404
Reproduction, Fertility and Development
A. Menchaca and E. Rubianes
et al.
1999). Follicular waves have also been demonstrated inewes during both the seasonal anoestrus (Souza
et al.
1996;Bartlewski
et al.
1998) and the transitional period to the breeding season (Ravindra and Rawlings 1997). A wave isdefined as the emergence of a group of small antral folliclesfrom which commonly one or two follicles reach a diameter of 5mm or more.According to different authors the number of follicular waves per cycle ranges between two and fivewith large variability in both sheep and goats.The most com-monly found pattern in ewes is three follicular waves duringa cycle (Leyva
et al.
1998; Gibbons
et al.
1999; Evans
et al.
2000; Viñoles
et al.
2000) and 4 follicular waves in goatswithaninter-ovulatorycycleofnormallength(19to22days)(Ginther and Kot 1994; de Castro
et al
. 1999; Schwarz and Wierzchos 2000; Menchaca and Rubianes 2002). The inter-wave interval is the period between the emergence of thelargest follicle (3mm in diameter) of two consecutive waves,whichisaroundfourtosevendaysinbothspecies.Thedayof emergence for each wave is variable in both ewes and goats,anddependsonthenumberofwavesineachcycle.Predictionof the day of emergence of each follicular wave is thereforeverydifficult,withtheexceptionofthefirstwaveofthecycle(Wave 1). All studies in sheep and goats agree that Wave 1emerges around the day of ovulation (Day 0) of the previouscycle.The characteristics of follicular waves have been recentlyreviewed (ewes: Evans 2003; goats: Rubianes and Menchaca2003)andsomeofthemorefrequentlyobservedaspectscom-mon to both species are: (
1
) at least one follicle attaining
≥
5mmdiameterisobservedperwave;(
2
)thelargestfollicleofeachwavegrowsfor5–7days,withagrowthrateofaround 1mmday
−
1
;(
3
)themaximumdiameterofthelargestfollicleof a wave differs between waves; (
4
) as the luteal phase pro-gresses and the serum progesterone concentrations increase,follicular turnover is facilitated and the inter-wave intervalsare shorter than during the early luteal phase; (
5
) during themid–late luteal phase the follicles that do not grow beyond 4 mm are often not part of the wave phenomenon and it has been suggested that they represent an underlying dynamic pool; (
6
) most of the follicles that are the largest on the dayof luteolysis are the ones that ovulate; (
7
) in most doubleovulatory cycles the ovulatory follicles emerge as part of thesame follicular wave, but in a few cases also as a part of dif-ferent waves; and (
8
) the double ovulations occur within 24hfrom each other.
Progesterone and follicular growth
The insertion of an intravaginal progesterone device (CIDR-G
®
) in anoestrous goats results in a sharp increase in serum progesterone concentration (
>
5ngmL
−
1
) for three or four days, which is higher than those observed during the physio-logicalmid–latelutealphase(Rubianes
etal.
1998).Aftersixdaysoftreatment,serumprogesteroneconcentrationsdeclineto subluteal levels (2ngmL
−
1
) and remain low until thedevice is withdrawn (i.e. Day 12). Similar pharmacokineticresults have been reported using intravaginal sponges con-taining medroxiprogesterone (MAP) (Greyling
et al.
1994)and fluorogestone (FGA) (Gaston-Parry
et al.
1988) in ewes.Therefore, the progesterone serum profile induced by thesetreatments is the opposite of that observed during the nor-mal oestrous cycle, when progesterone concentrations are atfirst low and then increase until luteolysis. The effects of progesterone on follicular dynamics have been studied insheep (Johnson
et al.
1996; Rubianes
et al.
1996; Leyva
et al.
1998; Viñoles
et al.
1999) and goats (Menchaca and Rubianes 2002). Special attention has been given to theeffects of induced subluteal progesterone concentrations onfollicular health. In ewes, subluteal progesterone levels pro-motedexcessivegrowthandpersistenceofthelargestfollicle(Viñoles
et al.
1999), increasing the age of the ovulatoryfollicles(Johnson
etal.
1996).Twelve-dayprogestogentreat-mentsinduceovulationofagedfolliclesinthisspecies(Flynn
et al.
2000;Viñoles
et al.
2001). In cattle, the ovulation of anaged follicle is followed by low fertility (Savio
et al.
1993;Stock and Fortune 1993; Mihm
et al.
1994; Austin
et al.
1999), apparently because an early resumption of meiosisoccurs in the oocyte (Revah and Butler 1996). Although asimilar detrimental effect of a long progestogen treatment(Viñoles
et al.
2001) or the use of a depleted progesteronedevice(UngerfeldandRubianes1999)ontheconceptionratehas been observed in the ewe, this issue is still a matter of debate (Evans 2003).Progesterone has a negative feedback influence on LHsecretion and pulsatility (Goodman and Karsh 1980) and LH pulses regulate the final growth of antral follicles (McNeilly
et al.
1991). A consequence of low levels of serum pro-gesterone is an increase of LH pulse frequency, which isassociated with an increment in the diameter of the largest(dominant) follicle. Normally, when the progesterone con-centrations are very low (i.e. after luteolysis), positive feed- back is established between oestradiol from the growingdominantfollicleandGnRHandLHfromthehypothalamus– pituitary-axis. This positive loop ends with the preovulatoryLHsurgeandovulationofthefollicle.Sublutealprogesteroneconcentrations increase the LH pulses frequency, but theLH surge does not occur. In consequence the largest follicle persists and its dominance is prolonged.Highprogesteroneconcentrations,incontrast,haveapos-itive effect on follicular turnover increasing the number of young large follicles with the potential to ovulate. Supra-luteal progesterone levels affect the dominance of the largestfollicle of Wave 1, inducing early regression and accelerat-ing the emergence of the next follicular wave, which resultsin the ovulation of a healthy young follicle (ewes: Rubianes
et al.
1996; goats: Menchaca and Rubianes 2002). Overall,thesefindingssuggesttheconceptthatahigh-levelshort-term progestogentreatmentcouldbebetteratcontrollingfollicular
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