BIOLOGY
OF
8,
REPRODUCTION
62-73
Comparative
Fine
GERTRUDE
Structure
W.
Marine
Institute
Coral
(1973)
Gables,
HINSCH2
Biological
of Molecular
Florida,
and
primitive
type
sperm
of
1956).
nucleus
The
and
of the
occurs
head
an
fertilization
when
the
with
the
undergoes
ture
egg
(for
from
or egg
well-defined
review
species
Fine
sperm
contains
The
coat
and the
changes
see
Colwin
1970).
These
studies
and the
Hinsch
and
Clark,
1970;
Summers,
exception
1970;
Colwin,
Franz#{233}n’s study
zoan
Nausitho#{235}.
The
and
1971;
1971;
‘Contribution
Miami.
grant
No.
and
This work
T1-HD-26-08
Physiology
Training
Laboratory,
Woods
‘NIH
Career
212
Cellular
from
and
was
supported
to the
Program,
Hole,
in
tion
head
NIH
Marine
Awardee.
62
Copyright
All rights
©
of
1973
by
reproduction
The
Society
in
any
for
form
the
Study
reserved.
of Reproduction
well
developed,
other
Clark
probably
for
the
Summers,
i.e.,
1969;
of
most
act
as
distal
1970;
Lunger,
the
phyla
and
Anderson,
function
though
structure
of the
vesicles
associated
and
of the
pericentriolar
A comparative
using
sperm
of all three
Biological
they
Subsequent
pericentriolar
these
invesan
an-
centriole
Afzelius
1971).
study
was initiated
in order
more
fully
the spermatozoa
coelenterates;
by
so
and
Franz#{233}n, 1971;
of
and Gamete
1964;
in gametes,
the sperma-
from
1967;
Longo
device
1964;
of
Mass.
Development
feel
Institute
part
not
1971).
The
is unknown,
University
Fertilization
gregarium.
similar
animals
Austin,
1969;
tigators
O’Rand,
the
Evolution,
of
and
Hinsch,
This
derstand
Molecular
(Szollosi,
and Clark,
1971;
Hinsch
and
Lunger,
1971).
Szollosi
(1964)
though
in sperm
(Fallon
1970;
Summers,
1971;
Dewel
Lunger,
during
Hinsch
and
Clark,
1970;
Afzelius
and
Franz#{233}n,
structures,
choring
(Szollosi,
Clark,
1970;
1970;
Phialidium
have
shown
1970;
1970;
and
vesicles
to date
rids
of
reports
Jespersen,
structures
Clark,
Franzen,
concerning
first
described
such
processes
which
he termed
satellites,
in
of the
primitive
scyphoThe
above
studies
indi-
and
and
these
studies
Dewel
1971;
cated
that
coelenterate
sperm
may lack an
acrosorne.
It is interesting,
however,
that
while
no discrete
acrosome
has been
described
numerous
investigators
(Hanisch,
IIinsch
Afzelius
have
noted
the nucleus
is raised
of
sperm
1971;
Clark,
Lucchi,
1970;
Lunger,
1971).
to the above
is Afzelius
a question
Hanisch,
Summers,
of the
et al.,
1970;
and
thus,
terate
vary
of coelenterate
majority
of these
Stagni
1971)
between
Of further
interest
are elaborate
pericentriolar
processes
noted
in association
with
the distal
centrioles
of many
of the coelen-
to species.
structural
are sparse,
1972
the
function
fertilization.
acrosome
in struc-
and
9,
sors;
species
contact
studies
have
been
done
on sperm
hydrozoans
(Szollosi,
1964;
Burnett
1966;
Weissman
et al., 1969; Hanisch,
only
the
changes
JR.
and plasma
membrane
in many
coelenterate sperm.
Such
vesicles
may
be of evolutionary
significance
as acrosomal
precur-
initiation
process
in many
sperm
first makes
Franklin,
1967;
usually
acrosome.
CLARK,
personal
communication;
small
vesicles
situated
characteristically
has a short
round
or conical head,
a midpiece
usually
containing
4-5
mitochondria
and
a long
flagellum
(Franzen,
H.
Woods
Hole,
Massachusetts,
Evolution,
University
of Miami,
of Biology,
University
of Houston,
Texas
February
metazoans
Spermatozoa’
WALLIS
AND
Laboratory,
and Cellular
the Department
Houston,
Received
The
of Cnidaria
and
to unof the
func-
with
the
processes.
approach
has
been
taken
from
animals
representative
classes
of the
Cnidaria.
CNIDARIA
MATERIALS
Animals
the
used
Supply
tory,
for
Woods
and
as
sperm
Testicular
from
anthozoan
aquaria
was
Karnovsky’s
obtain
were
from
the
sperm
ob-
dianihus.
incomplete
were
mature
animals.
suspensions
were
(1965)
mixture
in
in
embedded
in
with
phosphate
fixed
or
in
stained
and
1965),
HS-8,
citrate
viewed
or
acetate
(Wat-
and
Cogges-
300
AEI-EM6B,
microscope.
OBSERVATIONS
Though
resentative
been
placed
sperm
from
several
species
of each
class
of Cnidaria
studied,
on one
obvious
major
sperm
differences
described.
Class
Hydrozoa.
both
Tubularia
have
placed
been
on
exist,
sp.
differences
scribed.
The
of Tubularia
those
described
will
Though
sperm
and
Hydractinia
studied,
major
the sperm
of
obvious
they
exist,
conical
they
and
will be
Where
will
be
elongate
by Tuzet
studies
(1929,
1950)
of Pennaria
mers,
and
panularia
flagellum
anterior
1970)
and
end
vesicles
of the
studies
show
that
with
the nucleus
tron-dense
material
resemble,
Cain
Hydrozoan
sperm
4-5
mitochondria,
in
size
(Sum-
(Lunger,
associated
nucleus
(Fig.
a
appearance,
envelope
the
end
1 and
3).
com-
to
(Figs.
Vacuolated
condensed
of
the
1 and
nucleus
is
The
is pyra-
closely
of the
midpiece
associated
midpiece
is composed
(Figs.
proximal
variation
to l)e
sampling
The
error.
mitochondria
contain
several
which
are
tubules
with
flattened
(Figs.
3-fl).
of adjacent
apposed
but
in
real
4 and
5)
and
distal
This
appears
each
mitochondrial
rather
than
The
paired
mitochondria
do
not
cristae
widened
outer
are
appear
to
a
niemclosely
fuse
with
other.
The
neath
proximal
the nucleus
centriole
is situated
and is orientated
bealong
the sperm
axis
(Figs.
3 and
4).
The
triole
is composed
of nine
triplet
sets
tubules.
While
there
are no pericentriolar
structures
associated
is a
triplets.
with
concentric
the
are
nine
distal
same
(Figs.
1).
Our
centriole
and
a matrix
3, 6-8).
its
matrix
of pericentriolar
these
processes
from
centriole,
circles.
inner
and
electron-dense
in
material
centriolar
of
Extend-
outer
dense
spokes
(Fig.
centriolc,
which
is orientated
plane
as the proximal
centriole,
is embedded
arise
this
ecu-
core-like
structure
within
the
This
core is composed
of two
between
circles
4).
The
in the
tem
laths
an
areas
are
nucleus.
centrioles.
number
the
vesicles
Golgi
comparable
shape
and
mitochondria
with
the
vesicles
associated
are filled
with
an elec(Figs.
1 and 2). These
and
in
the
ing
and
typically
have
two
centrioles,
posterior
midal
with
electron-dense
and
Hydractinia
resemble
in the light
optical
studies
the
at the
anterior
and
posterior
the membranes
of the envelope
apposed
to each
other
as well
The
there
nine
de-
sperm
of Tubularia
ultrastructural
1971).
nucleus,
be
from
sp.
emphasis
Tubularia.
nuclear
the
nucleus.
present
in
ends
branes
rephave
emphasis
will
be
each
class.
Where
in
structure
of 4 or 5 mitochondria
which
surround
the
MT-2
uranyl
an
a loose
(Figs.
were
(Venable
with
Philips
and
Sections
a Porter-Blum
with
lead
and
on
post-fixed
dehydrated
Araldite.
knife
ultramicrotome,
Hitachi
buffer,
Maraglas
a diamond
1958)
buffer,
phosphate
by
as to
often
collected
with
of the typical
spermatozoa
was
The dense
nucleus
is surrounded
2),
except
ends where
are closely
paraformaldehyde-glutaraldehyde
in 1% 0s04
hall,
sperm
a single
acrosome
observed.
the
to
Metridiuin
dissected
sperm
from
association
in
plex in developing
spermatids
(Clark
and
Hinsch,
unpublished
data;
Lunger,
1971).
No evidence
of fusion
between
vesicles
to
form
the
animals.
Spawned
and
from
unable
mature
containing
Tissues
were
these
the
septa.
sperm
dissected
We
and
tissue
secondary
sp.
tissue
from
by
Labora-
Tubularia
sp. were
et al. (1957).
Costello
were
Aurelia
Testicular
tained
by
tissues
spawned
collected
Biological
sp. and
described
scyphozoan
were
nature
Hydractinia
Germinal
son,
study
found
Massachusetts.
tissues
obtained
METHODS
Marine
Hole,
hydrozoans
cut
this
Department,
Germinal
from
AND
63
SPERM
plaque-like
matrix
between
of
electron-dense
Associated
is an
with
elaborate
processes.
(primary
depressions
triplets
this
sys-
In Tubuprocesses)
of
(Fig.
the
6).
Fic.
1.
and
compact.
cleus
are
acteristic
Fic.
of
electron
Small
two
9 +
2.
2 tubular
micrograph
a Tubularia
sperm.
3.
and
micrograph
vesicles
mitochondria
A
Fic.
(PC)
An
An
the
electron
distal
of
a mature
surround
of
pattern.
the
x
illustrating
the
sperm
anterior
midpiece.
Tubularia
from
region
Also
seen
of
are
the
cross
sp. The
nucleus
is extremely
nucleus
(arrows).
Posterior
sections
of flagella
exhibiting
dense
to
the
nu-
the
char-
of the
head
36,000.
the
vesicles
(arrows)
associated
with
the
anterior
region
x60,000.
micrograph
centriole
showthg
(DC)
are
the
found
midpiece
between
64
of
the
a
Tubularia
mitochondria.
sperm.
The
x 60,000.
proximal
centriole
CNIDARIA
The
primary
processes
extend
the
centriole
and
secondary
processes
nate
at the
In Hydractinia
give
cesses
dna
extend
out
and
plasma
form
a thickened
from
tip
(Fig.
6).
processes
matrix
between
The
primary
pro-
between
membrane
these
processes.
from
smaller
to termi-
appear
plasma
membrane
similar
primary
from
the
centriolar
(Figs.
7 and 8).
secondary
with
each
Both
the
to
rise
which
arise
triplets
tending
secondary
away
the
(Figs.
mitochonwhere
they
8 and
thickened
The
9).
Ex-
tips are
first
and
processes
of adjacent
other
at their
extreme
primary
and secondary
three
third
arms
fuse
(Fig.
8).
processes
exhibit
a distinct
periodicity.
The nine
outer
doublets
of the flagellum
arise from
the A and B tubules
of the centriolar
exhibit
triplets.
The tubules
of the
the typical
9 + 2 pattern.
Class
sp.
Scyphozoa.
have
long
flagellum.
rior end
projection
studies
In
of
this
veal
microtubules
extending
Scattered
and
mature
area
sperm
in
late
with
among
the
profiles.
Behind
anterior
to the
midpiece
mitochondria,
associated
13 and
of adjacent
electron-dense
slight
the
rehelical
projection.
microtubules
are
structures
and
tation
in its anterior
end
of fibrous
electron-dense
The
shaped
ante-
sev-
membrane-bound
vesi11). Sections
of the vesiringor donut-shaped
nucleus
membrane-bound
a very
the
spermatids
a
these
is not
Aurelia
forms
a long
Ultrastructural
throughout
electron-dense,
(Figs.
10 and
show
elongate,
and
(Figs.
of
heads
above
just
an inden-
is a condensation
material
which
(Figs.
consists
the
11).
10 and
of four
wedgedistal
centriole,
pericentriolar
14).
The
outer
mitochondria
plates
and
structures
membranes
The
distal
centriole
to the mitochondria.
cesses
extending
are also found
they
differ
from
in the
somewhat
is found
just
Pericentriolar
the centriolar
Scyphozoa;
from
those
postepro-
matrix
however,
of
from
The
the
triplets
extend
primary
centriolar
processes
matrix
arise
between
(Fig.
14). These
primary
radially
away
from
the
and
terminate
from
each
in thickened
thickened
ary processes.
minate
on
an
circumscribes
within
the
the
plasma
15-17).
The
tal centriole
are
Extending
three
secondary
membrane
second-
processes
band
terwhich
processes
(Fig.
just
14).
the
midpiece
a long
surrounds
the
anterior
long
flagellum
(Figs.
flagellum
(Figs.
electron-dense
anterior
tubules
cytore12,
extends
from the disand
15).
A ring
of
12
material
doublets
two inner
the
processes
centriole
tips.
tip
The secondary
electron-dense
Posterior
to
plasmic
sleeve
gion
of
the
is
seen
to the region
arise
(Figs.
within
the
where
15 and
the
16).
flagellum
exhibits
the classic
9 + 2 pattern
with
neither
A nor B tubules
of the doublet
appearing
filled
with
electron-dense
material
(Fig.
central
doublets
Class
possess
(Fig.
rounded
lope.
17).
Vesicles
the
do
as
Scyphozoa.
shaped,
vesicles
was
are
found
and
of the
nucleus
the
of
anterior
in
Hydrozoa
the
the
the
which
to the
and
nuclear
near
cytoplasm
head
and
midpiece
21).
consists
and the
to each other
(Fig.
tron-dense,
lipid-like
a limiting
membrane
head
surenve-
en-
the
base
18-21).
in
of
centriole
the
membrane
the
(Fig.
is
numerous
donutelectron-dense
(Figs.
present
region
The midpiece
mitochondrion
proximal
between
appear
between
plasma
granules
centrioles
not
seen
However,
membrane-bound,
velope
Also
posterior
extend
an elongate,
wedge-shaped
18).
The
condensed
nucleus
by a loose-fitting
nuclear
nucleus
like
Fibers
tubules
and doublets
and
between
and
flagellar
sheath
(Fig.
17).
Anthozoa.
The
Metridium
sperm
cytoplasm
appear
fused
into
22 nm in width
(Figs.
11-13).
rior
Hydrozoa.
In this region
small
fibers
extend
from
the
doublets
and terminate
in a “Y”-like
junction on the flagellar
sheath.
Posteriorly
the
flagellum
sperm
of the spermatozoan
(Fig.
10).
bend
eral
des
des
The
elongate-shaped
65
SPERM
are
glycogen-
of a large
complex
proximal
and distal
are
at
21).
with
900
Several
spheres
are found
and
of the
the free
the
respect
large,
which
around
anterior
eleclack
the
portion
66
HINSCH
Fic.
sperm
4.
rounded
Fic.
centrioles.
A
by
5.
cross
section
through
Tubularia.
from
four
A cross
Note
Note
mitochondria.
section
there
proximal
intricate
centriole
substructure
CLARK
in
the
within
anterior
the
region
of
centriole.
The
the
midpiece
centriole
of
is
sur-
x59,300.
through
are
the
the
AND
five
the
midpiece
mitochondria.
of
a
x45,000.
Tubularia
sperm
between
the
proximal
and
distal
a
NIDAEIA
of the
distal
centriole
distal
The
centriole
electron-dense
is
matrix,
embedded
and
this matrix
triole
are
at the posterior
pericentriolar
(Fig.
Extending
23).
and
gion
the flagellar
of the tail
scribed
end
primary
the
24).
and
metrical
complex
mitochondrion
the anterior
portion
of the
21 and
24).
The
flagellum
9 + 2 tubular
sical
rede-
the
Stagni
been
the
Aurelia
(Tuzet,
nucleus.
DISCUSSION
type
a head,
show
may
Aloia
the
be
from
odic
acid-silver
the
Golgi
reactions.
Clark
et a!.
proacrosomal
a flagellum.
spermatids
and
consists
which
a
midpiece
containing
mitochondria
and
The head
in
most
spermatozoa
of the
nucleus
and
a membranevesicle,
the
acrosome,
which
caps
bound
are
PAS
the
nucleus.
As reported
here,
the
spermatozoa
of Cnidaria
do not possess
such
a vesicle.
However,
the marine
hydrozoans
and
undergo
tal
tract.
Tubularia
another
similar
(Afzelius,
C lava
(O’Rand,
Campanularia
1971),
personal
(O’Rand,
1971)
several
and Pennaria
small
vesicles
gion.
Like
tozoa,
the
these
origin
and
6.
sperm.
from
a cup-shaped
process
Ftc.
The
three
7.
A high
(large
mary
FIG.
chondrion
three
A
join
primary
matrix
longitudinal
a sperm
are
view
of
and
centriole
at
are
illustrating
pericentriolar
at
their
extremes.
of
a primary
Hydractinia.
in
The
region
female
geni-
vesicles
in
nature.
In
in
events
vesicles
while
tissue.
preceding
and
near
of
from
a
thickened
first
and
tip.
third
from
each
primary
x 50,000.
membrane.
of Hydractinia
Portions
x135,950.
with
Extending
a Tubu-
Extending
matrix.
(arrows).
triplets
secondary
of
matrix.
a sperm
associated
between
fusion.
midpiece
plasma
an electron-dense
between
triplets
matrix
the
Extending
the
centriole
membrane
electron-dense
arrows).
processes
centriolar
terminate
arrow).
a distal
matrix
are
enzymes
1971)
the
of changes
gamete
in an
(large
in
origin,
reproductive
of
posterior
terminate
of
are embedded
from
this
(small
the
embedded
which
micrograph
the
similar
female
subsequent
processes
centriole
emanating
from
in the
the
suum
fertilization
may
clarify
the
role
by the vesicles
during
penetration
during
played
arrow)
processes
other
the
primary
body
donut-shaped
be
have
in
Ascaris
dehiscence
in contact
with
An investigation
centriole
(small
processes
each
distal
distal
Extending
sperm.
The
from
the
seen
micrograph
secondary
processes
9.
be
be
electron
of
Golgi
1972)
of
Cam panularia
(O’Rand,
seem
to undergo
a series
acrosomal
in fresh
the
processes
magnification
can
the
absent
of the
in this
of
thus
through
tubules
Electron
arrow).
are
of
secondary
a Hydractinia
from
tip
8.
may
tubules
origin
unpublished
/3-gly-
anthozoan,
Bu nodoso
ma cavernata,
rings,
donut-shaped
vesicles,
are
and
appear
to be of Golgi
body
(Dewel
and
Clark,
1972).
In
present
have
re-
sperma-
to be
are
section
triplet
fossa
are
Note
the
triplet
primary
processes
Ftc.
and
forerunners
vesicles
A cross
lana
Lunger,
other
Hinsch,
1971)
evolutionary
vesicle.
Such
of
appear
vesicles
Lunger,
Ftc.
1971;
(Summers,
1970)
in a comparable
acrosome
(Clark
data;
Hydractinia,
peri-
and
vesicles
hydrolytic
may
vesiwhich
positive
(1967;
Golgi
in
acrosomal
contain
The
Metrklium
communication),
of
those
reported
rotifers
have
sperm
also
positive,
only
to the
like
methenamine
Likewise,
demonstrated
the
presence
anterior
has
of
cerophosphate
in
of
primitive
and
studies
studies
vesicles,
(1970)
the
acrosome
the
spermatozoa
arise
find
and
conby Tuzet
light
an
vesicles
Hydrozoa,
in
Lucchi,
to
our
These
structures.
des
of
sperm
sperm
Weiss-
and
Although
1950),
donut-shaped
1966;
unable
presence
of the
al.,
centrosome
filament
seen
Tubularia.
in
indicate
the
clas-
structure.
Cnidaria
sperm,
like
all primitive
sperm
(Franz#{233}n, 1956;
1970),
have
et
1969;
have
scyphozoan
surrounds
(Figs.
19,
tail
We
(1950)
scyphozoan
by the asym-
exhibits
(Burnett
al.,
acrosome-contained
nected
intranuclear
doublets
sheath
in the anterior
are Y-shaped
fibrils
as
hydrozoans
A sleeve
formed
et
1970).
from
of the denprocesses
between
Hydra
man
an
in
extending
in the
(Fig.
water
18, 19, 21-23).
(Figs.
67
SPERM
the
are
distal
from
processes
each
of
sp.
some
centriole
primary
processes
of
thickened
adjacent
pri-
X 63,840.
process
x95,760.
passing
between
the
plasma
membrane
and
a mito-
08
HINSCH
AND
CLAE1
#{149}
c-_s
-
0
_____
Ftc.
10.
pletely
An
condensed.
Just
tubules
are
noted
the
midpiece
gate
of
micrograph
Just
arrow).
anterior
anterior
to
in
I
___
electron
the
is
the
of
to
fibrous
anterior
just
posterior
a late
the
spermatid
nucleus
region
region
of
to
the
is
Aunelia
from
a moderately
are
numerous
the
sperm
nucleus.
sp.
dense
vesicles
head
x24,000.
(small
The
region
filled
arrow).
with
nucleus
of
fibrous
a dense
The
is
almost
material
material.
mitochondrial
com(large
Microaggre-
CNIDARIA
In
a recent
(1970)
sperm
study
Afzelius
have
described
of the primitive
They
emphasize
is short
and
the
contains
lumps.”
Tubularia
In
1971)
fact
and
that the midpiece
four
“mitochondrial
an electron-dense
material.
may
act as a cementing
(Afzeliiis,
(Summers,
1970)
midpiece
pears
to be
The
chondria
though
short
a common
spermatozoa.
in
there
sperm
this
such
are
of the
feature
of four
midpieces
exceptions,
sea
urchin
is common,
e.g.,
in
the
and
of some
teleost
mitochonwe have
in the
complexity
of the
mitochondrial
the midpiece
in various
cnidaria
those
hydrozoan
sperm
examined
form
sperm.
by
and
a!.,
other
workers
(Burnett
et
1969;
Hinsch
and
1970;
al.,
Clark,
1970;
Summers,
midpiece
tinct
Stagni
1970;
is composed
of
exhibit
Fio.
11.
Aurelia.
Ftc.
an
A
12.
micrograph
An
electron
x
13.
A
mitochondria
chondria
14.
of
cross
section
and
the
seen
in
dis-
15.
Portions
These
the
the
region
the
both
anthozoans
which
the
mitochondrial
surrounds
the
proximal
anterior
region
of
in
the
sperm
B.
1972).
and
However,
are
sperm
Clark,
from
of
bodies
of
and
spermatozoa
In the
inclu-
anterior
lipid-like
midpiece
(Dewel
a
This
complex
of mitodhon-
spermiogenesis.
large
lipid-like
the
of
has
likewise
mass.
product
in
a
sperm
cavernata
1972)
Similar
centrioles.
in
showing
extending
through
which
the
hydro-
we have
exaggregate
or
and
distal
the
sperm
of
head
of a sperm
Au-
from
of
the
a
midpiece
anterior
portion
of
a
of
the
sperm
from
sperm
tail
midpiece
of
a sperm
between
from
Aurelia.
Note
outer
membranes
apposed
the
four
wedge-shaped
of
adjacent
mito-
the
pericentriolar
(arrow).
in
in
region
of the
midpiece
of
a dense
matrix.
Extending
thickened
processes
through
tips.
Extending
terminate
the
processes
tubules
Cross
the
posterior
embedded
on
from
an
a
the
sperm
from
the
thickened
electron-dense
band
from
Aurelia.
matrix
are
nine
tips
sec-
which
are
The
three
circumscribes
x53,500.
section
and
section
through
material
lies
inside
plasma
the
the
posterior
(large
membrane
anterior
doublets
region
arrow)
of
the
region
of
of
can
most
of
flagellum.
be
the
midpiece
seen,
as
anterior
sperm
tail
Y-fibrils
well
region
and
of
Aurelia.
of
Y-fibrils
(small
of
the
x41,000.
cytoplasmic
extend
sperm
a
as
from
tail.
collar.
doublets
arrow)
A
to
ring
of
flagellum
x43,000.
Cross
are
section
surrounding
material
secondary
tangential
electron-dense
17.
longitudinal
collar
the
terminate
The
between
16.
a
cytoplasmic
electron-dense
processes.
A
of
Fic.
the
Cross
section
through
the
distal
centriole
are
pericentriolar
flagellum
found
In the
zoans
amined,
mass
The
the
X43,200.
processes.
Ftc.
Clark,
fusion
during
of Metridium
cavernata
from
contains
Bu nodosoma
and
are
Aurelia.
sperm
usually
18,300.
processes
ondary
sheath
micrograph
and
(arrow).
primary
FIG.
anthozoan
midpiece.
Aurelia
than
mass.
the
vesicles
primitive
mitochondrial
sions
five
more
single
1966;
by Afzelius
possible
explanaNausitho#{235} is con-
the
1970;
the
feature.
far
mathe
in
Nau.sithoe
(1971).
A
above
is that
the
main-
mitochondrial
an inter-mitochondrial
been
demonstrated
jellyfish
is
material
substande
the
The
midpiece
of
anthozoan
Metridium
drial
sperm
midpiece
illustrating
Mitochondria
evident.
Ftc.
triplets
interesting
for
memthere
x52,600.
Aunelia.
Ftc.
of the
scyphozoan
tion
outer
This
of
single
mitochondrial
mitochondrion
is the
Lucchi,
or
of the
(Dewel
of
In
us
1971)
four
Such
not
Franz#{233}n
the
mitochondria.
sp.
the
and
Lunger,
The
four
mitochondria
from
the sperm
of the
are
et
Hanisch,
integrity
and
sidered
mito-
fish the
midpiece
contains
one
drial
mass.
In the present
study,
demonstrated
a definite
progression
Weissman
sperm
ap-
of primitive
appearance
the
aggregate.
terial
has
the
and
apposed.
Between
the
of apposed
mitochondria
taining
ther
that
wedge-shaped
closely
branes
to the larger
They
fur-
out
are
the midpiece
of the
jellyfish
Nausitho#{235}.
midpiece
is asymmetrical
due
size of one of the mitochondria.
point
mitochondria
Franz#{233}n
larynx
as in Pennaria
69
SPERM
section
present.
through
x53,500.
cytoplasmic
collar
and
sperm
tail
at
level
where
central
tubules
of
HINSCH
70
CLARK
AND
I
.
I
24
Ftc.
18.
wedge-shaped
An
electron
micrograph
and
exhibits
a distinct
proximal
centriole
chondrion
is
present
(arrow),
in
which
the
midpiece.
of a
fossa
is
mature
at its
surrounded
x24,000.
sperm
posterior
by
lipid-like
from
end.
Met r-idium
Just
inclusionS
dianthus.
posterior
(1).
to
A
the
large
The
nucleus
nucleus
is
complex
is
the
mito-
relict
the
centriole
distal
is
in
posterior
We
have
to the
mitochondrial
been
unable
to find
centriole,
describe
although
one
in
Afzelius
the
sperm
with
all three
are
pericentriolar
tures
were
first
in
(1964)
the
and
gregarium
spermatids
have
more
the sperm
of
(Fallon
and
Clark
son,
and Hinsch,
1969; Longo
1969;
Hanisch,
1970;
Clark,
1971;
Summers,
Franz#{233}n, 1971;
Dewel
1967;
and
Clark,
1971;
Jesperson,
1971;
Afzelius,
1971).
Szollosi
lieved
these
structures
to
(pericentriolar
Bessis
and
bodies),
Breton-Gorius
hard
de
and
(Hanisch,
apparatus
Afzelius,
(Lunger,
1971)
Similar
fibers
Fm.
19.
sperm
An
20.
Ftc.
21.
High
centriole
collar
around
Fic.
22.
x20,000.
Ftc.
The
the
23.
primary
Ftc.
The
tubules
Cross
cytoplasmic
the
be
seen
electron
in
section
collar
flagellum
of
lipid-like
through
(arrow).
time
in
been
responsible
the
satelstruc-
spermatids
has
for
which
of the
some
exists
satellite
Summers
in
of
the
com-
(1972)
has
the
use
of
the
of
in
we
which
de-
of
a term
the
function
We
have
nucleus
region
out that
process
and
the
of these
further
midpiece
nucleus
of
(arrow).
Gly-
x28,000.
vesicles
(small
Metridium
of
in
same
the
(Dewel
subse-
of both
further
be pointed
term
pericentriolar
posterior
Metridium
centriole
midpiece
embedded
with
in Metridium
sperm.
x61,200.
showing
proximal
centriole
(PC),
found
sperm
of
niitochondrial
complex
which
forms
x42,000.
of
distal
ecuor
associated
time
during
presence
spermatozoa
region
the
Metridium
and
portions
(1)
midpiece
of
are
differentiation
confusion
In his model
avoided
posterior
of
tail.
associated
associated
a function,
since
structures
is unknown.
cytoplasm.
midpiece
the
spermatozoans.
It should
using
the
illustrating
sperm
through
portion
a dense
matrix.
sperm
illustrating
arrow)
and
sperm
in
the
proximal
posterior
Extending
region
from
inclusions
lipid-like
centriole
(large
of
matrix
are
distal
nine
arrow).
centriole.
pericentri-
x40,000.
through
contains
to
free
seen
notes
the
inclusions
of
are
the
in
have
around
micrograph
portion
section
centriole
processes
24.
of
section
anterior
Cross
of
in
called
and
found
the
Hanisch
not
shown
the
secondary
processes
associated
with
the
pericentriolar
processes
similar
to those
seen
here
in the cnidarian
Franz#{233}n,
spikes
showing
are
section
(DC),
Cross
triplets
olar
power
anterior
around
plex
processes
Vesicles
Longitudinal
distal
these
at
the present
literature.
anchoring
micrograph
can
particles
Fic.
1)
electron
Metridium.
from
cogen-like
for
with
by
are
which
The
undoubtedly
by
Bern-
processes
have
been
in flagellates
(Gibbons
be
disappear.
tures
Subsequent
and
centriolar
of
have
pro-
of the
distal
from
satellites
bodies,
spermatid
lites
satellites
1970),
(Afzelius
1971)
and
agrees
structures
may
quent
have
used
the term
satellite
Anderson,
1969; Hinsch
and
1971;
Summers,
1970),
cen-
triolenfortsatz
transition
(1960).
well
cells
pericentriolar
distal
centriole
at the same
and
Clark,
1972).
However,
1971;
be-
as described
(1958)
and
Harven
investigators
(Longo
and
Clark,
1970,
be
as
We
1965).
as used
These
cells
sensory
the
posterior
end
and
are
distinct
processes
and
1971;
Hinsch
Lunger,
somatic
1970),
with
a more
anterior
region
of the centriole.
In young
spermatids
both
satellites
(pericentriolar
bodies)
and
pericentriolar
inverte-
Afzeius
Clark,
in
in translation
pericentriolar
and AnderHinsch
and
1970;
and
and
Stubblefield,
centriolenfortsatz
(1970).
of Phiarecently
other
Austin,
which
with
triole
Similar
strucin gametes
by
noted
in
species
sperm.
and
as spokes
in the utricular
fish
(Flock
and
Duvall,
termed
these
structures
term
the
distal
centriole
in
classes
of the Cnidaria
processes.
described
1960)
(Brinkley
and
Franz#{233}n
of Nausitho#{235}
been
brate
fiber
1971;
Grimstone,
position
aggregate.
a proximal
cesses,
(1971).
Associated
sperm
from
Szollosi
ligium
a
71
SPERM
CNIDARIA
the
cytoplasmic
portions
flagellum
collar
of
the
sheath
and
anterior
mitochondrial
(arrow).
portion
complex.
x
28,200.
of
Y-fibrils
tail
of
extend
Metridium
from
sperm.
the
outer
a
72
AND
HINSCH
avoided
this term
tion
the term
transition
was initially
used
(Gibbons
and
been
used
(Brinkley
1970)
we
the structures
adult
life
of
are
in
oocyte
(Longo
and
The
be
feel
the
appearance
in
anneid
in
anterior-most
the
flagellar
As
the
ooplasm
of
and
Anderson,
the
processes
should
“Y”
as initially
fibers
in
by
the
sperm
Colwin
and
initially
fibers
are
portion
of
Colwin
found
of
the
the
function
pothesis
is the
the pericentriolar
close
It
sized
that the above
as
is demonstrated
other
possibilities,
pertaining
to
mers
(1970)
proximity
should
be
to the
empha-
structures
should
not
such
sperm
suggested
symmetry
resulting
movement
may
pre-
as functional
motility.
Sumthey
contractility
in the
structure
in the
hy-
is not always
the case,
in
the
scyphozoan
clude
roles
Such
this
many
instances
appear
to ter-
while
these
role, one
has
support,
the distal
for
that
in
structures
at or within
membrane.
contractile.
for
may
could
of
be
cause
pericendirectional
the
sperm.
des
R.,
and
and
centriole
vances
M.,
F.
A.
E.
A.
(1971),
study
of
cells
arising
L.,
Fine
Coelenterata).
I. Ultrastruct.
B.
AFzEuus,
spermatozoon
struct.
Res.
A.,
AND
FRANzEN,
of the jellyfish
37, 186-199.
structure
larynx
Res.
of
the
A.
Nausithoe.
(1971).
The
I. Ultra-
“Ad-
In
meiosis.
I.”
(Prescott,
E.,
cdi.),
L.
E.,
and
AND
differentiation
of
of
cells
120, 1-8.
H.,
Ja., AND
interstitial
Hydra
in
HtNscH,
C.
sperm-egg
lacteus.
RUFFINC,
ultrastructural
viridis.
W.
(1969).
interaction
Biol.
Bull.
in
137,
395
(abstr.).
CLARK,
MoilErrI,
R. L.,
W.
W.
W.
(1967).
Electron
microscopic
for
the
presence
of
H.,
JR.,
an
var.
W.
W.
of
the
and
spermatids
lumbricoides
suum.
L.,
AND
C0LwIN,
of
the
spermatozoon
evidence
in
Exp.
Cell
Res.
AND
THOMSON,
ultracytochemical
and
var.
THOMSON,
reaction
R. L.,
Histochemical
studies
AND
acrosomal
suum.
Ascaris
lumbricoides
47, 643-647.
CLARK,
W. H., JR., MonErrt,
sperm
Biol.
Ascaris
of
Reprod.
7.
145-
159.
COLWJN,
A.
structure
hexagonus
to
(Annelida),
the
acrosomal
Cytol.
COLWIN,
L.
brane
of
with
special
I.
Biophys.
region.
“Fertilization.”
cdi.),
Vol.
I.
(1961).
Fine
Hydroides
H.
10, 211-230.
L. H., AND COLWIN,
A. H.
fusion
in relation
to sperm-egg
reference
Biochem.
(1967).
Mem-
interaction.
(Metz,
C.
B.
Chapt.
7.
Academic
and
Monroy,
Press,
York.
M.
D.
P.,
H.,
AND
obtaining
M.
DAVIDSON,
C.
HENLEY,
and
handling
Lancaster
E.,
(1957).
ECGERS,
marine
Press,
A.,
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Inc.,
and
Lancaster,
Penn.
DEWEL,
37, 679-689.
(1970).
kinetochore
McConkey,
gland
aspects
New
(Hydrozoa,
the
Vol.
histological
from
Cerebratulus
E.
and
DAVIS,
A
et
elec-
363-386.
of
and
J.-P.
golgi
N.Y.
germinal
W.
Some
for
Tubularia
of
L.,
(1966).
de
microscope
13,
Biology,
Con-
THIERY,
STUBBLEFIELD,
Century-Crofts,
embryos.”
B.
au
mitosis
Cell
Goldstein,
BURN1-r,
AND
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Intl.
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corps
Etude
AND
(1960).
elements
B-2,
J.,
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B.
E.
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spermatozoon
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(1958).
aster
DE
centriole
Electron
M.,
BES5IS,
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REFERENCES
of
spermato-
Asplanchna
achromatique.
on
In
the
in
of
AND
l’appareil
gress
CLARK,
not
sug-
have
in
support
fact
Aureli.a.
Thus,
play
a supportive
function
device
is
the
dissertation,
du
Appleton
of
on
of
rotifer
Ph.D.
W.,
BERNHARD,
D.
tail.
from
the doublets
appear
to insert
previously,
female
Ultrastructure
the
differentiation
transport
Riverside.
J. Morphol.
Apparent
changes
triolar
changes
not
sheath.
centriole.
the
tronique.
that
they
may
as an anchoring
minate
plasma
Clark
1968;
pericentriolar
processes
Several
investigators
gested
acting
early
Cytological
and
Cosse.
de
mentioned
of
the
known.
into
welli
the
their
They
extend
radially
the
tail,
branch
and
the
zoan
present
throughout
cell as indicated
by
gametes
Hydroides
The
“Y”
(1970).
spermatozoan
L’ultrastructure
the
(1961).
to denote
a temand
Stubblefield,
the
since
with
described
it has
R.
ALcoA,
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