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Evaluation
of Frozen-Thawed Semen
T Katila, E Koskinen* and M Andersson
Department of Clinical
Veterinary Sciences, Saari Unit, Finland Saarentaus
*Department of Animal Science, Finland, Helsingin Yliopisto
University of Helsinki
Introduction
There
is considerable variation between individual stallions in how well their
semen retains its fertilizing capacity after freezing and thawing. It has
been estimated that only 20% of fertile stallions produce sperm that
survive the freezing and thawing processes (Tischner
1979). Although our knowledge and techniques have improved within
the last 20 years, a considerable proportion of stallions are still not
suitable for semen freezing. Development of freezing methods requires in
vitro tests that correlate with in
vivo fertility, but controlled breeding trials with horses are
expensive (Loomis 1999). The slow progress in the development of freezing
techniques is partly explained by the lack of reliable laboratory methods.
Some in vitro methods work
reasonably well in the assessment of fresh but not frozen semen, the best
example being motility evaluation. In spite of its limited applicability,
motility is the most commonly used parameter in the evaluation of
frozen-thawed semen, in both laboratories and stud farms, because it is
easily accessible and quick to perform. A combination of laboratory tests
should enable better assessment of the fertility of cryopreserved stallion
semen (Blach et al. 1989).
There
is very little data about the correlation between semen evaluation tests
and pregnancy rates after insemination with frozen semen. Although good
correlations have been found between different tests, it does not mean
that they would correlate with fertility.
The purpose of this study was to find out which tests used to
evaluate stallion semen would correlate with foaling rates of mares
inseminated with frozen semen. Our previous study indicated that
incubation of the semen sample at +37oC might be useful in the
evaluation of frozen-thawed semen (Katila
et al. 1999). Therefore, the evaluation tests were performed
immediately after thawing and after 3 hours incubation at +37oC.
Materials and Methods
Frozen semen doses were available from 31 commercially used
stallions. The semen had been frozen between the years 1988 and 1998 in 5
different countries: 22 in Sweden, 5 in Finland, 2 in Italy, 1 in Denmark,
and 1 in Germany. Semen from 23 stallions had been frozen in 2.5-ml straws
and 8 in 0.5-ml straws. Thirteen stallions were American Standardbreds,
one stallion was an English thoroughbred, and the remaining 7 were various
types of riding horses. Sufficient foaling data – at least 5 mares and
10 cycles – were available from 23 stallions. Foaling data originated
from Finland and Sweden from the years 1989 to 1999. The average number of
mares/stallion was 35 (range 5 – 121). The average foaling rate was 59%
(range 11 – 91%). Seventeen stallions had a foaling rate >50% and 6
had a foaling rate <50%.
Semen evaluation
The
following tests were done from 1 to 3 straws on one day immediately after
thawing and after 3-hour incubation at +37oC extended in
Kenney’s skim milk extender (Kenney
et al. 1975): 1) resazurin reduction test by a fluorometer, 2)
plasma membrane integrity by CFDA/PI (carboxyluoresceindiacetate/propidium
iodide) staining and counting of cells in a fluorescence microscope, and
3) plasma membrane integrity by PI staining and by an automatic
fluorometer. In addition, 4) concentration was determined in a Bürker
counting chamber. On another day the following tests were done immediately
after thawing and after 3-hour incubation at +37oC extended in
Kenney’s skim milk extender: 5) motility by a light microscope and 6)
motility parameters by a Hamilton Thorn Motility Analyzer, 7) HOST
(hypo-osmotic swelling test) by light microscope and 8) HOST by a
fluorometer. In addition, concentration was determined in a Bürker
counting chamber.
The
0.5-ml straws were thawed for 30 sec in +37oC and the 2.5-ml
straws in +50oC for 40 sec. Semen concentration was measured in
a Bürker counting chamber, and the total number of spermatozoa/straw
calculated. An insemination (AI) dose was one straw when using 2.5-ml
straws and from 1 to 10 straws for the 0.5-ml straws. The sample for the
longevity test was prepared by extending semen with skim milk extender to
a concentration of 40 to 100 x 106 spermatozoa/ml. The sample
was kept in a water bath at +37oC for 3 hours.
For motility evaluation, semen was extended with warm (+30oC)
skim milk extender (Kenney et al.
1975) to a concentration of 40 x 106 spermatozoa /ml and
incubated at +37oC for 10 min. An aliquot of 7 ml
of extended semen was placed on a slide and covered by a cover slip.
Motility was evaluated with a light microscope: percentage of
progressively motile spermatozoa, percentage of total motility and a
velocity score from 1 to 3. Motility was measured also with an automatic
sperm analyzer (Hamilton Thorn Motility Analyzer, HTM-S, version 7.2). A
10-ml
semen sample was placed onto a Makler chamber, and two chambers were
prepared from the same sample. Six fields/chamber were videotaped for 15
sec/field. The videotapes were analyzed for total motility, progressive
motility, and path velocity.
For
CFDA/PI staining, semen was extended with skim milk extender to a
concentration of 50 x 106 spermatozoa/ml. An aliquot of 20 ml
of CFDA stock solution consisting of 0.46 mg CFDA in 1 ml of DMSO (dimethylsulphoxide)
and 10 ml
of PI stock solution (0.5 mg PI in 1 ml of 0.9% NaCl-solution) were taken,
mixed with 950 ml
of skim milk extended semen and incubated for 8 min at +30oC. A
drop of 5 ml
was placed on a slide and overlaid with a cover slip (Harrison and Vickers 1990). The proportion of fluorescent cells
was counted in 200 cells in a fluorescence microscope (Olympus BH2 with
epifluorescence optics) using oil immersion and a fluorescein filter set.
An
automatic fluorometer (Fluoroscan Ascent) which reads a 96 well
microtitration tray and has an incubation compartment was used for the
fluorometric measurement of plasma membrane integrity. The interference
filter at the exitation path and that of the emission filter had a maximum
transmission at 544 nm and 590 nm, respectively. For the fluorometric
assay, 20 mg of PI was dissolved in 1 liter of BTS (Beltsville Thawing
Solution) and dispensed in 3 ml aliquots. Equal aliquots (50 ml)
of skim milk extended semen sample (40 x 106 spermatozoa/ml)
and PI solution were dispensed into the well plate, and the well was
shaken gently for 2 min. Spermatozoa from the same samples were killed by
unprotected rapid freezing-thawing to have internal control samples
consisting of only non-viable cells (100% fluorescence).
The control sample was immersed into liquid nitrogen for 1 min,
thereafter it was allowed to stand in a room temperature for 30 sec and
then 3 min in a water bath (37oC). Blanks containing 50 ml
of diluted extender and 50 ml
of PI were analyzed separately for every experiment in 4 replicates. The
incubation time was 8 min. The percentage of fluorescence was calculated
from the ratio of fluorescence intensities of the rapidly frozen control
sample and the sample to be analyzed, taking into account the blank values
(Juonala et al. 1999).
For
the resazurin reduction test, 400 mg of resazurin was dissolved in 1 liter
of distilled water. One part of this solution and 9 parts of 0.9% NaCl
were mixed. An equal volume of this mixture and skim milk extended sperm
(40 x 106 spermatozoa/ml) were combined and shaken for 2 min.
After that it was incubated for 30 min at 32oC and measured
with the fluorometer using the same fluorometer settings as in the plasma
membrane viability test (Eriksson
et al. 1998).
Hypo-osmotic
solution of 100 mOsmol/kg for HOST was prepared by dissolving 4.5 g
fructose and 2.717 g sodium citrate in 500 ml of distilled water. An
aliquot of 0.5 ml of this solution was mixed with 0.125 ml of semen and
incubated at 37oC for 45 min. A 5 ml
drop was placed on a slide and overlaid by a cover slip. A total of 200
spermatozoa per sample were evaluated for bent tails in the microscope.
For the fluorometric determination of HOST 0.5 ml of the same hypo-osmotic
solution (100 mOsmol/kg) was mixed with 0.125 ml of skim milk extended
semen (concentration 100 x 106 spermatozoa/ml). The
fluorometric method was the same as for PI-stained semen.
Data
analysis
Raw
correlations were determined by analysis of least squares. The level of
significance was set at p<0.05. Only the results of the 23 stallions
having sufficient foaling data were included in the statistical analysis.
Results
None
of the tests had high correlations with foaling rates. Only HOST performed
at 0 (p = 0.03) and 3 hours (p = 0.04) correlated with fertility. When two
tests were combined, HOST and CFDA/PI staining at 0 and 3 hours showed
significant correlations (p<0.05).
Conclusions
None
of the parameters measured correlated well with foaling rates. It seems
that tests evaluating plasma membrane integrity (HOST, fluorescent stains)
might be more useful in the evaluation of frozen-thawed stallion semen
than motility parameters. However, many factors may have influenced the
foaling rates. Sperm had been frozen over many years in different
laboratories. The number of mares inseminated/stallion was small and
inseminated in various conditions over many years. Futhermore, pregnancy
rates/cycle would have reflected fertility more accurately than foaling
rates, but these figures were not available.
References
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RAP, Vickers SE 1990. Use of fluorescent probes to assess membrane
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