NOTES

A note on measurements of Onsager coefficients in maple petioles and

maple woody xylem by electroosmotic techniques
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D. J. URSINO~
AND D. S. FENSOM~
Departnlent of Biology, Queen's University, Kingston, Ontario
Received February 13, 1969

Dedicated to the memory of the late Dr. G. P. Krotkov and the late Dr. C.D. Nelson
URSINO,
D. J., and D. S . FENSOM.
1970. A note on measurements of Onsager coefficients in maple petioles
and maple woody xylem by electroosmotic techniques. Can. J. Bot. 48: 1289-1292.
Measurements on petioles and on 2- and 4-year old xylem of Acer saccharurn have shown that the
specific conductivity (LEE) measured after they were soaked in 10-4 N KC1 was about 2 x 10-4 ohm-1
cm-1 and did not change significantly through the growing season in these plant tissues. On the other
hand the electrokinetic coefficient (LPE) varied seasonally in the petioles with a value of about 3 X 10-6
in spring and 3 X 10-5 cm2 s-1 volt-1 in summer. The electrokinetic coefficient in the woody xylem
measured about 6 X 10-5 cm2 s-1 volt-1. From the values of these Onsager coefficients the contribution of electroosmosis to phloem transport would seem to be negligible in these cut sections.

Introduction

Materials and Methods

Studies of electrokinetic effects in plants have

been made for some time by one of us (Fensom
1957; 1958; 1963). From the measurement of
Onsager coefficients, it is now known that the
magnitude of internal pressure gradients in
woody plants may be determined (Tyree and
Fensom 1968). Certainly the expanding use of
Onsager coefficients in plant tissue (Dainty,
Croghan, and Fensom 1963) makes it important that relevant measurements be made
available in the literature. The present work was
undertaken to improve our knowledge of biokinetic theory in relation to transport, to check
with some precision the measurements made by
Fensom (1962) on electroosmosis in freshly
thawed twigs of maple (Acer saccharum), and
to extend the work on Lens culiizaris roots
(Fensom, Meylan, and Pilet 1965) into living
tissues of maple. Maple petioles were chosen
for this work to find whether electroosmotic
flow was possible in vascular tissue of this
tree.

The petioles used in all experiments were obtained

from a single branch of a maple (Acer saccltarum,
Marsh) tree located on the Queen's University campus.
Once removed from the tree, the base of the petiole was
immersed in 1.0 X 10-4M KC1 and a section of the
petiole, 2.0cm in length midway in the petiole and
about 1.Omm in diameter, was obtained for use in the
experiment. The woody xylem came from 2- and 4-year
old woody stems. The outer layers of bark, cambium,
and xylem were removed by razor blade and knife and
samples were taken from the inner xylem region. Like
the petioles, the xylem sections were 2.0cm in length,
1.O mm in diameter and were retained in 1.0 X 10-4 M
KC1 for about 30 minutes before they were placed in the
electroosmometer.
The magnitude of induced electroosmotic flow in the
maple petiole and woody xylem was determined using
an electroosmometer, first described by Fensom and
Dainty (1963). The sides of the samples were greased with
vaseline over about 15 mm to hold the sections firmly in
the split rubber stopper between chambers and to reduce
current flow along the lateral surfaces.
Electrical potential differences were established using
a 7.5-volt battery and currents were measured using a
d-c. microammeter. Temperatures were kept constant
during the experiments by immersion of the entire
electroosmometer apparatus in a constant-temperature
bath. At all times the solution used in each chamber of
the electroosmometer was 1.0 X 10-4M KCI, the same
solution in which the petioles and woody xylem were
maintained before the experiments. After the assembled
electroosmometer containing the experimental material
was placed in the water bath, about 30 minutes was
allowed for the internal KC1 solution to reach the same
temperature as that of the water bath (21-22.5 "C).

lThis work was carried out in collaboration with the

late-6r. C. D. Nelson.
zpresent address: Department of Biology, Brock
University, St. Catharines, Ont.
3Present address: Department of Biology, Mount
Allison University, Sackville, N.B.

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1290

CANADIAN JOURNAL O F BOTANY.

Calculations
The amount of induced solution flow per
unit of electrical current applied is known as
electroosmotic efficiency and was calculated
from the relation
e-o efficiency = J/I(,p = 0)
where J is the induced solution flow in cm3
s-1 cm-2.
I is the applied current in A cm-2
A P is the pressure difference across the
tissue.
Electroosmotic efficiency is measured in cm3
coulomb-1, or alternatively in moles of water
transported per Faraday.
Electrical resistance (R) of the plant tissue
(either petiole o r woody xylem) was calculated
from measurements of the applied voltage (V)
and current (I) across or through the solution
tissue
alone or the solution
Rtissuc = Rso~ution+ tissue - Rso~ution
and the specific resistance was obtained from
the equation:
Rspecific= RtissueX (area of cross section) /
(length of section).
The Onsager coefficient of electrical conductance
(LEE) in ohm-' cm-1 is the specific conductivity
where A P = 0 and is thus the reciprocal of the
The electrokinetic
specific resistance (RspeCific).
cross coefficient (LpE) = J /I(P
= 0) X LEE and
was calculated from the prior measureme1:ts of
e-o efficiency and specific conductivity. Its units
are cm3 cm-2 s-1 volt-1 cm = cm2 s-1 volt-1.

Results
Table I summarizes the results obtained on
petioles during the season of their development
and function and Table I1 summarizes the data
from the 2-year-old and 4-year-old woody
xylem. That electroosmosis occurred is evident
and since water flow always moved toward the
cathode when current was applied, we may conclude that the net zeta potential on the walls of
maple xylem o r petiole is negative. This is
similar to other plant tissue studied to date and
confirms the previous findings of Fensom (1962).
It may be seen that the specific conductivities
(LEE) of both the petioles and woody xylem
were relatively uniform around 2.2 X 10-4
ohm-1 cm-1. This value is intermediate between
the specific conductivity of Nitella cell walls
(Tyree 1968) and pure KC1 solution 10-4N as

VOL. 48.

1970

might be expected when cell wall channels are

large. This value also compares reasonably well
with the value of 3.0 X 10-4 f o u i ~ dby Tyree
and Zimmermann (private coinmunication) for
30-year-old sapwood xylem of Acer rubrum.
With the electrokinetic cross coefficient (LPE)
the results were different. In petioles a marked
seasonal variation was evident (Table I) with the
values in mid or late summer very significantly
different from those measurements in spring and
autumn. The mid o r late sunlmer measurements
approached those of xylem but were still
significantly lower than them. Our values for the
electrokinetic cross-coefficient found for the mid
and late summer petioles are in reasonable
agreement with those reported for Nilella cell
walls (Tyree 1968), for vascular strands of
Nynlphoides (Fensom and Spanner, in press),
and for Heracle~irn (Tyree and Fensom, in
press). Furthermore, it looks highly probable
(Marr and Fensom, private communication)
that the magnitude of LPE is small in relation
to the other Onsager coefficients of electrical or
hydraulic conductivity (LEE and Lpp) SO that
significant electroosmotic contributions to phloem transport would seem improbable in these
petioles.
The seasonal variation of LpE in petioles may
possibly be explained by considering that in
these tissues phloem and xylem pathways are
in parallel (for parallel electroosmotic pathways,
see Kedem and Katchalsky 1963). Let the fluxes
and coefficients pertaining to phloem be marked
with a prime, and those pertaining to xylem
with a double prime. Then

A E = difference in electrical potential:

electrical gradient.
Therefore where A P = 0,

or

but the new coefficients must be weighted in

terms of the appropriate cross section areas:
let a be the fraction of xylem and (1-a) be
phloem, then

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TABLE I
Electroosmotic efficiency and the Onsager coefficients, LEE and LEP,in maple petioles: seasonal changes

--

Electroosmotic efficiency
Time of experiment

Number of
samples

Spring 1964
(23 15-10 16)
~ u k i e 1964
r
( 1 7 18-22 18)
Late summer 1964
(419-619)
Autumn 1964
(9110-15/10)

Mean current,
FA

Moles water
per Faraday

cm3 coulomb-1

LEE, ohm-1 cm-1

LPE (= LEP),
cm2 s-1 volt-1

4.4(?0.4)

64(+ 6)**

12(+ 1 .O)xlO-3

2.6(?0.3)XlO-4

3.14(+0.3)XlO-6**

NOTE:
Slandard deviations o f the means given in parentheses.
Difference between mean and next seasonal mean,

V]

* significant, ** very significant.

TABLE 11
Electroosmotic efficiency and the Onsager coefficients, LEE and LPE, in maple woody xylem
Electroosmotic efficiency
Age of xylem

Number
of
samp!es

Mean current,
PA

Moles water
per Faraday

2 years
4 years

6
3

3.3(?0.3)
4.5(+0.2)

1770( 350)
1230(+ 120)

NOTE:Standard deviations o f the means gi\.en in parentlleses.

cm3 coulomb-1

LEE, ohm-1 cm-1

LPE = (LEP),
cm2 s-1 volt-1

330(+ 66)X 10-3

240(+ 23) X 10--3

1.7(+0.3)XlO-4
2.7(+0.3)x10-4

56(+ 1O)X 10-6

65(+ 9)XIO-6

1292

CANADIAN JOURNAL O F BOTANY.

In our work LEE(for petiole) was very nearly =

LrtEE (for xylem) and in summer (1-a) LtEE
< aLttEE.

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Therefore LpE N (I-a) L t p ~ ~L"p,y,

which means that since the fraction of xylem
(a) is large, particularly in petioles in mid and
late summer, the Onsager electrokinetic coefficient (LpE) at that time will very nearly be
that of the constituent xylem tissue, though in
the spring a larger amount of phloem might be
expected. It seems probable that most of the
seasonal variations found in the coefficient
(LPE) may be due to a changing charge on the
xylem, i.e. changing LttPE. It would also appear
that the value for the Onsager electrokinetic
coefficient for phloem is less than that for
xylem, an observation similarly made in Heracleum (Tyree and Fensom, in press).
DAINTY,J., P. C. CROGHAN,
and D. S. FENSOM.1963.
Electro-osmosis with some applications to plant
physiology. Can. J. Bot. 41 : 953-966.
FENSOM.
D. S. 1957. The bio-electric uotentials of vlants
and their functional significance. I.' An electro-kinetic

VOL. 48, 1970

theory of transport. Can. J. Bot. 35: 573-582.

1958. The bio-electric potentials of plants and
their functional significance. 11. The patterns of bioelectric potentials and exudation rate in excised sunflower roots and stems. Can. J. Bot. 36: 367-383.
1962. The bio-electric potentials of plants and
their functional significance. IV. Changes in the rate
of water adsorption in excised stems of Acrr sacc/~arunl
induced by applied electro~notiveforces: the "flushing
effect". Can. J. Bot. 40: 405413.
1963. The bio-electric potentials of plants and
their functional significance. V. Some daily and seasonal changes in the electrical potential and resistance
of living trees. Can. J. Bot. 41: 831-851.
FENSOM,
D. S., and J. DAINTY.1963. Electro-osmosis in
Nitella. Can. J. Bot. 41: 685-691.
FENSOM,D. S., S. MEYLAN,and P. E. PILET. 1965. Induced electro-osmosis in root tissues. Can. J. Bot.
43: 453467.
FENSOM,
D. S., and D. C. SPANNER.1969. Planta Arch.
Wiss. Bot. 88: 321-331.
KEDEM,D., and A. KATCHALSKY.
1963. Permeability of
composite membranes. Faraday Soc. Trans. 59: 19181951

TYREE,M. T. 1968. Determination of transport constants of isolated Nitrlla cell walls. Can. J. Bot. 46:
3 17-327.
TYREE,M. T., and D. S. FENSOM.1968. Methods of
measuring hydrokinetic pressure gradients in the
xylem of plants in situ. Can. J. Bot. 46: 310-314.
1970. Some experimental and theoretical observationsconcernine mass-flow in the vascular bundles
of Heracleum. J. EX;. Bot. In press.

Sensitivity of the infrared COz gas analyzer to 14COi1

N. R. BULLEY~
Department of Biological Sciences, Simon Fraser University, Buriraby, British Cohtmbia
AND

E. B. TREGUNNA
Botany Department, University of Britislt Colnmbia, Vancouver, Britislt Colrtrnbia
Received October 3, 1969

Dedicated to the memory of the late Dr. G. P. Krotkov and the late Dr. C. D. Nelson
BULLEY,N. R., and E. B. TREGUNNA.
1970. Sensitivity of the infrared C 0 2 gas analyzer t o 14C02.
Can. J. Bot. 48: 1292-1294.
The infrared C 0 2 gas analyzer (Beckman model 215) is only 5% as sensitive t o 14C02 as it is to 12C02.

Radioactive carbon dioxide (14C02) is used in

many biological experiments related to photosynthesis, photorespiration, translocation, and
the products of photosynthesis. Rates of photosynthesis are normally measured by the rate of
1Supported by the National Research Council of
Canada.
2Present address :Department of Agricultural Engineering, University of British Columbia, Vancouver 8, B.C.

14C02 uptake as measured by a Geiger tube or

the rate of 12C02 as measured by an infrared gas
analyzer. To the authors' knowledge, the sensitivity of the infrared gas analyzer to 14C02 concentrations is not known. Most experiments use
very low concentrations of 14C02 and any effects
that the 14Co2might have on the readings are not
discussed. At high specific activities the error
would be sigdcant. The following experiments