Morphological
changes
in
organelles
of
Pinus
II.
and
the
quantitative
population
during
sylvestris
Morphological
I
prometaphase
of
cell
microsporogenesis
L.
changes
until
the
from
tetrad
stage
M.Th.M.
Botanisch
Willemse
Laboratorium, Universiteit, Nijmegen
SUMMARY
A
morphological study
Pinus
sylvestris
breakdown
with
and
made
was
L. from
rebuilding of
the transfer
with
prometaphase
the nuclear
of ribosomal
to the
respect
changes during microsporogenesis
I until the tetrad stage. Described
material,
membrane,
the nuclear
the
mixing
of
invaginations,
and discussed
and
karyoplasm
the kinetochore
are
of
the
cytoplasm
and micro-
tubuli.
A decrease
The
of starch in the
“lipid-complex”
during
the cell
divisions.
during interphase II
produces
small and
culum. These
membrane is
between the
A relation
and
postulated
in
telophase
Ribosomes
vesicles and
vesicles fuse
interphase II.
II.
may
The mitochondria
vesicles
Many
observed
are
as
flat
long
in
all
cisternae,
and form the
for the
necessary
primary
synthesis
morphological events during callose
exist between
“lipid complex”,
present
are
stages,
but
tetrad
a
the callose wall
relation
stage
with
wall
of callose.
and
is absent. The
difference
of the
the
and
polysomes
reticulum
the formation of the callose wall
proceeds
found
formation.
lipid granules
is not clear.
in the
reti-
plasma
has been
and cellulose wall
endoplasmic
visible
Golgi body
endoplasmic
The presence
No
change.
cytoplasm
become
polysomes
to smooth
wall.
formation
formation
smooth
similar
cell
do not
in the
same
During
way.
INTRODUCTION
During
meiotic cell
similar to those
No
or
it
same
tochore
didum
for
(Esau
variability.
or
a
a
during metaphase
Nuclear
already
A
polar
I in
has the
reported
Need.
a
20(4), August
as
1971
The centriole in
appear in
(Wilson
by
or a
rebuilding
the
1968)
the
plant
cell
(Tanaka
“polar
1969).
hollow”
The kine-
and in Lilium
structure as
pine during
can-
has been found
early
tetrad stage,
Dickinson & Bell (1970) and
Aldrich & Vasil
of the
1969),
& Jentzsch
morphological
are
organisms.
& Gill
(Robbins
in Pinus banksiana
mixing
which
occur
may sometimes be absent
Tradescantia
same
Podocarpus macrophyllus by
of the nuclear membrane
Acta Bot.
centre
events
the breakdown and the
1969).
polar body (Esau
invaginations,
been
concerning
& Gill
vesicle aggregate
(Dietrich 1968)
pine.
exists
opinion
may appear as
(Burgess 1970),
have
of
many
the mitotic divisions in other
nuclear membrane
shows the
1970)
division in Pinus sylvestris
during
consensus
of the
in
in
telophase II. Roughendoplasmic reticulum
large
Golgi
diplotene and early
1.
and
starts
plastids
appears
(1970).
karyoplasm
and
After the breakdown
cytoplasm
takes
place.
411
M.
M. TH.
In Lilium the
ribosome
Schrauwen
1968 and
increases
population
(Mackenzie,
I
taphase
Dickinson
&
bosomal material from the nucleus
is
supposed
meiosis
Willemse &
by
in
change
mitosis
the
During
Linskens
No
paludosa.
(Roth,
the
of Pinus
in
change
tetrad stage
early
bodies
Golgi
callose wall
microspore.
cell-plate
During
In
1968).
Godwin
the
of
Golgi
the formation of the
microtubules and tubular
been
suggested
contains
Baranova
of callose
Angold
by
callose,
because
ER
is
future cell
wall,
microtubules
in which many
Northcote & Pickett-Heaps
1968 and
cell
wall
Heaps
Golgi
1963; Sassen
(Sievers
(1966) supposed
The
only.
the
in
multiple enzymatical
Pickett-Heaps
by
that
(1967)
cisternae of the
(Werz
a
where the
glucolipid,
D-glucose
membrane
fragments
1969).
body
a
which
is
many
place
acts as a
polymerized
are
1967;
necessary
Villemez
vesi-
present. It has
are
generative
&
cell wall
Petrovskaya-
in
the
phragmoplast
the
to
of the
plane
(Frey-Wyssling
al.
et
1964;
1967; Hepler & Jackson
during
the
function of the
the
formation of the
Northcote &
synthesis
Golgi
by
Wise
particulate
on
primary
Pickett-
the
of
pectic
bodies has been
& Flickinger
enzymes
have
to
supposed
(1970).
been
membrane and is
et
transport of cellulose by the
(Hassid
for cellulose
al.
not retrans-
1969).
cell,
Chemical intact
synthetase activity (Pinsky
particles
vesicles
present
are
1968). However,
Golgi
In
found
1970).
plasma
cellulose
and
substances
carrier and is translocated outside the
Polysaccharide-synthesizing
membrane (Ray
that
to
and demonstrated
Golgi
1968).
separate way cellulose should synthesize
Golgi body
is
&
forming
distinct cell organelle. During cellulose formation, D-glucose is
to a
Ordin
general
& Kellner 1970; Staehelin & Kiermayer
to a
ferred
In
of the
starts
1964; Jordan 1970).
Cellulose formation may take
lated
vesicles fuse
1967).
vesicles
contain and transport material for the
vesicles
restrict the function of the
they
Golgi
(ER)
right angles
at
1966; Pickett-Heaps
Hepler & Newcomb
cellulose wall the
oriented
the
(Echlin
connected with the formation
are
cell-plate
of
microspore
callose
Tsinger
In cellulose walls the formation of the
consists
the
known
cell of the
(Heslop-Harrison
structure
present.
odoratus.
which
tubes of
are
Endymion non-scriptus,
Lathyrus
pollen
occurs
Martens,
vegetative
contain
reticulum
that the
and
purpurella
D.
report that sphaerosomes
(1965)
plugs
in
(1968),
tubular
bodies
place.
separating
cell wall in
endoplasmic
no
callose wall surrounds the whole
generative
and
generative
cles,
Golgi
formation takes
a
callose precursors
or
found
1968)
the meiotic divisions
1966).
which
vesicles,
Dactylorchis fuchsii
contain callose
(1965,
during
In Helleborus foetidus the earliest wall
by aggregation
faba
sylvestris during
the tetrad stage. Some studies
sylvestris during
about the callose wall formation between the
formed
mitosis in Vicia
mitochondria and
showed that
(1967)
&
Transfer of ri-
1970).
and in Pinus
Maruyama
(1968).
me-
1967; Linskens
cytoplasm during
Wilson & Chakraborty
Waterkeyn & Huyskens
microspore
Dickinson
Heslop-Harrison
to
mitochondria and
plastids,
of Tradescantia
during
&
Lafontaine & Chouinard (1963)
by
diakinesis and
mainly during
Heslop-Harrison
WILLEMSE
Brown
takes
place
on
the
&
plasma
(1969) suggests
in
Pleurochrysis
scherffelii.
412
Acta Bot.
Neerl.
20(4), August
1971
MORPHOLOGICAL
CHANGES
the presence
is
ported
membrane. The
2.
MATERIAL
The
same
callose
cell wall materials
is
the
ER
formed
at
3
applied
plays
role in
a
Jackson
H-glucose
cell
and
wall
the pore sites of the sieve
The
in the
produced
are
callose
is
probably
Golgi body
synthesized
and
the
on
necessary for the formation of the callose wall
products
lipid granules
AND
that
and Hepler &
of
1962).
or
and in the
“lipid complex”
trans-
plasma
are
pro-
plastids.
METHODS
material and methods
of this paper
3.
al.
(1967)
labeling pattern
cell-plate,
where
et
vesicles.
stored in the
bably
the
near
(Esau
pine, primary
by Golgi
basis of the
present
in Cucurbita
plate
In
of ER
ER
synthesis.
the
on
suppose,
II
MICROSPOROGENESIS IN PINUS SYLVESTRIS
Hepler & Newcomb
(1967),
Pickett-Heaps
(1968)
DURING
were
used
is described in the
as
preceding part
(Willemse 1971).
RESULTS
3.1.
Prometaphase
During
surrounded
by
and
of
meter
near
the
are
lation
to
30
chromosomes.
electron
nm
and 15
attached
are
I.
smooth
to
Many plastids
contain
an
not
ER
granular
vesicles of the
same
Telophase
polar
a
large
starch
a
which is
marked
polar
lying together
are
in
a
dia-
situated
visible
as
a
i). During
more
from
direction
zone
granu-
pole
point
in
to
re-
ends of the cell
several
distinct
type
They
produce
brane
(fig. 7).
found
during
Acta Bot. Neerl.
The
The mitochondria have
lipid granules
are
contain
situated
an
mainly
diakinesis
are
present. A small popula-
cytoplasm.
I
a
close
contact
of chromosomes and
the
Golgi
small vesicles which fuse
centre
content.
with the chromosomes (fig. 2). In the cytoplasm many
as
ends of the cell
mass
granule (fig. /).
bodies with few vesicles
Golgi
markable is the presence of many
In the
or
are
(fig.
extending
be observed. On the
(SER)
tion of ribosomes is situated in the
geneous
kinetochore,
karyo-
with
kinetochore is
chromosome
chromosomes
A centriole
electron transparent
electron dense material. The
In the
the
granules
The microtubuli
the
the
the border between
contains
(fig. 3,4).
on
on
on
5).
few cristae and
3.2.
nm
structure
the microtubules could
elements similar
groups (fig.
(fig. I).
karyoplasm
The microtubuli between the
present in anaphase
around the
callose wall
During prometaphase
dense
I microtubules
(fig. 4).
a
I
the volume of the cell increases. The cell is
The
cytoplasm (fig. I, 2).
less
anaphase
pole
thin cell wall and
approximately
globular
lar
a
telophase
division,
of the nuclear membrane are visible
Fragments
plasm
I until
these stages of cell
of the cell many
20(4), August
1971
granules
exists between the somewhat homo-
new
nuclear membrane
(fig. 6).
bodies around the chromosome
and
locally
are
present,
form
most
a
new
nuclear
of them
are
Re-
mass.
mem-
approxi413
M. TH.
mately
15
in
nm
diameter. In
the
equatorial
accumulation of small vesicles which
{fig. 8,
microtubule
Interphase
3.3.
The cell has
a
I
taphase
in
the
in
an
end of the
club-shaped
decreases.
cytoplasm
clear membrane remain visible.
a
volume and
large
the
on
each
at
lies
pole
ends and has
polar
The nucleus contains
has few
grown
nucleus. The callose wall
a
in
centre
of about 30
the starch
cytoplasm
nucleoli and
homogeneous
granules
Golgi bodies, producing
In the
of vesicles
a
comparison
with
prome-
{fig. 10).
karyoplasm
In the
number
the microtubuli end
plane
represent
II
rather
thickening
shows
The
9).
vesicles with
Mainly
seem to
M. WILLEMSE
nm
in the
granule
few small vesicles,
of the cell many
sometimes inclusions.
decreases in size
plastids
are
spread throughout
ribosomes
polysomes,
The
in diameter.
and
the
{fig. 10).
cytoplasm.
microtubuli
some
are
present {fig. 11).
3.4.
Prometaphase
The
breakdown of the
dilatations between
membranes {fig.
II
many
or
The
approximately
has
not
found
are
cross
in these
in the
material.
a
II
the
the
at
pole.
The
13).
form {fig.
a crescent
Vesicles and
appearance of
of
fading
In
14).
the
metaphase
A centriole
elements of SER
continues. Some
appears
region
of the
lipid granules
lipid granules
fragments
of
contain
an
mem-
electron
connected with electron
are
{fig. 15). Golgi bodies, producing
dividing
tains many vesicles
vesicles
Golgi
some
this
are
increases
3.5.
are
dense
trans-
The
early
unit
lying
cell division the
during
close
com-
to
cytoplasm
diakinesis. Ribosomes
around the
mainly
II and
are
telophase
the
and
including
The number of
plastids.
remarkable
the
con-
po-
helical polysomes
II small concentric
in
mem-
cytoplasm (fig. 13,15).
arranged opposite
are
homogeneous
possibly
an
of the nucleus.
surrounded with
In the
are
tetrad stage
membranes,
periphery
vesicles,
clear membrane and dilated cisternae of SER
those found
during telophase II;
The nuclei contain
414
a
observed
observed in the
The four nuclei
are
to
{fig. 12). During
stage {fig. 16). During metaphase
branes
the
with
similar
polysomes
lysomes
nucleus
or
Plastids without starch have
granule.
parent vesicles and the whole group is surrounded by dark dots: the “lipid
plex”
two
karyoplasm
regions {fig. 13).
plastids
bowed appearance. The
During telophase
by
in diameter {fig.
nm
been observed.
only
decrease of starch
15
branes become visible around the starch
sometimes
followed
section
with the
locally
starts
Here the breakdown initiates
of
point
SER
to
tetrad stage
early
envelope
membranes,
two
microtubules show in
direction
a
similar
until the
nuclear
the
12).
granules
has many
II
cytoplasm
each other
{fig. 17).
invagination
These vesicles
karyoplasm
the
to
nucleoli
or
of the
near
the
plasma
membrane.
Vesicles surrounded
nuclear
membrane,
by
two
occupy
contain thin fibrillar material and
chromatin material {fig.
decrease of starch in the
plastids
18).
continues. The
Acta Hot. Neerl.
lipid
20(4), August
1971
MORPHOLOGICAL
have
granules
The
CHANGES DURING
a
different electron
bodies
Golgi
ribosomes
are
MICROSPOROOENESIS IN PINUS SYLVESTRIS II
electron dense threads
into
more
3.6.
The callose
thin
threads
vesicles
cell-plate
of the
the
near
telophase
II and
During anaphase
sent
chromosomes. In the
ribosomes have been observed
vesicles
SER.
cytoplasm
which
chromatin,
and
con-
converts
direction
centripetal
visible
are
everywhere
the
large vesicles,
long
flat cisternae
Except
vesicles, large vesicles
cisternae in
like SER,
cisternae
the
well
from
are
In the
places
where the
nucleus
{fig. 33).
the
at
same
The fused vesicles
the
are
direction
flat
the
Golgi
cisternae
vicinity
bodies.
electron transparent
electron
more
contact
now
bodies
cell-plate
vesicles and
other,
the callose
During
fusing
wall,
the
near
plane
Golgi
sometimes
visible in
The
of the future
Remarkable is
the
some
the
two
rem-
persistence
A very thin
somewhat
content
becomes
centre
of the
cell-plate {fig. 28).
in the electron transparent
the membrane of the callose
cell-plate
Golgi
cell-plate (fig. 26).
now
{fig, 25, 27).
which surrounds the whole cell
vesicles
flat
These
long
{fig. 30).
Thereafter the callose becomes visible as
transparent line {fig. 31, 32).
particles mainly
flat
long
cell-plate, looking
with their membranes. The
becomes
of the vesicles
disappears
small round
The
transparent, the volume increases and in the
(fig. 29,30).
to
clearly distinguishable {fig. 25, 26, 27).
vesicle fine fibrillar material appears in the
vesicle
the
on
small round
produce
cell-plate {fig. 24).
the first elements for the
The fine fibrillar material
pre-
similar
cisterna,
(fig. 22, 23).
Many Golgi
membranes touch each
centre
or
of the future
of the future
by making
of the
The ribosomes
irregular shaped vesicles,
of these membranes remain visible in the vesicles
line in
large
{fig. 20).
are
connected with
present in the early tetrad stage {fig. 21). The
long
region
vesicles
vesicles
stage
dilated vesicle
very
However,
of the unit membrane become
layers
then
as
Golgi
situated in the
cisternae and vesicles fuse
nants
a
and in the
cytoplasm
originate
are
as
tetrad
many small
flat cisternae look like SER.
long
irregular shaped
II many
early
the
near
and the result is
disappear
vesicles and
of
containing
content
a
very
of the
electron
lead-containing
vesicles and
near
{fig. 21, 30).
formation microtubules
are
found
perpendicularly
to
{fig. 34).
DISCUSSION AND
CONCLUSION
The nucleus
A local
and
the
{fig. 17).
time
4.1.
in
wall formation
First the
4.
zones
of
between the four cells grows in
the
present.
{fig. 19, 34).
The callose wall
fusing
is also
“lipid complex”
cytoplasm {fig. 17). Many polysomes
Some electron transparent
present.
material, resembling
tain fibrillar
the
density,
in the
dispersed
are
breakdown,
prometaphase
anthus
(Bajer
visible in the
Acta Bot.
Need.
possibly enzymatical,
II
starts at
& Mole-Bajer
cytoplasm.
the
1969).
After
20(4), August
pole,
1971
of the nuclear membrane in diakinese
which has also been
reported
for Haem-
Remnants of the nuclear membrane remain
diplotene
no
rough
ER
(RER),
but
only
short
415
M. TH. M.
cisternae of SER
the
during
present, therefore the membrane bundles,
are
division
membrane. In
pine
have
stages,
the
nuclear
new
of the nuclear membranes and of
while also
nucleus,
fusing vesicles, forming
In HeLa cells Robbins & Jentsch
sent.
nuclear
membrane from
rebuild the
to
plasma
Phaseolus
roots
in the cell
coat
of several types of
I the
During prometaphase
structure
the
on
visible
on
in
as
the
postulates,
ends,
In
but
a
interphase
II
inclusions
invaginations
those described
(1970)
and for Pinus
synthetized
the
polar
also
which
observed in
for
stage
to
the
a
starts
may
This material possibly
after the
In
(1970).
granules
originate
with the
ed in
cytoplasm
of
and
become
mitotic cells
nm,
nm
by
in
the
Wilson
a
In
pine
(1971)
the
similar
visible,
Vasil
to
& Aldrich
the
Podocarpus
invagina-
exine precursors which
contain and
in the
zones
by
cytoplasm
means
wall
pollen
are
of the nucleus
out
nucleus
trans-
the vesicles
sylvestris
of material
the
of the
formation
direct relation has
again
of Vicia
karyoplasmic
centre.
faba
a
not
been
spindle
of
karyoplasm
nuclei.
During
II
resemble ribosomes.
and
telophase
production
with the
& Gill
dividing mesophyll
of
cytoplasm,
II after
(1969)
cells
granules
which
show the
are
diplo-
and 15
Lafontaine & Chouinard (1963)
nucleolar
granules
in the
new
probably ribonucleoproteins,
visible during interphase
in the cell
granules
in the
granules strongly
resemblance with ribosomes. Esau
somes
As
early
role in the
a
envelope,
30
approximately
cytoplasm
of these
centriole
a
been observed in
invaginations
However,
included
not
from nucleoli. The 15
polysomes
electron
Tradescantia
clearly.
After breakdown of the nuclear
mixed in the
less
a
in
the nucleus.
some
In Pinus
transport
plays
tetrad stage.
early
as
polar
be delivered from the
possibly
the
pine
SER is found in the
Podocarpus
cytoplasm.
If there is
In
in
and
here,
I the microtubuli be-
1970).
pine
cell-plate
1969).
nuclear membrane become
within the nucleus. In Pinus the
sylvestris
demonstrated
416
In
vesicles
present.
channels for the transport of
invaginations.
invaginations.
ture
not
reform
to
Golgi
resembling
not
thin fibrillar material present in electron transparent
of Pinus
tene
(Tanaka
Dickinson & Bell
by
as
has
pre-
vesicles.
visible
structure
1970)
centre.
have been
tetrad
port probably nucleic acids
probably
a
of the
the
during
tions may function
are
as
al.
e?
metaphase
organized
& Gill 1969; Burgess
ER may function
tetrad stage
In
1968).
relation with microtubules is
is
pine
are
aggregates.
as
is described
as
has been observed
as
ranarum
of the
fusing Golgi
kinetochore in
Basidiobolus
fungus
pine,
(Rambourg
from
chromosomes
the kinetochore. An
polar body (Esau
or
cells
rat
partly
membrane,
vesicular
polar
capacity
in
or
telophase
the re-formation of the
the formation of the
by
1967)
1968) and Lilium (Dietrich
(Wilson
come
also
originates
The
1969).
& Newcomb
from
the
partly
surround the
nuclear
new
nuclear
remnants
rebuild
to
visible
are
of the
have also been described
membrane is shown
(Helper
nuclear membrane
dense
& Gill
(Esau
a
derived
spheroids
bodies
(1969) supposed
Fusion of vesicles and cisternae of the ER
the nuclear membrane
which fuse
Golgi
Many
which
remnants
as
may be formed from the
envelope
Golgi vesicles,
nuclear membrane parts.
lacking
be considered
to
WILLEMSE
mixture
describ-
and
have
presence
nm
Many
a
a
mix-
strong
of ribo-
of Nicotiana and suggest
Ada
Bot. Neerl.
20(4), August
an
1971
MORPHOLOGICAL
CHANGES DURING
MICROSPOROGENESIS IN PINUS SYLVESTRIS
II
entry of ribosomes in the nuclear region. In pine the karyoplasmic granules
of ribosomes.
precursors
and
are
included in the
not
Heslop-Harrison
and
Trillium
by
may
be restricted
structures
to
I and II
(Tucker
in the
and
metaphase
interpreted
as
they
a
situated
are
of microtubule
stage
& Jackson
in the
4.2.
The
with the
content
mitochondria have
Roth
et
Golgi
bodies
al.
microsporogenesis
body changes
No
in
the
in
an
no
electron
no
in the
Tradescantia,
are
Dickinson
but in
changes
cross
(1969),
pro-
can
be
bridges
Hepler
1967).
prometaphase
1968)
as
an
as
in Lilium has
As in zygotene and
II.
Many
in
in
the division stages many
surround the
rat
dividing
hepatic
Ribosomes
in
are
always present.
pollen
observed
the “lipid
in
that
during
Golgi
telophase
II
In all
may
inter-
stages
absence of the
(Heslop-Harrison
complex”
&
appears again
content
and
a
development. Mainly during
vesicles and cisternae of SER,
1967)
polysomes
as
has been found in
could not be observed. The
are
The presence of helical
mother cells of Ipomoea
and
cristae.
microspores,
electron transparent
& Lee
RER is absent. After the nuclear divisions
is in
in the cell. The
Golgi body stage.
vesicles, mainly Golgi
(Dougherty
few
but the
occur,
lipid granula during
been
an
possible
II and
nuclei. A local accumulation of SER
cells
spherical
and
position
Tradescantia
early
pachytene,
vesicles with
both
content
It is
place.
changes
of
not
interphase II,
of Tradescantia. The
pine
in the mitochondria
changes
multilayered rings
(1966,
in
in the number of mitochondria and
clear membrane have been observed in all stages of
4.3.
Wilson
plastids
transparent
present .An aggregation
nuclear membrane
reported
or
during
They
1968). During microsporogenesis
of vesicles and
production
Maruyama
lipid granules
dividing
links
plastids starting
content
mitotic division take
pine
with the
compared
preted by
in
form.
Nassula
development stage
a
ofthe nucleus.
region
small concentric membranes found during metaphase
be
ciliate
observed
are
reported by
(Maruyama
that
(1966) supposed
during
in
disappear
rod-like mitochondria persist
the
or
crescent
a
the
(1964).
of the
starch
unchanging
rod-like mitochondria
pine
cytoplasm
phenomenon
cytoplasm
The decrease of the starch
contrast
mainly
found in
development.
and Krishan & Buck
(1968)
those
as
in Lilium
only.
either artefacts
between microtubuli have been observed
This
1968).
the C-filaments
mainly
the
provide
a way to
microtubuli have
are
pine
Dickinson &
and
(1968),
outburst of ribosomes derived
& Linskens
C-filaments like
These C-filaments
1967).
This
(1967).
some
microtubule formation. In
are
nucleus
diplotene
reported by
Schrauwen
the meiotic division
the
are
has been
&
the meiotic cell divisions is
during
During prometaphase
These
al.
the
I nucleus. A rise in the number of riboso-
meiocytes
nuclear information (Willemse
new
not
et
visible in
clearly
and Linskens
(1970)
Mackenzie
from the nucleus
with
telophase
I in Lilium
during metaphase
mes
become
They
found in the
polysomes
cytoplasm.
has also been
(Echlin 1965).
Callose wall formation
Callose wall formation takes
Acta Bot. Neerl.
20(4), August
place during diplotene
1971
and the
early
tetrad stage.
417
M.
I and II microtubuli
During telophase
to
the
icles
plane
absent.
are
The
pine,
cell-plate.
microtubuli
surrounds the cell. In
the
are
I
standing perpendicularly
diplotene
WILLEMSE
cell wall is formed and
no
fibres
phragmoplast
&
(Bajer
ves-
Jensen
found like those
are
early
for Haemanthus. In the
(1968)
M.
observed, standing perpendicularly
were
telophase
material and end knobs
amorphous
Hepler & Jackson
some
In
microtubuli,
lie in clusters in
1969)
reported by
of
of the future
TH.
tetrad stage
the callose wall which
to
relation between microtubuli and the callose
no
wall formation has been found.
the
During
formation of the
appear. These vesicles and the
ced
by
the
bodies. The
Golgi
has been
Golgi body
similar
nae
SER
to
(Brown
has
be
to
coated membrane of
fuzzy
Hepler & Newcomb
In the
of the
plane
of the
Golgi body
ducts becomes
plasma
1969).
(1967)
membrane (Hepler
is
non
gested.
not
or
thesis of
during
particulate
for the
reason
et
al.
(or
of callose
necessity
growth
within fused
of the
presence
the
1967)
and the
contact
formation of the
wall formation
starts
long
cell-plate;
are
on
the
may be
vesicles
Golgi
cisternae.
the
case), they
not
callose
of the
observed: small and
418
Golgi body,
syn-
Golgi
to
which
Werz
&
of the
enzymes
present for
function
on
since the appear-
the
transport. The
from the
vesicles
further
(Hepler
&
membrane with the callose wall
the
tetrad stage,
when
will be described in the
becomes
large Golgi vesicles,
on
layers
are
have
appears
visible
transparent central region, which subsequently changes
cisternae derived from the
the
to
sug-
of the
sign
particulate
during
membrane
in
two
If enzymes
first
into
parent line in the plane of the cell-plate. During diplotene the
are
(1967)
a
1970;
plasma membrane,
plasma
as
in the
phenome-
Golgi body
Kiermayer
located between the
disappearance
protrusions
not
that this
supposition
&
vesicles and
plasma
of the
pro-
1969; Ray 1967). This may be also
In the
Golgi
(Willemse 1971a)
thereafter
Golgi
but
cell-plate,
SER,
possibly
then outside the
Acta
an
the
next
In the fused vesicles fine fibrillar material appears in line with the
future
A
by
flat cisternae
Hepler & Newcomb
as
cellulose in another
of the cell wall after the
Newcomb
during
starts
of the
Phaseolus
these accentuations of
pine
In
of the
the
(Staehelin
or
Golgi body.
these membranes with relation
1969).
the membranes of fused vesicles
ance
on
the enzymes
(Wehrli
of callose
to
elal. 1968; Hassid
present derived from the
synthesis
rise
sectioning
present
are
1970). Probably
membrane
could be
1967).
plane
dark stained membranes of the
always
enzymes
vesicles and
large
membrane of the vesicles
enzymes
callose(ViLLEMEZ
Kellner
the
in the unit
flat cister-
long
for
reported
as
on one
the fusion. This membrane is the future
& Newcomb
Golgi body, give
a
change
of
particulate
unit
near
of
production
features of the
produ-
the transport
as
unit membrane of these
surrounding
the result of normal
The
presence
the
or
well
as
are
vesicles
observed.
not
the small
the unit membrane of the vesicles in the
cytoplasm
the
to
SER-like elements
the SER
to
large Golgi
(1967),
the
pine
vesicles
Golgi
was
The
In
adjointed
accentuated
more
of small and
Pickett-Heaps
the
cell-plate
fuse.
flat cisternae similar
production
reported by
of the whole cisternae
many vesicles and
cell-plate
long
as
plane
an
Bot. Neerl.
trans-
phenomena
also the
plasma
of the
electron
electron
same
pollen
part.
long
flat
membrane first
20(4), August
1971
MORPHOLOGICAL CHANGES
MICROSPOROOENESIS IN
DURING
the fine fibrillar material and
wall. Callose wall formation
for the
cell wall
primary
wall formation does
the
II
the electron transparent line of the callose
finally
depends
and
PINUS SYLVESTRIS
the
on
vesicles
Golgi
of the
presence
materials
containing
membrane. Callose
plasma
differ from the cellulose wall formation in its
not
morpho-
logical description.
In
the role of the SER is
pine
tetrad stage,
may
be
as
analogous
Newcomb
also
reported
ribosomes.
ing
ER is
1967),
the
pine
bodies.
Golgi
mains
is
difficulty
distinguish
to
cells
al.
(Angold
long
of
presence
1968)
and of
1966).
callose wall
during
and the
re-
callose
lipid granules
Tsinger & Petrovskaya
as
&
and SER
flat cisternae of the
polysomes
“lipid complex”
cells,
(Hepler
1962)
the formation of the
to
the
et
early
disappear-
1967 ; Heslop-Harrison
the SER from the
their relation
plastids,
plasmodesmata
plates (Esau
the
HeLa
dividing
because of the
SER,
generative
Remarkable is the
in
(1967)
formation of
storage for polysaccharides
a
to
microspore (Angold
wall formation. Besides the
may
the
sites of sieve
callose wall of
Therefore,
hypothetical.
be
to
pore
SER before
to
coated with ribosomes in
Robbins & Jentz
by
related
the
to
the callose wall around the
In
clear. A transition of RER
the transition from RER
to
the formation of the
to
not
has been described. The vesicles
diplotene
(1965)
sug-
gested.
The
ume
cell
and the
centripetal ingrowth
connected with the
of
cytoplasm
is great
on
the
the presence of microtubuli
on
the
during
membrane. The pattern of
jacent cytoplasm
as
growth
of the
Hepler & Newcomb
on
cell-plate
the
the border of the
vesicles may
1967).
position
may be
depend
The orientation of
and in the
during diplotene
depend
at
Golgi
& Newcomb
(Hepler
formation may
cell-plate
the onset, because the vol-
at
ends of the cell and
polar
the fine fibrillar material around the cell
icles
vesicles
Golgi
after the divisions. The orientation of the
centre
of the callose wall may be
polar thickenings
number of
high
Golgi
of the
regulated by
ves-
plasma
the ad-
(1967) suggested.
ACKNOWLEDGEMENTS
The
M.
author is much
M,
A.
Sassen
G. W. M.
Barendse
Dicke
his
for
indebted
for
skilful
for
his
to Prof.
critical
the translation
technical
Dr.
reading
and
F.
H.
of the
Linskens for his
stimulating interest, to
manuscript,
Dr.
correction
of the
to
P.
van
manuscript,
Gijzel
and to
Dr.
and Dr.
Mr. A. W.
assistance.
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Cell.
fracturing. Expl.
Protoplasma 69:
(1971a): Morphological
P.
& P.
218: 878-880.
organelles during microsporogenesis
from
—
McNab
(Land.)
K. Mühlethaler
method
replica
Werz,
J.
L.,
saccharides.
Personal
E.
Amoeba
&
C.
communication.
J.
proteus.
Acta Bot. Neerl.
Flickinger
Planta
(in press).
(1970): Cytocheraical staining
of the
Golgi
apparatus
in
J. Cel! Biot. 46: 620-626.
20(4), August
1971
421
M.
I.
Fig. 1. Prometaphase
(Nm),
Fig.
Golgi
bodies
cytoplasm. Golgi
Plastids
bodies
with starch
3. Kinetochore
during prometaphase
4. Kinetochore
during anaphase
Fig.
5. Detail
6.
Telophase 1; rebuilding of
nucleus,
7. Detail
9. Detail microtubule
II.
Interphase
with
Cell
11.
Interphase II.
Fig.
12.
Piometaphase
nules
41,000
x
karyoplasm,
SER
cell,
and
51,800.
x
microtubuli
bodies
Golgi
many
Golgi body producing
centre
in the cell
(arrow),
club-shaped end (arrow),
with
(g)
and
In the cell
callose
wall
and
(C)
x
(m),
surround the
(G)
vesicles
(v)
which
fuse
21,600.
x
108,000.
starch
decreasing
vesicles
Many
in the
(A)
plastids,
Fig.
14,
Microtubules
Fig.
15.
Telophase
near
II.
in the
the
16.
Helical
Fig.
17.
Early
with
polysomes
tetrad stage.
in
Note
18. Nuclear
invaginations,
Fig.
19. Electron
transparent zone with
Fig.
21.
Golgi
with ribosomes
bodies
(arrow),
x
(G)
Fig.
22.
Golgi body
23.
Golgi
with
Fig.
24.
Cell-plate
Fig.
25.
Fusing Golgi
bodies
a
near
the
granular
have
No
zone.
distinct
polar
centre.
a
form
crescent
(arrow),
X
32.200.
Concentric membranes
(c)
and
47,300.
x
visible
between
the cells.
“lipid complex” (Lc),
fine
fibrillar
the nucleus.
material,
Polysomes
with
large vesicle,
thin
vesicles
x
are
x
In the cell
light
zones
8,640.
43,750.
visible
vesicles
many
(v)
(p),
x
and
29,400.
long
flat
cisternae
and
Fig.
28. Detail
clear
a
is visible
cell-plate,
content of the
x
x
vesicles,
more
becomes
30.
Detail of the
31.
In the vesicles the callose becomes
Fig.
32. Detail of the callose
Fig.
33.
Centripetal ingrowth
Fig.
34.
Microtubules
x
44,400.
Within
the
39,900.
48,900.
transparent
content and fine
more
electron
fibrils
in
plane
Note
the
black,
visible,
X
46,000.
transparent.
in the vesicles remnants of membranes
visible
containing vesicle,
of the callose
perpendicular
x
centre,
clearer.
lead
21,900.
Fig.
(arrow),
x
the cell
becomes
39,900.
x
electron
Fig.
vesicles,
x
membrane
36,800.
vesicles
containing particles,
the
(arrow),
membrane,
of fused vesicle with
50,600.
x
vesicles and cisternae in
cisternae,
in the
35,000.
x
cisternae,
line becomes visible
27. The thin line
of the future
flat
Many Golgi
formation.
26. Fused vesicles with
mentioned
gra-
31,500.
the
cytoplasm
small and
Fig.
threads
with
44,400.
producing long
Fig.
29. The
21,900.
21,900.
Fig.
vesicles
in
x
x
(Nm). Karyoplasm
5,400.
cell,
becomes
(arrow).
polysomes (p),
cytoplasm (Lc).
x
II
the
x
Some
the
telophase
Fig.
20. Vesicles
in
plastids,
Cell-plate
fibrillar material
Fig.
(c),
the chromosomes.
“Lipid complex”
decreasing starch (A)
Fig.
around
(v)
and
membrane
24,800.
x
present
are
(m)
of the nuclear
Golgi bodies (G),
II.
Anaphase
centre microtubules
II. Breakdown
Concentric membranes
422
Granular
3,060.
Fig.
Unless
karyoplasm,
of the
pole
granules (g).
many
15,000,
membrane,
with
(Nm)
Telophase I; ending microtubuli
x
Fig.
WILLEMSE
membrane
28,800.
x
8.
13.
the
near
with
karyoplasm
x
microtubuli.
the nuclear
of nuclear membrane
Fig.
Fig.
TH. M.
nuclear
17,400.
x
Fig.
10.
the
(M),
I. Granular
I with
I
cytoplasm anaphase
(arrow),
Fig.
of the
21,900.
new
Fig.
round
(G)
Fig.
Fig.
(G)
and mitochondria
(A)
Fig.
x
and remnants
4,770.
x
2. Detail
Note the
to
wall.
the
x
(arrow),
x
are
18,900.
47,500.
Note the
Golgi bodies (G),
coming cell-plate (m).
Note
x
the
14,700.
electron
dense
22,000.
otherwise,
the line
in the
figures represents
a
length
of 1 jxm.
Acta Bot. Neerl.
20(4), August
1971
MORPHOLOGICAL
CHANGES
Acta Bot. Need.
20(4), August
DURING
MICROSPOROGENESIS
1971
IN
PINUS
SYLVESTRIS
II
423
M. TH.
424
Acta
Bot. Neerl.
M.
WILLEMSE
20(4), August
1971
MORPHOLOGICAL
CHANGES DURING
Acta Bot. NeerL
20(4), August
MICROSPOROGENHSIS IN PINUS SYLVESTRIS
1971
II
425
M.
426
Acta 801.
Neerl.
TH.
M.
WILLfcMJrE
20(4), August
1971
MORPHOLOGICAL CHANGES
Acta Bot.
Neerl.
DURING
20(4), August
MICROSPOROGENESIS IN PINUS SYLVESTRIS
1971
II
427