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OSMOTIC PHENOMENA.
BY CHARLES
A. SHULL,THE UNIVERSITY
OF KENTUCKY,
U.S.A.
(Presented by Mr. R. A. KEEN.)

A thoughtful consideration of the various relationships which exist

between plants and their environment, leads one to the conclusion that
none is of greater importance than the relationship established with the
moisture of the soil through the roots and root hair mechanism. The
fundamental character of this relation is seen clearly in the facts that soil
moisture is the dominant edaphic factor in the geographic distribution of
vegetation, and that some of the most striking modifications in the form
and structure of plants are produced by physical or physiological dryness of
the environment. The deserts of the earth have a sparse vegetation not
because of infertility, but from low soil moisture content. And the succulent types of plants found in xerophytic and halophytic habitats have been
developed through the physiological conditions set up inside the plant, incident to low availability of water.
As has been shown recently by MacDougal and others, the course of
carbohydrate metabolism is determined in certain succulents by the amount
of water existing in the cells, and the probability is that most succulent
plants have been modified by the osmotic conditions in their cells. Whenever the water content exceeds the critical amount, the carbohydrates form
polysaccharides of low imbibitional capacity. But if the saturation deficit
becomes great enough, the tendency is for the carbohydrates to go over
into pentosans which are hydrophilous. I n such cases, if in addition there
is the production of much free acid during the carbohydrate transformations,
succulence will probably result from the combination of circumstances.
Thus low availability of soil moisture may fundamentally change the metabolism of the plant, and modify its whole form and structure.
Whether we consider the influence of the plant upon soil moisture, or
vice versa, the influence of the quantity of soil moisture upon the plant,
these influences must act through the same mechanism, the osmotically
active membranes of the plant. Usually these are living membranes, but
osmotic membranes of cellulose, and dead cells, as in seed coats, are now
well known.
When dry seeds with osmotically active coats are placed in moist soil,
they absorb moisture with great power, such forces as imbibition, capillarity,
surface force, hydration power of colloids, and osmotic pressure from
internal salts being involved. The total power of the seed to absorb water
is the resultant of all these forces working together. Opposed to these
internal forces are external forces which tend to prevent intake of water.
The surface forces of soils, capillarity, surface tension, adhesion, cohesion
of water in thin films, osmotic action of solutes, etc., are pitted against these
internal forces; and water movement into or out of the seed will take place
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OSMOTIC PHENOMENA
in accordance with the lack of equilibrium between the two resultant forces.
I t is an interesting fact that seeds will actually germinate in a soil that has
a moisture content below its wilting coefficient.
As soon as the seed begins to germinate the characteristic osmotic
mechanism for soil water intake is established. T h e view point is general
that the intake of water, its transfer through the plant body, and its distribution to the living cells, are processes involving osmotic action at various
places in the organism. Although some suggestions have been made
assigning to the root a rather passive rdle in water intake, it seems to me a
mistake to adopt the idea that the root is mainly a n anchorage organ, and
that water merely filters through the root under the negative pressure of
atmospheric evaporation and the cohesion of water. The osmotic delivery
of water by the root to the lower end of the cohesive water columns probably goes on at a rate determined by all of the physical and chemical conditions within the soil-root system, including the cohesive pull upon the
root cells. And if this does not keep pace with evaporation, the deficit in
the aerial portions becomes greater and greater, and the tension stronger,
until breakage of the column and permanent wilting ensues.
There is reason for believing that the drying power of the atmosphere,
which fluctuates greatly with changes in relative humidity, averages not far
from 1000 atmospheres. And this is the tremendous force which is responsible for the disturbance of moisture equilibrium in the leaves, generating in the cell walls bounding intercellular spaces, in the protoplasm itself,
and in the vacuoles of leaf cells, those imbibitional and osmotic forces
which lift the water column of the transpiration stream to the tops of even
the tallest plants.
T h e root hair itself is very well adapted to its function as an absorbing
organ. As Miss Roberts has shown, the wall of the root hair is usually the
outgrowth of the middle lamella of the epidermal cell. I t is lined inside
by a cellulose layer deposited upon it by the living protoplasm. T h e external layer, therefore, is largely calcium pectate, and a pectin mucilage
brings it into most intimate contact with the particles of soil minerals. As
this pectic material is a hydrophilous colloid, it is highly adapted to imbibe
surface moisture from the particles of soil with which it is in contact. I t is
worth while to note in passing that the whole problem of water intake and
water outgo from the plant begins and ends in imbibition, imbibition of
water from the protoplasm and vacuole by the exposed cell walls in leaf
interspaces, and imbibition of water from the surface of soil particles by the
pectic walls of absorbing root hairs.
There are several main problems connected with the osmotic phenomena
of plant life which are worthy of consideration. These are, the nature of
osmotic pressure itself, the cause of semipermeability in membranes, and the
cause of unilateral movement of water across the semipeyeable septum.
Before considering the nature of osmotic pressure, the power of the plant to
adjust itself osmotically to its environment should be mentioned. I t has
been fully established by the work of Hill, Drabble and Lkabble, and Miss
Roberts that the plant increases or decreases the osmotic concentration of
its cell sap part. passu with changes in the environment which determine the
scarcity or availability of water for the plant. Miss Roberts work is
especially significant as related to soil moisture, since she made her determinations on root-hair producing cells. She found that the osmotic pressure
of the root hairs is maintaineda few atmospheres in excess of that of the
surrounding medium. Such changes in the root cells are accompanied by
corresponding changes in the osmotic mechanism more or less throughout

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the entire plant, so that appropriate Gsmotic gradients are constantly maintained. With plants growing in the soil, a similar gradient for moisture
intake exists. At the wilting coefficient the soil withholds water from the
plant with a force of about four atmospheres, while the usual osmotic concentration of the sap of root cells of land plants is seven or eight atmospheres.
I n desert regions, of course, these pressures would run much higher. Under
normal field conditions, therefore, the pressure gradient in ordinary mesophytic plants should run from four to eight atmospheres, as the fluctuation
of the water withholding power is from zero to four atmospheres. At the
wilting coefficient the effectiveness of the intake gradient is lost because the
rate of transfer of moisture from soil particle to soil particle toward the
plant becomes entirely inadequate. And now, even though the plants
osmotic pressure rises rapidly, it does not succeed in securing the needed
moisture, because it is trying, as it were, to draw water from a dry well.
Permanent wilting must quickly ensue in most cases after the soil reaches
the wilting coefficient.
T h e main criticism to be made of the current discussions of the nature
of osmotic pressure is that they attempt to simplify too much a process that
must, like other life processes, be complex. We may attribute osmotic
pressure to the difference in free energy of the solvent and solution, or to
the kinetic energy of the solute acting on the semipermeable membrane. I t
seems to me more appropriate to consider osmotic pressure a complex
force, the resultant of numerous factors. T h e free energy of the solvent,
the kinetic energy of the solute, the chemical forces of hydration, especially
in more concentrated solutions, and the attraction of ions in solution for
particles of water charged oppositely while passing through the differential
septum, are all factors in producing osmotic pressure. I n the plant this
complex force is smallest in the root hairs, and progressively increases to the
farthest limits of water transfer.
T h e most important problems in connection with osmosis centre in the
nature and causes of semipermeability in membranes. T h e sieve theory,
the various solubility theories, the surface tension theory, and the hydrone
theory are all well known. Every one of them has some supporting
evidence, but all of them have their limitations, and prove inadequate to
account for the known facts.
Armstrongs hydrone theory has come particularly to my attention in
connection with my own work. The assumptions which are made in this
theory as to the character of semipermeable membranes and the salts which
they exclude from passage are of such a nature that they seem to me to
necessitate the fundamental likeness in behaviour of all semipermeable
membranes. If only hydronated membranes can exhibit semipermeability,
and if hydronated membranes exclude all hydronated or hydrolated salts,
then all membranes of this kind should have identical behaviour. The
variability in behaviour of the natural semipermeable membranes, then, is
a challenge to the hydrone theory.
Again, it is not possible to attribute semipermeability to some particular
chemical substance in the membrane, like tannin, or suberin, or cutin, etc.
Many different kinds of membranes are semipermeable, and no single substance, or class of substances, is common to them all. When copper
ferrocyanide and other chemical precipitates, cellulose cell walls, or suberised cellulose, gelatin tannate, celloidin, cutinised tissue, parchment paper,
and living protoplasm all exhibit semipermeability, it is certain that this
peculiar property does not depend on chemical composition, but must be
related rather to the structure of the membrane.

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OSMOTIC PHENOMENA

There is one property which all of these membranes have in common.

They are all colloidal gels. I t is my opinion that any theory of semipermeability should frankly recognise this property, and be couched in the
language of the physics and chemistry of colloidal matter.
Water can undoubtedly penetrate both phases of the colloidal gel ; but
salt molecules attempting to penetrate the membrane would probably be
prevented from travelling through the disperse phase by surface tension and
other surface forces on the contact surfaces between the disperse and continuous phases. If so, they would have to travel in the continuous phase.
Recently Free has proposed a theory of protoplasmic permeability based
upon this conception. H e assumes that protoplasm is made up as a twophase system in which the phases differ mainly in their percentage of water
content, arranged as colloidal globules dispersed in a colloidal medium.
These two phases are supposed to be so related that interchange of water
between the two can occur accordingly as changes in the physics or chemistry
of metabolism, or of the environment, necessitate. By such changes, the
colloidal globules may increase in size while the continuous phase decreases ;
or, vice versa, the globules may become smaller while the continuous
phase separating them increases in thickness at their expense. Conceivably
the globules might reach such a size that the continuous phase would remain
as the thinnest possible films between them, or they might enlarge until the
gel is reversed, the discontinuous phase becoming a continuous medium,
and the former continuous phase becoming discontinuous.
Changes in permeability and semipermeability would be related to these
phase interchanges. Semipermeability would exist when the continuous
phase existed in such thin films that molecular diffusion through it was
impossible. Adequate provision is found here for natural differences in
membranes, for they could hardly have identical behaviour unless the
colloidal material were identical in kind. Two different kinds of membranes
could exhibit differences in reaction toward the same solution, and one
would not expect to find semipermeability behaviour identical in all membranes.
I t seems clear to me that we must have a theory of semipermeability that
recognises the nature of the membrane as Frees theory does. Living
protoplasm is an amphoteric substance with an isoelectric point; and it
may react as an acid or base depending on hydrogen ion concentration. As
it is played upon by various solutes, or by temperature, or light, changes in
the permeability or semipermeability relations would undoubtedly occur
according to the nature of the solutes, and the effects of other factors on the
aggregation in the colloidal system. In non-living membranes like seed
coats, the gels are much firmer, more difficult to change, and the characters
more stable than in living protoplasm. But even here one might expect
changes from permeability toward semipermeability, or vice versa, with
reagents having powerful effects on colloidal aggregation. In the problem
of soil-moisture intake by roots, of course, we are dealing with a very delicate
protoplasmic gel.
Finally, the movement of water through the membrane offers an interesting problem. When the osmotic cell is completely surrounded by the
solvent, like an enclosed sack, and if the membrane is elastic, one can
think of the pressure moving the membrane outward in every direction to
include more water, rather than water moving in through the membrane.
I t is the membrane that moves. But in the living plant osmotic and imbibition forces set up a unilateral movement which has not been satisfactorily
explained. The suggestions of Pfeffer as to protoplasmic differences on

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opposite sides of the cell, and of differences in concentration on opposite

sides of the vacuole are too well known to need discussion. Rhythmic
pulsations of the osmotically active cells cannot be assumed.
A suggestion which merits attention has recently been made by Loeb.
In amphoteric membranes like the protoplasm of root hairs, and of vacuolate
cells generally, the opposite sides of the membrane may be oppositely
charged. If they were, Loeb points out that positively,charged water particles
would be driven through the membrane from positive toward negative side,
where they would lose their charge after completing passage. The loss of
charge allows other positively charged particles to follow; and if a whole
series of cells one after the other had such properties, the series would set
up a unilateral current of water; in the case of roots let us assume toward
the open tracheze of the vascular system. At least this is one of the
possible factors in addition to osmotic pressure gradients, and might be responsible in part for the removal of water from the cortical cells into the open
tracheae. Electric exosmose has been suggested to account for the glandular
secretion of water from cells of high osmotic concentration into ducts with
lower concentrations. It can just as easily be a factor in the secretion of
water into the tracheze of roots.
The plant and its whole environment may be looked upon as a system
in which atmospheric evaporation is the chief disturber of moisture
equilibrium. In response to this disturbance, imbibitional and osmotic
forces are set up which reach back from leaves to roots through cohesive
water columns. These water columns are fed from below by an active root
system which delivers its water by osmotic action and possibly electric
transfer of water. The water is imbibed by the hydrophile colloids of the
root hairs, from the surface of contiguous soil particles, which in turn
exert surface tension, surface, and capillary forces to draw moisture from
particle to particle toward the plant. This disturbance finallp reaches to
the ground water surface, which slowly lowers as the result of this upward
movement of water during periods of no precipitation, but is maintained by
precipitation from the same atmosphere which, when dry, disturbs the water
equilibrium.
I am fully aware that there are many points which need investigation
before an entirely satisfactory picture of the osmotic phenomena of plant
life can be drawn. They are very complex phenomena, perhaps much
more complex even than the considerations here brought forward indicate.
But with a working hypothesis such as I have briefly outlined, we should
be able by appropriate investigations to throw much light on those more or
less obscure regions of the problem.