Managing Reproductive Disorders in Dairy Cows
Twinning
Twinning is an unavoidable outcome of
reproduction in dairy cattle and is undesirable in a dairy operation because it
reduces overall profitability and reproductive efficiency (Eddy et al., 1991;
Beerepoot et al., 1992). One study estimated that every twin birth incurs a
$108 economic loss compared with a singleton birth (Beerepoot et al., 1992).
Twinning also reduces reproductive performance by increasing average days open
and services per conception of the dam during the subsequent lactation (Nielen
et al., 1989). In addition, the etiologies of many periparturient diseases in
cattle are associated with twin births. Cows calving twins have higher risks
for stillbirth, retained placenta, metritis, displaced abomasum, ketosis, and acidurea
(Pfau et al., 1948; Markusfeld, 1987; Nielen et al., 1989). Incidences of
abortion (29.3% vs. 12.0%), neonatal calf mortality (15.7% vs. 3.2%), reduced
birth weight (43.5 vs. 30.6 kg), and retained placenta (34% vs. 7%) also are
greater among twins compared with singleton calves, probably due to the reduced
gestation length and increased incidence of dystocia among cows calving twins
(Pfau et al., 1948; Erb and Morrison, 1959; Neilen et al., 1989; Day et al.,
1995). Culling rates are also greater for cows calving twins (Eddy et al.,
1991). One impact of twinning is a reported reduction in the number of fertile
heifers available for use as replacements in the dairy herd (Table 3). This
decrease arises from increased neonatal calf mortality of twins and a skewed
gender ratio resulting in more homozygous male pairs.
Table 3. Effect
of twinning on the number of replacement heifers per pregnancy.
|
|
Replacement Heifers per
Pregnancy |
|
|
Reference |
Single Births |
Twin Births |
|
Erb & Morrison, 1959 |
0.43 |
0.34 |
|
Nielen et al., 1989 |
0.48 |
0.42 |
|
Day et al., 1995 |
0.42 |
0.29 |
Mechanisms of Twinning
Cattle are a monotocous species meaning
that, under most circumstances, a successful pregnancy results in the birth of
a single calf. The physiologic mechanism responsible for regulating the number
of follicles that become dominant within each follicular wave usually results
in selection of a single dominant follicle capable of ovulation (see Figure 2).
Initiation of an induced or naturally occurring ovulatory stimulus causes release
of a single oocyte from the dominant follicle at ovulation. If the subsequent
events from fertilization to parturition occur normally, the pregnancy will
result in birth of one calf. On occasion, however, two follicles are selected
to continue growth from among the group of growing follicles in a follicular
wave resulting in a phenomenon termed codominance. If the appropriate stimulus
for ovulation occurs naturally or is induced when codominant follicles are
present, two oocytes, one from each follicle, will be released. If the
subsequent events from fertilization to parturition occur normally for both
oocytes, twins will result. The single cell that is formed after an oocyte is
fertilized by a sperm is called a zygote. Thus, twins that occur as a result of
ovulation and fertilization of two oocytes are called dizygous twins.
Most twins in cattle are of the dizygous
type (Erb and Morrison, 1959; Johansson et al., 1974; Ryan and Boland, 1991).
Dizygous twins can be the same or opposite in sex and are no more alike
phenotypically or genetically than siblings with the same parents born during
different gestations. Ovulation of a single dominant follicle also can, on rare
occasions, result in twins. Twins resulting from ovulation and fertilization of
a single oocyte are called monozygous twins. Monozygous twins are genetically
and phenotypically identical and, therefore, are always of the same sex. The
mechanism by which monozygous twinning occurs is not clearly understood, but
monozygous twinning can be considered a natural cloning of the original zygote
in vivo. The rate of monozygous twinning in cattle is low, with estimates
ranging from 7.4% (Erb and Morrison, 1959) to 13.6% (Ryan and Boland, 1991) of
all twin births or less than 0.3% of all births.
Twinning Rates in Dairy Cattle
Risk factors for twinning in cattle
include effects of breed type, parity. (Nielen et al, 1989; Ryan and Boland,
1991). The percentage of twins born also varies among seasons of the year, with
a trend toward more twin births during the summer months. This seasonal effect
on twinning has been attributed to an increased plane of nutrition during the
fall when cows calving during the summer would have conceived, a decreasing
light period, and a decrease in the viability of early stage embryos conceived
during summer months compared with those conceived during cooler fall months
(Cady and Van Vleck, 1978; Nielen et al., 1989). High cumulative milk
production and previous twinning are additional factors that increase the risk
of twinning (Nielen et al., 1989; Kinsel et al., 1998). In general, the
twinning rate for most beef breeds of cattle is less than 1% (Rutledge, 1975).
The reported incidence of twinning in dairy cattle ranges from 2.5 to 5.8% and
is dramatically affected by parity, ranging from 1% for first parity to nearly
10% during later parities (Table 4).
Table 4.
Effect of parity on twinning rate (%) or relative risk (R)a of
twinning in dairy cattle.
|
|
No. of |
Parity |
All |
|||||
|
Refb |
Calvings |
1 |
2 |
3 |
4 |
5 |
6 |
Parities |
|
1 |
937 |
0.7 |
5.0 |
4.2 |
5.0 |
7.0 |
6.7c |
4.2 |
|
2 |
7,387 |
1.3 |
4.4 |
5.3 |
4.6 |
5.8 |
6.0 |
4.6 |
|
3 |
11,951 |
0.8 |
2.7 |
4.1 |
4.5 |
4.9 |
4.8c |
3.2 |
|
4 |
19,755 |
0.9 |
2.1 |
3.5 |
3.4 |
3.7 |
3.2 |
2.5 |
|
5 |
24,843 |
1.0 |
7.0 |
7.5 |
7.9 |
9.1c |
- |
4.2 |
|
6 |
19,497 |
1.3 |
6.0 |
9.4c |
- |
- |
- |
- |
|
7 |
52,362 |
1.0 |
2.9 |
3.2 |
3.9 |
3.3 |
4.1c |
2.4 |
|
8 |
8,521 |
R=0.2 |
R=1.5 |
R=1.6 |
R=1.6 |
R=2.0c |
- |
5.8 |
aRelative
risk (R) of twinning for each parity group is calculated against all others
pooled together.
bReference: 1=Pfau et al., 1948;
2=Erb & Morrison, 1959; 3=Nielen et al., 1989; 4=Eddy et al., 1991; 5=Ryan
& Boland, 1991; 6=Berry et al., 1994; 7=Kinsel et al., 1998; 8=Markusfeld,
1987.
cIncludes
all cows ³
the parity listed.
The effect of parity on twinning rate is
not clearly understood but may be explained by an increased ability of older
cows to support twins throughout gestation, an increase in the rate of double
ovulation, or an interaction of both of these factors. Increased uterine
capacity of cows calving twins has been reported (Ryan and Boland, 1991).
Furthermore, the incidence of double ovulation in lactating dairy cows is
around 14% (Kidder et al., 1952; Fricke et al., 1998), and, as with the
incidence of twinning, increases with parity (Labhsetwar et al., 1963). Kinsel
et al. (1998) reported an increase rate of twinning over a 10-year period. The
single largest contributor to this increase was the increase in peak milk
production that occurred over that period. They also suggested that feeding
higher energy diets to high producing cows may be increasing the incidence of
double ovulations, and hence the rate of twinning. This nutritional effect is
similar to the practice of "flushing" in ewes (Dunn and Moss, 1992),
but further research is required in dairy cattle.
Increasing levels of dietary bypass
protein can increase ovulation rate and incidence of twinning in ewes (Nottle
et al., 1988). Therefore, the high levels of bypass protein fed to lactating
cows may partially account for the increased twinning rate. The overall
twinning rates reported for dairy cows in recent studies are greater than those
reported in many earlier reports (Day et al., 1995), indicating that twinning
rate may be increasing over time in the dairy cattle population as a whole. If
twinning is related to nutrition and/or milk production (Nielen et al., 1989),
this increase in twinning would not be unexpected considering recent trends in
feeding practices and yearly increases in milk production per cow.
Freemartinism
Freemartinism in heifers results from
twinning when embryonic membranes of a male and female conceptus fuse during
gestation resulting in exchange of blood between the male and female fetuses.
Endocrine factors or cells from the male calf cause abnormal development of the
reproductive organs of the female calf resulting in infertility. Freemartinism
occurs in about 92% of heifers born as a result of heterosexual twin
pregnancies (Buoen et al., 1992). Thus, about 8% of heifers from heterosexual
twin pregnancies will be fertile, presumably because the fetal membranes fail
to fuse or because membrane fusion occurs after the critical period of
reproductive organ differentiation (Buoen et al., 1992).
The earliest developmental abnormalities
of the female reproductive tract resulting in freemartinism occur between 49 to
52 days post fertilization (Jost et al., 1972). Interestingly, freemartinism
has been documented in singleton female calves, which probably results due to
loss of a male twin after fusion of the embryonic membranes but before
parturition (Wijeratne et al., 1977). In addition, many bulls born twin to a
heifer exhibit various degrees of impaired reproductive function including
inability to produce semen, reduced sperm production, or increased incidence of
abnormal spermatozoa (Dunn et al., 1979). From a practical perspective, bulls
born twin to a heifer can be used for breeding purposes after passing a
breeding soundness and semen quality examination (Long, 1979).
Management of Twins
Management of cows carrying twins depends
on accurate identification of the presence of twins early during gestation.
Cows carrying twin pregnancies can be accurately identified at 40 to 55 days
post AI using transrectal ultrasonography (Echternkamp and Gregory, 1991; Davis
and Haibel, 1993; Dobson et al., 1993). Palpation per rectum between 50 to 70
days post AI also results in an acceptable degree of accuracy among experienced
bovine practitioners (Day et al., 1995).
Several management scenarios could be
considered upon diagnosis of a twin pregnancy in a dairy cow. Continued
management of the cow carrying twins could be avoided either by culling the cow
or by aborting the twin pregnancy, usually through administration of an ecbolic
agent such as PGF2a. Several factors would argue against
aborting a twin pregnancy with the intent of rebreeding the cow. First, the
estimated average lactation length of cows subjected to induced abortion and
rebreeding would approach 500 days (~18.5 month calving interval) based on
average reproductive performance and management indices for lactating cows
(Table 5). Second, the risk for a twin pregnancy during the subsequent
gestation is increased because cows calving twins are at greater risk for
subsequent twinning (Nielen et al., 1989). Based on these considerations and
depending on the value of the dam and calf, culling may be a better alternative
to aborting the pregnancy.
Table 5. Estimated
intervals and cumulative days in milk associated with events after a management
decision to terminate the pregnancy of a cow carrying twins.
|
|
Interval |
Cumulative |
|
Mean
interval from: |
(Days) |
Days in Milk |
|
Calving to twin pregnancy diagnosis
and induction of abortion |
184a |
184 |
|
Induction of abortion to second
conception |
84b |
268 |
|
Second conception to dry off |
232c |
500 |
aAverage
days open (144 days; voluntary waiting period = 60 days, conception rate = 40%,
and service rate = 40%)+ day of gestation at diagnosis of twins (40 days).
bMedian
days to second conception using AI breeding (84 days; conception rate = 40%,
and service rate = 40%).
cAverage
gestation length (282 days) - average dry period (50 days).
If a cow carrying twins is to
be maintained in the herd until parturition, several management practices
should be considered. First, based on research in beef cows (Wheeler et al.,
1979; Koong et al., 1982), dairy cows carrying twins should be fed a higher
plane of nutrition, especially during the last trimester of gestation (Nielen
et al., 1989; Day et al., 1995). Second, because the gestation length of cows
calving twins is reduced by 7 to 10 days (Pfau et al., 1948; Nielen et al.,
1989; Ryan and Boland, 1991), most cows calving twins would miss the transition
diet feeding period beginning about two to three weeks before the estimated
calving date. Earlier dry off and feeding of a transition diet may reduce
the incidence of postpartum metabolic problems associated with cows calving
twins (Ryan and Boland, 1991). Finally, assistance at calving for cows carrying
twins may reduce complications associated with dystocia and may reduce economic
losses by reducing the rate of neonatal calf mortality.