Frontiers in sheep reproduction - making use of natural responses to environmental challenges to manage productivity.

This review addresses advances, directions and opportunities for research on sheep reproduction in the context of the global challenges of food security and climate change, and demand for 'clean, green and ethical' (CGE) animal management. The foundation of CGE management is an understanding of the physiological processes through which the reproductive system responds to changes in the animal's environment. These days, to the main environmental factors (photoperiod, nutrition, pheromones), we need to add stress from extreme weather events. With respect to nutrition in rams, we now have a deeper understanding of the responses of the brain centres that control gonadotrophin secretion (the kisspeptin system). At testis level, we have found that nutrition affects non-coding RNAs in Sertoli cells and germ cells, thus affecting the balance between cell proliferation and apoptosis. This proliferation-apoptosis balance is also affected during prenatal development, when undernutrition or stress in pregnant ewes seems to elicit epigenetic changes in developing gonads that could affect offspring fertility in adult life. With respect to nutrition in ewes, metabolic signals act directly on ovarian follicles, and thus change ovulation rate, but the variety of signals now includes the adipokines. An early concern was that nutritional supplements that increase ovulation rate would also increase embryo mortality but we now know that embryo survival is improved under field conditions. Finally, we had always thought that the efficiency gains from early puberty in lambs could only be achieved by accelerating fat accumulation, but we now know that faster muscle growth will achieve the same goal, offering two advantages in meat production systems. With respect to pheromones ('ram effect'), we have a deeper understanding of the brain responses (kisspeptin system) but, most importantly, a realization that the response of ewes to the ram signal involves cell division in memory centres. Many opportunities remain.