contribution
A
mental
towards
gynoecium
Engelhardia
develop-
the
morphology
spicata
Lechen.
of
Blume
ex
(Juglandaceae)
H. Verhoog
de
Hugo
of Amsterdam
Vries-Laboratorium, University
SUMMARY
interpretative gynoecial morphology of
The
their
pistillar
structures
Lechen.
Engelhardia spicata
and is
in
structure
of the
derived
mainly
of cells
the
lower
of E.
ex
the
Bl. shows
of the
not be
A
that the
ovarial
cells
study
of
and the
the
intralocular
of the
formed
is
parenchyma
the
pistil
of dual
cells
superficial
means
to
of
origin
thin
remainder, a
the
by
in the
and the vascularisation
anatomy
satisfactorilyexplained by
terminology applied
histogenesis
pistil wall,
cavity, being
the text. The
“septum-2” in
as
spicata can
Juglandaceae
discussed.
critically
from the inner dermal
portion
to
referred
pistil
are
layer
of the
pattern
of the conventional
foliar
carpel theory.
1.
INTRODUCTION
Publications
tributions
and
and
amara
&
genera
by
(1932:
of the
the
employed
older
so
that,
to
He
also the first
was
greater
spicata
start
all
detail,
nous
a
on
over
Braun
(1909:
the
(1872:
con-
Juglans
Juglans regia);
”
=
pecan
Carya
Wood-
olivaeformis );
mandchurica and
(1939: Juglans
by Hjelmqvist (1948:
attempted
is
one
30-63)
p.
and
by
comprehensive theoretical explana-
a
observational data.
and
again
to
resulting
re-examine
rapide
1’etude
inconsistencies
semantic
as
of
examen
anatomique
de
fleurs
cette
d’un interet fondamental”. However,
(1964)
pointed
from
out
the
tried
a
studying
the
as
ovarial
possible phylogenetic
Leroy
possible.
Engelhardia
des
in the
confusion,
cases
many
gynoecial anatomy
“un
que
confronted with
of the
discuss the
to
nous
often
account
Meeuse & Houthuesen
spicata
de
Eng.
ne
espece
this did
not
of
structure
importance
of
structure.
Both
Leroy
morphology
foliar
the
and
concluding:
Engelhardia spicata. They
this
Hicoria
Langdon
pecan”);
convaincu
serait pas pour
keep
“
A.
deal
usually
ones are
Juglans, Pterocarya fraxinifolia, Carya
of
Benson & Welsford
1927:
Juglandaceae
important
Juglans regia);
previously accumulated,
literature
decided
in
of
most
The latter author reviewed the various genera, gave definitions
he
terminology,
(1869:
(1926,
Hicoria
tion of his and of the
In
The
Carya.
and the extensive surveys
(1955).
terms
“
and
(1902: Species
tomentosa);
C.
Carya glabra),
Leroy
Tieghem
van
Woodroof
Shuhart
gynoecial morphology
Juglans
Karsten
Carya);
roof
the
treating
with the
only
and
of the
Meeuse
Juglandaceae
carpel theory.
Croizat
carpel theory altogether,
points,
the
latter
Acta Hot. Need.
c.s.
especially
17(2), April
1968
by querying
(1966)
whilst
on
different
even
approach
gynoecial
of the
considerably
farther and
rejects
goes
of
the
applicability
criticising Leroy
account
to
the universal
their
and Meeuse
putative
c.s.
in several
interpretations
of
137
H. VERHOOG
the
gynoecial
it
only
desirable
thought
was
of
morphology
and Houthuesen had
a
Engelhardia
species
re-investigate
taking
to
E.
spicata,
general
structure, the vascularisation pattern,
and the
development
2.
MATERIAL AND
The available
ted
by
of the intralocular
material,
in FPA. It consisted of
relatively large
a
the initial
male and
in the
of
an
A
some
material,
no
wax
technique,
stigmatic
lobes
in
sectioning
and
were
seventeen
flowers
in
the
“septum”
Only
were
a
much
and fixed
were
development
not
a com-
immature
more
development, apart
female flowers
collec-
was
number of female flowers with
of floral
were
used for
plane
transverse
were
India,
Oktober, 1966,
few flowers in
a
of
Survey
Shillong
from numer-
used in the inves-
being represented
the presence
regarding
integument (= tubillus).
microtome-sectioned
(i.e., transversely
for
account
by
of the
means
standard
the average thickness of the serial sections made
the median
six
Botanical
at
integument
ones.
about 10 micra, and the slides
of the older flowers
Meeuse
disposal,
pistil.
of the
conclusive evidence could be obtained
number of flowers
lobes,
of
phases
bisexual
tubular extension of the
apical
paraffin
and
sac
The intermediate stages
tigation.
into
the orientation of the
parenchyma
received from the
pletely differentiated embryo
ous
their
at
METHOD
H. Deka in the Botanical Garden
stage showing
that
Considering
spicata.
few female flowers of this
longitudinal
sections
in
diagram),
the floral
perpendicular
sections.
used for
being
stained with Saffranin-Fast Green. Nineteen
Of
the
to
the
longitudinal
for
seven
six for
of the
plane
longitudinal
direction of the
young
and
the
through
stigmatic
developmental
transversely
stages,
serial
cut
sections.
In
addition,
pattern
of
1958) proved
pistils
the
larger
female flowers
vascularisation.
to
be inferior
The
to
were
the
of Nelumbo. About ten flowers
cleared in
method used
were
treated
Fig.
order
recommended
procedure
1.
by
van
Leeuwen
according
Various
to
(1948,
(1963)
for
the lattermethod
interpretations
morphology
the
study
to
Sporne
by
of
the
floral
of
Engelhardia spicata
A.
EICHLER’s
B.
NAGEL’s
interpretation
C.
LEROY’s
diagram
D.
A combination
of
“perianth”,
stigma,
p:
(“midrib
138
Ada
of
floral
si:
carpel”),
Bot. Neerl.
diagram
of the
(B)
sp:
pistil
and
(C)
su:
suture
septum.
17(2), April
1968
DEVELOPMENTAL GYNOECIUM
MORPHOLOGY
but the time of
hydrolysis
duration of the
bleaching
four
days
way
were
in
OF ENGELHARDIA SPICATA
HC1
N/l
examined
under
incident illumination
at
binocular
a
dissecting
stained with saffranin and differentiated again
of
intensity
strands
layer
this
layer
outer
envelope
Eichler’s
and
b
as
the
names
lateral
most
off under the
diagram
is
and
position
carpels
-
(1914)
as
terms
intended
genetic
the
carpel,
and also the
in the
beween the
ditional
et
by
the
sense.
two
It
in
a
of
of
a
for
to
or
In my
not
(N.B.
Acta Bot. Neerl.
:
a
of
17(2), April
1968
1
a'ß
given
carpels
mor-
here,
tends
the
much
detail,
a
one
or
etc.
can not
a
semantically
Juglandales
gymnosperms,
the
to
avoid
to
the
traas
a
The “5-lobed
possible
“cupule” by
different
one
reasons.
“perianth”
chlamys).
among the
distinguish
to
the
and refer
to
and
accept the
for semantic
possible
and
sense
possible phylo-
in the conventional
consider
but in order
“Cupuliferae”
dissipiment
typological
of any
cycadopsid
as
plane
which lie in the
importance
to
integument
(median)
the
carpels
or
cupule (or gnetalean
presumably
the
followed
are
perianth
of essential
one
as
outer
discussed in
this is
cupule
two
discussion of the
opinion,
and of
advanced
principle
pteridospermous
has
of symmetry.
septal plane
carpels
If
as
the bifid lobe
also lies the
purely descriptive
be
wall)
envelope
prophylls
two
(b).
two
plane
it should be mentioned that this bract is called
so-called
a
plane being
of the
edges
to
this
(ovary
bract” is
Shuhart
not
Leroy
“flowers”, “perianth”, “carpels”,
adhere
“flowers”
outer
and
which shows
“bract”
in which
meanings.
appears
different
terms as
al.
homologue
fusion,
Juglandaceae,
carpels
persistent
used
the
(b)
topographical indications, irrespective
primitive Angiosperms
be very
avoids such
Meeuse
here
of
sutural
principal
the coalesced lateral
are
as
morphological
to
that
being:
morphogenetic interpretation.
or
presence,
so
without
A combination of the last
stigmas.
terminology
that it is sometimes called the
so
These
wall
increased.
diagram (fig. IB). Leroy
starting point
a
I A:
but
(see fig. 1C)
commissural
is used
fig.
of the floral bract
of the “fruit”
intercarpellary (i.e., transverse) plane,
only
observation,
red
as
outer
outer structure
stigmatic lobes,
in Nagel’s
structure
in
reproduced
shows cleft
midrib of each
-
both
Commissure
septum
bundles
The
dissecting microscope
of this
knowledge
the
important circumscriptions
Suture
vascular
were
appropiate
called the female juglandaceous
usually
as
The definitions and the
bisecting
An
hypochlorite.
cells which obscure the
concrescence
represented
median
phology.
oblique
STRUCTURE
of what is
diagrams (fig. ID)
two
in
transparent background.
scraped
interpretation
a
of the
diagram
in
as
The
ß).
opposite
a
(1878)
interpreted
(a
at most
with
microscope
of the
recognition
more
crystal
GYNOECIAL
has received various
is
to
the vascular strands.
GENERAL
The
be
carefully
must
the
and
paler
a
of the flower contains
damaging
3.
enables
staining
against
reduced
The flowers treated in this
of about 45 °C. Some bleached flowers
angle
an
down to 10 min and the
cut
was
hypochlorite (1:3)
of the flowers.
prevent desintegration
to
60 °C
at
in commeicial
con-
Benson and
“cupule”,
Amentiferae),
viz.,
and
a
139
H.
involucre”
“cupular
the
views
personal
employ,
the
be
to
A floral
The
descriptive
of
diagram (fig.
terms
terminology
theory,
for
they
unless
“flowers”, “perianth”
as
definite choice
a
that
in favour
and
matter)
and
suggests
of
simplified representation
a
is
Engelhardia spicata
ID)
of such
use
and
background
of
only
are
purposes.
diagram being only
morphology
determine the
that I have made
any other
carpel theory (or
intended for
workers
mentioning morphological interpretations
imply
not
the theoretical
Manifestly,
various
elucidating.
does
“carpels”
foliar
the
that I have shied of
so
they proved
and
by Langdon.
of
VERHOOG
a
considerably
detailed
more
flower, the floral
a
than the
complicated
more
description
of
parts is
some
indicated.
The septum:
which
ced,
in
2
results
number
a
The classical
-
in
dissenting
blätter
ausgehende
oder
im
stets
durch
Accordingly,
Langdon
who
do
of the
ovule
called
as
subsequent
is
a
turned
margins”
Meeuse and
of
1895;
cium
be
to
an
of the
and of
Houthuesen call
the
ovary
which
has
“false”
a
originated
juglandaceous
time
indicating
its
ovarial
plane
of
auf dessen
as
de
as
primary septum,
a
Candolle,
Shuhart) is
of the
even
1862; Karsten,
that the
opinion
apex and that the
carpels
that the
septum is
of
speaks
not
out-
an
should
consequently
not
be
and “in-
“carpels”
“primary” septum.
a
septum “false”,
integuments
also
they
the
because
and
“false
term
from the
dorsal
My
a
had
of
conclusion is that
we
a
better
wall
in
should refer
septum”,
the
simply
orientation,
and thus evade all theoretical
at
an
and
carpel
dissipiment
as
be of
gynoe-
perhaps
dividing
region
“the
interpre-
possibly
juglandaceous
septum”
(median)
in their
can not
consider the
septum is traditionally
develops centripetally (fide Eichler).
the
it
to
and
carpels
two
ecarpellate structure,
because
dropped
and
implies
structure
derivation. However,
be
sich
formation of the ovule. This interpretation
tation, it divides the space between
carpellary
der
Begriff
vereinigen
Mittelsäulchen,
authors refer
the
margins
carpels,
sind
der Frucht-
den
diese
also
variabelerHöhe und bilden
zu
Art
septum. Nevertheless Langdon
primary
and
regelmässig
unter
fallen;
derivative of the floral
a
“terminal”
a
of the fused
growth
somit
accordance with the views of C.
Juglandaceae
“Ganz
for-
decidedly
workers
ist”.
befestigt
“contribute” towards
not
stated:
die
eine
Nicoloff, 1904-1905; Nawashin,
ovule of the
straightforward
a
Verwachsungslinien
Teile der Frucht bis
most
(in
provide
thus becomes
among the older
Scheidewände
zusammenstossen
der Same
Gipfel
den
vorhanden,
primären
unteren
das
Eichler
von
not
explanation
opinions
of inconsistencies.
Scheidewandbildungen
ächten
does
carpel theory
of the septum and the
interpretation
to
same
implica-
tions.
As regards the
primaire
in
his
in the
fig.
Psilocarpae
septum-like
140
72
situation
in
Engelhardia
intracarpellary (i.e..
a
and
transverse
draws
outgrowth
an
section
the
of
Leroy
transverse) plane.
a
excroissance
perpendicular
spicata,
to
fruit of
septale
the
a
In
species
tabulaire
plane
of the
speaks
of
addition,
of
a
he
cloison
depicts
Engelhardia
supposed
to
sect.
be
a
“primary” septum.
Acta Bot. Need.
17(2), April
1968
DEVELOPMENTAL
Fig.
2.
OYNOECIUM MORPHOLOGY
Diagrammatic threedimensional
and
of its
accrescence
OF ENGELHARDIA SPICATA
representation
(“septum
2”;
(right) semidiagrammaticcross-sections
pt:
packing
tissue
vb:
ventral
bundle,
ib:
stigmatic lobe,
(The
same
cw:
“carpel” wall,
are
used
From my observations of the
in the
made
are
is also
the septum
appears
fig.
more
reconstruction
drawing
or
other
Shillong
lobe
of
septum,
to
the
ovarial
material I
depicted
by Leroy.
17(2), April
1968
and
perspective.
i:
cavity,
integument,
b:
lobe
of
bract,
si:
suture.
finally managed
actually
out
broader than it
in the
This
outgrowth
In my fig.
3
and
is
one
not
can
to reconstruct
diagrammatic
transverse
lie
much closer
A
together,
so
(so
longitudinal
with
comparable
strongly
clearly
see
three-
sections used
other directions
actually is).
of the septum
3. The
Engelhardia
Leroy)
bundle, sb: septal bundle,
“perianth”,
su:
less distorted because the
plane
fig.
in
of
of
figures)
drawn apart and
relatively
perpendicular
Acta Bot. Neerl.
the
p:
somewhat drawn
72 A is shown in my
in the fruit
in
oc:
db: dorsal
septum
tabulaire
drawn
levels,
strand,
sp:
pistillar
septale
septum-like outgrowth (see fig. 2).
dimensional figure is
and the
v.s; vascular
integument bundle,
symbols
the septum and the
at four
(intralocular parenchyma),
nucellus, sp-2: septum-2,
n:
of the
excroissance
that
section
Leroy’s
developed
that the
as
“out-
141
H.
Fig.
3.
Part
of
exactly
locular
growth”
142
longitudinal
section
possible through
parenchyma (pt)
the
“winged
of
a
were
evaginations”
packing tissue (in
the
“female
septal plane.
which is indicated
consists of cells which
resemble the
called
a
as
sense
flower”
Cellular
by
a
of Benson and
partly
shade of
more
(Nawaschin),
Engelhardia spicata,
structure
darker
formed in
of
VERHOOG
or
cut
in
as
intra-
grey.
less distinct
“horns”
others)
shown
rows
(Nicoloff)
in the
pistil
Acta Bot. Neerl.
of
or
and
so-
Juglans.
17(2), April
1968
DEVELOPMENTAL
GYNOECIUM
MORPHOLOGY
However, these outgrowths
than
spicata
by
they
less
equivalent
tissue
what
to
de
it parenchyme
tissue”
“packing
idea of the
what
oblique
believes
structure
with the
tissues and it
bicarpellate
the
that
represents
the
they
is
der Same
in
called
of these
of the
it is
As
“carpels”.
If
problem.
the
to
one
ist.
which
the
the
(if
septum, and
median but
oriented and the
transversely
ral. This would
only
certain
theory requires
does
unequivocally.
add
As
intralocular
not
a
to
the
of
explain
parenchyma.
of the
the
the
When the
intralocular
endocarp”
(as
17(2), April
1968
by
that,
of the
pistil
be
to
seem
“septa”
are
conspicuous.
provide
the
at
interpreted
plane,
the
septa,
as
it
appears
con-
there is
in
Gipfel
derivative of
not a
in order
two
as
that
dessen
auf
a
septal
become
to
to
save
be
to
carpels
point
are
morphology
the
the
not
of the
the
carpel
obser-
Juglandaceae
it is untenable.
structures
were
Engelhardia
Shuhart and
of fact the
variance with
floral theory
of
satis-
a
centripetal
carinal in stead of commissu-
are
septal
of
“septum-2”
however. In
parenchyma
defined
corresponds
primary septum,
a
ovule is
so
consider
to
gynoecial
is
ovary
transverse
Mittelsdulchen
which
general interpretation
not
not
Engelhardia,
stigmas
confusion,
interpretations
the
suppose that
to
it
because
gynoecial
derivative of the
a
only ultimately
thought,
would have
one
so
the
orientation of the
that forms the
In this train of
more
question
definition of
structures
of
endocarp”
portion
does
no
in the
within,
regards
applies
septal
“septum-2”
from
with
tissue” is
differential growth of the various
a
There is
present
primary (= carpellary)
Acta Bot. Need.
the
and hence
some-
nor
the excroissance
between
which is
carpel theory
carpels
already
are
by
structures.
of
margins
interpretation,
chymatous
being,
call
histologically they
developed
classical
appears that the
befestigt
equivalence
to
a
calls
adequate
was
“packing
Typologically
primary (and assumedly carpellary) dissipiment,
vations and
sectioning
flowers reveals that the
mature
extensively
an
stigmatic plane
case
Juglans,
fused and
clearly accomplished
structure)
another
Pterocarya.
Shuhart,
Leroy
Houthuesen
give
not
“parenchymatous
intermediate
completely
are
but far less
with the
Engelhardia
The
of
plane
2” for the time
an
([sensu Benson)
developing centrifugally
classical
Meeuse and
73),
all the tissue
not
by
ovarial wall.
with the
exactly coinciding
so
tissue”
given by Eichler,
a
it
explanation
“outgrowths”
still
because
An examination of less
outgrowth
crescent
fig.
pistillar cavity
parenchyma being
endocarp”
which the ovule is inserted. In the lowermost
on
already present
tissues
his
e.g.,
been used
“filling tissue”,
a
fill the
not
their figs. 2 and 2a, which do
of Oreomunnea and that of
contiguous.
factory
see,
is
really
of the
Engelhardia
tissue” has
the intralocular
“parenchymatous
Leroy “septum
that
types of septa
two
does
structure
Juglans,
only partly represents
“packing
“pedestal”
Their
plane,
tabulaire of
septale
the
remplissage,
it
Juglans,
centripetal proliferation
(compare
and
In
in
developed
“packing
term
To evade this semantic confusion I propose
Shuhart).
Leroy
called the
a
and neither
kind
one
it does in
as
gynoecial morphology
the commissural
of
is
as
conspiciously
sense.
homologous
completely
originating
different
a
the
Engelhardia
more or
far less
are
in the walnuts. The
are
different authors in
but in
OF ENGELHARDIA SPICATA
others)
discussed,
and
was
the
the
“paren-
mentioned.
143
H.
It
develops
as
a
more
less
or
simultaneous
dermal cells of almost the entire inner
cation of this
in the inner
there is
carpels
outgrowth
epidermal
hardly
and that of the
centripetal
formed
only
chymatic
one
by
rows
of
the
two
formed,
pistil
The fact
septum-2
as
an
wall and
that there is
and
the
of cells which
dermal cells
transverse
the
no
right
region
of the
7 the inner
cell
outer
of
layer
of the
inner
sections
is
from parenwall.
up
produced by
to
the
tissue
thin
a
centripetally
clearly
discern-
the whole inner surface of
of the
apical region
of the inner
This is
2). Only
is
“septum-2”
has
septum-2
carpel
(see fig.
wall has
carpel
indistinguishable
are
demarcation line between
sharp
superficial
basal
inner walls
but the demarcation line between the
parenchyma
extends
in the
Particularly
In fig.
the
division walls
periclinal
structure of the
tissue and the tissue derived from
ible. The intralocular
the
the
of cells whereas
layers
examination of
of tissue is thus
developed filling
“carpels”.
developing septum-2.
cells derived from the
confirmed
layer
layer
of the
of
proliferation
4 and 5 the first indi-
figs.
visible in the form of
clearly
difference between
any
formed
is
centripetal
wall. In
pistil
VERHOOG
the
ovarial
wall could be
pistil
cavity.
of
layers
outer
adduced
argument pleading in favour of the carpellary interpretation of the pistil,
if other considerations did
alternative
chlamydote
interpretations
ovule
Integuments
theory)
not
render this
(such
were
as
unacceptable (see
Leroy’s
“unitary”
and
no
Meeuse’s
available.
Meeuse and Houthuesen believed
tubular connection between
and if
59)
p.
theory
ovule
and
“stylar
to
have demonstrated that
canal” is
Fig.
4.
section
Longitudinal
as
accurately
the
through
a
the
hardia
lobe
septal plane
gynoecium
spicata.
initiation
/:
made
possible
as
of
developmentalstage
young
of
the
extension
apical
an
has
of
of
Engel-
Integument
just begun.
bract,
i:
(young)
integument.
144
Acta Bot. Need.
17(2), Apnl
1968
DEVELOPMENTAL GYNOECIUM MORPHOLOGY
OF
ENOELHARDIA SPICATA
5.
Fig.
Detail
of
first
fig.
wall”
formation
of the
cisely
and
integument
The
tum.
they
made
figures they published
oriented in the median
their evidence is conclusive. The
of the
rim-like apex of the
of
integument
is bilobed
Engelhardia
compare Boesewinkel and Bouman
unfortunately
ment to
it
is
of
takes
were
decide this
no
considerable
place
parenchyma
in
pistils
at
at
and
the
apex
the
of the
ascertain
to
in
(as
1967). Among
if the
were
study
of younger
icises
Meeuse and
not
possibly
pistils
tubular
by
Juglans
the
reason,
conclusions
My
that
an
aside
gives
on
(and
or
to
has
collected,
the
the
my
question
different
fig.
but he did
It
were
Pterocarya,
of
develop-
also because
fertilisation process
and
9
not
figures!)
(which
the
intralocular
can
figures
is
what
17(2), April
Croizat
originate by
Croizat
definitions!),
1968
an
is
not
a
Croizat
disposal.
by
(1966)
our
crit-
examine any material and could
a
and
even
ovule is
of
plane
also
no
by
markedly
question
the
of the
young
altogether
excentric
instead of
of
an
(and
two).
excentrically
peculiar
suggestion
integuments. Leaving
“obturator”
examination of
an
of fig. 2a of Meeuse
symmetry
makes
an
that Croizat based
draws
reproduction
shortening
means
Engelhardia
confirmed
somewhat oblique and their evidence,
show that there is
ovule.
this
is, therefore, deplorable
of their
longitudinal
one
asymmetrical
Acta Bot. Need.
elongation
studied there
proper stage
be
to
how
their
at
“demonstrate” that the
only
“obturator”
two
some
line “m” in his
pistil
slides
inserted
had
they
unequivocal.
not
Houthuesen)
that the
than
Houthuesen,
certain
and
pre-
developing
and
specimens
structure
part of the tubillus;
know that their sections
for that
imaginary
not
all involved.
derivative of the basal
a
and Gne-
Juglandales
Meeuse and Houthuesen assumed that the intralocular tissue of
is
intralocu-
integument).
young
whereas the
integument,
Engelhardia
importance
Engelhardia
is
of
to the
tubillus is formed
More material will have
point.
ultimately leading
direction and it is doubtfulwhether
gnetalean
young inner
the
inner
made from sections which
transverse
or
of
of the bulk of the
between
and
indicate
the
parenchyma (i:
comparison
a
were
ovule
showing
divisions
periclinal
“carpel
lar
4,
parts. The arrows
adjacent
(Jackson
1949,
developmental stages
145
H.
of the ovule
tation,
the
Croizat
makes
which
“and
writes,
exactly
the
deliberate
a
or new
of my
some
of tegumentary
not
or
mistake that he
strobile
the
cavity being
origin.
professes
“Drop
but it
to
not
are
theory”,
every
would
that
seem
condemn. His
based
decapi-
a
parenchyma-
and the “creation” of
(his fig. 5)
of integuments
decapitation
integument initiation, according
this
slides,
in the
begin
seems to
Boesewinkel and Bouman
(see
carya
is
such
undergoes
ever
he
hypothetical
obturator
an
observational data
on
“facts”.
regards
As
integument
investigate purely for facts ”,
same
female inflorencense
by
that the
other tissue present in the ovarial
tissue
filling
tous
showed
never
only
VERHOOG
opment may well be
a
characteristic
to
same
F.
Bouman, who examined
Juglans
in
as
way
and this form of
1967)
feature of all
and Pterodevel-
integument
family
of the
representatives
Juglandaceae.
Vascularization. The
determination of the
ning
a
number of cleared
different
at
I
levels,
cularisation which
struction of the
of the vascular tissues very
must
make
to
come
vascular
and
specimens,
managed
accurate
small size of the flowers renders the
relatively
courses
the
to
by
transverse
reconstruction of the
a
close
pretty
supply
guided
difficult,
the actual
to
nucellus
is
but
sections
made
of
pattern
situation. The
particularly
a
by exami-
vas-
recon-
tedious
job.
-
The
places
of the
that it is
pistil
the
bundles,
pattern is
The
more
not
and
Leroy.
not
give
their
Other workers
the
can
“ventral”
or
used
only
that
“septal”
however.
of vascular
describe
decide what
individual
is
Shuhart
strands but do
difficult
to
assess
the bundles is
a
a
al-
not
certain author
bundles after
by.
taxa
Langdon,
it
region
vascularisation
juglandaceous
figures
to
the
details,
other
course
of the
origin
Woodroof,
accompanying
one
the
concern
&
describe the
terminology
Usually
“dorsal”,
terms
only
trace
impression
only
Woodroof
Without
because the
to
the
studied in
was
in the basal
closely together
so
The differences
Welsford,
unequivocal.
have
I
vascularisation
&
lie
impossible
because
illustrations.
findings
ways
almost
so
constant.
pistillar
Benson
e.g.,
by
where bundles branch off
means
of his
perusal
or
her illustrations.
The
term
“septal
off in the basal
integument.
median
which
itself,
are
as
of
symmetry
approximately
vascular
The bundles
trace
refer
in the
in
the
to
a course
pistil
wall
can not
forming
The bundles
they really
always
be
the vascular
running
are
of rather
septum, but
for
(see fig. 6)
clarity.
In
followed
more or
ones,
not
to
the
less in the
and
to
in the
those
septum
the dorsal bundles
reality they
closely adjacent
individually
supply
run
the “dorsal”
as
of the
the septum
through
the bundles which
plane
consisting
off from the stelar system
rows
along
form
their full
of the 5-lobed bract very
one
of tracheidal
soon
course.
branch
of the floral axis.
towards the
fairly regular phenomenon
146
and follow
“ventral” strands. In the reconstruction
elements which
A
to
drawn farther apart than
complex
be retained for those bundles which branch
can
gynoecial region
It is customary
plane
run
bundles”
is that
“perianth”
two
show
bundles
run
some
interesting
into each of the
Acta Bot.
Neerl.
features.
‘perianth
17(2), April
1968
DEVELOPMENTAL OYNOECIUM
Fig.
6.
Reconstruction
(For
the
of the
meaning
follow
a more
Acta Bot. Neerl.
vascular
of the
lobes” situated in the
in the median
MORPHOLOGY
less
septal
plane
1968
but
Those
independent
17(2), April
system
of the
symbols SI, S2,
(dorsal) plane.
or
OF ENGELHARDIA
only
course
gynoecium
D 1 etc.;
running
SPICATA
one
see
of
Engelhardia spicata.
text)
into each of the
towards the
lobes
dorsal lobes
in respect of the other dorsal
placed
usually
strands,
147
H.
i.e., they branch off"
in the
of
transverse
these
off
they
appear
tried
to
such
to
theii
if connected with
as
they
reported
only
this
is
originally
dorsal
As
vascular
of the
one
have
bundles.
septal
become
already
ventral
In
traces.
the ventrals
be
to
strand.
to
At
the
form
a
1 did
not
young
advanced
ently
sal” is
a
stages
more or
less
derivation.
“innervation”,
qualification,
some
stages
to
Usually
divide
(such
ment
SI
section 8
it is this
septal
cance
as
or
run
there
case
10
is
other
independent
a
can
principal
origin
traces
off from
could
not
are
other bundles which appar-
under discussion, the
(such
as
Dl) being
term
“dor-
situated in
a
a
the
through
trace
is
but he
intricate and
“cycle de
the
base of the
narrow
(fig. 7).
so
that
in
one
of the
integu-
Bundles
a cross-
be counted. The
clear, unless
the
signifi-
interprets
an
S2
bundle which sends off a branch towards the
usually
consists
than the
rather
not
some
septum (S2).
integument,
can
that
continue in the
to
to
goes
branch of a ventral
“dividing” meaning
-
tegumentary bundles
bulkier
branch
a
an
of
single
offshoot
An extensive
a
and
of
group
often
running
to,
parallel
traces
tenuous strand
the
e.g.
S2,
pistil wall,
of thevascular-
comparative analysis
ontogeny of the pistils of the various genera of the Juglandaceae
recognised.
Engelhardia,
148
at
related families will be necessary
be
more
the
SI
However,
considerably
isation and the
and
ones run
principal septal
which suggests that S2
than
branching
turning point
of bundle from which
type
the function of the S2 bundles is
the continuous
which
the mode of
closely together.
too
towards the base of the nucellus
SI) and
as
integument.
an
bundles which
the tegumentary
bundles
but their
bundles
below the nucellus
again
type divide again
to
of
lobes”
intermediate position between the dorsal bundles and of uncertain
vascular bundles
of the
to
impres-
‘perianth
parallel
septum and continues upwards in the septum together with
bundle
managed
I
question
no
the
similar
(Shuhart)
connection with the dorsal and
no
the
off from the dorsals. In the
relative
of
establish
to
manage
“inversely oriented”,
from the ventral
branch
carefully
substantiated,
but I have the
of
pair
development
be located because the ventral bundles run
Apart
level
the
more
inwards that
have very
Juglandaceae.
section,
transverse
a
initiated, but there is
more
seem
In
of other
pistils
less
or
much
are
the
at
having
stigmas.
two
previously mentioned,
of the
S2. I
oriented bundles”
“reversely
bifurcate
usually
traces
continue into
more
bend
strongly
so
bundles
connection and that there is
secondary
a
continuous
in
on
The dorsal bundles
septal
drawn
trace
indeed link up; for, ifthis could be
in the
occur
run
lying
branch off at the level
The ventral traces, after
sometimes
the
the ventral lobes
to
independent
case.
stigmas.
direct connection in
one
sion that
to
going
hand, only
“perianth lobe”,
a
branches
vascular strands would be
find
the
and
establish whether
those
altogether
exceptional
terminate in the
to
ramified
freely
The
an
the other
on
branch which enters
a
direction
same
lobes”.
“perianth
whereas those
level,
septal plane,
left side in fig. 6 is
extreme
given
low
at a
or
VERHOOG
Leroy briefly
gives
not
faisceaux
a
so
before any
clear and
consistent pattern
mentions the vascular pattern in the
rather
simplified description.
constant
as
Leroy
carpellaires-sépalaires”
pistil
of
The situation is much
suggests,
consists
Acta Bot.
e.g.,
by
of 8
Need.
or
stating
that
10 bundles.
17(2), Apnl
1968
DEVELOPMENTAL GYNOECIUM
Fig.
7.
Longitudinalsection
as
of the “female
possible through the
(For
the
This is rather
the various
Acta
MORPHOLOGY OF ENGELHARDIA
meaning of
Bot. Neerl.
the
meaningless
pistillar
median
one
1968
51
and 52:
does
elements. The
17(2), April
of
Engelhardia spicata,
plane (perpendicularly to
symbols
if
flower”
SPICATA
not
see
the
as
accurately
text)
know how the
majority
cut
septal plane).
vascular
traces run
in
of the bundles situated between the
149
H. VERHOOG
median and
the
septal
neither the number
of symmetry
planes
the ultimate
nor
are
offshoots of dorsal
of these branches is
course
and
traces
constant.
ACKNOWLEDGEMENTS
The
of
author
wishes
to thank
Engelhardia, Mr.
for
technical
Prof.
A.
assistance.
D. J.
the Botanical
F. Bouman
The
was
for
the collection
of the material
and Messrs. J. Houthuesen
advice,
version
English
of India
Survey
for his critical
from
adapted
the
and J.
Vuijk
original manuscript by
Meeuse.
REFERENCES
M.
Benson,
&
E.
Juglans regia
Boesewinkel,
C.
18:
& F.
Bouman
of the
morphology
Ann.
genera.
ovule
and female
flower
of
Bot. 23: 625-633.
of
initiation
(1967): Integument
in
Juglans
and
Pterocarya.
86-101.
16;
den inneren Bau
Mémoire
(1862);
de
The
(1909):
allied
der Frücht
la
sur
familie
der
des
Juglandeen.
Juglandées.
Bot. Zeit.
Sei.
Ann.
30: 371-375.
Nat., Bot.,
4me
5-48.
L.
Croizat,
F. D.
of a few
(1872); Über
A.
Candolle,
Sér.,
Welsford
Neerl.
Acta. Bot.
Braun,
J.
and
(1960); Principia
Weldon
Botanica.
&
Wesley Ltd., Codicote, Hitchin, England
(2 Vols.).
—■ (1966): Observations
A.
Eichler,
W.
H.
Hjelmqvist,
Botan.
Jackson,
(1948):
Notis.,
B. D.
Karsten,
Studies
A
(1949):
on
2:
Vol.
Suppl.
Juglandaceae.
the floral
die
32-40.
Southwest.
morphology
and
11
Natural.
72-117.
(1):
Engelmann, Leipzig.
Wilhelm
of the
phylogeny
Amentiferae.
1—171.
glossary of botanic
(1902): Über
G.
the ovary of the
on
(1878); BUithendiagramme II,
Entwicklung
Gerald
terms.
der weiblichen
Duckworth
Blüthen
bei
& Co.
London.
Ltd.,
einigen Juglandaceen.
Flora 90: 316-333.
Kershaw,
E. M.
Langdon,
L. M,
ceae and
Leroy,
J. F.
ceae)
and its
ceae.
L.
—
47
—
(1895);
über
Neues
34
18:
(Paris)
:
The
:
France
—
van
16:
opment
150
Faga-
Gynoeciumof Nelumbo
lutea
Mus. Hist.
of the
Nat.,
N.S.B.
6:1-246.
(Bot.)
Juglandaceae. I.
The
Inflores-
(1964):
die
The
Acta Bot.
Familie
der
J.
Bot. 27: 839-852.
gynoecium
der
Neerl.
Engelhardia spicata (Juglanda352-366.
Juglandaceen.
Chalazogamie.
Sur le type floral
of
13;
et le
développement du
19;
134-152, 380-385;
Jahrb.
Bot.
Bot. Centralbl.
50:
459-530.
353-357.
fruit des
Juglanda-
69-84.
63-68,
morphological differentiation
63:
of the
pistillate flowers
of the pecan.
687-696.
(1948): A
Some
P.
Woodroof,
J.
23: 336-337.
and fruit of
note
on a
of the fruit of Hicoria pecan. Bot. Gaz. 93:1-20.
rapid clearing technique
of wide
application.
New
Phytol.
290-291.
(1958);
&
of the
of the flowers
and fruit. Amer.
Beispiel
(1904-1905):
(1927):
Res.
K. R.
Tieghem,
-—
V.
flowers
Houthuesen.
Studien
of Bot.
407-419.
25:
pistillate
Ann.
of the flower
84-97.
morphology
(1932): Morphology and anatomy
Sporne,
structure
of the
12:
phylogenetic significance.
M. Th.
Agric.
the Julianaceae.
anatomical studies
101: 301-327.
study
Neerl.
The
Bot.
Journ. Bot.
Shuhart, D.
J.
J.
(1914):
Nawaschin,
Nicoloff,
Bot.
Gaz.
A
(1963):
(1938):
A. D. J. & J.
K.
Nagel,
van
Acta.
II. The
(1940):
Meeuse,
E.
Amer.
cence.
relationship of
(1955); Étude surles Juglandaceae. Mém.
Manning, W.
—
A.
Pers.
(Willd.)
the
on
(1939): Ontogenetic and
Juglandaceae. Bot.
W.
Leeuwen,
Note
(1909);
aspects
(1869):
of floral
de
la
systems.
fleur
Proc. Linn.
femelle
et
Soc. London
du fruit
169: 75-84.
du noyer.
Bull.
Soc.
Bot.
412-419.
J. G. & N. C.
of the flower in
(1927):
Agric.
vascular
Anatomie
The
Res.
34:
Woodroof
(1926):
the Hicoria pecan.
development
Fruitbud
J.
of the pecan
Agr.
nut
differentiation
and
subsequent
devel-
Res. 33: 677-685.
(Hicoria pecan)
from flower
to
maturity.
1049-1063.
Acta Bot.
Need.
17(2), April
1968