Mating behavior in commercial broiler breeders: Female effects
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©2010 Poultry Science Association, Inc. Mating behavior in commercial broiler breeders: Female effects J. R. Moyle, D. E. Yoho, R. S. Harper, and R. K. Bramwell1 Department of Poultry Science, University of Arkansas, Fayetteville 72701 Primary Audience: Broiler Breeder Managers, Flock Supervisors, Researchers SUMMARY Downloaded from http://japr.oxfordjournals.org/ by guest on November 4, 2015 Commercial broiler breeder managers have reported differences in fertility among strains of birds, with the assumption that less feathering on the backs of hens results from more frequent mating. To determine if mating frequency is associated with back feathering, a study was under- taken to compare the mating behavior of 2 different lines of broiler breeder hens housed with a single strain of roosters. Each pen used in this study contained between 80 and 85 hens and 9 males. Mating behavior activities recorded included aggression, neck flares, male approaches, male waltzes, hen crouches, attempted mounts, completed mounts, attempted matings, and completed matings. Mating observations took place in the evening because this was the most active time for mating. Perivitelline layer sperm penetration, fertility, and hatchability were de- termined on eggs collected at each observation period. The hen strain with greater feather loss had fewer mounts by males as well as fewer completed matings, indicating that feather loss is not necessarily a good indicator of mating activity and, in fact, that it may deter such behavior. Sperm penetration, fertility, and hatchability were not different between the hen lines. In con- clusion, although mating behavior varies in some genetic strains of broiler breeders, it does not fully explain the differences in feather loss, indicating that feather loss on the backs of hens is not a good indicator of mating activity. Furthermore, even though mating activity varied in the 2 hen strains, it did not result in differences in fertility. Key words: mating behavior, broiler breeder, hatch 2010 J. Appl. Poult. Res. 19:24–29 doi:10.3382/japr.2009-00061 INTRODUCTION factors play a role in broiler breeder fertility, in- In the broiler breeder industry, it has long cluding the physical condition of the birds [2, 3], been assumed that hens with more feather loss the ratio of males to females [4], nutrition, the on their backs are mating more frequently than growth curve, and environmental factors such as hens with less feather loss. Jones and Prescott disease and temperature [5]. Siegel [6] observed [1] proposed that hens with less feather cover- a negative relationship between breast yield and ing may be those that more readily acquiesce to fertility, whereas Hocking and Duff [7] found males for mating. Thus, it is assumed that hens that a decline in fertility was caused by a reduc- with more feather loss, particularly as they age, tion of overall reproductive fitness. Bowling et will have higher fertility than hens with more al. [3] also reported a negative relationship be- feather covering on their backs. Many known tween sperm mobility and BW. 1 Corresponding author: bramwell@uark.edu
Moyle et al.: MATING BEHAVIOR IN BROILER BREEDERS 25 A study was initiated to determine if a dif- tained throughout production. New spare or re- ference in mating behavior and fertility existed placement males were not introduced during the between 2 strains of broiler breeder hens when observation periods. A total of 216 males and a single male strain was used. This study was approximately 2,000 hens were observed during undertaken when it was noticed in commercial the study. Mating activity was observed hourly flocks that strain 1 (S1) hens had fewer feath- throughout the day, and it was determined that ers on their backs than strain 2 (S2) hens, with mating activity peaked in the evening hours. The the actual cause of this difference not known. results of these observations are consistent with It was therefore postulated that greater mating the findings of other authors [10–13]. Therefore, activity in S1 hens caused the increased feather all observations for this study were made in 1-h loss. It was further postulated that if the S1 hens periods from 1800 to 1900 h, which was 14 to 15 mated more frequently, they would have higher h after the onset of light (16.75L:7.15D). fertility and greater sperm activity at the site of A total of 15 paired observations were made, fertilization. Bramwell et al. [8, 9] reported that with the same individual observing a pair of increased mating activity and increased num- adjoining pens containing the same male strain Downloaded from http://japr.oxfordjournals.org/ by guest on November 4, 2015 bers of sperm inseminated result in increased housed with 1 of 2 hen strains. All observations sperm penetration of the perivitelline layer (PL) were made during a 14-d period of time. Hens and increased fertility. Therefore, this study was were 49 wk old at the beginning of the trial. conducted to evaluate actual mating activity and Males were maintained with the same line of the behavioral patterns of broiler breeder males hens for the duration of the trial. All males used when housed with 2 different hen strains, as well were of the same strain and age and were reared as their fertility and PL sperm penetration val- sex separate at the same facility as the hens so ues. that any difference in mating behavior could be attributed to the hens. Both female strains were MATERIALS AND METHODS reared in the same facility and were reared to meet recommended BW. All observations took place at the University At the conclusion of all the observation peri- of Arkansas Broiler Breeder Research Farm. ods, eggs were sent to a commercial hatchery by This project complied with the provisions of hen strain group to determine fertility, hatchabil- the Institute Animal Care and Use Committee ity, and embryo livability through embryo diag- as specified by the Animal and Plant Health nosis of unhatched eggs. Additionally, 2 random Inspection Service, USDA, in 9 CFR Part samples of eggs (n = 120) were collected from 1(1–91). Birds used in this trial were housed in each hen strain and subjected to the sperm pen- a production facility that contained 48 experi- etration assay procedure to determine the PL mental and 5 spare pens measuring 4.3 m2 (14 sperm penetration rates [8, 9]. ft2). The birds were reared on the research farm Sperm penetration assays was performed us- in a separate pullet-rearing facility according to ing the technique described by Bramwell et al. industry standards and were housed in the pro- [8] to determine fertility because the number of duction house at 21 wk of age, at which time holes has been shown to be closely correlated the males and females were commingled. Each with fertility [8, 9]. Initially 120 eggs per treat- pen observed contained between 80 and 85 hens ment were used; however, some data were not and 9 males. Although this female-to-male ra- obtained because the PL tore or egg yolks broke tio (9:1) may by lower than that recommended before the PL could be removed. by the primary breeder, our past experience has Breakout analysis was performed on a mini- shown that in pen trials, more males are needed mum of 770 eggs per treatment. Early dead em- to obtain adequate fertility. Each pen contained bryos were those that died before 8 d of incu- separate feeders for the males and females to bation. Those that died between 8 and 14 d of control BW, water was provided ad libitum, and incubation were classified as midterm deaths, nests were provided to facilitate egg collection. and those that died after d 14 were recorded as Spare males were housed in the same facility late dead embryos. Fertility was also evaluated so that a constant number of males were main- at the time of breakout.
26 JAPR: Research Report Table 1. Results from the observation of mating behavior1 Item M-F MW HC AM CM ATM SM Strain 1 7.73 0.47 1.67 12.00 8.40a 7.93a 6.26a 2 5.07 0.33 3.60 16.07 12.60b 11.13b 9.26b P-value 0.1569 0.6615 0.1069 0.0562 0.0023 0.0145 0.0120 a,b Means within a column with different superscripts within a column differ significantly (P < 0.05). 1 M-F = male-hen aggression (male aggression not directly related to mating activity); MW = male waltz (dancing around or up to the hen in a courting manner); HC = hen crouch (when a hen crouches to the ground with wings at the shoulder elevated in an attempt to allow the rooster to mount); AM = attempted mounts (when a male makes an attempt to mount the hen for mating purposes); CM = completed mounts (when the male obtains a position on top of the hen); ATM = attempted mating (when a male mounts and tries to mate the hen); SM = successful mating (when a male makes cloacal contact with a hen). The results are reported as number of observations per hour. Characteristics observed were male-male sulted in the male obtaining a position on top Downloaded from http://japr.oxfordjournals.org/ by guest on November 4, 2015 aggression, male-hen aggression, hen-hen ag- of the female where mating activity could occur. gression, male waltzes, hen crouches, attempted Unsuccessful matings after a successful mount hen mounts, completed hen mounts, attempted occurred as a result of balance lost by the male hen matings, and completed hen matings. Male- during the act of copulation, by aggression by male aggression included fighting, aggressively other males, causing the male mounted atop the pecking the head of another male, or disrupting female to lose his position, or by the male appar- the mating action of another male. Male-hen ag- ently losing interest and voluntarily dismounting gression included aggressive attacks to the head after mounting the female and gaining a mating or body of a hen in such a way as not to be as- position. sociated with an attempted mating, or when the Attempted matings were recorded each time male attacked the hen in an aggressive way and a male had successfully mounted the female and tried to force the female into mating. Hen-hen then made an actual attempt to complete the aggression included any aggressive confronta- mating process through copulation. The caus- tion and pecking. ative factors for incomplete matings after an Incidence of a male waltz was the approach initial attempt are similar to those listed in the of a male to a female while the male dropped a previous paragraph. Successful matings were wing to the ground and performing a semicircu- recorded as the number of times males success- lar movement around the hen in a courting man- fully completed copulations (i.e, a male made ner [14, 15]. This type of courtship behavior is contact with the cloaca of the female). From this commonly observed among feral or backyard- contact, it was assumed that ejaculation took type domestic chickens, in which courtship of place and that semen was transferred. Each of the hens is more common [1]. In broiler breed- the previous events that occurred was recorded ers, this behavior is seen less often and may not be exhibited as vigorously. Hen crouches were recorded when a hen crouched to the ground Table 2. Sperm penetration analysis on 2 random with wings at the shoulder, elevated in an at- samples of eggs (n = 120 per treatment) from strain tempt to allow the rooster to mount for mating 1 (S1) and strain 2 (S2) hens taken at the time of activity. Hen crouches were recorded only if the observations1 crouch was independent of male mating activity Holes, no. S1, % S2, % and not after the male had attempted to mount 0–10 1.6 2.5 the hen. 11–30 3.2 2.5 Attempted mounts were recorded when a 31–60 4.8 5.0 male made an attempt to mount the female for 61–100 17.7 20.0 mating purposes, usually including grabbing the 100+ 72.6 70.0 female at the back of the neck behind the head 1 None of these treatment means are significantly different or comb. Completed mounts were those that re- (P > 0.05).
Moyle et al.: MATING BEHAVIOR IN BROILER BREEDERS 27 Table 3. Results of embryonic breakout from the study (n > 720 per treatment)1 Strain Fertile, % Hatch, % Early, % Mid, % Late, % a 1 96.25 86.25 7.27 0.20 6.73 2 96.39 85.28 7.80 1.40b 6.80 a,b Means with different superscripts differ significantly (P < 0.05). 1 Early = number of dead embryos between d 1 and 7; mid = number of embryos that died between d 8 and 15; late = number that died after d 15, including those that pipped. independently of other events that occurred in because of sexual rejection by the hens, or the the process. increased aggression may have been an attempt All data collected were evaluated by ANOVA at intimidating the hens into mating. This ag- using JMP statistical software, and means were gressive behavior had been noticed in field compared using the LSD method [16]. Differ- studies conducted by researchers at the Univer- Downloaded from http://japr.oxfordjournals.org/ by guest on November 4, 2015 ences were deemed to be significant when the sity of Arkansas (unpublished data). When the P-value was less than 0.05. males matured more quickly than the females, they became more aggressive and attacked the RESULTS AND DISCUSSION females. This aberrant behavior occasionally led to forced copulations. Millman et al. [18] report- The results of the behavioral observations ed that male broiler breeders displayed a greater from this study are shown in Table 1 and are re- than normal aggression toward females and that ported as the number of observations per hour. it could lead to injuries to the females. At the The S2 hens were mounted, attempted to mate, time of the study, males attempted to mount the and mated more frequently than the S1 hens. S1 females less frequently but did tend to peck The S2 hens may have been more receptive to aggressively at the S1 females more often. This, males, or males may have responded more read- however, would not account for the feather loss ily to hen signals or cues or were simply more on the backs of the females. We have observed interested in these hens, and the hens were there- male aggression toward females peaking at ap- fore mated more often than S1 hens. Marin and proximately 28 wk of age and then declining Satterlee [17] found that males mated more of- rapidly thereafter (unpublished data). Therefore, ten with hens that had been selected for low lev- previous injury to the hens would have taken els of stress compared with hens that had high place 20 wk before the observation and the fe- levels of stress. Therefore, it may be that the S2 males would have had time to recover. hens had inadvertently been selected for some Table 2 contains the results of the PL sperm other traits or other characteristics that the males penetration analysis performed on samples of preferred that were undetectable to the human eggs laid by both strains of hens [8, 9]. There eye. This is in line with observations by industry were no differences in the percentages for each personnel who have reported that S2 birds are category. Results showed both strains to have better adapted to varied environments than S1 very good sperm activity at the site of fertil- birds. Therefore, the increased loss of feathers ization. Although the S1 hens were mated less in the S1 hens was not a result of more frequent frequently than the S2 hens, they were able to mating or mating behavior and must therefore be maintain PL sperm penetration values as high as attributed to some other factor. Feather loss may those of S2 birds. Brillard and Bakst [19] report- be genetic, the feather strength of the S1 hens ed a strong correlation between the number of may be inferior to that of the S2 hens, or the S1 sperm in the sperm storage tubules and the holes hems may simply be less densely feathered. in the PL. Therefore, a possible explanation for Although not statistically significant, there the higher than expected sperm activity at the was a trend (P = 0.16) for more male-female site of fertilization is that the S1 hens were able aggression from the males housed with the S1 to store sperm for a longer duration [20] or that hens (Table 1). The trend toward increased ag- they had a greater capacity for oviductal sperm gression appeared to be caused by frustration storage. Last, there is also the possibility that
28 JAPR: Research Report a difference existed between the strains of fe- 4. The role of the hens is often overlooked males in their physiological ability to select and when searching for reasons for low fer- transport sperm [21]. If sperm in S1 hens had tility in broiler breeders, and this needs increased selection and transport, then S1 hens to be investigated more thoroughly. would need to be mated less frequently than the S2 hens. The question may then arise whether REFERENCES AND NOTES the males are able to detect the need for indi- 1. Jones, E. K. M., and N. B. Prescott. 2000. Visual vidual hens to be mated based on their fertility cues used in the choice of mate by fowl and their potential or sperm availability. importance for the breeder industry. World’s Poult. Sci. J. 56:127–138. Results of the residue breakout from the 2. McDaniel, G. R., J. Brake, and M. K. Eckman. 1981. study are shown in Table 3. The only signifi- Factors affecting broiler breeder performance. 4. The in- cant difference between the strains was in the terrelationship of some reproductive traits. Poult. Sci. number of embryos that died in the middle of 60:1792–1797. the incubation period (between 8 and 15 d). One 3. Bowling, E. R., D. P. Froman, A. J. Davis, and J. L. Wilson. 2003. Attributes of broiler breeder male character- possible reason for this difference is the low Downloaded from http://japr.oxfordjournals.org/ by guest on November 4, 2015 ized by low and high sperm mobility. Poult. Sci. 82:1796– number of embryos that die during the middle 1801. period of incubation, so any outliers have a large 4. Hazary, R. C., H. J. Staines, and G. J. Wishart. 2001. Assessing the effect of mating ratio on broiler breeder per- effect on the results. There was no significant formance by quantifying sperm:egg interaction. J. Appl. difference in fertility, hatch, or early and late Poult. Res. 10:1–4. dead embryos. 5. McDaniel, C. D., R. K. Bramwell, J. L. Wilson, and B. Howarth. 1995. Fertility of male and female broiler breeders following exposure to elevated ambient tempera- CONCLUSION AND APPLICATIONS tures. Poult. Sci. 74:1029–1038. 6. Siegel, P. B. 1962. A double selection experiment for 1. There are differences in mating activity body weight and breast angle at eight weeks of age in chick- ens. Genetics 47:1313–1319. between different strains of hens; thus, 7. Hocking, P. M., and S. R. I. Duff. 1989. Musculo- management strategies, such as the fe- skeletal lesions in adult male broiler-breeder fowls and their male-to-male ratio, need to be based on relationships with body weight and fertility at 60 weeks of age. Br. Poult. Sci. 30:777–784. these differences to maximize fertility. 8. Bramwell, R. K., H. L. Marks, and B. Howarth. 1995. 2. It was observed that although a differ- Quantitative determination of spermatozoa penetration of ence in mating behavior existed in the the perivitelline layer of the hen’s ovum as assessed on ovi- 2 strains of hens, this difference did not posited eggs. Poult. Sci. 74:1875–1883. equate to a difference in fertility or PL 9. Bramwell, R. K., C. D. McDaniel, J. L. Wilson, and B. Howarth. 1996. Age effect of male and female broiler sperm penetration values overlying the breeders on sperm penetration of the perivitelline laver over- germinal disc. One possible reason that lying the germinal disc. Poult. Sci. 75:755–762. the S1 hens maintained higher levels of 10. Lake, P. E., and D. G. M. Wood-Gush. 1956. Diurnal rhythms in semen yields and mating behaviour in the domes- fertility while mating less frequently is tic cock. Nature 178:853. they may be able to store sperm more 11. Duncan, I. J. H., P. M. Hocking, and E. Seawright. efficiently or in greater quantities than 1990. Sexual behavior and fertility in broiler breeder domes- the S2 hens. Another reason may be tic fowl. Appl. Anim. Behav. Sci. 26:201–213. that these hens have a greater number of 12. Pizarri, A. R., and T. R. Birkhead. 2001. For whom does the hen cackle? The function of postoviposition cack- sperm-binding proteins in the PL overly- ling. Anim. Behav. 61:601–607. ing the germinal disc. In addition, it is 13. Bilcik, B., and I. Estevez. 2005. Impact of male-male possible that although these hens mated competition and morphological traits on mating strategies and reproductive success in broiler breeders. Appl. Anim. less frequently, they were still mating of- Behav. Sci. 92:307–323. ten enough to maintain high fertility. 14. Wood-Gush, D. G. M. 1954. The courtship of the 3. Hens play an important role in the mat- Brown Leghorn cock. Br. J. Anim. Behav. 2:95–102. ing behavior, and although males can 15. Wood-Gush, D. G. M. 1956. The agonistic and court- ship behaviour of the Brown Leghorn cock. Br. J. Anim. physically dominate the hens, hen be- Behav. 4:133–142. havior can still affect the overall mating 16. SAS Institute. 2009. JMP Version 8.0. SAS Inst. Inc., activity. Cary, NC.
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