UTERINE INVOLUTION: ITS ROLE IN REGULATING POSTPARTUM INTERVALS G. H. KIRACOFE J ANIM SCI 1980, 51:16-28.

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U T E R I N E I N V O L U T I O N : I T S R O L E IN R E G U L A T I N G POSTPARTUM INTERVALS
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G. H, K I R A C O F E Kansas State University, Manhattan Summary AN involuting uterus is a temporary barrier to fertility in cows, ewes and sows. Parturition is followed by a period when conception is not possible: about 1 week in sows and about 3 weeks in cows and ewes. Estrus and ovulation seldom occur together during this period and, if fertilization occurred and the embryo reached the uterus, placentation would be virtually impossible. T h e period of no fertility is followed by 2 to 3 weeks when fertility is possible, but not optimal. T h e extent to which the involuting uterus contributes to infertility during the second period is difficult to determine. Embryonic mortality in sows bred during this latter period appears to be a major cause of reduced litter size, so changes in uterine environment associated with involution may be essential for optimum fertility. Conception rate is lower up to 4 0 days after parturition than later in cows and ewes. Uterine involution appears not to be a barrier to fertility after 3 to 4 weeks postpartum in sows or 5 to 6 weeks postpartum in cows and ewes unless delayed b y inflammation or infection. (Key Words: Uterine Involution, Postpartum Intervals, Postpartum Conception, Short Estrous Cycles, Cattle, Sheep, Swine.) Introduction Processes involved in the return of the postpartum uterus to a state capable of supporting another conceptus after parturition are complex and the interrelationship of the uterus with the ovaries is not understood. That the time required for uterine involution limits postpartum fertility under current livestock management systems is questionable. Considering only the basics of normal uterine involution, the uterus rarely is unprepared for pregnancy when ovulation and estrus occur together, fertilization has taken place, and the embryo is ready for placentation. Contribution 79-379-J, Dept. of Animal Sciences and Industry, Kansas Agr. Exp. Sta., Manhattan.
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Uterine size has not been a good indicator of subsequent fertility, but more subtle anatomical changes may continue well beyond the dramatic changes in size noted shortly after parturition. Subsequent fertility is impaired when uterine involution is complicated by pathological conditions induced by dystocia, retained fetal membranes, metritis or other disorders. However, whether these factors directly affect postpartum endocrinology or reduce fertility by providing an improper milieu for spermatozoa and embryos has not been established. Lack of a precise definition of uterine involution prevents direct comparisons of many data. Despite voluminous data concerning uterine involution and postpartum fertility, specific studies correlating the two are lacking. This review describes normal processes of uterine involution in cows, ewes and sows. Factors related to normal uterine involution are considered to determine if the uterus limits postpartum fertility. Bovine Species The Process of Uterine Involution. Before 1950, postpartum changes in the bovine uterus received little attention. Studies were limited primarily to pathological situations and to palpation per rectum to determine uterine size, tone and positional changes; a few uterine weights and measures were recorded. Pioneering studies included those of Hilty ( 1 9 0 8 ) , Boyd ( 1 9 2 5 ) , Casida and Venzke ( 1 9 3 6 ) and Casida and Wisnicky ( 1 9 5 0 ) . Rasbeck ( 1 9 5 0 ) reviewed much of the early research and later reviews on uterine involution were done by Casida et al. ( 1 9 6 8 ) , Morrow et al. ( 1 9 6 9 ) and Moller ( 1 9 7 0 a ) . Since 1950, regression of uterine size, factors affecting postcalving uterine changes, and descriptions of tissue sloughing, vascular changes and tissue regeneration have been reported. T h e complexity of the involutionary process and the difficulty in determining exactly when uterine involution is complete has

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U T E R I N E I N V O L U T I O N AND P O S T P A R T U M made comparison of data among investigators difficult. Conditions that signify complete uterine involution are nebulous. Casida et al. ( 1 9 6 8 ) defined uterine involution as complete when the uterus returned to its normal nonpregnant position and when the two horns were similar in diameter and showed normal consistency and tonus. Palpation estimates differ widely because of methods used to measure the uterus and inherent variation among individuals. Gier and Marion ( 1 9 6 8 ) reasoned that a variety of histological and physical changes must take place in the uterus before involution could be termed complete. They described uterine involution as having three overlapping processes: ( 1 ) reduction in size, ( 2 ) loss of tissue, and ( 3 ) repair. Reduction in Size. Estimates of the time required for complete reduction of uterine size after calving range from 18 (Rasbeck, 1 9 5 0 ) to 5 6 days (Wiltbank and Cook, 1 9 5 8 ) . Estimated intervals from parturition to complete uterine involution, ovulation and estrus for various breeds are listed in table 1. Palpable changes in uterine size, tone and position after calving were described by Rasbeck ( 1 9 5 0 ) , Buch et al. ( 1 9 5 5 ) , Morrow et al. ( 1 9 6 6 ) , Bhalla et al. ( 1 9 6 6 ) , Moller ( 1 9 7 0 b ) , Mohamed ( 1 9 7 4 ) and El-Fouly et al. ( 1 9 7 6 ) . Early reduction of uterine size after calving results from vasoconstriction (Morrow, 1 9 6 9 ) and peristaltic contractions that persist for several days (Jordan, 1952; Venable and McDonald, 1 9 5 8 ) . Weight and size of the uterus after calving decrease in similar patterns. Riesen ( 1 9 6 8 ) and Gier and Marion ( 1 9 6 8 ) , who weighed uteri from cows after calving, reported the uterine weight at calving was approximately 9 kg and was reduced to approximately 3 kg by 10 days, to 1 kg by 20 to 3 0 days and to 7 5 0 g by 5 0 days postpartum. Fluid is present in the uterus after calving (Johanns et al, 1967; Gier and Marion, 1 9 6 8 ; Morrow, 1969; Morrow et al, 1 9 6 9 ) . T h e maximum amount of lochia ( 1 , 0 0 0 to 1,600 m l ) is present during the first 4 8 hr and then decreases to 4 0 0 to 100 ml by day 8 and is absent by 14 to 24 days postpartum. Loss of Tissue. Gier and Marion ( 1 9 6 8 ) described early uterine changes. Shedding of fetal membranes appeared to be associated with peristaltic muscular contractions. Reduction in uterine size progressed in a negative logarithmic manner so that the postgravid horn diameter decreased by 50% by day 5 . Loss of tissue fluids appeared to be a major factor in early size reduction. Endometrial edema,

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which occurred during the first day postcalving, receded b y day 8. Rasbeck ( 1 9 5 0 ) , Gier and Marion ( 1 9 6 8 ) and Wagner and Hansel ( 1 9 6 9 ) described sloughing and regenerative changes in histological preparations from excised uteri at various times postpartum. J o hanns et al. ( 1 9 6 7 ) , through radiographic studies of postpartum uteri, showed degenerative changes in the epithelium as well as changes in the circulatory system. Morrow ( 1 9 6 9 ) reported that the caruncular stalk disappears, the superficial layer of the caruncle dissolves and uterine lochia form. T h e bulk of the caruncle had dissolved by 10 days after calving. Johanns et al. ( 1 9 6 7 ) described constriction of the caruncular arteries and decreasing caruncular blood supply during these 10 days. Endometrial changes resulting in edema, then regression and folding, and a general "stalked" appearance of the caruncle were described by Rasbeck ( 1 9 5 0 ) and Gier and Marion ( 1 9 6 8 ) . Sloughing of the necrotic tissue progressed to an extent that material of the caruncle came off in chunks b y day 5 and most of the necrotic layer was gone by 10 days. By 15 days all of the caruncular mass involved in the placentome was sloughed, and by day 19 the arterioles within and beyond the stratum compactum had disappeared leaving the caruncles relatively smooth (Gier and Marion, 1 9 6 8 ) . T h e deciduum of the caruncle sloughed just basal to the crypts and regeneration of the epithelium occurred from the margin of the slough (Wagner and Hansel, 1 9 6 9 ) . Repair. Several descriptions of tissue repair have been reported. T h e intercaruncular areas never were devoid of epithelium, and both degenerative and regenerative epithelial cells were observed in the intercaruncular areas before day 15, but after day 15 no degenerative cells were observed (Archbald et al, 1 9 7 2 ) . Epithelium was reestablished over the caruncle 9 to 2 0 days after parturition (Archbald et al, 1972; Rasbeck, 1950; Roberts, 1 9 5 6 ) . Gier and Marion ( 1 9 6 8 ) found that epithelial regeneration was not complete b e fore day 2 5 ; or for at least 10 days after sloughing had ceased. Caruncle sites were covered with epithelium b y postpartum days 10 to 2 0 in the previously nongravid horn, but not until 20 to 3 0 days in the horn which had been gravid (Riesen, 1 9 6 8 ) . Gier and Marion ( 1 9 6 8 ) considered involution to be complete and the uterus to be restored to a cycling condition by 4 0 to 6 0 days when the caruncle had regressed to a

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KIRACOFE T h e net effect of nutrition on uterine involution appeared to be minimal in the absence of specific deficiencies. A remarkable feature of uterine involution in the rat is the loss of collagen (Harkness and Harkness, 1954; Harkness and Moralee, 1956; Woessner, 1 9 6 9 ) ; 85% of the total collagen of the term uterus was removed in the first 4 days postpartum. Similar research with large animals on this aspect of involution is lacking. Since relaxin can induce breakdown of collagen in some circumstances (Frieden and Hisaw, 1 9 5 3 ) , it would be of interest to determine if relaxin plays a role in uterine involution. Kiracofe et al ( 1 9 7 3 a ) did not alter the involution rate in postpartum cows by intrauterine infusion of collagenase. Treating the uterus with enzymes, antibiotics or antibacterial compounds failed to hasten uterine involution, but they frequently prolonged the time to involution (Homan, 1969; Kiracofe et al, 1973b; Oxender and Seguin, 1976; Seguin et al, 1 9 7 4 ) . A profound effect on the size of the postpartum uterus resulted from injecting cows with GnRH (gonadotropin releasing hormone) 14 days after calving (Britt et al, 1974a; Fernandes et al, 1 9 7 8 ) . T h e GnRH accelerated reduction in uterine size between 14 and 24 days postpartum. No beneficial effect on conception was associated with decreased uterine size after GnRH. An increase in serum progesterone concentrations after GnRH, noted by Fernandes et al. ( 1 9 7 8 ) , may have accelerated uterine involution by affecting uterine blood flow and water imbibition. Riek et al. ( 1 9 7 7 ) showed that GnRH partially synchronized cows in the early puerperium, but the condition of the postpartum uterus appeared to affect the ability of GnRH to initiate cyclic progesterone production. Other postpartum hormone treatments have hastened onset of estrous cycles and subsequent conception. Shorter intervals from calving to conception after administration of progesterone or synthetic progestogens during the postpartum period have been reported by Fosgate et al. ( 1 9 6 2 ) , Norwood ( 1 9 6 3 ) , Foote and Hunter ( 1 9 6 4 ) , Saiduddin et al. ( 1 9 6 8 ) , Boyd ( 1 9 7 0 ) , Britt et al ( 1 9 7 2 ) , Huertas-Vega et al. ( 1 9 7 2 ) and Britt et al. ( 1 9 7 4 b ) . Likewise, Casida and Wisnicky (1950), Norwood ( 1 9 6 3 ) , Ulberg and Lindley ( 1 9 6 0 ) , Foote and Hunter ( 1 9 6 4 ) and Brown et al. ( 1 9 7 2 ) reported some beneficial results with either estrogens or synthetic estrogens alone or in combination with progestogens. However, ef-

smooth, oblong, epithelial covered, avascular knob. Caruncles which had been in the proximity of the fetus required the longest interval to return to normal. Obviously, involution continues past the time that uterine regression is detectable by palpation, but the degree of involution required to support another pregnancy has not been established. Many factors delay the normal process of involution: uterine infections, retained fetal membranes (Boyd, 1 9 2 5 ; Rasbeck, 1950; Buch et al, 1 9 5 5 ; Norwood, 1 9 6 3 ; Gier and Marion, 1 9 6 8 ; Marion et al., 1968; Tennant and Peddicord, 1 9 6 8 ; Morrow, 1 9 6 9 ) and other periparturient diseases (see reviews by Morrow et al, 1969; Moller, 1 9 7 0 a ) . Factors Influencing the Time to Involution. Suckling, in contrast to nonsuckling, has been reported to promote uterine involution (Casida et al., 1 9 6 8 ; Riesen, 1 9 6 8 ) , delay uterine involution (Wiltank and Cook, 1 9 5 8 ) or have little effect on the rate of uterine involution (Wagner and Hansel, 1969; Moller, 1970b; Oxenreider and Wagner, 1 9 7 1 ; El-Fouly et al., 1 9 7 6 ) . However, limited or no suckling hastens resumption of estrous cycles (Wiltbank and Cook, 1 9 5 8 ; Moller, 1970b; Oxenreider and Wagner, 1 9 7 1 ; Short et al., 1972; Randel and Welker, 1977; Radford et al, 1 9 7 8 ; Wettemann et al., 1 9 7 8 ) . Generally, uterine involution takes a few days longer in pluriparous cows than in primiparous cows (Rasbeck, 1 9 5 0 ; Buch et al., 1 9 5 5 ; Quevedo et al, 1 9 6 5 ; Morrow et al, 1966; Marion and Gier, 1 9 6 8 ; Araujo et al, 1 9 7 4 ) . However, Tennant et al. ( 1 9 6 7 ) and El-Fouly et al. ( 1 9 7 6 ) reported that age or number of previous pregnancies had no effect on involution. Several investigators have reported seasonal effects on rate of uterine involution. Norwood ( 1 9 6 3 ) , Marion et al. ( 1 9 6 8 ) , ElFouly et al. ( 1 9 7 6 ) and Gospodinov and Rostov ( 1 9 7 6 ) found slower involution in the fall than in the spring. Energy and protein level of the diet only slightly affect the rate of uterine involution but drastically affect the interval from parturition to resumption of estrous cycles (Wiltbank et al, 1962; Dunn et al., 1969; Clemente et al, 1 9 7 8 ) . Uteri of cows fed high energy diets after calving involuted 3 days sooner than uteri of cows fed moderate energy diets, and uteri of Angus cows involuted 3 days sooner than uteri of Hereford cows (Dunn, 1 9 6 4 ) . Kiracofe et al. ( 1 9 6 9 ) found no significant difference in uterine involution rates in cows fed low or high levels of energy or protein.

UTERINE INVOLUTION AND POSTPARTUM fects of exogenous steriods on conception vary widely and the beneficial effects of exogenous steriods cannot be related to hastened uterine involution. Marion et al. ( 1 9 6 8 ) showed that progesterone actually lengthened the period of uterine involution. Therefore, if exogenous steriod hormones prove beneficial to postpartum conception, it is unlikely that success will be attributed to an effect on uterine involution. The postpartum uterus contains many types of bacteria (Dawson, 1950; Wulf and Dracy, 1952; Lindley, 1954; Gunter et al, 1 9 5 5 ; Eliot et al, 1 9 6 8 ; Griffin et al, 1 9 7 4 ; Studer and Morrow, 1 9 7 8 ) . Bacteria of the early postpartum uterus do not appear to affect involution rate and may or may not b e associated with subsequent fertility (Graden et al., 1 9 6 8 ; Fuquay et al, 1 9 7 5 ) . Kiracofe et al ( 1 9 7 3 a ) showed no effect on uterine involution and fertility from innoculating the early postpartum uterus with lactobacillus acidophilus. Involution and Resumption of Cycles. F e w direct relationships can be made between the rate of uterine involution and subsequent fertility. Albrechtsen ( 1 9 1 7 ) found that if uterine involution required more than 4 weeks the onset of first estrus was delayed. However, Buch et al. ( 1 9 5 5 ) found only a slight association between interval to first estrus and degree of uterine involution. Other researchers have found no relationship between uterine involution and interval from calving to first estrus or conception of clinically normal cows (Foote et al, 1960a; Perkins and Kidder, 1963; Dunn, 1964; Tennant and Peddicord, 1 9 6 8 ; Kiracofe et al, 1 9 6 9 ) . Some conditions can delay uterine involution, resumption of cycling and conception (see review by Morrow, 1969; Studer and Morrow, 1 9 7 8 ) . An abnormal uterus may delay follicle development and ovulation, but it is doubtful that a normal involuting uterus plays a role in reestablishing estrous cycles. Pituitary gonadotropin content was not altered in cows hysterectomized within 2 4 hr after calving (Foote, 1 9 7 1 ) . Vital information in regard to the role of the uterus in initiating postpartum estrous cycles might be obtained by observing the intervals to ovulation and estrus in cows hysterectomized at calving. It had been proposed that the corpus luteum of pregnancy might delay the onset of the first postpartum estrus (McNutt, 1 9 2 7 ) ; however, Morrow ( 1 9 6 9 ) showed that the corpus luteum of pregnancy is nonfunctional after parturition. Apparently the ovaries do not affect the rate of uterine involution because Marion et al.

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( 1 9 6 8 ) showed uterine involution time was unaltered in ovariectomized cows. When beef heifers were subjected to hypophysial stalk transection in middle or late pregnancy, pregnancy was maintained and live calves were born, and then the uterus involuted to a size similar to that of a prepuberal heifer (Anderson et al, 1 9 7 9 ) . Cows can conceive before uterine involution would b e expected to have been complete (Buch et al, 1 9 5 5 ; Perkins and Kidder, 1 9 6 3 ; Moller, 1 9 7 0 a ; Odde and Kiracofe, 1 9 7 8 ) . Olds and Cooper ( 1 9 7 0 ) found no difference in conception rates of dairy cows bred 4 0 to 6 0 days postpartum and those bred later, although uterine involution would not be complete in all cows by day 4 0 . Tudorascu ( 1 9 6 9 ) and Casida et al, ( 1 9 6 8 ) found no relationship between the postpartum interval to breeding and embryonic mortality in animals bred after day 3 0 postpartum. Conception rates appear to b e rather low at less than 3 0 days postpartum (Olds et al., 1949; Van Demark and Salisbury, 1950; Norwood, 1 9 6 3 ; Perkins and Kidder, 1 9 6 3 ; Odde and Kiracofe, 1 9 7 8 ; also see review by Casida et al., 1 9 6 8 ) . Perkins and Kidder ( 1 9 6 3 ) reported conception rate to be unaffected b y involutionary state after directly comparing conception rates of cows in which uterine involution was complete and cows in which uterine involution was not complete. Another approach to determine factors limiting early postpartum conception is to induce ovulation and inseminate at various postpartum intervals. Henderson ( 1 9 7 8 ) bred cows after progestogen and estrogen treatment ( a 6 mg norgestomet implant for 9 days and an injection of 6 mg estradiol valerate and 3 mg norgestomet at the time of implanting). Conception rates did not differ among cows bred less than 4 2 days postpartum and those bred later. Further attempts to breed cattle or transfer fertilized ova at specific postpartum intervals and relate pregnancy maintenance to the degree of involution at the time of insemination or transfer are needed.
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Follicular activity begins soon after parturition (Morrow, 1969; Wagner and Hansel, 1969; Moller, 1 9 7 0 b ) . T h e corpus luteum of pregnancy is palpable for about 2 weeks postpartum but does not prevent follicular development (Moller, 1970b; Wagner and Hansel, 1 9 6 9 ) . Small corpora lutea, similar in size to Progestogen (17a acetoxy 11/3 methyl 19 norpreg 4 ene 3, 20 dione) used in the Syncro-Mate B treatment of G. D. Searle and Co.
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KIRACOFE dairy cows than in beef cows; however, this difference has not been studied independent of nutritional level. Many dairy cows have an ovulation without estrus before their first overt estrus (Morrow et ol., 1966; Marion and Cier, 1968; Callahan et al., 1971; Whitmore et al., 1974). In both beef and dairy cows, the corpus luteum resulting from the first ovulation has a short life span (Menge et al., 1962; Morrow, 1969; Kiracofe et al., 1969). Perhaps the involuting uterus could affect both the ovulatory pattern (i.e., which ovary is involved in ovulation) and the life span of the first corpus luteum after calving by a mechanism similar to that shown for uterine dilatation (Hansel and Wagner, 1960). However, Ward et al. (1979) found that short cycles were frequent when calves were weaned from anestrous postpartum cows as late as 68 days after calving when uterine involution should have been completed. Many attempts have been made to elucidate ovarian function and reproductive hormone levels prior to resumption of estrous cycles after calving. An increase in serum or blood progesterone level occurs in postpartum cows before the first estrus (Donaldson et al., 1970; Erb et al., 1971; Henricks et al., 1972; Humphrey et al., 1976; Echtemkamp and Hansel, 1973; Tribble et al., 1973; Arije et al., 1974; Corah et al., 1974; Prybil and Butler, 1978; LaVoie et al., 1980). LaVoie et al. (1980) found that when data from cows that had an ovulation without estrus were removed, no cow had a cyclic luteal phase pattern of preestrus

follicles during the first 3 weeks postpartum, occasionally are huried in the ovarian stroma (Moller, 1970b). Moller (1970b) suggested that the early developing small corpora lutea were inactive and speculated that ovulations leading to their formation were infertile. Evidence indicates that the postpartum uterus may influence ovulation. A high percentage of ovulations in the first 20 days postpartum occurs on the ovary opposite the previously gravid hom (Saiduddin et ol., 1967; Foote and Peterson, 1968; Kiracofe et al., 1969); however, Wagner and Hansel (1969) were unable to confirm these observations. Ovulations which occur on the ovary opposite the previously gravid horn are more numerous in dairy cows than in beef cows, and conception at first service is lower if ovulation occurs on the ovary ipsilateral to the previously gravid horn (Foote and Peterson, 1968). Table 1 lists estimates of the time required for uterine involution in early studies in which breeds were identified and the time of first ovulation and estrus were determined. The average postpartum interval to the first ovulation is less than that to the first estrus. Data presented from these early studies indicated first ovulation occurred from 35 to 60 days postpartum. Other studies, especially with dairy cows, have shown that ovulation occurred approximately 14 to 22 days postpartum (Menge et al., 1962; Morrow et al., 1966; Marion and Cier, 1968; Callahan et al., 1971; Morrow, 1971; Whitmore et al., 1974). The first postpartum ovulation occurs earlier in

TABLE 1. EARLY STUDIES ON POSTPARTUM REPRODUCTIVE FUNCTION IN THE BOVI~E SPECIES

Days postpartum Reference Casida and Wisnicky (1950) Wiltbank and Cook (1958) Wiltbank and Cook (1958) Foote et al. (1960a) Foote et al. (1960b) Foote et al. (1960b) Ulberg and Lindley (1960) Fosgate ei al. (1962) Fosgate et al. (1962) Foote and Hunter (1964) Foote and Saiduddin (1964) Saiduddin et al. (1968)
eNureed, bMilked.

Breed Holstein Shorthorn" Shorthorn" Hereford Angus Shorthorn Hereford Holstein Jersey Hereford Hereford Hereford

Uterine invol ution 28 56 44 44 41 43 46 28 27 47 46 46

First ovulation 35 53 36 38 61 62 38 45 3.5 44 38 38

First estrus 63 84
M

59 86 76 46 51 45 49 46 46

U T E R I N E I N V O L U T I O N AND P O S T P A R T U M plasma progesterone, but all cows with normal postpartum estrous cycles had a preestrus increase in plasma progesterone level. T h e progesterone increase was not associated with a palpable corpus luteum. W a r d ( 1 9 7 8 ) demonstrated that most postpartum anestrous cows that were suckling a calf had a short estrous cycle (7 to 11 days) if they exhibited the first estrus within 12 days after the calf was weaned. It appeared that ovulation did occur, but the corpus luteum in most cases was not palpable during the short cycle and the corpus luteum regressed before it was fully formed ( W a r d et al, 1 9 7 9 ) . Corpora lutea induced by G n R H at 30 or 4 8 days postpartum in anestrous cows also had a short life span (Lishman et al., 1 9 7 9 ) . Cows with a short estrous cycle did not have a preestrus blood progesterone peak in data reported b y Corah et al ( 1 9 7 4 ) and LaVoie et al. ( 1 9 8 0 ) . T h e preestrus blood progesterone measured by many researchers before the first postpartum estrus of a normal cycle may have come from a corpus luteum formed from the first postpartum ovulation, since the first ovulation in lactating cows is sometimes unaccompanied by estrus and the resulting corpus luteum has a short life span (Ward et al, 1 9 7 9 ) . A source of preestrus progesterone appears to be important to the resumption of normal cycles; however, the source of this progeterone in the postpartum cow has not been determined and may or may not be different in postpartum cows and prepuberal heifers. T h e source of preestrus progesterone in prepuberal heifers appears to be from small stromal follicles that luteinized (Berardinelli et al, 1 9 7 8 ) . Connor et al. ( 1 9 7 4 ) postulated that luteinization of follicles may be necessary to initiate the first postpartum estrus with ovulation, and others have suggested that luteal function may be deficient during early estrous cycles (Short et al., 1972; Edgerton and Hafs, 1 9 7 3 ) . Since the short life span of the first corpus luteum occurs independent of the time postpartum, uterine involution appears to have no role in early regression of the first corpus luteum unless a different mechanism is involved during the early postpartum period than is involved later. Lack of an overt estrus at the first postpartum ovulation is related to deficient ovarian function (Whitemore et al, 1 9 7 4 ) . Nonstanding estrous behavior decreases as the postpartum interval increases (Saiduddin et al, 1968; LaVoie et al, 1 9 8 0 ) . Lack of uterine involution could be a factor in suppressing ovarian function and, thus, estrus;

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however, suckling intensity or milk removal appears to be the primary influence (Ward et al, 1 9 7 9 ) . Perhaps depressions in conception rate at the first postpartum estrus have been affected by formation of abnormal corpora lutea, failure to maintain luteal tissue and other factors rather than by delayed uterine involution. Under normal conditions, the involuting uterus appears to be a contributing factor to lowered fertility for approximately 4 0 days postpartum. Lack of early large follicle development, ovulation in the absence of behavioral estrus, the requirement of preestrus progesterone secretion to establish cycle patterns, failure of the first ovulation to result in a normal corpus luteum and other factors may supercede the effects of the involuting uterus. In dairy cattle, in which ovulation frequently occurs before day 2 0 , uterine involution certainly could contribute to infertility. However, a low percentage of cows displays an estrus accompanying ovulation before day 2 0 and conception before uterine sloughing would be difficult. Once the uterine milieu returns to near the nonpregnant condition (i.e., relatively sterile, repaired, cleared of lochia, e t c . ) , there appears to be no direct evidence that the uterus interferes with conception. Unless complications develop, the uterus is apparently repaired b y 4 0 days after calving. Ovine Species Uterine Involution and Associated Factors Affecting Fertility. Limited data are available on uterine involution in the ewe. However, the process appears to be similar to that in the cow, in that necrotic tissue over the caruncle was very loosely attached b y day 15 postpartum and was sloughed b y day 2 0 to 2 5 (Uren, 1 9 3 5 ) . Kohl ( 1 9 7 1 ) found shrinkage in uterine size relatively complete and necrotic tissue absent from the uterus by 25 days. Epithelium had also regenerated from the uterine glands and intercaruncular areas. Caruncles were covered with epithelium by 2 5 to 3 0 days. Investigators agree on description of involution processes and the time they occur. T h e uterus appears to be near its nonpregant state by 2 5 days after lambing (Uren, 1 9 3 5 ; Foote et al, 1967; Foote, 1 9 6 8 ; Hulet, 1968; Kohl, 1 9 7 1 ; Jennings and Tisdall, 1974; Call et al, 1 9 7 6 ) . Time of first ovulation after lambing has not been adequately determined in the ewe. Thus, the role of the uterus in regulating the first postpartum ovulation is unknown. T h e first postpartum ovulation usually is not as-

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KIRACOFE uterine weight remained fairly constant, except that Svajgr et al ( 1 9 7 4 ) reported that involution time was slightly longer, with endometrial repair essentially complete beween 13 and 17 days. T h e number of muscle cells of the myometrium declines with involution, and muscle cell size and amount of connective tissue decreases (Graves et al., 1 9 6 7 ) . Palmer et al. ( 1 9 6 5 b ) found that the uterine epithelium, which is degenerated during the early postpartum period, started regenerating 7 days postpartum and appeared complete by 2 1 days. T h e uterus involutes more rapidly in the sow than in the ewe or cow; the sow's lack of necrotic tissue to be sloughed appears to be the primary difference. One cannot consider uterine involution and its relation to postpartum fertility in the sow without considering weaning effects. To determine how quickly the uterus can support a new pregnancy, one must eliminate the suppressing effect of suckling on the reproductive endocrine system. Morphologically and histologically, uterine involution requires approximately 3 weeks. In the sow, however, uterine size has decreased considerably and epithelial repair is well underway by 1 week postpartum. There is no evidence to suggest the extent of repair or reduction in size required before conception can occur in the postpartum sow. According to Polge ( 1 9 7 2 ) , uterine involution may not limit reproductive ability after parturition because blastocyst attachment does not occur until 12 to 13 days after ovulation. Hays et al. ( 1 9 7 8 ) found at 3 days postmating that ovulation rate, ova recovered and fertilization rate were not affected by length of lactation ( 6 , 12, 18, or 24 days). Generally, ovulation and fertilization rates have been similar in sows bred before 2 1 days postpartum and those bred later (Baker et al, 1953; Moody and Speer, 1 9 7 1 ; Svajgr et al., 1974; Rampacek, 1974; Varley and Cole, 1 9 7 6 ) . Farrowing rates were slightly lower in sows bred after 6- and 12-day lactations than in sows bred after 18- or 24-day lactations (Hays et al, 1 9 7 8 ) . Pay ( 1 9 7 3 ) and Svajgr et al. ( 1 9 7 4 ) found that reductions in litter size were more pronounced when sows were weaned less than 7 days postpartum than in those weaned later. Reduced litter size from sows bred less than 3 weeks postpartum has been attributed to increased embryonic loss by Varley and Cole ( 1 9 7 6 ) and Svajgr et al. ( 1 9 7 4 ) . T h e involuting uterus may limit reproductive efficiency if the sow is bred less

sociated with estrus (Call et al, 1 9 7 6 ; Hulet, 1 9 7 9 ) . Estimates of the interval from lambing to ovulation have ranged from 18 days (Hulet, 1 9 7 9 ) to approximately 3 5 days (Foote et al, 1967; Hunter and Lishman, 1 9 6 7 ) unless lambing occurred during seasonal anestrus. A review of postpartum anestrous intervals and factors that influence those intervals in ewes was reported by Hunter ( 1 9 6 8 ) . T h e average interval from lambing to first estrus was found to b e 3 5 days (Hulet, 1 9 7 9 ) ; however, the interval to conception was 44 days. Barker and Wiggins ( 1 9 6 4 ) found the postpartum interval to conception to be about 8 0 days for Rambouillet and 5 5 days for Dorset ewes. T h e single earliest postpartum conception reported has been 9 days, but average time of conception was greater than 4 0 days postpartum (Hulet, 1 9 6 8 ) . Suckling inhibits estrus (Shevah et al., 1 9 7 5 ) and ovulation (Foote et al, 1 9 6 7 ) . Since no difference in uterine involution rates was found between lactating and nonlactating ewes (Foote, 1 9 6 8 ) , the suckling effect on resumption of estrous cycles apparently was not mediated through delayed uterine involution. Follicles larger than 3mm in diameter began appearing on the ovaries after 10 days postpartum, and the number of follicles exceeding 3mm remained constant after 14 or 2 0 days postpartum (Kohl, 1 9 7 1 ) . B y 2 0 days postpartum, follicle growth and turnover were normal. Ovaries appear capable of responding to gonadotropins relatively early postpartum (Hulet, 1 9 6 8 ) . However, the estrogen-induced luteinizing-hormone-release mechanism appears to recover relatively slowly after parturition (Wright and Findlay, 1 9 7 7 ) . Seasonal anestrus has been a complicating factor in determining the influence of uterine involution on early postpartum fertility. T h e uterus appears to be completely involuted b y 30 to 3 5 days after lambing, and no direct evidence is available to attribute low fertility after lambing to a lack of involution of the uterus. Data concerning fertility at the first ovulation after lambing, a time when uterine involution might be incomplete, is lacking. Porcine Species Uterine Involution and Associated Factors Affecting Fertility. Numerous investigators have reported similar decreases in uterine size until 3 weeks after farrowing (Palmer et al., 1965a,b; Graves et al, 1967; Smidt et al., 1969; Svajgr et al, 1974; Varley and Cole, 1976; Hays et al, 1 9 7 8 ) . After 3 weeks,

UTERINE INVOLUTION AND POSTPARTUM than 2 1 days postpartum. Embryo mortality rather than fertilization failure appears to cause smaller litter size in sows bred early postpartum. Hays et al. ( 1 9 7 8 ) concluded that weaning pigs when they are 1 8 days old results in maximum sow productivity in terms of the number of pigs produced per year. Other data also indicate that weaning less than 3 weeks after farrowing increases the interval from weaning to estrus, reduces litter size, and increases both frequency of cystic follicles and embryo mortality (Baker et al., 1953; Self and Grummer, 1 9 5 8 ; Smidt et al., 1965; Moody et al, 1 9 6 9 ; Peters et al, 1969; van der Heyde, 1972; Cole et al., 1975; Hays et al, 1 9 7 8 ) . Attempts to accelerate normal postpartum uterine involution in the sow have been unsuccessful. Hayes et al. ( 1 9 7 8 ) found no change in involution rate after administration of prostaglandin or aspirin. If uterine involution is a limiting factor in reproductive efficiency in the sow, the limiting factor is an unsuitable milieu for embryo survival for approximately the first 3 weeks after farrowing. Segerson and Murray ( 1 9 7 8 ) have shown that the uterus may not be completely functional until 3 to 4 weeks postpartum because the percentage and quantity of uterinespecific acidic proteins secreted before this stage were lower than at later stages. Total uterine protein per corpus luteum, percentages of uterine serum-like protein, total uterinespecific acidic protein were all significantly affected by postpartum interval. Whether these secretions contribute to embryonic survival or mortality during the early postpartum period or are associated with involutionary changes in the uterus has not been established. There seems to be little doubt, however, that reproductive efficiency is impaired in sows bred early postpartum even though the role of the uterus is not clear. Literature Cited Albrechtsen, J. 1917. Sterility of cattle and methods of treatment. Cornell Vet. 7:57. Anderson, L. L., D. L. Hard, L. S. Carpenter and E. E. Awotwi. 1979. Pregnancy, parturition and ovarian function after hypophysial stalk transection in beef heifers. The Endocrine Society 61st Annu. Meet. 126. Araujo, P. C , G. N. Pizelli, M. R. de Carvalko and C. A. Menegnelli. 1974. Postpartum uterine involution and ovarian activity in crossbred dairy cattle. Pesq. Agropec. Boras. Ser. Vet. 9 : 1 . Archbald, L. F., R. H. Schultz, M. L. Fanning, H. I. Kurtz and R. Zemjanis, 1972. A sequential

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histological study of the post partum bovine uterus. J . Reprod. Fertil. 29:133. Arije, F. R., J . N. Wiltbank and M. L. Hopwood. 1974. Hormone levels in pre- and post-parturient beef cows. J . Anini. Sci. 39:338. Baker, L. N., H. L. Woehling, L. E . Casida and R. H. Grummer. 1953. Occurrence of estrus in sows following parturition. J . Anim. Sci. 12:33. Barker, H. B. and E. L. Wiggins. 1964. Estrual activity in lactating ewes. J . Anim. Sci. 23:973. Berardinelli, J . G., R. L. Butcher and R. A. Daily. 1978. Source of progesterone prior to puberty in beef heifers. Beltsville Symp. in Agr. Res. (BARC) III. Anim. Reprod. (Abstr.). Bhalla, R. C , B. K. Soni and D. P. S. Sengai. 1966. Studies on reproduction in Murrah buffaloes. II. Involution of the uterus. Indian Vet. J . 43:892. Boyd, W. L. 1925. A study of the physiological and pathological changes occurring in the reproductive organs of the cow following parturition. Minnesota Agr. Exp. Sta. Bull. No. 23. Boyd, L. J . 1970. Effects of feeding melengestrol acetate (MGA) on occurrence of estrus fertility and milk yield in dairy cows. J . Anim. Sci. 3 1 : 751. Britt, J . H., E . Huertas-Vega and L. C. Ulberg. 1972. Managing reproduction in dairy cattle. 1. Progrestogens for control of estrus in dairy cows. J . Dairy Sci. 55:598. Britt, I. H., R. I. Kittok and D. S. Harrison. 1974a. Ovulation, estrus, and endocrine response after GnRH in early postpartum cows. J. Anim. Sci. 39:915. Britt, J . H., D. A. Morrow, R. J . Kittok and B. E . Seguin. 1974b. Uterine involution, ovarian activity, and fertility after melengestrol acetate and estradiol in early postpartum cows. J . Dairy Sci. 57:89. Brown, J . G., D. W. Peterson and W. D. Foote. 1972. Reproductive response of beef cows to exogenous progestogen, estrogen and gonadotropins at various stages of postpartum. J . Anim. Sci. 35:362. Buch, N. C , W. J . Tyler and L. E . Casida. 1955. Postpartum estrus and involution of the uterus in an experimental herd of Holstein-Fresian cows. J . Dairy Sci. 38:73. Call, T. W., W. C. Foote, C. D. Ecker and C. V. Hulet. 1976. Postpartum uterine and ovarian changes, and estrous behavior from lactation effects in normal and hormone treated ewes. Theriogenology 6:495. Callahan, C. J., R. E. Erb, A. H. Surve and R . D . Randel. 1971. Variables influencing ovarian cycles in postpartum diary cows. J . Anim. Sci. 33:1053. Casida, L. E., W. E . Graves, J . W. Lauderdale, J. W. Riesen, S. Saiduddin, E . R. Hauser and W. J. Tyler. 1968. Studies on the postpartum cow. Res. Bull. 270. Univ. of Wisconsin, Madison. Casida, L. E. and W. G. Venzke. 1936. Observations on reproductive processes in dairy cattle and their relation to breeding efficiency. Proc. of Amer. Soc. Anim. Prod. 36:221. Casida, L. E . and W. Wisnicky. 1950. Effects of diethylstilbestrol dipropionate upon postpartum changes in the cow. J . Anim. Sci. 9:238. Clemente, P. F., R. E . Short, R. B. Staigmiller and

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R. A. Bellows. 1978. Effect of precalving nutrition, early weaning, CB-154, and antiprolactin treatment on postpartum interval length in beef cows. J . Anim. Sci. 47(Suppl. 1 ) : 3 5 1 . Cole, D. J . A., M. A. Varley and P. E. Hughes. 1975. The effect of lactation length on the subsequent reproductive performance of the sow. Anim. Prod. 20:401. Connor, J . S., R. L. Tribble, T. L. Woodward, J. L. Fleeger, J . R. Beverly and A. M. Sorensen, Jr. 1974. Postpartum ovarian activity in firstcalf Hereford heifers. Beef Cattle Research in Texas. Texas Agr. Exp. Sta. Progress Rep. 3216. Corah, L. R., A. P. Queely, T. G. Dunn and C. C. Kaltenbach. 1974. Prepartum and postpartum levels of progesterone and estradiol in beef heifers fed two levels of energy. J . Anim. Sci. 39:380. Dawson, F . L. M. 1950. The microbial content and morphological character of the normal bovine uterus and oviduct. J . Agr. Sci. (Camb.) 40:150. Donaldson, L. E., J . M. Bassett and G. D. Thornburn. 1970. Peripheral plasma progesterone concentration of cows during puberty, oestrus cycles, pregnancy and lactation and the effects of undernutrition and exogenous oxytocin on progesterone concentration. J . Endocrinol. 4 8 : 599. Dunn, T. G. 1964. Influence of energy intake upon reproductive performance and milk production of 2-year-old beef heifers. M. S. Thesis, Univ. of Nebraska, Lincoln. Dunn, T. G., J . E. Ingalls, D. R. Zimmerman and J. N. Wiltbank. i969. Reproductive performance of 2-year-old Hereford and Angus heifers as influenced by pre- and post-calving energy intake. J . Anim. Sci. 29:719. Echternkamp, S. E. and W. Hansel. 1973. Concurrent changes in bovine plasma hormone levels prior to and during tire first postpartum estrous cycle. J. Anim. Sci. 37:1362. Edgerton, L. A. and H. D. Hafs. 1973. Serum luteinizing hormone, prolactin, glucocorticoid and progestin in dairy cows from calving to gestation. J . Dairy Sci. 56:451. El-Fouly, M. A., E . A. Kotby and H. E. El-Sobhy. 1976. Effect of suckling on uterine and cervical involution in post-partum Egyptian buffaloes. Indian J . Anim. Sci. 46:221. Elliot, L., K. J . McMahon, H. T. Gier and G. B. Marion. 1968. Uterus of the cow after parturition: bacterial content. Amer. J . Vet. Res. 29:77. Erb, R. E., A. H. Surve, C. J . Callahan, R. D. Randel and H. A. Garverick, 1971. Reproductive steroids in the bovine. VII. Changes postpartum. J. Anim. Sci. 33:1060. Fernandes, L. C., W. W. Thatcher, E. P. Call, C. J . Wilcox. 1978. Responses to PGF and GnRH in postpartum dairy cows, J . Anim. Sci. 47 (Suppl. 1):359. Foote, W. C. 1968. Control of reproduction with or without hormones. Proc. Symp. on Physiol, of Reprod. in Sheep Amer. Sheep Prod. Council and USDA, Beltsville. Foote, W. D. 1971. Endocrine changes in the bovine during the postpartum period. J. Anim. Sci. (Suppl. I ) 32:73. Foote, W. C , J . W. Call and C. V. Hulet. 1967. Effects of lactation and hormone treatment on
:

ovulation, estrus and uterine involution in the ewe. J . Anim. Sci. 26:943 (Abstr.). Foote, W. D., E. R. Hauser and L. E. Casida. 1960a. Some causes of variation in postpartum reproductive activity in Hereford cows, J . Anim. Sci. 19:238. Foote, W. D., E. R. Hauser and L. E. Casida. 1960b. Influence of progesterone treatment on postpartum reproductive activity in beef cattle. J. Anim. Sci. 19:674. Foote, W. D. and J. E. Hunter. 1964. Postpartum intervals of beef cows treated with progesterone and estrogen. J. Anim. Sci. 23:517. Foote, W. D. and D. W. Peterson. 1968. Relationship between side of pregnancy and side of subsequent ovarian activities in beef and dairy cattle. J . Reprod. Fertil. 16:415. Foote, W. D. and S. Saiduddin. 1964. Hormone treatment of postpartum beef cows. J. Anim. Sci. 23:592 (Abstr.). Fosgate, O. T., N. W. Cameron and R. J. McLeon. 1962. Influence of 17-alpha-hydroxyprogesteronen-caproate upon postpartum reproductive activity in the bovine. J . Anim. Sci. 21:791. Frieden, E. H. and F. L. Hisaw. 1953. The biochemistry of relaxin. Rec. Prog. Horm. Res; 8:333. Fuquay, J . W., R. H. Harris, W. H. McCee, J . F. Beatty and B. L. Arnold. 1975. Routine postpartum treatment of dairy cattle with intrauterine neomycin sulfate boluses. J. Dairy Sci. 58:1367. Gier, H. T. and G. B. Marion. 1968. Uterus of cow after parturition: Involutional changes. Amer. J . Vet. Res. 29:83. Gospodinov, G. and L. Rostov. 1976. Clinical and endocrinological studies on the puerperium in cows. Veterinarnomeditsinski Nauki 13:93. Graden, A. P., O. Olds, C. R. Mockow and L. R. Mutter. 1968. Causes of fertilization failure in repeat breeding cattle. J . Dairy Sci. 51:778. Graves, W. E., J. W. Lauderdale, R. L. Kirkpatrick, N. L. First and L. E. Casida. 1967. Tissue changes in the involuting uterus of the postpartum sow. J . Anim. Sci. 26:365. Griffin, J . F. T., P. J. Hartigan and W. R. Nunn. 1974. Non-specific uterine infection and bovine fertility. Theriogenology 1:91. Gunter, J. J., W. J. Collins, J. Owen, A. M. Sorensen, J . W. Scales and J. A. Alford. 1955. A survey of the bacteria in the reproductive tract of dairy animals and their relationship to infertility. Amer. J. Vet. Res. 16:282. Hansel, W. and W, C. Wagner. 1960. Luteal inhibition in the bovine as a result of oxytocin injections, uterine dilation, and intrauterine infusions of seminal and preputial fluids. J. Dairv Sci. 43:796. Harkness, M. L. R. and R. D. Harkness. 1954. The collagen content of the reproductive tract of the rat during pregnancy and lactation. J . Physiol. 123:492. Harkness, R. D. and B. E . Moralee. 1956. The time course and route of loss of collagen from the rat's uterus during post-partum involution. J. Physiol. 132:502. Hayes, S. H., L. A. Edgerton, G. L. Cromwell, T. S. Stahly and R. H. Dutt. 1978. Prostaglandin Fi effect upon rate of uterine involution in postpartum gilts. J . Anim. Sci. 47(Suppl. l ) : 365.
a

U T E R I N E INVOLUTION AND POSTPARTUM Hays, V. W., J. L. Krug, G. L. Cromwell, R. H. Dutt and D. D. Kratzer. 1978. Effect of lactation length and dietary antibiotics on reproductive performance of sows, J . Anim. Sci, 46:884. Henderson, K. A. 1978. Conception rates of beef cows synchronized at different post-partum intervals. J . Anim. Sci. 47(Suppl, 1 ) : 3 6 5 . Henricks, D. M., J . F. Dickey, J . R. Hill and W. E. Johnston. 1972. Plasma estrogen and progesterone levels after mating and during late pregnancy and in postpartum cows. Endocrinology 90:1336. Hilty, H. 1908. Untersuchungen iiber die evolution and involution der uterus mucosa vom rind. Schweiz Arch. f. Tierheilk. 50:353. Homan, A. E . 1969. The effect of nitrofurazone and proteolytic enzymes on the postpartum bovine uterus. M. S. Thesis, Kansas State Univ., Manhattan. Huertas-Vega. E., J . H. Britt and L. C. Ulberg. 1972. Svstem managing reproduction in dairy cattle, j ' Dairy Sci. 55:401 (Abstr.). Hulet, C. V. 1968. Factors affecting fertility in the ewe and ram. Proc. Symp. on Physiol, of Reprod. in Sheep. Amer. Sheep Prod. Council and USDA, Beltsville. Hulet, C. V. 1979. Reproduction in Sheep. In Reproduction in Farm Animals. E. S. E . Hafez, Ed. 4th Ed., Lea & Febiger. (In press). Humphrey, W. D., D. R. Koritnik, C. C. Kaltenbach, T. G. Dunn and G. D. Niswender. 1976. Progesterone and LH in postpartum suckled beef cows. J . Anim. Sci. 43:290 (Abstr.). Hunter, G. L, 1968. Increasing the frequency of pregnancy in sheep. I. Some factors affecting rebreeding during the post-partum period. Anim. Breed. Abstr.36:347. Hunter, G. L. and A. W. Lishman. 1967. Postpartum ovulation and estrus in spring lambing ewes. J. Reprod. Fertil. 14:473. Jennings. J. and G. Tisdall. 1974. Uterine involution in the postpartum ewe. An Foras Taluntais Research Rep., Co. Meath, Ireland. Johanns, G. J., T. L. Clark and J . B. Herrick. 1967. Factors affecting calving interval. J . Amer. Vet. Med. Assoc. 151:1692. Jordan, W. S. 1952. The puerperium of the cow: A study of uterine motility. J . Comp. Path, and Therapy 62:54. Kiracofe, G. H., G. R. Brower and R. R. Schalles. 1973a. Effects of various uterine treatments on calving-to-conception interval. Kansas Agr. Exp. Sta. Bull. 568. Kiracofe, G. H., A. E. Homan, G. B. Marion, H. T. Gier and R. R. Schalles. 1973b. Post-calving intrauterine infusion of nitrofurazone and enzymes. J . Anim. Sci. 37:318 (Abstr.). Kiracofe, G. H., R. R. Schalles and G. B. Marion. 1969. Effect of wintering ration on reproductive phenomena in beef cows on range. Kansas Agr. Exp. Sta. Bull. 529. Kohl, L. E. 1971. Uterine involution and postpartum conception in the ewe. M. S. Thesis, Kansas State Univ., Manhattan. LaVoie, V., D. K. Han, D. B. Foster and E. L. Moody. 1980. Suckling effect on estrus and blood plasma progesterone in postpartum beef cows. J. Anim. Sci. (In press). Lindley, D. C. 1954. Intra-uterine antibiotic ther-

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apy postservice in infertile dairy cattle. J . Amer. Vet. Med. Assoc. 120:12. Lishman, A. W., S. M. J . Allison, R. L. Fogwell, R. L. Butcher and E . K. Inskeep. 1979. Follicular development and function of induced corpora lutea in underfed postpartum anestrous beef cows. J . Anim. Sci. 48:867. Marion, G. B. and H. T. Gier. 1968. Factors affecting bovine ovarian activity after parturition. J . Anim. Sci. 27:1621. Marion, G. B., J . S. Norwood and H. T. Gier. 1968. Uterus of the cow after parturition: Factors affecting regression. Amer. J . Vet. Res. 29.71. McNutt, G. W. 1927. The corpus luteum of pregnancy in the domestic cow (Bos Taurus) and a brief discussion of cyclical ovarian changes. J. Amer. Vet. Med. Assoc. 72:286. Menge, A. C , S. E. Mares, W. J. Tyler and L. E. Casida. 1962. Variation and association among postpartum reproduction characteristics in Holstein-Fresian cattle. J . Dairy Sci. 45:233. Mohamed, A. A. 1974. Physiological changes in the reproductive organs of buffaloes from parturition to conception. Ph.D. Thesis, El-Azhar Univ., Cairo, Egypt. Moller, K. 1970a. A review of uterine involution and ovarian activity during the postparturient period in the cow. New Zealand Vet. J . 18:83. Moller, K. 1979b. Uterine involution and ovarian activity after calving. New Zealand Vet. J . 18; 140. Moody, N. W., D. S. Baker, V. W. Hays and V. C. Speer. 1969. Effect of reduced farrowing interval on sow productivity. J . Anim. Sci. 28: 76. Moody, N. W. and V. C. Speer. 1971. Factors affecting sow farrowing interval. J . Anim. Sci. 32:510. Morrow, D. A. 1969. Postpartum ovarian activity and involution of the uterus and cervix in dairy cattle. Vet. Scope Vol. 14 No. 1 The Upjohn Co. p. 2. Morrow, D. A. 1971. Effects of periparturient disease on post-partum reproduction in dairy cattle. J . Anim. Sci. 32 (Suppl. I ) : 1 7 . Morrow, D. A., S. J . Roberts and K. McEntee. 1969. A review of postpartum ovarian activity and involution of the uterus and cervix in cattle. Cornell Vet. 59 (No. 1):134. Morrow, D. A., S. J . Roberts, K. McEntee and H. G. Gray. 1966. Postpartum ovarian activity and uterine involution in dairy cattle. J . Amer. Vet. Med. Assoc. 149:1596. Norwood, J. W. 1963. Factors affecting postpartum regression of the bovine uterus. Ph.D. Thesis, Kansas State Univ., Manhattan. Odde, K. G. and G. H. Kiracofe. 1978. Effect of postpartum breeding interval on conception rates in beef cows, Kansas Agr. Exp. Sta. Bull. 320. Olds, D. and T. Cooper. 1970. Effect of postpartum rest period in dairy catde on the occurrence of breeding abnormalities and on calving intervals. J . Amer. Vet. Med. Assoc. 157:92. Olds, D., H. B. Morrison and D. M. Seath. 1949. Efficiency of natural breeding in dairy cattle, Kentucky Agr. Expt. Sta. Bull. 539. Oxender, W. D. and B. E. Seguin. 1976. Bovine

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KIRACOFE pregnancy on postpartum function in the cow. J . Dairy Sci. 50:1846. Segerson, E . C. and F. A. Murray. 1978. Uterine protein secretions in postpartum sows. J. Anim. Sci. 46:1730. Seguin, B. E., D. A. Morrow and W. D. Oxender. 1974. Intrauterine therapy in the cow. J . Amer. Vet. Med. Assoc. 164:609. Self, H. F . and R. H. Grummer. 1958. The rate and economy of pig gains and the reproductive behavior in sows when litters are weaned at 10 days, 21 days, or 56 days of age. J . Anim. Sci. 17:862. Shevah, V., W. J . M. Black and R. B. Land. 1975. The effects of nutrition on the reproductive performance of Finn x Dorset ewes. II. Postpartum ovarian activity, conception and plasma concentrations of progesterone and LH. J. Reprod. Fertil. 45:289. Short, R. E., R. A. Bellows, E . L. Moody and B. E. Howland. 1972. Effects of suckling and mastectomy on bovine postpartum reproduction. J . Anim. Sci. 34:70. Smidt, D., B. Schwven and J . Steinbach. 1965. The influence of lactation on sexual functions in sows. Zuchtungdkunde 37:23. Smidt, D., O. Thume and W. Jockle. 1969. Investigations on sexual regeneration postpartum in suckling and non-lactating sows. Zuchtungskunde 41:36. Studer, E. and D, A. Morrow. 1978. Postpartum evaluation of bovine reproductive potential: comparison of findings from genital tract examination per rectum, uterine culture, and endometrial biopsy. J . Amer. Vet. Med. Assoc. 172:498. Svajgr, A. J., V. W. Hays, G. L. Cromwell and R. H. Dutt. 1974. Effect of lactation duration on reproductive performance of sows. J. Anim. Sci. 38:100. Tennant, B., J. W. Kendrick and R. G. Peddicord. 1967. Uterine involution and ovarian function in die postpartum cow. A retrospective analysis of 2,338 genital organ examination. Cornell Vet. 57:543. Tennant, B. and R. G. Peddicord. 1968. Influence of delayed uterine involution and endometritis on bovine fertility. Cornell Vet. 58:185. Tribble, R. L., A. M. Sorensen, Jr., T. L. Woodard, J . S. Connor, J . R. Beverly and T. L. Fleeger. 1973. Serum progestins and luteinizing hormone levels in non-suckled primiparous heifers. Nature 246:495. Tudorascu, R, 1969. Investigations concerning the frequency of embryonic mortality in cows inseminated at different postpartum intervals. Anim. Breed. Abstr. 37:55. Ulberg, L. C, and C. E. Lindley. 1960. Use of progesterone and estrogen in the control of reproductive activities in beef catde. J . Anim. Sci. 19:1132. Uren, A. W. 1935. Involution of the uterine mucosa in the ewe. Michigan State Coll. of Agr. and Appl. Sci. Tech. Bull. No. 144. Van Demark, N. L. and G. W. Salisbury. 1950. The relation of the post-partum breeding interval to reproductive efficiency in the dairy cow. J . Anim. Sci. 9:307. van der Heyde, H. 1972. A practical assessment

intrauterine therapy. J . Amer. Vet. Med. Assoc. 168:217. Oxenreider, S. L. and W. C . Wagner. 1971. Effect of lactation and energy intake on postpartum ovarian activity in die cow. J . Anim, Sci. 33:1026. Palmer, W. M., H. S. Teague and W. G. Venzke. 1965a. Macroscopic observations on the reproductive tract of sows during lactation and early weaning. J . Anim. Sci. 24:541. Palmer, W. M H. S. Teague and W. G. Venzke. 1965b. Histological changes in the reproductive tract of the sow during lactation and early postweaning. J . Anim. Sci. 24:1117. Pay, M. G. 1973. The effects of short lactations on the productivity of sows. Vet. Rec. 92:255. Perkins, J . L. and H. E. Kidder. 1963. Relation of uterine involution and postpartum interval to reproductive efficiency in beef cattle. J . Anim. Sci. 22:313. Peters, J . B., R. E. Short, N. L. First and L. E . Casida. 1969. Attempts to induce fertility in postpartum sows. J . Anim. Sci. 29.20. Polge, C. 1972. The endocrinology of the sow in relation to early weaning. Proc. Symp, on Improvement of Sow Productivity. Rowett Res. Inst., Aberdeen, Scotland. Prybil, M. K. and W. R. Buder. 1978. The relationships between progesterone secretion and the initiation of ovulation in postpartum beef cows. J . Anim. Sci. 47(Suppl. 1 ) : 3 8 3 . Quevedo, M. M S. Saiduddin and W. D. Foote. 1965. Influence of estradiol on reproductive activity in postpartum dairy cows. J . Anim. Sci. 24:587 (Abstr.). Radford, H. M., C. D. Nacarrow and P. E . Mattner. 1978. Ovarian function in suckling and nonsuckling beef cows postpartum. J . Reprod. Fertil. 54:49. Rampacek, G. B. 1974. Investigations into reproduction in the pre-puberal gilt, the earlyweaned sow and uterine capacity in the female pig. Ph.D. Thesis, North Carolina State Univ., Raleigh. Randel, R. D. and G. A. Welker. 1977. Effect of energy intake and once daily suckling on postpartum interval in Brahham x Hereford heifers. J. Anim. Sci. 45 (Suppl. 1 ) : 1 9 8 . Rasbeck, N. O. 1950. Den normale involutio uteri hos koen. Nord. Vet. Med. 2:655. Riek, P. M., N. Butendieck, E . A. B. Oltenacu and R. H. Foote. 1977. Response of early postpartum dairy cows to GnRH. J . Anim. Sci. 4 5 (Suppl. 1 ) : 2 0 1 . Riesen, J . W. 1968. The effects of suckling on reproductive function in postpartum dairy cows pituitary prolactin, corpora lutea, and uterine histology. Ph.D. Thesis, Univ. of Wisconsin, Madison. Roberts, S. J . 1956. Veterinary obstetrics and genital diseases. P. 123. Published by the author. Ithaca, NY, Saiduddin, S., W. D. Foote and M. M. Quevedo. 1968. Response of beef cows to exogenous progesterone and estradiol at various stage postpartum. J . Anim. Sci. 27:1015. Saiduddin, S., J . W. Riesen, W. J . Tyler and L. E . Casida. 1967. Some carry-over effects of

U T E R I N E INVOLUTION AND POSTPARTUM of early weaning. Proc. Brit. Soc. Anim. Prod. 1:33. Variey," M. A. and D. J . A. Cole. 1976. Studies in sow reproduction. 5. The effect of lactation length of the sow on the subsequent embryonic development. Anim. Prod. 22:79. Venable, J . H. and L. E. McDonald. 1958. Postparturient bovine uterine motilitynormal and after experimentally produced retention of fetal membranes. Amer. J . Vet. Res. 19:308. Wagner, W. C. and W. Hansel. 1969. Reproductive physiology of the postpartum cow. J . Reprod. Fertil. 18:493. Ward, H. S. 1978. The effects of Syncro-Mate B and/or calf management on reproductive performance in the beef cow. M. S. Thesis, Kansas State Univ., Manhattan. Ward, H. S., K. C. Odde, C. H. Kiracofe and R. M. McKee, 1979. Short estrous cycles after weaning in anestrous beef cows. J . Anim. Sci. 49(Suppl. 1):345. Wettemann, R. P., E. J . Turman, R. D. Wyatt and R. Totusek. 1978. Influence of suckling

INTERVALS

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intensity on reproductive performance of range cows. J . Anim. Sci. 47:342. Whitmore, H. L., W. J . Tyler and L. E. Casida. 1974. Effect of early postpartum breeding in dairy catde. J . Anim. Sci. 38:339. Wiltbank, J . N. and A. C. Cook. 1958. The comparative reproductive performance of nursed milked cows. J . Anim. Sci. 17:640. Wiltbank, J . N., N. W. Rowden, J . E. Ingalls, K. E . Gregory and R. M. Koch. 1962. Effects of energy level on reproductive phenomena in mature Hereford cows, J . Anim. Sci. 21:219. Woessner, J. F. 1969. Inhibition by oestrogen of collagen breakdown in the involuting rat uterus. Biochem. J . 112:637. Wright, P. I. and J . K. Findlay. 1977. LH release due to LH-RH or oestradiol-17 beta in postpartum ewes. Theriogenology 8:191. Wulf, F. F. and A. E. Draey. 1952. A survey of the microflora obtained by flushing the normal bovine uterus. Proc. South Dakota Acad. Sci. 31:116.

DISCUSSION Question: Dr. Bellows Your paper was very interesting, but I am not sure if I followed one point that you made regarding the reason for short cycles. Specifically, in relation to the size of the follicle from which the C L was derived. If I understand you correctly, you were implying that the size of the ovulating follicle was involved here? Dr. Kiracofe: I do not think size of the ovulating follicle is playing a role in short cycles in our data. In that respect, I was referring to the research of Moller in Australia, who observed luteinized stromal follicles in postpartum cows. I think they were similar to the luteinization noted b y the West Virginia investigators in prepuberal heifers. T h e data of Berardinelli et al. at West Virginia indicates that these are stromal follicles that are small and luteinized. However, in postpartum cows we do not feel that it was a small follicle that ovulated. Question: Dr. Pickett D o you have any information on the effect on subsequent fertility of those animals that were bred early, say quite soon after parturition, and did not conceive? Dr. Kiracofe: I do not have any data on the effect of early postpartum insemination on subsequent fertility. I think there are some in the literature, particularly the earlier research of Van Demark and Salisbury ( J . Anim. Sci. 9 : 3 0 7 ) and that of Olds and Cooper ( J . Amer. Vet. Med. Assoc. 1 5 7 : 9 2 ) . I think data can be found both ways; some have reported no effect and others indicate a detrimental effect. Question: Dr. Pickett Do you think it has an effect? Dr. Kiracofe: In my opinion there is no effect, although I do not have the data to support that opinion. I do not feel that breeding cows during the early postpartum period effects subsequent fertility. Question: Dr. Pickett Is there a possibility that there is a difference between dairy and beef cattle in this regard? Dr. Kiracofe: There could be. B y putting semen in the early postpartum uterus there may be a chance of causing an inflammatory response or low grade bacterial infection. Since the dairy cow tends to ovulate earlier, many times before uterine involution is complete, she could be more susceptible. Question: A Questioner

On the basis of our knowledge of follicular development, is it not true that follicles have to pass through a certain stage before they will ovulate and that the size of the follicles that ovulate really is fairly consistent within species? Dr. Kiracofe: That may be true in cycling animals where definite endocrine patterns

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KIRACOFE exist, but in animals that are ovulating for the first time in their life or after a prolonged period of anestrus, such as postpartum animals, it could be different. In the case of prepuberal heifers, the follicle is probably luteinizing without ovulation; however, our limited data in the postpartum cow indicate she does ovulate, but we do not know anything about the size of follicle. matter of observing small luteinized follicles or corpora lutea in the ovary. Question: A Questioner In these luteinized follicles, were ova trapped in them or were they actually small corpora lutea? Dr. Kiracofe: Maybe there is someone here who could comment on the West Virginia research. Question: Dr. Lauderdale You are talking about short cycles, therefore, the corpus luteum ( C L ) has obtained its lifespan and the discussion so far has indicated what I would refer to as an inherent decrease in its lifespan. Does the CL have an inherent lifespan, and with short cycles is the inherent lifespan of the C L altered or is there a "superluteolytic" factor in the postpartum animal? Dr. Kiracofe: W e have investigated this to some extent. T h e data are incomplete so far, but I think that in this respect an inherent lifespan might be a possibility. I think that the inherent lifespan would be due to a lack of gonadotropin or lack of a luteotropic substance rather than a luteolytic factor. However, we have not tested the luteolytic aspect.

Question: A

Questioner

T h e evidence is that the corpora lutea are smaller that develop from these ovulations. Did you find any of these small follicles or follicles of large size prior to the ovulation? What kind of evidence is there for smaller follicles ovulating? Dr. Kiracofe: I may be misinterpreting your question. All data that we have with respect to the postpartum cow indicate the corpus luteum is normal size at day 4 of the short cycle and that short cycles are a result of failure to maintain the corpus luteum, not a result of small follicles that luteinize. T h e only evidence where there is a possibility of ovulation of small follicles, is in the cases I indicated, i.e., the prepuberal heifers in the research at West Virginia or the research of Moller in Australia, and this is just a

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