¹) § perhaps discriptions they develop starting point, protostele

On
the
foliar
origin
of
the
structure
stelar
of
Marattiaceae
internal
the
¹)
by
J.C. Schoute
.
§
The
peculiar
has
ceae
gators,
elucidation
the
a
of
of
stem
The
the
What
they
20),
(2),
have
same
important
adult
show in
starting
elaboration
the
investi-
devoted
stelar
point,
chiefly
due
to
to
the
structures
in
forms
of
acquainted
most
us
(4),
Gwynne
West
(7),
the
10), Shove
(9,
Hill
vascular
in
good
detail
the
stem
in
parts
Marattiaceae.
these
in
discriptions
different
different
develop
the adult
more
genera,
protostele,
the
same
in
stems
or
less
and
(21)
stems, whatever
the solid
a
Kuhn
and
us
they follow exactly
differences between
are
made
strikes
perhaps
that the
may
the
Charles
arrangement of the
most
the fact
many
has been
(12),
Farmer
(3),
(19,
Blomquist
the
of
morphology
in the Marattia-
stem
attention of
work
of Mettenius
papers
the
deal of
the
these ferns.
Vaughan
with
of the adult
drawn
good
Brebner
(18),
of
structure
long
since
and
Introduction.
1.
and
the
from
that in
rules,
is
structures
so
the
their
that the
different
advanced stage
forms
in
the
development.
The
J
)
course
Extended
Association for
of
form
the
development
of
paper
read
may
therefore be
before
Advancement of Science,
Section
Hull
K
of
described
the
British
Meeting, Sept. 13, 1922.
18
270
general
in
done
The
starting
referred
adventitious
hurried
the
same
or
ferns;
internal
stelar
troduced,
of
which
the
by
the
Hill
wards
in
the
from
In
outgoing
p.
cortex
earlier
t,
were
crescentic
once
of
gaps
and
stelar
the
that
which
single
a
in
the stele
a
normal
so
point
to
that
of
the
element
new
is
ferns
among
the
from
elaboration
a
of
structure
different
rare
particular
structure,
in-
and
interest.
the
in
descriptions
deal before
viz.
describe
the
their
stele
I
come
of
the
to
the
the direction in which
from
these
the
and
descriptions
which
leaf-trace,
tissue
376);
the
protostele;
e s
leaf-traces,
remarkably
have
generally
arose
leaf.
as
I
at
cause
and
grow.
vascular
4,
is
begins
one
an
initiated.
when
our
in
structure
instead
ferns,
comparatively
internal
The authors
the
is
to
general
way
structure
the leaf-traces
begin
only
is
however
the
leaf-traces
no
strands
other
the
deserves
is
with
authors
it
which
fully
There
in
stelar
have increasing
in
point
is
the
leaf-trace.
dictyostele
a
this
to
cause
investigated by W
the first
strands
as
way
of other
of
yet
the
gaps in
lateral
been
even
leaf-traces,
simple
protestele
two
that
generally
of
sporeling
already
present.
was
became
of
these
Marattiaceae
into
not
has
as
observed
377)
p.
leaves
of
one;
Up
this may be
in the
structure
protostele,
protostele
the
insertion
solenostele
origin
stelar
have
do
and
following
median
which
the
elaboration
consist
the
the
traces
over,
The
soon
of
(21,
leaves
the
in
above
Marattiaceae and
the adventitious bud
in
stages
in
bud
first
these weak
only
the
solid
West
to;
The
all
point
the
invariably
is
for
terms
follows.
as
the
traces
axile
are
phenomena
developed
leaf-gap
the
takes
away
strand
simple
at
as
if the
acropetally
is
a
caused
portion
(Farmer and
first, but after-
plant produces stronger leaves they dichotomise
(Blomquist
2,
p.
186)
and
finally they
are
271
often
given
in
some
off
In
some
simply
real
of
the authors
formed
are
the
his
below
all
in
coming
clearly borne
dermis
(this
any
induced
in
adhere
to
far
as
I
the
of
fact
leaf-traces
ward ;
if
tribution
or
the
even
317,
which
has
I
pass
follows
consider
the
true.
as
formation of
and
the endo-
this
to
as
be
to
proved) and
inclined
16, p.
has
it
From
is
this
spoken of
from
arise
525).
direction,
take
that
the
17
156) I
p.
leaf-traces
still
am
been
never
development of
check
which
place
in
is
the
truth,
its
of
is
inclined
to
refuted
as
of
procambial
as
the
order
the subse-
as
differentiation:
basipetal order of the induction
the
traces
were
from
given
insertions
regular
of
regularity
than
off
should
the
the
the
be
is
distribution
leaf-insertions
from
the
quite
phyllotaxis
first
invisible, does
course
same
processes
diminishes
these
more
the
to
visible
the
often
of
of
strand,
following
only proof
by
be
time
p.
the
about
described from
is
definitively
leaf-traces
explanation.
an
will
are
about the
writes
same
roots
(3,
that
sufficient
not
necessarily
quently
the
(Schoute
opinion,
observation
is
induction of
not
leaf-traces
know.
Direct
strands
this
the
at
been
the
prove
basipetal
a
that
roots
Brebner
papers
to
be
n e r
is
the
to
as
is clear that
it
the
development
but
yet
regards
previous
phenomena
cases
to
b
r e
hence the
downwards”
as
tried
not
that
rate
description
the
mind
in
outwards and
In
and
has
370).
the
that
considered
is
the stele,
to
believe
to
Danaea B
on
cases
upwards,
other
or
laterals
two main
presumption
any
In
527);
six
as
reality and the expression
in
leaf-gap
paper
“In
point:
have
inclined
the
describing
without
3, p.
n er
stem-stele
21, p.
es t
of
structures.
acropetally
of
origin
the
(W
way
metaphorical
origin
In
this
cases
b
r c
the
which
among
easily be recognised
may
at
they
species
(B
initio
leave
even
separate strands,
more
ab
separately
stele
as
of
the
downthe dis-
regular
itself.
the
given
as
272
If
read
we
the
the Marattiaceae
which
does
which
is
We
that
find
inserted
is
natural
the
for
if
of
at
the
the
strands
levels
is
even
of
nothing
fact
one
for
Danaea alata
(21,
usually
are
This
370).
p.
down from
come
the leaf
the stele may be somewhat
to
strands,
two
is
leaf-trace
one
different
their insertion
stem :
different
the
the
on
quite
into
stele
West
by
strands
lateral
there
it;
structures
there
of it.
indicated
namely
two
favour
in
stelar
of view,
point
harmonize with
not
decidedly
the
this
the
of
descriptions
from
according
the
form
of
the origin
as
to
stele.
If
the
two
or
leaf-trace
the
origin
the
fact
on
truth
leaf-traces
the
of
however
in
the
Marattiaceae.
be
inclined
are
of
conception
we
To
then
to
this
answer
Posthumus
if the
to
in
a
the
I
the
illustrating
growth of the
traces
question
from
(13)
facts
more
downward
ask:
situated
are
understand
to
reconciled
easily
cortex.
treat
the
might
not
understand, and
to
strands
the
in
way
is
two
of
leaf-gaps?
paper
shall
we
how
ward,
up
symmetrical
Later
One
higher
levels
the
the stele,
difficult
already
is
different
at
that
perfectly
the
strands
more
from
springs
down-
grow
formation
may
refer
of
to
a
Groningen botanical
laboratory.
Posthumus,
formed
for
in
that
a
basipetal
(p. 277—278)
mesarch.
panied
If this
a
of
on
c.
p.
that the
The central
that
leaf-traces
comes
by
is
a
certain
poorly
tube in the strand,
strength
or,
to
in Ferns
protoxylem-strand
leaf-traces
fresh
bringing
1 -45 —148),
parenchyma strand
diminishes in
opinion
and
way
its adaxial side
cylindrical
itself
(1.
view
the
sharing
as
conclusion
were
of
developed,
the
arguments
the
originally
is however
amount
are
accom-
parenchyma.
it
will
form
protoxylem-strand
in downward direction,
a
funnel
parenchyma.
This
funnel
is
stronger
developed
in
traces
of
stronger
273
leaves, and
ward
of
xylem elements,
These
forming
form
may
These
in
medulla,
a
inner
I
seems
In
the
first
of
cases
a
all
the
it
proved
From
stele,
only
is
to
be
that
me
internal
more
small
a
extremely useful
they
may
be
of
its
the
from
precisely
the
strand,
meristele,
which
composed
534),
p.
from
runs
in
of
this
view
the later
formed
over
way
the
be
must
the
on
a
a
and
coming
trace
the lower strands
back of these
this
on
which
being
on
and their
only
can
parenchyma strands
these
subject
Mettenius
since
commissural
as
p.
first
at
interior
strands
it
of
top
higher leaves, which
strands,
parenchyma
the literature
partly
some,
medulla.
quist2,
The
upper
others
in
strands
are
strands
compensating
as
compensating segments (Ersatzbiindel, Ersatzschicht)
times
correct.
takes its origin
system
(Brebner 3,
only
point
our
accompanying
In
the funnels
leaf-traces.
leaf-gaps.
defined
one
cribed
the
description of the origin of the internal
continued downwards abuts
into
already reduced
present,
to
and
xylem
inward.
leaf-gap
pursue
stem-xylem.
the
was
was
insertions of
genera the
phloem
well
from
the
stem-xylem
complex anatomy of many Fern families,
the
to
normal
of
margins
At
stem-xylem
probable
very
come
structure.
is
the
funnel in the
hollowing
medulla
a
suppositions have
now
from
or
if the
its down-
in
prevent the formation
the
groove in
a
even a
thus
the stele
must
cause
unite,
above
gaps
understanding
and it
will
it
and
parts
reach
parenchyma
the leaf-traces, and
funnels may
its
it will
most cases
As this
course.
opposite
in
in
186),
term
least
literature
as
conveys
by
system;
close
or
strands
(West
central strands
the
and
leaf-gaps
(Brebner 3,
the
idea
aid
of
the
direct
recurs
p.
a
p.
the
statement
more
West
21,
p.
p.529).
is
gap
closed
from
that
than
Blom-
3,
arising
or
or some-
369,
(Brebner
that
strand
even
536;
21,
des-
once
370,
the
internal
in
the
Blom-
274
2,
quist
if
Instead
of
The
blindly
the
described
of
88)
medulla.
by
endings
were
facts
are
quite
some
the
where
the
internal
continued leaf-trace
a
One
single
of
case
Vaughan
Miss Charles
length
would
desig-
(20,
system
may
end
condition
this
for
261)
p.
describes
0.3
to
up
the
strands
(7,
with blind basal
mm
indications
I
*);
will
support
towards
the
in
revert
the
take
not
stem
literature
this
to
if
views
our
did
reverse
free
ending
vague
case
strongly
that
certain
internal strands
be
somewhat
the medulla.
in
These
used
are
therefore choose the
stems
Gwynne
and
certain
a
those
developed,
Archangiopteris;
p.
which
shall
leaf-traces”.
In
cases.
in
be abandoned
course
these continued leaf-traces may be different
of
weakly
very
of
must
terms
I
confusedly,
course
in different
is
above
“continued
nation
idea
good.
the
and
vaguely
is
holds
view
our
this
186):
p.
place:
There
apex.
that this
in
again
the
we
may
second
paragraph.
In
most
with
In
normal
ends
find
we
central
these
such
we
fuse
distinguish
may
second
such
it
way
cases,
of
fact
is
be
two
though
the
"It
paper
however,
is,
of
ends
inferred
in
the
the
are
from
strands
the view
turned
the
paper
of
is
not
not
lower
while
found
Farmer
ending blindly
as
to
their
is
these
and
Hill
unusual
very
upwards
to
in the
secondary
not
directly
context.
Miss
medullary system
description
Hill
with
up
a
leaf-gaps
and
of
continued
joins
above
some
Farmer
the
commissural
the
found
origin
at
which supports
That
may
indication
that
sentence:
rudimentary
side
opposite
read
we
the
system
medulla and
occur
may
when
the
a
internal
the
indicates
534)
cases."
though
Another
to
p.
way;
parenchyma,
in
expressed
her
(3,
387
p.
of
some
origin
continued leaf-traces
and
the
on
strands
this
in
weak
crosses
cylinder
The
Brebner
!)
blind
the
tissues,
a
generally
leaf-gap.
find
with
stems
leaf-trace
(4)
vascular
here.
cases
the
however
cases
other
quite
Charles
may
clear.
(7,
p.
88);
probably pertain
275
others
devoid
are
{Charles
7,
what may
be
if
they
forming
the
in
acropetally
internal
leaf-traces
together
This
in
leaf-traces
would
it
do
not
hard
the
cross
protostele
a
be
just
protostele
is
elaborated
soon
the
did
(West 21,
becomes
of
postponed
and
The
the
resembling
In
in
this
second
with
stele,
the
In
ued
consequence
indicated
second
this
stele)
ture
development
the
stems,
stele but
also
far
abut
may
of
what
simply
be
gaps in the
built
is
second,
said
third
the
this
traces
strands,
and
begin with
I will call
above
stele, and the
etc.
portions
(between
continuation,
of
contin-
them
the roots, which
and which
the
are
formed
different
etc.
are
not
struc-
yet mentioned
abundantly formed
not
internal
may
the normal and
what is known about the stelar
of
of the Marattiaceae I have
on
if
repeated;
to
up,
one.
downward in
medulla
has been
continuation
second
of
is
interior systems.
parenchyma
formed
that
trace
lower, earlier formed
dorsal
newly
means
the
exterior
enough
on
of
cause
normal
process
only
dictyostele
where needed the
account
the
the
to
remarkable
very
It
now.
higher
the leaf-traces in
as
third system.
first
as
of the
in
unite,
the continued leaf-
commissural strands of the authors
or
stem
first
bundles
begin
second
extend
a
internal steles
leaf-traces;
In
same
the
into
compensating
leaf-traces
of the
one
soon
respect
will
they
in
a
accompanying
in
turn
these
so
every
the elaboration of
the
main lateral
place
leaf-traces
their
then
and
stele
continued
be
quite natural
two
takes
stele,
the
the
369, 386); this
p.
continued leaf-traces
internal
strength,
but
as
leaf-traces become double, just
the fusion
explain.
to
in
medulla,
as
continued
phenomenon
exactly
become stronger and stronger. The
themselves
cortex
is
basipetally;
grow
increases
system
succession
distribution
irregular
the
irregular
an
did.
sporeling
traces
if
expected
the
continued
them,
This
89).
p.
grew
When
of
only
ones.
on
the
outer
276
The
roots
leaves
idea that
stelar
and
their
the
that
need
we
understand
to
Blomquist
of
phyte
younger
strand
may
Holle
in
few
a
is
of
the
the
above
by
a
the
leaf-gap,
continuation of
In
Kaulfussia
are
inserted
but
usually
insertion of
The
a
the
arise
of
the
root,
displaced
as
the
case
varying conditions;
on
leaf-trace
every
p,
506)
that
the
roots
in
the
inner
majority
meristeles
dorsiventral
nearer
stem
almost
through
between
West
roots
arise
the
on
West
(21,
for
leaf-trace
root
the
make
the whole
p.
from
369)
goes
the
outer
where
struc-
Danaea and
p.
262);
stelar system
internally
(21,
p.
383)
upon any of the
stele
cauline
for
a
first
inward.
bears
continuation of
between
and
Archangiopteris (20,
spot
found
second
the first
relation
roots
from
indiscriminately
the
central
stem.
mentioned
plants
two
surface
another relation
these
in
but
outer,
which
lower
to
the
corresponding
individuals
grown
Gwynne Vaughan
by
the
may be
trace
“In
Angiopteris and Shove (18,
under side
the
related
condition of the central strand.”
analogous,
full
in
roots
the
to
way
Quite
ture
of
stems
than
thickness
of
exist
attempt
our
to
of its
insertion
found
217)
not
follows:
corresponding
spiral
specimens
Those
upper
their
col.
(8,
originate,
steles.
the
to
the
authors mention
young
found
of
in
definitely
as
joined
is
trace
the
does
roots
is
older stages the leaf
left
or
due
Other
in
above
regular.
about the young sporo-
leaf
described
leaf
The
structures.
writes
is
the
to
shape of the cauline
influence
with the
“Each
:
each
the
in
right
be,
188)
p.
rather
naturally; I shall however
an
stelar
their relation
directly
the
cauline
(2,
stages
root; but
to
reckon
Angiopteris
root”;
one
that such
not
the
may
itself quite
presents
demonstrate
be
influence the
might
related
undoubtedly
cases
many
distribution
roots
system
to
try
in
are
and
leaf-traces
roots
the
and
that
most roots
stem
meristeles,
just
under
the
leaf-traces.
roots
therefore
is
277
a
varying
arise
are
the
present:
of
a
the
roots
trace
second stele
from
probably
It
any
of
the
the
that
early stage
get
(21,
meanwhile,
erably
In
es t
the
roots
which
number
distribution
be
never
explained
irrugularly
have
forms
the
on
root
a
about
the
cauline
in
inner
a
of
the
shall
roots
therefore
far
as
of
crosses
leaf-gap above;
near
addition of those of
I
very
and may
system
of its vascular elements
the
by
the
by
distributed
tracheal elements
end of
observations
own
continuation.
may
more
the upper
augmented
my
can
more
wrote
the
and
trace
“The commissural strand then
371).
p.
towards
over
the
W
stronger.
Danaea
where
from
or
regular shape of the different
much
develop
insertion
stele
trace,
be
may
the
first
continued
a
the
system
meristele
a
may
of
that the
influence
the
continued
points
the
only
is
system
cauline
from
action
roots;
clear
just under
insertion of
usually
roots
leaf-traces
from
stele,
as
the
farther
however
is
steles
inward
which
descending
originate
may
under
from
places
the
the first
on
bends
trace
the
but
one;
those where
a
consid-
are
root-trace.”
only
point
required
as
to
the
by
above considerations.
After
this
this
paper;
hope
of
development
idea
any
of
the
other
lowed
of
plant.
to
from
West
great
prove
leaf-traces
stelar
the
indications
system
to
is
I
now
that
to
apt
morphology
the
of
of the
us
give
the
the
to
come
idea
a
aim
of
basipetal
much better
Marattiaceae than
conception.
In order
ution
introduction
long
I
of
this
that
gives
an
a
old
writes:
will
In
specimen
the
"An
is
show that the distribso
as
the literature
be the case, but
figure (21,
to
must
leaf-traces
conception.
this
resemblance
He
I
bring this about
continued
p.
of
367)
of
no
interesting
we
have
fol-
some
sufficient
proof.
internal
stelar
an
Danaea,
external
might be
which
system
parallel
of
a
may
shows
a
younger
also
be
drawn between the leaf-traces and the compensating strands.
278
which
respectively
stages
with
through
pass
simple
single
intermediate condition
where
a
with
ending
and
from
not
the
What
of
point
second
the
view
stele
and
I
this
is
the
not
living
material
plant
Garden.
al
pruinosa,
J
filicum
The
a
leaves:
genetic
of
in
lack
the
on
our
the
gaps in
phyllotaxis. From
the
is
out.
consisted
of
one
single
Groningen botanic-
Nr
11101
Marattia
as
of
surface
the
rhachis
habit
four
the
of
a
sketched
living
from
four
Induces
As
numbers
remaining
a
with
left-
and
nowhere
sure
me
yet
that
to
the
it
23—26. After
from
part,
The
fresh
one
leaves, counted from
leaf bases
duck-egg.
in
was
living
the
the
recurrentes
made
Sm.
leafbases and four
Fibonacci
regular
removed
which
of
roughly
were
scar,
venulae
fraxinea
garden.
a
the
spiral;
I
first
phyllotaxis.
imbedding,
general
Marattia
our
this
under
number
be
shape
any
The
the
by
gathered and that
disposal
was
sectioned
and
without
a
the
visible
partly
the form
Kze:
it
to
determining this,
paleae
the
of
mentioned in Christensen Index
not
determine
lowest
!)
find
continued
Investigation.
my
showed
proved
handed
further
their
).
to
and
to
cultivated
was
of
required
the
from
be
that
is
places
Marattia sp. from the
species
stem
This
the
It
a
expanded
sides
of
at
the
distribution
to
§ 2.
The
which originate,
margins
however,
follows
myself
set
lateral
an
its base,
near
372).
found in
as
through
passing
strands
the
prove
that
such
is
that
from
p.
to
really
are
literature
reason
(21,
have
we
leaf-traces
independent
two
base, but
gaps.”
respective
strand,
the strand forks
of
series
ontogenetic
an
which
sections
state;
the
of acuminate
believe
species
not
that
has
object
the
of
was
were
apices
and
pruinose,
was
the stem,
of
cut
was
leaflets,
presence
of
Angiopteris pruinosa
it
was
probably
not formed
a
form
fructifications
279
fixed
in
of 250 fj-
with
the
contained
1
of
the
stem,
more
and
more
first,
the
central
1■—42
showed
others
of
been
The
drawn
low
nifying
From
these
1
2;
the
left
The
white.
concentric
and
possible
fixed
being
and difficult.
camera,
to
were
lucida
camera
mag-
and
one
roots,
on
the
of
their
the
the
stele
54
three
inner
we
on
the
ring.
and
roots.
the
inner
arbitrary
avoid
to
way
is
in
of
is
three
include
an
outer
indicating
confusion between
drawings.
in
drawing
Where
less
outer
roots
than
ring
it was
not
always
xylem of the meristeles
continuous
no
by
one
cortex
therefore also
was
see
the
xylem parts
represented
the
between the
the
in
xylem parts of the
these
are
divides
rather
in
rootstele
In section
of
roots
chosen
only
the
surrounding
one
distinguish
the
represent
outermost
This
reproduction
in
give part
magnification used
the steles
as
the
meristeles
to
meristeles
areas
middle
was
At the low
long
all
numerous
part.
roots
and
a
13
stem
had then
quite whole,
with
/.,)
the
stem
more
not
were
of
contains sections 54—50. In every
phloem
the
inner
I
1
Fig.
lines:
cortex,
roots
the
got
21
(X
Sections
sections
parts
As
glass from spectacles;
a
section
the black
drawing
the
of
drawings
and
meristeles;
and
which
the
became
evidently
was
apex
bigger
sectioning
of
over
peripherical leaves
these 42
After
sections.
magnification
and
stem
cut
leafbases
stand.
figures
the
sections
monocotyledons.
the
cut;
farther
the aid
glass
itself.
thicker
some
the
many
stem
were
/*
sections,
and with
same
the
350
at
42
was
the
those of the
last;
in
as
halfway
cut
Shanze micro-
a
razor
it
as
2—6
sections
ones
somewhat
requiring
microtome
leaves,
only
in
contiguous,
hollowed
slightly
ordinary
clasp of
obtained.
were
Section
apex
manufactured
specially
a
tome, and
spring
given
with
six
from
in black
the
the
as
meristele.
of such insertions
of meristeles
and
three
Fig.
1.
Marattia
sp.
Four
consecutive
sections
of
6
stem,
/t
nat.
size.
281
first
At
reveals
In
of
distribution
closer
irregular;
54
section
the
of
the
belonged
to
the
from
comparison
edge
the
of
part
partly
cut
one
The
both
From
this
cut
both
we
the
the
its
20
still
to
have
nearly
the
in
The
in
sides.
For
sight
which
which
do
of
the
only
was
of 19 and 20
first
unite
this
18—20
22,
18
by
is
to
53.
22 is
placed
equal and
single
then
one,
simplified
is
equal
two
only
in
trace
as
leaf-trace
way
may
with
21,
which
a
we
state
all rather
to
plant,
section.
which is
whole
single;
trace
the
in
recognise
a
are
that the
in
half of the trace;
double in the
trace
in
traces are
lateral
the
following
same
19 has four strands in section
sections:
for
54
the
rest
If
being
the
one
ring
of
strands
not.
the
one
soon
and
confirmed
thus
shown
half
really
46.
section
section
18
traces
traces
but
than
and
is
higher
of
strand
course
idea
on
the unsectioned
leaf
contained
we
54,
strands
strands,
leaf-traces
be
that
18,
left.
were
were
insertion
right
This
in
traces
right
conceive
downward
its
51,
to
left
trace
and 20,
section
and
only
were
spiral;
the
to
succession
19
in
left
two
two
strands.
18
same
like
present
lower,
in
in
nearer
that
regular
the
strands,
Here
two
strands
inner
with
the
leaf
under,
easily be inferred
of
way
of
that both consist of three strands,
see
of
to
soon
strands
two
could
scar
same
right and
strands
the
continue
partly
we
the
that
is
these
the section
the
seems
however,
right side
seen
18
leaf
right and the left
only
three
the
two
the
and
In
away.
to
of
where
stem,
determined;
were
viz.
the
meristelcs
leaf-trace
be
That
section.
of
the
examination
to
are
trace
of
regularity.
lowest
strands
two
of
degree
great
a
which
the
the
sight
rather
be
the
dictyosteles.
we
look
the
two
smaller
it
not
ring
belong
In
to
for
is
the
this
part
stele
on
clear
both
at
dictyostele
at
we
occupying
ones
quite
outer
has
same
first
great meristele,
very
and
the
quite regular,
section
first
and
53,
Fig.
2.
Marattia
sp.
Six
consecutive
sections
of
stem,
%
nat.
size.
283
which
whole
is
there,
leaf-trace
22
strand
situated
is
have
we
that
see
approached
between
to
them,
the
the
strands
two
ring and that
but
much
of
third
a
towards
more
the internal system.
We
of
a
to
the
leaf,
make
xylem
the
50
of
at
even
sections
section
50
which
left
shifted
still
free
from
right
the
is
of
to
the
xylem strands
disappeared.
the
to
moves
the
the
in
unite
observe
this
dictyostele
first
union
54;
we
the strands
we
may
of
much
sections
two
the
to
this
section
on
our
stele.
outer
first
inward of the
will
than
in
the
situated
one
strand
places
To
for
ring
but
22,
in 49 and 48 the
22;
describe
traces
cut
in
done about half the way.
how
gaps.
consists
four
of
free.
The
it
25
has
is
united
to
still
to
is
the
36
the
nearer
to
external
in
the
to
clear
quite
advanced
in
shown
leaf-trace
24
the
system;
united
strands
is
unites
Leaf-trace
the first
more
in
continued
system
already
lateral
two
strand
dictyostele.
attached
of
strands
right
its
the internal
second
side
Leaf-trace
trace
being
to
inside
and
when
just
the
parts,
this
has
sections
are
leaf-trace
trace
higher
their insertions.
23
49—38
has
left
trace
the first
external
in 51 these have
at
well look
which
the
In 50 the three
it
These
formed
as
just
Leaf-trace
of
clearly
see
22.
so
leaves
higher
trace
from
somewhat
three
going
am
have had lower
however
the
54
in
I
we
trace
the
in
or
continued
way
Here
of
separated:
formed just
are
gaps
strands
to,
and
however,
section
should
its
go
2.
into
there.
is
part
system,
First,
fig.
by phloem-parts,
middle
internal
must
in
strands
two
uninterrupted
is
So
stele
making
we
divided
already
is
still united
two
the
is
one
neighbourhood
between the
xylem
last
is
50
section
the
in
are
In
clear
this
to
stem,
just
this
which
second stele.
closed
is
that
see
higher
To
the
shall
stele,
which
and still
is
also
on
continued
the second system.
in
section
system,
and
50;
the
its
gap
284
between
this
the
no
to
continued
it
see
have
deal
models
easily
stelar
in
have
line
both
figured
a
given
not
of these models
photos
to
discern.
one
55
than
the
the
outer
the
sections,
of
the
circles
the
such
to
a
wax:
I
longitudinal
a
scale.
were
way
3
10
gives
the
2
so
times
of
in
cm
much
to
that
scale.
represented
the
original
the
gaps
the
way
to
of
dictyostele
in
these
set
relation;
thickness
distance
sections.
of
of the
the
drawing
1/3
the
and
centrum
lengths
this
of
the
that the
the
From
the
drawing
to the
required
thickness
distance
in
I noted
circumference,
the
got
in
dictyostele
arc
nearer
according
the
circles
meristeles;
regularly upwards,
as
of
every
the
chosen
good
arcs
form
). On
was
drawn
ten
magnification
the
l
distributed
being
was
rather
that the
55
circumferences
were
rather
tapered
sector
Fig.
their
them
dictyostele
arcs
represent the
to
where
was
the
as
concentric
—
section
drawing
10-fold
this
in
conditions
circles
same
In
that
the
46
10
inmost circle
the
dictyostele
enlarged.
in
fan-shaped.
is
and
were
one
in
intermediate
sections
between
of
took
after
open
dictyostele:
outer
intended
one
dictyostele
cm.
circles
the
the
gaps
the
7,2
44
of
the
As
1)
though
survey,
modelling
to
cut
as
better
a
equal
figure
centrum
from
true
These
are
especially the distribution
gives
not
the
the
conical,
drawn, every
where
is
dictyosteles
represents
was
From
which
it
were
1
they
steles is difficult
successive
though
structures.
unrolled.
3
Fig.
at
Most
paper.
requires
this topic
on
useful
most
a
wax,
disadvantage:
one
a
two
respects
and
itself
As
section 44
drawings
authors
most
the reader, and
to
in
in
of
series
energy,
have
structures,
many
the
50
in section
to
have therefore tried another, easier way of
representing
the
in
25
upwards
follow
53.
section
trace
elucidating the complex
however
leaf-gaps
such
models
of
means
from clear
of the
of
mental
reproduced
far
in
of
should go
in
we
state.
constructed
laborious
I
free
in
inwards of
we
if
be closed;
to
closing
it
see
trace
the examination
good
ar
we
discernible,
more
As
a
strands is going
two
gap downwards
As
is
its
of
section
circles
became
the
circles
was
pretty
diminuation.
285
being united the figure
points
only
fill
to
more
roots
the
In
marked
leaf
or
with
the
taken
of
the
insertion
of
the
to
the
cross.
placed corresponding
of
trace
which
inserted
was
dictyostele
the
the
in
has
have
which
The
in 26
hardly
the
that of
is
reverse
simple
the
at
the
each
the
of
is
left
insertion;
on
tissue.
shapes;
somewhat
a
23,
in
the left
two
the
of
parenchyma
varying
(e.g.
In 28
its
that every
vascular
often
one
case.
is
strands
dorsal
a
remarkably
strand
the
figure
lateral
two
the formation
right
as
24,
25,
strand is
components
are
discernible.
All
this
is
leaf-traces
The
quite
are
of
the
the
binary,
therefore
which
choice;
in
the
two
leaves
it
gap
can
27
six
gaps
The
with
quite
the
facts
found
the view
and
all
the quinary
the
traces
as
running
regular
parastichies
ones;
the
strands.
every
come,
dictyostele
If
we
we
have
ask
not
be any other than that of leaf
inward
by
that
gaps.
leave
lateral
incoming
regular,
conspicuous
adjacent other
the continued
the
is
most
leaf-traces
hardly
with
over
basipetally.
the ternary
continued
between
more
af the
leaves.
has
The
accordance
developed
phyllotaxis
that
in
and
authors,
previous
gap
than
level
The
otherwise of
the
was
considered
was
elliptical figure,
an
feature in
apparently
of
insertion
by
counted
a trace
off there.
cut
wings.
prevents
are
gaps
higher
are
two
places
incoming
be
where
where it
or
rendered
were
conspicuous
strand
as
stele
most
leaf-gap
still
dictyostele
an
should
trace
The
not.
or
had been
leaf-trace
27),
continued
strand
The
the
the
into
up
rather arbitrary whether
was
departing
a
leave
So
were
the
a
had
sections where hurt,
gap.
trace
if
the
centres
with
numbers
the
In the sections it
to
the
dictyostele
gaps
of
places
some
I
the
on
number
the
in
over
I
nearly completed.
was
where
strand
must
be
n
just
from
much
+
5.
that of
19
286
leaf
32,
the
two
their downward
would
with
of
is
be
the
it
left
strand
cases
traces,
or
the
leaf-gaps
course,
the
If
continued
Crosses
belongs;
two
we
are
numbers
it
and
represent
those
at
of
take it
were
for
the
e s t
which
dictyostele
of
of
of
of
figure
the
times
the above
cut
as
their
open
incoming
as
of
in
men-
continued
free
and
Numbers
numbers
improbable,
traces
different
during
stem
are
combined
two
double
unite
leaf to which
more
at
of root-insections.
centres
right edge
becomes still
strands
by
think
we
W
traces
Outer
sp.
as
of
union
in
explanation
5,
+
n
formed
are
united
having
it
of
The
8.
+
n
which
described
Marattia
3.
only
of
other possible
right strand
of
probable,
very
unrolled.
the
leaves,
tioned
Fig.
is
strands
The
course.
that
different
not
lateral
in
leaf-trace
sections.
those
cases
quite equal.
granted
that
the inward
running strand
287
the
is
of
trace
this
strand
the
place
shifted
all
other
At
the
the
that
time
and 30
strands
sented
by
the
shifted
to
the
is
of
27
and
strands
In
is
the
as
the
was
following.
the
two
the
two
repre-
partly
phyllotaxis. For
than
divergence
A2
.
formed leaf-trace
cushionlike
lateral
two
paren-
are
already
secondary
previously
have
we
the
those
formed,
the
is
of the
smaller
in
seen
of 32 in growing
trace
Thus it
as
is
gutterlike
parenchyma
there,
traces
it
can
depression between them, and
the
the
In
gaps
in
they
now
left
are
all
in
the
bears
second
root
way
we
the number
as
insertion is
one
in
only
irregular. In
higher than the
hardly
be measured
to
great
outer
devoid
placed
in
consider
causing
figure 4.
the
in
and
32.
them,
of
strand. But 28 has
is inserted
dictyostele
as
irregular distribution
(trace 32).
has been drawn for the second
here
as
simple,
first
right
has been indicated
two
extraordinary
have been
If
have
sometimes
given
same
the
figure
a
the
the
roots
height of
right
cases
which is
above.
expressed
back
the
the
way
same
numbers
which
and
in other
dictyostele
gaps
one to
the difference is
27)
the
now
left;
leaf-traces
root;
(trace
leaf-traces
most
the left and
to
to
one,
left
30.
A3
of
that it has
be
to
is
between
go
have mentioned. Their
we
conspicuous;
all
The
drawing the position of the
our
crosses
relation
of
backs
32
but
be
may
consequence
on
as
gap;
the
relation
quite below
not
inward.
run
by
abuts
is
where
place
30, which afterwards
than 30,
32
it
follow
only
in
the
in,
came
leaf
to
29 and
right,
its
on
and
parastichy
Furtheron
of
forced
of 29
to
nearer
ternary
a
been
gaps
of
existed.
already
chyma
same
trace
27
gap
32
explanation
the
that
see
in
g.
The
the
traces;
e.
we
leaf-trace
right.
downward has
29
inward
runs
5,
+
n
where the
to
of 29
leaf
wings.
accordance
to
the continued
in
a
of
We
leafgap
new,
a set
see
dictyostele,
27,
simple
In
but
these
the views
trace
we
gap
32,
get
it
which
288
The
distribution
the
quite
left-handed
a
term
are
in
The
for
a
stele
the
first
In
the
show
second
meristeles;
quite
analogous
the
4.
unrolled.
the
lines
stele
by
the
first,
use
of
gaps
this
gap
n
+ 3.
the
in
gaps
and
that
corresponding
a
the
was
n
to
with absolute clearness that
principal
Black
in
figure
that
to
Marattia
represent
there
sp.
in
second
every
in
one
gap
the
aim of my investi-
the
of third
leaf-gaps
first.
the
the
with
springing from
roots
that their
spots
the
distribution
Every
place
dictyostele of
represent
communicate
strands
the
Second
points
also
were
shows
somewhat under
third system
Numbers
stele
our
from
Fig.
this
and
of
the
in
represented
adjacent
n—3
2,
shows
that
to
explained.
as
the
as
there is
owing
gaps:
parastichies,
the
but
n+
the
three
second stele the quinary para-
distribution
same
is
To
second.
gation,
the
of
after
is
and
Yet
recognized.
“contact”
sense,
n—2,
however,
rest
is
the
in
n+1,
figure,
the
longer
no
the
second stele
right-handed
two
easily
are
of
size
modified
n—1,
are
this
in
gaps
the mutual position
in
much smaller
stichies
the
parastichies
difference
the
of
phyllotaxis;
same
stem
root
where
as
where
cut
the
system running through
are
those
leaf-traces.
of
the
springs
a
open
contf-
and
strands
second dictyostele:
the
is
of
dotted
medulla.
incoming continued
289
nucd
leaf-trace
tinued
leaf-trace,
stele,
bears
low,
so
be
side
of
the
The
we
the
in
figure
31
of
under
its
insertion
lower
down
first
stele),
in
traces
in
one
the
traces
the
first
they
are
In
our
meristeles
the
as
higher
after
is
present, and
53
the
at
left
here still
is
of
is
between
low,
very
more
37
trace
of insertion with those found
these
As
have
this
tried
accurately
the
of
third
the
so
the
in
comprised
continuation of
that
as
some
that
vascular
tissues.
strands of
deserves
the
were
a
begin-
our
full
of
its
permitted;
in
course
sections
the
or
dictyosteles
system
thick
the leaf-
leaf-traces
where
adjacent
strands
tiny
the sections,
the
determine
to
as
them
in
also
were
be
may
the
on
see
also
large parts
there
the
28
(the other leaf-
impression
we
but
under
steles.
strong
a
not
just
root
one
(leaf-trace
cases
arise
different
stele),
leaf-trace
a
relations
most
leaf-trace,
a
root,
one
to
All
to
second
lateral
roots
here
root
the
in
but
leaf-traces,
to
have
we
roots
get
parts
boiling
was
root
34
trace
ibidem),
the
system.
I
attention,
the
one
only where
sections
third
ning
34
the
in
united
of
37
stele).
therefore
continued
as
It
section.
and
insertion
stele;
276)
p.
lateral
not
in
first
(gap
description;
are
height
leaf-trace,
two
54
the second
inserted very
was
last
it
yet
sections
authors:
(gap
the leaf-traces
We
in
that
(see
by
a
4,
figure
in
30
trace
under the
these ways
compare
of
of
lay
con-
every
in
32.
described
middle
reversely
course
stele.
high,
and
literature
and
it,
one
1,
the
in
very
the gaps
The
internal
than
inserted
its
indicated
in
to
downward
centre
difference
striking
If
in
its
not
seen
united
root.
a
that
therefore
to
has
only
starch
discernible
grains
were
dissolved.
The
which
As
sections
are
contained
represented
these
strands
in
did
five
fig.
not
4
yet
internal
with
meristeles,
dotted
form
a
all
of
figure
of
lines.
stele,
a
290
the
third
those
black
stele
and
If
a
with
28
try
must
+ 3.
For
leaf
the
left
side
Besides,
of
second
it
where
the
not
28
one
the
three
come
from,
gap.
leaf-traces,
and
we
leaf
from
come
strand,
its
it
which
the
on
was
dictyostele
+ 3
Only the
gap,
remote,
the mutual
n
34,
37,
strands
they
dictyostelc
strand
as
1, would rather abut on
+
surely
second
where
is
that
n
and
first
the
to
of
continued
more
and
gap,
as
the upper
as
presently.
these
are
here
4- 2
way
noted
left the second
from
arose
middle
they
is
n
dictyostele,
another,
is
+ 5
the
run
deal
conclusion
the
in
as
which
+ 5
n
strands
right part of their
out
that
between
passing
only
denoted
gaps
will
the
the
the
to
come
clearly
the
I
in
make
being
opinion
our
fifth
same
attached.
were
of
the
therefore
where these
viz.
from
to
drawn in
have
strands
originated
sprang
we
they
the
I
two:
five
gap
be
not
places
the
of
27;
others
n
the
of
gap
in
outer
and where
margin
n
the
points
Four
of
could
system
of
probably
at
gap
of
position
is
right, half.
the
in
trace
in
n,
the
the
gaps
analogous
position.
There
In
section
the
As
our
in
cortex
one
of
trace
of
a
leaf-trace
was
the
continuation
of
second
supposition,
this
not
higher
itself
the
higher leaves
the
stele,
third
this
system
strand
were
any
also
be,
be
e.
g.
clearly
have
been
certainty.
came
must
easily
sections
might
them with
see
the numbers
could
highest
32—40
leaves
this view.
from
As
leaf-
according
the second continuation of leaf-trace 40.
the
last
it
is
found.
If
stem,
was
favour of
27—31
and in
still
of
could
the
of
in
leaves bore
leaves
object,
strand
highest
37
gap
to
I
the
traces
traces
but
there,
the
argument,
expanded
them
the
in
6,
The
seen.
the
Between
distinguished
in
another
plant,
our
23—26.
still
is
leaf
would
it
not
it
less
developed
unnatural that
was
would
be
clearly
one
be
its
in
the
second
the continuation of
the
that
the
more
clear
than
strange,
its
continuation.
291
Of
the four strands mentioned,
straightway
and attached
of
leaf-gap:
the
next
attached
In
be
not
them
conical
This
lower
leaf-gap
other
One
the
much
being
apparently
or
right
leaf-trace
30.
I have,
way;
however,
by dotted lines along
sake,
fuse
may
found
which
developed
in
and
poorly
together,
Miss
by
unite
may
is
to
a
in accord-
Charles
and
above.
from
run
lower
its
devoid
of
developed
by the
strengthened
strands
inner
reported
strands,
very
half had been
the
been
as
thinner
gap
).
margin
well
28, which
1
what has
of
only
was
in the
that
investigators,
margin
upper
of the strands of the third system
clearness’
for
dictyostele
with
ance
course
condition,
from
continuation of
represented
represented
the
the
the
to
strand
the
second continuation of leaf-trace 31,
second
4
fig.
our
could
the
to
themselves
only
the
therefore be
may
three crossed the medulla
leafgap
half,
the
there and
strand
only
with
contact
33,
to
half
upper
the
xylem;
34
was
its lower
the
second
system.
I
have still
This
one
half
in
of
our
As
strand
a
The
which
44
present
with
analogy
the
would have
!)
A
out
and
the
better
the
form of
sented
strands
by
from
gap
as
28,
the
as
the
with
a
and
cone;
threads,
left
the
side
strands
spanned
lower
indicated
lost
be
united,
the
the
in
its
obtained
that
third
inner
34
however
been
upper
system
duly
half.
basipetally
basal part and
figure actually
space.
might
sections.
if the drawing
the
upper
the
formed
developed
so
in
having
are
the
half
upper
leaf-gap
aborted
well
of
but
from
bundles
across
contained
the
the strand,
representation would
right
a
strand
wholly
that
theory
two
sections
there,
plant,
that
possibility
induced,
these
the
would have
the
in
the
a
in
come
damaged
suggests
For
had
which
seriously
been
present
4.
figure
was
have
only
was
section
half
mention the fifth strand of the third system.
to
could
then
was
a
cut
assumed
be
repre-
292
badly
developed
seem
form
to
blind
literature
there
apical
wanting
from
endings
that
a
We
difficulty.
serious
a
downward
the
in
perhaps
or
the
should
theory;
part,
now
found
we
indications of blind
are
may
only
expect
upwards
endings.
According
much
too
well
is
to
importance
developed,
Dicotyledons
of
the
the
xylem
Now
it
is
than the
strands
the
first
if
of
strands
and
I should
be
to
This
the
of
we
is
a
the
made
direction
third
can
is
of
the
parts
may
has
an
end
second stele,
strands
this
two
is
to
strand
is
always
not
the
that
that
blindly
the
internal
both
been done
not
infer
as
down-
yet,
direction of
then
the
which induced the strands
is
this.
out in
being
that
see
Blind ends
goes.
induced
another way; the
much less than that of
only conclude that the regularity of the
we
the
transmitted farther
and it need
not,
where
causing
three
other
quite natural,
internal
internal
in
strands
strands;
acropetal
it
regularity
the leaves,
third system
weakened consequence of the regularity of the
Here
of
reduction.
extreme
be
when
the
stimulus
afterwards
to
from
made
system
some
that differentiates.
stimulus
out
single
may
partly
which
only be inclined
of
that
vascular tracheid:
afterwards
arise,
Marattiaceae
upwards,
a
that
proved
were
the
propagation
not
may
unite
capable
induced part
it
observe
parts
strand
differentiated and
Even
wards
these
differentiation
actual
vascular
partly
Yet
vascular
a
attach
certain strand
have considerable parts without either
probable
very
a
not
aborted. In leaves
or
sometimes
may
must
we
whether
developed
apparently
are
formation of
be
to
fact
represented only by
is
meristeles
of
we
xylem.
or
the
poorly
smallest veinlets
phloem
however,
opinion,
my
phyllotaxis.
unite
for
to
the
basipetal
would be
in-
comprehensible.
There
the
is
still
one
internal strands
point that
should
must
have
a
be discussed here: why
strong
tendency
to unite
293
the
to
dictyostele
the
only
not
the
in
this fact.
tissue,
we
we
lower
end
of
sides
of
the
would
for
§
2
the
The
question
structure
distinction
the
all
strands
cauline
could
the
are
try
common
be
a
of
of
may
in
be the top
interest
would
leaf-traces
the
the leaf-
only part,
this
in
dictyostele
leaf-gaps, and then
strands
given
only
leaf-gaps.
that the
some
unite
the
the
strands
than
decurrent
the stele.
for
only
not
behave.
and
evidence
the
the
in
thesis
normal
strands”
“compensating
of
the
must
whether
rise
now
stem
the
strands
since
the
apply
strands
this
stelar
and
de
to
course
distinction
would
be
the
whole of
leaf-traces,
or
The distinction
strands
Bary;
leaf
to
the
different elements
structures.
belong
their
of
two
cauline
time of
which
the
composed
is
be made between
restricted in
to
of
dictyosteles.
those
are
of
margins
other
also
may
common
botanists
might
have
but
dorsal
generally
Marattia
internal meristele
gaps
are
constitution of
between
to
hope
of
conceive
between
I
a
side
inner
Meristeles
the internal
leaf-traces
the inner
in
stelar
I
dictyostele
outer
a
a
The relation
§ 3.
whole
Marattiacea, where the last
have accessory
determine how
the
parenchyma
an
As
account.
have
to
the formation of vascular
the
are
It
leaf-gap.
a
look
to
respect
in
of
explanation of
into
unappropriated for the attachment
therefore
may
full
a
seems
dictyosteles
the
gaps
the union with
margin
and
was
described
plants,
following
meristele.
the
in
we
pass;
for
that
give
can
the
that
very
a
meristeles
these
Through
traces
In
leaf-gap. This
other
in
preventing
expect
probably
their sides;
if
may
may
is
with
to
I
take
parenchyma,
dictyostele
fit
that
pretend
not
Yet
strand of
a
here,
lower
a
also
meristele of the Marattiaceae
every
of
but
literature.
shall
I
the top of
at
case
the
here in
is
the
and
familiar
common
the
stem,
stem; and
this
continued
way
we
that
leaf-traces
294
and
the
to
seem
of
stripes
pass
cauline strands
is
a
e.
and
clearly
g.
solid
the
xylem
different stelar
In
recent
a
place
xylem,
the
is
stem-xylem
the
not
the
by
leaf-traces
the Ferns
descriptive
of
different
protoxylem
is
is
derived from
made
the
fully
is
in
both,
same.
either
out;
the
view
stem-xylem
differentiation is
the
in
views
point
stem-xylem
be
the
way
arise.
and
leaftrace-xylem
of
this
102) he puts his
p.
a
In
not
to
and
the leaftraceis
case
as
likely
as
other.
The
first
of
The
second
xylem,
already
such
it is
the
in
in
any
hopeless
there
in
is
only
remnants
a
of
what
up
old
strong
from
cauline
hope,
Ferns
with
before
in
existed
the
views
To my
leaf-traces
strands
solid
a
the
later
For
as
central
xylem
there where
in
was
leaf-
whether
or
ancestors
or
offspring.
will
opinion
indication
or
found.
shows,
Rhynia
parts
sooner
be
corresponding
a
never
these
is doubtful.
built
of
ancestors
has
of
we
two
is
of
description.
separate
a
as
Posthumus
between
criteria may
whether
disappeared
either
rather
of
will,
the
way
case
prove
deserve
instances
ask
of
sentences
differences
perhaps
stem-xylem
a
To
may
existent
traces,
are
sentence
several
we
two
parts
validity:
its
have
we
has
these
xylem,
loose
these
there
if
quite clear;
the
is
the
Osmundaceae with
reduced itself.
of
(14,
course
the
Whether
be
From
between
of
fossil
perforated
is
paper
way.
the
dictyosteles.
parenchyma strands; besides the central
may
distinction
justified:
Ferns;
in
the
in
seen
structures
following
the
the remainder of the
the
xylem.
stem-xylem
of
arise,
may
leaf-traces
whereas
277) generally distinguishes between
p.
be
their accompanying
part
be
roots
leaftrace-xylem
to
central
This
(13,
where these
dictyosteles
where
would
Posthumus
stem-xylem
the
and
that
and
not
be
easy:
that
in the Marattiaceae
the stelar
that
whatever.
structure
there
are
no
295
To
The
make
this
idea of
solid
a
united.
are
of
dictyostele
stele
formed.
is
the
way;
parts
the
in
Marattiaceae,
are
also
united
will
must
become
leaf-traces
form
the
they
this
the
leaf-traces
repeated
different
Wholly
cauline
Marattiaceae in
been
carried
The
outcome
is
no
So
with
out
in
of
built
up
to
theirs.
however,
this
up
The
on
in
stelar
a
very
suppose
where
leaf-traces:
so
his
laboratory
studies
the
results
system
as
foliar
morphology
traces
of
the
sporelings
of
similar research has
byBaasBecking(l).
was
leaf-traces
whatever in
of
developed by Campbell.
afterwards
them
to
stele and
every
investigated
6);
by
phenomenon
other
the relation
have been
internal
the
formed.
the
on
our
but
between
enlarge,
to
with
way
(5,
point
described
are
from
traces
stem-xylem
simpler
tendency
same
these
cauline part
the
repeatedly
ferns
Eusporangiate
is
has
foliar
apparently
is
much
connection
and
general
Campbell
sporeling
is
views
strands
formed
is
and hence
complicated when the continued
and it
parts
this
in
remnants
usual way,
part
And this
This
the
in
vascular
the
the
in
steles.
have
into
come
uniting
and
more
hypothesis
may
is
more
stele
form
original
cylindrical
disappeared.
and
the
are
stele
the
medullated
a
normal soleno-
a
same
the second
and
have
more
complicated
that
be
part
will
exactly
in
the
stem-xylem
second
a
the
traces
by the
explained
traces
stem-xylem;
not
central
a
steles
two
by
foliar
however,
has
that
suppose
therefore
only
If
it
the
satisfactorily
the
and
leafgaps
of
part
foliar
which
to
of
following.
Posthumus,
by
whole form of
be
may
stem-xylem.
must
The
uniting
of
we
central
the
remember the
strand
development
dictyostele
or
to
advocated
as
xylem
the
By
disappearance
have
we
stem-xylem,
a
from
starts
clear
that the
only,
stele
are
of
of
quite
the
stele
and
young
in
of
the
that there
plants.
accordance
Marattiaceae is,
strictly cauline;
reasons
for
296
this
assumption
that
I
not
can
Neither
not
of
the
I
Eusporangiatae
the
should
stem
we
may
of
fictitious.
The
there
leaf-traces
clearly
the
forms
not
the
the
a
stele
of
the
stem
itself
(1,
stem
does
be
(5
213):
p.
the
least,
at
the
stem
apex
be
so
far
But
as
new
the
proved
that the
granted
the
that
between
has been
this
which
in
true
seen
himself.
cannot
this
reminds in
theory
of D
evidence
embryological
view
only
is
to
apex;
tissue
order.
higher
stem
the apex
leaves with
originate.
phyllopodium
phytonic
part
meristematic
Becking
Baas
the
useful
composed exclusively
is
and roots,
sentence
strands
and
very
of the vascular skeleton,”
up
Campbell
play
leaf-traces
their
of
by
of
entity
considerable time
a
building
the
being wholly
a
Marattiaceae
sporophyte
cauline
is
surrounding parenchyma
an
the
stele
a
the leaves
to
of
part
no
does
apex
the
in the
part
last
are
quite
of
belonging
highest
for
exception,
no
instances,
of
as
up
the
always
the leaf-traces and the
on
purely
Angiospermae
strands,
assumption
writes
is
study
stem
several
built
In
of
tissue
cited
is
a
existance of cauline
the
system
playing
vascular
Yet
stele
theory
by
that the stele
notion
Bary.
only,
Campbell
tissue
de
Bary
the
vascular
the
stelar
revealed
should consist of
even
de
“Without
of
or
of
sense
the
as
false
the
of
out
one;
the
leaf-traces
which
tissues
that
say
of
consist
say
304),
p.
on
contain
the
cauline strands in
to
unnecessary
that
the facts
(6,
based
is
is
opinion.
the view
reconciled with
This opinion
of
share
It
given.
this
support
can
be
to
not
are
p.
is
not
334);
e
mainly
is
his
idea
unimportant
constitute
an
(1,
connection
1 p in
or
is
o
in
335)
of
the
that
the
favour
evidently
does
p.
and remarks that
not
as
the
part of
the
exist
independent
plant body.
I
is
am
not
still
inclined
among
the
to
believe
successful
that the phyton
ideas
in
plant
conception
morphology:
297
if
held
it
which
propagation,
the
be
the
true
laws
of
leaves
should
not
evidence which reveals
the
in
apical cell,
It
leaf
assumes
that
dons,
decidedly
is
only
is
the
stele”
a
the
of
tubular
origin of
included
and
the
the
bundles
This
the
of
of
stem
from
the
cited
in
from
of
other Ferns,
the insertion of
paper
too
§
4.
is
are
the
wholly
leaf-traces;
of
Comparison
the
would
ground
if
tissue
the
by
up
like
13) but
the other
authors
as
leaf-gaps
are
the
right
were
built
Posthumus,
and
the
it
be
clear that,
quite
the
sense
single
a
formed
erroneous;
in
no
in
leaves.”
true
tubular
also follows
in
the
outer
leaf-gaps
stele
as
in
caused
by
element in the present
new
gaps
the internal
Marattiaceae
the
which
group
placed
of
Mettenius
breaks
breaks
between the coalescent
plants (confer
it is
structure
in
the
inner systems
leaf-gaps.
true
Among
first
314): “The “dictyo-
network
Marattiaceae
e.
an
Monocotyle-
the stelar
on
not
part
being that the analogous
the
are
i.
where the
many
p.
are
spaces
investigation,
the
with
Marattiales is in
most
gaps”
from
introduction,
own
my
(6,
Eusporangiatae
of
meristem,
even
such conceptions.
to
in
views
cylindrical
seed
works
dictyostele
all
and
pith
is
the
the
the
his
merely
derived
description
stem
the
as
are
apical
sporophytes
follows
The ..foliar
but
an
Marattiaceae
leaf from another leaf is simulated.
summarizes
within
leaf-traces,
united
a
would
and the anatomical
explain,
as
a
of
that
way
how this
kept;
favourable
not
Ophioglossum
stele,
of
kind
a
in such
the presence of
youngest
Eusporangiatae
monostcle.
be
terminal position,
a
Campbell
of
the
in
to
easy
us
sporophytes
young
multiply by
regulated
would
phyllotaxis
about is
brought
should
be
is
by
ferns
so
with
with
internal
much akin
nearly
all
that
to
stelar
of
structure
other
of
ferns.
there
is
one
the Marattiaceae that
they
structure
investigators
in
the
same
order
298
Marattiales,
terest
to
that
the
Marattiaceae
good for Psaronius
The
is
is
the
Marattiaceae,
recent
and
clear.
Rudolph
wiirden
einfache
aufwarts
Reihen
parallel
den
in
der
of
und
Wege
laufen,
seitlich
our
(15,
and
“Die
Verhalten
stelar
description
than any
of
perfectly
Blattspurstrange
schliessen,
zu
tangential
als
Kreisen
treffen
jedem der
in radialen
stehenden Bundel,
und
die offenbar
wiirden
verschmelzen,
Zylindern
der
und
an
oben sich wieder
u.s.f.
eintreten
derselben erweitert, als
die
Achse
Zeitlang
eine
verlaufen
weiter
Zylinder
nachsten
Verlauf nach
geradem
nach der Anastomose mit den
die Arme
this
purpose
178):
p.
gebildeten
den
hold
views
our
the relation between the different
derselben teilnehmen,
in
schliesslich
in-
dem Innern des Stammes kommen und
ihrem
loslosen,
if
the Psaronius
ziehen, hiebei wiirden sie mit
Bildung
durch
aus
diesen
of
of Psaronius with
see
(15),
for
diesem
schiefem
von
therefore
is
vascular strands is made
und konzentrischen
auf
sie
writes
nach
Strange
radial
in
as
the connecting
also,
to
Rudolph
suitable
more
steles
and
description
by
given
much
even
). It
structure
too.
detailed
most
structure
sp.
the internal
compare
of
l
the Psaronii
viz.
und
peripheren Stelen,
fertige Blattbiindel
austreten."
A
1)
strong
is
Marattiaceae
Ferns.
among
of
the
indication
given
„ebenso
for
the
the
multicellular
near
who
Solms-Laubach
root-traversed
multicellular
by
pseudo-cortex
as
die
ist
hier
Erzeugung
doch
einfache
von
ist
sehenen
Cyatheaceenstamm
bis
Campbell
not
179)
yet
between
hairs,
first explained
formed
by
a
Psaronius
the
and
been
jetzt
nichts
describes
Kaulfussia
pointed
at
eine
peculiar origin
strong development
ahnliches
waren
nur
however
(ibid.
as
far
p.
as
Besonderheit
erwarten
bekannt,
einfache
multicellular
187); this
I
know.
wiirde.
an
and
feature
unique
a
(11,
p.
mehrzelligen Fiillgewebshaaren,
Wurzelhaare
Farnen
p.
root
hairs from the stem surface and from the roots, wrote
wurzelbiirtigen Systemen wenigstens,
man
relation
der
of
755):
an
den
Psaronien,
da
Unter den lebenden
einem
von
Wurzelhaare
root-hairs
of
mir
zu
einge-
flnden.”
Danaea
(5,
peculiar correspondence
has
299
From
that
this
the
description
the
In other
which
first
fern
with
follows
it
second
leaf-traces
the gaps in
species
in
gaps
continued
the
stele,
families
internal
and
698)
p.
phenomenon
viz.
the
in
there
and
five
of
is
investigate
If
ferns.
get
a
within
the
systems
ment,
the
all
in
ferns
xylem-ring
the
on
inward,
in
our
read
that
the
system
lower
the
first
but
from
in
Vaughan
the
all
these
begins
leaf-gaps
description
and
of
an
authors,
to
we
the internal
cases
(19,
accessory
stelar
local
of
in
p.
704)
develop-
cylinder
by
of
the
thickening
As
abutting
703).
leaf-traces.
ordinary
margin.
531),
paper
the conclusion
a
p.
of these many
above cited
to
with the
:
Pos-
p.
present
structure
continued
of
Asplenieae
Mettenius
of
the
the
leaf-gap
224,
view
our
higher
the
leaf-traces
the continuationof these traces
Gwynne Vaughan
fits
well
conception.
origin
is
(in
Chrysodium, Posthumus
stelar
came
elaboration
Posthumus
of
232,
scope
else
derived
at
Thyrsopterideae,
Polypodiaceae
and
the internal system is
progressive
internal
p.
of the
we
nothing
be
to
families,
subfamilies:
Pterideae: Pteris,
Gwynne
Gwynne Vaughan
that
find that the
other
three
258);
(Ceratopteris,
impression
are
the
p. 701,
219
p.
the
origin
however
strong
we
four
Davallieae: Saccoloma and Denn-
(Matonia,
not
202)
p.
least
p.
240, Acrosticheae:
Matoniaceae,
It
217,
p.
Parkedaceae
247);
(13,
at
subfamilies, Aspidieae: Dipteris coniugata,
Posthumus
thumus p.
of
together what
put
we
each of
(in
Posthumus
Polybotrya,
p.
in
Posthumus
Posthumus
the
top
(12), Gwynne V aughan
Posthumus
the nine
steadtia,
if
GwynneVaughan
Cyatheae,
by
the
however, also scattered
are,
structure:
Cyatheaceae
Dicksoniae,
from
etc.
discernable
is
caused
were
down
run
is written about by Mettenius
(19,
with absolute clearness
stele
of
the
distinguishes
the internal
one
described
between three
stelar
by
structure
Gwynne
(13
different ways
p.
202).
Vaughan:
The
the
300
second
with
the
the
into
same
way
different
have
the
of
an
leaf-gap
be
the
as
strands
explained
In many
In
unity.
united, and
ferns
in the
the
strand
enters
the composing
Cyatheae
the
in
composed
these strands
forms all
other
internal system
the
in
decurrent from
minor strands
of all
the
parenchyma
and
to
We
will
run
prove
the
origin
stelar
a
number
structure
of
that
of
this
many
theory,
these
The
1.
of
The
basipetal
stelar
groups
related
the
by
inward
that
a
detailed investigation
will
be
the
same
is
natural
quite
more or
structures
the
in
as
phenomenon
according
to
less decurrent
plants.
Summary and conclusions.
that
the
stimulus
transmitted
is
direction, might
explain
the
of the Marattiacean
literature
been
for
forced
bend
to
distribution of the
peculiar
leaf-trace
important
yet
structures
conception
structure
the
most
of
strands
the leaf-traces may be
as
a
other
presume
remote
§ 5.
course
and part
of
room
them is
in other ferns with internal stelar
closely
in
even
sufficient
not
the medulla.
therefore
Marattiaceae.
so
strands,
strands
into
may
has
margin
the
not
same
running
petiole and the resulting
a
uniting
leaf-gap
petiole.
leaf.
yet
free
petiole.
insertion
in
the
essentially
is
varieties may
of
the
as
not
form
The
in
these
the
in
stele
have
them
the
in
number
parts
the
into
third way
of
the
through
in
run
instead
the first mode. The leaf-trace itself is
as
united
strands
of
free
that
great
a
and
this difference that the inward
clear
is
they
of the leaf-trace
parts
body,
main
stele,
The
pith.
lateral
two
the
normal
already
It
of
from
with
second,
were
that
is
way
themselves
all
point
deficits
made
in
out
to
our
the
a
complicated
direction;
the
in
stem
The facts recorded
knowledge
whether
the
determining
peculiar
stem.
that
in
was
gaps
one
of
that
it
in
the
the
had
inner
301
steles
In
the
single
of
the
of
the
be
from
strands
that
leaf-traces
2.
the
second
of
the
point
after
the
causes
the
fusion
the
The
robust
only
but
in
normal
the
tiaceae
the
is
is
way
leaf-trace
the gap
gives
an
lies
the
at
of
the
dorsal
stelar
Marattiaceae
of
type
same
image
side of
structure
it
only
is
more
Marattiaceae
in
one
a
in
the natural
is
the
that
as
origin
into
that
of
some
in
the
form
the
other
ferns;
single
stele,
variation of
the
the
Rudolph
on
of
the
the
ferns
the
owing
strand
or
be
its
one
by the
structure
established
in
Marattia
too.
other
fern
in
decidedly
leaf-trace
while part of
must
to
in
Marattiaceae; what
the
normal stele,
This
in
structures
in
that topic points
these
medulla.
single
as
exist
to
internal
same
corroborated
is
correspondence
which
leaf-trace,
a
the
demonstrated
literature
In
Rudolph
as
steles,
here
inserted
to
same
individuals with
too
by
probably
found in
direction.
able
the stele;
in
the
relation between Psaronius and the Marat-
shown
as
The
directly
insertion
strands
monostely.
Psaronius
partly
should
separate lateral strands
two
which
the
ones
investigation,
is
they
basipetal development of
their
the
young
adult
different
5.
distribution
as
stelar
essentially
of
families
the
form of
of
The
present
the
and
were
development of stronger leaves with
The close
4.
one
distribution
elaboration
leaf-traces.
in
of
two
the
stems
the
therefore
not
in
the
system
at
strand
gradual
of
that
given
strand.
individual
outcome
is
(fraxinea?)
The
view.
the
gap:
parenchyma
leaf-trace
3.
sp.
third
gap
of
simple
a
the
proof
dictyostele
consisting
two-topped
a
must
thus confirmed.
is
Leaf-traces
cause
the
Marattia
of
places where they
view.
paper
a
in
gaps
those
at
that
to
present
stem
of
is
found
according
occur
of
be
to
are
explained
complication,
single
to
is
it
in
is
tangential
that
only
runs
this
not
row
20
302
of
which
strands,
of
part
the
it,
outer
theory
of
the
upon
the
stele
is
plant
with
of
taken
a
Pteridophytes
7.
way
of
the
the
would
stele
If
distinction
the
not
plants,
and
On
of
wholly
mostly
by
sense
Marattiaceae
between
revealed
should
a
be
leaf-traces
idea
be
of
the
single
a
the
among
the lower
only
by
based
is
the contrary,
stem-xylem
Posthumus
stelar
describing
not
is
do
not
is
structures.
yet
point
be
to
the
ends
higher
of
unit
to
that
the
1926.
it
as
has
conditions
a
a
in
direction.
Laboratory
State
July
and leaftrace-
accepted
Whether
certain;
Botanical
Groningen,
the
would
rare.
is
as
that
it.
seed
be
composed
the
of
is
that the stelar
stele
stem
a
ferns
opinion.
conclusion
and
Further
these
facts
This
there
way
among
they
made
phylogenetic
the
structures
of
this
inward
run
row.
conclusion
that
leaf-traces,
this
in
stele
of
simple
a
of strands;
them.
by
The
xylem
the
as
rows
outer
view
reconciled with
generally
as
of
Eusporangiatae.
the
leaf-traces;
formed
to
two
the
correctness
came
subsidiary
be
is
of
point
stele
the inner will
stele,
misunderstanding
independent
should
stele
be
cannot
study
the
the
Campbell
6.
a
this
check
to
the
to
least
at
parenchyma
from
investigation
desirable
it
unite
may
forms
the
into
enter
may
that
dorsal
the
on
but
University.
of the
303
Literature.
1.
L.
G.
lar
3.
evecta.
G.
b
Gazette.
On
n e r,
the
Marattiaceae. Annals
4.
J.
and
Angiopteris
Ibid.
D.
16,
evecta,
1902, p.
The
,
Americ.
G.
M.
H. G. H
R.
11
o
e,
11.
,
H.
the
some
16,
1902,
the
vascular
other
strands
73,
Psaronius
Zeitschr.
Gaz.
51,
of
1911,
the
sporeling
p.
col.
die
of
81.
Vegetationsorgane
anderer
theory.
303.
1921, p.
iiber
der Marat-
215.
Anatomic der
Gefasskryptogamen.
Flora
457.
den
1890, p.
anatomischen
Solms —Laubach,
Bot.
Bau
von
Danaea.
147.
Haidingeri
fur
compar-
and Marat-
ferns and the stelar
1876,
zu
in
Marattiaceae.
Ophioglossaceae
The anatomy
Zeitung,
Ueber
Graf
517.
arran-
1911.
Ueber die
und
p.
Hill, On the
34,
1889, p.
Ibid.
Angiopteris
181.
p.
Untersuchungen
Marattiaceen
10.
on
need.
371.
Bot.
Botan.
Kuhn,
72,
of
Journ. of Bot. 8,
Charles,
tiaceen.
9.
1922,
Botany vol.
Eusporangiate
Marattia alata.
8.
hot.
travaux
anatomy of
73,
and
Washington
tiaceae.
vascu-
notes
anatomy of Danaea and other
of
structure
morphology of
6.
with
Campbell, The Eusporangiatae. The
H.
ative
7.
des
Bretland Farmer and T. G.
gement
5.
Recueil
Vascular
Blomquist,
r e
Botrychium;
333.
Botanical
B
genus
anatomy.
1921, p.
L.
the
in
general
18,
H.
The origin of the
Baas—Becking,
structure
the
2.
M.
3,
Der
von
Manebach
1911,
721.
p.
in
tiefschwarze
Thiiringen.
304
12.
G.
d.
1864,
13.
O.
p.
Rccueil
On
,
and
the
stelar
on
K.
J.
C.
Die
p.
morphology.
der
23,
additional
mor-
111.
p.
remarks
94.
p.
Marattiaceen.
Ver-
Denkschr.
Untersuchung.
d.
math.-naturw.
Wien,
in
stelar
1924,
Hymenophyllaceae
1926,
und
Wissensch.
165.
Schoute.
Beitrage
Recueil
Beitrage
,
F.
D.
d.
Blattstellungslehre,
zur
bot.
trav,
need.
10,
im
T.
allgemeinen.
Shove,
Gwynne
,
and
other
West,
tiaceae.
Ibid.
the
Ann.
the
Ibid.
Bot.
anatomy
1917,
of
14,
of
Ibid.
p.
to
p.
the
1900,
der
I.
1913,
II.
Ibid.
stem
19.
361.
Ptcrop-
p.
of
on
17, 1903,
1905,
p.
An-
497.
Archangiopteris
the
Ueber
93.
Observations
Vaughan.
contribution
31,
1914,
structure
of
Marattiaceae.
A
Verastelung
11,
solenostelic ferns,
On
20.
On
evecta.
II.
Blattstellungslehre
zur
Baumfarne und die
anatomy of
C.
the
some
Psaronien
anatomische
of
need. 21,
of
anatomy
1906, p.
giopteris
21.
6,
153.
siden
19.
Cl.
math.-phys.
principles
bot.
and
vcrastelte
R.
Angioptcris.
von
Wiss.
some
trav.
Ibid.
Theorie.
17.
18.
the
Akad.
78,
d.
Schizaeaceae
gleichend
Kl.
On
d.
Rudolph,
kais.
Bau
den
Ges.
499.
Posthumus.
14.
16.
Sachs.
k.
phology.
15.
Ueber
Mettenius,
Abh.
the
p.689.
Henryi
259.
study of the Marat-