Idea Transcript
Stomatal ontogeny
and
phylogeny.
I.
Monocotyledons
G.S. Paliwal
of
Department
1.
of
Botany, University
India
Delhi, Delhi,
introduction
The
study
of the cuticle in
clear that stomatal and in
only
the
of allied
systematic
first
Odell
by
two
parts
work. the
(1932) describes
of the
Contrary
leaf
do
provide
He,
has accumulated
a
the
it
prehensive account, information
on
the first
will deal with the
this
is
cuticle.
early with
angiosperms.
The
living angiosperms
features of the
epidermal
distinguishing
that
with
as
must
for
vege-
diagnostic
some
be taken into account. considerable
a
related
closely
other features in classi-
of data
body
However, in the absence of any
worthwhile
bring
to
complete
to
the
dealing
appeared
and fossil
not
fragmentary In
1958).
really satisfactory
publications,
I propose
aspect.
1933,
works
living
characters
two
thought
was
of
argued
epidermal
the
stated that the structural differences in
(1935) means
however,
regarding
1931,
of the
none
angiosperms
Since the appearance of these
one
Florin
abundantly
often of great value
distinguishing
comprehensive
that
fossil
or
total of the
sum
in
are
the cuticle in 84 genera of the
this Edwards
to
epidermis
a
reached
living
taxonomic groups. fication
also
value of these characters in the
and the conclusion is tative
(see
species
thirties of the present century, the
characters
epidermal
delimitation of genera but
remains
fossil
and fossil gymnosperms has made it
living
other
it under
and
monocotyledons
together
com-
the available
separate articles,
two
the second with the
dicoty-
ledons. A “stoma” the
plants
size and
contents
tically they the
comprises
epidermal
may
and or
“stomatal As
are
may
subsidiary cells,
two
cells
cells
guard
adjacent
then known not
subsidiary
as
be related
to
when present, is termed
Florin’s
earlier,
by
in the
between
major
recent
subsequent
works tend
regards
to
appreciated,
work
gymnosperms and
some
shape,
Ontogenealong
stoma
apparatus”
with the
or
plants.
on
the
a
that
features in
leading
to
the
there
and
is
much
was
the appre-
the
rela-
opinion
arrangement of cells a
more
have stimulated
ontogeny
to
interpreting
fixed character in
investigations
morphology
led
gymnosperms
The basis for this idea
relatively
suggest his
on
of
leaf. Although variation than a
good
stomata
in
deal of
ferns,
dicotyledons.
the
Stebbins & Khush
654
groups of
stomatal complex is
what Florin had
As
accessory cells.
cells. The
the “stomatal
epidermal
him that the sequence of divisions
mature
several
or
guard
as
work
pioneering
ciation of the fundamental value of the
held
the
In
pore.
also become modified in
stoma
complex”.
stated
tionships
surrounding the enclosed
the
to
monocotyledons (1961)
the
most
comprehensive
who have attached great
work
is
that
phylogenetic significance
Acta
Bot.
Need. 18(5),
Oct.
of to
1969
STOMATAL
this
ONTOGENY
feature based
assumptions:
two
on
from organ
constant,
I
AND PHYTOGENY
organ within
to
families exhibit constancy for their of the
survey Ananas nus
still
ideas
prevalent
this
on
to
The
those of
that
1966) and
(Hamann
Stebbins
Khush.
The
of the
some
demarcate between observations and
to
bases of
between
comparisons
purely morphological,
are
1961), Panda-
deserves main consideration is
understand the
to
whether these
different families,
& Roy
evaluate and reconsider
point
have failed
is unable
one
are
even
A brief
particular complex.
a
(Richharia
with
to
attempt
an
subject.
that these authors appear inferences and
sativa
variance
at
are
therefore,
present paper is,
that the genera and
1965), members of the Philydraceae
results
the
others,
1948), Oryza
(Krauss
(Tomlinson
spp.
of
possession
modes
developmental
(2)
and
literature, however, indicates that in several instances, e.g.
recent
comosus
that the
(I)
plant,
a
developmental,
or
or
both.
2.
CLASSIFICATION AND
attempts
Although
and
nosperms
1966),
have so
and
1950;
no
serious
works
later
We
lateral subsidiary
two
made towards
thought
monocotyledons.
in
gans
been
dicotyledons (see
& Chalk
calfe
Guyot
have
TERMINOLOGY
of
along
paid
of the
the
1962)
ceae.
the
gave
for such
tetracytic
term
polar subsidiary two
However, these
cells
terms
as
In view
of the above
these stomata
diary
cells.
subsidiary
on
In
in
remaining
two
helped
of any suitable
particular
would, therefore,
like
1
asahkoshik
‘A’
chatur
=
Acta 801.
=
type to
-
to
propose
the
First
letter
of
Asahkoshik
(1961)
order
or
four
+ sahkoshik;
18(5),
Oct.
(adj.)
cells and
shat
1969
becomes
looking
on
Without
=
Six
+
stomata
used
to
sahkoshik;
cells,
bahu
=
certain extent,
for the stomata
so
(p. 664).
formed
=
Many
to
familiar with the
without dwi
difficult
preparation.
from it
are
denote
subsidiary
and Pandanaceae. to a
more
terms
The
represented
are
given
a
no
sur-
Gramineae,
extremely
at
plants. Being
These
are
cells
Xanthorrhoeaceae.
the situation
it
there
subsidiary
Cyperaceae,
subsidiary
I have derived these
=
2
made 4 of groups
Liliales,
more
following classification
Devnagari Script;
by
the subsiarrangement of
and
remedy
immediately
(Aperigenous).
cell.
Need.
the
two
terminology
language Sanskrit,
native ancient
Sans:
in
more
far recorded among the monocotyledonous
Subsidiary
and
members of the
or
are
demarcate the various
Agavaceae, Araceae, Commelinaceae, Palmae,
absence any
to
& Khush
Pontederiaceae,
have four
categories
this attempt has
Although
imagine
found in
Juncaceae,
and
Metcalfe (1961,
accompanied
are
or-
stomata
members of the Commelina-
some
sufficient
with
category
which
and
1965;
of these
which
gramineous type in
number, shape
represented
group
second
guard cells is
in the families
the
first
The
Haemadoraceae,
1
grouping
monocotyledons.
realization, Stebbins
basis of the
the
the
cells. the
rounding
I.
Pant
Metcalfe;
gymMet-
1931;
dumb-bell-shaped guard cells
hardly
are
of
stomata
Florin,
the proper
to
stomata
seen
encountered among the types of stomata
I
the
1889;
characteristic of Cyperaceae and Gramineae. More recently
lateral and 2
in
of
publications has been
only know
cells
classifying
Vesque,
-
by
direct
sahkoshika
two
+
=
sahkoshik;
4- sahkoshik.
655
G. S. PALIWAL
divsions involved No.
of
1) 1)
1) 1)
(+ (+
1)
(+
(+
1
4
2(+ K+
2
1
1) 1)
variable
4(+
subsi- cells
ofsubsi-
No.
(+
variable
1)
1) 1) 1)
4(+l) 2(+ 6(+ 4(4(4-+l) (+
(+ (+ 2
1
6
4
2 2
0
0
4
2
0
3-7
2
6
4
Jain
(1961)
Khush &
(unpblished)
Steb ins (1960)
(1965)
Steb ins Paliwal Shanks
investigaetd
lancasteri;
in
Narcis us
Strasburge r
1) 1)
variable
(+ (+
2(+
6
2
3
Z.
Kraus
rosea
sp.
4
6
com su
variable 3(+ 3(+ (+ (+ 3
3
2 2
2
1)
1)
(+
(+
1
3
3(4-
Jain &
or
2-6
2
(186 6);
Campbel (1960)
Strasburge
~'*
2 2
0
Haber &
Foard
1962)
& &
0 0
2 2
(1962); Haber
(1959; (1961) (1946); (1960);
(186 ); (1892) Jain (1966) (1937) Roy (1937) Steb ins Moreland (1905);
com unis
sanguinea
ajax;
1) 1) 1)
Shah
&
(1966)
Northcote &
(1881)
& &
Benecke Paliwal Paliwal Haman Drawert Steb ins Strasburge Steb ins Porsch Tomlinso Portefi ld Shah Ric hari Portefi ld Flint (1961) Steb ins Picket-H aps Campbel
umbeilatus
p s e u d o n a r c i s u Sagitari Amarylis Zephyranthes Pathos Dief nbachi Ananas Butomus sp.
stomat Genera
vit ata;
(unpblished) (unpblished) (1963) (1941); (1948)
(1866)
Author/s
and
6(+
(18 81);
& &
species
(+
)
2 2
or or
(1960)
Monctyledons.
1) 1)
more
of
diary
1) 1)
Z. Z.
arista
sp.,
edulis
quadrngularis
vulgaris
C.
discol r
Can a Centrolepis Com elina Rhoeo Tradescanti
pendiula genera Zebrina Several
&
vulgare
& &
pubescens ofjlcinarum saliva
Arundiar Hordeum Oryza Phylostachys Sacharum
vulgare
Tritcum
mays Zea
of
Ontogeny 1.
Table Family 656
Alismatce Amarylidcea 1.
2.
Araceae 3. 3.
Bromeliac e Butomace Can cae Centrolpidace Comelinac
Graminea
8. 8.
9.
4. 4.
5. 5.
6. 6.
7. 7.
Acta
Bot. Need.
18(5),
Oct.
1969
STOMATAL
1) 1) 1) 1)
ONTOGENY
AND PHYLOGENY I
1) 1)
1)
1)
1)
1)
1)
(+ (+ (+
(+
(+
(+ (+ (+ (+
(+
K+
1 1
1
3 3
1
1
1) 1)
5(+
1 1
1 1
1 1
5(+
variable variable variable
5
5
more more 0
0
2
0 0
2
(1953);
(1961) (1866)
(unpblished) Khush (unpblished)
0
(1960);
Biegert Jain
& &
& & &
(unpblished) (unpblished) (unpblished) (1965) (1965) (unpblished) (1866)
(1965)
0
(1866)
0 0
(1968)
0 0
(1866)
4 4
4 4
2-6
or or
or or
4
2
2 -
(1966) (1965)
(1966) (1870) Kidwai
Strasburge Paliwal Steb ibns Paliwal Bain ing Steb ins Shanks Paliwal Paliwal Paliwal Shanks Strasburge Shanks Paliwal Strasburge Inamdar Strasburge Pfiltzer Tomlinso Pant Haman &
Pant
Pandanus capense
chine si cepa
Iris
Belamcnda Juncus Agrostcinum Allium
bar densi por um sativum por um
A. A. A.
Aloe
candicans uniflorum
Acta
11.
Bot.
H.
4
Marntcea Orchidae Pand cae Philydrace 13.
12.
Need. 18(5),
zeylanica marginat
Chlorphytum Galtonia Hyacinthus Ipheion Ornithogalum Sansevira Mar nta Habenari Orchis Pandanus unamed Pandanus Helmhotzlia novguine si Ortho ylax Philydrum Philydrela
Iridacea e Junca e Lilacea 10.
graminfolius species fasicularis acorifolia glaberimus lanuginosum pygmae of
Oct.
1969
14.
15.
16.
657
G. S. PALIWAL
division of the e.g.
Richharia &
Roy
presence The
side). cells.
cells. and 3.
similar
a
Such
(Biperigenous).
-
cells
subsidiary of
of this
the
their
owe
families
and
situation has
stomata
subsidiary
cells;
Orchidaceae.
In
been
by
noticed
the
by
(one
adjacent
Gramineae
their characteristic
also been found in
They have, however,
cells
guard
the
to
and
recognized
are
the
to
origin
Cyperaceae
with
category,
either
on
protodermal true
are
repre-
dumb-bell-shaped
guard
Alismataceae, Centrolepidaceae,
Philydraceae.
chatushsahkoshik
group have 4
laterals,
two
(Tetraperigenous).
-
cells. These may be
subsidiary
polars surrounding
Two laterals and 2
(a)
sativa
subsidiary cells, placed laterally
Members
sentatives
Oryza
Liliaceae
Araceae,
(1961).
dwisahkoshik
of2
of
devoid of
completely
are
Amaryllidaceae,
leaves
cotyledonary
2.
meristemoid and
members of the
located
being
either
along
Stomata
included
arranged
in
stoma
a
guard cell,
as
seen
two
under
in Rhoeo;
(b)
in members of the
e.g.
this
different ways: Four
family
Zingiberaceae. 4.
shatsahkoshik
Musaceae and
and 2 in
being placed laterally 5.
bahusahkoshik
members more
possess of
a
of
ring
Families
(Hexaperigenous).
-
such
polar
a
the
Agavaceae,
than 6
Such
stomata
Palmae
Araceae,
cells which
subsidiary
and
3.
so
far for
DIFFERENTIATION
DEVELOPMENT
studies
on
Differentiation
A.
the
through
as
parallel-veined and
THE
leaf
-
light
LEAF
may either be
arranged
some
These
in the form
well
type of ontogeny has
EPIDERMIS AND
epidermis
as
leaves
period mature
the
of
developing
several some
dicotyledonous
interesting
as
points.
well
These
as
are
the last 10-15 years.
during
that in the net-veined leaves of the dicotyledons
differentiate
not
of
revealed
of the results obtained
It is known
considerable
mental stages
young
in
seen
monocotyledons.
families have
do
stomata
a
4
STOMATA
the
in the
below,
IN
OF
monocotyledonous analysed
-
irregularly.
or
been recorded
Recent
are
Philydraceae.
As indicated in table I the syndetocheilic (mesogenous) not
cells
subsidiary
fashion.
(M ultiperigenous).
-
Commelinaceae,
as
Palmae include members which posses 6
of
simultaneously
growth
stomata
of the leaf
occur
together. where
monocotyledons
stomata,
whereas
those
but
on
to
continue
that different
so
This is in
the
the
contrast
basal
older
arise
developto
the
regions
bear
portions
have
acquired maturity. According
to
decreases after arise later
These
are
period
cells
and
the cells which
(1952)
of
by resumption
protodermal
1
Bunning a
of
these
have
rapid
the
activity give
meristematic
cell at
division. one
rise either
again acquired
some
activity
However,
pole
of the
to
pair
a
of
the young leaf
new
meristemoids
1
protoplasm
of
guard
of the characteristics
cells
of certain or
a
hair
of the meristematic
cells.
658
Acta Bot. Need.
18(5),
Oct.
1969
STOMATAL
(also
ONTOGENY
AND PHYLOGENY I
Sinnott & Bloch
see
pattern. This arrangement,
primordia developing
on
hairs
to
cell
a
has reached
further division is
region,
bution.
According cells
bouring
divide and
to
comparable
a
regular
that of the
to
of
a zone
very
leaf
towards unlimited growth
certain distance from the
a
this
Beyond
possible.
not
inhibition
regular patterns of
to
in several
Biinning
to
in
epidermis
tendency
suppress any
meristemoids originate and give rise
new
is
says,
surrounded by
are
in the
develop
Biinning
as
Thus, until
vicinity.
in their
or
the shoot apex.
These meristemoids tend
ristematic
The meristemoids
1939).
inhibition so that the stomata
these may
instances,
differentiate into
subsidiary
stomatal distrithe
cause
(cf
cells
me-
zone
neighin-
title;
next
formation available). idea of an “inhibitionzone”
Although Biinning’s of stomata and
ment
Millingtonia Vicia
a!.
et
in
arranged
about division of the as
of
of
stomata
contiguous
cells
placed
1
also Esau 1965b
;
together by
the
cells
subsidiary
the
Although
Yarbrough
calycinum,
(Basellaceae)
it
divide
transversely
It is
interesting
becoming
the
either into
the
such
1
According
of 2
lo Esau
works
vascularization
Pant
(1965)
sequences
a
due
to
of
of
bring
(1951) observed grown in
faba
Hence the
an
occurrence
protodermal
cells
not
Iatis
see
any
tinctoria 1
or
division of the
(Cruciferae) of the
2
cells
the
by
Running
cells.
influence extended by
cells
which
seen on one
(1965b) conclusions
of
are
side
Basella
1969).
cells differentiate
Blinning (1952)
the
(1960)
guard-cell-mother-cell
to
divide. The effect
do not find
and
Jain
that differentiation of the
bilaterally (depending
Hagemann (1957), Reinhardt (1960),
rubra
cells sometimes
Stebbins &
stimulated
or
and
cells.
cells in
series of divisions before
a
1952)
of
subsidiary
neighbouring
subsidiary
series
a
subsidiary
1965a, b;
whereas the
the
or
family Crassula-
off
cut
support
Torrey (1957)
of
whether the
on
from the
ex-
the induction
in roots.
writes
“one
stomata,
Acta Bot. Need.
Contiguous
row.
in members of the
that
protodermal cell undergoes
epidermal
Vicia
behaviour of the
longitudinally (Paliwal
induction is either
perimental
did
also indicated earlier
adjoining
an
in
epidermal
cells is
subsidiary
or
of
meristematic and
become
been found that
that
a
Kropfitsch
seedlings
be
to
2
guard-cell-mothcr-cell
the
suggested (as
on
has
found
Zimmermann &
which may sometimes
ripening apples.
&
nucifera
1965) .
(1934)
Bryophyllum
in
in
as
sativum,
(Maheshwari Nelumbo
stomata were
stomata
fungus
peculiar
In 1866-67 Strasburger observed ceae
6
for the such
and under the influence of external agents
nature
of this
Pant
or
a
(Gertz 1919a, b).
off
both in
explanation
recorded 5
the attack of
ethylene given
calls for fresh
(see
guard
stomata
6
as
many
atmosphere
by
ula
1961) and the
Pisum
Pulsatilla albana where
in
arrange-
species
anomala.
Gnetum
japonica
regular
in
triplets
Paeonia
observation),
also
as
(1962)
also induced
are
stomatal
damascena,
the
explains
adequate explanation
tabacum (Wehrmeyer
of 5
groups
offer any
and
In fact in Lonicera
1968).
Bachmann-Schwegler stomata
not
stomata
personal
Nicotiana
1961),
(Gupta
does
hortensis, Nigella
faba (unpublished
Vasil
it
of twin
occurrence
frequent
hairs,
and
18(5),
is,
however,
sporangial
Oct. 1969
unable
to
meristemoids
explain on
the
the same
simultaneous
and
gradate
basis”.
659
G.
cells
subsidiary before
ever, it is
cells
obviously
cells
activity
B.
suggested
Development changes
metrical
mitoses,
between the also
which
occur
a
1966).
the members of the
basis
of the
of
syndetocheilic
families Acantha-
and
Theligonaceae
several
others,
initial. This
same
induction mechanism.
publication (Paliwal
inherent
the
that in
1967)
to retain its
capacity
of
development
ultimately
polarity gradient cells
daughter
two
Jensen
in the
How-
pair
a
In
such
meristematic cells
produces subsidiary
guard-cell-mother-cell.
During
-
the
on
an
about the formation of
after the other from the
one
be manifested
may
guard-cell-mother-cell.
guard ceils. Moreover,
earlier
an
and
sides)
duration by virtue of which it first
longer
a
becomes the
finally
in
meristemoid has
the
for
explained
the
brings
Magnoliaceae,
produced
are
be
cannot
view of this I
instances
both
or
nucleus of the
encountered in
stomata
Cruciferae, Labiatae,
subsidiary
and
one
either side of the
on
of
development ceae,
on
known how this induction
not
subsidiary
the
formed
are
after the division of the
or
S. PALIWAL
at an
The smaller cell
result in is
set
up
a
its
cell,
more
within the cell
During
asym-
differences
causing
& Biegert
early stage (Sunning fails
irreversible
less
or
specialization.
1953;
see
differentiate and remains meriste-
to
matic. That
there
out
the
by
considerable
occur
and
epidermis
the
during
study
of Shanks
(1965)
the formation of the
compared
nuclear
Galtonia candicans
on
theprotodermal cell)
place during
production
that the stomatal mother appearance
mother-cell)
has
has
to form
equally
cells divide
a
a
large
a
with the
unequally
and
nucleolus,
small nucleolus. These
simple
clearly
stoma
with 2
to
the
brought
He has also
(Liliaceae).
the asymme-
He found
produce cells quite
unlike in
smaller
larger product,
distal
cell
the
(guard-celllater
guard-cell-mother-cells
cells. The latter becomes
guard
take
mitoses which
Ipheion uniflorum.
identical. The
genetically
leaf
differentiating
been
symmetrical
of the bulliform cells in
although presumably
epidermal cell,
the
stomata has
guard-cell-mother-cells (produced by
trical division of the
in
changes
formation of the
divide
speciali-
zed in form and function. Nuclear
occurred
changes
vacuolation and
tion,
of the
nuclear size of all cell types took
ovoid
spherical
or
cubical
to
with the increase in DN
exception, that
DNA
high
as
as
where the cell
cells grew very
usual
amount
when in
resulted,
ploidy
elongation
guard
of
rather
like
be
may
with
when cell
change
a
more
up
to
the
epidermal
10 times the
uniform in
Where
mature
in
shape
are
differentiation. It
during
found in
length
mitoses
failed, with
guard cell,
cells
original
and had
and DNA and their nucleolar volume
single,
elongathe
An increase in
from
associated
and nucleolar volume. Almost without
extent
likewise
for mitoses. a
taking place.
pyramidal, and these changes
or
were
growth period
were
place, frequently
to some
was
little,
protein
preparation
wall
A, nucleoprotein,
increased
20n
the
throughout
growth
a
was
of
volume. The
nearly
was
more
diploid
noted
Galtonia,
twice the
also doubled
elongated cell to
tetraploid
values.
Guard cells grew from
shape, 660
were
pulled apart
a
to
length of form
a
16 [j,
to
40
p,
the pore, while
at
maturity. They changed
wall thickness
Acta
Bot.
in
had increased
Need. 18(5),
Oct.
1969
ONTOGENY
STOMATAL
and 10
chloroplasts and
[x,
AND
were
DNA also
volume. The
appeared
to
material,
or
PHYLOGENY I
of
rate
more
both.
It
nucleolar material
there
and
growth
be associated
Nuclear size
numerous.
increased,
with
increased from about 6
level
of
to
little increase of nucleolar
being relatively
reached
polyploidy
in
the initial supply of nucleolar
these
cells,
cytoplasmic
or
found that those cells with larger initial supplies
was
nuclear
gained
and DNA
protein
more
and
rapidly,
of to
grew
greater length.
According
Shanks, the
to
ciated with
its initial
cells have
larger supply
a
only finally reach rate
than their
the
is
phenomenon of
cells Shanks
subsidiary
suggested by
as
in the
polyploidy
asso-
Epidermal
guard cells. They
also
does
develop
not
Stebbins & Jain cell
epidermal
be related to the may
formed)
a
at
not
faster
at an
of
development
to
seem
agree
to
but feels that
(1960)
early stage (before the
simple stomata, lacking
cells.
accessory
4.
but
level,
be
paired partners (epidermal cells).
development
stoma
or
to
appears material.
cytoplasmic
guard-cell-mother-cells
higher polyploidy
About the formation of the the induction
of the cell
development
of nucleolar and
than
much
a
of
rate
supply
AVAILABLE
INFORMATION
ON
DEVELOPMENT
STOMATAL
IN
MONOCOTYLEDONS
Campbell
vulgaris.
(1881)
In
surrounded five
or
the
an
elongated.
unequal
and
enlarges.
A pore In
the
of
consists
in
stomata
Tradescantia
semilunar guard
two
lateral) subsidiary
two
two
lateral and Next
cells.
cells.
develops
Zea mays
they
as
only
two
the
stoma
mature
is
septum
mother
are
a
stoma,
cut
cavity
a
some-
off from
down
cell) dividing
and the air
subsidiary cells
are
laid
cells
Occasionally,
becomes
two
cells
polar
two
vertical
a
of the
smaller
The
guard-cell-mother-cell (also
guard cells.
of
ontogeny
stoma
division.
Meanwhile,
protodermal
adjacent
stoma
the
each
cells may be found. At the time of initiation of
six subsidiary
centre of the two
leaves
by four (two polar
cell undergoes what
investigated
mature
in
the
it into
beneath the
produced
from the
adjacent protodermal cells. Porterfield (1937) studied the
development
of the
epidermis
Arundinaria quadrangularis. The protoderm pubescens and is
sheaths
cells. The to
the
dense
form the
cell and
longitudinally
The observations confirmed clear
by
to
Flint & Moreland
that in Arundinaria,
initial but from may
Acta Bot. Need.
cells
form the
the
the surrounding
thus be
18(5),
cells. The
1969
having
a
large
guard-cell-mother-cell
next
itself
guard cells.
(1881)
and
(1946)
in
Porterfield
Saccharum
Saccharum,
guard cells,
epidermal
quite misleading (see
Oct.
long,
off lenticular segments which lie
cut
Phyllostachys, to
broader than
mostly
Phyllostachys
of these cells function as the stoma-mother-
subsidiary
of Campbell
cells, although lying parallel
maturity
cells,
cytoplasm. Some
adjoining protodermal
guard
divides
of small
composed
nucleus and
in
of the culms and leaf
do
and
not
cells and
(1937)
officinarum. Zea,
the
have
been
Thus it is
subsidiary
arise from the stomatal
that
their appearance
also Ziegenspeck
at
1944). 661
G.
Working of
zone
cells
dividing
cells.
small, cytoplasm-rich Stebbins &
cells
the
nucleus then divides
divisions also
divisions
of the
vicinity
A similar
cell-mother-cell;
two
or
becomes
of the
more
the
development
of
of
triad
a
cells.
In
and
the
Commelina
subsidiary
vulgare
cells.
The
cells in the
Shah & Steb-
by
and other members
formation of the
(a)
cells
of the
consisting
division of the
(ii)
Later,
in certain
distal end
conducted
main steps:
subsidiary
zone
formed in Allium.
are
was
in Hordeum
(I960)
wide
cytoplasmic gradient.
guard
two
cells.
Commelina,
the
across
mm
cells and
guard
the
at
3
a
Cells of this
surrounding epidermal
cells
subsidiary
cutting-off
cells; (d)
polarized
form the
to
mention five
They
appeareance
subsidiary
form
to
and result in the formation of
cells. No
(b)
young
mitotic figure oriented
of stomatal
of the Gramineae.
cells; (c)
in
guard
study
latter divide
and Stebbins & Shah
(1959)
bins
a
asymmetrical
are
The
distal divides
occur
leaves.
produce (i) large, cytoplasm-poor
to
cytoplasm
by
cells the
two
base of the
also observed that in Allium and
(I960)
Jain
protodermal
Of the
at
occurs
differential divisions
undergo
the
found that
(1953)
Allium cepa, Bunning & Biegert
on
S. PALIWAL
the
by
lateral
guard-
epidermal
guard-cell-mother-cell
guard-cell-mother-cell;
and
and
(e) completed
stomatal complex of four cells. Some abnormalities in the formation of
subsidiary
two
stomata; and
of
presence
cell flanked
epidermal
cells
of
pair
a
by
a
of the
organization
the
cells
subsidiary
extra
on
the
four-celled complex
side of the
same
include
the stomatal apparatus; twin
adjoining
and
guard-cell-mother-cell
guard-cell-mother-cell;
a
undifferentiated
short
cell.
large subsidiary
A detailed study of the stomatal ontogeny of 4 unidentified species of Pandanus has
been
Pfitzer
made
innermost
next
below
the
hypodermal layer, by
by
once
confirmed the
earlier work
different from the
The
above
guard-cell-mother-cells a
cells of the
neighbouring
wall which
but
chamber,
the
produces
file.
Such
divisions
guard-cell-mother-cell produce never
the
synchronous
and may
which delimits
division
each side of the
in
the terminal
occur
early
lateral
or
of
Development
stomata
does
different stages of
at
general,
however,
divisions
not
which
but
lateral
follow
produce
are
cells.
subsidiary
development
situated
at
a
in
a
relatively
guard cells chronous
so
Another
662
wide
are
region
usually
that
point
only
are
last. one
completed second;
division
figure
of interest is that the
per
a
small
area
subsidiary
single
division in
to
is
the
as
the
divisions
are
before
files
on
cells
occur
complex usually
and
of the leaf. In are
are
com-
throughout
and divisions which
stoma
of
acropetal succession, a
terminal
Divisions within
function
pole
cells.
pleted first; divisions producing lateral subsidiary cells, which in
not
belonging
subsidiary
strict
occur
Transverse
usually completed
Cells
divide
They
each
subsidiary cells. These
late,
guard-mother-cell produce
stomata
cells
are
other-
not
are
guard cells.
divisions may continue in those cells of the stomatal file which do
guard-cell-mother-cells.
of
of substomatal chambers
development
position immediately
longitudinal
a
the
guard-cell-mother-cells.
their
cytologically
only
He has
(1965).
originate from epidermal cell-files directly above the
stomata
future
recognizable by wise
Tomlinson
by
that
(1870)
produce
rarely
syn-
seen.
guard-cell-mother-cell
Acta Bot. Neerl.
18(5),
Oct.
is
1969
SIOMATAL
AND
ONTOGENY
associated
with
further
internal
the stomatal pore opens, there is
soon
are
of the substomatal
hypodermal layer. Later,
communication with the internal
a
cells
guard
as
enlargement
within the second
by separation
occurs
As
development.
but before the stomatal pore opens,
produced, chamber
PHYTOGENY I
when
atmosphere
of the leaf.
According main it
to
Tomlinson this
types recognized
resembles the
ment
by
(1931)
This
mode of ontogeny has
the
available
clearly brings
ONTOGENETIC STUDIES STOMATAL
who
have
often
may
first
a
the file as
also
also
(see
VERSUS
more
compared
One
point
al I
two
to
sah
Pant’s offers
which
cannot
be
sound
employed
remedy
and
for
cells
borne
cells
originating
Neerl.
lateral subsinext
18(5),
cells
those
appear
of
to
situation is
Oct. 1969
could operate in
664).
p.
1963)
in such instances
Further,
identical
leaving
related
to
evolve
consideration between such
when one
to
to
the
types of cells
the
category be
latter
only
for
the
the
guard
adjacent epidermal
simple
The gener-
hardly satisfactory.
koshika
and
investigations
this
guard-cell-molher-
designated
sah
Mesogenous
comparative
as
seems
precise
a
from
cells of the
special structural relationship.
the
parents
of
distinguish
Perigenous,
same
from
quite
worthy
of the first
of
pathways
(see
situation would be
monocotyledons
of the
several
complex
to
file
same
examination
parallel
saying, therefore, that
point
a
has
ontogeny.
the
have
basis for
Subsidiary
Acta Bot.
of the
A somewhat
ontogenetically
are
simply
Subsidiary
2
may
mode of
cell" for both
koshika
very
types
significant
(1965) terminology
a
that each
rather than those from the
involved,
should be able
we
those which
1
1
these
suggest that the accessory
dar
are
com-
complex
clear for members of the Gramineae
1961).
It goes without
most
“subsidiary
term
the
actually
development
stomatal
impression gained by superficial
an
divisions
attempt
The
complex
against
how
mature
of epidermal cells which is placed
& Vasil
particular
a
of view is that
stomatal
ON
stomatal
of the
have revealed
row
guard-cell-mother-cells
maturity
to
way
terminology.
of
picture
of the mature stomatal
about
only
plants.
monocotyledonous families, Centrolepidaceae(HAMANN
than
at
develop-
suggested
concerning
aspect
OBSERVATIONS
of cells in the
abundantly
table I)
Mahhshwari
organization
guess
wrong
from that
good example.
one
where
cell
the
in other
seen
being
the
bearing
this
on
far for this group of
so
hand knowledge
guard-cell-mother-cell,
the
alone
a
(see
originates
cells
diary
provide
works
where several
the
to
COMPLEX
This has been made
developed.
although
two
structure
truly syndetocheilic (mesogenous)
that
would readily agree that the arrangement
plex
in
that in Tradescantia as
information
been recorded
not
MATURE
those
gymnosperms,
not to
out
ACTUAL
All
neither of the
to
(1961).
summarizes
I
monocotyledons.
5.
in
described in Juncus and Sagittaria,
Table
corresponds
stoma
type. Further, it is almost similar
amphicyclic
Stebbins & Khush
by
type of
Florin
on
2
.
as
saho-
Although
Meso-perigenous dicotyledons,
reason
that
the
it
leaf
cells.
cells.
663
G.
development terms
to
be
in the
two
employed
It is well established
guard to
a
this ceae
664
cells
can
and do
cell associated
groups is very different. I
in
distinguishing
now
vary
with
a
that the
of
As
are
such it is as
a
provided
not
always
subsidiary within
a
new
monocotyledons.
configuration of cells lying adjacent
guard cells,
variation in the monocotyledons
PALIWAL
have, therefore, suggested
the various types in
considerably.
pair
S.
cell.
easy
the
to
to
refer
Examples
of
family by Philydra-
(Hamann 1966) and within genera by Oryza, Pandanus and Philydrum (see
Acta Bot. Neerl.
18(5),
Oct.
1969
STOMATAL
also
ONTOGENY
might
1
PHYLOOENY
Arrillaga-Maffei
different types the
AND
It
1966).
seems
original pattern is completely lost,
probable,
very
in the
developed
have
same
way
to
a
type. In
particular
in
stomatal complex
monocotyledons
is still
FOR
COMMENTS
FUTURE
that has
The brief survey
ledons. A survey of the
are
a.
been
offer
a
few
a
cells
the
final
during ontogeny
difference in the the
one
other ( Ipheion
organization
hand The
).
level of
topic These
investigations.
complex
only author
The
Pfitzer
of
an
(1870)
becomes the Tomlinson
an
who has
intercellular
(1965)
linear
acropetal
provide
continuous et
relatively Acta Bot.
a
this aspect
the
cells.
hypodermal and
(1965)
that
of cells
The
provide
a
grow
uniform
28(5),
as
length
of
compared
Oct. 1969
the
with
guard the
isolated detail is
the meristemoid
subsequent-
Tradescantia).
cell files exercise the
by
one
of material.
generally
intercalary
development leaf. The
in
of
by intercalary
source
an
some
position
leaves which
cells
the
an
This
perhaps
monocotyledonous
on
some
not
in
1881;
suitable
series is usually present in the
layers
is another aspect
position of
Campbell
epidermal files
of
mind that the proto-
convenient material for studies of stomatal
more
Need.
-
hypodermal
underlying
sequence of cell divisions and
developmental
al.
plants,
degree
tissue.
bear in
to
the
by
purpose such leaves which grow
rows
Sequence of differentiation
Dunn
some
space in the
argues, in Pandanus the
meristems and possess
tem
to
versus
that the
unescapable
meristem within and
substomatal chamber (see also
the meristemoid itself. For this
an
always
cells
bulliform cells
versus
influenced
of the
that there
(ii)
epidermal
cell
epidermal
attention
paid
the arrangement of the
on
have
have
(1965).
supply
and
subsequent development
extensive
who found that in
is determined by
control
layer
initial
cell,
a
seems
The influence of
-
the
between
greatly
reached in the various cells of the
derm is the surface
of
epidermal
is
significant
two
of Shanks’ study
result
ploidy and
others
Among
-
a
conclusion, therefore,
of the stomatal
Influence of underlying layers
entity.
as
development
(Galtonia)
which deserves consideration. We
c.
for future
correlation between
some
initiation of the meristemoidand its
As
of this
personal experience
remarks
general
in monocoty-
development
stomatal
on
literature and
material and
cytoplasmic
on
polyploidy
ly
draw any
to
us
it
the
the foregoing pages clearly reveals
in
presented
have emerged
to
me
that there is
(a)
are:
marked
guard
to
appear
nuclear and
b.
to
me
current
of polyploidizalion
Effect
These
on
of
development
allow
to
assigning
summarized below:
points
is
in
WORK
the need of detailed and extensive work
have enabled
little
too
and
completed
difficulty
about the
knowledge
our
real
a
these often
sound conclusions.
fundamentally
6.
fact
a taxon.
after cell division has been
elaboration of the complex has taken place, posing
that
Very
therefore,
within
merissince
a
findings
of
monocotyledons
is
dicotyledons
can
be
explained 665
G.
this
on
basis
Araceae, appear
without much
follow
to
this variation in
d. Formation indicated
clearly brought
division of the
unequal
identified
an
the
to
Jain
The
do
not
significance
of
cells
and Shanks
meristemoids
cells.
compared
As
-
been
has cells
which
meristemoids. The reports
(I960)
protodermal as
Agavaceae,
devoid of such
are
Liliaceae members the
its smaller size
by
”
subsidiary
relation
as
question.
plants
Stebbins &
elongated
such
guard-cell-mother-cells
open
“
and
developmental
that in
out
the
acropetal development.
monocotyiedonous
Biegert(1953),
several families
others,
the merislemoids
special
may bear any
few
studies is
comparative
earlier
In
difficulty. a
strict sequence of
a
of
of Running &
easily
and
Philydraceae
S. PALIWAL
A
its
to
formed
meristemoid
sister
cell.
have
(1965)
are
may
by be
Suggestions
have been made that the guard-cell-mother-cell (meristemoid) may influence the
subsequent ability mechanism”.
1969),
of the associated cells
Although
suitable
explanation
bers of the families where
ceae,
a
this has
provide
Agavaceae,
ordinary
subsidiary
of “induction
are
Such
none.
Inamdar
1967;
exhibited
Pandanaceae,
by
whereas in
developmental differences
of Stebbins Khush
mem-
Philydra-
would
rather
ultimately
plants.
are
investigations
required
advocated
approach
conclusions it is cases
complexes
Bromeliaceae, Palmae,
interpretations
careful and detailed
speculations (?) tious
kind
some
(Paliwal
useful guideline regarding the distribution of stomatal types in
more
monocotyledonous Thus
by
earlier
large number of associated divisions take place {table I),
“phylogenetic”
a
divide
to
negated
is needed for stomatal
members of the Liliaceae there than the
been
necessary
to
by to
the
meet
follow the
cells,
holds
good
and
1924,
that
in order
the
fully
draw
to
agree with
cau-
definite
since in
sequence,
guard cells
not
stomata,
In this connection the
developmental
to
1
monocotyledonous
challenge.
Parkin in
epidermal cells parallel still
on
some
simulate the may
true
him.
ACKNOWLEDGEMENTS
I wish
to
express
ment and
to Dr.
from
Lalita
Miss
my
gratitude
Manohar
Lai
Kakkar and
to Professor
for
a
B.
M. Johri
critical review
Miss Kanan
Nanda
of
for his the
during
keen
interest
manuscript.
the
The
preparation
and
encourage-
help
I received
of this paper is
also
gratefully acknowledged.
REFERENCES
Arrillaga-Maffei, B. J.
Benecke,
W. E.
Sunning, —
R.
(1966):
Anomalous
stomata of
Stipa neesiana (Gramineae). Canad.
Bot. 44: 845-846.
(1892): 1952.
& F. Biegert
Die
Nebenzellen
Morphogenesis
(1953):
Die
in
Bildung
der
Spaltdffnungen. Bol.
plants. der
Surv. biol.
Progr.
50: 521
Ztg.
et
seq.
2: 105-140.
Spaltdffnungsinitialen
bei
Allium
cepa. Z. Bot. 41:
17-39.
Campbell, Amer.
D. H.
Nat.
Drawert,
H.
(1881):
15;
(1941);
development
Beobachtungen
Tradescantia virginica
666
On the
of stomata
of
Tradescantia
and Indian
corn.
761-766.
L. Flora
an
(Jena)
den
135:
Spaltöffnungen
und
den
Blatthaaren
von
Oct.
1969
303-318.
Acta Bot.
Neerl. 18(5),
STOMATAL
ONTOGENY AND
D.
Dunn,
G. K. Sharma & C. C.
B.,
Amer.
monocotyledons. Edwards,
W.
Biol.
—
K.
Campbell
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