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

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on average in Holsteins (Dransfield et al., 1998). The percentage of cows actually standing to be mounted has also declined from 80% to 50% (Dobson et al., 2008). This results in only 50% of cows being observed in oestrus (Van Eerdenburg et al., 2002). Hence detection of oestrus is increasingly more difficult and to improve the decline in fertility oestrous detection rates must be improved. 1.3.1.1 Endocrine, Neural and Genomic Changes Associated with Oestrous Behaviour Oestradiol is the key regulator that synchronizes the endocrinological and behavioural events to drive oestrus; resulting from the action of ovarian steroids on behavioural centres in the brain (Roelofs et al., 2010). The production of oestradiol from the ovary synchronises mating and ovulation; rising above its threshold in an all or nothing response (Allrich, 1994). During follicle development increasing concentrations of oestradiol are produced and secreted mainly from the dominant follicle (Staigmiller et al., 1982). This increases follicular oestradiol 3-4 days before oestrus, causes circulating concentrations of oestradiol to increase (Roelofs et al., 2010), acting at the level of the hypothalamus to trigger a series of programmed neurological events that result in behavioural oestrus (Reames et al., 2010). Other centres in the brain also trigger the closely related LH surge which is required for ovulation and occurs 28-32 hours after oestrus (Walker et al., 1996). In contrast progesterone from the CL also controls oestrus by inhibiting GnRH and LH pulses which reduces oestradiol concentration (Smith and Jennes, 2001). Often in the post partum period there is a silent oestrus which involves ovulation without overt oestrous expression. This is thought to be caused by high oestradiol levels following gestation inducing a refractory period (Allrich, 1994). Oestrus can be affected through the duration of progesterone and progesterone amplitude during the luteal phase which can influence the increase in levels of oestradiol. It is suggested that this occurs by the influence of progesterone on the neural mechanisms controlling release of GnRH, influencing the elements targetted by oestradiol to induce the preovulatory LH surge in the ewe (Skinner et al., 2000). It has also been reported in the ewe that previous progesterone exposure can affect the intensity of oestrous expression (Fabre-Nys and Martin, 1991). However, in the cow these mechanisms differ and oestrus can occur without progesterone exposure. Even with low levels of 16

oestradiol, the LH surge and ovulation can still occur; demonstrating that hypothalamic sensitivity to LH and oestradiol differs and could also differ between individual cows (Reames et al., 2010). Oestradiol is thought to alter neuronal networks, including dendritic connections between cells and receptors, and neurotransmitter release in order to facilitate oestrous expression (Boer et al., 2009). The shift to oestradiol stimulation causes elevated GnRH receptor gene expression in gonadotroph cells which can result in increasing LH pulses stimulated by synthesis and secretion of GnRH (Boer et al., 2009). The increased oestradiol has a self amplifying effect, stimulating the expression of oestrogen receptors in the brain (Pfaff, 2005). Therefore oestradiol indirectly synchronises mating and ovulation. Oestradiol affects certain areas in the brain in order to regulate female sexual behaviour. Specific areas reported to be involved in behavioural oestrus are the arcuate nucleus, ventromedial nucleus, the preoptic area of the hypothalamus and in particular the hippocampus and amygdala are related to behavioural oestrus (Molenda-Figueira et al., 2006). Oestradiol and other hormones, for example IGF-1 and GnRH, can cause up and down regulation of a number of genes in these brain areas known to be involved in oestrous behaviour (reviewed by Boer et al., 2009). Some preliminary work carried out investigating gene expression in the brain at oestrus compared to luteal phase Holstein Friesian heifers has found that oestrous behaviour may be linked to different patterns of gene expression in the pituitary gland, hypothalamus, amygdala and ventral tegmental area (Beerda et al., 2008). The majority of the research into the genomic control of oestrus has been carried out in rodents where increased oestrogen receptor expression in hypothalamic areas at oestrus have been reported (Pfaff et al., 2008) resulting in expression of genes to facilitate oestrous behaviour, stimulating behavioural oestrus and mediating neurotransmission resulting in oestrus (reviewed by Boer et al., 2009). However, parallels can be drawn between the brain areas involved in behavioural oestrus between rodents and ruminants (Stormshak and Bishop, 2008). Increased oestrogen receptor expression has also been linked to an increase in locomotion (Smith and Jennes, 2001), which is a similar oestrus response to that seen in cattle (Kiddy, 1977). 17

on average <strong>in</strong> Holste<strong>in</strong>s (Dransfield et al., 1998). The percentage <strong>of</strong> <strong>cows</strong><br />

actually st<strong>and</strong><strong>in</strong>g <strong>to</strong> be mounted has also decl<strong>in</strong>ed from 80% <strong>to</strong> 50%<br />

(Dobson et al., 2008). This results <strong>in</strong> only 50% <strong>of</strong> <strong>cows</strong> be<strong>in</strong>g observed <strong>in</strong><br />

<strong>oestrus</strong> (Van Eerdenburg et al., 2002). Hence <strong>detection</strong> <strong>of</strong> <strong>oestrus</strong> is<br />

<strong>in</strong>creas<strong>in</strong>gly more difficult <strong>and</strong> <strong>to</strong> improve the decl<strong>in</strong>e <strong>in</strong> fertility oestrous<br />

<strong>detection</strong> rates must be improved.<br />

1.3.1.1 Endocr<strong>in</strong>e, Neural <strong>and</strong> Genomic Changes Associated with Oestrous<br />

Behaviour<br />

Oestradiol is the key regula<strong>to</strong>r that synchronizes the endocr<strong>in</strong>ological <strong>and</strong><br />

behavioural events <strong>to</strong> drive <strong>oestrus</strong>; result<strong>in</strong>g from the action <strong>of</strong> ovarian<br />

steroids on behavioural centres <strong>in</strong> the bra<strong>in</strong> (Roel<strong>of</strong>s et al., 2010). The<br />

production <strong>of</strong> oestradiol from the ovary synchronises mat<strong>in</strong>g <strong>and</strong> ovulation;<br />

ris<strong>in</strong>g above its threshold <strong>in</strong> an all or noth<strong>in</strong>g response (Allrich, 1994).<br />

Dur<strong>in</strong>g follicle development <strong>in</strong>creas<strong>in</strong>g concentrations <strong>of</strong> oestradiol are<br />

produced <strong>and</strong> secreted ma<strong>in</strong>ly from the dom<strong>in</strong>ant follicle (Staigmiller et al.,<br />

1982). This <strong>in</strong>creases follicular oestradiol 3-4 days before <strong>oestrus</strong>, causes<br />

circulat<strong>in</strong>g concentrations <strong>of</strong> oestradiol <strong>to</strong> <strong>in</strong>crease (Roel<strong>of</strong>s et al., 2010),<br />

act<strong>in</strong>g at the level <strong>of</strong> the hypothalamus <strong>to</strong> trigger a series <strong>of</strong> programmed<br />

neurological events that result <strong>in</strong> behavioural <strong>oestrus</strong> (Reames et al.,<br />

2010). Other centres <strong>in</strong> the bra<strong>in</strong> also trigger the closely related LH surge<br />

which is required for ovulation <strong>and</strong> occurs 28-32 hours after <strong>oestrus</strong><br />

(Walker et al., 1996). In contrast progesterone from the CL also controls<br />

<strong>oestrus</strong> by <strong>in</strong>hibit<strong>in</strong>g GnRH <strong>and</strong> LH pulses which reduces oestradiol<br />

concentration (Smith <strong>and</strong> Jennes, 2001).<br />

Often <strong>in</strong> the post partum period there is a silent <strong>oestrus</strong> which <strong>in</strong>volves<br />

ovulation without overt oestrous <strong>expression</strong>. This is thought <strong>to</strong> be caused<br />

by high oestradiol levels follow<strong>in</strong>g gestation <strong>in</strong>duc<strong>in</strong>g a refrac<strong>to</strong>ry period<br />

(Allrich, 1994). Oestrus can be affected through the duration <strong>of</strong><br />

progesterone <strong>and</strong> progesterone amplitude dur<strong>in</strong>g the luteal phase which<br />

can <strong>in</strong>fluence the <strong>in</strong>crease <strong>in</strong> levels <strong>of</strong> oestradiol. It is suggested that this<br />

occurs by the <strong>in</strong>fluence <strong>of</strong> progesterone on the neural mechanisms<br />

controll<strong>in</strong>g release <strong>of</strong> GnRH, <strong>in</strong>fluenc<strong>in</strong>g the elements targetted by<br />

oestradiol <strong>to</strong> <strong>in</strong>duce the preovula<strong>to</strong>ry LH surge <strong>in</strong> the ewe (Sk<strong>in</strong>ner et al.,<br />

2000). It has also been reported <strong>in</strong> the ewe that previous progesterone<br />

exposure can affect the <strong>in</strong>tensity <strong>of</strong> oestrous <strong>expression</strong> (Fabre-Nys <strong>and</strong><br />

Mart<strong>in</strong>, 1991). However, <strong>in</strong> the cow these mechanisms differ <strong>and</strong> <strong>oestrus</strong><br />

can occur without progesterone exposure. Even with low levels <strong>of</strong><br />

16

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