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novel approaches to expression and detection of oestrus in dairy cows

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ecomes the ma<strong>in</strong> <strong>in</strong>hibi<strong>to</strong>r <strong>of</strong> FSH <strong>and</strong> dependency switches <strong>to</strong> LH (G<strong>in</strong>ther<br />

et al., 1997). Growth <strong>of</strong> the dom<strong>in</strong>ant follicle cont<strong>in</strong>ues, <strong>and</strong> the <strong>in</strong>crease <strong>in</strong><br />

oestradiol causes <strong>in</strong>creased GnRH pulse frequencies, promot<strong>in</strong>g LH<br />

secretion. This <strong>in</strong> turn stimulates oestradiol production by the granulosa<br />

cells (Fortune, 1994). This has a local effect on follicle development, but<br />

also a systemic effect, act<strong>in</strong>g upon the hypothalamic-pituitary axis <strong>to</strong><br />

<strong>in</strong>crease LH production further, by positive feedback (Aerts <strong>and</strong> Bols,<br />

2010). This ultimately results <strong>in</strong> <strong>oestrus</strong>, controlled by oestradiol, <strong>and</strong> the<br />

LH surge result<strong>in</strong>g <strong>in</strong> ovulation.<br />

1.2.1.2 Oestrus & Ovulation<br />

It is this rise <strong>in</strong> oestradiol; enhanced by LH, stimulat<strong>in</strong>g production <strong>of</strong><br />

<strong>and</strong>rogen <strong>in</strong> the theca cells (Garverick et al., 2002) <strong>and</strong> the subsequent<br />

<strong>and</strong>rogens be<strong>in</strong>g converted <strong>in</strong><strong>to</strong> oestradiol by aromatase enzyme from<br />

granulosa cells, which causes <strong>oestrus</strong> (Fortune, 1994). The positive<br />

feedback mechanism between oestradiol <strong>and</strong> LH, causes LH pulse<br />

frequency <strong>to</strong> <strong>in</strong>crease <strong>to</strong> about 1 pulse per hour (Roche, 2006). The<br />

<strong>in</strong>crease <strong>in</strong> LH concentration causes a cascade <strong>of</strong> events that <strong>in</strong>duce the<br />

release <strong>of</strong> the oocyte <strong>in</strong><strong>to</strong> the oviduct, by an <strong>in</strong>flamma<strong>to</strong>ry response;<br />

<strong>in</strong>volv<strong>in</strong>g prostagl<strong>and</strong><strong>in</strong>s, particularly prostagl<strong>and</strong><strong>in</strong> E (PGE), produced by<br />

the follicle (Aerts <strong>and</strong> Bols, 2010). Prostagl<strong>and</strong><strong>in</strong>s stimulate the<br />

proliferation <strong>of</strong> cells <strong>and</strong> production <strong>of</strong> proteolytic enzymes <strong>to</strong> disrupt the<br />

follicle wall, releas<strong>in</strong>g the oocyte (Espey, 1980). This process is ultimately<br />

under the control <strong>of</strong> the follicle itself, tim<strong>in</strong>g when it is appropriate <strong>to</strong><br />

trigger the LH surge for ovulation by production <strong>of</strong> oestradiol (Roel<strong>of</strong>s et<br />

al., 2010), usually about 10-14 hours after <strong>oestrus</strong> (Forde et al., 2011).<br />

1.2.1.3 Luteal Phase<br />

LH is the key hormone stimulat<strong>in</strong>g lute<strong>in</strong>isation <strong>of</strong> the theca <strong>and</strong> granulosa<br />

cells post ovulation, form<strong>in</strong>g the CL from the cells <strong>of</strong> the ruptured cavity<br />

(Alila <strong>and</strong> Hansel, 1984). The CL consists <strong>of</strong> small <strong>and</strong> large luteal cells,<br />

which have steroidogenic properties (Smith et al., 1994), which secrete<br />

progesterone, along with a range <strong>of</strong> other cell types. The function <strong>of</strong> the CL<br />

is <strong>to</strong> produce progesterone, <strong>in</strong> order <strong>to</strong> ma<strong>in</strong>ta<strong>in</strong> pregnancy if a conceptus<br />

is present (Forde et al., 2011). Susta<strong>in</strong>ed production <strong>of</strong> progesterone<br />

suppresses GnRH pulse frequency <strong>and</strong> hence LH secretion <strong>to</strong> prevent<br />

ovulation, but does allow enough LH for the cont<strong>in</strong>uation <strong>of</strong> follicular waves<br />

<strong>and</strong> dom<strong>in</strong>ant follicle growth (Savio et al., 1990).<br />

7

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