the presence of significant QTL for body weight (P < 0.01) and residual feed intake (P < 0.05) on chicken chromosome 4. Furthermore, evidence was found for QTL affecting the relative weight of bone and muscle in the thigh. Four more markers were added to increase resolution of the QTL positions. This increased the significance of the QTL for body weight (P < 0.001) and residual feed intake (P < 0.01) and showed evidence (P < 0.05) for additional QTL affecting carcass weight and conformation score. This study showed for the first time that a QTL that explains differences between broilers and layers was segregating in lines that have been selected for body weight over 50 generations. A possible explanation could be a pleiotropic or closely linked effect on fitness-related traits that are not part of the present study. The results demonstrate the feasibility of QTL detection and the potential for marker-assisted selection within a commercial broiler line without altering the existing breeding scheme.

Animals↗

PubMed2002

Quantitative trait loci affecting fatness in the chicken.

An F2 chicken population of 442 individuals from 30 families, obtained by crossing a broiler line with a layer line, was used for detecting and mapping Quantitative Trait Loci (QTL) affecting abdominal fat weight, skin fat weight and fat distribution. Within-family regression analyses using 102 microsatellite markers in 27 linkage groups were carried out with genome-wide significance thresholds. The QTL for abdominal fat weight were found on chromosomes 3, 7, 15 and 28; abdominal fat weight adjusted for carcass weight on chromosomes 1, 5, 7 and 28; skin and subcutaneous fat on chromosomes 3, 7 and 13; skin fat weight adjusted for carcass weight on chromosomes 3 and 28; and skin fat weight adjusted for abdominal fat weight on chromosomes 5, 7 and 15. Interactions of the QTL with sex or family were unimportant and, for each trait, there was no evidence for imprinting or of multiple QTL on any chromosome. Significant dominance effects were obtained for all but one of the significant locations for QTL affecting the weight of abdominal fat, none for skin fat and one of the three QTL affecting fat distribution. The magnitude of each QTL ranged from 3.0 to 5.2% of the residual phenotypic variation or 0.2-0.8 phenotypic standard deviations. The largest additive QTL (on chromosome 7) accounted for more than 20% of the mean weight of abdominal fat. Significant positive and negative QTL were identified from both lines.

Abdomen↗

PubMed2002

Role of dietary calcium and available phosphorus in the aetiology of tibial dyschondroplasia in growing turkeys.

1. Large White male turkeys from a heavy commercial male-line were fed 16 diets containing 4 concentrations of calcium (6, 10, 14 and 18 g/kg) and available phosphorus (3, 5, 7 and 9 g/kg) in a 4 x 4 factorial experiment. There were three replicates (pens) of each treatment and the skeletal health, morphology and mineral status of 4 turkeys from each pen were assessed at 7, 10 and 13 weeks of age. 2. The prevalence of tibial dyschondroplasia increased after 7 weeks of age and was present in 50 and 71% of turkeys respectively at 10 and 13 weeks. The lesion was localised in the caudal aspect of the proximal tibiae. Dietary calcium and available phosphorus did not affect the prevalence of the lesion except in turkeys on the diet containing 6 g calcium/kg, where body weight and the incidence of tibial dyschondroplasia were low. 3. Histological investigation showed no evidence of rachitic changes. 4. Low dietary calcium was associated with lower tibial plateau angles at 10 and 13 weeks of age. Tibial torsion and the angle of rotation were not affected by dietary treatments or age. Tibial torsion and the angle of rotation were not affected by dietary treatments or age. 5. Increasing dietary calcium increased tibial radiodensity, cortical density and the widths of the cortex and proximal tibiotarsus. Radiodensities increased to 10 weeks and were significantly lower at 13 weeks of age. 6. Bone ash, calcium and phosphorus declined with age, particularly between 10 and 13 weeks, whereas bone calcium: phosphorus ratios were not affected by dietary treatment or age. 7. Dietary calcium was positively associated with blood calcium and calcium ion concentrations and was without effect on blood phosphorus. Available phosphorus was associated positively with increased blood phosphorus and lower calcium ion concentrations but had no effect on total calcium. Alkaline phosphatase activity was low at high concentrations of dietary calcium with low available phosphorus and there was higher activity on diets containing low calcium and high available phosphorus.

Age Factors↗

PubMed2002

Welfare assessment of broiler breeders that are food restricted after peak rate of lay.

1. Broiler breeder females were reared on one of three growth curves (ad libitum, conventional or modified restriction) and given rations containing a high or low concentration of crude protein. After the peak rate of lay they were fed ad libitum or a decreasing quantity of food in response to declining egg production. The welfare of the birds was assessed by determining the changes in indices of welfare at 36, 48 and 60 weeks of age. 2. Body weight increased rapidly in restricted birds fed ad libitum post-peak and water intake declined. 3. Post-peak food restriction was associated with a decrease in resting and increased drinking and spot-pecking activities. Birds that were food restricted during rearing spent more time foraging and spot-pecking at 36 and 48 but not 60 weeks of age. 4. Immune function increased with age but was not affected by the experimental treatments. The heterophil-lymphocyte ratio in birds fed ad libitum during rearing was numerically lower at 36 and higher at 48 and 60 weeks of age compared with restricted birds. 5. There was no effect of treatment on plasma corticosterone concentration. Creatine kinase activity was high at 60 weeks in treatments that were characterised by poor reproductive status and the activities of other enzymes reflected differences in reproductive status and mortality. 6. There was no long-term welfare or production advantage from feeding low protein rations or more generous feeding during the rearing period compared with conventional food restriction programmes.

Animal Husbandry↗

PubMed2002

Effects of low dietary protein and different allocations of food during rearing and restricted feeding after peak rate of lay on egg production, fertility and hatchability in female broiler breeders.

1. A 3x2x2 factorial experiment was conducted with boiler breeder females to determine the effects of body weight gain (three levels) and conventional or lower crude protein concentrations in the rations during rearing, and restricted or ad libitum feeding after the peak rate of lay on egg production, fertility, hatchability, mortality and food consumption. 2. Compared with ad libitum feeding, conventional food restriction resulted in a decrease in average daily food consumption of 0.6 during rearing, 0.2 during early lay and an increase of 0.4 after the peak rate of egg production. Mortality was decreased by more than half. 3. Restricted birds had higher total and settable egg production, fewer defective or damaged eggshells and higher fertility and hatchability than those fed ad libitum. The modified (more generous) rearing programme resulted in lower rates of egg production and higher rates of mortality compared with the conventional food restriction programme. 4. Low-protein rearing rations were associated with higher rates of food intake, higher mortalities and lower rates of egg production than the conventional protein rations. There were no differences in the fertility or hatchability of eggs between birds fed on the two concentrations of dietary crude protein. 5. Ad libitum feeding post-peak was associated with higher rates of mortality to 60 weeks of age. Post-peak feeding had little effect on the rate of lay or egg weight in conventionally restricted birds fed high protein rations or in birds fed ad libitum. Restricted feeding post-peak decreased the rate of lay and egg weight in birds on the modified restriction programme and in conventionally restricted birds fed on the low protein rations. There was no effect of post-peak feeding on fertility or hatchability of eggs. 6. The total numbers of saleable chicks per kg food consumed were 1.83, 1.72 and 0.52 for conventional, modified and ad libitum feeding during rearing; 1.56 and 1.15 for restricted and ad libitum feeding post-peak: there were no differences associated with protein concentrations of rations fed during rearing.

Animal Welfare↗

PubMed2002

Effects of dietary calcium and phosphorus on mineral retention, growth, feed efficiency and walking ability in growing turkeys.

1. Large White male turkeys from a heavy commercial male-line were fed on 16 diets containing 4 concentrations of calcium (6, 10, 14 and 18 g/kg) and available phosphorus (3, 5, 7, 9 g/kg) in a 4 x 4 factorial experiment with three replicates (pens). Turkeys were weighed and food intakes recorded from 4 to 7, 8 to 10 and 11 to 13 weeks of age. 2. The diets containing 6 g/kg calcium and 5, 7 or 9 g/kg available phosphorus concentrations were associated with lower body weighs at 10 and 13 weeks of age. 3. Fewer than 6% of the turkeys had an abnormal gait at 13 weeks of age. 4. The optimum dietary concentrations were 10 g/kg of calcium and 3 g/kg of available phosphorus. It was concluded that these concentrations should be fed at least to 13 weeks of age. 5. Retentions of dietary calcium and phosphorus averaged 300 g/kg of intake. 6. Utilisation of dietary phytate ranged from 5 to 11 g/kg and it is recommended that organic phosphorus should be ignored in the formulation of diets for growing turkeys unless they are supplemented with a phytase enzyme.

6-Phytase↗

PubMed2002

Mapping of quantitative trait loci for body weight at three, six, and nine weeks of age in a broiler layer cross.

An F2 chicken population was established from a cross of a broiler sire-line and an egg laying (White Leghorn) line. There were two males and two females from both lines in the base population. The F1 progeny consisted of 8 males and 32 females. Over 500 F2 offspring from five hatches were reared to slaughter at a live weight of 2 kg at 9 wk of age. Body weights at 3, 6, and 9 wk were recorded. The DNA was extracted from blood samples, and genotypes for 101 microsatellite markers were determined. Data of 466 individuals from 30 families were available for analysis. Interval mapping QTL analyses were carried out. The QTL significant at the genome wide level that affected body weight at two ages were identified on chromosomes 1, 2, 4, 7, and 8 and a QTL on Chromosome 13 influenced body weight at all three ages. Genetic effects were generally additive, and the broiler allele increased body weight in all cases. The effects for significant individual QTL accounted for between 0.2 and 1.0 phenotypic standard deviations and the sum of the additive effects accounted for approximately 0.75 of the line difference in body weight at 6 wk of age. The largest single additive effect was on chromosome 4, and the effect of substituting one copy of the gene was an increase in weight of 249 g. Interactions of the QTL with sex or family were unimportant. There was no evidence for imprinting or of two or more QTL at the same location for any of the traits.

Aging↗

PubMed2002

Comparative changes in plasma concentrations of progesterone, oestradiol and LH during the ovulatory cycle in a multiple ovulating male line and a single ovulating traditional line of turkeys.

The aim of this study was to compare the profile of circulating concentrations of LH, progesterone and oestradiol in a multiple ovulating male line with that of a single ovulating line of traditional turkeys. Plasma samples from seven traditional and 12 male-line turkeys were obtained every 3 h for 36 h. Male-line and traditional turkeys had single peaks of LH and progesterone that were of similar duration in both lines. The mean height of the plasma peaks of LH and progesterone were similar in the two lines and there was no detectable peak plasma oestrogen concentration. Mean plasma concentrations of LH and oestrogen were higher in single compared with multiple ovulating turkeys, whereas there were no differences in mean plasma progesterone concentrations. The results indicate that the multiple ovulation state in genetically selected high-growth lines of turkey may be the result of a correlated response in the steroidogenic capacity of ovarian tissue associated with low plasma concentrations of oestrogen rather than of a disturbance in the hormone profile of the ovulatory cycle.

Animals↗

PubMed2001

Effects of anti-inflammatory steroid drugs on pain coping behaviours in a model of articular pain in the domestic fowl.

The optimum dose rates of betamethasone, dexamethasone (sodium phosphate) and methylprednisolone (acetate salt) were determined for the treatment of inflammatory pain in domestic fowl using the microcrystalline sodium urate model of articular pain. The response criteria were the changes in behavioural profiles and pain-related activity over 60 minutes commencing 1 hour after an intra-articular injection of sodium urate and a course of intramuscular injections of a range of doses of each drug on the previous 2 days. The optimum dose rates for betamethasone, dexamethasone and methylprednisolone respectively were 0.04, 0.06 and 2 mg kg(-1)and are in the range of the recommended doses for these drugs in mammals.

Animals↗

PubMed2001

Effects of ambient temperature and restricted feeding on the growth of feathers in growing turkeys.

1. Male turkeys were reared to 6 weeks of age at 15 degrees C and 26 degrees C and fed ad libitum or restricted to 0.5 of the body weight of birds fed ad libitum. Basal metabolic rate was determined by indirect calorimetry at an ambient temperature of 20 degrees C. 2. Turkeys at 15 degrees C were lighter than those kept at 26 degrees C. Feather lengths and weight were similar in both groups. Fasting heat production corrected for both metabolic body size and activity was greater in turkeys reared at 15 degrees C than those at 26 degrees C. 3. Cranial breast feathers were significantly longer in restricted birds than in those fed ad libitum in contrast to a proportional decrease in the lengths of other feathers of 0.1 to 0.3. Feather weight as a proportion of body weight was 0.072 in restricted turkeys compared with 0.046 in birds fed ad libitum. There was no difference in basal metabolic rate between ad libitum and restricted turkeys. 4. It was concluded that feather growth was maintained in preference to body and muscle growth and that rearing birds at 15 degrees C did not improve breast feather cover. It is suggested that the growth of breast feathers in turkeys fed ad libitum is impaired.

Animals↗

PubMed2001

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.

Aging↗

PubMed2001

Age-related changes in fear, sociality and pecking behaviours in two strains of laying hen.

1. Two lines of commercial hybrid layers (Tetra and ISA Brown) were reared from hatch to 30 weeks of age in groups of 8. The objectives of the experiment were to evaluate the significance of the different selection practices involved in the development of the lines and to assess the potential association between selected behavioural states and the potential for feather damage and cannibalism. 2. Behavioural tests related to fear (tonic immobility, novel object, open field), sociality (runway, proximity in the home pen) and pecking (feather bunch and focal observations of inter-bird and environmental pecking) were conducted at 0 to 2, 5 to 7, 12 to 14, 19 to 21 and 29 to 31 weeks of age. Scan sampling of general behaviour was also conducted at these ages. 3. Underlying sociality was greater in Tetras than in ISA Brown hens. 4. There were no apparent overall strain differences in fearfulness although ISA Brown hens showed significantly longer tonic immobility fear reactions than Tetras at 31 weeks of age. 5. ISA Brown hens gave and received more gentle pecks than Tetra hens whereas preening was commoner among Tetras. There were no strain differences in the number of pecks at litter or food. 6. As birds aged they showed less avoidance of novel objects and Tetras, though not ISA Browns, showed progressively shorter tonic immobility responses. Birds of both strains spent less time resting, more foraging (pecking and scratching) and pecked more at the feather bunch at older ages. 7. Measures taken in the TI, open field, runway (social affiliation) and feather bunch tests were stable over time whereas focal observations of pecking at hens and the environment were not. 8. Factor Analysis showed that 3 factors defined by the novel object test, environmental pecking and pecking ata feather bunch explained most of the variation in the correlation matrix between summary measures for the 8 behavioural traits.

Age Factors↗

PubMed2001

Acute heat stress-induced alterations in blood acid-base status and skeletal muscle membrane integrity in broiler chickens at two ages: implications for meat quality.

The effects of acute heat stress (AHS) on indices of respiratory thermoregulation and skeletal muscle damage (myopathy) were examined in broiler chickens at two ages (35 and 63 d of age); the relationships of these responses with changes in meat quality were assessed. Exposure to AHS significantly increased deep-body temperatures, panting-induced acid/base disturbances, and plasma creatine kinase (CK) activities, reflecting heat stress-induced myopathy (HSIM). The extent of the hyperthermia and disturbances in acid/base status and myopathy was significantly (P < 0.05) higher in the older birds. Consistent with AHS-induced alterations in thermoregulatory indices and muscle membrane integrity were changes in breast muscle glycolytic metabolism as indicated by lower muscle pH immediately postslaughter (pHi), increased water loss, and increased incidence of breast muscle hemorrhages. Values of pHi were lower and hemorrhage scores greater in the AHS birds at 63 d; drip losses were significantly higher in the 35-d-old birds. Exposure to AHS did not affect breast meat eating quality, although overall reductions in flavor attributes were observed in the older birds. We concluded that exposure to AHS induced disturbances in blood acid/base status and had a detrimental effect upon skeletal muscle membrane integrity. Muscle from broilers exhibited an increased sensitivity to AHS exposure with age. Alterations in antemortem blood acid/base status and muscle membrane integrity induced by AHS were associated (though not necessarily causally) with adverse effects upon breast meat quality. It is recommended that preslaughter exposure of broiler chickens to AHS should be avoided in order to reduce alterations in muscle metabolism and membrane integrity and undesirable meat characteristics.

Acid-Base Equilibrium↗

PubMed2001

Ovarian steroid hormone production in a multiple ovulating male line and a single ovulating traditional line of turkeys.

Ovarian production of oestradiol and progesterone was investigated in a single ovulating traditional line and a multiple ovulating male line of turkeys. The oestradiol output from small follicles in response to LH stimulation and the aromatase activity of the residual ovary were also compared. The multiple hierarchy of follicles in the male line was shown to consist of a series of follicles of a similar size and stage of physiological maturation. The follicles of the traditional line produced significantly greater quantities of oestradiol than those of the male line. Impaired oestradiol production may have resulted in the lower plasma oestradiol concentration observed in the male line. There was no difference in follicular progesterone output between the traditional-line and male-line turkeys. The oestradiol output of whole small white follicles in response to LH stimulation was similar in the two lines and the depressed steroidogenesis of the male line was not changed. The aromatase activity of the small follicles was also lower in the male line, and there was no evidence that the male line ovary produced oestradiol in quantities proportional to its size. The results demonstrated that the hierarchy of follicles in the male line consisted of groups of follicles of similar mass and hormone output and indicated that selection for increased meat yield may have resulted in reduced ovarian steroidogenesis in male-line turkeys, in comparison with traditional, unselected turkeys.

Animals↗

PubMed2000

Effects of food restriction or delayed photostimulation on ovarian follicle number, plasma oestradiol concentration and vaginal collagen content in male-line turkeys.

1. An experiment was conducted to test the hypothesis that severe food restriction throughout the onset of lay or delaying photostimulation by 5 weeks would decrease the number of ovarian hierarchical follicles in male-line turkeys. It was proposed that a decrease in the number of hierarchical follicles would lower plasma oestradiol concentration and be associated with higher vaginal collagen content and a lower propensity to prolapse of the oviduct. 2. Five weeks after photostimulation, neither restriction to 63% of ad libitum body weight nor delayed onset of photostimulation had any significant effect on follicle number, plasma oestradiol concentration or vaginal collagen content. 3. It was concluded that neither food restriction nor delayed photostimulation can be used to control the over-development of the ovarian hierarchy in male-line turkeys.

Animals↗

PubMed2000

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.

Adipose Tissue↗

PubMed2000

Effects of the age of male and female broiler breeders on sexual behaviour, fertility and hatchability of eggs.

1. The effects of the age of male and female broiler breeders on fertility, hatchability and sexual behaviour were determined in two 2x2 factorial experiments. In the 1st, male and female birds 27 to 29 and 35 to 37 weeks and in the 2nd at 35 to 37 and 55 to 57 weeks of age were compared. 2. Fertility was not affected by the age of the males or females. 3. Early embryo deaths were higher and hatchability of fertile eggs was lower in eggs from females aged 27 to 29 and 55 to 57 weeks compared with eggs from birds aged 35 to 37 weeks. 4. There were no differences in male display behaviours at any age. The number of observed matings were higher in young males 27 to 29 weeks old compared with males 35 to 37 and 55 to 57 weeks of age. There were no detectable differences in the responses of the females to the males at different ages. 5. Plasma testosterone concentration in males increased from 24 to 30 weeks of age and was relatively constant from 36 to 60 weeks. Plasma oestrogen in females was high at 24 and lowest at 30 weeks of age after which it increased. 6. The results suggest that there may be little difference in the fertility of male and female broiler breeders at different ages when body weight is adequately controlled to at least 60 weeks of age.

Age Factors↗

PubMed2000

Development of the reproductive system in turkeys with a high or low susceptibility to prolapse of the oviduct.

Lines of turkeys selected for rapid growth and high meat yield have an increased incidence of prolapse of the oviduct compared with unselected or traditional strains of turkeys. The development of the reproductive system and changes in plasma estrogen concentrations were compared in sire line and traditional turkeys with the aim of identifying any morphological or hormonal differences that could be associated with the high incidence of prolapse in the male line. Four turkeys from each strain were killed weekly from 0 to 7 wk postphotostimulation, and samples from prolapsed birds were obtained from field cases. There were no differences in the rate of development of the ovary, oviduct, uterus, vagina, sphincter ani muscle, or muscular cord of the ventral ligament between the two strains that could predispose the sire line to prolapse. Histological investigation of the uterus, vagina, muscular cord of the ventral ligament, and sphincter ani muscle 5 wk postphotostimulation in traditional, sire line, and prolapsed sire line turkeys did not reveal any differences that could be associated with prolapse. No prelay peak in plasma estradiol concentration was observed in either strain, and there was no evidence to suggest that plasma estradiol was higher in the sire line compared with the traditional turkeys. It was concluded that prolapse of the oviduct in sire line turkeys was not associated with any anatomical abnormalities or high plasma estradiol during reproductive development.

Anal Canal↗