See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/315589534

Early Embryonic Death in Bovines: An Overview

Article · September 2016

CITATIONS READS

8 12,695

4 authors, including:

Sanjay C. Parmar K. K. Hadiya

Anand Agricultural University Anand Agricultural University
51 PUBLICATIONS 146 CITATIONS 43 PUBLICATIONS 191 CITATIONS

SEE PROFILE SEE PROFILE

Cehtan Parmar
Nirma University
8 PUBLICATIONS 30 CITATIONS

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Nutritional and physiological interventions to enhance reproductive performance of dairy animals View project

All content following this page was uploaded by Sanjay C. Parmar on 24 March 2017.

The user has requested enhancement of the downloaded file.


Early Embryonic Death in Bovines:
Sanjay C. Parmar*, A. J. Dhami, K. K. Hadiya and C. P. Parmar,
Department of Animal Reproduction, Gynaecology and Obstetrics, College of Veterinary Science and
Animal Husbandry, Anand Agricultural University, Anand, Gujarat. *Corresponding author.
Abstract
Embryo mortality is a major cause of economic loss
in dairy production systems. Direct effects of
embryo mortality are reflected in reduced
conception rates with consequent effects on
efficiency, production and profitability. The major
component of embryo loss occurs before day 16
following breeding with emerging evidence of
greater losses before day 8 in high producing dairy
cows. Late embryo loss causes serious economic
losses because it is often recognized too late to
rebreed females. More balanced breeding
strategies with greater emphasis on fertility, feed
intake and energy must be developed. There is a
range of easily adoptable management factors that
can either directly increase in embryo survival or
ameliorate the consequences of low embryo
survival rates. The correction of minor deficits in
several areas can have a substantial overall effect
on herd reproductive performance.
Introduction
Embryonic mortality refers to the losses which
occur in the period between fertilization and the
completion of the stage of differentiation at
approximately day 45. Embryonic death or mortality
denotes the death of fertilized ova and embryo up to
the end of implantation. The reproductive efficiency
is affected by very well known factors like
fertilization failure and embryonic mortality, the later
being more significant. Early embryonic mortality is
a major source of embryonic and economic loss
through repeat breeding and increased cost of
Vol.VI | Issue 1 | Sep. 2016
LARGE ANIMAL SECTION
An Overview
artificial insemination. EED also leads to extended
calving intervals and prolonged dry period resulting
in reduced life time milk production and reduced net
calf production (1). Embryonic mortality (2) is
classified as Very Early Embryonic Mortality (day 0
to 7), Early Embryonic Mortality (day 7 to 24), Late
Embryonic Mortality and Early Fetal mortality (days
24 to 285). Mortality is more common during early
than the late embryonic period, i.e. from day 8th to
16th at the hatching of blastocyst and initiation of
elongation and commencement of implantation (3).
About 80 per cent of this loss occurs before days 16
to 17; 10 to 15 per cent between days 17 to 42 and 5
per cent after day 42.
Normal Embryonic Development
An embryo is a product of fertilization characterized
by growth and differentiation leading to the
establishment of different organ systems that make
up the individual. Fertilization in the bovines occurs
at ampullary isthmus junction of the fallopian tube
and the embryonic period can be divided into
zygote, cleavage, morula, blastocyst, implantation
and post-implantation. The zygote being the first
structure formed as a result of successful
fertilization cleaves mitotically into 2, 4, 8 and 16 cell
stages. The cleaved embryo enters the uterine horn
at the morula (mass of cells within the zona
pellucida) stage about 4 to 5 days after fertilization.
The blastomeres on the surface of these mass form
tight junctions to create an outer membrane,
therefore partitioning embryo into two major classes
of cells; inner-inner cell mass and outer
trophoectoderm later on trophoblast. The morula
6
RAKSHA TECHNICAL REVIEW
then develops into blastocyst (having distinct
blastocoele, trophoblast and embryonic disc) at
days 6 to 7. The zona pellucida ruptures, resulting in
hatching of the embryo after 9 to 10 days. The
hatched blastocyst begins a process of elongation
from about day 12 to 13, which is accompanied by
the secretion of embryonic interferons. Early
attachment (apposition) of the conceptus to the
endometrium takes place from about day 19 and
actual adhesion occurs by day 21 to 22.
Causes of Embryonic Mortality
1. Genetic Factors
Genetic abnormalities account for approximately 10
per cent of the embryonic losses and generally
result in pregnancy failure within the first two weeks.
Expression of lethal genes can cause death of the
embryo within the first 5 days of pregnancy. Another
genotypic factor contributing to embryonic death is
an abnormal chromosome number in some or all of
the embryonic cells that results in abnormal growth
of the embryo, and usually death within the first
trimester of gestation
2. Chromosome Abnormalities
2.1 Numerical abnormalities
2.1.1 Aneuploidy
The chromosome number is almost diploid but
there are one or two chromosomes too many or too
few. Aneuploidy arises if there is non-disjunction
during meiosis so that the chromosomes do not
separate in a balanced fashion. In females, X
chromosomal aneuploidy occurs like Turner's
syndrome (XO) and Triple X syndrome (XXX).
Aneuploidy of the autosomes results in either too
many (trisomy) or too few (monosomy) copies of a
particular chromosome and its associated genes.
2.1.2 Polyploidy
There are whole multiples of the haploid (i.e. half the
diploid) chromosome number in excess, e.g.
triploidy is three times the haploid number, and
tetraploidy is four times the haploid number.
Polyploidy arises when there is a failure of the block
to polyspermy or if there is retention of the first or
second (or both) polar bodies during oogenesis.
2.2 Structural Abnormalities
It depends upon whether genetic material has been
lost (deletions) or just rearranged (insertions,
inversions and translocations). The commonest
chromosomal abnormality in cattle is a structural
anomaly known as a centric fusion translocation.
Two chromosomes fuse together near the
centromere, resulting in a reduction in the
chromosome number but little or no loss in genetic
Vol.VI | Issue 1 | Sep. 2016
LARGE ANIMAL SECTION
material. The best characterized chromosomal
abnormality in cattle is the 1/29 Robertsonian
translocation, found in various breeds worldwide. It
is characterized by a fusion between two non
homologous chromosomes (1 and 29), resulting in
a single chromosome. As for other Robertsonian
translocations, carriers produce six different types
of gametes, only two of which would produce viable
offsprings, while the other four types would produce
chromosomally unbalanced embryos. Death of
these embryos would occur within the 1st or 2nd
week of development.
3. Endocrine Factors
Failure of release of LH results in delayed ovulation.
Since there is delay, the sperm cells have become
poorly viable as they do in the cow within 24 to 48
hours and because of the aged conditions of the
sperm cells, early embryonic death may result.
Injections of estrogen at the time of estrum or within
several days after ovulation will affect the transport
of the fertilized ova in the oviduct resulting in too
rapid transport or tubal locking of the ova and death
of the zygote. It has been recently proven that the
relationship between progesterone and estradiol in
circulation during the first two week post
insemination play a crucial role in the maintenance
of luteal function and therefore pregnancy itself. A
new wave of ovarian follicles grows during the luteal
phase, even if fertilization has been successful. If
estradiol production of these growing luteal phase
follicles is not diminished early enough, luteolysis is
triggered and corpus luteum resolved. A decreased
level of progesterone also results in EEM. There
are two major reasons for a lack of progesterone.
Corpora Lutea (CL) have a short lifespan (6 to 12
days). Thus, luteolysis occurs before the embryo
has time to signal its presence through secreting
bTP-1. The second category includes those CLs
that have a normal lifespan (more than 14 days) but
secrete low levels of progesterone, which does not
suppress the luteolytic affects of the prostaglandin.
4. Nutritional Factors
4.1 Effects of Energy and Protein
Dietary energy and protein levels play a role in
pregnancy success. Cows will have less embryonic
mortality if they are gaining condition, while those
losing condition will tend to have higher embryonic
loss. Decreased progesterone levels following
breeding may be responsible for the decreased
fertility/ increased embryonic mortality among cows
bred during a negative energy balance. Feeding
fishmeal has also been demonstrated to suppress
oxytocin induced prostaglandin secretion in heifers
with low progesterone concentrations suggesting it
may improve an embryo's ability to signal maternal
recognition of pregnancy.
7
RAKSHA TECHNICAL REVIEW
4.2 Effects of Toxins
Some of the more common plant toxins that can
cause reproductive problems include mycotoxins,
endophyte infected fescue, nitrates, locoweed, and
ponderosa pine. Mycotoxins can occur in moldy
feed and mycotoxin 'zearalenone' is also suspected
to cause abortions in cattle by decreasing
progesterone concentrations. Avoiding or limiting
the amount of moldy feed in the diet of pregnant
cows is the simplest way to avoid this problem.
Nitrate itself is not particularly toxic to animals.
Nitrates consumed by ruminants are normally
reduced to ammonia and then absorbed and
excreted, thus converted by bacteria into bacterial
protein. Nitrate poisoning occurs when nitrate
consumption in feed and water is sufficient such
that nitrates are converted to nitrites. Nitrites bind to
hemoglobin and decrease its oxygen carrying
capacity, which could lead to embryonic death in
less severe cases. Elevated nitrates are often found
in plants that are stressed by poor growing
conditions such as drought, cool and cloudy
weather, hail damage and frost. Plants such as
oats, millet, sorghum and corn are especially
susceptible to high nitrates and should be tested for
nitrate content before grazing/feeding. Diets of
pregnant cows should not exceed 5000 ppm
nitrates on a dry matter basis.
4.3 Excesses of Protein
Crude protein in the total diet greater than 17 to 20
per cent has been implicated in lowering conception
rates with increases seen in the number of services
per conception and days open. These observations
have not been consistent and may depend more on
the amount and type of protein and amount of
energy supplied in the diet than the amount of crude
protein. When an excess of degradable protein
and/or a deficiency of energy is fed, ammonia not
incorporated into microbial protein is absorbed into
the blood stream. In turn, this excess ammonia and
urea in the blood stream may decrease fertility, at
the same time energy is diverted away from milk
production and/or reproduction. Some studies have
indicated that blood urea nitrogen (BUN) above 20
mg/100 ml may decrease the chances of
pregnancy.
5. Macro - Minerals
Calcium deficiency in young calves prevents
normal bone growth and retards their general
growth and development. In cows with milk fever,
the uterus returns to normal size more slowly after
calving. These cows also have an increased
incidence of dystocia (trouble giving birth), retained
placenta, and prolapsed uterus. The ratio of calcium
to phosphorus in milking cow diets should be kept
between 1.5 to 2.5. However one must remember
Vol.VI | Issue 1 | Sep. 2016
LARGE ANIMAL SECTION
that the amount of each of these minerals is more
important than the calcium to phosphorus ratio.
Phosphorus deficiencies decrease fertility, feed
intake, and milk production. At the same time, cattle
may be lethargic and unthrifty. Usually other
deficiency symptoms are seen before infertility. In
particular, phosphorus deficiency results in a lower
conception rate, a decrease in ovarian activity,
irregular estrus cycles, anestrous (inactive ovaries)
and an increased incidence of cystic ovaries.
6. Trace Minerals
Iodine has an indirect effect on reproduction
through its action on the thyroid gland. Iodine
deficiencies may indirectly cause early embryonic
death, abortion, stillbirths, prolonged gestation
(time the calf is carried in the uterus), and an
increase in the incidence of retained placenta.
Retained placentas are the most common
reproductive problem associated with selenium
deficiency. But early embryonic deaths, increased
metritis, poor fertility and the birth of dead or weak
calves also are associated with low levels of
selenium. Rations fed to dairy cows should be
supplemented with the legal limit of 0.3 ppm
selenium in the total ration dry matter. Dairy farmers
should remember that a selenium deficiency is not
the only mineral and vitamin deficiency that can
cause retained placentas. Deficiencies of calcium,
copper and iodine increase the incidence of
retained placentas. A deficiency of copper is
associated with early embryonic death, reduced
ovarian activity, delayed or reduced estrus activity,
decreased conception rate, increased incidence of
retained placenta and increased difficulty in calving.
The availability of copper is reduced by excesses of
calcium, sulfur, iron, zinc, and molybdenum in the
diet or water.
7. Vitamins
Vitamin A is the most commonly deficient vitamin in
cattle. Vitamin A is necessary in maintaining the
health and integrity of epithelial tissue (tissue that
lines the reproductive tract, intestinal tract, urethra,
kidney, mouth, respiratory tract, salivary glands,
eyes, and tear glands). These tissues become hard
and crack during a Vitamin A deficiency. Deficiency
of Vitamin A and its retinoid derivatives cause early
embryonic death. Deficient cattle are highly
susceptible to infections and colds, and pneumonia
often occurs. Reproductive problems associated
with a Vitamin A deficiency include delayed sexual
maturity, abortion, and birth of dead or weak calves,
retained placenta, metritis, and shortened gestation
periods. Supplementation with vitamin A should be
considered when feeding poor quality forages or
low amounts of forage and corn silage. Beta-
carotene is a precursor of Vitamin A. Early research
showed that reproductive performance was
8
RAKSHA TECHNICAL REVIEW
improved when cows were supplemented with
beta-carotene. Deficiency of Vitamin E and
Selenium leads to embryo loss at the time of
implantation.
8. Importance of Energy
Energy is the most common nutrient limiting
reproduction. During early lactation, the peak in
feed intake lags behind the peak in milk production.
Milk production usually peaks approximately six
weeks into lactation, whereas feed intake peaks
four weeks later at ten weeks after calving. During
this ten-week period, the cow is in a negative energy
balance (i.e. more energy is being put into the
production of milk than is taken in through her diet).
To meet their energy needs, cows rely on their body
stores of fat (known as body condition). Cows that
lose an excessive amount of body condition or fat
stores during early lactation have longer intervals to
first ovulation and first estrus (heat period), lower
first service conception rates and more days open.
Cows should not lose more than 1.0 unit of body
condition when scored on a scale of 1 to 5. On this
scale, cows should calve with a body condition
score of 3.5 to 4.0 and milk down to a 2.5 or 3.0.
Improvements in a cow's energy balance may be an
important signal to the ovaries to start cycling. Cows
may start cycling when they are still in negative
energy balance but are starting to return to a
positive value. When a cow is returning to a positive
energy balance, the magnitude of the negative
energy balance may be important in determining
the amount of time it takes for her to start cycling
again. Preliminary results from research trials
indicate that energy balance may also influence
developing ova (eggs).
9. Temperature or Heat Stress
Among all environmental stressors, the
temperature and relative humidity are the major
factors, which affect the reproductive performance
of dairy cows. Incidence of lowest conception rate,
longer calving to conception and calving intervals
are more during summer months. Percentage of
abortion and retained placenta were highest for
cows calving during summer. Fertility of dairy
animals is markedly declined during summer
season (4). Heat stress can act in more than one
way to reduce fertility in lactating dairy cows. Heat
stress can reduce dry matter intake to indirectly
inhibit GnRH and LH secretion from the
hypothalamo-pituitary system. However, it is not
clear if heat stress can also directly influence the
hypothalamo-pituitary system to reduce GnRH and
LH secretion. Heat stress can directly compromise
the uterine environment to cause embryo loss and
infertility.
Vol.VI | Issue 1 | Sep. 2016
LARGE ANIMAL SECTION
10. Genital Infections
Infection of the embryonic environment can be
caused by specific and non-specific uterine
pathogens. Specific uterine infections are caused
by a number of viruses, bacteria and protozoa that
enter the uterus by the haematogenous route or via
the vagina. Non-specific pathogens are mainly
bacteria that enter the uterus by ascending
infection. Uterine pathogens may cause EM by
changing the uterine environment (endometritis) or
by a direct cytolytic effect on the embryo. Non-
infectious causes of EM such as chromosomal
aberrations, external factors (e.g. high ambient
temperature and nutritional factors) and maternal
factors (e.g. hormonal imbalances and age) are
multifactorial and difficult to diagnose.
11. Venereal Diseases
11.1 Trichomoniasis (Trichomonas fetus)
In the cow, T. fetus colonises the uterus, cervix and
vagina, but it survives poorly on the vulva. Within
the uterus, the organism produces a catarrhal
endometritis and vaginitis, with edema of vulva,
perivaginal tissue and uterine wall. It does not
generally invade through the epithelial surface. The
disease does not prevent fertilization, but causes
embryonic death at an early stage of gestation.
Typically, embryonic death occurs after the
maternal recognition of pregnancy (day 16),
causing an irregularly extended return to estrus,
although some animals exhibit normal, or even
short, returns to estrus. Embryonic death is not
infrequently (up to 10 % of cases) accompanied by
the development of pyometra, in which the uterus is
filled with enormous quantities of trichomonad and
vaginal discharge of thinnest pus is common.
11.2 Campylobacteriosis (Campylobacter fetus or
Vibrio fetus)
Vibrio fetus is responsible for infertility and causes
early embryonic mortality in cows. The sites of
infection in the cow are the vagina, cervix, uterus
and uterine tubes. The organism causes no lesions
of the vagina, but can persist in that site for some
time. In a much smaller proportion of infected cows,
later abortion occurs between 4 and 7 months
causes a mild endometritis.
11.3 Corynebacterium
Corynebacterium pyogenes is the most common
bacteria found in the uterus of cattle. This bacterium
is the largest cause of endometritis in cows. Coryn.
pyogenes is responsible for aborted embryos,
fetuses, placenta and vaginal discharge.
11.4 Viral Agents
Bovine herpes virus-1 (BHV-1) is a group of viruses
9
RAKSHA TECHNICAL REVIEW
that includes IBRV and IPV. This group is
responsible for more abortions than any other
infectious agent. BHV-1 also causes early
embryonic death, cystic CLs and some pathological
changes to the uterus. Bovine viral diarrhea (BVD)
has been shown to cause early embryonic loss, but
is not a major cause of embryonic mortality in cattle.
12. Uterine Environment
After fertilization, embryos cleave at different rates,
sometimes causing the maturity of the embryo to
differ from that of the uterus. The uterine
environment may be toxic to these embryos that are
out of phase, resulting in the death of the embryo.
Maternal recognition occurs around days 15 to 17 of
pregnancy. The embryo secretes bovine
trophoblast protein-1 (bTP-1) which alters the
production of prostaglandins and prevents
luteolysis from occurring. If the embryo does not
signal its presence adequately to the mother, the
mother will continue to cycle as if open.
13. Immunological Factors
If the mechanism of immunosuppression is not
going well, then the antibodies will interfere with the
development of the embryo in the uterus. The feto-
placental unit can be considered as a foreign body
in the uterus, paternal antigens are normally not
rejected by the maternal immune system.
Therefore, the immune system is involved in the
successful outcome of the pregnancy by creating
conditions that prevent rejection of the conceptus.
Inappropriate immune responses could lead to
rejection of the conceptus and cause embryonic
mortality.
14. Effect of Palpation
Generally, rectal palpation does not affect the
survivability of embryos if palpated gently. Palpation
between days 34 and 41 of pregnancy using the
fetal membrane slip technique did not affect
embryo/fetal viability (5). Through palpation,
producers can identify open cows or problem
breeders and can cull cows that do not meet herd
requirements.
15. Improper Timing of AI
Poor timing of insemination in regard to ovulation
and aging of ova or spermatozoa or both occurs as
a result of improper timing of heat detection.
16. Effect of the Male
There are two areas that the male can contribute.
The first area is genetic, where the male transmits
possible lethal genes, abnormal chromosome
numbers or some genetic mutation. This area
probably cannot be detected from a management
standpoint, while the second area can be controlled
Vol.VI | Issue 1 | Sep. 2016
LARGE ANIMAL SECTION
through vaccination. Bulls that are infected with
some infectious agent like vibrio can spread this
infection throughout the herd.
17. Other Possible Factors
Heifers are generally considered to have higher
pregnancy rates and this increase seems to be
associated with less embryonic mortality than cows.
Older cows nearing the end of their reproductive life
will also have an increase in embryonic mortality.
Among dairy cattle, both early and late embryonic
losses increased among cows with increasing age
(6). Little difference in embryonic mortality across
breeds has been shown. Cattle that are line bred or
inbred have been noted to have an increased rate of
embryo mortality.
Diagnosis
1. Examining Embryos
Examining embryos collected by in vivo flushing of
reproductive tract at different days after breeding.
2. Determining Progesterone in Blood, Milk and
Saliva
Determination of progesterone in blood and milk, 21
days after estrus is the most common method used
in the pregnancy diagnosis of ruminants until the
early nineties. A study (7) on monitoring of early
pregnancy and embryonic mortality using blood
progesterone concluded that it is impossible to
determine embryonic mortality alone on the basis of
progesterone profile while pregnant and non-
pregnant cows can be easily distinguished 21 day
post AI. They also concluded that it is very easy and
accurate to distinguish non-pregnant cows from
cows that have suffered early embryonic mortality.
CL dysfunction in reproduction is identified by either
of two criteria namely estimation of plasma or
salivary P4 levels or endometrial biopsy. Two or
three plasma samples could be processed for the
assay. Progesterone levels should not be less than
10 ng/ml during the luteal and mid luteal phase.
3. Pregnancy Associated Glycoprotein (PAG)
Test
The main advantage of Pregnancy Specific
Proteins (PSP) for pregnancy diagnosis in cattle is
their ability to prove the existence of placentation
and the presence of live, vital embryos, while
progesterone only proves the existence of corpus
luteum. The most commonly used pregnancy
specific protein for pregnancy diagnosis in cows is
PAG (pregnancy-associated glycoprotein).
Pregnancy-associated glycoprotein has been
found in the serum of pregnant cattle and used as a
pregnancy marker. As pregnancy failure occurs,
PAG concentrations drop and disappear from
maternal blood. The Pregnancy Associated
10
RAKSHA TECHNICAL REVIEW
Glycoproteins (PAG) is synthesized by the
ruminant's trophectoderm. A part of it is released
into maternal blood circulation which can be
assayed by RIA and ELISA. RIA methods are very
precise for measuring PAG concentrations in the
maternal blood and milk of the ruminants. The
sensitivity and specificity of this method are very
high. The results are encouraging and use of milk
and in blood for PAG test is helpful in detection of
embryonic mortality in the ruminants.
Treatment
Various methods have been tried using different
preparations for improving pregnancy rates by
reducing embryonic mortality.
1. Supplementing Progesterone/Progestogen
Research studies have reported that low
concentration of progesterone can result in the
development of a stronger luteolytic signal and
hence it might be concluded that cows with lower
plasma concentrations are apparently more prone
to embryo loss. A significant increase (8) in
conception rate among cattle was recorded when
Crestar ear implants (Norgestomet) were given on
day 7.
2. Use of hCG
An alternative approach to increase the
progesterone levels is by use of Human Chorionic
Gonadotropin (hCG) to enhance the production of
progesterone by the animals own corpus luteum.
Administration of human chorionic gonadotropin
(hCG) induces ovulation with the subsequent
formation of a functional accessory CL which in turn
increases progesterone and may enhance embryo
survival. Studies have supported that conception
rate is better in cows with three follicular waves after
insemination as compared to cows with two
follicular waves and hCG induction of three-wave
cycles may also contribute to higher pregnancy
rates. It was demonstrated that injecting 3300 IU of
hCG in lactating cows 5 days after AI resulted in
increased number of CL and higher plasma
progesterone concentrations (9). Conception rates
on days 28, 42 and 90 were improved by hCG
treatment. The findings of (9) were supported by
findings of (10) as hCG administered on day 6
increased the pregnancy rates (67.50 %) with
formation of accessory corpora lutea as compared
to control cows (45.0 %) or cows receiving hCG on
day 1 (42.50 %).
3. PMSG Administration
Luteotropic effect of PMSG in cattle was studied
and it was reported significant increase in
progesterone concentration on administration of
500 IU of PMSG on day 7 after estrus (11). They
Vol.VI | Issue 1 | Sep. 2016
LARGE ANIMAL SECTION
concluded that PMSG treatment increases
progesterone secretion and luteal function without
excessive follicular development.
4. GnRH Treatment
Administration of GnRH (250 μg) at the time of
insemination increases pregnancy rates by 12.5 per
cent and effect was more pronounced (12) in repeat
breeder cows. Administration of GnRH at estrus
increases serum progesterone levels and the
proportion of large luteal cells in the corpus luteum.
Conclusion
Efforts should be made to provide appropriate
protection against high or low temperatures,
effective vaccination and provide clean
environment and proper timing of insemination. The
use of highly fertile semen, appropriate semen
thawing techniques and good semen handling and
placement are also critically important to optimize
fertility and to reduce embryonic mortality.
References
1. Maurer, R.R. and Chenault, J.R. (1983). Fertilization
failure and embryonic mortality in parous and non-
parous beef cattle. J. Anim. Sci., 56: 1186-1189.
2. Walsh, S.W., Williams, E.J. and Evans, A.C.O. (2011). A
review of the causes of poor fertility in high milk
producing dairy cows. Anim. Reprod. Sci., 123: 127-
138.
3. Dunne, L.D., Diskin, M.G. and Sreenan, J.M. (2000).
Embryo and fetal loss in beef heifers between day 14 of
gestation and full term. Anim. Reprod. Sci., 58: 39-44.
4. Ghuman, S.P.S. and Singh, J. (2009). Proceedings of
Interactive meet on buffalo reproduction. CIRB, Hisar,
Haryana. P. 25-31.
5. Romano, J.E. and Magee, D. (2001). Applications of
trans-rectal ultrasonography in cow/heifer
reproduction. Proceedings of Annual Food Conference
Conception to Parturition: Fertility in Texas Beef Cattle,
Texas A and M University, USA. P. 99-104.
6. Humblot, P. (2001). Use of pregnancy specific proteins
and progesterone assays to monitor pregnancy and
determine the timing, frequencies and sources of
embryonic mortality in ruminants. Theriogenology, 56:
1417-1433.
7. Prvanovic, N., Tomaskovic, A., Grizelj, J., Kocila, P. and
Samardzija, M. (2009). Monitoring of early pregnancy
and early embryonic mortality by ultrasound and
determination of pregnancy-associated glycoproteins
and progesterone in cows. Vet. Arch., 79: 259-267.
8. Broadbent, P.J., Sinclair, K.D., Dolman, D.F., Mullan,
J.S. and McNally, J.R. (1992). The effect of a
Norgestomet ear implant (Crestar) on pregnancy rate in
embryo transfer recipients. Proc. 12th Int. Congress
Anim. Reprod., 2: 782-784.
9. Santos, J.E.P., Thatcher, W.W., Pool, L. and Overton,
M.W. (2001). Effect of human chorionic gonadotropin
on luteal function and reproductive performance of
11
 RAKSHA TECHNICAL REVIEW LARGE ANIMAL SECTION

high-producing lactating Holstein dairy cows. J. Anim.
Sci., 79: 2881-2894.
10. Nishigai, M., Kamomae, H., Tanaka, T. and Kaneda, Y.
(2002). Improvement of pregnancy rate in Japanese
Black cows by administration of hCG to recipients of
transferred frozen-thawed embryos. Theriogenology,
58: 1597-1606.
11. Hirako, M., Karmomae, H. and Domeki, I. (1995).
Luteotrophic effect of pregnant mare serum
gonadotrophin in cattle. J. Vet. Med. Sci., 57: 317-321.
12. Morgan, W.F. and Lean, I.J. (1993). Gonadotrophin-
releasing hormone treatment in cattle: A meta-analysis
of the effects on conception at the time of insemination.
Aust. Vet. J., 70: 205-209.

1 to 1½ month of bovine embryo Fetus contained within the discrete spherical

amniotic sac

Bovine conceptus at 60 days Fetus and fetal membranes formation

Formation of cotyledons and

2 ½ month of bovine embryo
surrounding structures

Vol.VI | Issue 1 | Sep. 2016 12

View publication stats