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Amdam, G. V., Csondes, A., Fondrk, M. K., & Page, R. E. J. (2006). Complex social behaviour derived from maternal reproductive traits. Nature, 439(7072), 76–78.
Abstract: A fundamental goal of sociobiology is to explain how complex social behaviour evolves, especially in social insects, the exemplars of social living. Although still the subject of much controversy, recent theoretical explanations have focused on the evolutionary origins of worker behaviour (assistance from daughters that remain in the nest and help their mother to reproduce) through expression of maternal care behaviour towards siblings. A key prediction of this evolutionary model is that traits involved in maternal care have been co-opted through heterochronous expression of maternal genes to result in sib-care, the hallmark of highly evolved social life in insects. A coupling of maternal behaviour to reproductive status evolved in solitary insects, and was a ready substrate for the evolution of worker-containing societies. Here we show that division of foraging labour among worker honey bees (Apis mellifera) is linked to the reproductive status of facultatively sterile females. We thereby identify the evolutionary origin of a widely expressed social-insect behavioural syndrome, and provide a direct demonstration of how variation in maternal reproductive traits gives rise to complex social behaviour in non-reproductive helpers.
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Belonje, P. C., & van Niekerk, C. H. (1975). A review of the influence of nutrition upon the oestrous cycle and early pregnancy in the mare. J Reprod Fertil Suppl, (23), 167–169.
Abstract: Attention is drawn to the beneficial effect of improved nutrition during winter and early spring on the ovarian activity of mares. Furthermore, the necessity of an adequate plane of nutrition during early pregnancy to prevent embryonic resorption is stressed.
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Keiper, R., & Houpt, K. (1984). Reproduction in feral horses: an eight-year study. Am J Vet Res, 45(5), 991–995.
Abstract: The reproductive rate and foal survival of the free-ranging ponies on Assateague Island National Seashore were studied for 8 years, 1975 to 1982. Most (52%) of the 86 foals were born in May, 13% were born in April, 22.6% in June, 10.4% in July, and less than 1% in August and September. The mean foaling rate was 57.1 +/- 3.9% and the survival rate was 88.3 +/- 3.6%. Forty-eight colts and 55 fillies were born (sex ratio 53% female). Mares less than 3 years old did not foal and the foaling rate of 3-year-old mares was only 23%, that of 4-year-old mares was 46%, that of 5-year-old mares was 53%, and 6-year-old mares was 69%. The relatively poor reproduction rate was believed to be a consequence of the stress of lactating while carrying a foal when forage quality on the island was low. The hypothesis was supported by the higher reproductive rate (74.4 +/- 2.4%) of the ponies in the Chincoteague National Wildlife Refuge on the southern part of the island. Their foals are weaned and sold in July each year. Despite the low reproductive rate on Assateague Island National Seashore , the number of ponies increased from 43 to 80, a 90% increase in the 8-year period or greater than 10%/yr. There were 24 deaths and 8 dispersals from the study area.
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Kirkpatrick, J. F., & Turner, A. (2002). Reversibility of action and safety during pregnancy of immunization against porcine zona pellucida in wild mares (Equus caballus). Reprod Suppl, 60, 197–202.
Abstract: Contraceptive management of publicly valued wildlife species requires safeguards to ensure that these populations are preserved in a healthy state. In addition, reversibility of contraceptive effects and safety in pregnant animals are major concerns. A population of wild horses has been immunized against porcine zona pellucida (PZP) over a 12 year period on Assateague Island National Seashore, MD (ASIS). Mares initially received one or two 65 microg inoculations and once a year 65 microg booster inoculations, all delivered by dart. All young mares aged > 2 years were treated with PZP for 3 consecutive years regardless of whether they have bred successfully and they were then removed from treatment until they had foaled. All mares vaccinated for 1 or 2 consecutive years became fertile again and 69% of mares treated for 3 consecutive years returned to fertility. All five mares treated for 4 or 5 consecutive years have also returned to fertility, but over longer periods of time. Mares treated for 7 consecutive years have not returned to fertility, but several, while still infertile, have started ovulating again. There was no difference in survival rates between foals born to treated and untreated mares, and PZP treatment of pregnant mares did not affect subsequent fertility of their female offspring.
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Lee, J., & Paik, M. (2006). Sex preferences and fertility in South Korea during the year of the Horse. Demography, 43(2), 269–292.
Abstract: Since antiquity, people in several East Asian countries, such as China, Japan, and South Korea, have believed that a person is destined to possess specific characteristics according to the sign of the zodiac under which he or she was born. South Koreans, in particular, have traditionally considered that the year of the Horse bears inauspicious implications for the birth of daughters. Using monthly longitudinal data at the region level in South Korea between 1970 and 2003, we found that in the year of the Horse, the sex ratio at birth significantly increased while fertility decreased.
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Lee, R. D. (2003). Rethinking the evolutionary theory of aging: transfers, not births, shape senescence in social species. Proc Natl Acad Sci U S A, 100(16), 9637–9642.
Abstract: The classic evolutionary theory of aging explains why mortality rises with age: as individuals grow older, less lifetime fertility remains, so continued survival contributes less to reproductive fitness. However, successful reproduction often involves intergenerational transfers as well as fertility. In the formal theory offered here, age-specific selective pressure on mortality depends on a weighted average of remaining fertility (the classic effect) and remaining intergenerational transfers to be made to others. For species at the optimal quantity-investment tradeoff for offspring, only the transfer effect shapes mortality, explaining postreproductive survival and why juvenile mortality declines with age. It also explains the evolution of lower fertility, longer life, and increased investments in offspring.
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Ransom, J. I., Cade, B. S., & Hobbs, N. T. (2010). Influences of immunocontraception on time budgets, social behavior, and body condition in feral horses. Appl. Anim. Behav. Sci., 124(1-2), 51–60.
Abstract: Managers concerned with shrinking habitats and limited resources for wildlife seek effective tools for limiting population growth in some species. Fertility control is one such tool, yet little is known about its impacts on the behavioral ecology of wild, free-roaming animals. We investigated influences of the immunocontraceptive porcine zona pellucida (PZP) on individual and social behavior in bands of feral horses (Equus caballus) in three discrete populations and used 14 hierarchical mixed effect models to gain insight into the influences of PZP treatment on feral horse behavior. A model of body condition was the strongest predictor of feeding, resting, maintenance, and social behaviors, with treated females allocating their time similarly to control females. Time spent feeding declined 11.4% from low condition to high condition females (F1,154 = 26.427, P < 0.001) and was partially reciprocated by a 6.0% increase in resting (F1,154 = 7.629, P = 0.006), 0.9% increase in maintenance (F1,154 = 7.028, P = 0.009), and 1.8% increase in social behavior (F1,154 = 15.064, P < 0.001). There was no difference detected in body condition of treated versus control females (F1,154 = 0.033, P = 0.856), but females with a dependent foal had lower body condition than those without a foal (F1,154 = 4.512, P = 0.038). Herding behavior was best explained by a model of treatment and the interaction of band fidelity and foal presence (AICc weight = 0.660) which estimated no difference in rate of herding behavior directed toward control versus treated females (F1,102 = 0.196, P = 0.659), but resident females without a dependent foal were herded 50.9% more than resident females with a foal (F3,102 = 8.269, P < 0.001). Treated females received 54.5% more reproductive behaviors from stallions than control mares (F1,105 = 5.155, P = 0.025), with the model containing only treatment being the most-supported (AICc weight = 0.530). Treated and control females received harem-tending behaviors from stallions equally (F1,105 = 0.001, P = 0.969) and agonistic behaviors from stallions equally (F1,105 < 0.001, P = 0.986). Direct effects of PZP treatment on the behavior of feral horses appear to be limited primarily to reproductive behaviors and most other differences detected were attributed to the effects of body condition, band fidelity, or foal presence. PZP is a promising alternative to traditional hormone-based contraceptives and appears to contribute few short-term behavioral modifications in feral horses.
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Turner, J. W. J., & Kirkpatrick, J. F. (1982). Androgens, behaviour and fertility control in feral stallions. J Reprod Fertil Suppl, 32, 79–87.
Abstract: This field study of feral stallions in Montana and Idaho examines and correlates the seasonal pattern of plasma androgens and specific sociosexual behaviour and reports the effect of a long-acting androgenic steroid on this behaviour and on fertility. Plasma testosterone was measured by competitive protein binding assay in samples obtained by jugular venepuncture from captured animals. In samples taken from 34 sexually mature stallions in 6 different months during the year, a definite seasonal pattern in testosterone was present, with a peak in May (3.04 +/- 0.63 ng/ml) and a nadir in December (1.55 +/- 0.34 ng/ml). Values were less than 2.0 ng/ml in non-breeding months and greater than 2.4 ng/ml in breeding months. Behavioural endpoints measured were (1) stallion scent marking in response to elimination by mares (elimination marking), (2) mounting and (3) copulation. The frequencies of each of these endpoints followed closely the seasonal pattern seen for plasma androgens. In the fertility study microcapsulated testosterone propionate (microTP) was administered i.m. to 10 harem stud stallions 3 months before the 1980 breeding season. In these stallions and in 10 control harem studs, the above behavioural endpoints were examined in the 1980 and 1981 breeding seasons, and foal counts were made in 1981. There were no direct inhibitory or stimulatory effects of microTP treatment on any of the behavioural endpoints in either year. In 1981 foals were produced in 87.5% of the control bands and 28.4% of the microTP-treated bands. These results indicate that microencapsulated testosterone propionate can provide effective fertility control in feral horses without causing significant alterations in sociosexual behaviour.
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