BIOLOGIA PLANTARUM (PRAHA)

1 (1) :39--47, 1959

Gradients of Osmotie Pressure of {Jell Sap in the Area

of one Leaf Blade

BOHDAN SLAVIK
Department of Plant Physiology, Institute of"Biology, CzechoslowtkAcademy of Sciences, Praha
Received February 24. 1958

Souhrn

Nejen podle v:~ky inseree (z~:konZalensk6ho), n~br~ tak6 v rg~mcijedn6 listov6

6epele existujl rozdfly v hydrataei. Byly zji~t~ny zna~n6 d61kov6, transversAlrd
i plo~n6 gradienty osmotiek6ho tlaku bun66n6 ~$Avy (stanoveni nep~imo indexem
lomu podle empiricky zji~t6n6 korelace mezi indexcm lomu a osmotick~m tlakem
kryoskopicky zm6~en:~m[obr. 1]) v rozsahu jedn6 listov6 6epele cukrovky. Gradient
base 6epele-vrchol se pohyboval kolem -4-40~ osmotick6ho tlaku bun66n6 $~Avy
na basi. Tento a ostatni gradienty ani jejich rozlo~enl se zf'eteln6 nezm6nily ani p~i
um616m dosyceni list6 vodou ani p~i vzniku nebo vyrovnAni vodnlho deficitu
aktuAlnimi zrn6nami vodnl bilanee. (P~i t6chto pokusech byla zachovAna polarita
transpira6niho proudu.) Taro zjiht6nA stabilita rflzn6ho stupn6 hydratace jednotli-
v:~ch 6Asti dosp616 6epele je dflkazem toho, ~.e toto rozrflzn6ni je ds p~edehozim
ontogenick:2m v:~vojem listu jako polArniho orgAnu, a ~e se v dosp61osti listu
udr~uje v podstat6 bez ohledu na aktuAlnl zrn6ny vodnlho provozu listu. ZjiSt~nA
heterogenita listu, pokud so tp6e uveden6 veli6iny, je v:Yrazemfysiologick6 hetero-
genity listov6 6epele.

Summary

W i t h i n one leaf blade there are considerable differences in h y d r a t i o n of

its parts. Marked longitudinal, transverse a n d area gradients of the osmotic
pressure of cell sap were ascertained within the leaf blade of sugar beet. The
osmotic pressure was d e t e r m i n e d indirectly b y the refractive index according
to the empirically established correlation refractive index: osmotic pressure
(measured cryoscopically) cf. fig. 1. The gradient from the base of the blade
to the tip r a n g e d a r o u n d plus 4 0 % of the osmotic pressure at the base. This
a n d o t h e r gradients m e n t i o n e d were not n o t i c a b l y changed even when the leaves
were artificially s a t u r a t e d with water or when water deficit arose or was
a d j u s t e d b y actual changes in t h e w a t e r balance. (In all these experiments
the p o l a r i t y of the t r a n s p i r a t i o n s t r e a m was maintained.) The s t a b i l i t y fi)und
in various degrees of h y d r a t i o n of individual p a r t s of the a d u l t leaf blade indi-
cates t h a t t h e variability is a result of t h e previous ontogenetical d e v e l o p m e n t
of the leaf as a polar organ a n d o f the fact t h a t the a d u l t leaf remains essentially
u n c h a n g e d b y t h e actual changes in the water exchange. The h e t e r o g e n e i t y
f o u n d in the leaf with regard to the values here dealt with is a n expression
of the physiological h e t e r o g e n e i t y of the leaf blade.

39
40 B. SLAV~[K

Introduction

Within the plant organism there exist progressive changes in the physio-
logical (and as a consequence also in the morphological) characters of organs,
their parts, tissues and cells, as a function of their relative positions (P~AT, H.
1948 and 1951). These physiological and morphological gradients are prima-
rily determined by the previous development of these parts. Many of them are

28

26

24

2Z

ZO

18
o/

18
I I
1"347. 1"345 1-~50

Fig. 1. Relation of osmotic pressure of cell sap (ordinate in atm.) of sugar beet leaves to their
refractive index (abscissa) in the experimental period (October 1957).

closely and directly connected with the water relations and their polarity.
The existence of axial gradients of the anatomical and physiological features
of xeromorphism in leaves, according to the level of insertion is known (Za-
lensky's law). Recently their physiological investigation has been linked
with the study of the physiology of xerophytes (recent work by GENKEL'
1946, SIMONIS 1941 and 1952, ZNAMENSKY 1948, MAXIMOV 1952, STOCKER
1954, 1956, KILLIAN and LEM]~E 1956, FARKAS and RAJttITHY 1955, CETL
1957, and others).
In view of the polarity of leaves during ontogenesis the existence of physio-
logical gradients within a single leaf blade may be anticipated. Heterogeneity
within the leaf blade has not yet been investigated, although it is theoretically
 OSMOTIC PRESSURE IN THE AREA OF ONE LEAF BLADE 41

probable (HU~ER 1956). As an indicator of this anticipated heterogeneity,

connected primarily with the character of water state and balance, the quanti-
tative distribution of the hydration of leaf tissue within the leaf blade of sugar
beet was examined. In seeking the cause of this heterogeneity the fundamental
question was whether with varying water balance it is connected solely or
primarily with the actual varying supply of water to different parts of the
leaf blade, or whether there is a relatively stable heterogeneity already determin-
ed by the ontogenetical development of the leaf.

Methods
T h e correlation g r a p h (fig. l), d r a w n u p ad hoc on t h e basis o f 36 c o n c u r r e n t d e t e r m i n a t i o n s
of t h e refractive i n d e x ( A b b e ' s u n i v e r s a l r e f r a c t o m e t e r ) a n d t h e cryoseopie m e a s u r e m e n t of
t h e o s m o t i c p r e s s u r e of cell s a p f r o m killed (100 ~ C) leaf tissue of s u g a r beet, s h o w s a practically
linear relation b e t w e e n t h e two v a l u e s w i t h i n t h e limits of t h e e x p e r i m e n t a l period (October).
T h i s relation c a n be r e g a r d e d as reliable for practical application.*) T h i s m a d e it possible to
m a k e a sufficiently a c c u r a t e e s t i m a t e of o s m o t i c p r e s u r e f r o m t h e refractive index, i. e. osmoti(:
p r e s s u r e could be m e a s u r e d in a m a n n e r w h i c h e n a b l e d t h e r e q u i r e d large n u m b e r of s m a l l s a m p l e s
o f leaf t i s s u e to be d e a l t with. I n t h i s w a y t h e os-
m o t i c p r e s s u r e of cell s a p f r o m i n d i v i d u a l killed
sectors of t h e leaf b l a d e o f s u g a r beet (Beta vulgaris
ssp. esculenta [SALISB.] Gff~KE var. altissima R o E s s m )
v a r i e t y Dobrovickd A w a s indirectly m e a s u r e d . F o r
t h e s e e x p e r i m e n t s leaves of m e d i u m i n s e r t i o n level
were g a t h e r e d f r o m t h e e n d of S e p t e m b e r to t h e
e n d of October 1957. T h e o s m o t i c v a l u e m e a s u r e d
was r e g a r d e d as a q u a n t i t a t i v e e x p r e s s i o n of t h e
h y d r a t i o n of t h e tissue. E a c h h a l f of t h e leaf blade
w a s d i v i d e d in e a c h case into eleven sectors (fig. 2
a n d 3), t h e a b s o l u t e size of w h i c h w a s d e t e r m i n e d b y
t h e a b s o l u t e size of t h e leaf blade. T h i s v a r i e d a r o u n d
a n a v e r a g e of 27 18 cm. T h e v a l u e s g i v e n were
d e t e r m i n e d w i t h different controlled s t a t e s of w a t e r
17.6 16.8 16.8 17.6
s a t u r a t i o n a n d w i t h different w a t e r b a l a n c e s in
n a t u r a l a n d artificial e x p e r i m e n t a l conditions. T h e
r e s u l t s r e c o r d e d r e p r e s e n t a v e r a g e s f r o m a larger 16.2 154 15-7 16.2
u u m b e r of d e t e r m i n a t i o n s (n a b o u t 20 to 30).
A t o t a l of 1,300 s a m p l e s w a s d e a l t w i t h i n t h e
period s t a t e d .
I w i s h to e x p r e s s m y sincere t h a n k s to Mrs.
M. K r e j c a r o v h for h e r t e c h n i c a l a s s i s t a n c e .

Fig. 2. S y m m e t r i c a l d i s t r i b u t i o n of o s m o t i c p r e s s u r e of cell s a p (in arm.) over

t h e a r e a of s u g a r b e e t leaf blades ( n = 16).

Results
With regard to the refractive index of cell sap, both halves of the blade of an

*) T h e e x i s t e n c e of t h i s p r a c t i c a l l y linear r e l a t i o n of t h e v a l u e s is l i m i t e d to a q u i t e definite
d e v e l o p m e n t a l s t a g e a n d to a definite p a r t of t h e v e g e t a t i v e period. I t is in f a c t o n l y v a l i d insofar
as t h e relative q u a n t i t a t i v e c o m p o s i t i o n of t h e o s m o t i c a l l y active s u b s t a n c e s dissolved in t h e
cell s a p does n o t u n d e r g o a n y m a r k e d c h a n g e (SLAviK 1959). T h e relation i l l u s t r a t e d in fig. l
is n o t q u i t e linear. U n d e r t h e s i m p l i f y i n g a s s u m p t i o n of l i n e a r i t y its correlation coefficient is
r ~ ~ 0"98, r e g r e s s i o n coefficient b y ~ 1-37 ( a t m . to t h o u s a n d t h u n i t s of t h e refractive index)
42 B. SLAVIK

adult sugar beet leaf were mutually symmetrical, both relatively and absolute-
ly (fig. 2). Corresponding values differed from each other on an average b y
less than 1%, maximally b y 1.5%. This result is especially important for
observation of the relative changes in osmotic pressure from corresponding
places, when the water balance is active or passive and of varying intensity.

A B C D

18"6 19'6

F i g . 3. D i s t r i b u t i o n o f o s m o t i c p r e s s u r e o f cell s a p o n h a l f t h e s u r f a c e o f t h e l e a f b l a d e o f s u g a r
beet in two sets of experiments (A a n d ]3 i n a t m o s p h e r e s , C a n d D i n p e r c e n t a g e o f
o s m o t i c p r e s s u r e a t t h e b a s e o f t h e b l a d e ; A a n d C : n = 36, B a n d D : n ~ 31).

These changes have been studied in another paper. Here we were concerned
only with establishing the changes in gradient within the leaf blade, i. e.
with the comparison of values within the same half of the leaf blade. The
symmetry of the two halves of the blade gives a guarantee that the gradients
in the two halves behave similarly under similar conditions.
The distribution of the average osmotic pressure of cell sap within one half
of the sugar beet leaf under natural field conditions in the morning (8 a. m.)
state of water saturation (resaturation at night) for two sets of experiments
(n z 36 and 31) provides a basic survey (fig. 3). The graphic illustration of
the axial gradient base of the blade tip along the main ribs (figs. 4, continuous
lines) as a percentage of the osmotic pressure at the base shows that osmotic
pressure increase is roughly linear according to position in relation to the
longitudinal axis of the blade, so that the gradient remains essentially the same
along the whole length of the blade. In this and other similar graphs it is
possible to observe a weak, but in almost all cases clear reduction in the
longitudinal osmotic gradient in the middle of the upper half of the leaf blade
(between points 3 and 4).
 OSMOTIC PRESSURE IN THE AREA OF ONE LEAF BLADE 43

A similar distribution of osmotic

pressure of cell sap was ascertained 140
in leaves which had been artificially
saturated with water (20 hours with
petioles in water in an atmosphere of
100% humidity and in the dark). The
longitudinal gradient was the same 130
(fig. 4, dashed lines) only the gradient
at the tip being smaller.
Cut leaves (without petioles) allowed
to wilt freely also maintain the same 120
type of gradient (fig. 5). In comparison
with the control (the second half of
the leaf saturated at the start of wilting)
this gradient does indeed show a slight 110
change, but no relation was found
between these changes and the magni-
tude or intensity of the water deficit
of the leaf due to wilting. I
In another set of experiments the B 2 3 4. T
influence of a passive water balance (i. e.
"wilting"), without interruption of the Fig. 4. R e l a t i o n of t h e o s m o t i c p r e s s u r e of
cell s a p (ordinate in per c e n t of o s m o t i c pres-
supply of water to the blade through sure a t t h e base of t h e blade) to t h e position
the petiole, was studied. Water deficit a l o n g t h e l o n g i t u d i n a l axis of t h e blade
in leaves with their petioles in water (abscissa: B base of blade, T tip). G r a p h i e
r e p r e s e n t a t i o n of axial g r a d i e n t .
was caused by artificial experimental
conditions (ventilation, radiation with C o n t i n u o u s lines: leaves picked in t h e
an infra-red lamp). This set of experi- m orning after normal overnight saturation
w i t h w a t e r (n = 31 a n d 36). D a s h e d lines:
ments was particularly designed to show artificial s a t u r a t i o n of leaves (n -- 16 a n d 16).
whether the area and longitudinal
gradients of osmotic pressure of cell sap
are noticeably changed by the assumed variation of water supply to the different
parts, i. e. by varying water balance. Not even here. however, did the distri-
bution and gradients change to any noticeable extent. Changes observed again
concerned only the gradients at the tip (fig. 6) and their relation to the absolute
magnitude of the water deficit and to the rate of its increase are clear from
the values given in the description. It can be seen from the results that
very small and medium deficits and their rate of increase did not change the
gradient as compared with the control, while deficits of about 5% and large
deficits (over 10%) reduced the tip gradient. But not even these changes
disturbed the general course of the gradients.
Neither did an active water balance change this situation. With leaf
blades in which the water deficit was artificially produced and which were
then saturated with the petioles in water in an atmosphere saturated with
water vapour and in the dark, the area distribution of the values under ob-
servation was also little changed as compared with the control (wilted half of
leaf blade before resaturation) (fig. 7).
44 B. SLAVfK

[ In general it can be said t h a t neither

- 140 /"' saturation of the leaf with water, nor-
passive, nor active water balance (the
occurrence or adjustment of total water

130
deficit), with water supplied through
the petiole or not, changed the relative
- distribution of osmotic pressure of cell
sap over the area of the leaf blade,
because all changes in these values took

//
place in parallel over the whole area.

- 120
/// Discussion
/
Considerable differences were ascer-
tained in the hydration of different
-m / parts of the sugar-beet leaf blade and

/
O
//

1
I
2
I
3
I
4 T
I
these corresponded to the natural gra-
dient along the main axis and trans-
versely (from edge to centre). Both
halves of the blade behaved quite sym-
metrically. The positive gradient of the
osmotic presure of cell sap from the
Fig. 5. Similar r e p r e s e n t a t i o n of the axial base of the leaf blade to the tip and
gradient as fig. 4. Continous line: wilting from the mid-rib to the edge is an
leaf blade w i t h o u t w a t e r supply t h r o u g h
petiole. D a s h e d line: axial gradient o f the expression of a decreasing degree of
second half of t h e blade before wilting
(n = 26).

140

Fig. 6. Similar r e p r e s e n t a t i o n of axial grad- 130 H..' /

lent as fig. 4 a n d 5; passive w a t e r balance
// t f'

e~lll"*~'llll
I
w i t h w a t e r supply t h r o u g h petiole.

120 /, . '/". /
w a t e r deficit 0 to 2 % , average
l~o, average rate o f increase o f
0-6~o per hour;
w a t e r deficit 2 to 6%, average
5'1~o, average rate of increase
2.2% p e r hour; 110 ~,//
. . . . . w a t e r deficit 6 to 10~o, average
8.3~o, average rate of increase r 1
8.8~o per hour;
....... w a t e r deficit > 10~o , average
14.4~o, average rate of increase
15~/o per hour.
100
l
/I I
I
2
I
3
I
4
I
T
 OSMOTIC PRESSURE 1R~ T H E A R E A O F O N E L E A F B L A D E 45

hydration of the leaf tissues. The grad-

ient from the base to tip in full grown
leaves of sugar beet in the autumn was
around 40~/o of the osmotic pressure at
the base.
These gradients were not changed
140
///+/
to any extent with actual changes in
the state of water saturation as a result
/+/
of different water balances. This in-
dicates t h a t these gradients (hydration 130
gradients) are relatively permanent
characters ensuing from the polarity of
growth and the ontogenesis of the leaf,
because when the leaf blade is mature lZ0
/+ /'
they are maintained essentially un-
changed without regard to actual
changes in the water balance. It follows
logically t h a t this more or less perma- II/+
nent hydration degree of various parts 110 //
of the area of the leaf blade undoubtedly
arose as a result of permanently differing i
water balances of those parts in the
early stages of development and growth 10D I I J I
of the leaf blade. I t is, however, very B 1 2 3 4 T
probably connected both as cause and
effect with differing intensities of other Fig. 7. Similar r e p r e s e n t a t i o n of a x i d a l grad-
physiological processes. It may be ex- ient as in t h e p r e v i o u s g r a p h s . S a t u r a t i o n of
wilted leaf blades. C o n t i n u o u s line: originally
pected t h a t similar differences such as wilted, n o w s a t u r a t e d h a l v e s of leaf blades.
were ascertained for leaves at different D a s h e d line: second h a l v e s of leaf blades
levels of insertion and of different de- w i t h original deficits (average 8.5O/o).
grees of xeromorphism exist analogically
within a single leaf blade. The magnitude of the gradients observed is consider-
able and corresponds to similar gradients according to height of insertion. At
the same time it provides the opportunity of following such changes conse-
cutively on the identical material.
The significance of these gradients of osmotic pressure of the cell sap of leaf
tissues for processes in the leaf itself can lie both in influencing the transpir-
ation stream (HU]3ER 1956, LEVITT 1956) and in influencing the intensity
of the transpiration of different parts of the leaf blade. As far as the first case
is concerned it must be remembered t h a t the factor which determines the
osmotic absorption of water by the tissues is not the osmotic pressure of the
cell sap (which is an indicator of hydration), but DPD (diffusion pressure
deficit), i. e. the so-called "suction power" of the tissue (HuBER 1956). The
gradients of osmotic pressure of cell sap determined need not, therefore,
quantitatively correspond to the DPD gradient of the tissue. Qualitatively,
however, this gradient of osmotic pressure of cell sap is a proof of the existence
of analogical DPD gradients determining the osmotic absorption of water
in the course of the transpiration stream. As far as the influence on transpir-
46 B. SLAV~[K

ation intensity is concerned, it has been theoretically established that only

after the reduction of turgor to nought does increased osmotic pressure of
cell sap start to act as a factor in reducing transpiration, and that this reduction
is under normal conditions comparatively small (~ENNER 1915, STALFELT
1956).

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Address: D r B o h d a n Slavik, I n s t i t u t e of Biology of t h e Czechoslovak A c a d e m y of Sciences,

N a cviSihti 2, P r a h a - D e j v i c e .
 OSMOTIC PRESSURE IN THE AREA OF ONE LEAF BLADE 47

~pa~HeHT~ OC~OTHqecKorO ~ B A e H H ~ K~eTO~HOrO c o K a

B npe~eaax H ~ C T H H ~ H ~HCTa
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