ACTA
BOTANICA
Secretion
The
Vol.
NEERLANDICA,
of
the
(3),
4
Salt
1955
of
Glands
Ktze
Limonium latifolium
BY
W.H.
H.
Arisz I.J. Camphuis
The
(.Botanical Laboratory, Groningen,
Tooren
van
Heikens A.J.
Netherlands)
(;received May 27th, 1955)
Introduction
In
a
found
great
in
number of
the
leaves.
Frankeniaceae and
as
Spartina,
glands
in
some
genera
the
(1910)
salt
glands
rich
in
in the
glands is
of the
glands
Vacuoles either do
not
On the outside the
membrane with
each
is
a
are
bounding
cation between
is
(the cupcells)
cells
are
and
on
of
complex
in
extrude the
present,
bounding
the
off
a
at
found
were
on
644
or
with
some
on
the
glands
are
a
by
The
sq.
cm.
small size.
Cutinized
(P),
one
in
the fluid emerges. The pressure
cytoplasm through
which closes
the
adjoining
gland
on
cells
in
the
since
the cell
of the tissue there
mesophyll.
of these
passage spots
pores,
all sides. Four
the side of the
means
these
opening
outer
collecting
Communi-
gland cells
collecting cells.
These
lying in the cutinized walls
gland.
isolated
Schtscherback’s
glands
on
water
the conclusion that under
observation
are
capable
fairly high
that
higher
than
the
NaCl
pressure
concentration in
leaf-
of secretion.
the
fluid of which the sodiumchloride concentration is
somewhat
com-
upperside
a
four pores
are
a
1).
per
but of
covered
gland cells is
places
of
large nuclei (Fig.
gland cells
and the
only possible by
segments and also
give
gland
corroborated
Ruhland
He arrived
the
Townsendii.
Spartina
with
which there
At these
gland cells
of
the underside
in the
of
were
Skelding and Winterbotham
There
/x.
leaves
Statice Gmelini
glands of
mesophyll and the cytoplasm of
provided
the
demonstrated salt
of Statice Gmelini consist
cutinized wall which encloses the
cells
or
and
kind of membrane is
wall. Between the
43-46
cuticle,
cannot
glands
occur
secreting cell (A).
in these cells
a
a
are
Tamaricaceae,
in excised
secretion
leaves
cytoplasm
diameter of the
leaf 722
the
studied
function of the salt
Ruhland showed that the
16 cells
glands
Plumbaginaceae,
of the Gramineae and
by Ruhland (1915).
fully investigated
plex of
the
are
( Avicennia marina).
Structure and
described
them
Aeluropus and Tamarix. Faber (1923)
Mangroves
Sghtscherback
Statice.
in saline habitats salt
plants living
Among
the
glands
equal
to
leaf cells.
THE
An
OF
SECRETION
THE
the
by
He
be
The
Barger method.
fairly large. This
1.
Fig.
Gland
secretion
LATIFOLIUM
After
organs
Pore;
Ruhland’s
any
of
B.
Our
sq.cm.
150
leaflength
mg.
Ruhland’s
a
short
laboratory
A.
(iStatice
hot
used
plant
for
Limonium
house
were
J.
I.
some
salt
being
the
tissue.
from
hour
this.
hours.
been
each
1946, Miss
working
H.
Heikens
obtained have
observations of my
Limonium
potplants
autumn
kept
in that
own.
Ktze
latifolium
showed secretion
and
repeated
students,
in
was
per
with
correspond
have
some
fully
been
not
attention,
1954. The results
In
which still
used,
in 24
J. Camphuis
researches
epidermal cell;
of 0.287-0.861 mg
younger
together with
these
deserve
removing
experiments
by
pseudolimonium).
also
mg
D.
glands have
latifolium
Tooren in
van
here
been summarized
70
secretes
Ruhland, slighdy altered.
to
mesophyll.
liquid
Limonium
period, Miss
1950 and Mr
The
for
fundamental
extended in my
during
values
concen-
organic substances.
collecting cell;
salt
in
not
salt solution
he sometimes found
content
however,
efficiency
of
C.
F.
the
do,
They
secretion
a
cells;
cup
publication
more.
found
according
adjoining cell;
extraordinary
Ruhland
per
gland he could
be due to the presence of
must
323
KTZE
difference between the osmotic
of Statice Gmelini
P.
cell;
examined
in
LIMONIUM
gland and the NaCl
E.
A
OF
determined the osmotic value of the secreted
tration of the sap of the
A.
GLANDS
increase of the chloride concentration in the
prove.
to
SALT
in
a
season.
Method
If
a
leaf
or
leafsegment
leaf and
it
will
whether
evaporation
watervapour
a
change
in
secretes
therefore
from
of
depend
water
the air takes
a
liquid,
on
from
place
the
the
on
drops
3
mm
height
were
remain
tension
or
of
on
the
the air
condensation of
the leaf. In order
concentration of the secreted fluid
inside diameter and
drops
the vapour
glass rings
to
of
fixed with Leicks’wax
prevent
21
on
mm
leaf
324
disks 3
cm
obtained
a
H.
in
diameter, punched from the leaf.
by fixing
slight
leaf
same
J.
glass plate
a
H.
CAMPHUIS,
Leafdisks
degree.
the
on
3
in
cm
A closed chamber
with vaseline.
ring
drops could
diameter
water
After the
The disks
compared.
was
salt
or
experiment
total
weight
which
HNOj in
AgN0
after
a
on
the
dish
a
a
sucked
was
dried and
weighed, and
weighing bottle.
The chloride
of
destruction
been
had
the chambers
previously
determined in
was
determined
was
lying
filter paper in
on
in
from
punched
solution.
the secreted fluid in
into filter paper that had been
up
content
put
were
way
only
occur
were
was
In this
either side of the midrib. The behaviour of disks
on
level
containing
the
HEIKENS, A. J. VAN TOOREN
in concentration of the secreted
changes
very
I.
ARISZ,
W.
solved.
the
with
filterpaper
Chloride
determined
was
3
by the Volhard method.
The determination of the osmotic
from
expressed
the
leaves
osmometric method worked
As
fluid
comparative
obtained after
was
hand press
By
leaves
in
used.
was
the
expressing
suitable leaves and
experiment
one
The
orientation about
the
be
influence of various
a
(1952).
sap
hydraulic
a
great variability.
the results
obtained
factors
sap
Johnson’s
expressed
leafdisks in
gathering
could
figures
fluid and
and
Van Andel
laboratory by
solution
and
killing
secreted
Baldes
by
definite pressure. The secretion shows
at a
choosing
our
in this
out
NaCl
a
value of
executed
was
of various
rendering
the
on
an
secreting
possible.
process
Pretreatment
Leaves
of
about 0.23
salt
proper
their
3
20
into
with
placed
after
hours,
In
6
of the
cut.
is taken
In
on
secretion,
fluid
so
leaf is
reduced,
which
can
This will
and
on
ment
in
cm
diameter
which
have
40
to
/ig and
are
have been for
which
and
a
with
after
next
Cl
fig
in
hours
22
the NaCl
through the edges
up
greased with vaseline, no more salt
it
now
that
in
increases.
and
In
owing
the
on
activity
that
If the
case
case
equal
NaCl
salt
no
the loss
to
second
the
secretion
secretion
of the
concentration
and
of
the
as
the
NaCl
leafdisks
takes
the
to or
salt
taken
up
on
0.72
0.08
M
during
absorption
is
continued,
weaker than the salt secretion.
So
M
fluid of 0.31
to
be
the concentration in the
concentration cf the external
leaf
secreted
amounts
of the
can
of salt
of the secretion process.
concentration
according
considerably smaller than if it takes place
the first
be stronger,
depend
the
appeared
in the pretreatment, the chloride concentration
water, the increase is
NaCl solution.
a
the
higher
secreted
on
obtain
beforehand
amounted
127
chloride,
to
up.
concentration is
place
little
order
take
If the latter have been
orientating experiments
of the
fluid
pretreatment
265 /ug per 20 hours. The leafdisks
3
leaves
NaCl,
M
0.25
the secreted
hours’
disks
from
In
placed
are
Next
disks
in
secretion.
leaves
solution.
petioles
water,
a
no
but
contain
garden
show
intact
leaves.
their
on
the
salt
a
the
the
They
%.
secretion
from
hours
been
0.37
petioles
punched
from
plant
a
to
on
we
found with
NaCl
M;
this
0.51
M
solution
rises
on
NaCl.
solution
pretreata
a
NaCl
higher
When the
THE
secretion
SECRETION
took
fluid
secreted
fluid
proved
concentrations of the
of low
concentrations
leaf
was
because
found
were
from
water
chambers
on
the air
concentration
high
respectively.
concentration of the
Without
important.
fluid
in
change
a
the concentration
M
0.23
the
325
KTZE
LATIFOLIUM
water,
on
0.16 and
to
prevent
be
OF LIMONIUM
external
M)
low
too
solutions
condensed
0.42
(over
found without chambers, because in this
were
from
to
to
of
solutions
evaporated
the
secreted
concentration,
On
drops.
GLANDS
amounted
of chambers
use
SALT
place while
of the secreted
The
OF THE
the
on
high
too
case water
drops.
TABLE I
Influence
of
pretreatment (24 hours)
on
the
secretion
at
25°
C
hours.
during 24
1951.
in
M
NaCl
%).
(0.5
0.171
„
„
(1.—)
„
„
(2.—)
„
„
(3.—)
„
„
(4.—)
0.342
0.513
0.684
.
.
.
.
.
.
.
.
.
Cl
M
NaCl
„
„
„
0.342
0.513
Cl
cone,
secreted
fig
in leaf
mg
on
0.47
M NaCl
53
931
0.49
0.17
31
381
0.34
0.18
40
656
0.46
0.28
31
596
0.54
0.39
171
0.35
0.16
13
0.171
„
in
fluid
Secretion
0.085
cone,
M
secreted
Secretion
0.085
M Cl
quantity
concentration
NaCl
during pretreatment
water
on
48
134
0.08
0.08
41
124
0.08
0.11
60
299
0.14
0.14
42
329
0.22
0.22
37
278
0.21
0.31
„
0.684
„
Table I
the influence of
gives
tions NaCl from 0.5
NaCl
(0.47 M)
is
or
to
a
pretreatment
on
various
%, secretion taking place
water.
on
Both the
optimal.
4.0
A
pretreatment
2
on
on
these
water
chloride concentration of the secreted
0.47
fluid is
M NaCl
the
treatment
in
ments
25° C
on
The
time
It has
so
has
an
place
a
in
after
In
pretreatment
on
effect
the
been standardized
artificial
already
up
takes
place
When secretion takes
place
of
light
the
at
20°
much
or
secretion
much chloride
amount
at
25° C
of NaCl
that
the
M) NaCl. So there
amount
as
for
place
been discussed that
an
% (0.51
subsequent
as
took
3
of the
external solution of 0.47 M
course
taken
fairly equal.
pretreatment
distinct
pretreatment.
1950
took
having
be
to
are
NaCl
fluid and
the concentration both in the leaf and in the secreted
after
greatest
appears
during
quantities
2.75 %
on
% (0.34 M)
that of the leaf have been determined. When the secretion
on
concentra-
either
experiments
possible.
24
hours
NaCl. In
25°
taken
the
In the
in
1954
0.34 M
on
of salt
the
up
pre-
experidark
at
pretreatment
NaCl.
process
a
leafdisk
during
Cl
the
placed
on
water
pretreatment,
concentration in
after
secretes
the
leaf is
326
W.
H.
reduced
considerably
H.
ARISZ, I. J. CAMPHUIS,
concentration in the
Cl
concentration
Pretreatment
a
fall in Cl
fluid.
0.42
II
secretion
during
on
VAN TOOREN
II). This is accompanied by
(Table
secreted
TABLE
in
Changes
A. J.
HEIKENS,
M
in
NaCl.
leaf
and
Secretion
M Cl
on
in leaf
cone,
fluid.
secreted
M
Cl
cone,
secreted
At
the
24
hours
beginning
pretreated
22 hours secretion
44 hours
secretion
66 hours
secretion
on
water
on
water
on
water
.
.
.
0.06
.
.
0.25
.
.
.
.
.
.
.
.
.
.
.
time
.
of secretion
course
24
0.17
0.15
0.16
0.06
0.13
0.06
.
on
Secretion
III
salt solution
a
25° G
hours, dark,
0.47
on
and
M
on
1950.
water.
M NaCl
0.47
on
Cl
quantity
first
.
day
.
second day
third
day.
.
.
Secretion
NaCl
M
/ig
22
517
0.66
28
324
0.32
,
25
508
0.58
31
196
0.18
.
.
.
22
410
0.53
32
76
0.06
cone,
gives the
at the
beginning
at
the
end
at
the
end
on
were
had
a
water
in the
the
much
was
amount
M
.
0.39
M
.
.
0.14
M
place
during 3 days.
0.47 mol
on
the salt
on
0.59
from
The
to.
successive
more
The
M.
0.39
the
The
Cl
average
of secreted fluid
Cl
reduced, from 0.59
of secreted fluid remained
content
to
of the
0.14 M.
fairly equal
the
day
with leafdisks
salt solution. The concentration of the secreted fluid
greatly
the end
present
on
reduced,
of this
in
significant,
the
from
leaf than
0.32
the
not
given off by secretion. Such
days,
was
not
already
besides
found in
reduced
most
on
has been examined
and
on
water.
The
a
secreted
regular
during
2
amounts
or
NaCl.
salt
It
seems
day.
3
on
this
of secretion
salt
Usually
solution
the
that
concentration
If
fluid.
rate
same
experiments.
the second
M
higher
experiment
rate
at
was
difference
all chloride present in the leaf
the
nearly
0.06
to
slightly
a
in
it indicates that
which is
slightly higher
was
experiment
than in the
Also
on
2nd and the 3rd
was
and
solution, the Cl
amount
days.
During
another
NaCl,
to
concentration than
concentration of the leaf amounted
fairly equal
0.59
.
placed
decreased
Cl
higher
Cl
water
When secretion took
fluid
in leaf
Na
on
of the secretion
course
secreted
case
cone.
.
of the leafdisks
in this
M Cl
fig
mg
.
water.
on
in
.
concentration
leafdisks
water
.
Table III
remained
Cl
quantity
Cl
cone.
mg
the secretion
of the leaves
part
on
on
(6 discs)
.
Cl
part
Pretreatment
solution.
(4 discs)
in
in
fluid
.
TABLE
The
1946.
water.
can
is
be
during three
as
on
water,
of secretion
is
In Tables IV and V the secretion
days
for leaf disks
of fluid exuded
on
on
salt
0.47
M NaCl
solution and
on
THE
water
SECRETION
secretion
SALT
and
the second
concentration of
the
third
the
secreted
being different
secreted fluids
salt
on
salt and
on
TABLE
The
time
of secretion
course
on
as
Secretion
on
M
on
in
Cl
first
day
second
the
.
day
.
.
.
.
.
.
.
.
water.
on
gland
NaCl
that
interesting
secretion
the
to
has
Secretion
MCI
/ig
quantity
cone.
in
on
water
Cl
MCI
fig
cone,
mg
66
1444
0.62
67
652
0.27
48
950
0.56
42
183
0.12
In
the
of
took
experiments
on
salt concentration
nearly all experiments the exuded
It should be borne in
periods.
in
place
exposure
the
dark.
light
to
that in
different from
course
of the
independent
course
know whether
a
Pretreatment
(5 discs)
of fluid is reduced in successive
amount
mind
sap.
1950.
III.
mg
of secreted fluid is
amount
of the
the
This indicates that
water.
(4 discs)
quantity
of the salt
concentration of
NaCl
and
in Table
0,47
they
water
on
day. This reduction of the
IV
salt solution
a
327
KTZE
day is independent
the
fluid,
LATIFOLIUM
solution and
the second and the third
on
on
GLANDS OF LIMONIUM
but both
fairly equal,
are
smaller
are
THE
OF
It
the
is
therefore
of the
strength
the dark.
TABLE V
The
time
course
of secretion
22
on
hours,
Secretion
first 22
second
third
hours
22
22
.
hours
on
solution
0.42
0.42
/ig
M
and
on
NaCl
M NaCl
M
1946.
Secretion
Cl
Cl
quantity
in
cone.
mg
water.
Pretreatment
solution.
on
water
M Cl
fig
cone.
mg
.
67
929
0.39
71
.
38
542
0.42
37
198
0.15
20
336
0.47
16
29
0.05
hours.
.
The influence of
experiment
dark,
Cl
quantity
in
salt
a
light
on
383
the secretion appears from
in which after the usual
pretreatment
a
0.15
comparative
half of the leafdisks
exposed to the light, the other half put in the dark on 2 % NaCl
(0.342 M). The average amount of secreted fluid in the light per disk
are
is
50
0.544 M
mg with
in the
Cl,
dark
16
mg
with
0.520
M
Cl.
TABLE VI
of
exposure
to
light
secretion.
on
0.342
Secretion
in
24 hours
amount
in
mg
light
97.6
(4)
dark
64.1
(4)
In
a
between
second
experiment
light and
dark
M
1954.
Pretreatment
24 hours,
25° C
NaCl solution.
25° C
on
0.342
M NaCl
M NaCl
cone.
osmometric
det.
j
Influence
j
0.384
0.379
M NaCl
0.393
0.413
M
a
slightly smaller
(Table VI). During the
difference
was
NaCl
found
first 24 hours 97.6
mg
328
W.
secreted
was
with
mg
in
the
0.393 M
this
with
the osmotic
case
the aid of
osmotic
concentration. In
the
the dark 0.413 M
The
with
differences,
known
osmotically working
determined
was
the
the
in
small
isotonic
the
0.379 M
was
NaCl
NaCl.
In
with
quite correspond
not
from the C1 determinations.
computed
too
are
fluid
was
28.9 mg.
in
light the osmotic value
do
64.1
concentration NaCl,
expressed
NaCl. These values
however,
dark
34.75 and
dark,
secreted
the
the secretion
concentration of the sap
been
the osmotic value which may be
The
a
and in
hours
24
in the
value of the
obtained have
values
and
light
being compared
osmometer
the second
In
osmometer.
an
NaCl
with 0.384 M
light
NaCl.
both in the
greatly reduced
In
H. HEIKENS, A. J. VAN TOOREN
ARISZ, I. J. CAMPHUIS,
H.
give
to
indication whether
an
substances other than NaCl
in the secreted
occur
fluid.
From
these
of secretion
experiments
is
Influence
promoted
of
The influence of the
vestigated
secretion
at
5° C and
5°
at
exposure
temperature
the
and
on
25° C
25° C of leafdisks,
at
C secretion takes
day.
solution
place
at
the tables it
From
the
amount
5° C
secretion process
(Table VII). Experiment
water, the other half on 2.75 %
on
secretion
the
on
day,
that when secreted
of secreted
fluid
at
25° C is
secreted
solution. In B and
25° C
at
on
the second
0.47
on
much
M
NaCl
greater than
5° C, but that the NaCl concentration of the secreted fluid is
at
the
at
two
in-
was
A shows the
half of which
one
(0.47 M) NaCl
the first
on
appears
rate
light.
to
the
temperature
at
conclusion that the
may draw the
we
by
equal
temperatures.
TABLE VII
Influence
of
temperature
secretion.
on
0.
A
1950
hours
24
Secretion
in
5° C
25°
B
C
0.47
dark, 25° C,
hours,
on
M NaCl
Secretion
amount
Cl
in
cone.
mg
on
water
M
Cl
Cl
ng
cone.
mg
6.6
(2)
132
0.56
13.6 (2)
268
0.56
58.0
(3)
1188
0.58
51.0
399
0.22
141
0.62
10.6
220
0.60
0.62
76.6
629
0.23
(3)
1950
first
sec.
day
5°
C
.
6.4
.
C
day 25°
951
43.6
.
.
Cl
C
on
24
solution.
M Cl
amount
temp.
Pretreatment
47 M NaCl
in leaf
0.47
M
after
the
two
days
on
NaCl
0.42
M
after
two
days
on
water
0.20
M
content at
beginning
1951
first
sec.
day
5° C
.
C
.
day 25°
The
salt
secretion
water,
297
0.60
22.0
320
0.41
1242
0.59
78.0
527
0.19
.
concentration
takes
the
14.0
59.0
.
place
on
in
a
the leaf is
salt
concentration in
solution.
the
but
slightly
If secretion
leaf after
the
reduced,
takes
second
day
when
place
has
on
got
THE
SECRETION
THE
OF
SALT
GLANDS
considerably reduced. At 5° C the
greater than
0.47 M NaCl
on
cells.
25° C
more
0.20
mol
The
of the
is
exuding
therefore has
and
sap
of
the
by binding
Fildes
and
The
secretion
the
anaerobic
on
hour.
The
M NaCl
the
fluid of 0.39
solution.
1.1
by
of
the added
to
mg.
of
gated in
1950.
arsenite,
NaF,
somewhat
the
blue.
Cl
fig
were
some
They
and
a
in
was
McIntosh
a
was
24
by
gas
oxygen
put
in
of
means
a
after about half
checked
by
of
means
with
hours’pretreatment
put in the
were
jar for 24 hours
on
mg
of
NaCl concentration in the secreted
the
not
secretion in
5 leafdisks
contain any demonstrable
ordinary air, but they
on
the
were
no more
weaker
a
were
and
KCN,
and
one,
NaF
variable
of inhibitors
lodoacetic acid
At
phloridzine.
give
whereas
distinct
a
azide
does
had
influence.
no
on
Secretion
secretion
not
24 hours
on
0.47
25°
C
1950.
on
0.47
5
X
IO“
x
10~
3
5
7.5-25
10-’
x
—
?
....
distinct
with
dini-
24
hours
dark
10
10'
4
6
solution
cone,
giving
of inhibitor
50
%
inhibition
M
7-8
X
IO" 3 M
M
2-4
x
10- 3 M
2.5-7.5
x
10-
x
10- 3 M
10 -3-5
x
10- 3 M
5
X
10- 4 M
Naazide
arsenite
results
Pretreatment
M NaCl
of inhibitor
0.25
NaF
a
inhibition of
M NaCl.
giving stimulation
I.A.A
The
give
an
concen-
arsenite
stimulation,
VIII
process.
cone,
KCN
low
a
investi-
(I.A.A.),
(Table VIII).
25° C
2,4-dinitrophenol
metabolism was
inhibiting
Naazide,
higher concentration they all give
Phloridzine
were
secretion
substances
TABLE
Influence
investi-
was
of the air
content
hydrogen
oxygen
After
a
2,4-dinitrophenol
I.A.A.
At
secretion.
trophenol
Cl
sap secreted.
Leafdisks in aerobic conditions gave 29
inhibitors
KCN,
stimulation.
the
of
hydrogen
free from
The secreted fluid did
The influence of
a
the
B from 0.47
secreting.
Influence
tration
in
the secretion
of
means
In the absence of oxygen
M.
Cl. Next the leafdisks
capable
result
a
the concentration
on
amount
on
oxygen
jar being
methylene
secreted fluid with 570
averaged
as
at
temperature
oxygen
0.47 M NaCl solution the leaf disks
0.47
this
in the
out
jar.
withdrawal of
reduced aqueous
the
Decrease in oxygen
oxygen
bound
was
oxygen
palladium catalysator,
an
at
off and
influence
no
only regulates
experiments.
some
obtained
the lower
flowing
considerably reduced,
The influence of withdrawal of
in
because
is
water
on
to
NaCl.
temperature
Dependence
gated
fluid
secreted
containing NaCl is given
sap
be due
must
329
KTZE
resulting stronger turgor
lower,
considerably
concentration in the leafdisk
to
the
LATIFOLIUM
of fluid secreted
amount
concentration of the
is
water
on
much
The
LIMONIUM
solution. This
osmotic value of the medium and
gland
OF
M
M
7.5
3
M
330
W.
On
H.
ARISZ,
of C1
amount
trations of KCN
fluid
and
Cl
correlation
J.
and
I.A.A.
that
the
given off by
0.91
does
affect the
rate
Fig. 2.
for
Influence
of
is
a
distinct
of
KCN
the
on
latifolium;
For
stimulation
at
the
fluid
under
that the
see
under the
From
-f- 0.96
to
this
it
follows
influence of
The inhibitors
liquid
amount
percentages
is
reduced
KCN
secretion.
on
1954.
therefore
the
only
KCN
Average value
fluid
expressed
shown
to
0.
M
the
on
the
salt
fluid
glands
expressed
by Fig.
KCN,
Table
2.
There
a
higher
at
IX shows the in-
rate
experiment
of secretion
ex-
without KCN.
IX
Pretreatment
in
by
unaltered.
of secreted
10~ 3
to
24 hours,
20° C,
0.342
on
0.342
percentage
NaCl
without
215
110
90
62
10
0
152
120
94
57
8
0
209
108
109
113
113
98
0.002
0.003
KCN
0.0005
191
0.001
M
of control
0.00025
189
M
light.
Secretion 24 hours, 20° C, dark,
of secreted
secreted
remains
X
concentrations
of the control
NaCl, exposed
cone.
we
quantities
amounted
concentration
TABLE
Quantity
KCN,
two
concen-
of secreted
amounts
+ 0.96.
0.25 and 0.5
KCN
of
of secreted fluid
KCN it
of
amount
the
on
fluence of various
Influence
the
of the control without KCN is
secretion
in
TOOREN
amount
the
to
without
in
Na-arsenite
concentration the
pressed
VAN
of secretion.
Limonium
percentage
the disks
great.
J.
certain time with various
significantly.
The influence of KCN
in
A.
HEIKENS,
of the secreted
alter
not
a
alterations
very
and
concentration
inhibitors
H.
these values
relating
between the
+
in
given off
influence of inhibitors is
for
CAMPHUIS,
the correlation between the
determining
and the
I.
0.004 M
12
60
10
0
THE
Relation
of
the
SALT
GLANDS OF LIMONIUM
sap
of
the
values
leaf
fluid which
on
is
have
water
on
equal
to
lower
or
salt solution have
a
the
of
external
the
than that
chloride
concentration of
from the fresh
does
strable
weight of
imply
not
chloride concentration
it
be
would
in cell
has
leaf
that
cytoplasm
in
the
know
To
whether
and vacuole
computed
of the
leaf
demoncells
this
concentration
chloride
high.
A
comparison
concentration in the leaf and in the secreted fluid therefore
only
limited value and
a
concerning
conclusion should be
no
accumulation in the
salt
possible
a
TABLE
Relation
the
that
phenomenon.
same
analytically
equally
was
Leaves
impression about this,
an
get
leaf.
quantity
a
the chloride
to
necessary
being
the leaf and
really exists.
wall, cytoplasm
of the Cl
the
the secreted
concentration in the
secreted fluid than in the leaf. Table X shows the
The
fluid
that leaves
appears
of the
higher chloride
a
solution,
secreted
concentration in
331
KTZE
LATIFOLXUM
and VII it
chloride
a
of
and
the data of Tables I, II, III
secrete
secrete
THE
OF
between the osmotic
expressed
From
that
SECRETION
between
the
concentration
Pretreatment
X
of NaCl
24
in
leaf
and
hours, dark,
0.42
M NaCl.
Secretion
drawn from it
glands.
on
0.42
in
secreted fluid.
1946.
M
Secretion
on
water
NaCl
in
M
Cl
in
cone.
secreted
in secreted
leaf
in leaf
fluid
at
the
fluid
0.06
beginning
0.06
....
after
the
after
22
hours
0.30
0.39
0.17
0.15
after
44 hours
0.28
0.45
0.16
0.06
after
66 hours
0.23
0.33
0.13
0.06
pretreatment
0.25
.
.
0.25
An osmometric determinationof the secreted fluid
a
result
good
with
In table VI it has
the
secreted
expressed
as
aid
of the
can
(secretion
0.34
on
NaCl
isotonic
osmometer
M
solution,
(Van
be made with
1952).
already been found that the osmotic value found for
fluid
an
the
NaCl
Andel,
solution
at
with the
corresponds
25°
C)
NaCl
concentration of the secreted
the dark and in
value of the
osmotic
It
determined. Both in
fluid analytically
light this value lies considerably higher than the
the
secreted
fluid also
external
desirable
thought
was
solution.
to
compare
with that of the
the
the osmotic value of the sap of the leaf rather
found,
but the osmotic
than that of the leaf
dence
fluid
between the
and
limits of
These
the
osmotic
divergent
value of the secreted fluid
sap.
In this
case
osmometrically
analytically
found
value
expressed leaf sap (Table
too
there
was
values
were
higher
great correspon-
determined value
one.
the
For
always
was
a
of
XI).
of the secreted
The differences lie
within the
error.
determinations had been
NaCl solution
(0.342 M).
of the secreted fluid
was
In this
made with
experiment
leaves
lying
the NaCl
higher than the NaCl
on
a
2
%
concentration
concentration of the
332
H.
W.
medium. The
with
ponds
osmotic
I.
ARISZ,
J.
osmotic
average
that
of
the
present in
solution.
in
higher,
depend
external solution. This will
Secretion
Leaf
of leaf
hours,
25°
determination
0.406
the
on
leaf sap
lower than
on
fluid
secreted
0.342
a
fluid.
M
NaCl
solution.
osmometric
secreted
M NaCl
0.463
NaCl
M NaCl
0.346
0.402
0.408
0.275
0.425
0.414
0.409
0.415
•
0.319
0.394
0.348
0.411
0.338
0.420
The osmotic
value of the
while this is
sap
evident
that
the
on
the
secretion
glands
of
equal
increasing
NaCl
+
99
Second
0.342
M
sucrose
„
outer
solution
by
to
light.
dark
Cl
cone,
m
ted
mg
secre-
fluid
37.2
771
0.58
24.8
593
0.67
21.8
655
0.85
22.2
457
0.58
16.9
380
0.63
11.9
398
0.97
M
„
period.
NaCl
+
99
To
0.071
0.142
+
99
M
99
NaCl, exposed
quantity
in
M
99
M
24 hours, 25° C,
period,
of the
1954.
secretion
0.342
work.
doing osmotic
osmotic value
0.342
of
periods
of
XII
sugar.
24 hours, 25° C,
Secretion 2
the
higher than that of
being
that of the external solution,
to
capable
are
adding
Pretreatment
fluid
secreted
about
TABLE
Influence
First
M
0.454
0.292
is
fluid
cone.
0.451
0.305
it
the
1954.
0.461
the leaf
the
that of
of other substances
amount
determination
0.411
Av.
corres-
experiments
some
XI
and
sap
C,
secreted
osmometric
sap
In
the sap.
cone,
24
J. VAN TOOREN
the others
TABLE
Osmotic
A.
HEIKENS,
concentration of the
external
concentration is
than NaCl
H.
CAMPHUIS,
0.071
+ 0.142
99
investigate
solution
M
sucrose
„
the
„
influence
the secretion
on
been added
to
the
added
sucrose
the
outer
the
process
solution
or
to
the
may
be
supply
So
concentration of
(Table
of
the
due
the
sucrose
solution
amount
concentration increases.
clearly shown. It
of
XII).
secreted
of sugar
the osmotic
as
a
external
source
Under the influence of
fluid
influence of
to
the
of various concentrations has
the
is
reduced
sucrose
and
addition
its
is
increase of the external
of energy
or
to
both in-
fluences. In Table XIII the influence of the increase of osmotic value
of the
on
the
outer
solution
NaCl
through addition of higher sugar concentrations
concentration of leaf and secreted fluid has been traced.
THE
As
is
SECRETION
already discussed
OF
SALT GLANDS
THE
OF LIMONIUM
lower
a
remains
the
about
increased
of secretion.
rate
with
The
that
same,
in
of
NaCl
the
the greatest addition
TABLE
Influence
increasing
the
in
Pretreatment
osmotic
value
leaf and
24
fluid
M
hours, 25° C, 0.342
25°
24 hours,
solution
outer
fluid.
99
99
0.342
„
„
MNaCl
„
„
+
0.150
„
+
0.250
„
M
NaCl
cone,
NaCl, exposed
light.
to
dark
C,
Cl
cone,
M
Cl
cone,
in secreted
in
mg
leaf
fluid
31.9
0.418
0.336
15.4
0.378
0.318
11.8
0.501
0.324
sucrose
„
the
on
1954.
M
99
considerably
XIII
of the
in
on
is
sugar.
quantity
99
Addition of sugar
cases.
concentration in the leaf
secreted
of
in secreted
Secretion
Secretion
333
KTZE
above the NaC 1 concentration in the secreted fluid
considerably higher than in the leaf in such
gives
LATIFOLIUM
„
TABLE XIV
Influence
on
the secretion
of
increasing
adding
Pretreatment
28
hours,
25°
Secretion
C,
24
the
value
osmotic
0.342
hours,
of the
solution
outer
by
1954.
sugar.
M
25°
NaCl, exposed
to
light.
dark
C,
osmotic
quantity
in
value
M
Cl
osmo-
cone.
metrically
mg
determined
Secretion
„
»
on
„
»
M NaCl
0.342
,,
„
,,
„
,,
,,
„
Table XIV
the external
of
the
0.086
+
0.172
sucrose
influence of
In
NaCl
this
of
0.412
37.0
0.558
0.613
17.6
0.688
0.750
sugar
of
the
in
osmotic value of
the osmotic
on
both
experiment
concentration
0.358
increase
an
through addition
fluid.
the
M
„
shows the
solution
secreted
mination of
+
38.2
secreted
fluid
value
deter-
analytical
an
and
an
osmometric determination of the NaCl concentration which is isotonic
with
the secreted
osmometrically
tration of the
a
addition of
Through
In
fluid
Table
added
substance
Here
amount
higher
MgS0 4
in
too
and
XV
an
sugar
so
appears
higher
that besides
have
here
been
too
that in
the
case
every
than the NaCl
NaCl
present
in
NaCl
a
small
the
the
concen-
amount
secreted
of
fluid.
concentration of the
greatly.
are
shown
in
which
MgS0
solution instead of sugar, in order
would
act
especially
different concentrations
osmotically.
as
to
has
been
4
investigate
NaCl
has
been
well.
addition of
NaCl
brings about a somewhat smaller
higher concentration of the secreted fluid, resulting in
amount
a
It
made.
experiments
which
a
fluid,
increases
two
was
must
the external
to
applied
a
secreted
different substance
secreted
a
fluid
determined value lies
of C1
considerably
given off.
smaller
On
enhancing the concentration with
amount
of
secreted
fluid is
obtained,
334
W.
while
H.
the Cl
result is
I.
ARISZ,
addition of
MgS0
In
in
the
first
place
the
their osmotic
VAN TOOREN
fluid
increases
influence of
of
less.
The
off than without the
being given
that the action
so
on
secreted
the
principle
.
HEIKENS, A. J.
of C1
amount
4
MgS0 4 corresponds,
H.
CAMPHUIS,
concentration of
smaller total
a
J.
these
and
NaCl
sugar,
substances
is
based
influence.
TABLE XV
Influence
on
the
secretion
of
the
increasing
osmotic
value
with
magnesium sulfate.
1954.
Pretreatment
28
hours,
20°
0.342
C,
M
NaCl, exposed
°
Secretion
28 hours, 20
C,
to
light.
dark
M
quantity
in
secretion
0.171
on
„
M NaCl
„
„
0.342
Mechanism
of
the
gland
The action of the
secreting
process
in
present
secreted fluid is
for
are
Our
them
data
i.e.
not
are
in
mainly
secretion
is
well,
as
as
the
on
0.429
the
by Ruhland
is
as
rich
is
gland cell
active
an
substances
organic
are
in salt,
in
poor
water, the secreted
the
salt, if
fluid is
glands work without selectivity.
but
they
squeeze
cells
gland
out
actively.
water
according
to
Ruhland.
found that the osmotic value of the
we
higher than
that of the
process dependent
proves
temperature,
the
but
rate,
that
the
expressed sap
work.
osmotic
perform
can
active
This
0.366
gland cell
if the
place
by
out
respect,
be
They change
unaltered.
326
647
and
salts
If the
opinion
demonstrated that
secretion.
25.0
42.4
yet adequate to give a full explanation of the
They supplement Ruhland’s results and differ from
glands
an
0.288
considered
hydathodes,
turned
fluid
516
not
one
can
sap
that the
and
In his
process.
secreted
is
quantity.
filtration-
no
in
secretion
which
rich in salt
Osmotic work is
secreting
in
water
poor in salt.
They
MgS04
the secretion takes
instance
also
M
gland cells
varying
a
0.081
+
„
cone.
NaCl
ßg
secreted
50.3
NaCl
M
„
Cl
mg
and
light
leave
was
of the
leaf,
that
proved
the presence of oxygen
on
secretion
It
the
inhibitors
NaCl
process
influence
concentration
depends
on
cell
metabolism.
In
fluid
a
is
leaf
on
a
higher
unknown,
cytoplasm
salt
the
whether the
or
vacuole,
concentration in
the
unknown whether
actually
solution the NaCl concentration of the secreted
than
higher
NaCl
nothing
NaCl
than in the
The osmotic value of the
pond
more
osmotic
or
value
higher, about
This
proves
less
of
be
in the
the
expressed
the
leaf is
M
secreted
sugar,
fluid,
leaf
cells
in
of the
sap
must
It
and
being
is
wall,
NaCl
therefore
fluid
is
leaf cells.
however,
can
seems
(Table
be
expressed
be done to
it
the secreted
of the leaf
than that of the
that osmotic work
leaf.
found in
stated about the actual
concentration
cytoplasm
of
with that of the external solution
of the
0.1
can
cytoplasm
the
concentration
chloride present
keep
to
corres-
XI).
The
considerably
sap
of the
the secretion
leaf.
going.
SECRETION
THE
The
the
of these
pressure
the
% of the dry
40
to
be
may
water
attributable
being reduced
has been
did
is
the
though
indicate
a
secretion
table XIV show that also
We
Limonium
there is
did
the osmotic
smaller than
of
when salt
than
of
the
by
can
fluid
of
the
in
from
be
sap
the
These
secretion
We have
be
is
XIV
content
and
are
dependent
In
gland cells.
in the
pores.
will be
the
The
relatively
suction
order
value of the
needs
gland
pressure
produce
to
has
sap
an
uptake
of
a
be
to
can
solution.
osmotically
solution.
that
the
a
of
quantity
higher
or
the
a
sap
exuded
is
on
the
gland
amount
assumption that
of the
of secreted
gland
cell
sap
(cf.
may vary,
under the influence of the
in NaCl.
results
osmotic value of the secreted
turgorpressure
discussed that the
poorer
considerable increase of
gland cells, which
in accordance with the
the salt concentration of the
expressed
richer
somewhat
or
cells till their suction pressure has become
the NaCl concentration in the
the sap
gland cells
the osmotic value of the external
external
experiments
already
pendent of
know
we
be further
to
the
to
and
equals
cells.
gland
the osmotic
Table
of
leaves
as
high concentration.
the
gland
of
small
since the osmotic
equal
substances
concentration of the sap in
higher NaCl
liquid.
a
enhancement of the osmotic concentration of the external
the osmotic
a
fresh
in
sap
be estimated
solution and that secretion is continued after
in
of
subject deserves
must
turgorpressure
that of the
appears
reduced
it
exuded
in
present
by enhancing
restoration
It
latifolium
gmelini.
concentration of the
active substances in the
higher
The
well.
as
higher than that of the external solution. The turgor
considerably
A
is
by
the leaves
continued,
The results
As far
permeability of the plasm
not
turgorpressure
reduced
experiments
be found in the secreted
secretion
administered.
was
of fluid
dry substance
being reduced
substances.
may
any
water.
secreted
turgorpressure
sufficient
however,
that
gmelini
destilled
on
two
or
of Limonium
case
out
cases
states
amount
in his
day
a
turgor
of Statice
leaves
water
be
flow
some
secretion, however,
organic
concentration
the
on
caused
The osmotic
the
the
in
the osmotic concentration of the sap of the
depend
difference is
be
of
directly determined,
concentration
plus
The
the
in Statice
especially
be
This will
find,
not
exudation of pure
no
Though
small
in
latifolium unless NaCl
examined
cannot
If the
that after
of other substances than NaCl
quantity
fluid.
third.
the concentration
decreased,
rate
This would
fluid
days. The secreted
assume
NaCl.
more
and
With
cells
will
naturally remain
the concentration of the
one
may
to
well. Ruhland
as
substance.
fourth,
than
we
off any
give
not
less
to
changed,
likely
will
of the secreted
organic
to
secretion
that sap
not
case
any
be secreted
to
matter
to one
fluid is
in
secreted
the
influence of the
under the
he obtained secretion for 34
then reduced
was
of
assumed
335
KTZE
LATIFOLIUM
wall
is
cytoplasmic
likely
are
It
molecular substances
NaCl
cytoplasm.
organic substances
25
The
LIMONIUM
outer
cytoplasm.
secretion cells
Great
OF
GLANDS
cutinized
the
cells.
quite unchanged.
in
the
leave
to
off from
given
in
apertures
enable fluid
SALT
OF THE
p.
it is
gland cells.
fluid is inde-
326). Seeing
evident that
turgorpressure may
also
336
H.
W.
Ruhland had
pressure
surface
the
of
the
outside,
pressed
of the
Here
the
in
pressure
the
to
the
cytoplasm
in the
pressure
cell.
place against
put
wax
the
of
the
comparison
a
those of
Spartina
the
that
a
on
fluid
is
In
Spartina the aperture
an
adjoining epidermal
be
must
transferred
through
cells.
not
requiring
cells
gland
of
coat
conclusion
gland
gland cell
epidermal
is the energy
take
can
a
TOOREN
of Limonium with
drawn the
pressure
VAN
A. J.
From
upwards.
in the
All these data indicate that
the
that
so
glands
has
(1955)
the
by
symplasm
up
HEIKENS,
which the sap exudes lies in the wall of
through
cell.
leaf is
Arisz
out
H.
CAMPHUIS,
from
structure
Townsendii
squeezed
J.
already shown that secretion
fairly high
anatomical
I.
ARISZ,
the exudation of the
from the
liquid
in the secretion but
process
by introducing
keeping
substances into these
cells.
The
in
problem therefore is
the
cells
gland
mechanism
must
substances into
dissolved
is
maintained
solution
a
carry
the
how this pressure and the salt concentration
being
of
and
simultaneously
is
considered whether the introduction of salt
or
that of
is
the
duction of
In leaves
is
sap.
gland
on
the
must
cells
the
pressure
the
cells
likely
secreted
Some
can
for
to
light
active
an
at
uptake
the pro-
maintained,
place
constant
a
rate.
the
in
as
leaves
on
a
sap
salt
concentration of the secreted
both
pressure
the
cases
pressure
is
secreted,
equally
NaCl will
especially
maintained
and
in
in
up,
leaves
be
on
in the
great.
pumped
water
by introduction of partly
when these other substances
inhibitors
the
be
have
water
passed
glandular
affect
cells
gland
assumed
uptake
this
and
with the aid
action
we
active
an
into the cell
of
assume
an
as
active
the
other
get
process
together
light
and inhibitors
of the
secreted
simultaneous active salt
energy
to
fluid
being
uptake, organized
equally influenced by the factors
The active salt
more
fluid
water
requires
with
a
lost
that
it
the
membrane that
process.
process
an
understand how
can
reduced. This
in
If
active
owing
be secreted without
such
a
way
mentioned as the active
uptake, however, would
into the
uptake
requiring
only active
uptake into the gland cells, it is hard
temperature,
the concentration
a
is
great
as
of fluid
in NaCl
especially
pressure
investigators
only
water
substance
organic
fluid.
An efficient
the
therefore
of outflow.
strength
cell.
the
be
NaCl
take
higher NaCl
keep
to
being
amount
rich
to
Temperature,
regulates
the
and
water
NaCl concentration of the secreted
the fact that in
to
regulating
substances than
in
much
a
be due
and
active
organic
process.
cells
gland
cells will
of sap
amount
gland
is
and
should,
well known. If by
are
the
the
water,
Whereas in leaves
into
in
pressure
which have
This
uptake
by the gland
sap
placed
reduced,
solution,
the
requiring
energy
Processes of active salt
It
unknown.
be
of substance
salts
gland cells. A mechanism that carries
substances
water
An
regulated.
with
water
requires
that it
make the active
is
uptake.
water
water
uptake
superfluous.
Until
above
we
have
conception
more
of the
data
at
our
mechanism
disposal,
as
a
it
system
seems
to
us
that carries
that the
salts into
SECRETION
THE
the
THE
SALT GLANDS
OF
gland cells actively and by doing
in
sulting
A
OF
an
definite
the
on
secretion
when the biochemical processes
Here
we
discussed
refer
may
secretion of
Arisz
by
the
to
salt solution
a
so
causes
LATIFOLIUM
a
pressure
of sap from the pores, is the
expression
opinion
LIMONIUM
mechanism
which it is
on
of this
correspondence
337
in them
re-
acceptable.
be
only
can
based,
the vessels with the
to
most
KTZE
given,
known.
are
with
process
of the
bleeding
the
root,
(1955).
Summary
The concentration of NaCl in the external solution
treatment
influences the secretion
with 2
to
%
% is
3
the salt
by
then both the
favourable,
most
concentration of the secreted fluid and also the
the leaf
The
greatest (Table
are
of
rate
reduced
secretion
higher
The
than
water
on
value of the external
decrease
on
salt
water
there is
When
always
exposed
a
owing
The concentration of the
If
is
Oxygen
In
a
times.
the
the
required
for
secretion
the
a
concentration
high
value is
equal
to
osmotic
value
of
osmotic
work
is
The osmotic
an
the sap
takes
place
they inhibit it.
value of the
secreted
of the secretion of
rate
(Table VII).
water
on
the
expressed
of the secretion
rate
The NaCl concentration
VIII and
medium
the
by
leaf sap
experiment
was
(Table
gland
(Table
IX,
Fig. 2).
XI). The
higher than the
(Table
is
For
XI).
computed
greater than the
This indicates
Increase of the osmotic
secreted
fluid
osmotic
value of
and
by NaCl (Table
On
there is
XII,
(Tables
sugar
as
the secretion
NaCl solution,
a
increased.
the
too
sugar
substances
in the
value of the external solution
decreases
NaCl
the
concentration
:
of
amount
and
the
fluid
but
fluid.
the
much
amount
diluted
amount
secreted
(Table
on
NaCl concentration of the
XIV)
secreted
decrease of the
concentration in the
has
XIII,
XV) alters
addition of
secretion
the presence of other
increases
the
with
XIV)
secreted
Cl
increases.
(Table VI).
leaf sap varies. The average osmotic
secreted fluid.
little.
fluid
change
not
25° C the
to
value of the secreted fluid,
other
fluid.
on
done.
plus salt solution it
by
either increase
with the NaCl concentration of the secreted fluid
corresponds
In
usually
osmotic
With secretion
secreted
unchanged (Tables
the osmotic
osmotic value of the
is
secretion.
The osmotic value of
expressed
(B)
of
amount
secreted fluid does
of the secreted fluid remains
(A)
fluid may
salt solution.
a
low concentration inhibitors increase
of fluid, in
the lower
to
or
The concentration of the secreted fluid remains
unless
fairly equal,
VI).
on
5
temperature is raised from
fluid increases 7
days
fluid is
decrease.
light
to
few
first
of secreted
solution.
the secretion
during
and the
amount
NaCl concentration in
the
during
amount
The NaCl concentration of the secreted
or
pretreatment
I).
constant
III, IV, V).
(Tables
somewhat
is
the pre-
during
A
glands.
fluid
of
secreted
salt solutions
of secreted
increases.
or
water
fluid. The NaCl
The
total loss
of
338
W.
(C)
by
H.
ARISZ, I. J. CAMPHUIS,
MgS0
(Table
4
secreted fluid and
secreted
XV)
H.
gives
fluid. The total
a
pressure
the
that results
cells.
gland
gland
external
The
into
actively
The
cells.
in
The
J.
VAN
reduction
a
loss of Cl
TOOREN
of
the
the
of
latter
gland
expressed
may
has
of
amount
decreased.
may be considered
squeezing of
a
amount
A.
increase of the NaCl concentration of the
an
The mechanism of the secretion
introduces salts
HEIKENS,
be
cells
depends
as
a
system
by doing
through
sap
sap
and
so
the pores
on
the
out
of the
pressure
influenced
osmotically
that
causes
by
in
the
solution.
composition
accumulated in the
of
the
expressed
cells and
gland
sap
on
depends
the
on
the
permeability
substances
the pores.
at
References
Andel,
Arisz,
root.
O.
W.
In
Ruhland,
M.
H.
van.
1955.
Proc.
Kon.
Significance
Ned. Akad.
of the
v.
Wetensch.
Symplasm theory
for
Ser.
C,
55:1.
transport
across
the
preparation.
W.
1915.
Schtscherback, J.
Schimper,
1952.
A.
F.
W.
Jahrb.
1910.
und
gischer Grundlage.
wiss.
Ber.
F.
d.
C.
Gustav
Botanik
D. Bot.
von
55:409-498.
Ges. 28:30-34.
Faber.
Fischer, Jena.
1935.
Pflanzengeographie
auf
physiolo-