novel approaches to expression and detection of oestrus in dairy cows
novel approaches to expression and detection of oestrus in dairy cows novel approaches to expression and detection of oestrus in dairy cows
1.2.1.1 Follicular Phase There are a fixed number of primordial follicles established during foetal development (Webb et al., 2004), which deplete thereafter as many follicles become atretic because follicle growth occurs continuously throughout the cow’s reproductive life (Fortune, 1993). Cattle are monovular therefore as primordial follicles are recruited most become atretic and do not progress through to selection of the single dominant follicle, and few follicles (4mm in diameter, until follicles reach 6-8mm diameter, when one follicle is selected for continued growth and becomes dominant and will either ovulate or undergo atresia (Webb et al., 2003). Follicular development is governed by a period of gonadotrophin independence followed by a period of dependence (Webb et al., 2004). During recruitment, each wave is preceded by increases of FSH secretion lasting 1-2 days, stimulating growth of smaller follicles,
ecomes the main inhibitor of FSH and dependency switches to LH (Ginther et al., 1997). Growth of the dominant follicle continues, and the increase in oestradiol causes increased GnRH pulse frequencies, promoting LH secretion. This in turn stimulates oestradiol production by the granulosa cells (Fortune, 1994). This has a local effect on follicle development, but also a systemic effect, acting upon the hypothalamic-pituitary axis to increase LH production further, by positive feedback (Aerts and Bols, 2010). This ultimately results in oestrus, controlled by oestradiol, and the LH surge resulting in ovulation. 1.2.1.2 Oestrus & Ovulation It is this rise in oestradiol; enhanced by LH, stimulating production of androgen in the theca cells (Garverick et al., 2002) and the subsequent androgens being converted into oestradiol by aromatase enzyme from granulosa cells, which causes oestrus (Fortune, 1994). The positive feedback mechanism between oestradiol and LH, causes LH pulse frequency to increase to about 1 pulse per hour (Roche, 2006). The increase in LH concentration causes a cascade of events that induce the release of the oocyte into the oviduct, by an inflammatory response; involving prostaglandins, particularly prostaglandin E (PGE), produced by the follicle (Aerts and Bols, 2010). Prostaglandins stimulate the proliferation of cells and production of proteolytic enzymes to disrupt the follicle wall, releasing the oocyte (Espey, 1980). This process is ultimately under the control of the follicle itself, timing when it is appropriate to trigger the LH surge for ovulation by production of oestradiol (Roelofs et al., 2010), usually about 10-14 hours after oestrus (Forde et al., 2011). 1.2.1.3 Luteal Phase LH is the key hormone stimulating luteinisation of the theca and granulosa cells post ovulation, forming the CL from the cells of the ruptured cavity (Alila and Hansel, 1984). The CL consists of small and large luteal cells, which have steroidogenic properties (Smith et al., 1994), which secrete progesterone, along with a range of other cell types. The function of the CL is to produce progesterone, in order to maintain pregnancy if a conceptus is present (Forde et al., 2011). Sustained production of progesterone suppresses GnRH pulse frequency and hence LH secretion to prevent ovulation, but does allow enough LH for the continuation of follicular waves and dominant follicle growth (Savio et al., 1990). 7
- Page 1 and 2: NOVEL APPROACHES TO EXPRESSION AND
- Page 3 and 4: TABLE OF CONTENTS TABLE OF CONTENTS
- Page 5 and 6: 2.2.3 Statistical Analyses ........
- Page 7 and 8: 5.3 RESULTS .......................
- Page 9 and 10: Furthermore, automated software was
- Page 11 and 12: LIST OF FIGURES Figure 1.1 Hormonal
- Page 13 and 14: LIST OF TABLES Table 1.1 Trends in
- Page 15 and 16: LIST OF ABBREVIATIONS ˚ ˚C μM 2D
- Page 17 and 18: CHAPTER 1 - Introduction & Literatu
- Page 19 and 20: the past 50 years and duration of o
- Page 21: Oestrus growing follicle (Staigmill
- Page 25 and 26: calf at 40-50 days post partum; inv
- Page 27 and 28: al., 2006). However, aged sperm hav
- Page 29 and 30: can occur within 2-3 days, but if t
- Page 31 and 32: educes the incidence of problem cow
- Page 33 and 34: oestradiol, the LH surge and ovulat
- Page 35 and 36: The secondary signs of oestrus can
- Page 37 and 38: There are also changes in normal be
- Page 39 and 40: engage in more natural behaviours i
- Page 41 and 42: 1983). Exact explanations and mecha
- Page 43 and 44: 2006) and disruption of LH secretio
- Page 45 and 46: 1.4.3.2 Milk Yield and Nutrition Di
- Page 47 and 48: influence the ability of the ovary
- Page 49 and 50: cyclicity can be delayed if dietary
- Page 51 and 52: indication for the optimal time to
- Page 53 and 54: calving), 3) pre-breeding heat date
- Page 55 and 56: 1.5.2 Physiological Changes Physiol
- Page 57 and 58: 1.5.2.3 Body and Milk Temperature T
- Page 59 and 60: physical activity and stage of the
- Page 61 and 62: caused by the general environment t
- Page 63 and 64: may be gained. This is because data
- Page 65 and 66: In summary the objective was to for
- Page 67 and 68: diet, with concentrates at milking.
- Page 69 and 70: oestrus was defined as 3 consecutiv
- Page 71 and 72: The interaction between parity and
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