Acta Bot. Neerl.
December
21(6),
1972,
Integumentary
p. 617-629
studies
in
the
Polycarpicae.
stellata
II. Magnolia
Magnolia
virginiana
R.
Bouman
de
Hugo
Boer
de
and F.
Vries-laboratorium, Universiteit
van
and
Amsterdam
SUMMARY
The
histogenetic development of
Magnolia
stellata
and M.
these two
species.
The two
ment
(o.i.)
initial
is
(i.i.)
wholly
is formed
or
cell
divisions,
which
the
younger
(in
region.
i.i.
only
After fertilization
forming the
1.
the
The
studied.
have
derivation,
either
the whole
o.i. is
integuments
of dermal
ring-shaped wall,
teguments.
was
representatives of
two
The mode
of
different form of
a
whereas
the
integument.The
stages
at
becomes
in
meristematic.
least) four-layered.
thicker
by
appreciable changes
sclerotesta and the outer
layers
The
take
place
mutual
The
cell
in the
the vascularized
rows
ofcells
no
apical
by repeated
to form
a
mainly three-layered,
is formed
divisions
o.i.,
integu-
integument
juxtaposition
i.i. is
in
same
inner
There is
layer.
micropyle
“secondary”
is the
of the outer
initials of the i.i. form
remaining
Magnoliaceae,
initiation; The
primordium
develop independently
rows
structure
the
development
the result of cell divisions in the first subdermal cell
as
apical growth in
periclinal
integuments of
the
virginiana,
the
in
both
by
the
in-
micropylar
innermost
cell
layer
sarcotesta.
INTRODUCTION
The
of the
family
and its
logists
to
study
the ovule
are
this
and
endosperm,
pyle being
ovule is
formed
by
the
the various
and the
ing
the
to
integument
the
the
Angiosperms
1967,
Liliiflorae (Bouman
instance,
are
formed in
(mature) stages
are
restricted
According
such
to the
to
to
the devel-
development
Davis
(1966)
the
a
(compare
and
some
indicate
Warming
b).
The
Illiciaceae,
the
1913,
integument
publications
quite clearly
integuments
deal-
that there
are
between different
Roth
1957,
Boese-
multi-layered integuments
of
Polycarpicae (e.g., present paper), for
fundamentally different
very similar
into the relations
improved by comparing
detailed. The few
of the
histogenesis
a)
insight
Schisandraceae,
as
However, reports concerning
sufficiently
Bouman 1971 a,
1971
Our
together.
development already
families of
the
characters of
embryological
mainly
Winteraceae could be
features.
and not
differences in the
& Bouman
the
gametogenesis,
embryogenesis.
Polycarpicae,
appreciable
winkel
as
theories,
phytomorpho-
microanatropous, bitegmic, and crassinucellate, the
embryological
scarce
phylogenetic
has induced many
were
features of the
Amonaceae and
testa are
with
to
As far
investigations
the i.i. and the o.i.
with other families of
Degeneriaceae,
particular
intensively.
taxon
concerned,
magnoliaceous
in
status
opment of the archespore,
of the
in various
Magnoliaceae figures largely
alleged primitive
(Huber 1969).
way,
although
their ultimate
618
There
The
initiation and
of
of
possible contribution
ses to
taxonomic and
generalizations
papers sometimes
quate
an
accurate
later stages
of the
“by
Maneval
the innermost
(1914)
Kapil &
to
also
as
yet
no
embryological
they
inade-
too
are
too
integuments
of
incomplete
and
be useful:
to
is from Asa Gray
of the ovule into
ovule,
the
but the
changes
Gray
a
(1858),
who
young seed.
histogenetic details
in the
of the
structure
noted that the hard sclerotesta
Magnolia virginiana
the i.i. is initiated earlier
Gray whose results he confirmed.
that
(1964) reported
state
stimulus of the
publications,
rule
taxon.
proces-
is recorded.
Magnolia
of the
stellata and of M. obovata the i.i. is 3
They
a
layer of young cells lining the primine {i.e., the o.i.)”.
found that in
Bhandari
principal
a
features of
within
illustrating
as
F. BOUMAN
histogenetic
descriptions
layers
particularly
more
than the o.i. He referred
layers.
The
development
development
only,
but
generally speaking,
after fertilization of the ovule.
integuments
is formed
AND
definite conclusions concerning the histo-
the genus
concerning
description
drawings
indications,
more
integuments.
are,
He described the whole
of the
certain
of the
growth
The first report
has been the
Line
the ultimate number of cell
mostly only
gave
up.
drawing of
genetic development
of their mode of
drawn
provide
the
permit
to
of these
greater knowledge
a
characters
embryological
the small number of relevant
to
be
can
other
phylogenetic inquiry
study. Owing
present
BOER
strong indications of the constancy of the histogenetic
are
integument
cell
DE
R.
that
to
in the
4 cell
the i.i. is
seed
mature
layers
of
Magnolia
thick and the o.i.
of dermal and the
o.i.
8
10
to
of subdermal
origin.
Hayashi
“the inner
(1964), who
integument
studied
arises
as
Magnolia liliiflora
rim-like
a
lus and the development of the primordium of the
Bhandari (1967) confirmed
The present paper aims
histogenesis
2.
of the
of
V,
of M.
wax).
Sections
7(jl
&
origin
description
paraffin
virginiana could
thick
were
of selected sections: since
may
give
of 488 slides
be followed
were
to
cut
Presumably
seeds,
not
and
perfectly
wax
not
used.
Integument
micropyle
or
be
in Ralwax
Safranin
older stages
paraffin
according
drawings
longitudinal
in
owing
cold for
mature
fixed in Craf
sectioned in
to
were
sections of
an
M.
to
stellata
seeds develop
only
up
incomplete
to
to
quite regularly.
ovule
total
could
the stage
seed
produce
Jo-
made
a
Magnolia virginiana
but in M. stellata
after fertilization
virginiana
virginiana L.,
the best sections selected from
development
too
was
(because
satisfactorily
median
maturity,
the Dutch climate is
whereas in M.
sarcotesta.
initiation and
and of M.
of Amsterdam,
stained with
misinterpretation, only
the stage of seed
of closure of the
of the
of the
hansen and counterstained with Astra Blue. Camera lucida
cause to
says:
the nucel-
integument follows
of the
Zucc.) Blackburn
Garden, University
and embedded in
development
surface of
METHODS
collected in the Botanical
or
Micheliafuscata,
the
Magnolia.
Material of Magnolia stellata (Sieb.
III, IV,
detailed
more
in
integuments
AND
MATERIALS
outer
Gray’s description
at a
and
outgrowth from
setting.
mature
As far
as
INTEGUMENTARY STUDIES IN
can
be
HISTOGENETIC
3.1.
Leinfellner (1967) the
to
vascular innervation of the ovules is
which is
fig. 1),
axis. In each
in
two
ANALYSIS
section
a
pistil
through
flower
a
The
juxtaposition.
ovules
section
the
through
about by
a
of the
growth
subdermal cells
The
the
in
i.i.,
the i.i. also
1
Fig.
a
the
required
of
in
occurs
a
transverse
longitudinal
of the ovule is brought
curvature
form
part
the ovules
the floral axis. A
to
side of the ovule
to
shows
showing
primordium {fig. 2).
at
ultimately
the
side
concave
circular raised
a
primordium
One of these
cell which in
turn
divide periclinally
and
cells, distinguished by
divides into
a
its
thus
larger
macrospore mother
etc.
contrast to
some
ovule
{fig. 3a).
archespore
parietal cell,
a
level.
floral axis
provides
laterally
The
{v.m.
trace
become anatropous
to
peltate
trace.
primordium.
of the
make the nucellus grow
cell and
at one
this
of
short time before the o.i. and initiates first
wall around the nucellus
size, becomes
the
to
plane perpendicular
of the ovule from where it extends
The
same
ovule
the
to
of vascular bundles in the floral
ring
the
(compare fig. 1). The
somewhat faster
a
The i.i. appears
at
flower, accordingly,
the ovules
through
a
branches
supplied by
perpendicular
bending
“horizontal” plane, i.e., in
section
develop
Magnolia belongs
ventral-median vascular
a
derivative of the inner
two
of
carpel
type. The only indicationis the presence of
a
in the
same
Generalities
According
in
development is the
integument
studied.
species
3.
the ultimate
ascertained,
619
THE POLYCARPICAE II
the
periclinal
o.i.,
is
entirely
divisions
occur
of dermal
but
origin,
in the subdermal
at
tissue,
the base of
so
that this
part (the chalazal zone) becomes slightly broader {fig. 5). The o.i. is initiated by
the
divisional
periclinal
ward the
rate
of
growth
Whilst the ovule is
cap
(see fig. 2).
funicle {fig.
features,
ty this
bending,
is
clearly
bright
protective
left
area
a
tissue forms
orange
on
sclerotesta
white).
in
a
a
micropyle
large
mass
like
This,
a
part of the
fleshy
parts
near
as
a
the
developmental
dispersal.
layer
of the
lies
parenchymatic
sarcotesta
the outside. The innermost cell
around the enclosed
of
other
future function in seed
At maturi-
with oil
cells
of the o.i. forms
seed
containing
the
embryo.
The funicular vascular strand bifurcates
ending
the
completed
on-
same.
the o.i. overgrows the distal part of the ovule
associated with
and the
endosperm
the
this stage
remains about the
integuments
On the chalazal side of the ovule
parenchymatic
which is
a
2d).
of these subdermal cells. From
When the curvation is
develops {fig. 3c,
tissue
activity
of the two
dendroid ramification in the
near
the chalaza,
hypostase
of its branches
one
where the vascular elements
become surrounded by tannin-containing tissue deeply staining with Safranin,
and the
other branch
o.i .{fig.
5).
At
{dg
in
maturity
fig.
1),
the
soon
fruit of
bifurcating
once
more
Magnolia virginiana
into strands
dehisces
which does not coincide with the closed
along
suture
innervating
a
dorsal
of the
the
groove
carpel
mar-
620
DE
R.
Fig.
1.
carpel
(d.g.
ring
Transverse
at the level
dorsal
=
of
outward
two
bundles
and enters the
bundles
0 o.s.b.),
of
the
to
the
split
and
cate
( d.m.).
The
v.m.
ovules. The two
off
run
branches
of the
has
of the
divide
2.
sends
of
of the
in
tegument,p.c.
developmental
procambial
stages
strand.
of
an
ovule: i.i.
=
also
carpel
condupli-
of initiation
phase
a
of
in which
initiated,
d
stages
drawn
represents
a
b the
inner
the
different
stage
and
=
o.s.b.
enter
development
Successive
off branches
development
outline;
3.
the
dorsal-
Longitudinal sections
of ovules
is
system
into
the
bundles
margin
between
pistil.
Fig.
Fig.
base
inner
moved
cortical
and
which
the
along
part
of the ovules
One
to
the
through
pistil
(v.m.)
F. BOUMAN
(i.s.b .)
outer
subsequently
ventral-median
median
section
of insertion
groove).
stelar
AND
BOER
c
young
integument,
the
of
the
in
the
i.i.,
the o.i.
further
the
ovule,
seed.
o.i.
=
outer
in-
INTEGUMENTARY STUDIES
gins.
did
The
IN
hanging
in the literature.
reported
protrude
on
Upon
a
zone
of abscission.
majority
light
only
one
supporting branch by
develop
seed matures, and in
into
seeds,
some two
We
repeatedly
as
touch the whole infructescence
from the
of the ovules
Only part
of the follicles
from the desiccated fruit valves.
thread-like funicular remains
because the stalk becomes separated
ed
621
POLYCARPICAE n
coloured seeds
brightly
observe seeds
not
THE
drops off,
a
so
seeds
preform-
that in the
or none at
all.
3.2.
The
inner
The i.i. is
outward
integument
of dermal
entirely
bulge (figs.
gument initiation
3 and
are
The
ally
such
outer two
as
a
a
rule,
way
but
that
tral
initial. The
and
8b).
Such
side of the
Fig.
some
during
cells of the
by
a
In
The subdermal tissue
these
to
three cells
divisions
of
do
a more
oblique
primordium
cellular arrangement is
patterns
than
in the
forms
most
c
on
are
the
produced
opposite
in
)
a
slight
which show
larger
growing integuments.
b,
c,
divide
is
leaning
highly
side
7a,
not
wall which
daughter cell is formed obliquely
resulting
(a, b,
(compare figs.
integument primordium
means
at
median section the first indications of inte-
peri-
variation in the number and in the orientation of
slanting cell walls
4. Various cell
a
confined
usually
clinal divisions. There is
the walls formed
origin.
4).
and
always
up
characteristic
(figs. 3c,
8).
oriented in
against
numbers
fig.
quite periclin-
on
the
cen-
(figs.
7b
the abaxial
4a and h,
7c).
The
622
5.
Fig.
seed:
Diagrammatic
f.b.
=
c.b
bundle,
embryo
sac
sents the
rate of cell division is
is
integument
the
On
usually
The
of the
transversely
and
a
for
4b,
instance).
The various
for
the
more
instance,
ment
in
initials
thus
b and
c
exhibit
ultimate
layered,
how
a
fig. 4c)
a
certain
appears
not
case
Although
two-
or
one
young
hypostase,
tissue,
dotted
area
c
—
en
=
repre-
side,
so
that the
the
with
primordium
cells
of
marked
The
only originate from
three
conclusion
a row
hold,
the
as
in
in
6.
Detail
cells
of the
divide
in
accom-
figs.
a,
b
and
c
2
be
recognised
in
fig. 4b,
of the integu-
of cells of the
rows
be
can
shown
drawn that
basal
the three
divisions. This
deduced from fig.
4a. The
is
for
the
sections.
a
of
two
of the inner two
dermal
periclinally.
the
8a demonstrates
condition
in
(cut
out
of the sclerotesta. The
become
hilum
of the o.i.
(as
divid-
periclinally
layers
formation
layer
Around
Fig.
their
three-
greater part
two-layered
means
4a
hypostase.
determination of
fig. 5) during the
periclinally.
s
the
some
initiation by
between the
in
integument
Fig.
a
d and
by periclinal
be
can
be initiated, but
an
lying
pattern
may
of cells
opportunism
four-layered
can
the
formed is
increases in size. This is
forms the
area
wall
periclinal
8b).
integument primordium
the
always
amount
four-layered pattern
can
first
development.
correspond
this
does
destination.
it
of
have each formed
simple description
cells
In
the
on
be traced back from fig. 7b. The cells
fig. 2).
,
a,
2) gradually
of the
stages
the
e.g., fig.
activity (of
configurations
can
(see,
The tissue in this
advanced
primordium
part (A
in fig.
mitotic
typical
=
a
tegumentary
=
micellar
(The
exostome.
=
of the ovule
zone
bases of the i.i. and o.i. (A x
panied by
ex
through
t.b.
h
=
BOUMAN
tannin-containingtissue).
primordium
oriented
narrow
very
bundle,
endosperm, n
F.
the nucellus.
pressed against
initially
chalazal
therefore, relatively, greater
adaxial side
more
bundle,
=
BOER AND
median section
funicular
with
endostome,
in
DE
R.
extended
the
and
subdermal
divide
cells
INTEGUMENTARY
STUDIES
IN
THE
POLYCARPICAE
623
II
7.
Fig.
Successive
initiation
of the
o.i.
of
=
outer
integument. Daughter
cells
of
the
ed
the
by
8. Initiation
initials
same
(“cell families”)
Fig.
stages
i.i.,;
are
indicat-
same letter.
of the inner
integu-
ment.
ed cells. Also from fig.
homogeneous,
cell,
as
4d
all cells
described for
The cells marked h
in the
l
micropylar region.
The
newly
that
Roth (1957),
figures
have
a
does
in the
primordium
cropylar part (A 4
nucellus
larger,
the
of
as
a
in fig.
cup and
4a is
b\
must
to
in this
clinally.
tends
is
9a).
to
2
infig.
stages
The cell
b\
of
an
The basal
The
as
is
apical
develop-
and
a
mi-
encloses the
the nucellus grows
in connection with
position
histogenesis
offig. 4d has
4d)
portion
narrower
fig. 2c).
of the
this stage of
cells b in fig.
2d,
become
in
means
struc-
growth
is, properly speaking,
at
girth appreciably
increase in
region
3
distinguished.
endostomium (A
4d &
produce
figure
towards the
be
can
reflected in the later
region (compare fig.
divisions
portion (A
the
great share in the formation of the
formed cell wall in this cell
basal
2d)
whereas the distal
formation of the
fig.
a
by
two-layered
cases
not occur.
anticlinal again, and this is rather characteristic. Already
ment
a
divide. In all these
a to
meristematic. Growth
remaining
Capsella by
reconstruct
easily
one can
by the failure of cell
develops
ture
of the cells b
1
of the
undergone
micropylar
several mitotic
the cell pattern shown in fig. 9a. The situation of the cells
typical. Fig.
formation of the
9b
shows
that the
inner cell
The cellular pattern of the closed
micropyle
the
layers
contribute
i.i.
by dividing peri-
is rather
irregular (fig. 9c)
micropylar portion of
;
624
Fig.
R.
9. Successive
developmental stages of
the various cell families
are
not
sion of the basal
of the i.i.
part
its
growth proceeds
at a
faster
the
micropylar part
of the
clearly distinguishable.
comes
rate
about
than the
The
the
rapid
outer
of the i.i.
ed. The other cell
layers
change appreciably (Jig.
Also
during
passively,
grow
The
longitudinal
nucellus,
changes
exten-
Sometimes
that
so
a
after fertilisa-
so
along actively
the final stages of seed
formation of the
during
seed
setting
seems to
stretch
that its cells become extended and flattenand the
of the cells does
shape
not
10).
ered. No vascular strands
the
integument.
of the
growth
F. BOUMAN
place.
increase in volume of the ovule
layer
BOER AND
by intercalary growth.
small gap appears between the i.i. and the nucellus. This
tion has taken
DE
develop.
testa;
it
setting
The i.i.
“persists
Fig.
seed,
as
10.
n
the i.i.
does
a
not
papery
Longitudinal
=
usually
nucellus.
remains
three-lay-
contribute substantially
layer
”
section
(Bhandari
of
the i.i.
of
to
1967).
a
young
625
INTEGUMENTARY STUDIES IN THE POLYCARPICAE II
3.3.
The
The o.i.
integument
outer
originates
the
on
convex
side of the ovule
its initiation the cells of the dermal layer
subdermal tissue below these cells forms
viewed under
ment
as
a
dissecting microscope,
ring-shaped
a
wall
interrupted
bulge
are
initiated
o.i.
are
on
visible in
process of
lus
as
parts,
a
a
rows
growth
as
a
by periclinal
concave
wall
horseshoe-shaped
ring-shaped
side
of
The
divisions. When
where
clear if these
quite
not
place
on
developseparate
a
two
parts, i.e.,
the dorsal
meristem. Both parts
side,
of the young
median section of the ovule (see figs. 2, 3 and 4). During the
of the ovule the dorsal part of the o.i.
bending
cap. The
or
histogenesis proceeds qualitatively
viz., by periclinal
parallel
cell
the inner side and the
separately
At the
o.i. appears in this
stage of
at the
outward bulge is formed (see fig. 11). It is
the
bulge
a
the
primordium.
dividing anticlinally (Jig. 7a).
start
of cells
occurs,
divisions
(fig. 12).
all cells
in the
Also
subdermal cell
during
later
overgrows the nucel-
in the
same
way
in the
which
layers
developmental stages
remaining meristematic.
In
fig.
13
produce
apical
no
shown
are
two
two
cells in the apical part of the growing o.i. dividing almost simultaneously.
In the
drawings showing
appreciable elongation
the
ed. The anticlinal cell divisions
frequently
than the
funicular side
as
a
periclinal
(fig. 12).
result of this
general aspects
of the
growing ovule (fig. 2)
of the funiculus above the horizontal line
responsible
for this
divisions of
the
growth
in
can
length
occur more
initiation
integument
The fold between the o.i. and the i.i. thus becomes
growth
process. The anticlinal mitotic
Fig.
11.
The
o.i.
forms
inner
a
Youngjovulejviewed
from
ring
bulge
develop
on the
as
a
the
wall
funicular side,
n
the
on
deeper
is stronger
activity
does not
an
be observ-
side.
micropylar
but as
=
a
cap
nucellus,
on
and
i.i.
=
integument.
Fig.
12.
of o.i.
Longitudinalsection
at the
funicular
side
(the “bulge”).
Fig.
13. Primordium
subdermal
layer.
of the o.i.
showing
mitotic
activity
in the
626
R.
the dorsal side and
remains
rate
growth
higher
is
gradually bend
so
that
the ovule
pace when
keep
layer
of
and
remains
endostome has been attained and it remains
The exostome proper
is formed
the circular
lying
the i.i. The
later
The
structure
raphal
not
of
into
development
two
at
Bhandari
bundle first
bulge
After
off
a
branch
innervating
fertilization has taken
o.i.
The
cells of the
the
o.i.
layer
a
cell divisions
time
same
superimposed
rate
(see
4.
growth
6).
s
The tissue
in fig.
we
have
over-
much
sec-
beyond
has been
micropyle
kind of
which is
Several
proper.
small
occur
of the
which
ovule
to
proceeds
pit-like perforation
ingoing
(d
occur
in
in
form
the
by
developed
hard
much faster
at a
of the sclerotesta
vascular bundle
is formed
fig. 6),
columns of
produce
sclerotic
direction
strongly
offshoots
changes
(figs.
the second cell
tissues of o.i. and funicle grow
more
o.i.
more
o.i. become stretched and
section
heteropyle
bifurcate
the chalazal side.
on
of the
which surrounds the
the
to
on
5
layer
form
out to
the chalazal
side,
(fig. 2, fig. 5).
seen
DISCUSSION
The
occurrence
of
to
has
suggested
tive
(Takhtajan
of
a
parenchymatic
sarcotesta
layer
longitudinal
protrudes
hypostase
becomes
layer
longitudinal
that
surrounding
The
6).
transverse
This
the
so
(“heteropyle”)
the massive
as
cells.
in
around the chalaza,
is formed
and
periclinal
cylindric
sclerotesta. The
partly
exostome
in the
considerable structural
place
inner dermal
but
the
o.i.
the
to
appear very much extended in
at
side.
dermal
“incipient obturator”. In these
an
penetrate into the massive parenchymatic tissue
the
antiraphe
the
at
the funicular side of the
(1964)
micropyle.
vascular bundles become differentiatedin the
splits
strands
form the
the funicular side. This differen-
T-shaped
that the
as
the
rates
developing nucellus,
cell divisions in the
a
region
cell thick until the level of the
one-layered
on
produces
exactly median)
thus formed
Kapil &
by
stages
distally
(if
BOUMAN
position.
endostome which suggests
elliptic
to
tions of the ovule
described
of the o.i.
development
F.
unequal growth
ultimately
one
by periclinal
of the lateral sides of the o.i. and of the
tial apical
The
overgrow the
they
juxtaposition
thedevelopingo.i.
AND
of the ovule. The dorsal
part of the o.i.
into its final anatropous
over
BOER
nucellus because also in this
the dorsal side of the ovule.
their rims remain in
The dermal
curvature
the i.i. and the
at
and the i.i.
The o.i.
the
causes
pressed against
DE
Magnolia
Capsella
many workers that
(both
a
subdermally
integuments
and in Lactoris
more
advanced
criteria.
of dermal
origin,
growth
a
2-
o.i.) clearly
origin,
the
two
4-layered
most
cell
i.i. of dermal
layers thick),
initiation,
integuments,
to
primi-
integuments
differs from the condition in
of dermal
appear
of the
to
of the
in Lilium
becoming “secondarily”
o.i.
from the number of
histogenesis
The
(viz.
Angiosperms
condition is the
bitegmic
(both integuments
initiation and the
phylogenetic
initiated
dermal
of
arily” three-layered). Apart
ument
the
1959, 1969; Eames 1961). The histogenesis
integuments
layered),
among the
single integument
described in the present paper
derivation and
(both
a
provide
integuments
the
multi-
i.i. “secon-
the mode of
integ-
useful taxonomic and
in
Capsella
is
brought
INTEGUMENTARY
about
STUDIES IN
cell
by apical
growth of
the
cerning
the
and
outer
all
is it
a
According
in the
in
reptilian,
assumes
this connection
these
of
cases
links
with
specialized palissade
cording
are too
in
in
a
the
to
be of
present
confident
to
a
“
is
is
the
of the
i.i.,
Endozoochory, possibly
course
one
the
opinion
Degeneriaceae,
occurs
taxa).
not
sense
rather
testa
examples
seed
of
seems
hard
a
with
Ac-
.
and the “sarcotesta”
sarcotesta
a
testa
and Van der Fiji’s ideas
the
of
Legu-
provided
improbable"
not
is
same
struc-
necessarily
integument development,
semophyletic significance
greater
and
diversity
morphology
coat
emerging
Magnolia-
are
(the
and
all
Paeoniaceae,
necessarily always
The differences in
suggestive
to:
Sapindaceae,
reversal
A
the
explain
roots
Dilleniaceae,
Meliaceae,
morphological
generalizations concerning
clearly
assuming genetical
common
in, &.g.,Welwitschia,
families is
and
noteworthy
could also
convergence, without
parenchymatous
some
It is
(1969) opinion concerning
Pul’s
Of
of seed coat ( testa)
dispersal.
(priv. comm.), Takhtajan’s
these
least
authors,
and
or
Angiosperms ”,
and seed
Euphorbiaceae,
divergent
at
o.i.,
treated in this chapter belong
strictly phytomorphological
considered
by
cases
anemochory
homologous throughout
con-
homologous
equivalent,
of the
follows:
(Lepidocaryoideae).
tissue
seventeen
integuments
Violaceae and Bixaceae. Further
J. Meeuse
one-sided:
reported
ture
A. D.
to
der
adaptive
as
,
and Palmae
that the
strongly developed peripheral
the earliest
structure
Winteraceae,
minosae, Sterculiaceae,
by Liliaceae,
of
as
“
Flacourtiaceae
Berberidaceae,
with its
sarcotesta;
a
clear. The
are
inner
implies
in discussions
account
homologue
However, in my
Gymnosperms.
all
whereas
instance,
This
“the basic primitive type
mention Van
to
Annonaceae,
ceae,
(1969)
characteristic
a sarcotesta
basic stock
from
the
taxa
cells". He continues
seeds with
angiospermous
are
Magnolia-type
the
relation between seed
a
for
the two?
Takhtajan
probably
was
be taken into
phytomorphologically
unitegmic
Angiosperms (is)
marginal meristem,
a
Magnolia is unlocalized.
must
also
product of
to
in
627
II
integuments:
integuments
layer of parenchyma
in
of
homology
fusion
by
or
integuments
solitary integument
or
growth
integuments
the
of the
histogenesis
POLYCARPICAE
THE
to
are
over-
be received
with diffidence.
The
coat
almost
needs
same
to a
made distinction between
discussion. In
organ, the younger stage
functional and
some
universally
some
ecological
protective function,
an
sense
it) often
pollen tube.
has
provisions
some
are
maturation,
are
as
After seed
show
a
It is
is
seed
for the
one testa.
however.
integument
the seed coat
In
a
Both
have
far
equal
by
not
integument being
setting, however,
made. The often very
profound changes
described here for Magnolia,
to
two names
the conduc-
(or
a
part of
function in seed dispersal (e.g., in Magnolia ) and the necessary
number of basically
cordingly.
are
the older
different,
very
and
integument(s)
these
integument,
it is true, but the soft
function. However, in different
a
called
they
sclerotesta in this respect, the function of the
tion of the
in
ontogenetic
being
sense
be
angiospermous
are
taxa
seed
different ways, and the testa
expected
greater uniformity in
that seeds with the
testa
development,
in the
dispersal
the
its
ultimate
is effectuated
development
same
during
testa
connected with
dispersal
differs
ac-
mechanism
whereas seeds with
a
different
628
R. DE
exhibit differences
ecology
development
differ
from
development
ations
based
the
regards
uniform type of
ble
exception
still
far
as
are
similar
as
the ovular
to
Kapil & Jalan
obviously
coat
of Schisandra
ovule
histogeneGeneraliz-
1964).
misleading,
be
can
certainly
as
of the ovular organs.
embryological
relating
papers
of detailed
means
to
within this
of Michelia. The available data are
by
but
instance,
very
1963;
integument development
verification
require
are
Hayashi
early development
Text and illustration of
to a
testa
illustrations the mode of initiation and
structure
the ultimate
on
for
Magnolia,
BOUMAN
F.
advanced stages of seed
related. The seed and
the stage of anthesis
to
Magnolia (Yoshida 1962;
the
during
closely
and
descriptions
up
especially
taxa are
from those of
appreciably
gleaned
sis in
ifthe
even
BOER AND
Magnoliaceae point
family
inadequate,
histogenetic
with the
possi-
and
however,
they
studies.
ACKNOWLEDGEMENT
The
assistance
for
sponsible
wish to
and critical
the
express
our
advice
received from
translation
English
thanks to Mr. J.
of the
Vuijk
Professor
manuscript,
for his
in
help
is
A. D.
J.
Meeuse,
who
is
gratefully acknowledged.
making the
illustrations
also
re-
We also
ready
for the
press.
REFERENCES
N.
N.
Bhandari,
(1967): Magnoliaceae, in:
Seminar
on
comparative embryology of Angio-
New Delhi.
sperms.
Boesewinkel,
D. &
F.
F. Bouman
(1967): Integument initiation
in
Juglans
and
Pterocarya.
ActaBot. Neerl. 16:86-101.
F.
Bouman,
(1971a);
problems.
—
The
Ber. Deal.
application
Bot. Ges.
(1971b): Integumentary
of
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studies
to taxonomic
and
phylogenetic
84:169-177.
studies
in the
Polycarpicae
I. Lactoridaceae.
Acta Bot. Neerl.
20:
565-569.
Davis,
G. L.
Eames,
A. J.
Gray,
A.
Soc. London
Y.
Hayashi,
female
Sci.
—
A short
(1858):
J. Linn.
(1963);
Tdhoku
exposition
of the
The
embryology
Univ.
(4)
of
female
Tdhoku Univ.
fuscata. Sci.
(1969): Die
Rep.
R. N. & N. N.
the
of
family
Magnoliaceae.
Schisandra
répandra
gametophyte and embryogeny
Samenmerkmale
Bot. Staatssamml. Miinchen
Kapil,
and seed-coats
of
Magnolia.
2.
and
Megasporogenesis,
Kadsura
japonica.
Biol. 29:403-411.
Michelia
H.
of the ovule
structure
embryogeny
and
(1964); Megasporogenesis,
Huber,
New York.
Bot. 2:106-110.
gametophyte
Rep.
New York.
(1966): Systematic embryology ofthe Angiosperms.
(1961): Morphology ofthe Angiosperms.
(4)
und
Biol.
of
Magnolia
liliiflora
and
30:89-98.
Verwandtschaftsverhältnisse
der Liliifloren.
Mitt.
8:219-538.
Bhandari, (1964); Morphology
and
embryology
of
Magnolia
Dill,
ex
Linn. Proc. Nat. Inst. Sci. India 30B: 245-262.
—,
& S.
Jalan
(1964):
Schisandra
Michaux:
its
embryology
and
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Bot.
Notis. 117: 285-306.
Leinfellner,
Michelia
Maneval,
W. E.
sion of the
Pul,
W.
L. van
(1967): Über
die
(Magnoliaceae). Osterr.
(1914):
The
primitiveness
der
Karpelle
development
of the
verschiedener
Magnoliales.
IV
Magnolia
und
Bot. Z. 114:73-83.
of
Magnolia and Liriodendron, including a
Magnoliaceae.Bot.
Gaz. 57:
(1969): Principlesof dispersal in higherplants.
1-31.
Berlin.
discus-
INTEGUMENTARY STUDIES
Roth,
I.
(1957):
Die
IN
THE
POLYCARPICAE
Takhtajan, A. (1959): Die
—
der
Histogenèse
phologische Bedeutung. Flora
629
[1
Integumente
von
Capselia bursa-pastoris
und
ihre mor-
145:212-235.
Evolution
der
Angiospermen.
Jena.
(1969): Floweringplants, origin and dispersal. Edinburgh.
Warming, E. (1913): Observations
tus
sur
la valeur
systématique
de l’ovule.
Mindeskriftfor Jape-
Steenslrup, Kjobenhavn 24:1-45.
Yoshda,
Sci.,
O.
(1962): Embryologische Studien
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Univ.,
Nat. Sci. Ser. 3:459-462.
über Schisandra
chinensis
Bailey.
J. Coll.
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