Puberty is defined as the ability to accomplish reproduction successfully (10). It can be defined in many ways in females such as the onset of estrus, the age of first ovulation, or the age at which an individual can support pregnancy without negative effects (10).
For sheep (ovine), the average age of puberty is similar between males and females with it happening at around 7 months of age (10). As seen from the chart, however, there is a large range in which it may occur, with females potentially showing their first signs between 4-14 months of age (10).
Feedback Loop
Negative Feedback: Estrogen and progesterone exert a negative feedback on the hypothalamus and pituitary most of the time. This suppresses the release of GnRH, FSH, and LH to prevent excessive hormone levels. This mechanism helps regulate the early follicular phase, where rising estrogen levels from developing follicles inhibit further FSH and LH release. (10)
Positive Feedback: Just before ovulation, when estrogen levels reach a certain threshold, estrogen temporarily exerts a positive feedback on the hypothalamus and pituitary. This leads to a surge in LH and a smaller rise in FSH, triggering ovulation. (10)
Hypothalamus: Secretes gonadotropin-releasing hormone (GnRH) in a pulsatile manner. (10)
Pituitary Gland: In response to GnRH, the pituitary gland releases two hormones—follicle-stimulating hormone (FSH) and luteinizing hormone (LH). (10)
Ovaries: FSH and LH act on the ovaries: FSH stimulates the growth and maturation of ovarian follicles. LH triggers ovulation and supports the production of sex hormones (estrogen and progesterone) from the ovaries. (10)
Hypothalamus: The hypothalamus is a small region of the brain that plays a major role in regulating various physiological processes, including reproduction, metabolism, stress response, and temperature control (10). It is located just above the brainstem, and it connects the nervous system to the endocrine system via the pituitary gland, which it controls through both neural signals and hormone release in the posterior and anterior sides respectively (10).
Hormones: The main hormone released by the hypothalamus is Gonadotropin-Releasing Hormone (GnRH) (10). This hormone stimulates the anterior pituitary to release luteinizing hormone and follicle-stimulating hormone (FSH) (10).
Pituitary Gland: It is the front portion of the pituitary gland, a small located at the base of the brain (10). The anterior pituitary is crucial for reproduction as it releases hormones that act directly on the ovaries in females, which influences fertility, sexual development, and reproductive cycles (10).
Hormones: The anterior pituitary is known to produce luteinizing hormone (LH) and follicle-stimulating hormone (FSH) (10). LH stimulates ovulation, and also promotes the formation of the corpus luteum, which produces progesterone necessary for maintaining pregnancy (10). FSH stimulates the growth and development of ovarian follicles, which support the developing egg (10). FSH also promotes the production of estrogen by the follicles, which helps regulate the menstrual cycle and prepares the body for ovulation (10).
Senger, P. L. (2015). Pathways to pregnancy and parturition (3rd ed.). Current Conceptions Inc.
Important Structures of the Hypothalamus
Surge Center: Full neural activity of the surge center must be achieved before ovulation can occur. When this happens, bursts of GnRH occur (10). in which the GnRH neurons fire frequently and release large quantities of GnRH causing a pre-ovulatory LH surge (10). The surge center is "activated" once estrogen reaches a critical threshold from hormonal changes during puberty (10).
Tonic Center: Must also reach a functional state since it regulates the tonic frequency of LH pulses. Frequency of GnRH pulses may be linked to “pulse generator” in the hypothalamus, which can read external and internal signals to influence the release of GnRH (10). This ultimately dictates the time of puberty. The tonic center is responsible for the basal release of GnRH 10, and it maintains a relatively steady and continuous pattern of secretion, which stimulates the release of small amounts of LH and FSH (10). This constant activity is crucial for the regulation of reproductive cycles and maintaining normal reproductive function in females and stimulates the ovaries (10).
Sheep have multiple estrous cycles during a specific season, making them seasonal polyestrous. This is controlled by changes in daylight, with breeding usually occurring in the fall and lambing in the spring.
Przygrodzka, E. (2024, October). Reproductive Cyclicity and Endocrinology. University of Maryland Department of Animal and Avian Sciences.
Przygrodzka, E. (2024, October). Reproductive Cyclicity and Endocrinology. University of Maryland Department of Animal and Avian Sciences.
Timing of the Estrous Cycle:
Length of estrous cycle = average of 17 days
Follicular phase (proestrus & estrus) = 20% of estrous cycle
Luteal phase (metestrus & diestrus) = 80% of estrous cycle (10)
4 Stages of the Estrous Cycle:
Proestrus -
Lasts between 12-14 hours for ewes (10)
Begins when progesterone declines due to luteolysis
Estrogen increases
Preovulatory follicle increases in size
Antral follicle matures for ovulation (14)
Estrus -
Also referred to as "heat"
Period of sexual receptivity
Average length of 29 hours
Estrogen levels peak and induce behavioral changes
Behavioral signals that she is ready for mating - standing to be mounted, more frequent urination, vocalization, etc. (10)
Surge of luteinizing hormone (LH)
Ovulation occurs 24-48 hours after the LH surge
High uterine motility (contractions)
Optimal conditions for sperm transport
Increase in cervical mucus (14)
Metestrus -
Occurs right after ovulation
Time between ovulation and the formation of a functional corpus luteum
Estrogen and progesterone levels are low in the beginning of this stage
Newly ovulated follicle forms the corpus luteum (luteolysis)
Corpus hemorrhagicum (bloody body) is present
The corpus luteum begins to secrete progesterone
Uterine contractions end
FSH increases which stimulates follicle growth (14)
Diestrus -
The longest stage of the estrous cycle
The corpus luteum (CL) is fully functional
High levels of progesterone secretion from the CL
The uterus prepares for early-embryo development
FSH increases to promote follicle growth within the ovaries
The CL regresses at the end of this phase if the female is not pregnant and Prostaglandin is released as a result (14)
Przygrodzka, E. (2024, October). Reproductive Cyclicity and Endocrinology. University of Maryland Department of Animal and Avian Sciences.
Przygrodzka, E. (2024, October). Reproductive Cyclicity and Endocrinology. University of Maryland Department of Animal and Avian Sciences.
Przygrodzka, E. (2024, September). Follicular and Luteal Phase of the Reproductive Cycle. University of Maryland Department of Animal and Avian Sciences.
Sheep Are Short Day Breeders
Increased darkness causes higher melatonin release from the pineal gland in the brain
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Decreases RFRP-3 inhibition of kisspeptin neurons in the hypothalamus
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High kisspeptin-10 stimulates the release of GnRH from the hypothalamus which initiates proestrus
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GnRH acts on the gonadotropin cells of the anterior pituitary gland to increase release of FSH and LH
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FSH and LH travel through the bloodstream to reach the ovaries
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FSH promotes the growth & maturation of ovarian follicles that begin to secrete estrogen and inhibin
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Increase in estrogen allows for behavioral changes in the ewe during estrus and stimulates a surge of LH that initiates ovulation while inhibin suppresses FSH secretion from the pituitary gland
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After ovulation, during metestrus, the ruptured follicle forms a corpus luteum which secretes progesterone
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During diestrus progesterone maintains pregnancy if successful implantation occurred or if there is no pregnancy, the corpus luteum regresses and progesterone levels drop
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A new estrous cycle begins with low progesterone levels
Key Hormones in the Estrous Cycle
GnRH:
Controls the release of FSH and LH from the anterior pituitary which then act on the ovaries to stimulate follicle synthesis
FSH:
Promotes the growth and development of follicles
Increases oxygen uptake of granulosa cells to initiate protein synthesis
Promotes formation of granulosa cells that secrete estrogen which are important for follicle maturation
Prepares the dominant follicle for ovulation in response to surge of LH
LH:
Aids in maturation of primordial germ cells (eggs)
Influences steroid hormone synthesis in the ovaries
Regulates length and order of phases in the estrous cycle
Estradiol:
Promotes maturation of growing follicles
Causes behavioral changes that occur during estrus (heat) - signal that the ewe is ready to mate
Enhances blood flow and glandular secretions to prepare the uterine lining for implantation of a fertilized egg
Progesterone:
Increases during luteinization in the metestrus phase of the estrous cycle
Produced by luteal cells in the corpus luteum (CL) to maintain pregnancy
Insufficent progesterone levels can lead to miscarriage during early pregnancy
Prostaglandin:
Receptors found on large luteal cells of the corpus luteum
Controls luteolysis in the ovary
Inhibin:
Produced by follicles once FSH and LH are released from the anterior pituitary
Reduced supply can lead to atresia (cell death) of the maturing follicle