BIOLOGY
OF
REPRODUCTION
286 -292
18,
Composition
of Rabbit
(1978)
and the
Semen
W. HOLTZ’
and
Department
Origin
of Several
Constituents
R. H. FOOTE
of Animal
Science,
Cornell
University,
Ithaca,
New
York
14853
ABSTRACT
Semen
was collected
twice
a day every
other
day (2X/48
h) for 36 days from
12 Dutch-belted
rabbits.
Sex drive,
ejaculate
volume,
microscopic
appearance
of the semen and sperm
counts
were
recorded.
Seminal
plasma
was analyzed
for it content
of fructose,
citric
acid,
glycerylphosphorylcholine
(GPC)
and the minerals
Na, K, P, Mg, Ca and Zn. Six males were exsanguinated
24 h
after
the last semen
collection
and 6 were
exsanguinated
5 weeks
later.
The
organs
of the
reproductive
tract
were
weighed
and analyzed
for their
contents
of spermatozoa,
fructose,
citric
acid and GPC. Rabbits
differed
in libido
and several
characteristics
of semen.
Second
ejaculates
had
lower
concentrations
of fructose,
citric
acid,
K and Zn than did the first ejaculates.
Fructose
was
found
chiefly
in the different
portions
of the prostate
gland and major
concentrations
of citric
acid
were found
in the ampulla
of the ductus
deferens
and in the vesicular
gland.
GPC concentrations
were highest
in the caput
and cauda epididymides.
Correlations
among
different
constituents
were
calculated
and their significance
is discussed.
INTRODUCTION
The
composition
has been
chemical
domestic
animals
(Mann,
1964),
been
made
of
despite
widely
but
the
the fact
used
in
of semen of many
studied
extensively
few systematic
chemistry
of
sperm
production,
(Macmillan
and
and
Desjardins
et
the
conditions
tained.
Except
seminal
and
1966;
Skinner,
al.,
perhaps
due
1968),
plasma
(Quinn
Kihlstr#{246}m, 1975)
Glover,
content
of
(Jones
mineral
and
of
reproductive
the
literature.
to
from
The
quantify
rabbits
et a!.,
and
1965;
epididymal
1973),
no
rabbit
seminal
purpose
constituents
ejaculated
tract
of
was
the
1964;
1967;
in
to
obin
Fjellstrom
fluids
following
data
to
stabilized
collection
156
first
study
on the
plasma
fluids
found
present
semen
on a controlled
Accepted
August
31, 1977.
Received
May 10, 1977.
‘Present
address:
Institut
f#{252}r
Tierzucht
riergenetik
der Universit#{228}t Gdttingen,
34
Albrecht-Thaer-Weg
1, West Germany.
in
study
the
of
male
these
repro-
AND
METHODS
Twelve Dutch-belted
male
rabbits
16-19
months
of age weighing
2.2 (2.1-2.4)
kg were caged individually at 20#{176}Cand exposed
to 12
h of light
daily.
All
animals
were trained
to ejaculate
into
an artificial
vagina
(Bredderman
et al., 1964),
which
was filled
and
lubricated
with
chemically
inert
“360
Medical
fluid”
(Dow
Corning
Corporation).
Animals
were
sexually
rested at the start of the experiment.
Bucks
were
ejaculated
in random
order
twice a day
every
other
day
(2X/48
h) for 36 days.
They
were
allowed
2 false
mounts
with
a teaser
doe prior
to
ejaculation.
Eagerness
of the bucks
to mate with
the
teaser
was judged
subjectively
on a scale of 0-5.
A
second
ejaculate
usually
was obtained
about
2 h after
the
first
one.
The
first
5 semen
collection
days
of
in part
several
in
Rabbits
1969a;
under
which
semen
was
for
limited
data on minerals
relate
origin
MATERIALS
maturation
Hafs,
1967;
Tomizuka,
to
their
organs.
studies
have
rabbit
semen,
Orgebin-Crist,
1968;
Paufler
and
Foote,
Bedford,
1974).
Seminal
concentrations
fructose,
citric
acid and glycerylphosphorylcholine
(GPC)
vary
considerably
(Mann,
Masaki
ductive
and
to
constituents
that
it is easily
obtained
and
reproductive
studies
such
as
those
involving
and
fertilization
schedule
tion
the
was
sexual
rest were excluded
from
the semen
provide
time
for sperm
output
to become
(Holtz
and Foote,
1972).
The remaining
13
days X 12 rabbits
resulted
in a potential
of
and 1 56 second
ejaculates.
A few ejaculates
were
lost or were too small to be processed.
Six of the bucks
were given sodium
pentobarbital
anesthesia
and exsanguinated
1 day
after
the last
obtained
collec-
semen
collection
and the
rested
for 5 weeks
before
tive
organs
were
removed
analysis.
Processing
und HausGottingen,
All
with
286
Semen
equipment
the semen
and
remaining
6 were sexually
being sacrificed.
Reproducimmediately
for chemical
Tissues
and glassware
or tissues
was
that came
thoroughly
in contact
cleaned,
It
ORIGIN
OF
RABBIT
with glass redistilled
water.
Immediately
following
semen collection,
umes of the liquid
and gelatinous
portions
ejaculate
were recorded
separately.
A small
SEMEN
was rinsed
the volof the
drop of
semen
was examined
microscopically
to determine
percentage
of motile
sperm,
rate of progressive
motility, frequency
of occurrence
of proximal
and distal
cytoplasmic
droplets,
degree
of head to head
sperm
agglutination
and density
of granular
materials.
These
were scored
subjectively
from
low to high on a scale
from
0-5.
Simultaneously
the nongel
fraction
was
centrifuged
at 9000
g for 2 mm. The clear supernatant
fluid,
essentially
devoid
of spermatozoa,
was transferred
by Pasteur
pipet
into
a small
plastic
vial and
frozen
in liquid
nitrogen
until
analysis.
The sediment
was resuspended
in physiological
saline,
pooled
with
any
gel
mass
present
and
sperm
rinsed
from
the
artificial
vagina
to make total sperm counts.
Addition
of 5% (v/v) “Glasol”
(Kihlstr#{246}m and Fjellstr#{246}m, 1967)
to the
saline
helped
to dissolve
the gel mass and
facilitated
subsequent
counting
of sperm
in the hemocytometer.
In a separate
trial with
21 ejaculates
from
7 bucks,
procedural
losses of the total
number
of
spermatozoa
in the ejaculate
were
estimated
to be
1.05 ± .17% (mean
± S.E.).
No significant
changes
over
time
were
observed
with
regard
to components
of seminal
plasma.
Therefore,
apart
from
a few
particularly
small
ejaculates
that had to be excluded
for technical
reasons,
all of
the 432 ejaculates
were used for the determinations
of
organic
components
and minerals.
Three
first
ejaculates from
consecutive
collection
days and 3 consecutive
second
ejaculates
for each
rabbit
were
pooled.
This
provided
enough
material
for chemical
analysis
and
permitted
differences
among
rabbits,
first
and
second
ejaculates
and
time
trend
to be analyzed
statistically.
Seminal
plasma
was deproteinized
before
determining fructose,
citric
acid
and
GPC content.
For determination
of citric
acid,
the
method
of Saffran
and
Denstedt
(1948)
was
modified
by extending
the
heating
period
preceding
the addition
of pyridine
from
10 to 40 mm. The GPC determination,
(Lambert
and
Neish,
1950),
was adapted
to small sample
size. Two 1
ml aliquots
of deproteinized
sample
were mixed
with
1 ml of a 4% cupric
sulfate
solution
and
.15 g of
calcium
hydroxide
(White,
1959).
After
centrifugation
1 ml of the supernatant
was treated
with .2 ml of each
CONSTITUENTS
with
concentrated
hydrochloric
acid and measured
by
the method
of Kitson
and Mellon
(1944).
Procedural
losses,
particularly
in the sediment,
were determined.
This
was estimated
to be 34 ± 2%, as determined
by
loss of 32 P-orthophosphate
added
to 10 samples
of
semen
prior to centrifugation
and the usual processing.
Mineral
data were corrected
for this loss. By preparing
solutions
containing
mixtures
of ions it was ascertained
that
there
was little
interference
by related
elements.
Upon
sacrificing
the
bucks
the excurrent
ducts
were ligated
between
individual
segments
to be investigated
(Table
3) to minimize
mixing
of their
contents
during
removal
of the intact
reproductive
tract.
The
tract
was removed,
placed
on ice and dissected.
Testes,
epididymides,
ductuli
deferentia
and
accessory
sex
glands
were weighed
and homogenized
in physiological
saline.
A small
amount
of homogenate
was removed
for
counting
the
number
of spermatozoa
and
the
remainder
was frozen
in liquid
nitrogen.
Later,
the
homogenates
were thawed,
deproteinized
and assayed
for fructose,
citric
acid and GPC content
exactly
as for
the seminal
plasma.
Means
and standard
deviations
were calculated
for
all
variables.
Sources
of variation
included
in the
analyses
of variance
were groups
of rabbits,
ejaculates
(first
versus
second),
collection
days
and
rabbits
within
groups.
Correlations
between
the concentrations of various
constituents
also were computed.
RESULTS
The
collection
day
had
volume
of
(Table
droplets
in second
were
volume
ejaculates
contained
proximal
and second
the
incidence
with
sperm
The
droplets
1).
always
more
of
was
less
consistently
numbers
ejacuand
than
432
than
20%
cells
with
than
10%.
ejaculating
of attached
cytoplasmic
droplets
should
probably
be
sidered
atypical.
Head to head agglutination
of sperm
cells
was
slightly
more
frequently
common
in first
second
ejaculates
(P<0.01).
Rabbits
differed
(P<0.01)
ment.
of
semen
The
and
average
in
progressive
daily
sperm
with
comparable
conof
and
octhan in
sexual
drive,
sperm
move-
output
the last 26 days of the 36 day collection
amcfunted
to 90.2
x 106 sperm,
which
closely
it
on
Cytoplasmic
proportion
Therefore,
any
rabbit
sperm
with
appreciable
the
ejaculate
teaser
more
often
in first
but
only
8 of the
present
them.
the
to
of
in Table
a first
rapidly
First
in
on
26 days
combined
produced
the second
ejaculate.
lates
differed
greatly
droplets
Ejaculate
last
are
at
significantly
combined.
the
more
plugs
of 6
sacrificed
differ
thus
period
gel
2 subgroups
were
not
during
1. Once a male
tended
to respond
The concentrations
determined
by atomic
organic
of each
Phosform
did
were
36
groups
curred
following
ashing
and nitric
acid
oxidation
of
material.
The method
permitted
determination
cation
without
interference
by related
elements.
phorus
was converted
to the orthophosphate
and
obtained
the
eventually
times,
(P>0.05)
DISCUSSION
from
that
results
An analysis
of sediments
and supernatants
from
20
ejaculates
revealed
that 10% of the fructose,
14% of
the citric
acid
and 18% of the GPC in the semen
remained
in the sediment after centrifugation.
All data
were corrected
for losses.
of Na, K, Ca, Mg and Zn were
absorption
spectrophotometry
each
different
ejaculates
5
13.3%
AND
obtained
data
rabbits,
N sulfuric
acid,
0.1 M sodium
periodate
and
sodium
arsenite
followed
by the addition
of 5
ml of chromotropic
acid reagent.
The samples
were
cooled
in ice and held at room
temperature
for 3-6 h
to
allow
excess
of
the
chromotropic
reagent
to
precipitate.
Prior
to the determination
of the optical
density
at 580 nm the samples
were centrifuged
for 10
mm at 1000g.
of
287
observations
during
period
agrees
on
AND
HOLTZ
288
TABLE
1. Libido
and
semen
characteristics
FOOTE
12 rabbits
of
ejaculated
twice
other
every
day
for
a period
of
26
days.
First
Item
measured
Sexual
drive
(0_5)a
aSubjective
coding
bEjaculates
differed,
included
Dutch-belted
by
b).
results
of
seminal
plasma,
corrected
(1968)
Amann
and
0.2
0.6
1.2
j0b
0.4
0.6
12b
0.8
0.8
1.0
1.3
1.0
1.3
the
in Table
chemical
for
2.
Des-
and
lower
Foote
are similar
to those
(1969)
and
Desjardins
analyses
procedural
concentraas previous1967;
Mac-
presented
found
et
of
losses,
Fructose
by
Mann
2
et al.
but
are
(1964)
ejaculates
had
fraction
of first
First
ejaculates
on,
g/ml
et
Mean
SD
Mean
SD
Fructose
1000
2100
sooa
75O
1530
710
1230
1910
1400
440
1400
380
380
1000
460
1090
230
1400
100
290
100
190
differed,
al.
man and bulls,
species.
In the
(1968).
acid
the
but
rabbit
and
dogs.
were similar
(1964)
and
First
ejaculates
40
5
P<0.01.
80
9
GPC
higher
most
values
reported
3110
tion.
bull
pg/mI
by
than
in most
of the seminal
are
lower
by Dawson
Masaki
and
than
2
other
citric
the
2860
et al. (1957)
and
Tomizuka
(1966).
concentrations
in the ejaculate.
was similar
2), as this
in stallion,
No
74
were
because
the
frequently,
of
epididymal
GPC concen-
in first and second
paralleled
sperm
ejaculates
concentra-
GPC in the rabbit
is lower
than
for the
and ram,
similar
to the boar
and higher
than
80
to
by
analyses.
tration
(Table
410
120
sheep,
is concentrated
in the gel plug which,
in
present
investigation,
was excluded
from
thus
reducing
luminal
materials
Second
ejaculates
Substance
aEjaculates
horses
and
However,
this
is not
surprising
rabbits
in our study
were ejaculated
Concentrati
Ca
Zn
in swine,
ejaculates.
cattle
about
twice as high as second
ejaculates
in citric
acid
concentration
(Table
2) and
in total
ejaculate
content.
Citric
acid levels in the liquid
fraction
of the ejaculate
were
lower
than
in
iig/ml
liquid
than
second
in man,
Citric
acid
concentrations
those
summarized
by Mann
The
in the
than
higher
the
TABLE
2. Constituents
and second ejaculates.
than
is lower
Desjardins
in Table
by Macmillan
al. (1968),
than values
summarized
earlier
literature.
First
300
270
88
10
3.4
concentrations
Fructose
variable,
Skinner,
millan
et a!., 1969).
The levels
of fructose
K
P
Mg
57
0-5.
but
2050
72
456
24
1.4
0.4
(1966),
Paufler
tion in seminal
plasma
was
ly reported
(Mann,
1964;
acid
0.08
of variance.
(1969a,
The
Citric
GPC
Na
1.1
0.17
94
427
20
1.0
0.5
0.3
1 from
3.5
0.26
0.10
15
64b
385b
3.3
of
in analysis
et
lower
from
10b
0.15
021b
SD
P<0.O1.
rabbits
a!.
3.2
58
ejaculate
Mean
0.27
86
387
in units
jardins
presented
SD
0.32
54c
Ejaculates
with gel(%)
Sperm/ejaculate
(X 10’)
Sperm/mI
liquid
semen (X 106)
Motile sperm
(%)
Rate of sperm
morility
0-5)
Proximal
droplets
0-5)
Distal droplets
0-5)
Sperm
agglutination
(0-5)
Seminal
granules
(0-5)
are
Second
Mean
Semen volume,
less gel (ml)
Gel volume
(ml)
cNO
ejaculate
first
and
literature.
(Table
agreement
seminal
(1965).
dog and
man
study
of
comparable
second
ejaculates
Concentrations
2)
in
with
plasma
K
was
first
ejaculates
(Mann,
1964).
mineral
content
was found
of Na, Mg
are
in
than
reported
the
Ca
excellent
those reported
for 5 samples
analyzed
by
Quinn
et
higher
of
in
and
by
of
al.
Fjell-
ORIGIN
strom
present
in
and KihlstrOm
in nearly
equal
rabbit
Glover
(1975).
Mg
concentrations,
epididymal
(1965).
plasma
No
previous
OF
RABBIT
and
P were
as found
by
Jones
values
for
SEMEN
strated
nation
and
Zn
were
(Lindahl,
of sperm
acid,
second
other
minerals
0.40;
P, r
ejaculates,
excepting
Comparison
several
of
species
Einarsson,
for
in second
1971)
was
ejaculates.
mineral
(Mann,
K which
concentrations
1964;
reveals
Saito
that
for
et al.,
Na and
1967;
P levels
that
Mg
acid
in
and
ratio
of
resembles
and gel
-0.49).
other
rabbit
species
Na predominates.
are particularly
high
semen
in most
Mg levels in the
while
Ca is lower
than
in many
species,
with
the exception
swine.
The Zn level found
was much
below
observed
in
occurs
in
baum
1967).
humans,
rats
and
extraordinary
amounts
cations
in the
Na and
K, but
than
Thus,
for
a
from
of
seminal
1964;
prostate
Zn.
The
plasma
of
concentrations
of
the
organic
seminal
plasma
it
(Birn-
bulk
the
sexually
are
ly
of
rabbit
to be lower
reported.
osmotic
to
appears
to come
components.
Correlation
binations
of
coefficients
for
19 characteristics
measured
Ejaculate
in each
variability
producing
larger
all possible
comor constituents
ejaculate
were
computed.
was greater
than in species
ejaculates,
such
as the
bull
(Seidel
and
Foote,
1970)
and contributed
to
low
correlations
found.
Among
the r values
significant
with
at
liquid
P<0.01
semen
and with
0.20
and
correlated
were
volume,
and
motility
correlated,
agglutination
r
r
=
be
ment
0.39,
0.27
with
may
surface
the percentage
0.14, 0.43, 0.22
=
of
motile
motility
and
and
spermatozoa
were highly
by
be
and
0.30.
or 0.01),
monovalent
the
fact
activators
reaction
the
epididymides
active
rested.
reserves
cantly
than
and
Appreciable
in
the
sperm
signifisperm
content
gland
in contrast
to
and Krichesky,
1943).
Concentrations
of
of the reproductive
in the
sexual-
of
gel
=
corpora
differ
numbers
with
=
(r
suggested
less
rabbits
not
(r
sum-
epididymal
did
=
nomenand
the
in
are
that
caput
group
mixed
of
a previous
several
organic
constituorgans also are given in
Table
3. Values
close to 0 may represent
slight
interference
by
tissue
components
with
the
assay
ents
rather
than trace
in
each
tissue.
arbitrarily
reported
amounts
However,
report
these
as measured.
among
organ
of the constiturather
than
values as 0, all values are
Correlations
were com-
weights
and
all constituents
measured.
As would
be expected
related
constituents,
not highly
to gland
excepting
correlated.
weight.
Concentrations
as noted,
generally
was
Fructose
occurred
These
fructose
glands
in
were
3)
Values
Mann
to
ATP-develop-
exception
has
been
they
found
of
to
with
ejaculate
agree
closely
and his coworkers
the
be the
semen.
associated
and
the
gland
content
of
were
concentrations
consisting
of proprostate,
and bulbourethral
glands.
appear
ejaculated
(Table
highest
in
in the gland
complex
prostate,
paraprostate
creases
demon-
to
of
significantly
(P>0.10).
numbers
concentration
were
output
to 0
may
divalent
correlated
(r
GPC concentra-
sperm
Weights
2 groups
found
who
FjellstrOm
organ
analyses
The anatomical
the
the vesicular
report
(Bern
ents
and
acid
Sperm
of
were
et al. (1965)
corresponds
(1972).
of
P<0.01),
who found
no relationfructose
concentration
with
followed
were
in contrast
cations.
This
that
of
which
These
correlated
Holtz
puted
motility
0.75.
The correlations
between
and Mg, Ca and Zn were respec-
explained
cations
sperm
=
(P<O.05
significant
correlations
correlated
movement
(r = 0.18,
semen
volume
was
of
gel,
with
total
with
(r
0.24).
The percentage
and rate of spermatozoal
tively
with
rate of sperm
0.25).
Liquid
with
volume
sperm/ejaculate
rate of sperm
libido
by Quinn
0.27)
and with
citric
0.46).
The
results
of the
marized
in
Table
3.
by
Cr=0.46,
relationship
(1975)
ejaculates,
Ca, r
citric
Concentration
calculated
volume
were negatively
In second
ejaculates,
clature
the
agglutinawith the
r = 0.60; Na, r = 0.47;
K, r = 0.23),
and with
negative
was
with
despite
prevent
correlated
(r=0.35).
were
aggluticitric
correlated
P<0.05),
generally
was
(Zn,
0.37;
and KihistrOm
ship.
In first
tion
positively
to results
a
Saito
et al.,
contributes
the
tend
most
common
mammals
substantial
contribution
pressure
where
concentrations
et al., 1961;
Mann,
In these species
the
substantial
dogs,
of
that
K
in contrast
found
rabbit
while
chelators
concentration
=
to lead to head
bull.
Surprisingly,
0.27,
=
concentration
Na
rabbit
semen
are considerably
lower
than
in
man, bull, ram and boar but are about
twice as
high as in stallion
semen.
The approximate
1:1
Na and
K found
in
that of stallion
semen
was
(r
fact
tion.
(P<0.05)
1973)
in the
a chelator,
agglutination
found
in the literature.
Mineral
concentrations
did not
differ
significantly
between
first
and
higher
289
CONSTITUENTS
major
source
Significant
sexual
sequence
of
de-
activity
(Table
2).
with
those
reported
(Mann,
1964)
with
the
bulbourethral
be essentially
gland,
devoid
by
which
of fructose.
HOLTZ
290
000000000000
C N N
N
N
‘0
X
N
N
‘0
Q
C
en
O
*
-
N
en
‘0
00N
-
N
N
-
Jones and Glover
(1973).
Citric
acid
was
found
throughout
the reproductive
000000000000
N
-
Os -
C
N
-
en
Vs
centrations
000000000000
-0,050550
enen,..en
ductus
C
Vs Ce’
en
en 0
ONC
000000000000
CO5OVsNOCOONN
enenen
*Nen.V5N
X
C
C
0’ X
Nn,N
C
50
0.
8
N
‘fl
X
N
X
‘5
.0.0
000000000000
N C 0 ‘00’
N
N
N 0
en-=
C
5’
000000000000
en X en vs
-
-
en eos
en
entenenNenN,..
N
N
en en
X
-
and
and
-
N
en -
N
N
en
-
N
Vs N
2
000000000000
C C VS N
0.
0
S
-
‘
C
.0
.0
0
en N 0
‘OVsN
ductus
filled
.0
N
en
00
vs
C
C
000000000000
en N vs N Oen
NN
N,..enN-enCX
Os 00
-
N
the
N00osenC
V500,..en
N
en
‘I
0
‘3
0
.5
.0
en
C.
0
O
‘3
C en
OC’n*C
NN
0’
C
00
00
0
C
N
N
0000
0
GPC
N en N en en C
C-.ONO*--N00=
I
I
I
0
C
‘00
en
0
00
C
-0-
C
O
*05500
C
en.
00
-
en
en
N
0
C
en en
en
and
-
.0.0
‘3
V
C
C
+1*1+1
05
CO
C
0
=C=ON,en=CCOen
8
.8
-E,
‘3
‘
‘3
C
50
0
#{149}
0.
5’
0
low
concentration
between
the
this
the
proprostate
assumed
to exist
by
could
not
be found
Davies
in our
opening
into
the
gland
and ductus
for
glands
contained
sperm.
the notion
that much of
of the vesicular
gland has
‘8
=Jv8e0
,E
2E
with
a mass of
remained
empty.
were
ejaculates
high
(Table
2).
gel,
while
in both
The
GPC synthesis
or
1963).
The level
first
highest
of GPC in the reproductive
caput and cauda epididymidis,
sites of
et al.,
the
tract
both
accumulation
in the cauda
epididymidis
was similar
to that
reported
by
Jones
and
Glover
(1965).
The
ampulla
also
contained
appreciable
amounts
of GPC, but it
cannot
be determined
from
present
studies
whether
GPC is synthesized
in the ampulla
or
V
enenONCN,.,CNOO.en
.0
C
The
the
proprostate
makes
source.
Furthermore,
05 CV’
-H+1+I+I*t.H+I+I*
Uvo
the
and
Lutwak-Mann,
1951;
that
the proprostate
concentrations
second
active
(Scott
O00O___.0.O._
0.
ampullae
gland
concentrations
were in the
0’O
Oen
0
.0
‘3
from
Mann
them
“vesicular
gland”)
released
its
into
the
vesicular
gland
(termed
accessory
support
content
luminal
of the
vesicular
V
C
‘0
N
flow
origin
in the ampullae
is supported
by an
observation
by Hodson
(1965)
who found
that
interference
with
the autonomic
nerve
supply
to the accessory
glands
resulted
in engorgement
.0
C
the
The
its
8
5’
0
other
the
en
Ex
C
of
gland.
gland.
Rather,
a common
from
the vesicular
Additional
‘S
0
ampulla
retrograde
connection
Os
X0V55fl0C0O00
X.’CC.
“SN
N
C.
the
forced
into
the urethra.
Also,
the vesicular
gland
gel contained
spermatozoa
(Table
3),
presumably
coming
from
the ampulla.
None of
‘3
(I
small
amounts
tract.
High
con-
was established.
Fluid
injected
into the
deferens
in a dead or anesthetized
rabbit
the vesicular
gland
before
it could
be
deferens
0
V
concenNegligible
vesicular
vesicular
vesicular
gland
Mann
(1947)
studies.
urethra
NenNen
000000000000
C
from
seminalis”).
anatomical
-
000000000000
CXen00sen0=CC
0
S
in
in the
the
of citric
acid
in
gland
an unlikely
Vs
.0
‘5
I
by
“glandula
.0
50N
result
into
(termed
secretion
j
8
found
and
in
coworkers
(Mann
and
Mann,
1964)
concluded
0
C
may
ampulla
000000000000
C
were
deferens
latter
2
in fructose
(P<0.01).
amounts
of fructose
originate
in the excurrent
ducts,
a finding
consistent
with
the reports
of
U
0.
0’enCenO,NenO--C
N O’fl
enCen
FOOTE
Differences
among
organs
tration
were
significant
N
flten,..NNen
Nflsen
000000000000
OXNONNC*0
NCNenO,C
N
en N
AND
L
22
i2
.0
carried
midis.
citric
with
spermatozoa
Positive
correlations
acid
content
in the
P<0.05),
vesicular
vesicular
gland
gel
mass
(r
interpreted
as an
indication
(r
=
from
the
epididybetween
GPC and
ampulla
(r = 0.64,
=
0.59,
0.74,
that
P<0.05)
P<0.01)
GPC
and
are
secretion
ORIGIN
takes
of
tration
midis
acid
place
citric
concomitantly
acid.
The
with
correlation
the
RABBIT
in the
SEMEN
formation
between
of sperm
and GPC
was 0.86 (P<0.01).
cauda
the
substances
the
sources
Furthermore,
epididymis
epididy-
and
of the
not
permit
However,
it
Foote,
1970).
components
proprostate,
to
citric
in
prostate
and
changes
which
result
of sexual
of
with
similar
an
the
total
series
defined
analyses
of
tract
of
long-standing
void.
The
the
relative
various
organs
to
and
possible
other
of
conditions,
various
the
under
by the
place
as a
chemical
ejaculates
portions
correlation
of
data
contributions
the composite
semen
significant
physiological
the
fill
rabbit,
a
assist
of
the
sample
rela-
tionships.
ACKNOWLEDGMENTS
The
authors
are grateful
to Linda
Pierro,
Elvira
Osirio
de Valdivia,
Janet
Wiebold,
Lynne
Deuschle,
Gary
Anderson,
Richard
Cole and Larry
Schaeffer for
valuable
technical
assistance.
The
senior
author
was
supported
by fellowships
from
the Kellogg
Foundation and the Population
Council.
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Mann,
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