Variables Influencing Ovarian Cycles in Postpartum Dairy Cows C. J. Callahan, R. E. Erb, A. H. Surve and R. D. Randel J Anim Sci 1971.

33:1053-1059.

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VARIABLES INFLUENCING OVARIAN CYCLES POSTPARTUM DAIRY COWS 1 C. J.

CALLAHAN,

IN

R. E. ERB, A. H. SURW 2 AND R. D. RANDEL3

Purdue University, La/ayette, Indiana 47907

may rupture OVARIAN follicles may form afterand corpora lutea (CL) calving without prior evidence of estrus in milked cows. Among studies reported, the average period after calving to rupture of the first ovarian follicle (ovulation) ranged from 14 to 35 days and only 14 to 35% of the cows showed estrus at the beginning of the first ovarian cycle (Casida and Wisnicky, 1950; Menge et al., 1962; Morrow et al., 1966; Saiduddin et al., 1968; Marion and Gier, 1968; Wagner and Hansel, 1969). Growth and persistence of the CL may be subnormal and time from the first to the second ovulation averages about 15 days as compared to about 21 days from the second to the third ovulation (Morrow et al., 1966; Marion and Gier, 1968). These studies suggest that hormonal balances established to support pregnancy and its subsequent termination are gradually readjusted after calving to reestablish normal estrous cycles. The time period required after calving for readjustment by individuals is variable and appears to be influenced by a variety of environmental and genetic factors (Erb, Ehlers and Morrison, 1958a; Erb et al., 1958b; Labhsetwar, Tyler and Casida, 1963; Morrow et al., 1966; Marion and Gier, 1968). The objective of the present study was to survey the influence of such variables as twinning, time from calving to release of the fetal membranes (placenta) and gross clinical symptoms of uterine infection (metritis) on ovarian cycles, cystic ovaries, reproductive efficiency and reproductive hormones. Results of this study were briefly referred to in a recent review (Erb, Randel and Callahan, 1971a). Quantitative data on reproductive hormones are reported in a companion paper (Erb et aL, 1971b). Experimental Procedure All heifers and cows calving between August 21 and October 21, 1966 at the Purdue Dairy Center were used in some phase of this study. There were 14 and 29 first calf heifers and 24 and 35 cows of the Red Danish and HoP stein-Friesian breeds, respectively. Eighty-nine of these animals were included in the prepartum study reported by Hunter et al. (1970) and the time from calving to release of the placenta was known for 82 following single births and for 10 following twin births. Based on clinical examinations, 65 cows scheduled for rebreeding were classified into postpartum groups consisting of 23 controls, 20 metritis, 10 cystic ovaries and 12 metritis plus cystic ovaries in combination. Each cow was assigned to only one group even though the abnormalities were not diagnosed at every rectal examination. The rationale for this approach was to isolate clinically normal cows (control group) from other conditions believed to adversely influence reproductive efficiency. The control group was never diagnosed as having metritis and CL were always detected after each estrus or after ovulation without estrus. The ovarian cycles all were 14 to 28 days in length. The metritis group consisted of cows showing fluid accumulation in the uterine horn(s) after 8 days postpartum and presence of pus in vaginal mucus discharged 13 or more days after calving. Such criteria would identify obvious cases of metritis or pyometra but would not detect all cows with low levels of infection. Cows not showing evidence of metritis were classified as cystic if ovarian follicles persisted in the absence of palpable CL for as long as 14 days, if ovulation was delayed more than 7 days after estrus or if CL were nondetectable for at least 7 days after ovulation. Marion and Gier (1968) have shown that diagnosis of cystic ovarian structures by palpation is accurate as compared to the micr0morphology of the ovary. Cows diagnosed cystic and showing evidence
JOURNAL OF ANIMAL SCIENCE, vol. 33, no. 5

1Journal-Paper No. 4387, Purdue Agricultural Experiment Station, Lafayette, Indiana 47907. Supported in part from a trust agreement between Purdue University and Normandy Farm, New Augusta, Indiana, Mr. and Mrs. Herman C. Krannert, owners. A contribution from Western Regional Project W-95. 2Present address: Sandoz Pharmaceuticals, Hanover, New Jersey. 3 Fellow, Purdue Research Foundation, Grant No. 5076. 1053

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CALLAHAN ET AL. levels of dietary urea as compared to a control ration containing only protein from plant sources. An effect of dietary urea, if any, in the present study should have been similar for all animals unless incorporation with a forage base of alfalfa-grass results in a different response as compared to corn silage. In this case, the 16 cows which calved between October 7 and 21 only received corn silage forage after calving. The data were analyzed by analysis of variance (Steel and Torrie, 1960) and group averages were tested for significant differences (Kramer, 1956). Results and D i s c u s s i o n

of metritis comprised a fourth group designated metritis-cystic. Beginning on day 8 after calving, rectal examinations were made at intervals to determine ovarian and uterine changes with special emphasis on follicular development and rupture, subsequent CL (corpus luteum) formation and evidence of metritis. The length and width of each ovary were recorded in centimeters as estimated by rectal palpation. Ovary size was summarized by multiplying length and width (cm2). Diameters of palpable follicles and CL were estimated and recorded in increments of 0.5 cm for those judged to be 1 cm or larger in diameter. Observations for estrous behavior were made at 2-hr. intervals. A cow was recorded in estrus based on mounting behavior and condition of the reproductive tract as determined by rectal palpation. Behavior characterized as estrus included attempts by the cow to mount other cows and partial acceptance of mounts by other animals as commonly seen during initiation of estrus. The cows were housed in two free stall barns, milked twice daily (0300 and 1500 hr.) in a milking parlor and fed outside from open feed bunks. All areas were well lighted and readily visible from working areas. Under the conditions available for observation, all cows showing signs of behavioral estrus for at least 2 hr. should have been detected. Before calving, the cows and heifers, managed as a group, were fed daily an average of 5 kg of concentrate containing 2% urea and low-moisture alfalfa-grass silage. After calving the same concentrate mixture was blended with the silage and fed ad libitum as a complete feed at a dry matter grain to forage ratio of 50:50 (high energy) for cows producing more than 22 kg of milk per day and at a ratio of 40:60 (medium energy) for cows producing less than 22 kg of milk per day. Corn silage was substituted for alfalfa-grass silage on October 7 and crude protein equivalent was maintained at earlier levels by addition of soybean meal. The complete high energy and medium energy mixtures, respectively, contained on a dry-matter basis about 14.5% and 12.6% crude protein and 42% and 35% of the crude protein was protein equivalent from urea. An earlier study had shown no differences in general health, milk production (Howard et al., 1968) and average services per conception (Howard, 1967) between least-cost rations containing similar

Twinning. Twinning rate was 7% for 42 heifers and 14% for 50 cows. None of three heifers and five of seven cows producing twins retained the placenta from 60 to 192 hr. after calving. The placenta was not retained by three animals producing twin females (/~--7 hr.) and was retained following birth of one of two sets of twin males and four of five sets of mixed sex. Among 43 cows producing single calves, the placenta was retained 12 or more hours following 21% of the male birth and following 13% of the female births. The difference associated with sex of calf is in agreement with other data (Erb et al., 1958b). All twin calves were born alive 269 to 284 days after breeding. Among the I0 cows producing twins, eight developed metritis and four of these also were diagnosed cystic. Placenta. Six percent of the heifers and 24% of the cows (table 1) retained the placenta 12 or more hours after calving
T A B L E 1. T I M E A F T E R C A L V I N G TO R E L E A S E OF F E T A L M E M B R A N E S B Y 39 H E I F E R S AND 43 COWS F O L L O W I N G B I R T H OF S I N G L E CALVES Fetal membranes released ( % ) Hr. after birth 2.0 to 2.9 3.0 to 3.9 4.0 to 4.9 5.0 to 5.9 6.0 to 6.9 7.0 to 7.9 8.0 to 8.9 9.0 to 9.9 10.0 to 11.9 12.0 1o 14.0 over 14.0 Heifers 10 36 27 3 3 5 8 3 0 3 3 Cows 5 16 23 14 2 5 2 7 2 5 19 Total 7 26 25 9 2 5 .5 5 1 4 11

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OVARIAN CYCLES IN POSTPARTUM TABLE 2. POSTPARTUM CLINICAL CONDITION OF COWS DELIVERING SINGLE CALVES AND RETAINING THE PLACENTA LESS THAN 7, 7 TO 12 AND MORE THAN 12 HR. AFTER CALVING Placenta release after calving (hr.) Groups A B C D E F ~7 7 to 12 >12

COWS

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No. of cows observed 58 14 10 Controls (%) 50 50 10 Cystic (%) 15 0 0 Metritis (%) 24 29 80 Metritis and cystic (%) 11 21 10 Totalcystic(B+D) (%) 26 21 10 Totalmetritis (C+D) (%) 35 51 9O Reproductive efficiency (% calving) 75 63 67 No. bred ~ 48 8 9 a Some cows were cuIled for low production, mastiffs, poor udder conformationor used for nurse cows and were not rebred.

(1274-20 hr.). Differences between heifers and older cows have been previously reported ( E r b et al., 1958b). Breed of animal and length of gestation were not significantly related to length of time the placenta was retained. B y 6 hr. postpartum, 76% of the heifers and 58% of the cows had released the placenta. Based on the accumulative frequencies b y hours after calving, a departure from a normal distribution was evident by 7 hr. p o s t p a r t u m (table 1). This suggests that retention of the placenta longer than 6 to 7 hr. m a y be abnormal. F i f t y percent of the animals releasing the placenta within 12 hr. p o s t p a r t u m were subsequently classified into the control group as compared to 10% of those retaining the placenta longer than 12 hr. (table 2). R a t e of metritis in the absence of ovarian dysfunction (cystic) was 80% in the latter group, as compared to 24 and 29%

for animals retaining the placenta less than 7 and 7 to 12 hr., respectively. Reproductive efficiency based on subsequent calving was 63 and 67% for the two groups retaining the placenta 7 to 12 hr. and more than 12 hr., respectively, compared to 75% for the less than 7 hr. group (table 2). These results do not show t h a t cows retaining the placenta 7 to 12 hr. had subsequent reproductive histories differing from those releasing the placenta within 7 hours. T h e major differences were a lower rate of metritis and a higher reproductive efficiency for the less than 7-hr. group. Reproductive efficiency for twinning cows was 43% as compared to 67% for nontwinning cows retaining the placenta more than 12 hr., and 90% for controls (table 3). Cows showing evidence of having cystic ovaries with no other genital complications conceived as efficiently as the controls. I n comparison, cows having metritis averaged 50% and those suffering a combination of metritis and cystic ovaries averaged only 22% (table 3). There is little evidence in the present d a t a t h a t twinning in combination with metritis and retained placenta reduces conception efficiency more than metritis alone. Morrison and E r b (1957) reported that placentae are retained longer than 18 hr. b y 4 3 % of the cows following birth of twins which compares favora b l y to 50% in the present study. Services per conception following birth of twins are comparable to single births when the placenta is not retained and post calving pathological conditions are minimal ( E r b and Morrison, 1959). W h e n the placenta is retained there is some evidence t h a t reproductive efficiency is reduced even when pathological conditions are not observed ( E r b e t al., 1958b). T h e

TABLE 3. EFFECT OF POSTPARTUM CLINICAL ABNORMALITIES ON REPRODUCTIVE EFFICIENCY First service conceptions (%) Months in calving interval (% of animals) <13 13 to 15 >15 Total % calving 43 67 90 50 90 22 67 Not conceiving (%) ~ 57 33 10 50 10 78 33

Group

No. bred

Twinning 7 14 14 14 14 Retained placenta ~ 9 22 56 11 0 Control" 20 55 70 10 10 Metritis b 10 30 40 10 0 Cystic b 10 40 60 30 0 Metritis and cystic b 9 0 11 11 0 Total 65 32 48 14 5 Twinning cowswere omitted. b Cowstwinningand having retainedfetal membranewereomitted. e Either diagnosedinfertileor nonpregnant followingmating 6 or moremond~s after calving.

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CALLAHAN E T AL. that of the cystic group. The metritis-cystic group did not form 1.5 cm CL by 7 days after ovulation until 49__+5.0 days after calving which was significantly ( P ~ . 0 1 ) longer than the control (25+__1.5 days) and metrifis (30 days) groups. Moreover, following 67% of the first ovulations in the metritiscystic group, no CL were detected or were less than 1.5 cm in diameter as compared to 22 and 35%, respectively, for the control and metritis groups. Presumably the incidence of CL less than 1.5 cm at 7 days after ovulations was associated more with cystic ovarian changes (70%) than with metritis (35%). Follicles 1 cm in diameter were detected earlier (15 days) in the cystic group as compared to the other three groups (20 to 24 days), but only 40 to 50% ruptured within 9 days in the cystic and metritis-cystic groups as compared to 100% of the controls and 90% of those having only metritis. Compared to controls, the most obvious effect of metritis was an extended interval to first estrus ( P ~ .01). Similarly compared, the cystic group showed earlier follicle development, delayed follicle rupture, a longer interval to first ovulation ( P < . 0 1 ) and delayed formation of 1.5 cm CL ( P < . 0 1 ) . No cow in the cystic and metritis-cystic groups demonstrated persistent estrus during the first 8 weeks after calving. The first ovarian cycle exceeded 25 days for 42% of the cows in the metritis group as compared to 4, 0 and 17% of the control, cystic and metritis-cystic groups, respectively. First ovarian cycles were less than 18 days in 43% of the control group, 60% of the cystic group, 26% of the metritis group and 58% of the metritis-cystic group. The second and third ovarian cycles each averaged 23 days and
CONDITION ON POSTPARTUM

increased rate of retained placenta by cows producing twins plus the markedly increased rate of metritis following retention of the placenta indicates that metritis is the primary cause of reduced fertility among such cows (table 3). Ovarian' Cycles. Initiation of postpartum ovarian cycles was not significantly different for cows producing twins or retaining the placenta more than 12 hr. as compared to those having metritis and cystic ovaries or only metritis, respectively. On this basis twinning and retained placenta were not included as variables for the summaries on ovarian cycles (table 4). This changed the number of cows in each subgroup and slightly altered the respective averages from results briefly summarized in a recent review paper (Erb et al., 1971a). Small, hard and firm CL of pregnancy were detected on the ovaries of five of 38 cows 8 days after calving. In the remaining cows both ovaries were not palpated at 8 days because they were out of reach. No effort was made to retract the uterus by use of instruments. Postpartum genital tract condition was associated with differences in time to first ovulation, first estrus, first 1.5 cm CL (all P < . 0 1 ) , first 1.0 cm follicle ( P < . 1 0 ) and length of the first ovarian cycle ( P < . 0 5 ; table 4). Time from calving to first ovulation by the control group was ( P < . 0 1 ) less as compared to the cystic and metritis-cystic groups. The metritis group showed the highest frequency of initial ovulations without detection of estrus (80%) and the longest interval to first estrus as compared to the control and cystic groups ( P ~ .01). The first ovarian cycle of the metritis group was longer than the other groups (table 4) and significantly ( P < . 0 5 ) different from
TABLE 4. E F F E C T OF CLINICAL

REPRODUCTIVE TRACT OVARIAN CYCLES

D a y s a f t e r c a l v i n g to o b s e r v a t i o n of f i r s t " Group N Ovulation Avg b Control 23 Metritis (M) 20 Cystic (C) 10 MandC ~ 12 Total 65 G r o u p differences 17+---1.2 g 2 1 - + 1 . 6 g' ~ 2 9 - + 3 . 9 ~''~ 33"+-4.4 i 23--+1.4 P~.01 % ~ 65 80 60 42 65 Estrus Avg 34 g 46-!---3.0~' 25-+3.0 ~ 37-----6.g g'h 37-+2.1 P~.01 CL--1.5 Avg 25-+1.5 30-+2.3 39+3.7 49+'5.0 33-+1.8 P~.01 g ~'~ u'~ i cm Follicle--1 cm Avg 20-I-2.3 24-+2.4 15-+1.2 20-+2.2 20-+1.0 P~.10 g'h g h g'h % '~ 100 90 40 S0

Days from first to s e c o n d ovulation Avg 18-+1.3 24-+2.4 15-+1.8 20-+4.0 g'h g h g'h % r 32 33 23 8 24

78

20-+1.2
P~.05

a Averages in columns with dissimilar superscripts (g, h, i) are significantly different ( P < . 0 5 to P < . 0 1 ) . i, Average+standard error. e Estrus not detected prior to ovulation. a Ovarian follicle 1 cm in diameter ruptured within 9 days after first detected by rectal examination. Diagnosed as having cystic ovaries subsequent to metritis.

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OVARIAN CYCLES IN POSTPARTUM COWS groups were not significantly different, though the abnormal groups had a smaller percentage of cycles in the 18 to 25-day range (50%) than the controls (83%). Estimated Size o/Ovaries. Ovaries increased in size between 8 and 20 days after calving (P<.005) and changes thereafter through 42 to 60 days were nonsignificant. Ovaries of the cystic group increased in size sooner after calving, were larger by 13 days and remained enlarged through 35 days as compared to the other three groups. The metritis-cystic group was intermediate, except on days 28 and 42 (table 5). The differences between postpartum groups were significant ( P < . 0 0 5 ) . Discussion Ovarian cycles are reestablished after calving in milked cows by follicle rupture without evidence of behavioral estrus in 65% or more of the animals (Casida and Wisnicky, 1950; Morrow et al., 1966; Marion and Gier, 1968; table 4). Estrus when detected is mild as observed in this study and by Morrow et al. (1966). The first ovarian cycle is shorter than normal (Menge et al., 1962; Morrow et al., 1966) as shown for the control and cystic groups but not for the two groups involving cows with metritis (table 4). First ovulations generally occur prior to 20 days, especially in cows not showing gross clinical abnormalities of the genital tract (Morrow et al., 1966; table 4) or cows not suckled (Graves et al., 1968; Wagner and Hansel, 1969). In contrast, first ovulation is delayed (29 to 33 days) in cows diagnosed as cystic or cystic in combination with metritis (table 4). Morrow et al. (1966) reported an average of 34 days to first ovulation for abnormal cows compared

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to 15 days for normal cows, and the normal cows had their second ovulation by 32 days which corresponds to an average of 35 days for controls in the present study (table 4). Absence of estrus or subnormal expression of estrus prior to the first follicle rupture after calving and subsequent decreased luteal life span suggests that at this time the endocrine balance mechanisms are incapable of establishing normal estrous cycles. Estrus behavior was observed less frequently preceding the first follicle rupture in cows with metritis and most frequently when cystic ovarian conditions were diagnosed. When cystic ovarian conditions were present, time to first ovulation and first 1.5 cm CL was increased (table 4) and ovaries were larger (table 5) as compared to the control and metritis groups. These differences suggest above average stimulation by FSH and below average stimulation by L H in the cystic and merifis-cystic groups the first 2 to 3 weeks after calving. Plasma L H was significantly lower in both the cystic and metritis groups at 8 days after calving, and somewhat lower in the metritis-cystic group than controls (Erb et al., 1971b). Further interpretations based on quantitative changes of hormones in these cows are discussed in a companion paper (Erb et al., 1971b). Differences in rate of involution and differences in the amount of debris in the uterus influenced, at least in part, by variable severity and duration of metritis could explain much of the variation noted for ovarian cycles. Uterine mucosal epithelium is reestablished in most cows by 30 days after calving (Rasbech, 1950; Gier and Marion, 1968; Wagner and Hansel, 1969) and approximates the time when uterine involution generally is diagnosed as complete (Casida and Wisnicky, 1950;

T A B L E 5. AVERAGE SIZE (ESTIMATED CM 2) OF B O T H OVARIES AS R E L A T E D TO DAYS A F T E R C A L V I N G AND POSTPARTU~r G E N I T A L C O N D I T I O N S Days postpartum 8 13 20 c 28 ~ 35 c 42+ " Total Day differences Group differences N.S." P<.10 N.S. P<.10 P<.05 P<.10 P<,005

Controls 11.6-----1.0(16) b 13.1+--0.8(19) 17.2-----1.8(21) 15.7-----2.1(11) 16.0-----1.6(11) 15.0+-0.8(22) 14.7-+0.9(100) P < . 05

Cystic 14.5 18.9-4-3.1(7) 22.0+-2.5(8) 25.0--+3.8(4) 23.8-+2.6(7) 15.8--+1.8(6) 20.2-+1.2(36) P < . 10

Metritis 12.6-----1.1(11) 12.7--+1.5(13) 17.5-+1.1(16) 18.2-+2.8(5) ...... 16.6+-2.4(5) 15.0-+0,7(50) P < . 01

M-4-C" 13.0-4-1.1(7) 13.6-----0.9(11) 18.4_____1.6(9) 15.9-----1.4(9) 17.9-----1.4(10) 19,6+-2.0(11) 16.6-+0.7(57) P < . 10

a Metritis and cystic for same animal. i, Average, standard error and number of observations,respectively. e Intervals of 16 to 24, 25 to 32, 33 to 38 and 42 to 60, respectively. a Nonsignificant.

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CALLAHAN ET AL. were classified as having metritis as compared to 24% of the cows retaining the placenta less than 7 hours. Percentages of cows ca~viag again within 15 months were 90% for the control and cystic groups, and 50% for the metritis and 22% for the M + C groups. Controls had their first ovulation at 1 7 ~ 1.2 days (65% had undetected estrus), first estrus at 3 4 z 3 . 3 days, first 1.5 cm CL detected at 25 days and first 1 cm follicle detected at 20 days. Differences between groups were significant ( P < . 0 1 ) except for the first 1.0 cm follicle ( P < . 1 0 ) . Compared to controls, the metritis group had the first estrus later, the cystic group had the first 1 cm follicle and first estrus earlier and the first ovulation and first 1.5 cm CL later, and the M + C group had longer intervals to first ovulation, first estrus and first 1.5 cm CL. Ovaries increased in size between 8 and 20 days after calving ( P < . 0 0 5 ) and were largest in the cystic group from 8 to 35 days ( P < .005). The first ovarian cycle averaged 2 4 ~ 2.4 days in the metritis group compared to 15~1.8 days in the cystic group ( P < . 0 5 ) and 18___1.3 days in the control group (nonsignificant). The longest ovarian cycles in the metritis group occurred when fluid was palpated in the uterus 13 or more days after ovulation in seven of nine cows. Early enlargement of ovaries and early growth of follicles with delayed rupture and decreased CL lifespan characterized cows subsequently developing persistent large cystic follicles on the ovaries. L i t e r a t u r e Cited Brinsfield, T. H. and H. W. Hawk. 1958. Unilateral effect of induced uterine infection on luteal development in the ewe. J. Anita. Sci. 27:1631. Buch, N. C, W. J. Tyler and L. E. Casida. 1955. Postpartum estrus and involution of the uterus in an experimental herd of Holstein-Friesian cows. J. Dairy Sci. 38:73. Casida, L. E. and W. Wisnicky. 1950. Effects of diethylstilbestrol dipropionate upon postpartum changes in the cow. J. Anim. Sd. 9:238. Erb, R. E., ]Yl. H. Ehlers and R. A. Morrison. 1958a. Environmental influences on frequency of estrous cycles. Wash. Agr. Exp. Sta. Gen. Bull. 583. Erb, R. E., P. M. Hinze, E. M. Gildow and R. A. Morrison. 1958b. Retained fetal membranes--the effect on prolificacy of dairy cattle. J. Amer. Vet. Med. Ass. 133:489. Erb, R. E. and R. A. Morrison. 1959. Effects of twinning on reproductive efficiency in a HolsteinFriesian herd. J. Dairy Sci. 42:M2. Erb, R. E., R. D. Randel and C. J. Callahan. 1971a. Female sex steroid changes during the reproductive cycle. Proc. IX Bien. Syrup. Anita. Reprod. J. Anita. Sci. 32(Suppl. 1):80.

Morrow et al., 1966; Riesen et al., 1968; Lauderdale et al., 1968). However, age of cow and other factors contribute to variability in the average times to involution (Buch, Tyler and Casida, 1955). Cellular debris in the involuting uterus may inhibit both neural and humoral feedback mechanisms regulating the endocrine control of normal estrous cycles. Estrous cycles are shorter than normal when the uterus is dilated during the early luteal phase (Yamauchi and Nakahara, 1958; Hansel and Wagner, 1960; Yamauchi et al., 1967), but increased if done during the late luteal phase (Yamauchi et al., 1967). Injection of sterile saline solution or suspensions of E. coli into one uterine horn inhibits CL development in both ovaries of sheep (Brinsfield and Hawk, 1968). Therefore, variable accumulations of fluid and debris in the uterus, as is generally observed during metritis, may result in seemingly incongruous effects on ovarian cycles depending on the stage of the ovarian cycle when such accumulations occur. The first ovarian cycles of nine cows in the metritis group ranged from 25 to 49 days, and accumulated fluid was palpated in seven cows 13 to 20 days after ovulation. No alteration in length of ovarian cycles was obvious when uterine fluid was palpated before the first ovulation or in two cows on the day of estrus. Fluid accumulation in the uterus was palpated on day 7, and on day 13 or 14 after estrus or ovulation in the remaining cases. The above observations appear to be in agreement with the controlled studies of Yamauchi et al. (1967).

Summary
Dairy cows, milked twice daily, were used to survey the relationships between postpartum genital tract condition and such traits as twinning, time to release of the fetal membrances (placenta), estimated ovary size, first estrus, first detectable 1 cm diameter follicle, first detectable 1.5 cm diameter corpus luteum (CL) and reproductive efficiency. Cows scheduled for rebreeding were classified into four groups, namely 23 control, 10 cystic, 20 metritis and 12 metritis and cystic ( M + C ) in combination. Within 7 hr. postpartum, 79% of the first calving heifers and 60% of the cows had released the placenta. Six percent of the heifers and 24% of the cows retained the placenta 12 or more hours after calving, and 80%

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CYCLES IN POSTPARTUM

COWS

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Erb, R. E., A. H. Surve, C. J. Callahan, R. D. Randel and H. A. Garverick. 1971b. Reproductive steroids in the bovine. VII. Changes postpartum. J. Anim. Sci. 33 : 1060. Gier, H. T. and G. B. Marion. 1968. Uterus of the cow after parturition: involutional changes. Amer. J. Vet. Res. 29:83. Graves, W. E., J. W. Lauderdale, E. R. Hauser and L. E. Casida. 1968. Relation of postpartum interval to pituitary gonadotropins, ovarian follicular development and fertility in beef cows. Wisconsin Res. Bull. 270, p. 23. 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. Dairy Sci. 43:796. Howard, W. T. 1967. The effect of least-cost computer formulated linear programmed rations fed ad libitum upon performance of lactating dairy cows. Ph.D. Thesis, Purdue University. Howard, W. T., J. L. Albright, M. D. Cunningham, R. B. Harrington, C. H. Noller and R. W. Taylor. 1968. Least-cost complete rations for dairy cows. J. Dairy Sci. 51:595. Hunter, D. L., R. E. Erb, R. D. Randel, H. A. Garverick, C. J. Callahan and R. B. Harrington. 1970. Reproductive steroids in the bovine. I. Relationships during late gestation. J. Anita. Sci. 30:47. Kramer, C. Y. 1956. Extension of multiple range test to group means with unequal numbers of replications. Biometrics 12:307. Lauderdale, J. W., W. E. Graves, E. R. Hauser and L. E. Casida. 1968. Relation of postpartum interval to corpus luteum development, pituitary prolacfin activity, and uterine involution in beef cows. Wisconsin Res. Bull. 270, p. 42. Labhsetwar, A. P., W. J. Tyler and L. E. Casida. 1963. Genetic and environmental factors affecting quiet ovulations in Holstein cattle. J. Dairy Sci. 46: 843.

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