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Clinical Biochemistry of Domestic Animals (Sixth Edition) - UMK ...

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646<br />

Chapter | 21 <strong>Clinical</strong> Reproductive Endocrinology<br />

through hysterectomy ( Stabenfeldt et al. , 1974b ). Removal<br />

<strong>of</strong> the uterus from these species during the luteal phase<br />

prolongs luteal activity. It is well established that the uterus<br />

in these species synthesizes and releases PGF 2 α , which<br />

causes the CL to regress ( McCracken et al. , 1972 ).<br />

The temporal release patterns <strong>of</strong> PGF 2 α ( Fig. 21-4 ),<br />

usually in a pulsatile mode lasting a few hours, have been<br />

described in the ewe ( Barcikowski et al. , 1974 ; Harrison<br />

et al. , 1972 ), sow ( Gleeson et al. , 1974 ), doe ( Fredriksson<br />

et al. , 1984 ), and cow ( Nancarrow et al. , 1973 ). Some <strong>of</strong><br />

the problems involved in determining PGF 2 α (e.g., a short<br />

half-life and formation by platelets at collection) can be<br />

avoided if the main blood plasma metabolite, 15-keto-<br />

13,14-dihydro-PGF 2 α , is determined. Data are available on<br />

the patterns <strong>of</strong> the metabolite during luteolysis in the cow<br />

( Kindahl et al. , 1976 ), ewe ( Peterson et al. , 1976 ), mare<br />

( Neely et al. , 1979 ), doe ( Fredriksson et al. , 1984 ), and<br />

sow ( Shille et al. , 1979a ).<br />

Regression <strong>of</strong> CL is usually accomplished within 48h<br />

following the onset <strong>of</strong> the prostaglandin release. It is likely<br />

that estrogens, presumably <strong>of</strong> ovarian follicle origin, initiate<br />

PGF 2 α release. Estrogen also initiates the formation <strong>of</strong><br />

endometrial oxytocin receptors ( McCracken et al. , 1984 ),<br />

which in sheep are important for pulsatile synthesis and<br />

release <strong>of</strong> PGF 2 α in that oxytocin can initiate the release <strong>of</strong><br />

PGF 2 α ( Sharma and Fitzpatrick, 1974 ). In ruminants, the CL<br />

is able to synthesize oxytocin and oxytocin release from the<br />

CL in response to PGF 2 α , which, in turn, initiates the synthesis<br />

and release <strong>of</strong> PGF 2 α by the uterus; this is the basis<br />

for the pulsatile secretion <strong>of</strong> PGF 2 α (for a review, see Flint<br />

et al. [1992] , and Whates and Denning-Kendall [1992] ). In<br />

the dog and cat, PGF 2 α does not appear to be involved in<br />

luteolysis, although its precise role is still uncertain.<br />

C . Early Pregnancy<br />

Modification <strong>of</strong> PGF 2 α release is essential for the establishment<br />

<strong>of</strong> pregnancy in the species (cow, ewe, mare, and sow)<br />

in which this compound serves as the luteolysin ( Kindahl<br />

et al. , 1976 ; Nett et al. , 1976 ; Shille et al. , 1979a ). The<br />

rapid elongation <strong>of</strong> fetal membranes, which precedes the<br />

critical time <strong>of</strong> the initiation <strong>of</strong> luteal regression by about<br />

3 days in the nonpregnant animal, appears to be important<br />

for modifying prostaglandin release. Fredriksson et al.<br />

(1984) in goats and Zarco et al. (1988) in sheep have shown<br />

that the main change in PGF 2 α synthesis and release, nonpregnant<br />

versus pregnant animal, involves a continuous<br />

mode <strong>of</strong> secretion, not pulsatile. In fact, PGF 2 α concentrations<br />

increase at the onset <strong>of</strong> pregnancy, but the release is<br />

continuous, which prolongs luteal activity.<br />

Maternal recognition <strong>of</strong> pregnancy in the cow, ewe, doe,<br />

and sow involves mechanisms that alter the prostaglandin<br />

release to protect the CL from luteolysis. In the cow,<br />

ewe, and doe, embryonic interferon tau (IFN- τ ), which<br />

is secreted between days 10 and 21 to 24 <strong>of</strong> pregnancy<br />

Estrone sulfate<br />

Mare<br />

Sow<br />

0 20 40 60 80 100<br />

Days <strong>of</strong> pregnancy<br />

Estrone sulfate<br />

Doe<br />

Ewe<br />

Cow<br />

0 40 80 120 160 200<br />

Days <strong>of</strong> pregnancy<br />

FIGURE 21-6 Blood levels <strong>of</strong> conjugated estrogens (mainly estrone<br />

sulfate) during early pregnancy in the mare, sow, doe, ewe, and cow. Data<br />

derived from Kindahl et al. (1982) (mare); Robertson and King (1974)<br />

(sow); Chaplin and Holdsworth (1982) (doe); Tsang (1978) (ewe); and<br />

Gaiani et al. (1982) (cow).<br />

by trophectoderm cells <strong>of</strong> the blastocysts ( Roberts et al. ,<br />

1990 ), inhibits uterine PGF 2 α pulses ( Spencer et al. , 1999 ;<br />

Thatcher et al. , 2001 ). IFN-τ is secreted into the uterine<br />

lumen and inhibits the expression <strong>of</strong> estrogen and oxytocin<br />

receptors, thus blocking the episodic uterine PGF 2 α secretion<br />

and luteolysis (for a review, see Spencer et al. [2004] ).<br />

In the sow, the maternal recognition <strong>of</strong> pregnancy is controlled<br />

by conceptus-derived estrogens ( Geisert et al. , 1982 ;<br />

Perry et al. , 1976 ) (Fig. 21-6 ).<br />

Modification <strong>of</strong> the release pattern <strong>of</strong> PGF 2 α (pulsatile<br />

to continuous) by the luteotropic products from the<br />

conceptus and uterus is probably the most important factor<br />

that allows luteal activity to continue. The net result<br />

is that luteal activity is extended in the cow, ewe, mare,<br />

and sow beginning at about 14 days following ovulation.<br />

Modification <strong>of</strong> PGF 2 α release appears not to be important<br />

for the establishment <strong>of</strong> pregnancy in the dog and cat.<br />

D . Pregnancy and Parturition<br />

The presence <strong>of</strong> a CL is necessary for the maintenance <strong>of</strong><br />

pregnancy in a vast majority <strong>of</strong> cows ( Estergreen et al. ,<br />

1967 ). The pig also requires luteal support throughout<br />

gestation (see Bazer et al. , 1979 ). In the ewe, the presence<br />

<strong>of</strong> CL is required for the first 50 to 60 days <strong>of</strong> gestation

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